rik) RA MALESIANA SERIES I - SPERMATOPHYTA Flowering Plants Vol. 7, part 2 INDEX TO REVISED FAMILIES Aceraceae. . . 4: 3,592 Epacridaceae . 6: 422 Pentaphragmataceae. 4: 517 Actinidiaceae s.str. . 4: 37 Ericaceae . 6: 469 Pentaphylacaceae. . 5: 121 Aizoaceae . . ». 43267 #£Erythroxylaceae . 5: 543 Philydraceae’ . .) >. asi Alismataceae . . . 5:317 #£Fagaceae . A 7: 265 Phytolaccaceae . . 4: 229 Amaranthaceae . 4: 69,593 Ficoidaceae 4: 267 Pittosporaceae . . 5: 345 Ancistrocladaceae . 4: 8 _ Flacourtiaceae 5: 1 Plumbaginaceae . . 4:3 107 Aponogetonaceae 4: 11, 7: 213 Flagellariaceae 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 Preece ite. 7h} 1172)! Bixaceae s. str. 4: 239 Goodeniaceae 5:.335. Primulaceae : . yy iGsalis Burmanniaceae . 4: 13,592 Haemodoraceae 5: 111 ‘Proteaceae... =e Burseraceae 5: 209 MHaloragaceae 73.239 ‘Punicaceae.’' .. \s.) Waleeereee 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. . 2. 22 Ss107 Campanulaceae 6: 107 Hydrocharitaceae. 5: 381 Salvadoraceae. . . 4: 225 Cannabinaceae 4: 223 MHydrophyllaceae . 4: 207 Sarcospermaceae. . 4: 32 Capparidaceae 6: 61 Icacinaceae Ts:, 1 “Saururaceae: .)) [aaa ua Caprifoliaceae . 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 Sparganiaceaae . . 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 it. Sasa Cochlospermaceae 4: 61 Martyniaceae . 4: 216. | Stylidiaceae, = <1) easel Combretaceae 4: 533. Molluginaceae 4: 267 Styracaceae . . . 4: 49 Connaraceae . . . 5:495 Mboringaceae . 4: 45 Thymelaeaceae 4: 349, 6: 1 Convolvulaceae . 4: 338,599 |Myoporaceae . 4: 265 Trigoniaceae . . . 4: 59 Corynocarpaceae 4: 262 #Myricaceae 4: 277 Turneraceae . .. (naeroso Crassulaceae . 4: 197 Najadaceae 6: 157 Typhaceae §. °') = Saeasa4s Datiscaceae 4: 382 Nyctaginaceae 6: 450 Umbelliferae . . 4: 113, 595 Dichapetalaceae . 5: 305 Nyssaceae. 4: 29 Valerianaceae. ~ 22 apeasi255 Dilleniaceae 4: 141 Ochnaceae 7: 97 Violaceae Pee ninth: Dioscoreaceae 4: 293 Oxalidaceae 7: 151 Xyridaceae . . 4: 366, 598 Dipsacaceae 4: 290 Papaveraceae . 5: 114 Zygophyllaceae . . 4: 64 Droseraceae 4: 377 _Passifloraceae . 7: 405 Elatinaceae 4: 203 Pedaliaceae 4: 216 ALL - FAGACEAE (E. Soepadmo, Bogor) Monoccious trees or rarely shrubs, in Mal. evergreen, sometimes buttressed or with stilt-roots; growth mode flushwise, with perular buds. Hairs simple or stellate or fasciculate, rarely with resiniferous colleters, or scales on pits on the underside of the leaf. Leaves simple, spirally arranged, rarely in whorls of 3 or distichous, sometimes crowded near the top of each flush, penninerved, in Mal. entire or rarely crenate or sinuate. Stipules present, caducous or rarely rather long persis- tent, rarely interpetiolar or peltately attached. Inflorescence a cyme or a simple or branched spike, bracteate, 3, 9, androgynous (with the 9 flowers borne on the lower part) or mixed. Flowers unisexual or functionally so. — 3 Flowers: solitary or in dichasial clusters of 2-30 along the rachis, sessile or pedicelled; perianth campanulate or tubular, 6(—9)-lobed, or irregularly incised; stamens (4-)6-12(—90), filaments filiform, long exserted, free or rarely connate at the base; anthers linear to reniform, dorsi- or basifixed, lengthwise dehiscent; pistillode absent or present, densely hairy. — 2 Flowers: sessile, solitary or in dichasial clus- ters of 2-15, surrounded by a cupule; ovary inferior, 2-6(—9)-celled, usually hairy; ovules anatropous, 2 per cell, apical and collateral; perianth usually regularly 6—-lobed, sometimes poorly developed; staminodes 6-12, or absent; styles as many as ovary cells, terete, rather short, conical or tongue-shaped; stigmas capitate, punctiform, or covering the inner surface of the styles. Cupules solitary or in dicha- sial clusters, often woody, rarely reduced or absent, from saucer- or cup-shaped to enclosing the fruit, indehiscent or splitting into 2-8 or more + equal segments, rarely consisting of 2 free segments, variously muricate, spiny, squamose, or with concentric or spiral lamellae, very rarely almost smooth. Fruit an indehiscent nut (achene), 1—3-celled, sometimes falsely multiseptate, rounded or sharply 2—3—an- gular. Seed one, exalbuminous; embryo large; cotyledons large, flat-convex, plicate or ruminate; germination epigeal or hypogeal. 266 FLORA MALESIANA [ser. I, vote? Recent distribution. Seven genera with possibly c. 700 spp., the majority on the northern hemisphere. In the Old World the distribution extends southwards from 62°N in Scandinavia southheastwards to Kashmir and then northeastwards to the Sea of Okhotsk at c. 55°N. In Africa, Fagaceae are confined to the northern rim in the western Mediterranean region. In Asia Fagaceae are absent from the dry parts of the Middle East, from the Deccan Peninsula and Cey- lon, from the desert and colder parts of China, from Manchuria, and from the extreme northern parts of Japan. In America, the distribution extends from Canada and the United States south- wards to Central America, as far south as a few scattered localities in Columbia, in South Ameri- ca. On the southern hemisphere, Fageceae are present in Malesia, in the scarce wet parts of East Australia, in Tasmania, New Caledonia, and in New Zealand (otherwise absent from Pacific islands); in South America they occur from Fuegia and Staten I. northwards to Argentina and on the western slopes of the Andes in Chile up to 33°S. Fig. 1. In Malesia: 5 genera (Castanea and Fagus absent) with c. 180 spp., from the Malay Peninsula eastwards to d’Entrecasteaux Is., New Britain, and the Louisiades to c. 153°E; they are rare in the seasonally dry parts of Central and East Java and absent in Madura and Kangean Is., the Lesser Sunda Is., the southern Moluccas (Tanimbar, Key and Aru Is.), New Ireland, the Solomons', and Woodlark I. Notable distributional features are that Castanopsis and Lithocarpus occupy almost exactly the same range in the whole world (fig. 10 and 16), and that Fagus and Nothofagus are exclusive northern and southern hemisphere counterparts (fig. 2). Fossil distribution. Many macrofossils (leaf impressions and fossil wood) have been described and in addition there are many fossil pollen records, especially of Nothofagus which possesses a characteristic pollen quite distinct from that of the other genera. These fossils occur through the Tertiary going down in the time-scale to the Upper Cretaceous (fig. 2). The oldest records on the northern hemisphere, based largely on macrofossils, are: Dryophyl- lum from Greenland, North America (Wyoming and Colorado) and Europe (W. Germany, Bohemia, Silesia) (see p. 403), Quercus antiquata from Utah, Q. ? fraxinifolia from Montana, a Fagoxylon from Japan, and two species of Fagophyllum from Vancouver I. On the southern hemisphere only Nothofagus occurs and occurred and is described from both macro- and micro- fossils, the Upper Cretaceous records being located in Seymour I. (Antarctica), New Zealand and SE. Australia. Tertiary records outside the recent range of living Fagaceae include southern Central Asia, the SE. Deccan Peninsula (Castanoxylon), almost the whole of Australia, McMurdo Sound in Antarctica (77°S), the western part of North America extending to 65°N, and Iceland. It is clear that the warmer and moister Upper Cretaceous and Tertiary climates permitted their growth in both hemispheres far beyond their present range and that the Pleistocene Ice Age has thrown them back to lower latitudes. Besides, desiccation of several continental areas were unfavourable for their survival. See also the discussion under the genera. Literature: BERRY, Tree Ancestor (1923) 119-145; CHANDLER, Publ. Brit. Mus. Nat. Hist. (1964) 1-151; CRANWELL, Ancient Pac. Fl. (1964) 87-98; CRANWELL c.s. Nature 186 (1960) 700- 702; Dawson, Trans. R. Soc. Canada 11 (1893) 53-73; LA Motte, Mem. Geol. Soc. Am. 51 (1952) 105-107, 150-151, 160-162, 205-206, 243, 285-301; SEWARD, Plant Life through Ages (1931); VAN STEENIS, Blumea 19 (1971) 65-98, with bibl. on Nothofagus; Stopes & Fumi, Phil. Trans. R. Soc. Lond. B, 201 (1910) 1-90. Ecology. Climate. — In Malesia most species of Fagaceae shun the seasonal climate and therefore they are almost absent from the large seasonal belt in Central and East Java and the Lesser Sunda Islands. Several species, e.g. Lithocarpus sundaicus, Quercus lineata and Q. gemelli- flora do, however, occur on some mountain summits or slopes in Central and East Java subjected 1) Quercus guppyi F. v. M. from the Solomon Is. has appeared to be a Litsea (Laur.), see FORMAN, Kew Bull. 19 (1965) 461. 1972] FAGACEAE (Soepadmo) 267 to a feeble seasonal climate. A similar tolerance of minor seasonal climatic conditions is shown by Castanopsis buruana in some Moluccan islands. Malesian Fagaceae do not possess resistance against fire and do not regenerate under pyro- genous conditions. In areas cleared by fire they can maintain themselves in the (sharp) forest borders surrounding such cleared areas; this has e.g. been observed in East Java, the Sula Is., Buru, and New Guinea. Soil. — Fagaceae grow on a great variety of soils, including clay, sand, etc. and most species seem to be rather indifferent to soil-type. However, several species are restricted to specialized soils. Species characteristic of peat swamps and wet kerangas (podsolized sands) are Castanopsis fulva and Lithocarpus dasystachyus. Species which seem to be confined to kerangas soils are Quercus kerangasensis, Q. percoriacea, Castanopsis foxworthyi, C.borneensis, and Lithocarpus luteus. Finally, two species seem to be char- acteristic of soils derived from ultrabasic bedrock, viz Quercus lowii and Lithocarpus hatusimae. Malesian Fagaceae generally shun calcareous soils; in passing it may be remarked that it is said that in New Caledonia Nothofagus is restricted to serpentine. Mycorrhiza. — All Fagaceae seem to live in symbiosis with ectotrophic mycorrhiza (of Agaricales), a feature which is generalized by SINGER & MorRELLO (Ecology 41, 1960, 549-551) and specifically studied by them in South America (Mycopath. & Mycol. Appl. 26, 1965, 129- 191).-They believe that the virility and social strength of Fagaceae, and other Amentiferae and Coniferae, may be due to these ectotrophs, facilitating settling of pioneers and providing intol- erance to other trees. Altitude. — In Malesia Fagaceae occur from sea-level up to 3350 m, predominantly between 600 and 1500 m. Species of Nothofagus occur between 750 and 3100 m; Trigonobalanus verticil- lata is found between 850 and 1500 m; and some species of Castanopsis, Lithocarpus and Quercus are known from lower elevations down to sea-level. Examples are: Castanopsis argentea is found not far from the shore in S. Bantam (West Java); C. acuminatissima occurs in Depok (West Java) at 90 m, in the Sula Is. and Buru at 100 m, and in the Buna Plain (E. New Guinea) at 300 m; C. costata is recorded from Lingga, Riouw and Banka at c. 50 m; C. fulva grows in the Lake Gardens and Weld Hill near Kuala Lumpur (Malaya) at c. 70 m; Lithocarpus elegans occurs in Templer Park (Selangor, Malaya) and Tasek Bera (Pahang, Malaya) at c. 100 m; L. platycarpus has been collected in Nusa Kambangan (South Central Java) at 25 m; Quercus subsericea is found on rocky sea-shores in Cave Rachado near Port Dickson (Negri Sembilan, Malaya) at 0-10 m, facing the mangrove. It is evident that the large destruction of Malesian lowland forests explains part of the present rare occurrence of Fagaceae in the lowland. Flowering. — Flowering occurs irregularly, but generally takes place soon after the dry season together with the young flush. There is a more or less strict periodicity in flowering, which is, although the individual flowers are small and insignificant, quite showy for its simultaneous profusion, particularly if coinciding with an outbreak of leaves, which are pale yellow-green, or red as in Nothofagus and Lithocarpus scortechinii. In Castanea, Castanopsis, Lithocarpus and Trigonobalanus verticillata with their erect 3 catkins pollination is effected by insects; a disagree- able, spermatic smell has been reported in Castanopsis and Lithocarpus. In Quercus and Trigono- balanus doichangensis with their pendulous 3 inflorescence and capitate or ligulate stigmas, pollination is by wind, as it is in Fagus and Nothofagus. The anthesis of 3 flowers is of very short duration and the ¢ inflorescences are soon shed as a whole. Fruiting. — In most species fruits ripen within 3-6 months after flowering. Though fruiting is often very profuse, seedlings are sometimes surprisingly scarce. This is in part due to the fact that the fruits, which are very rich in starch and/or oil, are greedily devoured by animals. Also an appreciable percentage of full-grown fruits appear to be barren. A third factor is that fruits fallen on or in the litter on the forest floor are quickly attacked by fungi. In Trigonobalanus a large percentage of fruits falls an early victim to insects. For successful germination it seems necessary that the nuts must be buried in the soil, in cracks, holes made by earth-worms, or covered by soil-wash, or carried by ground animals. 268 FLORA MALESIANA [ser. I, vol. 7? Hypogeal germination is found in Castanea, Castanopsis, Lithocarpus and Quercus, whereas in Fagus, Nothofagus, and probably in Trigonobalanus, germination is epigeal. Dispersal. — Since the fruits are not provided with any special means of dissemination and are generally fairly large, mostly rounded, and fairly heavy, dispersal in this family must be slow. The fruits are not capable to float either in freshwater or in the sea. Not far from river-mouths the tropical beach is often strewn with acorns, but these are barren, hence buoyant and drifted downstream to the ocean (cf. VAN STEENIS, Trop. Natuur 29, 1940, 74). This is in complete accor- dance with the experiments and observations made by PreEest on Nothofagus in New Zealand (in GressiTT, ed.: Pacific Basin Biogeography, 1963, 415-424); he found that the fruits do not travel very far from the stands and are poorly fitted for dispersal beyond the forest margin. A similar observation was made by KALKMAN & VINK in New Guinean Nothofagus (Blumea 18, 1970, 101-102). This slow dispersal makes the distribution of the family most interesting, suggesting that their present and fossil distribution is an indicator of prime importance for the presence of land in former geological epochs. See also VAN STEENIS, Blumea 19 (1971) 74-76, 91. Fagaceae in Malesian vegetation. — Generally the species are not emergent trees, but an often dominant or co-dominant closed canopy. When the emergents (Dipterocarps, Podocarpus, Agathis, Altingia) have been removed for timber, the great number of Fagaceae show up in the remaining forests; DE VooGp observed this on Castanopsis in West Java, 30 years after selective logging of Altingia. A similar observation is made on Fraser’s Hill and Ulu Gombak (Malaya), where Fagaceae (namely species of Castanopsis, Lithocarpus, Quercus and Trigonobalanus) become dominant in the forests between 900 and 1200 m elevation after the removal of the big timber trees. In some islands, Fagaceae show a remarkable tendency to grow socially over immense tracts, and in Malesia particularly in New Guinea. Among the phanerogamous trees of the globe, Fagaceae come, in term of biomass, as a single group probably second only to Conifers. In Malesia, all five genera are (co-)dominant in the montane and lower subalpine zones, together with Lauraceae, Altingia, Schima, Engelhardia, and Podocarpus in Java and Sumatra, and in other islands also with Agathis and Phyllocladus. Such stands often contain more than one genus of Fagaceae and this has induced JUNGHUHN and MIQUEL to call these forests correctly as ‘Fago- lauraceous’. The seeds of Fagaceae need shade for their germination as do all trees of the very mixed rain-forest. But we may guess that the litter of Fagaceae, during decay, produces (allelo- pathic) substances which may, together with certain fungi, prohibit or restrict the growth of other forest trees, or at least favour a better regeneration of their own kinship. Undergrowth in pure Fagaceous forests is often surprisingly poor. Trigonobalanus verticillata reinforces its stand by soon producing additional stems next to the first one, and in Castanopsis acuminatissima suckers are produced by the root system which surround the mother tree as a skirt in a circle with a radius of several meters; the Sundanese in West Java pungently call it ‘riung anak’, a parent surrounded by its children. Fallen trees of Nothofagus in mossy forest, e.g. of N. pullei, do produce side-shoots which grow vertically and also produce roots; this may lead to thickets of saplings but WOMERSLEY, who observed this, has no idea how many, if any, eventually reach mature tree size. As a whole, regeneration is almost entirely by seed. The main accounts of the gregarious growth are the following: Castanopsis. — In West Java C. acuminatissima, C. argentea, C. javanica, and C. tungurrut are the commonest constituents of the mid-montane rain-forests. JUNGHUHN recorded (Nat. Tijd. N. I. 15, 1858, 23-38) that C. acuminatissima occurred dominant on Mt Malabar (W. Java) and made up for 80 to 100 % of the stands, at c. 1500 m. A similar dominance of this species is report- ed by Steup (Trop. Natuur 23, 1934, 63) from Donggala and Gorontalo (Central and North Celebes), between 800 and 1200 m, and it is said that the species is in constant association with Agathis, Eugenia and Phyllocladus. In the Moluccas, it occupies a similar position, as recorded first by TEYSMANN (Nat. Tijd. N. I. 37, 1876, 97) and later by BLOEMBERGEN from the Sula Is. They stated that G. Aha near Fowata is even called after this tree. In the Sula Is. it often occurs 1972] FAGACEAE (Soepadmo) 269 in groves alternating with pyrogenous grassland (Ausu-kusu), subjected to occasional shifting cultivation. Whether the production of the edible fruits contributes to the sparing of trees or that this must be ascribed to its suckering capacity which would promote gregariousness is not known with certainty, but it would well agree with the situation found in many places in New Guinea (West and East) where C. acuminatissima is frequently found gregarious in conjunction with pyrogenous grasslands, according to Brass and LANgE-PooLr. BrRAss reported (J. Arn. Arb. 22, 1941, 291, 299, 303, 309) that Fagaceous forest forms practically pure stands on the broader ridge crests above Bernhard camp, ranging up and down the spurs in the rain-forest. The main constituents of this forest are: Lithocarpus celebicus, L. megacarpus, Castanopsis acuminatissima and Engelhardia. The forest is about 25 m tall, and the oaks grow well apart with a few substage trees; they produce a rustling ground litter which is slower to decay than that of most rain-forest leaves, accentuated by a characteristic undergrowth of the fern Syngramma hookeri. In the defor- ested area near the Balim valley, a gallery strip of forest (c. 10 m tall) sometimes consisted of a pure stand of C. acuminatissima with thick boles and spreading branches and a ground cover consisting of the fern Humata mecidioides. At lower levels on the Bele River camp chestnuts and oaks (Lithocarpus schlechteri, L. rufovillosus, and L. lauterbachii) and their associate Engel- hardia are common in the forest as subsidiary trees of 25-30 m tall, and survive as relics on cleared lands. At altitudes between 2200 and 2350 m, these trees form limited pure stands on the broad crests of spurs, but none is encountered much above the limit of cultivation. These forests may be distinguished by their open canopy dominated by species of Nothofagus, the dry appear- ance caused by the slippery ground litter of cutinous dead leaves and by the absence of terres- trial mosses. Due to the fact that the people of several village-groups extended their communal clearings, these forests were gradually disappearing. Most of the crop-gardens were established on new lands strewn with the trunks of large trees of Nothofagus, brought down through the burning of their bases. Other trees killed by fire or ring-barking stood as they had grown in the original forest. BRAss is convinced that Castanopsis and Lithocarpus are sure indicators of superior arable lands. On Mt Tafa and the Wharton Ranges, Brass found (J. Arn. Arb. 22, 1941, 335), above the mixed rain-forest, a similar Fageceous zone situated between 500 and 2200 m, in which he found nearly pure stands of Castanopsis acuminatissima associated with Lithocarpus, Engelhardia and Araucaria. In the Vanapa Valley, however, much of this Fagaceous forest had been cleared and converted into grassland. LANE-POOLE reported a similar situation from the Buna Plains and the Hydrographer’s Range (For. Res. Terr. Pap. & New Guinea, 1925, 22, 34, 79). In Malaya, C. acuminatissima, in association with C. lucida, C. megacarpa and several species of Lithocarpus, is the commonest constituent of the mid-montane forest, especially at Fraser’s Hill between 900 and 1200 m. Selective felling of the dominant trees (Dipterocarps, Agathis and Podocarpus) seems to favour the growth and spread of these Fagaceae. As in the other islands, it is only C. acuminatissima which produces numerous suckers around the base of its trunk. Lithocarpus & Quercus. — Species of these two genera are dominants or co-dominants in the mid-montane forests throughout Malesia, and the upper limit of their zone somewhat exceeds that of Castanopsis. On Mt Kinabalu (N. Borneo), L. turbinatus and L. lampadarius are dominants in the forests between 1800 and 2400 m, while L. havilandii is the only species surviving up to 3000 m, mixed with Leptospermum flavescens, Dacrydrium beccarii, and Phyllocladus hypophyllus. At this altitude it grows to not more than 5-10 m height, with spreading branches and rufous foliage, sticking out of the grey-green and glaucous crowns of Leptospermum and Phyllocladus. In New Guinea (Vogelkop) and adjacent islands (Normanby, Misima, Sudest and Rossel Is.), Lithocarpus vinkii occurs from sea-level up to 1800 m, commonly above 700 m, and on some ridges it may form pure stands. Nothofagus. — BRAss stressed the preponderant position which Nothofagus species occupy in the mountain forests of New Guinea (see VAN STEENIS, J. Arn. Arb. 34, 1953, 314; Blumea 19, 1971, 71). Its range lies between (750-) 1000 and 3100 m, but c. 90 % of the collections are made between 1750 and 2850 m, only N. flaviramea, N. rubra, and N. starkenborghii descending below 270 FLORA MALESIANA [ser. I, Von? 1000 m down to c. 750 m, the first even to 600 m in New Britain. The reddish flush is characteris- tic, but its appearance varies in time in different localities. Of almost all species dominance is reported, or co-dominance (often noted with Lithocarpus, Castanopsis, hoop pine), but VINK & KALKMAN Observed that the proper Nothofagus zone is often situated higher than that of the other Fagaceae. Mature size on broad ridges, gentle slopes and valleys is mostly 40 m or over by a diameter of c. 1 m, but on narrow and steep ridges dwarfing to stunted shrubs 2—5 m high are observed in N. perryi, N. carrii, and a few others. Fig. 8. Trigonobalanus verticillata. — On Mt Kinabalu in Sabah and the Hose Mts in Sarawak this species is gregarious at 1000-1500 m, in association with Agathis, Dacrydium elatum, and Podo- carpus imbricatus. In Malaya, it is so far only known from Fraser’s Hill, where it is also gregar- ious along the road leading to and near the waterfall, at 850-1200 m, associated with Agathis dammara, Podocarpus imbricatus, Schima wallichii, several species of Lauraceae and species of other genera of Fagaceae. Uses. In comparison with the importance of oakwood, oak tannin, beechwood, and chestnuts in the northern temperate regions, the economic value of Fagaceae in Malesia is slight, Notho- fagus obviously excepted. The wood of Castanopsis, Lithocarpus and Quercus is often very hard and difficult to work, although it is sometimes rather beautifully and evenly grained. It is very liable to splitting, and seems rather unfit for construction purposes, for which it is occasionally used in Indonesia. Records of durability vary with the species. Some New Guinean species are supposed to be promising for furniture and veneer, but tests have not been made, and species may differ considerably in properties. According to field notes several Nothofagus species, notably N. starkenborghii, N. perryi, N. pullei, and N. rubra, possess excellent very resistant hardwood, used for bridge constructions and locally used on a large scale also for general constructions under roof. The Papuans in East New Guinea plant various species (NV. grandis, N. pullei, N. perryi) in their native homesteads, around their villages, along tracks and around their garden lands, it is said for ornamental purpose, but probably also intended for later use. For this purpose they collect seedlings in the forest. In Borneo, Dr. J. A. R. ANDERSON informed me, that logs of some species of Castanopsis, Lithocarpus, Quercus, and Trigonobalanus verticillata are recently being tried for mushroom cultivation. This trial was apparently inspired by the success of mushroom cultivation on wood of Quercus species in Japan and made possible by the occurrence of closely allied edible mush- rooms on Mt Kinabalu. The fruits of some species of Castanopsis (e.g. C. argentea, C. costata, C. inermis, C. javanica, and C. tungurrut) are widely consumed after cooking or roasting, just like chestnuts in Europe and other temperate regions. The bark in many species of Fagaceae contains appreciable amounts of tannin, but perhaps because of the hardness of the wood, there seems to be no commercial exploitation of it. See HEYNE (Nutt. Pl. 1927, 535-543) and BurRKILL (Dict. 1935, 486-489, 1849-1859) for details. Cultivated. On the summit of Mt Pangrango, West Java, at c. 3010 m, above the mountain garden Tjibodas, a specimen of Fagus sylvatica was introduced by the famous Curator TEYSMANN, about 1840, probably grown from seed, together with other European vegetables and fruit trees for trial. The specimen was a densely branched shrub c. 114 m high; it was never bare and did not flower. Its morphology and anatomy was studied by CosTER (Ann. Jard. Bot. Btzg 35, 1925, 105-119, t. 7-10). He characterized it as a typically alpine dwarf form, the leaves having half the normal size, developing in flushes but not all simultaneously. The plant produced annual rings and had in the pith and xylem great quantities of starch which did not diminish noticeably when new leaves unfolded. The difference between ‘sun’ and ‘shade’ leaves was slighter than in Europe. Shortly after the war the plant died on account of a local fire. In Hakgalla Garden in Ceylon, at montane altitude, Fagus sylvatica also did not flower and remained poor; it seems not adapted to a tropical mountain climate. Castanea sativa has since 1894 been cultivated at Tjibodas mountain garden, West Java, at c. 1400 m; it was reported by HEYNE (Nutt. Pl. 1927, 537) to produce fruit satisfactorily. 1972] FAGACEAE (Soepadmo) 271 Morphology. Habit. — In Malesia most species of Fagaceae are trees of medium to large size, rarely shrubs of c. 2-5 m tall (Lithocarpus orbicularis, L. oreophilus, Nothofagus pullei). Buttresses and stilt-roots are often present in several species of Lithocarpus and Quercus. Bark. — Most Malesian species of Fagaceae are readily recognizable in the field by their char- acteristic light-greyish, smooth and mottled bark; only a few have bark which is grey-brown and rather rough. In Nothofagus the bark is often peeling in large flakes. Lithocarpus, Nothofagus, Quercus and Trigonobalanus verticillata are very easily recognized by the same typical slash characters: a reddish-brown bark, easily detached from the smooth whitish wood, with numer- ous longitudinal faintly bluish lines about 14-114 cm long and 1 mm wide and slightly depressed. The opposite parts on the inside of the bark are correspondingly slightly elevated. This typical pattern is not found in Castanopsis. Terminal buds. — In Malesia the genera of Fagaceae possess well-developed terminal buds, which are conferted in many species of Quercus. The scales are either spirally (Castanopsis, Lithocarpus, Trigonobalanus) or decussately arranged (Nothofagus and most species of Quercus). Glandular scales are found in Nothofagus and some species of Lithocarpus. Stipules. — Castanopsis, Lithocarpus, and Quercus have extra-petiolar, caducous stipules, and Nothofagus possesses stipules which are distinctly peltately attached and in the flush they develop earlier than the leaves. The stipules in Trigonobalanus verticillata are extra-petiolar in the first few leaves of the seedling, and later become interpetiolar. Indumentum. — The hairs are singly distributed or in fascicles. Simple hairs and armed stellate hairs generally consist of a single cell, but they may be uniseriate, e.g. Castanopsis and Trigono- balanus verticillata. In Lithocarpus the arms of the stellate hairs emerge from a usually dark- coloured, occasionally bulbous central cell. In Nothofagus only one species (N. pullei) pos- sesses hairs. In Malesian Nothofagus the undersurface of the dried leaves is dotted with pale brown scales which on removal leave a small pit; the scales are actually dried up glands and their resinous exudate. Small, gland-tipped hairs often occur in all other genera, with the gland itself being 1—-co-celled. Colleters are resiniferous sausage-shaped trichomes which are only found in Notho- fagus, namely at the insertion of the stipules and at the adaxial bases of the perulae and the cupular lamellae. Both the leaf-glands and the trichomes exude resin which may cover the under- side of the leaves, the flush and the young cupules with a pale yellow waxy resin. Phyllotaxis. — This is distichous in Malesian Nothofagus (consequently the perulae are in 4 rows). Some species of Castanopsis also possess distichous leaves. In Trigonobalanus verticillata the first few leaves of the seedling are spirally arranged, followed by a series of decussate leaves, and finally taking to a whorl of three phyllotaxis in the older plants. Lithocarpus and Quercus have the leaves spirally arranged, and in Quercus they are usually crowded at the end of each flush. Leaves. — The leaves of Malesian Fagaceae are only very rarely not entire; they are at most crenate or sinuate near the upper half in some species of Castanopsis and Quercus, and remotely, shallowy crenate in a few species of Nothofagus. Domatia are not found in Malesian Fagaceae. Inflorescence. — In Fagus and Nothofagus the inflorescence is morphologically a 1- to many- flowered dichasium or cyme, either subsessile or long-peduncled, and axillary. In Fagus the 3 inflorescence is 2—20-flowered, while in Nothofagus it is 1- or 3-flowered. The ° inflorescence, on the other hand, bears 1-6 flowers in Fagus and 1-7 flowers in Nothofagus; the central flower in Fagus is always missing. In Malesian Nothofagus the 2 inflorescence bears not more than 3 flowers. In Castanea, Castanopsis, Lithocarpus, Quercus, and Trigonobalanus the inflorescence is an unbranched or branched spike, either pendent (¢ inflorescence in Quercus and Trigonoba- lanus doichangensis) or erect (3 and @ inflorescence in Castanea, Castanopsis, Lithocarpus and Trigonobalanus verticillata); the flowers are sessile and rarely short-stalked, being arranged in dichasial clusters of 2-30 or solitary along the rachis. Apart from the 3 and 9 inflorescences just described, there is also a mixed type as found in Lithocarpus, and an androgynous type in Casta- nea, Castanopsis, Lithocarpus, and Trigonobalanus. In an androgynous inflorescence the 2 flowers are situated at the basal part of the rachis. 272 FLORA MALESIANA [ser. I, vali Flowers. — A pistillode is present in the 3 flower of Castanea, Castanopsis, and Lithocarpus, and replaced by a cluster of simple stiff hairs in Trigonobalanus verticillata and some species of Malesian Quercus. In the 2 flowers 6-12 staminodes are found in the above-mentioned genera. In some species of Lithocarpus (L. ruminatus and L. turbinatus) from Borneo, they are rather well developed and exceed in length the height of the perianth-lobes. The stigma is capitate in Malesian Quercus and Trigonobalanus, punctiform in Castanea, Castanopsis, and Lithocarpus, or forming a broad stigmatic surface on the inner side of the style-arms (Fagus, Nothofagus and extra-Malesian Quercus). Cupules. — Generally present, solitary either in the axil of the leaf (Fagus and Nothofagus) or along a rachis (Castanea, Castanopsis, some Lithocarpus, Quercus, and Trigonobalanus), or in many species of Lithocarpus they are in dichasial clusters of 2-24 along the rachis. In a few species of Nothofagus from New Guinea the cupule is reduced to two minute loose flaps or even may be entirely absent. In Castanea, Castanopsis, Fagus, Nothofagus, and Trigonobalanus the cupules are completely covering the nuts, and eventually they are splitting irregularly or into regular numbers of valves or lobes. In Quercus and the majority of Lithocarpus, however, the cupules are not completely enclosing the nuts, but either saucer- or cup-shaped, and never splitting. In Lithocarpus javensis, L. maingayi, L. turbinatus and several more related species, the cupule is also completely covering the nut and fused to the latter, but in L. coopertus, L. encleisa- carpus, L. wrayi and related species, though the cupule is also enclosing the entire fruit, it is completely free from the latter. See BRETT (New Phytol. 63, 1964, 96-118), FORMAN (Kew Bull. 18, 1966, 385-419), HJELMaQvisT (Bot. Notis. Suppl. 2, 1948, 1-171), LANGDON (Bot. Gaz. 108, 1947, 350-371), SoEpADMO (Gard. Bull. Sing. 22, 1968, 364-368; Reinwardtia 8, 1970, 205-212) and VAN STEENIS (J. Arn. Arb. 34, 1953, 309-313) for further details on the nature, anatomy, and systematic and phylogenetic significance of the cupule. Fruits and seeds. — In Fagaceae the fruit is an achene or nut, with the pericarp fused with the perianth tube. In Lithocarpus and Quercus the fruit is always rounded in cross-section, and that of the other genera is variously rounded-triangular or sharply triangular (Fagus and Trigonobalanus verticillata) or even winged (Nothofagus). In Malesia there seems to be a very high degree of sterility among the fruits produced by many species. For an account on the anat- omy of the fruit wall see SogEPADMO, Gard. Bull. Sing. 22 (1968) 369. Embryology. So far no study has been made on the Malesian species of Fagaceae. HJELM- QVIST basing his observations on Quercus robur, Fagus sylvatica, and Castanea sativa concluded that the development of the endosperm and the embryo in Fagaceae is rather similar to that of Juglandaceae, but differs very sharply from that of Betulaceae. Literature: CONRAD, Bot. Mag. 29 (1900) 408; HsELMaevist, Phytomorph. 3 (1953) 377; Bot. Notis. 110, 2 (1957) 173-195. Wood anatomy. Vessels. — Distribution solitary, ring-porous, or dendritic; scalariform perforation plates frequently occur in Fagus and Quercus; spiral thickening rather frequent in Nothofagus, but absent in the other genera. Parenchyma. — Distribution apotracheal, paratracheal or rarely concentric. Rays. — Exclusively uniseriate in Castanea and Castanopsis, uniseriate to many cells wide in the other genera. Tracheids. — Present except in Fagus, distribution paratracheal. Fibers. — Without spiral thickening, very rarely septate (Nothofagus); pits mostly simple, rarely bordered (Fagus). Interxylary phloem, intercellular canal, raphides, oil or mucilage cells absent. Tyloses and cristals constantly present. From an analysis on the 16 diagnostic characters selected by METCALFE & CHALK (1950), as to the wood, Fagaceae appear closely related to Casuarinaceae and Juglandaceae, differing from both only by the relative more frequent occurrence of dendritic patterns of vessel distribution. Literature: DEN BERGER, Determinatietabel Houts. Mal. Fam. (1949), 36; CUTLER, Kew Bull. 17 (1964) 401-409; ibid. 21 (1967) 332-334; DADSWELL & INGLE, Austr. J. Bot. 2 (1954) 141- 1972] FAGACEAE (Soepadmo) 273 153; Janssonius, Mikrogr. Holz. 6 (1936) 359-468; Anat. Bestim. Jav. Holz. (1940) 84-85, 108-109; Key Jav. Woods (1952) 87, 113; METCALFE & CHALK, Anat. Dicot. 2 (1950) 1309-1315. Palynology. Observed under the light-microscope, 3 main types of pollen grains may be distinguished in Fagaceae, viz the Quercus-type (Quercus, Fagus, Trigonobalanus), the Castanea- type (Castanea, Castanopsis, Lithocarpus), and the Nothofagus-type (Nothofagus). In the Quercus-type the pollen is more or less spheroidal, with average size 30 by 20 yp, 3-col- porate, 3-colpate, or 3-colporoidate, exine more or less scabrate. In the Castanea-type the pollen is generally ellipsoidal (perprolate), always 3-colporate, with an average size 16 by 10 wu, and the exine is more or less smooth. Pollen of Nothofagus differs from the two previous types by being peroblate (horizontally fiat), average size 15-20 by 30-45 yw, 4-9-colpate, and the exine is distinctly echinulate. Within the Nothofagus main type there are three types, the fusca and menziesii types for extra-Malesian Nothofagus, and the brassii type for all Malesian and New Caledonian species which together form the subsection Bipartitae. In the last case pollen morphology agrees almost with taxono- mic distinction; there is even a small distinction running parallel with Triflorae and Uniflorae within the subsection. However, the fusca and menziesii types run across the main taxonomic subdivision of the genus and do not agree with the morphological classification, with no possibil- ity towards reconciliation. According to ERDTMAN (1952) the pollen of Fagaceae belongs to a type widely known in the Angiosperms, Nothofagus excepted. For fossil pollen, see under the heading fossil distribution. Main literature: VAN CAMpo & ELHAI, Bull. Soc. Bot. Fr. 103 (1956) 254-260; Cookson & Pie, Austr. J. Bot. 3, 2 (1955) 197-206; CRANWELL, Rec. Auckl. Inst. Mus. 2, 4 (1939) 175-196; ERDTMAN, Pollen Morph. & Tax. 1 (1952) 176-177; Bot. Notis. 120 (1967) 324-333; HARRIS, New Zeal. J. Sc. & Techn. B 37 (1956) 731-765; NAKAMURA, Res. Rep. Kochi Univ. 5, 21 (1956) 1-5; PLANcHAIS, Pollen et Spores 4, 1 (1962) 87-93; SPoEL-WALvius, Acta Bot. Neerl. 12 (1963) 525-532; WALKER & WITTMANN, Pollen et Spores 7 (1965) 457-464; WoDEHOUSE, Pollen Grains (1935) 373-382; YAMAZAKI & TAKEOKA, J. Jap. For. Soc. 41 (1959) 125-130. Chemotaxonomy. All Fagaceae seem to be accumulators of polyphenolic compounds; flavonols, leucoanthocyanins, catechins and gallic and ellagic acid occur frequently in leaves, fruits, bark and wood. In wood of Nothofagus species flavanones (naringenin), flavononols (aromadendrin, taxifolin), stilbenes (pinosylvin, resveratrol) and dihydrochalcones (nothofagin, konnanin) have been observed in addition (HILLIs & INouE, Phytochemistry 6, 1967, 59). Besides these low molecular polyphenolic compounds most members of the family contain rather large amounts of tannins. Oak bark, oak wood, oak gall-nuts and chestnut wood represent important sources of vegetable tannins. Oak trees elaborate predominantly galli- and ellagitannins in leaves, gall-nuts and wood and predominantly condensed tannins in barks. The same may be true for other members of the family. Hamamelitannin, a simple gallitannin. accompanies condensed tan- nins in the bark of Castanea sativa and Quercus rubra. Turkish tannin is the gallitannin of Aleppo gall-nuts. Castalgin, vescalgin, castalin and vescalin are complex ellagitannins of the wood of Castanea sativa and Quercus petraea. Pedunculagin is an ellagitannin of known structure from gall-nuts of Cental European species of Quercus. Besides polyphenolic compounds members of Fagaceae contain rather large amounts of pentacyclic triterpenes in barks, leaves and gall-nuts. Friedelin and the two epimeric friedelanols have been observed most frequently but derivatives of a- and f-amyrin as well as still other triterpenes do occur also. All species of the large genus Quercus seem to store quercit in leaves, bark and seeds; this represents a character of the genus Quercus. In conclusion it may be stated that Fagaceae resemble other Amentiferous families in tannin-accumulation, in the patterns of low-molecular phenolic compounds and in the tendency to produce conspicuous amounts of triterpenes. Chemistry also favours a relationship with the Hamamelidaceous and Rosaceous stock. General reference: HEGNAUER, Chemotaxonomie der Pflanzen 4 (1966) 141-155. — R. HEc- NAUER. 274 FLORA MALESIANA [ser. I, vol. 7? Chromosomes. Data on tropical and subtropical species are unfortunately scant. In various extra-Malesian species diploid numbers have been reported to be 20 or 24 (Castanea, Fagus, Quercus) and recently 28 in Quercus castaneifolia CAMUS (TUTAJUK et al., ). ARMSTRONG & WYLIE reported for Nothofagus 2n = 26. Independently KwitonG JonG (in litt.) and Dinc Hou found for Trigonobalanus verticillata2n = 44 (DiNG Hou also n = 22). As to chromosome num- ber Fagaceae have a great affinity with that found in other Amentiferous orders (Betulaceae, Corylaceae). Literature: ARMSTRONG & WYLIE, Nature 205 (1965) 1940-1941; DARLINGTON & WYLIE, Chromosome Atlas FI. Pl. (1955) 181-182; Guimpu, Rev. Bot. Appl. & Agr. Trop. 9 (1939) 176— 179; Dinc Hou, Acta Bot. Neerl. 20 (1971) 543-549; Jaynes, Forest. Soc. 8 (1962) 372-377; Sax, J. Arn. Arb. 11 (1930) 220-223; SuGcira, Bot. Mag. Tokyo 45 (1931) 353-355; TuTasuK & TURCHANINOVA, Dokl. Akad. Nauk Azerbajdzansk. S.S.R. 24 (1970) 47-50; VON WETZEL, Bot. Arch. 25 (1929) 257-283. Taxonomy. Historical review. — DUMORTIER (Anal. Fam. Pl. 1829, 59-60, ‘Fagineae’) was the first to recognize the family Fagaceae, and A. DE CANDOLLE (Prod. 16, 2, 1864, 1) was the first to frame the family in its present circumscription, using the name Cupuliferae. OERSTED (Kongl. Danske Vid. Selsk. Skrift. V, 9, 1871, 351) accepted DE CANDOLLE’s classification and used characters of the stigma, cotyledons, and germination to subdivide the family into three subfamilies: Quercineae to include Quercus L. and Cyclobalanopsis OrRstT.; Castanineae: Pasa- nia (MiQ.) OErRst., Cyclobalanus (ENDL.) OERsT. (including Lithocarpus BL.), and Castanea MILL. (including Castanopsis SPACH); and Fagineae: Fagus L. and Nothofagus BL. PRANTL (in E. & P. Nat. Pfl. Fam. 3, 1, 1889, 52), who first used the name Fagaceae and recog- nized the importance of the inflorescence, divided the family into two tribes, viz Fageae: Notho- fagus and Fagus; and Castaneae: Castanea (including Castanopsis), Pasania (including Lithocarpus and Cyclobalanus), and Quercus (including Cyclobalanopsis). O. SCHWARZ (Notizbl. Berl.-Dahl. 13, 1936, 1-22), who combined the characters already mentioned by OrRSTED and PRANTL, and added those of the cupule and fruit, came to the same subdivision as OERSTED. However, he recognized more genera than his predecessors, namely by splitting the genus Quercus into four segregates: Quercus, Cyclobalanopsis, Erythroba- lanus O. SCHWARZ, and Macrobalanus O. SCHWARZ; and Lithocarpus into three: Cyclobalanus, Lithocarpus, and Pasania. Among the recent authors who basically accept OERSTED’s classification, though they used different combinations of characters, are: MELCHIOR (in Engler, Syll. Pfl. Fam. ed. 12, 2, 1964, 50) and FoRMAN (Kew Bull. 17, 1964, 381-396); those who agree with PRANTL are: BRETT (New Phytol. 63, 1964, 96-118) and LuoNG (Abstract Thesis, Leningrad, 1965). On the basis of her pollen-morphological study, KUPRIANOVA (Rep. Sovjet Palyn. Publ. U.S.S.R. Ac. Sc. Moscow, 1962, 17-25) recently suggested to accommodate the genera Notho- fagus and Trisyngyne BAILL. into a distinct family, Nothofagaceae, and place this new family in the Euphorbiales. Her opinion is, however, completely unacceptable to us, as we found that the resemblance between the pollen of Nothofagus (including Trisyngyne) and that of Longetia (Euphorbiaceae) is highly superficial. Furthermore, the presence of a cupule containing fruits which are nuts, is the most important feature characterizing the family Fagaceae, and is nowhere found among the Euphorbiales. Moreover, the morphology of Nothofagus fits admirably in with that of Fagaceae especially Fagus and not at all in with that of Euphorbiaceae. It is true that the pollen of Nothofagus is morphologically very different from that of the rest of the family. Morphologically it is even so isolated, that Mr. J. MULLER admits that if only the pollen was known, a palynologist would be at a loss to place it in any family! However, the problem in Nothofagus stands not alone, as there are many genera or families in which the pollen is very high- ly variable morphologically, and yet they are accepted as belonging to a single larger taxon. Affinity of the Fagaceae. — According to ENGLER’Ss system of classification, Fagaceae together with other Amentiferous families, are considered as the most primitive group among the Angios- perms, and their catkin-like inflorescence, simple unisexual flowers, chalazogamic mode of 1972] FAGACEAE (Soepadmo) 275 fertilization, and marked interval between pollination and fertilization are considered as primi- tive characters, linking the group with some Gymnospermous ancestor. However, in the more recent systems of classification as advocated by Bessey, CRONQUIST, HUTCHINSON, TAKHTAJAN, and others, Fagaceae is considered as a relatively advanced family, closely related to Betulaceae and Corylaceae, and most probably derived from a Hamamelidaceous ancestor; they consider the simple nature of the inflorescence and flower as advanced characters, resulting from reduc- tion. For further discussion see: Bessey, Bot. Gaz. 24 (1897) 145-178; Ann. Mo. Bot. Gard. 2 (1915) 109-164; Cronaquist, Evol. Class. Fl. Pl. (1968) 170-171; HUTCHINSON, Fam. FI. Pl. 1 (1959) 192-193; Evol. Phyl. Fl. Pl. (1969) 147-150; TAKHTAJAN, Orig. Disp. Fl. Pl. (1969) 95-107. Generic delimitation. — Though in Malesia there is no serious difficulty in placing any speci- men into the proper genus, there is in fact still some problem in delimiting some genera of Faga- ceae, especially within the subfamily Castaneoideae. For example, the distinction between Castanea and Castanopsis is obscure, and it is merely to avoid a nomenclatural instability that the genus Castanopsis is in this Flora regarded as distinct from Castanea. The only difference existing between these two genera is that in Castanea there is always a constant combination of the following characters: 2 flowers situated at the basal part of the androgynous inflorescence, and the presence of 6 or more styles in each 2 flower. For further discussion see FORMAN, Kew Bull. 18, 3 (1966) 421-426. Fagus and Nothofagus can best be distinguished by the following combination of characters (VAN STEENIS, J. Arn. Arb. 34, 1953, 326-327, slightly emended): Fagus Terminal buds elongate-acute, composed of imbricate, non-glandular scales. Stipules never peltately attached, ligulate, flaccid, firmer towards the apex; insertion not surrounded by resinous colleters. Leaf not glandular-dotted, thin. Male flowers in a 2-20-flowered, long-pedun- cled, capituliform dichasium. Anthers not apiculate, not glandular. Pollen grains spheroidal, 3-colporate; polar diam.: equatorial diam. = 5 : 4; exine scabrate to reticulate. Cupular appendages seta-like, irregularly placed, without resinous colleters. Female dichasium without the central or primary flower. Anatomy of fruit wall of the Fagoid-type’. Endocarp hairy. Wood-rays several cells wide; wood-fibres with bordered pits. Nothofagus Terminal buds ovoid, consists of glandular and decussately arranged scales. Stipules mostly peltately attached, very rarely ligulate, scarious towards apex; insertion surrounded by sausage-shaped resinous col- leters. Leaf distinctly glandular-dotted, thick coria- ceous. Male flowers borne in a sessile or short-pedun- cled, 1—3—flowered dichasium. Anthers apiculate, often glandular. Pollen grains peroblate, 4-9-aperturate; polar diam.: equatorial diam. = 1 : 3 to 1 : 4; exine echinulate. Cupular appendages regularly placed or in distinct lamellae, very often with resinous colleters at their adaxial base. Female dichasium always with the central flower present. Anatomy of fruit wall of the Nothofagoid-typet Endocarp glabrous. Wood-rays 1-2 cells wide; wood-fibres with simple pits. The most important distinguishing characters between Castanopsis and Lithocarpus and be- tween Lithocarpus and Quercus are summarized in the keys. For details see FORMAN (Kew Bull. 1) For details, see SOoEPADMO, Gard. Bull. Sing. 22 (1968) 369-372. 276 FLORA MALESIANA [ser. 1; viola: 17, 1964, 381-396; ibid. 18, 1966, 421-426) and SoePADMo (Gard. Bull. Sing. 22, 1968, 356-357; Reinwardtia 8, 1970, 202-212). Infrageneric subdivision of the genera of Fagaceae. — In this Flora no attempt is made to evaluate the different views dealing with subdivisions. Those who are interested are referred to works by BARNETT (Trans. Proc. Bot. Soc. Edinb. 34, 1944, 159-204), A. Camus (Les Chénes, 1936-1954, 3 vols.), O. SCHWARZ (Notizbl. Berl.-Dahl. 13, 1936, n. 116, 1-22), and TRELEASE (Mem. Nat. Acad. Sc. 20, 1924, 1-255). Subdivision of the family Fagaceae. — Apart from the rather unique position of the genus Trigonobalanus, Fagaceae can best be subdivided into three subfamilies, as has been done by OERSTED and FORMAN on the basis of characters derived from the inflorescence and the flower, as follows: 1. Inflorescence spike- or catkin-like. 2. 3 Flowers always with a pistillode and with 10-12 stamens, with the anthers dorsifixed and versatile. 2 Flowers always with 10-12 staminodes; stigmas terminal and punctiform.Casta- nea, Castanopsis (incl. Chrysolepis HyELMQvisT), and Lithocarpus (incl. Pasania, Cycloba- lanus, and Synaedrys LINDL.) . . NEA. : . .. . . Subfam. Castaneoideae 2. 3 Flowers without a pistillode, with 6 Sy amenss aedliers basifixed. 2 Flowers without stami- nodes or rarely with 6 staminodes; stigmas capitate or covering the inner surface of the styles. Quercus (incl. Cyclobalanopsis, Erythrobalanus, and Macrobalanus) and Trigonoba- lanus FORMAN. .. . . .. . Subfam. Quercoideae 1. Inflorescence consisting a a (rave flowered Gicnasiiin Ragas and Nothofagus (incl. Trisyn- Syne) eich Lins ¢ Bae oe a cee ee Geely. faye Te: aS ab Eee Acknowledgements. — I wish to express my sincere thanks to Dr. M. JAcoss who has helped me considerably with the redaction of the manuscript for this Flora and with the checking of literature and specimens; to Dr. VAN STEENIS who took care of the account of ecology and vege- tation, and who is responsible for the new revision of Papuan Nothofagus provided here; to Dr. R. HEGNAUER for the chapter on chemotaxonomy. Three precursory papers preceded this revision, viz on Quercus (Gard. Bull. Sing. 22, 1968, 355-427, 18 fig., 16 maps), Castanopsis (Reinwardtia 7, 1968, 383-410, 1 fig.) and Lithocarpus (ibid. 8, 1970, 197-308, 13 fig., 1 map). KEY TO THE GENERA (based on flowering specimens) 1. Inflorescence capitiform, axillary, consisting of 1—-3-flowered dichasial clusters. § Flowers with 10-20 stamens; anthers linear, 2-7 mm long, basifixed, apiculate; pollen grains peroblate, 4-9-colpate, exine distinctly echinulate. 9 Flowers dimerous, perianth poorely developed, styles 2, short, tongue-shaped; stigmas papillose, covering the inner surface of the styles; ovary 2-celled, flattened. Leaves glandular- dotted; stipules peltately attached, their insertion surrounded by colleters.. .. . 1. Nothofagus 1. Inflorescence a spike of condensed cymes, flowers solitary or in dichasial clusters of 3-30 along the rachis. ¢ Flower with 6-12 stamens; anthers 0.25-1 mm long, non-apiculate; pollen grains prolate or perprolate, 3-colpate, 3-colporate, exine scabrate or more or less smooth. 2 Flowers 3(—6)-merous; perianth well-developed; styles 3-6 (—9), conical or terete; stigmas terminal, punctiform or capitate; ovary 3(—6)-celled, not flattened. Leaves not glandular-dotted; stipules basally inserted; no colleters. 2. $ Flowers with the pistillode always present; stamens 10-12, anthers 0.25—-0.35 mm long, dorsifixed; pollen grains perprolate (ellipsoidal), c. 16 by 10, exine more or less smooth. 9 Flowers with 10-12 staminodes; stigmas punctiform. ¢ Inflorescence always erect. 3.1 Cupule-primordia already developed before anthesis, always solitary, with distinct vertical sutures, with 2-4(-8) separate growing-points; enclosing 1-3(-7) 2 flowers. .... . 2. Castanopsis 3.1 Cupule-primordia not yet developed before anthesis, solitary or in dichasial clusters, ring-shaped without vertical sutures and separate growing-points; enclosing 1 2 flower only. 3. Lithocarpus 1) To distinguish Castanopsis from Lithocarpus the 9 flower is needed. 1972] FAGACEAE (Soepadmo) TT 2. ¢ Flower without pistillode, stamens 5—6(-9), anthers 4-1 mm long, basifixed; pollen grains pro- late (spheroidal), 30-40 by 20 yw, exine scabrate. 2 Flower without staminodes or rarely with 5-6 staminodes; stigmas capitate. ¢ Inflorescence pendulous or erect. 4. Inflorescence always unisexual, simple. ¢ Inflorescence pendulous. 2 Flowers always solitary, sta- minodes sometimes present; ovary rounded in cross-section. Terminal buds conferted, the scales with a tendency towards orthostichy. Leaves spirally ahi att crowded at the end of the twigs; stipules not interpetiolar. . .. . . . 4. Quercus 4. Inflorescence unisexual or bisexual (androgynous), simple | or - much- branched. 3 Inflorescence erect. 2 Flowers in dichasial clusters of 3-15; ovary trigonous in cros-section; staminodes absent. Terminal buds not conferted, scales imbricate. Leaves in whorls of 3; stipules interpetiolar . 5. Trigonobalanus KEY TO THE GENERA (based on fruiting specimens) 1. Cupule open and 2-16-lobed or completely enclosing the fruits; not dehiscent, or irregular so, or into 2-8 + equal segments; always with vertical sutures. Nuts 1-15 in each cupule, rounded-angular or 2-3-gonous in cross-section. 2. Cupule lamellate or set with imbricate scales. Nuts 2—3-gonous in cross-section. Stigmas ligulate or capitate. 3. Cupule solitary, axillary, lamellate, 2-lobed, containing !—3 nuts. Fruits 2-gonous, winged. Stigmas ligulate. Leaves distichous, glandular-dotted. Stipules not interpetiolar, peltately attached 1. Nothofagus 3. Cupules many along a rachis, scales imbricate to concentrically set; 4-16-lobed, containing 3-15 trigonous but not winged nuts. Stigmas capitate. Leaves in whorls of 3, not glandular-dotted. Stipules interpetiolar, not peltately inserted... . . . .. . . 5. Trigonobalanus 2. Cupule spiny, muricate, or rarely almost smooth. Nuts rounded- angular in cross-section. Stigmas punctiform, terminal. Leaves spirally arranged. Stipules not interpetiolar. . . . . 2. Castanopsis 1. Cupule never lobed, saucer-shaped, cup-shaped, or almost completely enclosing the fruit, indehiscent, without vertical sutures. Nut always solitary in a cupule, circular in cross-section. 4, Cupules solitary or in dichasial clusters, sometimes almost completely enclosing the fruit; variously squamose, muricate, or lamellate. Complete acorn without a ringed umbo. Stigmas punctiform. Leaves usually not crowded near the end of the branchlet, entire. Terminal buds solitary, scales imbricate. ... . . . 3. Lithocarpus 4. Cupules always solitary, saucer- -shaped ¢ or ‘cup: -shaped, lamellate. ‘Umbo of complete acorn ringed. Stigmas capitate. Leaves crowded at the end of the twig, entire or crenate in the apical half. Termi- nal buds conferted, scales with a tendency towards orthostichy. . ........ -. 4. Quercus KEY TO THE GENERA (based on sterile specimens) 1. Leaves glandular-dotted beneath. Stipules peltately attached . . ....... .. 1. Nothofagus 1. Leaves not glandular-dotted. Stipules not peltate. 2. Leaves in whorls of 3. Stipules interpetiolar. ............. . 5. Trigonobalanus 2. Leaves spiral. Stipules not interpetiolar. 3. Leaves not crowded near the end of the twigs. Terminal buds not conferted, scales imbricate or rarely with a tendency to distichy. 4. Wood-rays (of older branches and mature wood) exclusively uniseriate . . . . 2. Castanopsis* 4. Wood-rays uniseriate and many-celled. . . . ‘ . . 3. Lithocarpus? 3. Leaves crowded at the end of the branchlets. Terminal buds conferted, scales with a tendency to orthostichy. i uh aT tL” 0) St a ea LR GLUE cpt 1. NOTHOFAGUS? BLume, Mus. Bot. Lugd. Bat. 1 (1850) 307, nom. cons.; OrRST. Vidensk. Selsk. Skr. V, 9 (1871) 354; STEEN. Blumea 7 (1952) 146; J. Arn. Arb. 34 (1953) 332, with full synonymy; ibid. 35 (1954) 266. — Fagaster SpAcuH, Hist. Nat. Vég. Phan. 11 (1842) 142. — Calucechinus Homer. & Jaca. in Dumont d’Urville, Voy. Pol Sud 1 See for the difference in slash characters under Morphology-bark, p. 271. 2 Elaborated by C.G.G.J. VAN STEENIS. 278 FLORA MALESIANA [ser. I, vol. 7? & Oc. (Astrolabe & Zélée) Bot. Atlas, Dicot. (1844) t. 6 0; (1845) t. 7 Z, 8 I. — Calusparassus Hompr. & Jaca. /.c. (1844) t. 6 E; (1845) t. 7 T, 8 y. — Lophozonia Turcz. Bull. Soc. Imp. Nat. Moscou 1858, I (1858) 396. — Fagus subg. Caluspa- rassus et subg. Calucechinus Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 103. — Fagus sect. Eufagus A.DC. Prod. 16, 2 (1864) 117, pro spp. 4-8; sect. Nothofagus A.DC. lc. 121; emend. B. & H. Gen. Pl. 3 (1880) 410. — Trisyngyne BAILL. Adan- sonia 11 (1873) 136; BAUMANN-BODENHEIM, Bull. Mus. Hist. Nat. Paris II, 25 (1953) 419. — Nothofagus subg. Lophozonia et subg. Molischia KRASSER, Ann. Hofmus. Wien 11 (1896) 162. — Parafagus OLIv. Trans. Proc. R. Soc. New Zeal. 66 (1936) 292, gen. foss. — Fig. 3-9. ae Sevmour I. Mc Murdo Sound Fig. 2. Distribution of the genera Fagus on the northern and Nothofagus on the southern hemisphere. Fossil localities indicated by crosses including also the Antarctic localities at Seymour I. and McMurdo Sound (Courtesy Blumea). Distr. From southern South America (incl. Staten I.) beyond 33° S to New Zealand, Tasmania, E. Australia, New Caledonia, and New Guinea (incl. the adjacent d’Entrecasteaux Is, Goodenough & Nor- manby, and New Britain), in all 37 spp. (South America 11, New Zealand 4, Tasmania 2 (1 endemic), Australia 2, New Caledonia 5, New Guinea c.a. 13). Fig. 2. Taxon. The genus is subdivided as follows (STEEN. 1953, l.c.): 1. Deciduous. Leaves plicate in bud, texture always thin. South America, Tasmania ...1. Sect. Nothofagus Calucechinus Homsr. & Jaca., Lophozonia Turcz., Fagus subg. Calucechinus Miq., N. sect. Calu- cechinus KRasserR, N. subg. Lophozomia KRASSER, N. sect. Deciduae STEEN. Blumea 7 (1952) 146, N. sect. Plicatae STEEN. I.c. 306, laps. cal. 2. Cupule 4-valved, 3—7-flowered. South America, Tasmania (8 spp.), type subsection. la. Subsect. Antarcticae STEEN. 2. Cupule 2-valved, with linear valves. 2 Flower 1. South America (1 sp.) . 1b. Subsect. Pumiliae STEEN. 1. Evergreen. Leaves not plicate in bud, flattish or folded along the midrib in bud, exposing the under- surface, with firm texture. Sy 5 ee 2. Sect. Calusparassus (HomBR. & JACQ.) KRASSER Calusparassus Hompr. & JAcQ., N. sect. Sempervirentes STEEN., N. sect. Planae STEEN. 3. Cupule 4-valved, lamellae subentire to lobed, parted or divided into appendages. Lateral 9 flowers normally 3-merous. South America (3 spp.), New Zealand (3 spp.), Tasmania & E. Australia (2 2b. Subsect. Tripartitae STEEN. 1972] FAGACEAE (Soepadmo) 279 3. Cupule 2-valved, the valves connate or free, with usually entire lamellae, or cupule reduced. 3 Flowers solitary or in triads, 2 flowers 1 or 3 per cupule. New Guinea (13 spp.), New Caledonia (5 spp.). 2c. Subsect. Bipartitae STEEN. Ecol. The ecological range of Nothofagus is defined by an everwet climate from the cold subantarctic and temperate-montane to tropical-montane and tropical thermo-ecological conditions; lowland tropi- cal species are found in New Caledonia and descent of Nothofagus to c. 600 m is observed in New Brit- ain and to 750 m in the mainland of New Guinea and d’Entrecasteaux Is. As other Fagaceae, Nothofagus shows a distinct social occurrence, past and present, mostly gaining dominance or co-dominance, as usual in the family, to which the occurrence in the montane forest in New Guinea makes no exception. The impression is gained that this is largely due to the symbiosis with ectomycorrhiza (cf. STEEN. Blumea 19, 1971, 72). Nothofagus is found susceptible to fire, and does not possess fire-resistant characters. Pollination is by wind and protandry seems to prevail. Anthesis occurs simultaneously and enormous quantities of pollen are produced which under certain conditions are observed (in New Zealand) to form clouds above the forest. Scattered pollen grains have been found at thousands of miles from the source area in windy austral areas. A symbiont of the crown, viz all species of the genus Cyttaria (Discomycetes), occurs throughout its range except in New Guinea and New Caledonia. Dispersal of the nuts of Nothofagus is very restricted and very slow. HOLLOWAy (1954) has shown that Nothofagus cannot stand transport by seawater. The nuts are compressed or triangular, possess mostly a sharp border, but are not provided with any special means of dispersal. They are rather small, but dis- persal by wind will not carry them far. PREEST (1963) estimated the absolute maximum dispersal distance at c. 2-3 km and concluded that long-distance dispersal, either by wind, by birds, or by ice-bergs is ex- cluded. In New Guinea KALKMAN & VINK (1970) found dispersal and regeneration confined to the stands. Consequently dispersal, past and present, seems to be bound to close distance of land. Wood anat. According to METCALFE & CHALK (Anat. Dicot. 2, 1950, 1309-1315) within Fagaceae the wood anatomy of Fagus and Nothofagus is most closely allied. Palyn. Within the family Fagaceae the pollen of Nothofagus is very characteristic and is not closely allied to that of the other genera. Within it, three main types are distinguished, the fusca, menziesii and brassii types. These types do not correspond with a single infrageneric taxon. As to the species, some of the latter type can be distinguished specifically by their pollen, but many of them cannot. Hence, one ‘pollen species’ may correspond with several botanical species. See CoOKSON & PIKE (Austr. J. Bot. 3, 1955, 197-206) and WALKER & WITTMANN (Pollen et Spores 7, 1965, 457-464). In Australia (incl. also New Guinea & New Caledonia), New Zealand and South America all three types are represented among the living species. Hybrids. In New Zealand and in South America hybrids and introgression has been recorded. As yet we have no indications of this to occur in New Guinea. Fossils. Both macro- and microfossils have with certainty been found in SE. New Guinea, Australia, New Zealand, Antarctica: McMurdo Sound (c. 165° E, 77° S) & Seymour I. (c. 64° W, 57° E) and Fuegia from the Tertiary onwards. In Victoria, New Zealand, Seymour I. and Fuegia fossil pollen is even found in Upper Cretaceous deposits, partly of the brassii and partly of the fusca pollen types. The menziesii pollen type appeared later, in the Eocene in Australia and in the Oligocene in New Zealand. In New Guinea fossil pollen is only known to occur as yet from the Upper Miocene and Pliocene; this belongs all to the brassii pollen type. In Australia and New Zealand fossil leaves and pollen of Nothofagus are so common that it is generally assumed that Nothofagus was abundant throughout (at least the eastern and southern parts of) the continent in pluvial Tertiary times. There are a fairly large number of subtypes or individual ‘pollen species’ known to be extinct in Australia and New Zealand, so that we may safely conclude that the genus has produced since the Upper Cretaceous a hundred species in austral Pacific lands. Records of fossil pollen from the northern hemisphere and from Africa are due to misidentifications. Among the living species the brassii pollen type occurs only in the New Guinean and New Caledonian species which form together subsect. Bipartitae and also in N. alessandrii Espinosa from S. Chile. This pollen type is one of the two ancient types and is found in fossil state throughout the known Nothofagus range; it occurs widely in the Australian and New Zealandian Tertiary and as in Metasequoia the fossils (in this case pollen) had been described before the living plants were known. See for the fossil sites fig. 2 and for the literature on it Pacific Plant Areas vol. 2 (Blumea Suppl. 5, 1966, 290-292, map 163, & ref- erences), and my recent paper in Blumea 19 (1971) 78-82, f. 97-98, tab. 1. Systematics & Phylogeny. Nothofagus is a strictly southern hemisphere counterpart of the nearest allied but quite separate genus Fagus of the northern hemisphere, of which the most southern stations are at present found in southern Yunnan and Formosa. These two genera form together the subfamily Fa- goideae. | have called Nothofagus a key genus for plant geographical and phyletic conclusions, as it meets the three criteria for safe reasoning, viz a sound taxonomy, an ample fossil record, and diaspores which require almost continuous land for dispersal. The ancestry of Nothofagus and Fagus cannot be divorced and they must have originated from a common ancestral matrix. Though their ranges do not 280 FLORA MALESIANA [ser. I, vol. 7? meet at present, the subtropical and tropical-montane area between southern Yunnan and Queensland seems the most likely place for the situation of their cradle. This area contained possibly the entire an- cestral Fagaceous matrix from which they differentiated, Fagus possibly at the northern end and Notho- fagus at the southern end. This cradle area harbours even at present, either inside itself or along its north- ern border the total range of morphological diversity of the family contained in the seven genera of Fagaceae. And in it is the highly interesting endemic genus Trigonobalanus which may be a palaeo-allo- polyploid. The spreading from the matrix area must have started before the Upper Cretaceous, as Faga- ceous fossils are known from the northern hemisphere at that time and also of Nothofagus in austral regions. Morphologically quite primitive species and appreciable diversity are still found in South Ameri- ca. In the warmer Cretaceous and Tertiary, when the major floral zones extended towards Antarctica Nothofagus could freely spread on available southern land areas. This wealth was brought to an in geolog- ical terms of time-scale abrupt end with the onset of cooler conditions at the end of the Tertiary followed by the havoc caused by the Ice Age: the retreat of the floral zones by the southern glaciation and the dramatic desiccation of most of the Australian continent. See for a more complete exposition of the theo- ry, VAN STEENIS (Blumea 19, 1971, 65-98). 1. Section Calusparassus (Homsr. & JACQ.) KRASSER, Ann. Hofmus. Wien 11 (1896) 163; STEEN. J. Arn. Arb. 34 (1953) 336. — Calusparassus HoMBR & JACQ. in Dumont d’Urville, Voy. Pol Sud & Oc. (Astrolabe & Zélée) Bot. Atlas, Dicot. (1844) t. 6 E; (1845) t. 7 T, 8 yw. — Nothofagus sect. Sempervirentes STEEN. et sect. Planae STEEN. Blumea 7 (1952) 146, 306. 1. Subsection Bipartitae STEEN. Blumea 7 (1952) 146, incl. ser. Triflorae et ser. Uniflorae STEEN.; J. Arn. Arb. 34 (1953) 338. Monoecious, evergreen, sometimes buttressed, trees or shrubs; bark grey, usu- ally coming off in flakes. Innovations usually varnished by (sometimes yellow) resin exuded by colleters and glands underneath the leaves. Perular bracts in 4 orthostichies, with colleters. Leaves (in Mal.) distichous, in bud conduplicate, exposing the lower surface, entire, rarely crenate, tip emarginate, beneath glandu- lar-dotted. Stipules peltately attached, at and near insertion with many colleters. — 3 Flowers in the lower part of the flush, axillary, the lower often between a pair of efoliar stipules, solitary or in ebracteolate triads, sessile or short-pedicelled or -peduncled; bud + club-shaped, perianth closed, often glandular-dotted, later rupturing at apex, basal part sometimes constricted. Stamens c. 12-18, filaments usually basally connate, not rarely lax-haired; anthers linear, basifixed, often glandular and/or laxly haired, dehiscence latrorsely; connective apiculate. — 2 Flowers in the upper part of the flush, axillary, 1 or 3, usually in a cupule. Ovary sessile, flat, sometimes with a narrow marginal wing, mostly with 2 shoulders (perianth), rarely (abnormal?) with a hornlet below the apex on the flat side, usu- ally glabrous; 2-celled; style short with 2 stigmatic arms. Cupule 2-lobed, rarely split to the base, usually with 1 or more lamellae, the latter at the adaxial side with colleters, finally often woody and gaping; cupule rarely reduced to 2 minute free elamellar thin flaps, or entirely absent. Nuts flat, + orbicular to ovate, rarely lined with a thin margin, 1-seeded, apiculate by the style-base. Seed with membranous testa; cotyledons thin, folded, with fatty reserve. Germination epigeal. Distr. New Guinea (13 spp.) and New Caledonia (5 spp.). Ecol. Throughout New Guinea in the tropical-montane everwet rain-forest, largely between 1000 and 3100 m, but in several places recorded from 900, 850 and 750 m (N. flaviramea, N. starkenborghii). In 1972] FAGACEAE (Soepadmo) 281 Normanby I. N. carrii and N. rubra occur down to 750 m, and in New Britain N. starkenborghii has been collected as low as 600 m. Some of the New Caledonian species descend or grow in the lowland hills or are even confined to the lowland forest, the lowest locality being 100 m. In New Guinea on deep soils Nothofagus trees grow to large straight trees, sometimes to over 45 m tall with a diameter of 1, or even 14 m. Fig. 3. On stony ridge crests and other localities with shallow poor soil trees may be dwarfed and represent gnarled poor shape or be shrubby, as observed in several species. Fig. 8. As to soils, there seems to be in New Guinea no preference; there are a few records of Nothofagus grow- ing on limestone (bedrock). The New Caledonian species are said to be restricted to serpentine. Dominance or co-dominance is reported for almost all species, especially for N. pullei (9 times), N. perryi and N. grandis (both 5 times), N. starkenborghii, N. rubra, and N. flaviramea (each 3 times). Dom- inance can easily be observed from the air when the trees are in reddish flush. Co-dominants consist, for a large part, of other Fagaceae, notably Castanopsis and Lithocarpus; other frequent genera are Arau- caria, Libocedrus, Weinmannia, Engelhardia, and Cryptocarya. BRASS, ROBBINS, and KALKMAN & VINK are of the opinion that co-dominance with Castanopsis and Lithocarpus is less manifest than collectors sometimes suggest, and observed that usually the Nothofagus zone is above the zone dominated by these two other genera. Wood anat. DADSWELL & INGLE (Austr. J. Bot. 2, 1954, 141-153, 3 pl.) found that the wood anatomy of the species of the subsection appears to be homogeneous and differs to some extent from other species of the genus. The wood structure of the twigs of New Caledonian species is similar to that of the New Guinean species, but not exactly so. Morph. In the New Caledonian species the phyllotaxis is not strictly distichous as in the New Guinean ones, but + spiral. In most species the ¢ flowers occur in either sessile or stalked triads, but some 4 Papuan species have the 3 flowers solitary. The 9 flowers occur in threes per cupule or are solitary; in some of the latter the cupule is merely a ves- tige or may even be quite suppressed. Solitary 2 flowers occur also in one or two New Caledonian species. The primitive state is obviously a well-developed, many lamellate cupule with 3 9 flowers and stalked triads of 3 flowers, from which the other states can be derived by reduction. Testimony of this derivation are very occasional aberrations; for instance I have found in one cupule of N. brassii the 2 lateral flowers not developed but aborted as rudiments. I have also found in one cupule of N. flaviramea 2 minute abortive lateral 2 flowers in addition to the central nut, testimony of its derived status. I have formerly attached taxonomical value to the occurrence of 3 versus 1 2 flower per cupule and distinguished ser. Triflorae and ser. Uniflorae. But new closer study of the New Caledonian species and new finds in Papua have induced me to view this distinction not as a phylogenetical cleft, and hence give it no value for natural distinction. Specific affinities point distinctly to parallel reduction from flowers in several species lineages. Therefore, I have not maintained these series. Specific delimitation. The five New Caledonian species are clearly distinct at first sight, even in sterile state; all possess coarse, hard leaves with characteristic shapes. Specific distinction by vegetative characters is for New Guinean species less easy, and I am not certain that all sterile specimens have been properly named. Great value is given to the characters of the mature cupule, but of some species these are not known in the fully mature state and in other specimens it is sometimes difficult to decide from the herbarium mate- rial whether the mature state is reached. I have allowed a greater variability in cupule structure in N. pullei than in my monograph. A problem remains with N. resinosa and N. pseudoresinosa, which almost only differ in the absence or presence of a cupule. Observations on development of cupular structure in numbered trees may provide a better insight in this problem. The ¢ flowers do not provide reliable characters, but 4 species have them solitary instead of in triads, viz N. pullei, N. crenata, N. resinosa, and N. pseudoresinosa. In N. starkenborghii and N. carrii the 3 peri- anth is more cup-shaped than in the others where it is tubular. In N. carrii and N. grandis triads appear more clustered than in other species. Uses. Some species provide obviously excellent timber of big dimensions, e.g. N. perryi, which seems durable also underground. Also the timber of N. starkenborghii and N. pullei seems good. Papuans use this timber; they also plant these species around their villages and fields for which they use seedlings taken from the forest. Notes. Collecting adequate Nothofagus material is far from easy, partially by the large size of the trees, partially by the short period of the flush in which ¢ and 2 flowers are produced, but also partially by the proportionally scarce and often very inconspicuous cupules or nuts. Since the cupules furnish the most important characters for identification, collectors should carefully examine many twigs of felled trees to search for them, as they are scarce on a single twig and easily over- looked. As in the forest more than one species may be present one should be very careful to correlate fallen cupules with freshly collected specimens, but it is certainly very worthwhile to pick them from the forest floor underneath the tree and state so in the field notes. Leaves may vary considerably in texture and size in conjunction with the condition of the tree during collecting and also on the age of the tree and furthermore with the habitat; in exposed sites they are gener- 282 FLORA MALESIANA [ser. E volig* ally smaller and more coriaceous. Sterile material from immature trees or from lower branches of mature trees is mostly impossible to identify. In 1953 only 36 collections were available; at present there are nearly 400; this has necessitated some reductions, but also yielded 2 new species. For using the key to the species one needs to have material with cupules. A number of species can be distinguished on vegetative characters (material from mature trees) and these have been listed separately for practical purposes. KEY TO THE SPECIES! 1. Cupule 3-flowered. 2. Leaves upwards crenate, usually ovate to elliptic-oblong, c. 5-10 by 24-4 % cm. Peduncle of cupule c. 144-3 cm. 3. Cupule well-developed, lamellate, c. 144-2 cm through, the nuts not or hardly exserted. Leaves usually ovate-oblong, more or less pointed to the apex; glands underneath less than or at most up to 1 mm spaced. Stipules early caducous, at least not present on fruiting twigs. Peduncle of 3 triads 10-25 mm os . . 1.N. perryi 3. Cupule hardly developed, less than Yo cm long, consisting of 2 poorly developed valves each with an indistinct lamella, the nuts almost entirely protruding. Stipules persistent, even on fruiting twigs. Leaves elliptic-oblong, not or hardly pointed. Glands underneath more widely (1-2 mm) spaced. 2. N. nuda 2. Leaves entire, elliptic in outline, up to 51% by 24% cm. 4. Midrib sulcate above, without trace of an elevated ridge over its entire length. Anthers c. 24% mm long. Peduncle of cupule 2— Asm) sar Yee . . 3. .N. starkenborghii 4. Midrib at least in the basal half with an 1 elevated ridge. " Anthers c. 5- 3 mm. Peduncle of cupule 5=lSamms 990 J smd TOeleadowky coer 2) aes 2 Flowers solitary, cupule present or absent. * Cupule entirely absent. 6. Leaves waxy beneath, elliptic, margin towards apex faintly crenate or minutely toothed, 444-10 by 2—5 cm; nerves c. 9 pairs, rather parallel and extending towards the margin; petiole 5-8 mm. Stipules 4-6 mm caducous. 3 Flowers 1, 6-7 by 344-4 mm; anthers 5-6 mm. Nut hairy, ovate- oblong ... . . 7.N. resinosa 6. Leaves not waxy ‘beneath, ay ovate- ‘elliptic, entire, 244-4 by Wty cm; nerves c. 6 pairs, looped at apex; petiole 2-3 mm. Stipules 3-4 mm, rather persistent... . - evs 5. IN. puilei 5. Cupule present, each valve consisting of at least an elamellate flap. 7. Cupule only represented by 2 small, free, elamellate flaps very much shorter than the nut. 8. Leaves + obovate to obovate-oblong, rarely also some + elliptic, 2—41% by 1-214 cm; nerves 5-7 pairs; midrib above sulcate with ridge to reas gine 244-5 mm. Ultimate twigs + thin. Nuts c. 6-7 by 4-5 mm. .. . . 9.N. carrii 8. Leaves ovate-oblong, flat, acutish towards apex, or even ale acuminate ‘(very rarely blunt and el- liptic), 644-13 by 244-5 Y, cm; nerves 8-10 pairs; midrib not sulcate, the ridge prominent at least halfway, often to apex; petiole 7-10 mm. Nuts c. 8-10 by 5-7 mm. . . . . . 10. N. flaviramea 7. Cupular valves at least provided with 1 lamella. 9. Cupules already in the young state distinctly stalked. 10. Leaves ovate-oblong, distinctly acutish towards the apex. 11. Leaves smallish, c. 244-5 by 114-2 cm, crenate towards the apex. Cupule with c. 3 lamellae, the valves oval. Nut c. 5 mm, narrowly ee about as large and of similar shape as the cupule halves) 2 2s Se GE OL eG ey erent Leaves much larger, C. 4Y,-9 by 2-4 cm, entire. aD, Cupule on a thin stalk, c. 10-15 mm, with 1 rather apical lamella, much smaller and narrower than the nut, c. 6 mm long. Nut ovate-acute, c. 10 by 6 mm, winged towards apex. 13. N. womersleyi 12. Cupule sessile to provided with a thick, obconical stalk-like extension c. 5 mm long, large, with 3—4 lamellae, finally woody, 12-13% by 744-10 mm. Nut rhomboid, c. 7-10 by 94%4-10 mm, smaller than the cupular valves. Twigs coarse, greyish... ... .. . I1.N. grandis 10. Leaves elliptic, not acutish towards the apex. 13. Leaves large, with distinct, somewhat prominent venation above, c. 444-7 by 244-4 cm. Twigs coarse, greyish. Mature Patios large, c. 12-20 by 7144-10 mm. Nut large, rhomboid, c. 8-10 Ohad el . . . II. N. grandis 13. Leaves much ‘smaller, c. 134-234 by rae 1y, cm, Teticulations fine-tessellate, not prominent. (Immature) cupules c. 3-4 mm, immature nut suborbicular, about the same size . 12. N. rubra + Leaf-sizes cannot be taken from young trees or watersprouts. Cupule structure is that of mature cupules, but this is sometimes difficult to ascertain and in some species unknown. 1972] 9. Cupules sessile or almost so. FAGACEAE (Soepadmo) 283 14. Leaves thin, (minutely) crenulate or minutely toothed towards apex, waxy underneath, c. 244—5 by 114-21% cm; nerves 7-8 pairs. Cupule c. 7-8 by 4-5 mm, with (1--)2 lamellae. 3 Flowers solitary ad ren ei ak senshi ace Wena eet 14. Leaves quite entire, not waxy, not thin. ee Se) Meee Shs ot he Oe IN. PSCUGOTesINOSA 15. Rather coarse. Leaves longer than 3 cm. Cupules woody, not split to the base in mature state. 16. Leaves large, c. 444-7 by 2%4—4 cm, rather thin-coriaceous (also on twigs with mature cupules), with distinct and -- prominent venation above. Twigs greyish, rather thick. Cupules c. 12-20 by 7144-10 cm, with 3—4 lamellae. Nut c. 7-10 mm 9. 11. N. grandis 16. Leaves mostly hard-coriaceous, sometimes convex or with recurved margin, smooth above, but the nerves (not the fine-tessellate veins) thickish and obtusely prominent, c. 244-5 by 114-3 cm. Cupules smaller, at most 12 by 6 (10-12) mm, with 2—3 lamellae. Nut c. 4-5 mm @ 15. Less coarse. Leaves less than 3 cm long. 12. N. rubra 17. Young twigs hairy to puberulous, at least minutely so on the innovations. Cupule with 1 lamella, not seldom irregularly incised-dentate, mostly split to the base, almost always nar- rower than the nut. d Flowers solitary . 5. N. pullei 17. Twigs glabrous. Leaves generally hard-coriaceous, smooth above, but the nerves thickish and obtusely prominent, sometimes convex or with recurved margins. Cupule with 2-3 la- mellae, woody, not split to the base, as broad as or broader than the nut. j Flowers in triads 12. N. rubra SOME CHARACTERS FACILITATING IDENTIFICATION of sterile, very young, inadequate or 3 material Leaves crenate towards apex: Nerves ending in the incision of the crenation: N. perryi, N. nuda, N. crenata. Nerves (or accessory vein) ending in the crenation or tooth: N. resinosa, N. pseudoresinosa. Young foliage with abundant resin: N. resinosa, N. pseudoresinosa. Leaf apex pointed: N. perryi, N. crenata, N. flaviramea, N. womersleyi, very rarely slightly in N. grandis. Midrib above, sulcation without trace of ridge: N. starkenborghii. Midrib above not in a depressed sulcation, but sharply prominent as a ridge: N. flaviramea. Innovations hairy, sometimes minutely puberulous: N. pullei. Very slightly so in N. resinosa where also the nut and the perianth of the 3 flower is puberulous. Twigs flattened with yellowish bark (not an exclusive decisive character): N. flaviramea. Cupules peduncled: N. brassii, N. nuda, N. crenata, some forms of N. rubra, N. womersleyi, shortly in N. starkenborghii. In N. grandis sometimes a short thick peduncle-like obconical attenuation. 3 Flowers solitary: N. pullei, N. crenata, N. resinosa, N. pseudoresinosa. 3d Triads rather crowded, forming pseudo-glomerules: N. carrii, N. starkenborghii, N. flaviramea, and often in N. grandis. 1. Nothofagus perryi STEEN. Blumea 7 (1952) 146; J. Arn. Arb. 34 (1953) 347, f. 3-2’, 6. — Fig. 4, 9b-b?. Large tree, (14-18—-)25—40 m and 60-160 cm go. Ultimate twigs + zigzag, -- applanate, sometimes yellowish. Stipules c. 7 by 3 mm, attached at lower third. Leaves ovate-oblong, coriaceous, 4144-11 by 2-4% cm (I = 2-249), shallowly crenate in upper half to -- pointed apex; ridge on midrib to %4 or almost to top; nerves 6-8 pairs, rather straight, prominent beneath, not or rather distinctly so above; reticulation above usually indistinct, be- neath often weakly prominent, a veinlet running to the toothlet; petiole 4-6 mm. — ¢ Flowers in 4-12 mm peduncled triads, sessile to 1 mm stalked; perianth + truncate-campanulate, 5 by 3 mm. Stamens c. 13-15; filaments connate beyond peri- anth, free for 12-15 mm; anthers 4-5 mm. — 2 Flowers: cupule c. 4 mm peduncled, consisting of two thin elamellar flabs c. 2 by 3 mm; stigmas just protruding. Fruiting cupule coarse, woody, on a straight or curved, 15-25 mm long, usually rather coarse, upwards obconically thickened peduncle, 3-lamellar, 10-15 by 11-18 mm. Nut ovate, api- cally + winged, + ovate, 5-8 by 5-6 mm. Distr. Malesia: East New Guinea (Western Highlands: Wabag, Lai R., Nondugl, Al River Mts, Jimmi Valley, Bleekep, Mt Hagen, Mt Oga, Nona Camp, Kubor Range, Giluwe, near Kuli; Central Div.: East Mt Tafa, near Nemodi; Morobe Distr.: Trist; Eastern Highlands: Upper Chimbu, near Raregigl, Mt Michael, Goroka). In all 41 collections of which 13 fertile. Ecol. Mountain forests, not rarely abundant or dominant, on Mt Elimbari at 2100 m also on limestone, 1600-2600 m. 3 Fi. Febr., July, fr. June- Sept., flush June-July, Dec. (once Sept., Febr.). Near Wabag dominant at 2400 m, near Goroka dominant in ridge forest with Lithocarpus co- dominant, at Baime Creek (Yamap, Wau) co- dominant with hoop pine, in Bena Bena Valley (Goroka) the dominant tree in ridge forest asso- ciated with Lithocarpus, on Mt Michael co-domi- nant with Castanopsis at 2000 m, on Mt Tafa to- 284 FLORA MALESIANA [ser. I, vol. 7? Se ae Fig. 3. Nothofagus grandis STEEN., some 40 m tall, along a clearing for coffee plantation, at Aiyura, T.N.G. (cf. NGF 3389)(WOMERSLEY, Sept. 1951). 1972] FAGACEAE (Soepadmo) 285 Fig. 4. Necklace of cupules of Nothofagus cf. perryi STEEN., adorning Miss J. VANDENBERG, Of the Division of Botany, Lae, at the Lae Show, 1968; an Aseki exhibit. wards Nemodi dominant as tall forest with Litho- carpus at 2100 m, at Lake Trist (Morobe) domi- nant tree on spurs and side-slopes at 1650 m. Uses. Apparently a very useful species for its excellent hardwood; on Mt Michael and on Mt Kubor used for building purposes. On Hagen and Chimbu recorded to be planted along roads and in Chimbu also on edges of coffee gardens. Vern. Tar(a), taro, t. karap, Wabag, Enga lang., iumba, suker, Chimbu, garaip, gripe, Hagen, sama, Mt Tafa, Waitape, vembeh, Minj, jufeta, Bena Bena. Notes. By its large, coarse, crenate leaves, peduncled triads of ¢ and @ flowers, and multila- mellar woody cupules the most primitive among the New Guinean species. An allied species is N. nuda (see there) and the New Caledonian WN. discoidea (BAUM.-BoD.) STEEN., but that species is crenate almost along the entire margin, with veins ending in the crenation- bays, -+ scattered leaves, sessile, few-lamellate cupules with only 1 nut. Even in fairly hard smallish leaves of old trees or from exposed habitats the leaf-shape is always somewhat pointed towards the apex. Of young trees, saplings or lower branches the leaves are thinner and much larger (up to 1714 by 61% cm in a tree 15 m high!) and resemble in shape those of N. flaviramea and N. womersleyi; but they have always some coarse crenulations towards the apex and are generally thicker in texture. In all three the ridge on the midrib above extends almost to the leaf apex. 2. Nothofagus nuda STEEN., nov. sp. — Fig. 5. Nothofagi perryi affinis, differt foliis ellipticis (index 2%-3), obtusis, subtus glandulis 1-3 mm sibi remotis, valvis cupularibus minutissimus, c. 2-3 mm longis liberis unilamellatis. — Typus: R. PULLEN 6582, holotype CANB, isotypes A, L, LAE. Tree, c. 20 m. Stipules 7-9 by 3-4 mm, peltate, rather persistent. Leaves distichous, elliptic (I = 214-3), coriaceous, towards apex shallowly crenate, 8-10 by 3-4 cm, apex blunt, emarginate; midrib above with prominent ridge except near apex, very prominent beneath; nerves 7-9 pairs, prominent on both sides; venation prominent on both sides, fine-tessellate above, a veinlet ending in the inci- sion of the crenations; petiole c. 10 mm, thick.— 3g Unknown. — 2 Peduncle 2-2% cm, thin. Cupular valves free, thin, c. 3-4 mm high, emarginate to 2-lobed, obviously with 1 lamella. Nuts 3 per cupule, ovate, c. 10 by 6-7 mm, 3-dentate at apex by the style base and 2 erect, perianth teeth. Distr. Malesia: East New Guinea (Upper Wenna Creek, branch of Tauri R., near Paina village, Gulf Distr. in Papua). One collection. Ecol. Mixed lower montane rain-forest, asso- ciated with Castanopsis. Vern. We-ukwe, Nauti lang. Notes. Allied with N. perryi, but differing in the extremely small, not woody, 1-lamellate cupule, the leaf-shape, the wider spaced glands, the thin peduncle, the prominent venation, and the veinlet ending in the bay of the crenation instead of in the tooth. A primitive species but with a reduced cupule. Also resembling the New Caledonian N. balan- sae (BAILL.) STEEN., but this differs by obovate leaves, large cupules and nuts. 3. Nothofagus starkenborghii STEEN. Blumea 7 (1952) 347; J. Arn. Arb. 34 (1953) 347, f. 7. Generally a large tree, 16-45 m and 24-over 100 cm go; bark shedding in hard, thin sheets or large scales, often defoliating in large, + rectan- gular plates. Leaves elliptic (rarely some leaves + obovate), subcoriaceous, 3-8 by 14-344 cm (I = 2%-31%); midrib sulcate above without a ridge; nerves 6-10 pairs, as the reticulations rather indis- tinct on both surfaces; petiole 44-1 cm. Stipules up to c. 6-7 mm, acute at both ends or rounded below, inserted at the lower 1/3. —d Flowers: 1-2 mm stalked triads, many together as in a glome- rule; buds at base stalk-like for 1-2 mm, then 3-4 mm ellipsoid, when open perianth wide-campanu- late cup-shaped, 3-4 mm high; filaments connate to above the perianth rim. Stamens c. 12-14; an- thers 244-3144 mm. — 2 Flowers 3. Cupule rather woody, 2—5 mm peduncled by an obconical stipe, with 3-4 lamellae, split about halfway or more, the 286 FLORA MALESIANA [ser. dp vals? Fig. 5. Nothofagus nuda STEEN. a. Habit, x 24, b. mature cupule carrying 3 nuts, c. nuts, d. cupule, all <2 (PULLEN 6582). halves obovoid, c. 11-15 by 10-11 mm. Nuts not seen. Distr. Malesia: West New Guinea (Ransiki; Wissel Lake area; Balim; Habbema Camp) and East New Guinea (Western Highlands: Mt Hagen, Wankl village, Mt Kum; Eastern Highlands: Aiy- ura Subdistr., Kainantu; Southern Highlands: Anga village, Lake Kutubu), New Britain (Torlu R. and Pomio area, 5°30’-6°N, 151°-151°30’ E). In all 24 collections, of which 12 fertile (5 3, 7 9). Ecol. Often common large tree in mountain forest, 1200-2400 m, but stands are found at Ku- tubu Lake at 810 and 900 m, and full-grown trees in New Britain at 600 m, the lowest locality of the genus in Papuasia. In New Britian (Torlu R.) reported from limestone in Eugenia ridge forest. Flush twice noted: Aug., Oct. 3 Fl. May, July, Aug. Sept., Nov., fr. July, Nov., Febr. ' Common on Mt Hagen and near Wissel Lakes, reported dominant at Habbema, at 2250 m, at Agunamura near Kainantu (Eastern Highlands) in some places in practically pure stands with a high volume per acre; dominant at Telefomin on middle slopes, and at Aiyura. 1972] Uses. At Telefomin the timber is used for house posts. The Lutheran Mission at Raipinka reported the timber to be very resistant, posts having been 16 years in the ground without deterioration. About Agunamura very abundant and used for bridge-decking, fence posts, efc. The timber is hard to cut, the largest trees being 45 m by 1 m plus @; older trees are inclined to be stag-headed. Vern. Senoko, Ransiki, Manikiong lang., West New Guinea; katula, New Britain, Pomio area. Notes. The species is characterized by a narrow, elliptic leaf-shape and absence of any ridge on the sulcate midrib above; furthermore by glomeruled, almost sessile 3 triads which are often found later- ally on twigs with mature leaves as in N. carrii (but that species has only 1 @ flower per cupule). As usual in the genus leaves of saplings and young trees are large: those of a tree 16 m high measured 11 by 4 cm, but the leaf-shape is con- stant. In ScHoppE 1544 I found an abnormal flower in which the stigmas were swollen and obviously not fertile; one of the 2 flowers was flanked by the bud of a 3 flower. The nearest allied species seems to be the New Caledonian N. aequilateralis (BAUM.-BOD.) STEEN. 4. Nothofagus brassii STEEN. Blumea 7 (1952) 146; J. Arn. Arb. 34 (1953) 350, f. 8, 9. — N. recurva FAGACEAE (Soepadmo) 287 STEEN. Blumea 7 (1952) 146; J. Arn. Arb. 34 (1953) 343, f. 4, incl. var. microphylla STEEN. I.c. 345, f. 5. — N. pseudoresinosa var. microphylla STEEN. Blumea 7 (1952) 147, sphalm. — Fig. 7. Generally a large tree, 24-45 m by 25-100 cm @, sometimes dwarfed to a shrub, 2-6 m, or small tree (Arfak). Leaves elliptic-oblong or ovate to ovate-oblong, coriaceous, not rarely hard, the margin often recurved, apex sometimes acutish, (2%4-)3%-9 by 1144-4 cm (I = 114-21); ridge on midrib at least halfway sometimes to apex; nerves 7-9 pairs, above sometimes depressed, sometimes elevated, venation indistinct; petiole 4-7 mm. Sti- pules 5-10 by 214-3 mm. — 3 Flowers: triads ses- sile or almost so; perianth -- tubular, 7-9 mm. Stamens c. 15; anthers 5-7 mm. — 2 Flowers 3; cupule c. 4-5 mm g; 1-2 laterals sometimes abor- tive or sterile; style 1-244 mm. Cupule on a 5-15 mm long curved or straight peduncle, with 4—5 lamellae, thick, c. 10-15 by 12-15 mm, sometimes smaller, 7-10 by 5-9 mm (always mature?). Nut 6-10 by 4-6 mm, sometimes only 4-5 mm and orbicular. Distr. Malesia: West New Guinea (Arfak: Angi Lakes; Wissel Lakes; Habbema Camp; Mt Cycloop) and East New Guinea (Morobe Distr.: Mt Rawlinson; Eastern Highlands: Chimbu, Go- roka). In all 16 collections, of which 15 fertile. Ecol. Mountain forests, sometimes frequent PINACe Fig. 6. Nothofagus crenata STEEN. a. Twig with cupules, nearly nat. size, b—-c. cupules, <3, d. nut, x3; all immature (BRAss 11335)(Courtesy Journal Arnold Arboretum). 288 FLORA MALESIANA [ser. I, val? 7? Fig. 7. Nothofagus brassii STEEN. Mature forest at Bele R. valley, a southern tributary of the Balim R., West New Guinea, at 2450 m alt., most undergrowh and smaller trees removed as a preliminary to fel- ling; big tree on left charred at base by fire (BRASS, Nov. 1938). 1972] FAGACEAE (Soepadmo) 289 very common or dominant, 1550-2700 m. 3 FI. Oct.-Nov., April, fr. Jan.-Febr., Sept.-Nov., flush Sept. (once), Febr. (once). At Arfak Mts and Wissel Lakes frequent or very common, sometimes in old secondary forest, at Habbema dominant tree 40 by 1 m at 2300 m, at Chimbu growing together with N. perryi. Notes. More material has shown that the dif- ferences between N. recurva and N. brassii do not hold: in one specimen fully mature cupule pedun- cles may be recurved or straight and this is also true in N. perryi. Also N. recurva var. microphylla cannot be upheld; there appears to be a rather great variation in the size of the cupules, from small ones as in this variety to large ones. Also in the leaf-texture and hence degree in ele- vation of the venation there is quite some varia- tion, from very stiff hard smooth leaves with recur- ved margin to rather flattish less hard leaves. In a few specimens some very faint crenations occur, but no vein is running towards the crena- tions as in N. perryi. 5. Nothofagus pullei STEEN. Blumea 7 (1952) 146; J. Arn. Arb. 34 (1953) 353, f. 10. — ? N. cornuta STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 359, f. 3-10’, 15. A fairly large, finally often flat-topped tree, 20- 50 m by 25-100 cm g, rarely (juvenile) smaller, or dwarfed to a 2-4 m high crooked shrub on ridges. Twigs in the typical form densely patent yellow- pubescent, but with gradations to minutely puber- ulous, also petiole and midrib underneath puber- ulous. Flush generally distinctly pale-dotted with scales underneath. Leaves small, coriaceous, smooth above and with recurved margin to flat and thinner and with prominent venation, broad-elliptic to elliptic-oblong, 1-44 by 34-23, cm; ridge on midrib about halfway; nerves c. 6 pairs; reticulation dense; petiole 1-3 mm. Stipules c. 4-5 by 2 mm, tardily ca- ducous, attached at lower 1/3.— ¢ Flowers solitary, perianth rather tubular 5—6 mm long; filaments c. 10-15, connation not emerging from the perianth; anthers 3-4 mm. — 2 Flowers sessile or | mm stalk- ed, with a distinct style and often 2 mucronate shoulders. Cupule split to the base, with a single lamella, apical part often irregularly laciniate, narrow, elliptic, 244-5 by 2 mm or shorter (3 by 1 mm), shorter and narrower than the nut at matu- rity only the lower part lignified and thick. Nut acute-orbicular to elliptic, 5-6 by 344-5 mm. Distr. Malesia: West New Guinea (Arfak: Angi; Mt Hellwig) and East New Guinea (Western Highlands: Sirunki on Mt Kabanunt, Mt Hagen, Mt Kum, Mt Oga, Wabag, Kepilam village; Southern Highlands: Mt Giluwe, Tari Subdistr. foot of Mt Né; Mt Ambua; Central Highlands: Chimbu, Nondugl, Kerowagi, Mt Karigomna; Eastern Highlands: Mt Otto, Goroka; Morobe Distr.: Edie Creek, base of Mt Saruwaged, Kaindi). In all 31 fertile and 35 sterile collections. Ecol. Often gregarious or dominant, on ridges sometimes dwarfed downto ashrub of 2-4 m (type), (1650—1950—)2000-3000 m. ¢ Fl. mainly July- Aug., fr. mainly July-Aug., flush mainly July-Sept. SLEUMER & VINK found a single surviving small tree of 9 m in a pyrogenous fern-sedge vegetation near Angi Gigi (Arfak) at 2500 m. Reported also from steep limestone at Mt Lu (Kuara River Val- ley) and at Wabag. Frequently on ridges (Mt Hellwig, Edie Creek, Goroka, Nondugl) or crest of spur (Mt Otto), but also on slopes, often dominant, e.g. at Kaindi the principal dominant on the upper part of the moun- tain above the N. grandis zone (BRASS), on Mt Otto forming pure forest between 2630-2680 m on crest of spur (BRASS), at Nondugl gregarious on ridge at 1650 m, on Mt Giluwe dominant at 2500 m, and at Kepilam village at 2400 m; also dominant on Mt Hindenburg mainly on ridges, at Wabag dominant over large areas, sometimes in pure stands, at 2700 m, at Laiagam Distr. domi- nant at 2700 m, and further reported dominant at Habbema. From Wabag regeneration is reported under Castanopsis, but also dominance in large pure stands. Also in mixed forests with Litsea, Pandanus, Cinnamomum, Xanthomyrtus, Phyllo- cladus, Weinmannia, and Podocarpus. Uses. Obviously a valued timber tree appreciat- ed by the Papuans. FRODIN reports it from the vicinity of Lei Camp on the SE. slope of Mt Ambua for local bridge construction; HOOGLAND & PUL- LEN reported from Mt Hagen, 2550 m, that it is planted along houses and creeks, and occasionally along the track from seedlings brought up from lower altitude; also at Chimbu planted in garden areas. Vern. Ta(r), Laiagam, taro pulau, Wabag, taro yam yam, t. yum yum, Poget, all Enga lang., garaip, graip, gripe, karaap, Hagen, Minj lang., garap, Kubor, Minj lang., jomba, yomba, Giluwe & Mairi, Mendi lang., pu, Giluwe: Ialibu, zopa, Durantina, asaro, Kefamo, jobbailua, jowara-aiwa, Chimbu: Masul, danda, Nondugl: Bleekep, igamuk, Hinden- burg Ra. Notes. The material is not homogeneous. The species is typified by a patent, yellowish, rather stiff and persistent hairiness on the twigs, the single Malesian species possessing this, and there are quite a number of collections agreeing with the type from both West and East New Guinea. How- ever, there is a grading towards a very fine white puberulence which is only well visible on the twig- ends. As could be suspected the variation of N. pullei is only now better understood with c. 65 collec- tions at hand (c. 40% fertile), as contrasted with the one collection it was based on. In exposed situations the leaf is thick and -+- convex, smooth above, and hairiness is distinct, often yellowish. However, in small fertile trees and in less exposed conditions leaves are flatter, thinner, often larger, with raised venation on both sides, and with the indument on the innovations reduced to an extre- mely short and also sparser white puberulence. These latter often have also somewhat smaller stipules (3-5 mm). But there is a full intergradation and one cannot find minor breaks. The cupule has only 1 lamella, which is usually irregularly incised and narrower than the nut, and 290 FLORA MALESIANA [ser. I, vol. 7? the valves free. But in a few specimens the cupule is almost as wide and the valves are connate at the very base, fallen cupules appearing 2—valved (BRASs 30894, HOOGLAND & SCHODDE 7532, PUL- LEN 5400); these specimens are all very fine-puber- ulous at twig ends. I have tentatively reduced N. cornuta to N. pul- /ei; it cannot be distinguished vegetatively; this is possibly a freak without a cupule, the type speci- men (EyMA 5122) besides being extremely poor; two specimens resembling ‘N. cornuta’ are VAN ROYEN & SLEUMER 7281 and 8013, with ¢ and young 9 flowers, both from the Vogelkop. 6. Nothofagus crenata STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 355, f. 3-6’, 11. — Fig. 6. Tree to 40 m by over 1 m go. Leaves flat, thin- coriaceous, ovate-oblong, pointed towards apex and distinctly crenate, 244-5 by 114-2 cm; ridge on midrib prominent almost to apex; nerves 5—6 pairs; venation above + prominent; petiole 4-5 mm. Stipules ovate, early caducous, c. 5 by 2 mm. — ¢ Flowers solitary (only loose ones found); anthers 5 mm. — 2 Flowers solitary; immature cupules split halfway, c. 5 mm stalked, with 3-4 lamellae, about as large as the developed 2 flower, 44-6 by 5-6 mm. Jmmature nut + orbicular, narrowly winged, style distinct, c. 5 by 4 mm. Distr. Malesia: Central West New Guinea (Bele R. near Habbema Lake), only the type collec- tion. Ecol. Common in forests, 2300 m, young 2 Nov. Vern. N’gi. Note. It is peculiar that of this rather charac- teristic species no additional specimens have be- come known. 7. Nothofagus resinosa STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 356, f. 3-7’, except NGE 5134 (which is N. flaviramea). — Fig. 9 d—d?. A large tree, (15—)20-S0 m by 40-104 cm ga, young parts characteristic by a conspicuous cover of pale yellowish waxy or resinous exudate. Leaves elliptic to broad-elliptic, coriaceous, flat, rounded at both ends, margin apically not quite entire, but with minute teeth causing a fine undulation, 4-10 by 2-5 cm; upper face smooth, the midrib ridge at least up to halfway; nerves 8-10 pairs, rather parallel towards the edge, prominent beneath; petiole 5-10 mm. Stipules early caducous, elliptic, 5 by 3 mm. — § Flowers solitary, almost sessile, ++ tubular, 7 mm, puberulous. Stamens 13-15; an- thers 3-5 mm. — 2 Flowers solitary, ecupular, ses- sile, ovate-oblong, puberulous, 6-7 by 344-4 mm. Nut broadly elliptic, puberulous, 9-10 by 64-74% mm, with persistent style base. Distr. Malesia: West New Guinea (Wissel Lakes, Habbema) and East New Guinea (Western Highlands: Kubor, Lagaip, Minj, Keglsugl; East- ern Highlands: Upper Chimbu; Morobe Distr.: pees: In all 12 collections of which 6 fer- tile. Ecol. Mountain forest, often common, report- ed as dominant between 2500-2800 m at Habbe- ma and Kubor, and completely dominant at Tam- bua—Keglsugl, 2300-2850 m. Flush Febr., July- Sept. Uses. Once said to be planted by Papuans at Kepilam village, Lagaip Valley, Western Highlands from seed said to be obtained from Wabag. Must be a good timber tree of large dimension. Vern. Garuwa, Wissel Lakes, Kapauku lang., ndak, yembeh, Western Highlands, Minj lang., garaip, Togoba. Notes. Young parts are studded with sulphur- yellowish waxy resin, which tardily disappears on the underside of the leaves, a conspicuous charac- ter in common with N. pseudoresinosa, and at first sight astonishingly alike that species. The main difference with the latter is the com- plete absence of a cupule and the puberulous nut; also the g perianth, the twig-ends, petiole and stipules are more or less finely white-puberulous (not to be confused with blackish ‘hairs’ which belong to a fungus); furthermore by the usually more acutish leaf apex. One might suppose the absence of the cupule to represent a freakish state, but the 5 9 fertile speci- mens are exactly matching. In 1953, /.c., 1 added that the cupule could have a basal appendage, on the strength of NGF 5134, but this was an error, the specimen belonging to N. flaviramea. 8. Nothofagus pseudoresinosa STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 358, f. 13, 16 d— h. — Fig. 9 e-e?. Tall tree, sometimes short flanged, 30-45 m by 90 to over 100 cm go. Leaves 24%-5¥% by 14%4-21% cm, rarely 8-9 by 4-5 cm, flat, elliptic-oblong, some rather narrow, undersurface and young parts with a pale or pale yellowish, rather persistent waxy or resinous exudate; ridge on midrib up to halfway; nerves 8-9 pairs only distinct beneath, venation indistinct; petiole 5-10 mm. Stipules early caducous, 5-6 mm, attached at lower third. — $ Flowers solitary, perianth tubular, c. 7 mm, slightly puberulous or glabrous. Cupule sessile, split to base, rather narrow, valves (S5—)6-7 by (2-)3 mm, with 1-2 lamellae, somewhat shorter and narrower than the nut or + equally long. Nut solitary, ovate, glabrous, 7-8 by 4-5 mm, the smallest (mature?) 5 by 3 mm. Distr. Malesia: East New Guinea (Morobe Distr.: Mt Sarawaket; Western Highlands: Wabag along Wapu R.; Eastern Highlands: Finisterre Ra.); 3 fertile collections (CLEMENS 5849, PULLEN 6083, HOOGLAND & SCHODDE 7204). Ecol. Mountain forest, 2300-3100 m, very common, dominant on ridges and valleys at Wa- bag and the Finisterre Ra., co-dominant at Lake Habbema. Flush & fl. April, July, fr. Oct. —Uses. None known, but by the large dimension it must be a valuable timber tree. Vern. Tart, Poio, Enga lang., mépa, Finisterre Ra., Naho lang. Note. In habit extremely like N. resinosa, but different by the presence of a distinct lamellate cupule about as long as the nut and absence of the minute puberulous hairs on nut, flush and stipules. 1972] 9. Nothofagus carrii STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 359, f. 3-9’, 14. Tree, 20-45 m (bole 18-24 m), dbh 25-130 cm. Leaves usually obovate, more rarely elliptic, coria- ceous, rather flat, 2-6 by 1—3 cm; ridge on midrib in lower half; nerves 5—7 pairs, rather indistinct on both sides; reticulations not prominent; petiole 2%-5 mm; very young flush often waxy. Stipules early caducous. — Flowers sessile in 1-3 mm pe- duncled triads -- crowded on very short (to 10 mm) efoliate twiglets in the axils of normal leaves; perianth hypanthium-like contracted at base, cam- panulate-urceolate, c. 344-4 mm. Stamens c. 10; anthers 2144-3 mm. — @ Flowers solitary, 1-114 mm peduncled, only sustained by an elamellar flap much smaller and narrower than the flower, 2-3 mm long; flower itself ovate-oblong, 5 by 3 mm; style c. 1-144 mm. Nut elliptic to ovate- oblong, 7-11 by 4-5 mm. Distr. Malesia: West New Guinea (Arfak, Wissel Lakes), East New Guinea (Ok Denim R.; West Sepik: Telefomin; Western Highlands: Laia- gam, Mt Giluwe, Minj-Nona Divide; Southern Highlands: Ialibu, Anga Valley, Habono; Morobe Distr.: Kaindi, Sarawaket, Goroka; Eastern High- lands: Kerowagi, Bundi Gap), and d’Entrecas- teaux Is. (Normanby & Goodenough Is.). In all 21 collections, of which 14 fertile. Ecol. Mountain forests, sometimes low mossy forest, or shrubby growths on summits or ridges, 1900-2850 m. Fi. Dec., Jan., also flush June-Aug. On Normanby I. at 900 m on summit shrubby growths the major constituent 2—5 m tall; on Mt Kaindi in low mossy forest also only 3 m. Forming extensive forests at Bundi Gap at 2700 m and at Laiagam (Kandep Road) the dominant tree at 2850 m, but also found dominant at Ialibu at 1950 m and Habono at 2100 m. Uses. None reported, but it must be a valuable timber tree of often great dimension. Vern. Dierie = didamé, Wissel Lakes, Kapau- ku lang., kundap, West Sepik, Telefomin, yomba, Southern Highlands, Mendi lang.; Eastern High- lands, Bundi Gap, Chimbu lang., taggiruba, South- ern Highlands, Habono, taro, Western High- lands, Laiagam, Enga lang., garaip, Hagen. Notes. Vegetatively some specimens show re- semblance to N. starkenborghii by the fine tessel- late-reticulate venation and sometimes an almost absent ridge on the midrib above, and + elliptic leaves; furthermore both species share the charac- ter of clustered ¢ triads. The cupule under the solitary nut is only repre- sented by 2 small, elamellate flaps, and this is consistent with all 2 fertile collections. 10. Nothofagus flaviramea STEEN. Nova Guinea, n.s. 6 (1955) 281, f. 1. — N. resinosa (non STEEN.) STEEN. J. Arn. Arb. 35 (1954) 267, f. 1 f. — Shorea sp., SLOOT. Reinwardtia 2 (1952) 61. Tree, 15-45 m by 25-150 cm g, sometimes but- tressed; bole 10-25 m. Twigs usually + flattish, and sulphur-yellow, rather coarse. Leaves ovate- oblong, acutish attenuate to short-acuminate, en- FAGACEAE (Soepadmo) 291 tire, flat, usually chartaceous, c. 5—12 by 234-5 cm, at base broadly acute to rounded, the sulcate midrib above with a ridge; nerves 8-10 pairs, rather prominent on both sides as are the reticula- tions; petiole c. 5-10 mm. Stipules early caducous. — §$ Flowers in + sessile triads; perianth c. 5 by 21%, mm, very truncate. — 2 Flowers solitary; cu- pule sessile, consisting of 2 free, tiny, suborbicular flaps without lamellae; in fruit c. 1-2 mm, cadu- cous. Nut obovate-apiculate, when mature c. 9-10 by 6-7 mm. Distr. Malesia: West New Guinea (Fak Fak, Arfak, Mt Cycloop, Bernhard, Habbema, Wissel Lakes, Star Mts) and East New Guinea (Sepik: Telefomin; Western Highlands: Aweta Bridge, Minj; Finschahfen; N. Div.: Hydrographers Ra.; Eastern Highlands: Kainantu). In all 35 collec- tions of which 10 fertile. Ecol. Mountain forest, sometimes mossy, on slopes and spurs, 750-2450 m (mainly 1000-2000 m). Fr. April, June, Oct., Dec., flush Oct.-Dec. Common on Mt Arfak (Genofa, etc.), dominant on Aifat R. (Arfak) at 1400 m, and near Trist (Morobe Distr.), on Mt Cycloop at Tamrau co- dominant with Castanopsis acuminatissima at 1350 m, on slopes of the Stars Mts dominant with Araucaria. Once reported on limestone at Chimbu (SIMONETT 69). Uses. Seems to be a good timber tree, often of large dimension. Vern. Essamene, Fak Fak, Argunilang., snokko Arfak, Anggi Gigi, gripe, Hagen, diedame, Wissel Lakes, Kapauku lang. Notes. In KALKMAN 4266 I found in a single cupule 2 aborted 2 flowers besides the central fertile one. Related to N. womersleyi which has, however, long-peduncled cupules, with a distinct large inner lamella. See further the note under N. grandis. 11. Nothofagus grandis STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 363, pl. 1, f. 17, 18. — Fig. 3, 9 c—c®. Large tree, (12—)25-45 m by (18—)45—150(—250) cm @ (bole 10-25 m), sometimes slightly buttressed. Twigs terete, often pale greyish, lenticellate. Leaves elliptic-oblong, often dull greyish above and pale brown beneath, 414-10 by 2-5 cm; ridge on mid- rib at least halfway; midrib red; nerves c. 7-9 pairs, as the veins somewhat prominent on both sides; petiole 3-10 mm. Stipules 5—7 by 3144-4 mm. — $ Flowers orange, in 2-9 mm peduncled triads, perianth sessile, tubular, c. 6-7 mm. Stamens yellow to red, 10-17; anthers 4-5 mm. — ° Flowers solitary, surrounded by a cupule as large as the flower already when in anthesis, orbicular, c. 9 mm 9. Cupule in fruit large, woody, 13-17 by 8-14 mm, provided with (2—)34 lamellae, sessile or rarely on a 1-4 mm long stalk. Nut rhomboid or mostly orbicular, 7-10 by 944-10 mm. Distr. Malesia: West New Guinea, rare or uncertain (twice: Kebar, BW 10414, Wissel Lakes, BW 3260), one of the commonest species in East 292 FLORA MALESIANA [ser. 1, veney- New Guinea; in all 52 collections of which 22 fertile. Ecol. Mountain forest, 1350-2600 m. 3 FI. mostly June-July, fr. almost in all months, flush May-Aug., Dec.-Jan. Not seldom very common and forming massive forests e.g. in Western Highlands (Wabag), Cen- tral Highlands (Goilala, Wahgi Divide), Southern Highlands (lalibu, dominant at 2000 m; Mt Gilu- we, S. slopes, dominant on dissected ash plains, with Pandanus, a single time as a shrub, 2 m, ScCHODDE 1450), Eastern Highlands (Al R.; Non- dugl in almost pure stands at 2100 m), Mt Hagen (common, Aiyura, Wau), Morobe Distr. (Edie Creek; Kaindi, overwhelmingly dominant, 2000- 2400 m), Milne Bay Distr. (Mt Dayman, dominant from 1800 m to summit), sometimes associated with Phyllocladus. Uses. Planted at Bomkan village (Upper Chim- bu) for ornament around villages, E. Mt Wilhelm, for ? ornament, seedlings brought from Bundi Gap, and Bismarck Ra., ditto near native homesteads and in garden lands. Vern. Peil, Mendi, ufoiya, Anona, ifoya, Aiyura, unuza, Kamano, graip, gripe, Hagen, jomba, Chim- bu: Masul, dzopa, Asaro: Kefamo, ififi, Fiyugi: Oriko, taro, t. korn, t. porgere, t. pulen, Wabag, Enga lang., kaarao, Nondugl, traf or trap, Telefo- min, iew, Kebar, Nettoti Mts, diri, Wissel Lakes, Kapauku lang., arape, membi, peti, Giluwe: Mendi. Notes. In 1953 I have added two sterile collec- tions from West Guinea (BRAss & VERSTEEGH 13553 & 13147) which I now believe belong to N. flaviramea; the latter has always rather large, ovate-acutish, flattish leaves with the ridge on the midrib running to the apex, and usually yellowish twig bark, not grey and lenticellate as in N. grandis. It appears, however, that certain specimens which certainly belong to N. grandis may have acutish leaves, although mostly of more elliptic shape; these are usually from pole trees or saplings (e.g. ANU 2814, NGF 18291). In N. grandis the midrib is sulcate with a prominent ridge, in N. flaviramea the ridge is on the flat leaf surface. Its scarcity in West New Guinea is remarkable. 12. Nothofagus rubra STEEN. Blumea 7 (1952) 147; J. Arn. Arb. 34 (1953) 368, f. 20. — N. eymae STEEN. lI.c. 147; l.c. 370, f. 3-15’, 21. — N. dura STEEN. lI.c. 147; I.c. 371, f. 22. — N. bernhardii STEEN. /.c. 147; l.c. 361, f. 16 a—c (excl. d—h). — N. decipiens STEEN. I.c. 147; l.c. 367, f. 19. Tree, 17-45 m, 25-65 cm g. Perules rather long- persistent. Leaves ovate-oblong to elliptic, round- ed or even notched at both ends,' coriaceous to hard-coriaceous, entire, flattish but often bullate, Fig. 8. Nothofagus cf. rubra STEEN. Poorly developed, small tree (see Papuan in front of it) in semi- devastated terrain with macchia-like heath, on exposed hillock at 1700 m, Wissel Lakes, West New Guinea (RAPPARD, Oct. 1955, to be related to BW 3302). 1972] FAGACEAE (Soepadmo) 293 Fig. 9. Nothofagus womersleyi STEEN. a. Habit, x 73, a1. cupule, with 1 nut, x2. — N. perryi STEEN. b. Young @ flower, b1. cupule, b?. nut, all x 2. — N. grandis STEEN. c. ¢ Mature bud, c!. triad in anthesis, c?-c+. mature cupules, c'-c®. mature nuts, all x2. — N. resinosa STEEN. d. Habit, with nut, x 4, d‘. nut (probably mature; without cupule), x2. — N. pseudoresinosa STEEN. e—e!. Cupule in lateral and dorsal view, e?. nut, all x2 (a-a! BW 10320, b Ropsins 511, b1-b? PULLEN 5420, c—ct HOOGLAND & PULLEN 5856, c2—c? NGF 20212, c*-c® NGF 6050, c& NGF 4823, d-d! SAUNDERS 765, e PULLEN 6083, el_e? HOOGLAND & SCHODDE 7204). 294 2%-91% by 1144-4% cm, midrib with a ridge well over halfway, upper surface rather smooth with no prominent tessellation but the c. 5—7 pairs of nerves bluntly prominent, undersurface with fine- tessellate but not prominent venation; petiole 2—5 mm. Stipules oblong, early caducous, 4-9 by 2-4 mm. — 3 Flowers in + sessile triads; perianth c. 5—6 mm long, the central flower sometimes pedi- celled and shorter. Stamens 10-12. — 2 Flowers solitary, slightly winged. Cupule usually sessile but sometimes shortly (up to 7 mm) stalked, at least halfway split; valves rather woody, sometimes unequal, with 2-3 lamellae, rather variable in size, gaping when mature 7-10 by 7-14 mm. Nut orbi- cular to broad-ovate, c. 4-6 mm @. Distr. Malesia: West New Guinea (Nettoti, Arfak, Cycloop, Bernhard, Habbema, Wissel Lakes, Star Mts), East New Guinea (Western Highlands: Minj, Kubor, Wabag; Southern Highl- lands: Tari, Onim, Giluwe, Kufubu, Mt Né) and d’Entrecasteaux Is. (Normanby). In all 36 collec- tions, of which 14 fertile Ecol. Mountain forest, sometimes in poor situ- ations a shrub (Enarotali, 4 m, in boggy heath- land at 1800 m), in boggy forest (Wabag, Ialibu, Onim), on exposed summits or ridges reduced to a stiff gnarled shrub 2-5 m and sometimes major component (Bernhard at 2150 m; Normanby in stunted open Dacrydium forest at 850 m), not rare in mossy forest (Normanby at 950 m; Nettoti Mts, main constituent at 1700 m; Idenburg R. at 2150 m), 1700-2850 m, but on Normanby I. 750-850 m. 6 Fl. mostly June-Aug., but also noted Febr., May, Oct., fr. mostly July-Sept., but also Jan.-March, flush May-Aug., also Febr. A common species often abundant (Bernhard, Arfak, Wissel Lakes), dominance noted on Mt Cycloop above the Castanopsis zone, on Nettoti Mts in submossy forest, at Habbema at 2840 m, on the Kutubu-Tari track at 1950 m. Uses. Obviously an excellent timber tree, found planted on the Upper Chimbu from seedlings brought from the Bundi Gap. Vern. Diedamé, diri, Wissel Lakes, Enarotali, Kapauku lang., snokko, Arfak, Manikiong lang., pémmeém, Arfak, Hattam lang., taro, Wabag, Enga lang., yomba, Giluwe, Mendi lang. Notes. Vink found the first leaves on a seed- ling spirally arranged, but already on the first lateral branchings distichous. Through the increased collections I felt com- FLORA MALESIANA [ser. I, volrg? pelled to combine five specific names, which were based on one or few specimens in different onto- genetic stages. Experience with N. grandis learned that a smallish stalk under the cupule is of no taxonomic importance. In N. rubra the cupule is usually sessile, in N. bernhardii and N. eymae short- stalked, and in N. decipiens (immature) distinctly stalked, and I have come to the conclusion that in N. rubra there is a situation similar to that in N. grandis. In sterile state N. rubra is hard to distinguish from N. brassii. 13. Nothofagus womersleyi STEEN. nov. sp. — Fig. 9 a-al. Nothofagi flavirameae affinis, differt: cupulae longe pedunculatae, dimidiis liberis e squama basali unilamellata unica compositis, lamella achenam ovato-oblongam semiaequante. — Typus: BW 10320 VERSTEEGH, holotype L, isotypes A, CANB, K, LAE. Tree, c. 20 m, dbh 40 cm 9; young twigs dark- red with elongate yellow dots; twigs terete, those of innovations + flattened; glabrous. Leaves ovate- oblong, rounded at base, pointed and + acumi- nate to apex, entire, c. 5-9 by 244-4 cm, rather flat, midrib -- convex, sulcate above with the ridge running rather high up; nerves 7-9 pairs, venation above not or little prominent but rather well so beneath; petiole 5-8 mm. Stipules peltate, ellipsoid, c. 5mm. — g Unknown. — 2: Cupule c. 10-15 mm peduncled, apart from the two basal bracts with only one rather apical lamella, immature rather thin, about half as long as and narrower than the single nut; valves almost free to the base, but some- times (mostly unilaterally) connate for 3 mm. Nut (not fully mature) flat, ovate-oblong, glab- rous, distinctly (1 mm) winged towards apex, 7-10 by 5-614 mm; style 0; stigmas 2. Distr. Malesia: NW. New Guinea (Vogelkop Peninsula: Watjetoni Mts near Kebar Valley), one collection. Ecol. Primary forest on peaty soil, common, 1200 m. 2 Nov. Vern. Jew, Kebar lang. Notes. General shape of leaf resembling that of N. flaviramea, which has, however, an obovate nut and a much smaller, sessile cupule consisting only of a tiny elamellate flap. Cupule and nut are described from the immature stage of the type. 2. CASTANOPSIS SPACH, Hist. Veg. Phan. 2 (1842) 185; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 103, 118; BENTH. & Hook. f. Gen. Pl. 3 (1880) 409; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 93; SCHNEIDER, Illust. Handb. Laubh. 1 (1906) 159; REHD. & WILS. in Sargent, Pl. Wils. 3 (1916) 97; A. Camus, Chat. (1930) 243; HickeL & A. CAMUs, Fl. Gén. I. -C. 5 (1930) 1007; SoppADMo, Reinwardtia 7 (1968) 384. — Callaeo- carpus Mig. Pl. Jungh. (1851) 13. — Chrysolepis HyELMQvisT, Bot. Notis. Suppl. 2, 1 (1948) 117; FoRMAN, Kew Bull. 18 (1966) 425. — Fig. 12-15. 1972] FAGACEAE (Soepadmo) 295 Trees of medium to large size. Innovations densely yellowish brown to fulvous pubescent or stellate-hairy. Terminal buds ovoid to ellipsoid, scales ovate to linear. Stipules extrapetiolar, deltoid to linear-acute, caducous. Leaves spirally arranged, rarely with a tendency to distichy, thin- to thick-coriaceous, concolorous to dis- tinctly discolorous, glabrous or beneath hairy or scaly, margin entire or rarely (in non-Malesian species frequently) serrulate in the apical half. Petiole more or less thickened at the base, terete or flat or shallowly furrowed adaxially. Inflorescence erect, male, female, androgynous, or mixed, including bracts and bracteoles dense- ly stellate-pubescent. Male rachis slender to rigid, solitary in the axil of a lower leaf or in dense paniculate clusters on the lateral or subterminal new shoots, mostly simple; 3 flowers solitary or in clusters of 3-7 along the rachis or in an androgynous inflorescence along the upper part of the rachis; perianth cam- panulate, (5—)6(-7)-lobed; stamens (10—) 12(-15), filaments filiform, glabrous, anthers dorsifixed, reniform, 0.25—0.35 mm long, 2-celled, 4-lobed, lengthwise dehiscent; pistillode rudimentary (usually less developed than in Lithocarpus), more or less rounded-triangular, densely woolly pubescent. Female, androgynous, or mixed inflorescence simple, rigid, solitary in the axil of a higher leaf or on the upper part of the paniculate cluster; 9 flowers solitary or in clusters of 3-7 along the rachis!, or in an androgynous inflorescence along the basal part of the rachis, perianth campanulate, (5—)6(—7)-lobed; staminodes 10-12, rudimentary; styles 3-5, conical to terete and slender, connate to recurved, densely hairy at the base, stigmas terminal and punctiform. Cupule completely enclosing the 1-7 fruits except for the persistent styles and perianth, at maturity splitting into a regular number of segments (valves) or irregularly, variously spiny or tubercled or with a few undulating ridges, the spines simple or in bundles or stellate, on the 2-4(-8) cupule segments with the sutures in between. Fruit more or less rounded with the adjoining sides flat, wall bony to woody; scar small to almost occupying the entire surface, glabrous, dull, rugose; the remaining part free from the cupule, densely yellowish brown to fulvous tomentellous or glabrous. Cotyledons flat-con- vex. Germination hypogeal. Distr. In her monograph, A. CAMus recognized not less than 120 species. The distribution covers NE. India (Nepal, Bhutan, Assam), Burma, China (except in the N. and NW. parts),Korea, Japan, Formosa, Hainan, Indochina, Siam, Malesia, and SW. North America (1 sp.). The greatest assemblage and most primitive forms are found in SE. Asia (Indochina and Malesia). In Malesia 34 spp. (see fig. 11); the genus has been recorded from most islands except in the eastern half of Java and in the Lesser Sunda Islands; the largest number of species is found in Borneo. The only representative of the genus outside Asia, namely C. chrysophylla from SW. North America, has been recognized by HyeLMavist (1948) and For- MAN (1966) as a separate genus, ChArysolepis, an opinion which [ cannot share. Fig. 10. Fossil records. Very little is known about the geological history of the genus. In Europe two doubtful records were made, viz a wood fragment recovered from the Tertiary Brown Coal deposits in Germany (SCHONFELD, Ber. Freiberg. Geol. Gesch. 10, 1925, 18-24), and several pollen grains discovered in the London Clay and Bembridge Floras (CHANDLER, Publ. Brit. Mus. Nat. Hist. 1964, 1-151). In Asia the fossil record of the genus is even more meagre. OGuRA (Jap. J. Bot. 14, 1949, 15-18) reported the finding of a wood fragment resembling Castanopsis from the Tertiary of Nagano Prefecture, Japan, and MULLER (Proc. 10th Int. Bot. Congr. Edinb. 1964, 271) recorded several pollen grains of Castanopsis-type from the Tertiary of North Borneo. In Australia and North America there have been several records (leaf- impression, pollen grains) from various Tertiary deposits. The identity of these records, however, should be considered with great caution, as the genera of Fagaceae are not always easily distinguishable ty the characters of the wood, pollen grains or leaf alone. a In species with more than 1 2 flower per cupule, there is a tendency that the upper cupules on a rachis have less or even only 1 2 flower (and of course fruit). 296 FLORA MALESIANA (ser. d-e@hay? Fig. 10. Distribution of the genus Castanopsis, the fossil records indicated by triangles. Ecol. In Malesia the genus occurs from the everwet lowland to the montane forest, between 0-2500 m, mostly between 1000-1500 m, and on various types of soil, except limestone. On the whole Castanop- sis shuns a seasonal climate. For this reason it is absent from Central Java, except in a few local everwet rain-forest complexes (Mts Muria, Ungaran, Telomojo), East Java, and the Lesser Sunda Islands. C. buruana is largely a rain-forest species, but has been observed to survive locally in a seasonally dry area in SW. Ceram which is subject to fire. Of the 34 spp. recognized, only 12, viz. C. borneensis, catappaefolia, curtisii, fulva, inermis, johorensis, lucida, malaccensis, motleyana, oligoneura, pedunculata, and wallichii, are apparently confined to lowland forest (below 500 m); the rest occurs both in the lowland and the sub- montane or montane forests. Most species start flowering during the rainy season, the ripe fruit being developed about six months later. In New Guinea, C. acuminatissima sometimes forms an almost pure stand, and occurs commonly as a codominant together with Anisoptera (Dipt.), Lithocarpus or Notho- fagus, between 1000-1500 m. KEY TO THE SPECIES? (based on fruiting specimens) 1. Cupule enclosing one fruit only. 2. Greater part of the fruit adnate to the cupule. 3. Cupule almost smooth or beset with short prismatic tubercles. 4. Cupule almost smooth; peduncle c. 1 cm. Leaves glabrous on both surfaces . . 1. C. curtisii 4. Cupule beset with short prismatic tubercles except on the adaxial side, sessile to almost so. Leaves densely adpressed stellate-tomentose beneath .............. 2.C.nephelioides 3. Cupule covered with various types of spines. 5. Spines 14-1 cm long, simple, sturdy, flat. 6. Spines densely but irregularly arranged. 7. Cupule more or less globose. Petiole 44-14% cm. 8. Leaves glabrous or sparsely stellate-tomentose. 9. Cupule sparsely set with short (c. 44 cm) spines. Petioles flat adaxially; reticulation dense, SCalarifOrm) — ocukty ve beacudear tis ier aoe Se TON) <) oleic cease Bcc Is 1 id a EEE 9. Cupule densely set with spines 1 cm long. Petiole adaxially sulcate; reticulation areolate or subscalariform. 10. Leaves glabrous on both surfaces; midrib and nerves thin on both surfaces; reticulation areolate. jj donies6 ps Rene Asatienaiéy Ue, kok anal SM SMa sy cotati Se ples gage tek Ome ECC ISES, 10. Leaves sparsely stellate-tomentose beneath; midrib and nerves strongly prominent beneath, impressed above; reticulation lax, subscalariform. . . ...... .. 5.C. johorensis (1) The term ‘tomentose’ is used for any indument which is continuous interwoven and does not show the parenchyma beneath, irrespective of its thickness and the size of the hairs; the latter may be of mi- croscopical dimension and the tomentum very thin. The size of the cupule is including the spines. 1972] FAGACEAE (Soepadmo) 297 8. Leaves densely reddish brown stellate-tomentose. ........... +. 6.C. wallichii 7. Cupule ovoid-ellipsoid. Petiole 2—3 cm. 11. Leaves elliptic- to ovate-lanceolate, ei pale aly: brown adpressed stellate-tomentose; nerves 5-7 pairs .. BS Micha laecisni) Ualara einla ee Pet Ge OUSOnenra 11. Leaves elliptic- oblong, glabrous; nerves qe 13 pairs cmt. Ate Ge & oh Rey 8s GC. endertii 6. Spines arranged in regular arching rows with gaps in between. 12. Leaves beneath densely set with reddish brown, adpressed, stellate and woolly simple hairs. 13. Leaves thick-coriaceous, se nerves 14-16 pairs, dense, reticulation distinct beneath. Pet- iole 2-3mm thick ... ‘ . . 9. C. densinervia 13. Leaves thin-coriaceous; nerves lax, 9- 10 pairs; reticulation obscure o on both surfaces. Petiole mom! thick . . . : CPSi 2 Fee areca. Me eee aOs: © pancispind 12. Leaves glabrous or almost s so. 14. Cupule 2-214 by 14% cm; spines 44-1 cm, simple or in bundles. Leaves with a more or less areolate reticulation .. . ified 26 AADAC. clemensii 14. Cupule up to 5 by 3 cm; spines rigid, flat, always simple. ‘Leaves with a scalariform reticula- tions aukedl Phe BE £6 625 GC: oviformis 5. Spines much branched, ‘needle- shaped, 1 AOU, cm long. 15. Cupule 4-6 by 3-5 cm. 16. Cupule ovoid-globose. Leaves ovate-elliptic, 344-10 by 214-5 cm, underneath densely set with reddish brown, adpressed, stellate hairs; nerves 7-9 pairs. . . . . 13. C. malaccensis 16. Cupule obovoid-ellipsoid. Leaves ovate-oblong or elliptic- oblong, 10-17 by 3144-6 cm, under- neath with a dense cover of sordid silvery grey, adpressed, stellate hairs, or subglabrous; nerves 11-19 pairs. 17. Leaves densely set with sordid silvery grey, as stellate hairs. Cupule 5—6 by 3-4 cm. Fruit wall thinnerthanimm. . . . 14. C. tungurrut 17. Leaves subglabrous. Cupule 44% by 3- A Vy cm. Fruit wall 2- 3 mm thick . =. lS: Cevansi 15. Cupule 7-10 by 5-7 cm. 18. Leaves densely set with yellowish to greyish brown adpressed, stellate hairs and erect, simple or stellate hairs beneath; reticulation scalariform, distinct beneath. Spines set in 6-8 regular, MELESPACEGCURVEG LINES) carta: Menno mrenneens . . . 16. C. hypophoenicea 18. Leaves with a dense cover of reddish brown or greyish brown adpressed stellate scales under- neath; reticulation obscure on both surfaces. Spines densely but irregularly set. 17. C. megacarpa 2. Greater part of the fruit free from the cupule. 19. Cupule ovoid-globose or discoid, See 144-5 by ak cm. Branchlets densely set with large, warty lenticels .. . = S18. C.yavanies 19. Cupule asymmetrically ovoid, maarially ‘flat, 0. cml 1%, by 1-2 cm. etameniee densely or sparsely set with minute lenticels. 20. Cupule sparsely set with simple, sharp spines. Petiole 1-114 cm, adaxially sulcate. 19. C. buruana 20. Cupule irregularly tuberculate. Petiole 14-1 cm, adaxially flat . . . . 20. C. acuminatissima 1. Cupule enclosing more than one fruit, usually three. 21. Cupule densely or sparsely set with various types of spines. 22. Leaves glabrous, reticulation irregular. 23. Spines slender, much-branched, regularly set. . ............ 21. C. foxworthyi 23. Spines sturdy, simple, densely but irregularly set. . . ..... .. .. 22. C. schefferiana 22. Leaves densely set with various types of hairs. 24. Leaves linear-lanceolate, 5-9 by 1-144 cm. . . . . 23. C. microphylla 24. Leaves ovate-elliptic or lanceolate-oblong, (7— )10-40 (50) by (Oy 5— ~)3- er 19) cm. 25. Leaves 35-50 by 14-19 cm; nerves 23-27 pairs; petiole 1-144 cm, %-34 cm @ 24. C. catappaefolia 25. Leaves 10-23 by 5-10 cm; nerves 9-21 pairs; petiole (4%—)1—2 %(-3) cm, (1-)2-3(-5) mm @. 26. Leaves densely set with silvery, glaucous, adpressed stellate hairs or scales. . 25. C. argentea 26. Leaves with a dense cover of yellowish to reddish brown adpressed, stellate and woolly or erect hairs. 27. Petiole c. 4% cm. Spines and cupule densely reddish brown tomentose . 26. C. scortechinii 27. Petiole 1-3 cm. Spines more or less glabrous. 28. Leaves elliptic-oblong, 9-17 by 3-6 cm; densely fulvous stellate and woolly pubescent be- neath; petiole adaxially flat; nerves fewer than 15 pairs . . . Pee C-TGen TE 28. Leaves ovate-elliptic, obovate-oblong, or oblong-lanceolate, 14-23 i 5-10 cm, densely set with yellowish to reddish brown adpressed stellate or simple hairs, or almost glabrous; nerves usually more than 15 pairs. 29. Leaves densely set with reddish brown, adpressed stellate hairs and woolly stellate hairs; reticulation fine, dense obscure on both surfaces. Cupule 2-3 (-4) cm through. 28. C. costata 298 FLORA MALESIANA [ser. I, vol. 7? 29. Leaves densely set with yellowish to greyish brown adpressed stellate hairs and simple or stellate erect hairs; reticulation lax, subscalariform, distinct beneath. Cupule 3-5 by 3-6 cm. 29. C. motleyana 21. Cupule set with short tubercles to almost smooth. 30. Leaves glabrous. 31. Cupule almost smooth, peduncle 1-1%cmlong..........- 30. C. pedunculata 31. Cupule short-tuberculate; the tubercles sometimes recurved; sessile or on short peduncle. 31. C. psilophylla 30. Leaves densely set with adpressed stellate hairs beneath. 32. Cupule not transversely ridged. Leaves thin-coriaceous; petiole shorter than 2 cm and thinner than 2 mm. Branchlets sparsely lenticellate. 33. Leaves elliptic-oblong or elliptic-lanceolate, apex rounded or abruptly acute; nerves 10-15 pairs. Cupule and tubercles woody . . . 32. C. inermis 33. Leaves ovate-elliptic to ovate-lanceolate; apex gradually acute to 2 cm acuminate; nerves less than 10 pairs. Cupule thin, tubercles soft, sometimes recurved . 33. C. philipensis 32. Cupule with 3—4 distinct, bold, rounded ridges, set with short but sturdy spines. Leaves thick- coriaceous; petiole up to 3% cm long, 2-3 mm thick. Branchlets densely set with large warty lenticels 1. Castanopsis curtisii KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 107, t. 103; GAmBLE, J. As. Soc. Beng. 75, ii (1915) 405; Ripi. Fl. Mal. Pen. 3 (1924) 392; A. Camus, Chat. (1930) 466, t. 68: 5-6; SOEPADMO, Reinwardtia 7 (1968) 389. Tree, c. 20 m by c. 30 cm g. Innovations densely set with short stellate hairs. Branchlets glabrous, with minute lenticels; terminal bud ellipsoid, 2-3 by 1 mm; scales linear. Stipules linear-acute, 5—7 by 1 mm, soon caducous. Leaves chartaceous, lanceo- late- or elliptic-oblong, 6-13 by 2-5 cm, entire; surfaces more or less concolorous, glabrous; base rounded and abruptly acute, slightly asymmetrical, apex 5-10 mm acuminate; midrib and nerves prom- inent beneath, slightly so above; nerves 10-13 pairs, parallel, ascending, at an angle of 60—70° with the midrib, arcuating towards the margin; reticulation fine, dense, scalariform, distinct be- neath; petiole 5-10 mm, slender, glabrous, slightly thickened at base, adaxially flat. Cupule excentrical- ly pear-shaped, adaxially flat, 2-3 cm 9, gradually tapering towards the 1 cm long peduncle; wall 4%4- 1 mm thick; surface sparsely set with patches of flattish tubercles except on the adaxial side, inde- hiscent. Fruit solitary, pear-shaped or reniform, —— ~ - ——s Fig. 11. Species density of Castanopsis in Malesia, above the hyphen the number of non-endemic species, below the hyphen the number of endemic species in the island or island group. 34. C. lucida excentric, c. 3 by 2 cm, wall woody, c. 1 mm thick, completely adnate to the cupule. Distr. Malesia: Malay Peninsula (Selangor, Penang). Ecol. In lowland forest up to 300 m. FI. fr. Aug.-Sept. Notes. Very close to C. nephelioides and C. rhamnifolia, but differing from both by its glabrous leaf and by its almost smooth, peduncled cupule. The inflorescence is still unknown. 2. Castanopsis nephelioides KING ex Hook. /. FI. Br. Ind. 5 (1888) 624; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 106, t. 102; GAmBLE, J. As. Soc. Beng. 75, ii (1915) 464; A. Camus, Chat. (1930) 467, t. 69: 1-2; SoppADMO, Reinwardtia 7 (1968) 396. Tree, 10-20 m by 20-60 cm go. Innovations den- sely set with yellowish brown woolly hairs. Branch- lets glabrous, densely or sparsely minute-lenti- cellate; terminal bud ovoid-ellipsoid, 3-4 by 2 mm; scales narrowly ovate-acute. Leaves thin-coria- ceous, Ovate-elliptic or ovate-lanceolate, 8-15 by 3-6 cm; surfaces more or less discolorous, above glabrous, dull greenish grey, underneath with a dense cover of adpressed stellate hairs; base round- ed and abruptly acute, margin entire, recurved, apex acute or abruptly 5 mm acuminate; midrib and nerves prominent beneath, flattish or impres- sed above; nerves 8-14 pairs, parallel, at an angle of 45-70° with the midrib, arcuating but not anas- tomosing towards the margin; reticulation fine, dense, subscalariform, obscure on both surfaces; petiole slender, 5-10 mm, glabrous, adaxially furrowed. Male rachises solitary or in paniculate clusters, 10-20 cm; bracts and bracteoles ovate- acute; d flowers in clusters of 3-7; perianth 6— lobed, lobes acute, 1-114 mm, densely rufous- tomentose outside, stamens mostly 12, filaments 2-3 mm, anthers c. ¥% mm long, pistillode subglo- bose, c. 1 mm @. Androgynous rachis 10-20 cm; 2 flowers solitary; perianth 6—-lobed, staminodes 12, rudimentary, styles 3, conical, 1-144 mm long. Infructescence c. 10 cm, rachis 2-3 mm @, densely minute-lenticellate, carrying 1-3 cupules. Cupule 1972) sessile, asymmetrically obovoid-globose or pear- shaped, adaxially flat, 2-344 by 144-2 cm; wall 3-4 mm thick; surface, except for the adaxial side, densely set with short, woody, pyramidal tuber- cles. Fruit solitary, obovoid or pear-shaped, 11% by 2% cm, wall 144-1 mm thick, rugose, completely adnate to the cupule. Distr. Malesia: Malay Peninsula (Kelantan, Kedah, Selangor, Perak, Pahang, Malacca, Jo- hore). Ecol. Forests, 60-1200 m. F/. Febr.-June, /r. July-Dec. Note. Close to C. rhamnifolia but differing by its cupule with shorter but more rigid tubercles, and by its thicker indument on the undersurface of the leaf. 3. Castanopsis rhamnifolia (M1g.) A.DC. Prod. 16, 2 (1864) 113; Hook. f. Fl. Br. Ind. 5 (1888) 624; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 105, t. 100B; A. Camus, Chat. (1930) 469, t. 69: 3-5; SOEPADMO, Reinwardtia 7 (1968) 402. — Quercus rhamnifolia Miq. F\. Ind. Bat. 1, 1 (1858) 853. — Callaeocarpus rhamnifolia (MiQ.) Mia. Sumatra (1861) 353. — Castanea rhamnifolia (Miq.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 378. — C. pachycarpa A. CAMus, Bull. Mus. Nat. Hist. Paris II, 6 (1934) 92. — Fig. 15 g-i Tree, 10-25 m by 15-40 cm @; bark surface smooth, grey. Branchlets initially densely set with short stellate hairs, later subglabrous, dark grey, finely fissured or lenticellate; terminal bud ovoid- globose, 2-3 by 1-2 mm, scales ovate or linear- acute. Leaves chartaceous, ovate-elliptic, (5S—)8—10 (-13) by (2—)3-4(-5) cm, entire; surfaces more or less concolorous, above glabrous, dull or glossy, underneath with sparse stellate hairs, acute or abruptly 44-1 cm acuminate; base rounded and abruptly acute; midrib and nerves thin, prominent on both surfaces, stronger beneath; nerves 8-11 pairs, parallel, ascending, at an angle of 60—70°, arcuating and anastomosing near the margin; reticulation fine, dense, scalariform, obscure on both surfaces: petiole 44-1 cm, 1 mm 9, adaxially flat. Male rachises 5-10 cm by 1-2 mm, in dense paniculate clusters; bracts and bracteoles ovate- acute, 1-144 mm, densely tomentose outside; 3 flowers in clusters of 3; perianth deeply incised, lobes 6, ovate-acute, 144-2 by 0.7 mm, densely tomentose outside, stamens 12, filaments 3-4 mm, anthers 44 mm long, pistillode subglobose, 1.2-1.5 mm 9. Female or androgynous rachises c. 5 cm by 1—2 mm; bracts and bracteoles ovate-acute, 1-1 yy, mm; 2 flowers solitary; perianth 6—-lobed, stami- nodes 12, rudimentary, styles 3, conical, 1-114 mm long. Young cupule ovoid-globose, densely set with short tubercles. Ripe cupule sessile, asymmetri- cally depressed-subglobose, 314-4 by 244-34 cm, indehiscent; spines sparsely set, rigid, flat, with sharp tip, c. % cm. Fruit solitary, of the same shape as the cupule, 3-314 by 2-3 cm, wall 2-3 mm thick, rugose, completely adnate to the cupule. Distr. Malesia: Sumatra, Malay Peninsula, Banka. FAGACEAE (Soepadmo) 299 Ecol. In lowland to submontane forests from sea-level up to 1500 m. In Banka sometimes small trees grow in poor sandy soils derived from gran- ite. Fl. July-Sept., fr. Sept.-June. Notes. The type of C. rhamnifolia (HORSFIELD HB 4268) from Banka, has very young cupules; it is matched by TEISMANN HB 4432, in fruit, from Lampong in S. Sumatra. Leaves 7-12 cm long, + concolorous, scales and hairs very sparse, cupule at least 234 cm through, subsessile with tapering base, densely tomentellous, spines 3-4 mm long, 2-5 mm apart. There is a great resemblance with later collected (non-fruiting) materials from Banka. Kurz 2202 from Burma, which has been refer- red to this species, belongs to C. armata SPACH. 4. Castanopsis borneensis KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 99, t. 90; A. Camus, Chat. (1930) 297, t. 29: 1-3; SoEPADMO, Reinwardtia 7 (1968) 387. Tree, 10-33 m by 15-50 cm g, sometimes with stout buttresses; bark smooth, pale ochre-brown, shallowly flaky. Branchlets glabrous, slender, bark finely fissured, greyish; terminal bud ovoid, 2—3 by 2 mm, scales linear-acute, 2-3 by 14 mm. Stipules linear-acute, 2-314 by 44 mm. Leaves chartaceous, 5-13 by 2-4 cm (index 2.3-—3.6), widest at the middle to slightly below; base rounded and abrupt- ly acute, top + abruptly acuminate (tip 44-2 cm); surfaces glabrous, + discolorous, above greenish and glossy, beneath green-brownish and less glossy; midrib slightly prominent on both surfaces, stronger beneath; nerves 5-8 pairs, thin, flattish on both surfaces, at an angle of 70-80°, subparallel, arcuating and anastomosing near the margin; reticulation irregular, sometimes obscure; petiole 0.8-1144 cm, blackish towards the base, adaxially sulcate. Male rachis 10-15cm by 1-2 mm; bracts broadly ovate-rounded, membranous, 0.7 by 1 mm, bracteoles lanceolate-rounded, membra- nous, 0.7—1 by 0.3-0.4 mm; 3 flowers in clusters of 3; perianth deeply incised, lobes 6, membranous, elliptic-acute, 1-144 by 0.7 mm, hairy outside, stamens 6-8, filaments 244-3 mm, anthers 0.2— 0.25 mm long, pistillode 1-14, mm go. Female rachis 10 cm by 1-2 mm; bracts and bracteoles ovate-acute, 0.7 by 1-114 mm; flowers solitary; perianth deeply incised, lobes 6, membranous, ovate-acute, 0.7—1 by 0.4-0.6 mm, densely hairy outside, staminodes 12, rudimentary, styles 3, conical, 1-144 mm. Ripe cupule sessile, subglobose, 2%-3 by 2-214 cm; wall 1 mm thick, sparsely hairy, mostly very densely set (except the narrow adaxial sector and abaxial suture) with slender to rather sturdy spines 6-12 mm long, in bundles, not branched, straight to slightly recurved, sub- glabrous; indehiscent. Fruit solitary, ovoid to compressed-reniform, 1144-2 by 1-1% cm, adaxi- ally flat, wall 1-2 mm thick, woody, completely adnate to the cupule, rugose and glabrous; cotyle- dons flat-convex. Distr. Malesia: Borneo (Sarawak, Brunei, and Sabah). Ecol. Heath or kerangas forest, also in disturbed primary forest, on black soil and deep yellow sands, 300 in hilly country below 250 m. Fi. March-July, fr. June-Dec. Notes. Records of this species for SE. China, Hainan, and Formosa could not be confirmed. Near C. johorensis and C. wallichii; see the key. 5. Castanopsis johorensis SOEPADMO, Reinwardtia 7 (1968) 393. Tree, c. 10 m by 10-20 cm g; bark smooth, grey- brown. Branchlets initially fulvous-hairy, soon glabrescent, slender, greyish black, with sparse minute lenticels; terminal bud ovoid-ellipsoid, 3-4 by 2 mm, scales ovate or elliptic, acute, 2-3 by 1 mm. Stipules linear-acute, 3-4 by 0.7—1 mm. Leaves 7-15 by 2-5 cm (index 2.5—3.1), widest about the middle; base attenuate, top gradually to abruptly acuminate with a tip of 1-2 cm; surfaces both glabrous and then concolorous, rather glossy, to underneath sparsely set with adpressed stellate scales (magnification 40!) and then pale dull brown- ish; midrib and nerves strongly prominent be- neath, sunken above, nerves 7-11 pairs at an angle of 60-80°, ascending, subparallel, arcuating and anastomosing near the margin; petiole 1—2 cm, dark-coloured towards the base, adaxially sulcate. Female rachis c. 5Y% cm, 1% mm 9g; bracts broadly ovate-acute, 1 by 1 mm; @ flowers solitary, perianth deeply incised, lobes 6, ovate-acute, den- sely hairy on both sides, staminodes 12, rather well developed and + exceeding the perianth, styles 3, cylindrical, recurved, 1-144 mm long. Cupule_ sessile, compressed-globose, 2-3 cm through; wall subglabrous, very densely spiny, less densely in the adaxial sector, the spines in bundles, 8-10 mm, unbranched, slender, -+ straight, subglabrous. Fruit solitary, ellipsoid-com- planate, 1144-2 by 1-114 cm, wall woody, 1-2 mm thick, completely adnate to the cupule, rugose, glabrous; cotyledons flat-convex. Distr. Malesia: Central Sumatra (east of Paja- kumbuh), Malay Peninsula (Johore). Ecol. Lowland forest, up to 500 m. Fi. Aug. (Sumatra), fr. July (Malay Peninsula). Note. Male inflorescences are still unknown. 6. Castanopsis wallichii KING ex Hook f. FI. Br. Ind. 5 (1888) 624; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 106, t. 101A; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 464; A. Camus, Chat. (1930) 470; CORNER, Ways. Trees (1940) 293; SoEPADMoO, Reinwardtia 7 (1968) 403. Tree, 12—15m by 45-60 cm o. Branchlets initially densely brownish tomentose, late glabrescent, slen- der, bark greyish, finely fissured, with minute lenti- cels; terminal bud obovoid-globose, 2-3 by 144-2 mm, scales ovate-linear, acute, 2—3 by 1 mm. Stipules triangular, 3-4 by 1144 mm. Leaves thin-coriaceous or chartaceous, 3144-9 by 1-344 cm (index 2.2-3.5), widest at the middle to sometimes below; base ++ rounded and acute, top acute to 44-11% cm acu- minate; surfaces discolorous, above greenish, more or less glossy, glabrous, beneath brownish, dull, not scaly but densely covered with short stel- late hairs, their branches sometimes partly ad- pressed; midrib and nerves + prominent beneatk_ FLORA MALESIANA [ser.. I, vote/* midrib sunken above; nerves 6-8 pairs at an angle of 45-60°, ascending, parallel, arcuating and anas- tomosing towards the margin; reticulation scalari- form, fine, rather obscure beneath; petiole 44-1 cm, adaxially sulcate. Male rachis 5-10 cm, 1-2 mm g; 3d flowers in clusters of 3, perianth deeply incised, lobes 6, ovate-acute, 1-1 mm long, stamens 12, filaments 244-3 mm, anthers 4% mm long, pistillode c. 1 mm g. Female rachis c. 4 cm, 1 mm 9, bracts and bracteoles ovate-acute, 0.7—1 mm; & flowers solitary, perianth lobes 6, rounded, 0.4-0.5 mm long, densely hairy, staminodes 12, rudimentary, styles 3, conical, recurved, 0.7—1 mm. Cupule subsessile, subglobose to obovoid with flattish adaxial side, 3-4 by 214-3144 cm; wall 1-3 mm thick, outside with dark brown pubes- cence, spines rather densely set except in the ada- xial sector, solitary and unbranched, sturdy, + straight, 5-6 mm, densely fulvous puberulous with glabrous tip. Fruit solitary, similarly shaped as the cupule, 1144-3 by 1-2 cm, wall 1-2 mm thick, almost completely adnate to the cupule, glabrous, rugose. Distr. Malesia: Malay Peninsula (Penang, Kedah, Perak, Maiacca, Singapore), few collec- tions. Ecol. Primary forest at 300-400 m. Fi. Dec. — May, /r. June—Oct. 7. Castanopsis oligoneura SCEPADMO, Reinwardtia 7 (1968) 397. Tree, 10-25 m by 20-25 cm g; bark smooth, lenticellate, or scaly, greenish brown. Branchlets initially with some reddish brown tomentum, glabrescent, slender, sparsely lenticellate; terminal bud ovoid-globose, 2—3 by 2 mm, scales ovate- linear, acute. (Stipules not seen.) Leaves stiff- coriaceous, 7-15 by 3-6 cm (index 2.4-3.4), widest at the middle; base attenuate, acute, top (sub)acuminate; surfaces discolorous, above more or less glossy greenish, glabrous, beneath dull pale brownish, more or less densely covered with adpressed stellate scales (magnification 30-60!) and virtually no hairs; midrib and nerves prom- inent beneath, thinly prominent or flat above; nerves 5—7 pairs at an angle of 45—60°, ascending, subparallel, arcuating near the margin; reticulation fine, scalariform, obscure above; petiole 1144-3 cm, slender, adaxially flat, base thick. Male rachis 5-10 cm, 1144-2 mm g, subglabrous, bracts ovate- acute, 1-144 by 0.7 mm, thick-coriaceous, sub- glabrous, bracteoles ovate-rounded, 0.7 by 0.4 mm, membranous, with dense tomentum outside; 3 flowers solitary or more commonly in clusters of 3-7, perianth deeply incised, lobes 4—6, ovate- acute, 1 by 1 mm, densely tomentose, stamens 8-12, filaments 3-344 mm, anthers 0.2-0.25 mm long, pistillode 1 mm @. Female rachis (sometimes androgynous) 5-8 cm; & flowers solitary, perianth 14-1 mm incised, lobes 6, ovate-acute, densely hairy outside, staminodes 12, strongly reduced, styles 3, cylindrical-conical, recurved, 1-2 mm; ovary rounded-triangular. Cupule subsessile, ellip- soid-globose, 2144-3 by 2-2% cm, adaxially slight- ly flattened; wall c. 2mm thick, with some fulvous 1972] puberulence but largely glabrous, except for the adaxial sector rather laxly but evenly covered with sturdy spines 6-10mm, mostly solitary but sometimes in bundles, somewhat recurved, spar- sely puberulous; dehiscence into 3 (?) parts or irregular. Fruit solitary, ellipsoid-globose, 2-24 by 2 cm, wall 1 mm thick, hard, almost completely adnate to the cupule; when ripe rugose and glab- rous; cotyledons flat-convex. Distr. Malesia: Borneo (Sabah), various collec- tions. Ecol. Lowland forest to 300 m. Fi. April-May, fr. June-Sept. 8. Castanopsis endertii HATus. ex SOEPADMO, Reinwardtia 7 (1968) 390. Tree, 15—20 m by 25-60 cm g. Branchlets initial- ly with a dense stellate tomentum, soon glabres- cent, inconspicuously lenticellate; terminal bud ovoid-ellipsoid, 3-7 by 2-3 mm, scales ovate- acute or linear-acute. Stipules linear, acute, c. 5 by 1 mm, soon caducous. Leaves thick-coria- ceous, elliptic-oblong, (8—)10-15(-19) by (31%) 5—6(—7) cm, entire; surfaces concolorous, glabrous; base rounded, apex abruptly acute or 44-1 cm acuminate; midrib and nerves strongly promi- nent beneath, slightly so or flattish above; nerves 7-13 pairs, subparallel, ascending, at an angle of 60-70°, arcuating but not anastomosing towards the margin; reticulation fine, dense, subscalariform, distinct on both surfaces; petiole 2-3 cm, 1-2 mm @, glabrous, adaxially flat. Male rachis 10-12 cm, 1-2 mm 9, solitary or in paniculate clusters; bracts and bracteoles ovate-acute; gj flowers in clusters of 3; perianth deeply incised, lobes 6, acute, 1-114 mm, densely stellate tomentose out- side, stamens 12, filaments 2-244 mm, anthers c. ¥, mm long, pistillode subglobose, c. 2 mm g. Young infructescence: rachis 15-20 cm, 2-3 mm @, carrying 1-2 cupules. Young cupule ovoid- ellipsoid, sessile, 144-2 cm through, adaxially flat and smooth; spines densely but irregularly set, 4-1 cm, glabrous, flat, tips sharp. Fruit solitary. Distr. Malesia: Borneo (two collections from W. Kutei in Kalimantan). Ecol. Forests up to 450 m. Fi. Sept., fr. Nov. Note. The female flowering rachis is not yet collected. 9. Castanopsis densinervia SOEPADMO, Reinwardtia 7 (1968) 389. Tree, 12-36 m by 35-50 cm go. Branchlets ini- tially with dense reddish brown adpressed stellate tomentum and woolly stellate hairs, glabrescent, sturdy with short internodes, greyish black, with sparse but large lenticels; terminal bud ovate, angular, acute, red-brownish, 3-10 by 3-4 mm, scales ovate, rounded to acute, 3-9 by 2-3 mm. Stipules (not known) leaving distinct scars. Leaves thick-coriaceous, 6-17 by 3-8 cm (index 1.9-3), widest mostly below the middle to rarely above; base rounded to subcordate to sometimes acute, occasionally asymmetrical, top rounded and sub- acuminate to + abruptly 14-1 cm acuminate, tip sharp or blunt; surfaces discolorous, above green- FAGACEAE (Soepadmo) 301 ish, glabrous, glossy, underneath brownish, dull, with a dense cover of more or less adpressed, stel- late scales (waxy when older), often interspersed with shorter or longer stellate hairs, on the nerves only the latter; midrib broad and strong, dark coloured like the petiole; nerves 12-18 pairs at an angle of 45-60°, distinct underneath, parallel, ascending, arcuating and disappearing towards the margin; reticulation fine, obscure, scalariform, rather dense; petiole 44-1144 cm, to3 mm g, adax- ially deeply sulcate. Male rachis 3-7 cm, 3-4 mm 9, bracts broadly ovate-acute, thick-coriaceous, 1-114 mm, bracteoles lanceolate-acute, membra- nous, 144 by 0.7 mm; flowers in clusters of 3, peri- anth deeply incised, lobes 6, ovate-acute, 114-2 mm deep, densely hairy, stamens 12, filaments 2144-3 mm, anthers 0.2-0.25 mm long, pistillode 2-3 mm g. Female rachis 2-3 cm, 3-4 mm a, bracts broadly ovate-acute, thick-coriaceous, 11% by 1144 mm; @ flowers solitary, perianth lobes 6, rounded or acute, 1 mm deep, densely hairy, sta- minodes 12, well-developed, exceeding the peri- anth, styles 3, conical, recurved, 2-244 mm long. Young infructescence: rachis 5-10 cm, 4-5 mm 9g; carrying 5-10 young ovoid-ellipsoid, subsessile cupules, adaxially flat and scaly, laterally with short reddish brown spines. Cupule (almost ripe) on stalk a few mm long and thick, excentrically compressed, obovoid, c. 3 by 4cm, black in colour; wall 1-2 mm thick, glossy, glabrous, spiny except in the adaxial sector, spines in bundles arranged in 3-4 curving bands, unbranched, rather slender, + straight or sometimes reflexed, 8-10 mm, glab- rous. Fruit solitary, reniform or ovoid, base rounded, top acute, 2-244 cm; wall woody, 2-3 mm thick, adnate to the cupule except the top, rugose, glabrous; cotyledons flat-convex. Distr. Malesia: Borneo (Sabah: Mt Kina- balu; Mt Lamaku in the Crocker Range; NE. Kalimantan: Mt Kemul in W. Kutei). Ecol. Primary forest, 1000-1800 m. FI. fr. Sept. —April. 10. Castanopsis paucispina SOEPADMO, Reinwardtia 7 (1968) 398. Tree, 12-30 m by 20-60 cm g; bark dark-grey, smooth or cracked. Branchlets initially with dense, brownish, short-stellate tomentum; glabrescent, slender, densely lenticellate; terminal bud ovoid- globose, 2-3 mm by 2-3 mm, scales ovate-acute. Stipules ovate-acute, 2 by 1 mm. Leaves 8-14 by 3-514 cm (index 2.4-3.2), widest below the middle to sometimes about the middle; base rounded acute, top attenuate and acute to 1 cm acuminate; surfaces discolorous, above more or less glossy, greenish, glabrous, beneath dull cinnamon-brown, with thick cover of adpressed, stellate scales (mag- nification 20!) and on the veins some (often acro- scopically bent) hairs often branched near their base; midrib and nerves prominent beneath, flat- tish above; nerves 9-10 pairs at an angle of 45-60°, ascending, subparallel, arcuating towards the margin; reticulation fine, dense, scalariform, some- times obscure above; petiole slender, 1-144 cm, adaxially shallowly furrowed. Male rachis 5-10 302 cm, 1-144 mm 9g; bracts and bracteoles ovate- acute; ¢ flowers in clusters of 3, perianth deeply in- cised, lobes 6, rounded-acute, 1 by 44-1 mm, with dense tomentum outside, stamens 12, filaments 214-3 mm, anthers 4% mm long, pistillode c. 1 mm g. Female rachis 5-10 cm, in the axil of an ovate-acute bract on the upper part of a new shoot; 9 flowers solitary, perianth deeply incised, lobes 6, rounded-acute, 1 by 1 mm, with dense tomentum outside, staminodes 12, rudimentary, ovary 3-locular, styles 3, conical, 1-144 mm. Ripe cupule chocolate-brownish, subsessile, obovoid- globose but adaxially flattened, 4-414 by 3-3\% cm, base attenuating into the short peduncle; wall 2—4 mm thick, with some sparse fulvous pu- berulence and very sparsely set with 3—4 curved rows of mostly solitary spines, sturdy, often somewhat flat and recurved, 5—7 mm long, subglab- rous, the adaxial sector not spiny. Fruit solitary, obovoid-globose, c. 4 by 3 cm, wall woody, 5-7 mm thick, completely adnate to the cupule, when ripe rugose and glabrous. Distr. Malesia: Borneo (Central Sarawak: Anap; Sabah: Keningau Distr.). Ecol. Mixed Dipterocarp forest on basalt de- rived soil, 700-1100 m. F/. Aug. — Oct., fr. Sept. 11. Castanopsis clemensii SOEPADMO, Reinwardtia 7 (1968) 388. Tree, 10-30 m by 15-30 cm g; buttresses up to 1 m tall; bark smooth, lenticellate, greyish brown. Branchlets initially with dense, yellow- ish brown stellate tomentum; glabrescent; in- ternodes sometimes short, slender, dark purplish brown-grey, with minute lenticels; terminal bud ovoid-globose or ellipsoid, 3-5 by 2—3 cm, scales ovate, elliptic, acute, 3-4 by 2 mm. Stipules ovate, triangular, blunt, 5-10 by 3-5 mm, rather long persistent. Leaves thin-coriaceous, 6-12 by 2-6 cm (index 1.8—2.6), widest mostly below to rarely abo- ve the middle; base rounded to subcordate, usually asymmetrical, top rounded and subacuminate to 1 cm acuminate with blunt to acute tip; surfaces green-brownish, above glabrous, rather glossy, beneath dull, nerves rather densely set with bunch- like stellate hairs, the branches of the latter straight to somewhat curly, scales non; midrib and nerves weakly prominent on both surfaces; nerves 8-10 pairs at an angle of 50-60°, subparallel, ascending, arcuating and disappearing near the margin; re- ticulation fine, distinct on both surfaces, more or less irregular; petiole 44-1 cm, terete or adaxially flat. Female rachis 3-7 cm, 1 mm @, bracts and bracteoles ovate-acute, 0.6-1 by 1-144 mm; 2 flowers solitary, perianth deeply incised, lobes 6, Ovate-acute, 0.6 mm long, densely hairy on both sides, staminodes 12, rudimentary, styles 3, co- nical, 1.2-1.5 mm, recurved. Cupule (not yet ripe) c. 5 mm stalked, obovoid-ellipsoid, 2-24% by 1% cm, tapering towards the base, dark-coloured; wall c. | mm thick, sparsely fulvous-puberulent, spiny except the adaxial sector and abaxial sutures; spines laxly set in bundles arranged in more or Jess parallel curving lines, 5-8 mm long, sparsely branched near the base or simple, rather slender, FLORA MALESIANA [ser. I, vol. 7? mostly straight, sometimes reflexed, subglabrous. Unripe fruit solitary, ellipsoid, 1144 by 1 cm, wall 1 mm thick, completely adnate to the cupule, rugose and glabrous; cotyledons flat-convex. Distr. Malesia: Borneo (Sabah: Mts Kinabalu and Trusmadi). Ecol. Primary forest, 750-1800 m, on black rocky soil. Fi. Sept., fr. Nov. — May. Notes. Close to C. oviformis; see the note under that species. The male inflorescence and the mature cupule are not yet known. 12. Castanopsis oviformis SOEPADMO, Reinwardtia 7 (1968) 397. Tree, 15-30 m by 20-45 cm 9; buttresses small, sometimes branched; bark smooth or shallowly fissured, greyish brown, lenticellate. Branchlets initially with brownish, adpressed stellate indu- mentum, late glabrescent; rather slender, grey- brownish or dark-purplish, sparsely set with mi- nute lenticels; terminal bud ovoid-globose, 2-3 by 2 mm, scales ovate, linear, acute. Stipules Ovate-acute, 3-5 by 2-3 mm. Leaves sometimes tending to distichy, more or less thinly coriaceous, 6-15 by 4-614 cm (index 2.2-3), widest somewhat below to somewhat above the middle; base + rounded and acute, top mostly rounded and ab- ruptly acuminate with a tip of 44-1 cm; surfaces green-brownish, above very glossy, glabrous, beneath sometimes dull, very sparsely to very densely set with adpressed stellate scales (mag- nification 30—60!), sometimes with a few stellate hairs on the nerves; midrib prominent beneath, flattish or slightly elevated above; nerves 8-13 pairs at an angle of 60—80°, ascending, parallel, arcuating towards the margin; reticulation dense, fine, scalariform, distinct underneath; petiole 1-144 cm, adaxially flat. Male rachis 5-15 cm, 1-2 mm 9, bracts ovate-acute, 1-144 by 1 mm; 3 flowers in clusters of 3-7, perianth deeply in- cised, lobes 6, 1-14% mm long, acute, with dense tomentum outside, stamens usually 12, filaments 1-114 mm, anthers 4, mm long. Female rachis 5-10 cm, androgynous inflorescence with § flowers in its apical part; bracts 144-2 by 1-14% mm, with dense tomentum; @ flowers solitary, perianth deeply incised, lobes 6, ovate-acute, 1 by 1 mm, with dense tomentum outside, staminodes 12, rudimentary, styles 3, cylindrical-conical, 2 mm. Cupule 34-1 cm stalked, ellipsoid to obovoid, with tapering base, 244-5 by 14%-3 cm, wall 14-1 mm thick, sparsely puberulous, spines ar- ranged in arching lines on both sides of the cupule except on the adaxial sector, solitary or in bundles, sparsely set and 2-3 mm long to rather densely set and 8-10 mm long, unbranched, sturdy, straight to somewhat recurved, densely fulvous- puberulous; dehiscence irregular or in two equal halves. Fruit solitary, 2-34% by 144-2 cm, obo- void-cylindrical, wall 44-1 mm thick, completely adnate to the cupule, rugose, glabrous; cotyle- dons flat-convex. Distr. Malesia: Borneo (scattered all over the island). 1972] Ecol. Primary forest, sometimes kerangas for- est, at low altitude (also montane? see note), on sandy loamy soil. Fl. June-Aug., fr. Sept.-July. Notes. In ENperT 3610 from Mt Kemul in Central East Borneo at 1200 m, the leaves are elliptic, to 11 by 6% cm, underneath densely set with both scales and stellate hairs. In S 20139 from Ulu Baram in NE. Sarawak at 1080 m the leaves are elliptic, to 9144 by 6 cm, with some scales and few hairs underneath. These two specimens, both in flower, are suggestive of C. clemensii, which is known from somewhat higher elevations. Most material of C. clemensii is not in fruit, the only cupules available being from the type, SAN 25315, from Sg. Mentaki in the Ranau District at 750 m. Here the leaves have underneath a few stellate, somewhat curled hairs and, unlike the rest of the clemensii material, have virtually no scales. The cupule, however, differs distinctly from that of C. oviformis in the smaller size, blackish colour, subglabrous surface, and branched spines. The leaves in KorTHALS s.n. from Martapura, are up to 141% by 5 cm and underneath thickly covered with scales and also have hairs on the nerves; those of S 18532 seem to be intermediate between these and the rest of the material. 13. Castanopsis malaccensis GAMBLE, Kew Bull. (1914) 178; J. As. Soc. Beng. 75, ii (1915) 445; A. Camus, Chat. (1930) 319, t. 32: 11-13; Corner, Ways. Trees (1940) 293; Soppapmo, Reinwardtia 7 (1968) 395. Tree, c. 15 m by c. 30 cm g; bark yellowish brown, fissured. Branchlets greyish black, with minute lenticels, initially densely set with fulvous stellate tomentum; terminal bud ovoid, 2-3 by 1-2 mm; scales ovate-acute, with dense stellate tomentum. Leaves thin-coriaceous, 314-10 by 24%4~ 5 cm (index 1.8-2), widest below or at the middle; surfaces strongly discolorous, above glabrous, glossy, greyish green, underneath with a dense cover of rufous, adpressed, stellate hairs; base rounded and abruptly acute, apex acute or abrupt- ly 4% cm acuminate; midrib and nerves prominent beneath, impressed above; nerves 7-9 pairs, paral- lel, ascending, at an angle of 50—-70°, arcuating but not anastomosing towards the margin; reticulation fine, dense, subscalariform, obscure on both sur- faces; petiole 1-3 cm, 1 mm 9, adaxially flat. Male rachises 4-7 cm, 1-2 mm 9g, in paniculate clusters, densely rufous stellate-tomentose; bracts and bracteoles ovate-acute; 3 flowers in clusters of 3, perianth deeply incised, lobes 6, ovate, 1-114 mm, densely stellate-tomentose outside, stamens 12, filaments 3-4 mm, anthers 0.2-0.3 mm long, pis- tillode subglobose, 144-2 mm g. Cupule ovoid- globose, 344-414 cm; wall 1-2 mm thick; pedun- cle 14-1 cm; spines much-branched, needle-shaped, 1-2 cm, densely set; dehiscence into two equal halves or irregular. Fruit solitary, ovoid-conical, adaxially flat, 14-21% cm; wall 44-1 mm thick, glabrous, rugose, adnate to the cupule except for the top which is set with a thin cover of silvery tomentum; cotyledons flat-convex. FAGACEAE (Soepadmo) 303 Distr. Peninsular Siam, Malesia: Sumatra (Ri- ouw: Kuantan Distr.), Malay Peninsula (Negri Sembilan, Selangor, Malacca, Johore, Singapore). Ecol. Forest up to 300 m. FI. fr. Jan.-May. Note. The female rachis and flowers are not yet known. 14. Castanopsis tungurrut (BL.) A.DC. J. Bot. 1 (1863) 182; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 97, t. 87; A. CAMus, Chat. (1930) 358, t. 44: 6-10; BACKER & Baku. f. Fl. Java 2 (1965) 4; SOEPADMO, Reinwardtia 7 (1968) 403. — Castanea tungurrut BL. Bijdr. (1826) 525; Fl. Jav. Cupul. (1829) 42, t. 22; K. & V. Bijdr. 10 (1904) 7; Koorpb. Atlas 1 (1913) t. 39. — Castanea tungurrut f. suma- trana Mia. Sumatra (1861) 353. — C. ridleyi GAMBLE, Kew Bull. (1914) 180, p.p. lectotypi GOODENOUGH 1479. — C. conspersispina MERR. Pap. Mich. Ac. Sc. 19 (1934) 152, t. 17. Tree, 20-25 m by 60 cm g; bark rough, fissured. Branchlets initially with fulvous tomentum, later glabrescent, rather sturdy, dull greyish black, smooth, sparsely lenticellate; terminal bud ovoid- ellipsoid, 3-5 by 2-3 mm, scales ovate, elliptic, acute, 3-4 by 2 mm. Stipules boat-shaped, ovate- triangular, 4-6 by 2-3 mm. Leaves (8—)12—15(—23) by (314-)5-6(-9) cm (index 2-3), widest about the middle to slightly below; base rounded-acute, sometimes asymmetrical, top tapering and subacu- minate; more or Jess discolorous, above rather glossy, green-brownish, glabrous, beneath dull with a faint greyish tinge, densely covered with a waxy mass in which stellate scales are very closely adpressed (magnification 60!), and also with some longer hairs (the latter in Sumatra almost wanting); midrib prominent on both surfaces, stronger be- neath; nerves 11-19 pairs at an angle of 60-70°, parallel, arcuating and disappearing near the mar- gin, prominent beneath, flattish above; reticula- tion fine, dense, scalariform, sometimes obscure above; petiole (144-)1-114(-2) cm, adaxially flat or shallowly sulcate. Inflorescences male, female, or androgynous. Male rachis 10-25 cm by 1-2 mm; bracts and bracteoles ovate-acute, 2-3 by 1-1% mm, hairy outside; ¢ flowers in clusters of 3, per- ianth deeply incised, lobes 5-6, acute, 2-2% by 1-14 mm, hairy on both sides, stamens 10-12, filaments 3-4 mm, anthers 0.2-0.25 mm long, pistillode 144 mm g. Female or androgynous rachis 5-15 cm, bracts thick-coriaceous, ovate-acute, 2— 21%, mm, densely hairy outside; 2 flowers solitary, perianth lobes 5-6, acute, 0.7—1 mm, densely hairy, staminodes 12, rudimentary, styles 3, conical, recurved, 2-244 mm. Young cupule sessile, ovoid, on the rounded lateral sides with spine-like tuber- cles, on the flat median sides scaly. Ripe cupule subsessile, obovoid-ellipsoid, 5-6 by 3-4 cm; wall 1—2 mm thick, covered partly with fulvous puber- ulence, and mostly all over with scattered bundles of slender spines branched in the basal half and easily reflexed 10-23 mm long, sparsely puberulous; dehiscence none or irregular. Fruit solitary, ovoid- ellipsoid, 3-4 by 114-2 cm, wall 1 mm thick, com- pletely adnate to the cupule, when ripe rugose and glabrous; cotyledons flat-convex. 304 FLORA MALESIANA [ser. I, vol. 7? NUR SSA SEES Fig. 12. Castanopsis hypophoenicea (SEEMEN) SOEPADMO. a. Habit, x 24, b. underside of leaf, nat. size, c. ripe cupule, x 24, d. longitudinal section of cupule and fruit, x 73 (a-c SAN 47196, dSAN 40316). 1972] FAGACEAE (Soepadmo) 305 a ——————__——____ Distr. Malesia: Sumatra (scattered from Tapa- nuli in the north to Lampong in the south), Malay Peninsula, also in Simalur and Banka Is. and West Java (rather common). Ecol. Forests, from swampy lowlands up to 1500 m. Fi. fr. + throughout the year. Vern. Kalimorot, tungeureut, S, with variants. Notes. In the type of C. conspersispina, the cupule is but sparsely spiny. In the past many specimens belonging to this species have been erroneously identified. 15. Castanopsis evansii ELMER, Leaf]. Philip. Bot. 5 (1913) 1778; SozPADMO, Reinwardtia 7 (1968) 391. — C. javanica (non A.DC.) MeErR. En. Philip. 2 (1923) 24. — C. woodii MeErRR. Philip. J. Sc. 29 (1926) 362. — C. elmeri Merr. Pl. Elm. Born. (1929) 42; A. Camus, Chat. (1930) 442, t. 76: 6-8. Tree, 5-24 m by 15-50 cm g; bark grey, smooth. Branchlets initially with some tomentum, soon glabrescent, slender, smooth or finely fissured, rather glossy dark purplish brown, sparsely lenti- cellate; terminal bud ovoid-globose 3-5 by 3 mm, scales ovate-acute, 3 by 2 mm. Stipules triangular, 5-6 by 2 mm, rarely persistent. Leaves 10-17 by 34%4-5'% cm (index 2.3-3.4), widest about the middle; base tapering to rarely rounded and abruptly acute, apex bluntly acute to %-1 cm acuminate with sharp tip; surfaces green to brown- ish, above more or less glossy, glabrous, beneath duller and mostly greyish with wax in which ad- pressed stellate scales are mostly discernible (mag- nification 60!) with very sparse longer hairs mostly only on the nerves; midrib strongly prominent beneath, slightly so above; nerves 11-14 pairs at an angle of 60-70°, parallel, ascending, arcuating towards the margin, slightly prominent beneath, flattish above; reticulation fine, dense, scalariform; petiole 1-214 cm, adaxially flat. Male rachis 10-15 cm, 1-2 mm g, bracts ovate-acute, 144 by 1 mm, bracteoles 1 by 0.7 mm; ¢ flowers in clusters of 3; perianth lobes 4-6, connate at the base only, 1-144 by 0.7-1 mm, ovate-acute, densely hairy, stamens 10-12, filaments 114-2144 mm, anthers 0.2-0.25 mm, pistillode 14% mm o. Female rachis 5-10 cm, bracts ovate-acute, 1 by 1 mm; @ flowers solitary, perianth deeply 6-lobed, the lobes 1 by 0.7 mm, densely hairy; staminodes 12, rudimentary, styles 3, recurved, conical, 1-2 mm long. Young infructescence c. 10 cm long, carrying numerous solitary cupules. Young cupule ovoid, bearing short, simple spines on the rounded lateral sides, and scales on the median sides. Ripe cupule Y¥ cm stalked, obovoid-ellipsoid, 4-414%4 by 3-344 cm; wall 2-4 mm thick, inside brown tomentose, out- side with some brownish puberulence, otherwise glabrous, spines sparsely to densely set sparsely puberulous, sturdy and blackish to slender and rather soft and brownish, in bundles, often bran- ched in the basal part with the branches reflexed, 10—23 mm long in all; dehiscence none or irregular. Fruit solitary, ovoid, adaxially flat or depressed, 3-314 by 2-3 cm, wall 2-3 mm thick, woody, ad- nate to the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Palawan), Borneo (Sabah: rather common; SE. Kalimantan: region of Balikpapan, Riko; also Banguey I.). Ecol. Lowland forests to 500 m altitude, on brown soil and clayey loam. Fi. Jan.-May, /r. May-Dec. 16. Castanopsis hypophoenicea (VON SEEMEN) SOE- PADMO, Reinwardtia 7 (1968) 392. — Quercus hypophoenicea VON SEEMEN, Bot. Jahrb. 23, Beibl. 57 (1897) 52. — Lithocarpus hypophoenicea (VON SEEMEN) BARNETT, Trans. & Proc. Bot. Soc. Edinb. 34 (1944) 171; A. Camus, Chénes 3 (1954) 890. — C. dispersispina MERR. Univ. Calif. Publ. Bot. 15 (1929) 41; A. Camus, Chat. (1930) 442. — Fig. 12. Tree, 12-30 m by 10-60 cm g; buttresses up to 1.8 m tall, 1 m out; bark surface grey-brown, smooth or finely fissured. Branchlets initially den- sely set with yellowish brown stellate and simple tomentum, later subglabrous, smooth or finely fissured, or sparsely lenticellate; terminal bud ovoid, 3—5 mm, scales ovate. Stipules ovate-acute, 3-5 mm. Leaves thin-coriaceous, elliptic-oblong, (6—)10-15(-19) by (2144-)4~-5(-7) cm, entire, dis- colorous, above glabrous except the midrib and veins, dull or glossy, greyish green or chocolate brown, underneath with a dense cover of yellowish brown, adpressed stellate hairs and simple long hairs; base rounded and abruptly acute or cuneate, sometimes asymmetrical, apex bluntly acute or 1-114 cm acuminate; midrib and nerves prominent beneath, flattish or impressed above; nerves (9—) 11-13(-15) pairs, parallel, ascending, at an angle of 60-70°, arcuating but not anastomosing near the margin; reticulation dense, fine, subscalariform, distinct beneath; petiole 5-15 by 2 mm, densely stellate-tomentose, glabrescent, thickened and ru- gose at base, adaxially flat or shallowly furrowed. Male rachises 5-10 cm, in lax paniculate clusters; bracts and bracteoles ovate-lanceolate, 0.7-1 by 0.7 mm; ¢ flowers in clusters of 3; perianth deeply incised, lobes 6, ovate, 114-2 by 1-114 mm, sta- mens 10-12, filaments 3-4 mm, anthers 0.25—0.30 mm long, pistillode subglobose, 144-2 mm go. Ripe cupule ovoid-ellipsoid, sessile, 7-10 by 5-7 cm; wall woody, 2-3 mm thick; spines much- branched, needle-shaped, 1-2 cm long, regularly set in more or less 6-8 curving lines with space in between the rows; dehiscence apparently irregular. Fruit solitary, ovoid-ellipsoid, 4-6 by 3—5 cm, wall woody, rugose, 3-7 mm thick, completely adnate to the cupule; adaxially flat. Distr. Malesia: Borneo (Sarawak, Kalimantan, Sabah). Ecol. Forests up to 650 m. Fi. Sept.-May, /r. June-Dec. Note. The female rachis and flowers are not yet known. 17. Castanopsis megacarpa GAMBLE, Kew Bull. (1914) 180; J. As. Soc. Beng. 75, ii (1915) 462; A. Camus, Chat. (1930) 440, t. 61: 1-3; CORNER, Ways. Trees (1940) 293, f. 94; SoEPADMO, Rein- wardtia 7 (1968) 395. Tree, 12-36 m by 30-90 cm g; buttresses up to 2 m tall; bark surface smooth or shallowly fissured, 306 FLORA MALESIANA [ser. I, vol. 72 pale greyish brown. Branchlets initially densely set with reddish brown adpressed stellate tomen- tum, later greyish brown, sparsely lenticellate; terminal bud ovoid-globose, 3-5 by 3-4 mm, scales ovate-acute. Leaves thick-coriaceous, lan- ceolate-oblong or elliptic-oblong, 10-25 by 3-10 cm; surfaces strongly discolorous, above glabrous, glossy, greyish green, underneath with a thick cover of reddish brown adpressed stellate hairs; base rounded or acute, sometimes asymmetrical, margin entire recurved, apex rounded or abruptly acute; midrib and nerves prominent beneath, flat- tish above; nerves 8-16 pairs, parallel, ascending, at an angle of 60—70°, arcuating but not anastomos- ing near the margin; reticulation fine, subscalari- form, obscure on both surfaces; petiole subglab- rous, thickened and rugose at the base, adaxially flat or shallowly furrowed, 1144-214 cm by 2-3 mm. Male rachises 10-20 cm, in paniculate clusters; bracts ovate-acute; g flowers in clusters of 3, peri- anth deeply incised, lobes 6, acute, 1-2 mm, stamens 12, filaments 344-4 mm, anthers 0.20- 0.25 mm long, pistillode rounded, c. 1 mm go. Fe- male rachis 10-15 cm; flowers solitary, ovary roun- ded triangular in cross-section, 3-locular, perianth 6-lobed, staminodes 12, rudimentary, styles 3, terete, c. 1 mm, recurved. Ripe cupule ovoid-glo- bose, subsessile, 8-10 by 6-7 cm; wall woody, 2-4 mm thick; spines much-branched, 1-114 cm long, with sharp tip, rigid, densely but irregularly set; dehiscence into two equal halves or irregular. Fruit solitary, ovoid, 6-8 by 3-5 cm, wall woody, 3-4 mm thick, rugose, adnate to the cupule; coty- ledons flat-convex. Distr. Malesia: Malay Peninsula (also Singa- pore), Borneo. Ecol. Forests, up to 1350 m. Fl. March-Aug., fr. Sept.-June. Note. The syntype of C. ridleyi GAMBLE, K1NG’s Coll. 6831, 1 refer to C. megacarpa. 18. Castanopsis javanica (BL.) A.DC. J. Bot. 1 (1863) 182; Prod. 16, 2 (1864) 111, incl. var. mon- tana (BL.) A.DC.; Hook. f. Fl. Br. Ind. 5 (1888) 620; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 97, t. 88; GAMBLE, Kew Bull. (1914) 180; A. Camus, Chat. (1930) 316, t. 32: 7-10; BACKER & BAKuH. f. Fl. Java 2 (1965) 4; So—EPADMO, Reinwardtia 7 (1968) 393. — Fagus javanica BL. Flora 7 (1824) 295. — Castanea javanica BL. Bijdr. (1826) 525, incl. unnamed variety /.c. 526; Fl. Jav. Cupul. (1829) 44, t. 23, incl. var. montana l.c. 45, t. 24, and var. fuscescens l.c. 45; K. & V. Bijdr. 10 (1904) 9; Koorp. Atlas 1 (1913) t. 38. — Castanea mon- tana BL. Bijdr. (1826) 526; Hassk. Cat. Hort. Bogor. Alt. (1844) 73. — Quercus discocarpa HANCE, J. Bot. 12 (1874) 242; Hook. f. Fl. Br. Ind. 5 (1888) 616; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 76, t. 70; CoRNER, Ways. Trees (1940) 302, f. 97. — C. discocarpa (HANCE) Hance, J. Bot. 16 (1878) 201; A. Camus, Chat. (1930) 356, t. 43: 4-9. — Quercus javanica DRAKE in Morot, J. de Bot. 4 (1890) 153. — Pasania discocarpa (HANCE) GamBIE, J. As. Soc. Beng. 75, ii (1915) 447. — Synaedrys discocarpa (HANCE) Koibz. Bot. Mag. Tokyo 30 (1916) 186. — C. lentiginosa E. F. WARB. Kew Bull. (1936) 20. — C. penangensis A. CAMUus, Bull. Soc. Bot. Fr. 94 (1947) 4. Tree, 10-40 m, trunk sometimes fluted, 20-100 cm g. Branchlets initially rather densely set with reddish brown, adpressed, stellate hairs and fim- briate scales, mixed with patent hirsute bundle- hairs, later glabrescent; rather slender with dark grey, sometimes brownish bark full of distinct lenticels, especially when older; terminal bud ovoid-globose, 3-9 by 2-4 mm, scales roundish, 3-5 by 2-3 mm. Stipules 3-4 by 144 mm, recurved, hairy, very soon caducous. Leaves thick-coriaceous, (6-)10-13(-18) by (2—)3-6(-8) cm (index 1.9- 3.6), widest about the middle to slightly lower or higher; base attenuate-acute to rarely rounded- acute, apex bluntly acute to 4-2 cm acuminate; surfaces with a livid tinge, discolorous, above greenish and more or less glossy, glabrous, be- neath light brown and more or less dull, some- times with a distinct cover of wax with scales hardly discernible (magnification 60!), often also with scattered hairs simple or in 2—3-fid bundles, sometimes glabrous; midrib prominent beneath, flat or impressed above; nerves (8—)9-11(—13) pairs, thin, ascending, at an angle of 45-70°, sub- parallel, arcuating towards the margin; reticula- tion scalariform, thin and obscure on both sur- faces; petiole 44-1 cm, adaxially flat. Male rachis 10-15 cm, slender, bracts ovate-acute, 1-1.2 mm; 6 flowers solitary or in clusters of 3, perianth lobes 6, connate at the base only, 1.2—-1.5 by 1 mm, den- sely hairy on both sides; stamens 12, filaments 3-4 mm, anthers 0.2-0.25 mm long, pistillode 1.2-1.5 mm g. Female rachis 5-10 cm, slender, bracts ovate, sometimes irregularly lobed, 1-144 by 2-244 mm; flowers laxly scattered, solitary, perianth 6-lobed, the lobes ovate, 0.8-1 by 0.3- 0.5 mm, staminodes 12, rudimentary, styles 3, conical-cylindrical, recurved, 2-3 mm. Young cupule 3-5 mm stalked, densely covered with irregularly placed spines or with sparse spines in 4-5 + concentric rows. Mature cupule on peduncle 4-5 mm g, globose or discoid-conical, 244-5 by 1.3-4 cm, base rounded or concave; wall 144 mm thick, inside silky with fulvous hairs, outside densely fulvous-velvety, spines 2-21 mm long, tree-like branched (sometimes twice) or sometimes in bundles, straight to very slightly recurved, or round the base the whole spine bundles reflexed, densely hairy to subglabrous, arranged with few sturdy ones in 4-5 + concentrical rows with much of the cupule practically smooth, to densely cov- ering the whole cupule surface with many slen- der ones; dehiscence into 4 equal segments or irregularly. Fruit solitary and hence round on section, depressed-conical, 1-144 by 2-2% cm; apex acute to rounded or depressed, umbonate; wall 2 mm thick, scar 144-1 part, flat or convex, the free part hairy. Distr. Malesia: Sumatra (from the north to the Palembang area; also Banka), Malay Penin- sula (Perak, Selangor, several collections; also Penang), West Java (common), and Borneo (not seen from western Kalimantan). 1972] FAGACEAE (Soepadmo) 307 Ecol. Primary, sometimes secondary forests, up to 1650 (—2000) m. F/. Sept.-March, fr. March- Dec. DOocTERS VAN LEEUWEN (Zoocecidia, 1926, 105) described a stem-gall caused by a psyllid, and (in Ned. Kruidk. Arch. 51, 1941, 134) a stem-gall caused by a Lepidopteron, both from Java. Vern. Kalimorot, ki hijur, S. Uses. Although a tree of poor shape, one of the most durable timbers in the Kelabit Highlands, central Sarawak, and widely used for house posts. Bark used for making kegs for storing rice. Notes. Young growth scurfy brown or red- dish. In all islands there is a continuous variation in the texture and pubescence of the leaf, and also in the density and arrangement of the spines. Spe- cimens below 500 m usually have thinner and less pubescent leaves and fewer and more regular- ly spaced spines, resembling C. discocarpa from Banka. Those from 500-1000 m match in all re- spects C. lentiginosa from Sarawak, whilst spe- cimens from higher altitudes agree with C. java- nica. In Java, the influence of altitude on varia- tion is particularly distinct. Plants from Depok (at 100 m), Janlappa (at 90 m), and Bogor (at 250 m), have thinner and less pubescent leaves and less spiny cupules than the many collections from 1000 m and above. In Sumatra, Malay Pen- insula, and Borneo, the same pattern of variation occurs and no distinctions appear to hold good in the considerable range of diversity. The species is quite different from C. echidno- carpa from India and from our C. acuminatissima with which HANCE compared it; rather it is related to C. ferox from India, with which E. F. WARBURG compared his C. /entiginosa, but it differs in its len- ticellate branchlets, fewer nerves, shorter spines, and a tomentose fruit. The cupule, whatever its outward resemblance with that of C. costata, con- tains 3 fruits in the latter. The leaves of C. java- nica are, on the average, smaller than in C. costata, with thinner veins and more obscure reticulation. Its brachlets, anyway the older ones, are strikingly rich in lenticels, and the scales of the terminal bud are broader. A malformed specimen labelled ‘‘in sylvis mon- tis Dieng 5—6000’ Hb. Waitz’’ (L) belongs here; B.iumeE identified this in sched. as Quercus angusta- ta. Since all other collections are from West Java, a mistake in the labelling is assumed. 19. Castanopsis buruana Mia. Ann. Mus. Bot. Lugd. — Bat. 1 (1863) 120; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 107, t. 101B; A. Camus, Chat. (1930) 475, t. 72: 13-14; SoEPADMo, Rein- wardtia 7 (1968) 387. — C. buruana f. grandifolia Mia. Ann. Mus. Bot. Lugd. — Bat. 1 (1863) 120. — Castanea buruana (MiQ.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 377. Tree, up to 20 m by 10-30 cm g; bark shallowly fissured or scaly, greyish brown. Branchlets ini- tially densely set with brownish erect stellate hairs, later subglabrous, greyish black, terete, densely minute-lenticellate; terminal bud ovoid-globose or compressed ovoid, 3-4 by 1144-3 mm, scale ovate-acute. Phyllotaxis with a tendency to distichy. Leaves thin-coriaceous, elliptic-lanceolate, rarely ovate-elliptic, (7—)10-15(-24) by (21%-)4~-5(-8) cm, discolorous, above glabrous, glossy, under- neath with a dense cover of chocolate-brown fimbriate scales and erect stellate hairs; base round- ed and abruptly acute or cuneate, margin entire, recurved, apex acute or more commonly 14-14% cm, acuminate, with a sharp tip; midrib and nerves thinly prominent beneath, flattish or impressed above; nerves 8-15 pairs, subparallel, ascending at an angle of 45-60°, arcuating but not anasto- mosing towards the margin; reticulation fine, sub- scalariform, obscure on both surfaces; petiole 1-114 cm by 1 mm, thickened and rugose at base, adaxially shallowly furrowed. Male rachises soli- tary or more commonly in paniculate clusters, 10-15 cm; bracts ovate acute, membranous, 1-144 mm; ¢ flowers in clusters of 3; perianth deeply incised, lobes 6, membranous, c. 1 mm, stamens mostly 12, filaments 244-34 mm, an- thers 0.2-0.25 mm long, pistillode subglobose, 1.2-1.5 mm @. Female rachis 10-15 cm; @ flowers solitary, perianth membranous, 6-lobed, stami- nodes 12, well developed and exceeding the peri- anth, styles 3, cylindrical, recurved, 2-244 mm. Infructescence 10-25 cm, rachis terete, densely minute-lenticellate, carrying 10-20 cupules. Cu- pule sessile, asymmetrically compressed ovoid- conical, 1-2 cm through, wall thin, completely enclosing the fruit except for the rather long per- sistent perianth and styles; spines sparsely set in 4-5 curving rows, sharp, 3-5 mm; dehiscence irreg- ular. Fruit ovoid-conical, more or less rounded- triangular in cross-section, 1-114 cm; free part occupying the greater surface, densely yellowish brown simple-tomentose, base rounded, scar flat or convex, rugose, glabrous. Distr. Malesia: Borneo (Sabah, once), Celebes (fairly common), Moluccas (Sula, Buru, Batjan, Morotai, Obi, Ceram). Ecol. In primary or secondary forest on low hills, up to 1000 m. In SW. Ceram it survives in secondary forest under the influence of constant burning (during the long dry period). Fl. Aug.- Jan., fr. Jan.-July. Note. Near to C. acuminatissima, but different by its somewhat larger leaf with entire margin, sulcate petiole, and by its cupule set with spines arranged in 4-5 curving rows. 20. Castanopsis acuminatissima (BLt.) A.DC. J. Bot. 1 (1863) 182; A. Camus, Chat. (1930) 433; HIcKEL & A. Camus, FI. Gén. I.-C. 5 (1930) 1012; SOEPADMO, Reinwardtia 7 (1968) 385. — Castanea acuminatissima BL. Mus. Bot. 1 (1850) 283. — Castanea sessilifolia BL. l.c. 284. — Quercus lineata (non Bu.) Mia. Pl. Jungh. (1851) 10. — Quercus varingaefolia Mia. I.c. 12. — Quercus junghuhnii Mia. FI. Ind. Bat. 1, 1 (1856) 853; OuDEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 205; Mig. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 117 (“‘iunghuhnii’’); Oupem. Natuurk. Verh. Kon. Akad. 11 (1865) 16, t. 9; WENziIG, Jahrb. Bot. 308 FLORA MALESIANA [ser. I, vol. 7? Fig. 13. Castanopsis acuminatissima (BL.) A.DC. Tree surrounded by a skirt of root suckers. G. Telaga, above Puntjak Pass, Mt Gedeh, W. Java, c. 1500 m alt. (VAN STEENIS, May 1939). 1972] FAGACEAE (Soepadmo) 309 a ———————————————————— Gart. Berl. 4 (1886) 237; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 72, t. 73; CRAIB, Kew Bull. (1911) 471; LANE-PooLE, For. Res. New Guinea (1925) 78. — Quercus fagiformis JUNGH. Bonplandia 6 (1858) 83, see Reup. J. Arn. Arb. 1 (1919) 122, note; K. & V. Bijdr. 10 (1904) 54. — Quercus acuminatissima (BL.) A. DC. Prod. 16, 2 (1864) 102, non MerRR. 1908; BACKER & BAKH. f. FI. Java 2 (1965) 6. — Pasania acuminatissima OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 84; GamMBLE, J. As. Soc. Beng. 75, ii (1915) 448. — C. schlenkerae BAILEY, Queensl. Agr. J. 22 (1909) 149; A. Camus, Chat. (1929) 436, t. 60: 18-20. — Synaedrys fagiformis Koibz. Bot. Mag. Tokyo 30 (1916) 187. — C. junghuhnii (M1Q.) MARKGR. Bot. Jahrb. 59 (1924) 62, f. 1: A-D. — C. nebularum Hicket & A. Camus, Bull. Mus. Hist. Nat. Paris 32 (1926) 398. — C. longispicata Hu, Bull. Fan Mem. Inst. Biol. Peiping, Bot. 10 (1940) 86. — C. bejaudii A. Camus, Bull. Mus. Hist. Nat. Paris IT, 13 (1942) 479. — Fig. 13. Tree, 10-36 m by 30-90 cm g; buttresses to 2 m tall, 144 m out; bark greyish brown, rough, fissur- ed, inner bark 1—2 cm thick, pale to reddish brown. Branchlets initially with a dense layer of rufous fimbriate scales and adpressed stellate hairs; slender, later glabrescent, with minute lenticels, greyish black, finely fissured, at the base with scars of the bud scales; terminal bud ovoid or flat-el- lipsoid, mostly found when having attained a size of 4-8 by 2-3 mm, apparently waiting for some time before developing further, the scales imbricate or mostly distichous, membranous, ova- te-acute, 1-144 by 1-1.2 mm, densely hirsutely ciliate, persistent for some time. Stipules ovate- acute, 3-4 by 1 mm, caducous. Leaves thin- coriaceous, 444-17 by 2144-6 cm (index 2.4-3.5 (-4)), widest at or below the middle; base rounded or attenuate, acute to sometimes decurrent, mar- gin entire and undulate or remotely serrate in the apical half, top tapering and acuminate with sharp tip 1-214 cm; surfaces discolorous, above glossy green, glabrous, underneath dull brown to sometimes silvery with a mostly thick cover of adpressed scales, no hairs; midrib and nerves prominent beneath, flattish or slightly sunken above; nerves 10-14 pairs at an angle of 45- 60°, subparallel, ascending, arcuating and dis- appearing towards the margin, reticulation fine, scalariform or irregular, obscure on both surfaces; petiole 44-1 cm, adaxially flat. Inflorescences male, female, or androgynous. Male rachis 5-10 cm, slender with sparse simple hairs, bracts ovate- acute, membranous, 1-2 by 1-144 mn, ciliate, bracteoles reduced to a cluster of simple hirsute hairs; g¢ flowers densely arranged but solitary, perianth lobes 6, almost free, ovate-rounded, 1-2 by 14% mm, membranous, ciliate, stamens 12-14, filaments slender, 2-314 mm, anthers 0.3- 0.45 mm long, pistillode rudimentary, 1-144 mm @. Female and androgynous rachis 5-10 cm, slender, with sparse simple hairs, bracts ovate- acute, 1144-2 by 1-1.2 mm, membranous, densely ciliate, bracteoles 1 by 0.7 mm; androgynous rachis with a few ¢ flowers in the apical part; 2 flowers solitary, perianth membranous, deeply incised, lobes 6, 1-1.2 by 0.7 mm, densely hairy, staminodes 12, styles 3, recurved, cylindrical, 1-144 mm, sparsely tomentose at the base. Young infructescen- ce 10-15 cm, carrying 10-20 young cupules more or less ovoid-globose, sessile, enclosing the fruit except for the persistent perianth and styles, sur- face covered all over with scale-like appendages. Ripe cupule 1-1% by 0.7-1.2 cm, (sub)sessile, more or less globose with the fruit partly emerg- ing; wall 14-1 mm thick, inside sparsely hairy, outside densely grey-fulvous puberulous, rather densely set with acroscopical firm scales or small flat-triangular spines, irregular or in 5—7 + regu- lar transversal rows, to 2 mm long with a sharp glabrous tip or obtuse; dehiscence irregular or medianly into 2 equal segments. Fruit solitary, ovoid-conical, pointed, 1-144 by %4-1 cm, lon- gitudinally ribbed (also in the fresh state?), scar flat or rounded, + % part, the remainder free, with dense rufous tomentum, glabrescent. Distr. India (E. Bengal), Burma (Upper Bur- ma, Tenasserim), China (Kweichow, Yunnan), also Taiwan, Siam (NE. and SE. part), Indo-China (Tonkin, Laos, Annam); in Malesia: N. Sumatra (Atjeh, one collection), Malay Peninsula (Kedah, Salangor, Pahang), Java (mountains between Gede and Wilis), N. Borneo (Mt Kinabalu), Celebes (scattered in the mountains), New Gui- nea (very common), also Japen, Misima, Fergus- son, and Goodenough Is., and New Britain. Ecol. In primary or relict forest, often on loamy sandy soil, 300-2500 m. In New Guinea it may form pure stands on ridges up to 1500 m, or grow together with Lithocarpus, Intsia, Anisoptera, or Nothofagus as co-dominants. Fertile almost throughout the year. Vern. Riung anak, S, derived from its peculiar capacity to produce a circle of suckers (young- sters) round the base of the stem. The record in the Kew Index of ‘C. junghuhnii in Hayata, Ic. Pl. Formos. VI Suppl. (1917) 72’ is enigmatic; the name is not in the General Index to that work, and the supplements are of 1925 and later. Notes. Close to C. tribuloides, C. echidnocarpa and C. ferox from India and Burma, C. brevi- spina and C. chinensis from Formosa and China, and C. cuspidata from Japan; it differs mainly in the cupule which is covered with short tuber- cles and not with spines, and to some extent also in the pubescence on the lower surface of the leaves. From its nearest ally in Malesia, C. buruana, it may be distinguished by its smaller leaves with incised margin in the apical half, and by its small- er cupule with shorter tubercles. In several specimens the trunk was recorded to produce many coppices round the base. 21. Castanopsis foxworthyi SCHOTTKY, Bot. Jahrb. 49 (1913) 358; SopEPADMO, Reinwardtia 7 (1968) 391. — C. kinabaluensis A. Camus, Bull. Soc. Bot. Fr. 75 (1928) 698; Chat. (1930) 357, t. 44: 1-5. — Fig. 14 h-i. Tree, (5—)15-36 m by 10-50 cm 9, occasionally 310 fluted or with stout buttresses; bark greyish brown, smooth or finely fiissured, (always?) flaky. Branch- lets initially densely set with fulvous, small, stellate hairs and larger bundle-hairs, or glabrous from the beginning; terminal bud ovoid, 3-4 by 2-3 mm, scales ovate-acute or linear. Stipules 6-8 by 1-2 mm. Leaves thick-coriaceous, 8-22% by 4-8 cm (index 2.1—3.6), widest about the mid- dle, sometimes above, or below; base attenuate or rounded-acute, apex somewhat rounded and acu- tish to rather abruptly acuminate with a sharp tip 14-2 cm; surfaces glabrous, + discolorous, above glossy and olive-greenish, beneath duller and brownish; midrib strongly prominent beneath, slightly so above; nerves 9-14 pairs at an angle of 60-80°, subparallel, ascending, arcuating and dis- appearing towards the margin, thin and subprom- inent on both surfaces, reticulation fine, sca- lariform to irregular, obscure on both surfaces; petiole 34-24% cm, adaxially flat or shallowly fur- rowed. Male rachis 10-15 cm, 1 mm 9; bracts and bracteoles ovate-acute; 3 flowers in clusters of 3, perianth lobes ovate-rounded, 1 mm long, with dense tomentum outside; stamens 12, filaments 2%-3 mm, anthers 0.25 mm long, pistillode 1-114 mm @. Female rachis 10-25 cm by 2-3 mm, bracts ovate-acute; 2 flowers in clusters of 3, perianth lobes ovate-acute, 1 mm long, with dense to- mentum outside, staminodes 12, sometimes con- siderably developed and producing good pollen grains; ovary rounded-triangular, styles 3, conical, 1-14% mm. Young cupule %-1 cm stalked, ovoid- globose, the segments between the 4 sutures cov- ered with spines. Ripe cupules on slender pe- duncle, obovoid-globose, more or less distinctly 2-4-lobed, 3-314 by 2-3 cm, wall 1-2 mm thick, outside velvety and more or less densely set with rather sturdy spines 8-13 mm long, simple or in bundles or tree-like branched and arranged in 4-5 concentric bands, fulvous puberulous with glabrous top; inside densely fulvous-tomentose. Fruit 3 per cupule, more or less ovoid-conical, 1%4-2 by 1-1% cm, the scar covering 1/,_1 part, convex; wall 1 mm thick, the free part densely ful- vous-tomentose; cotyledons flat-convex. Distr. Malesia: Malay Peninsula (Trengganu); Borneo (Sarawak and Sabah; also Nunukan I.). Ecol. Primary or secondary peat-swamp and kerangas forests, also montane forests, up to c. 2400 m. Fi. July—April, fr. May-Febr. Note. In the kerangas forest the twigs are rather straight and slender, in the mountains (“C. kinabaluensis”) they are rather sturdy with shorter internodes, sometimes richer in-lenticels, and with brown leaves in the dried state; in both habitats the leaves are smaller, thicker coriaceous, and with fewer nerves than under more favourable lowland conditions. 22. Castanopsis schefferiana HANcE, J. Bot. 16 (1878) 200; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 105, t. 99; A. Camus, Chat. (1930) 456, t. 66: 4-6; SopPADMO, Reinwardtia 7 (1968) 402. — C. andersonii GAMBLE, Kew Bull. (1914) 179; FLORA MALESIANA [ser. I, vol. 7? J. As. Soc. Beng. 75, ii (1915) 458 (‘andersoni’); A. Camus, Chat. (1930) 342. t. 39: 1-4. Tree, 10-24 m by 30-60 cm g; buttresses up to 1.2 m tall, 5 cm thick; bark smooth, grey. Branch- lets initially tomentellous or glabrous; slender to rather sturdy, sparsely lenticellate; terminal bud ovoid-globose or ellipsoid, 3-5 by 2-3 mm, scales ovate-acute. Stipules linear, 3-7 mm long, very early caducous. Leaves thick-coriaceous, (6—-)8-12(-18) by (1-)3-5(-7) cm (index 2.2-3), widest about the middle to rarely below; base attenuate to rounded, and acute, apex acute to 14,—1 cm acuminate; glabrous, mostly concol- orous, glossy above, dull beneath; midrib strongly prominent beneath, slightly so above; nerves 10-15 pairs, -- prominent beneath, flattish above, at an angle of 45—70°, parallel, ascending, arcuat- ing towards the margin; reticulation fine, dense, more or less regularly scalariform, obscure on both surfaces; petiole 1-214 cm, adaxially flat to deeply sulcate. Male rachis 10-17 cm, 1-2 mm g; bracts and bracteoles ovate-acute, 1 by 1 mm; 3 flowers in clusters of 3—7, perianth lobes mem- branous, 0.8—1 by 0.5—0.7 mm, densely hairy out- side; stamens 12, filaments 2 mm long, anthers 0.2 mm long, pistillode 1-144 mm @. Female or andro- gynous rachis 10-15 cm, 2-3 mm @, bracts and bracteoles ovate-acute, 1 by 1 mm; @ flowers in clusters of 3, perianth lobes 0.7—1 by 0.5—0.7 mm, membranous, densely hairy; staminodes 12, rudi- mentary, styles 3, cylindrical-conical, 1-14% mm. Young infructescence: rachis 20-25 cm, 3 mm @. Young cupule obovoid-globose, the segments be- tween the 4 sutures covered with spines. Ripe cupules scattered along the rachis, (sub)sessile, obovoid-globose, sometimes distinctly 4-lobed, 314-4 cm through; wall 114 mm thick, inside with long, ferruginous, silky hairs, outside more or less densely hoary-puberulous or felty, rather densely set with sturdy somewhat recurved spines 4-15 mm long, solitary or in bundles, hairy with glabrous top; dehiscence into 4 segments or irreg- ular. Fruits 2-3 per cupule, ovoid-conical, 1-14%4 cm through, scar occupying %4—% part, flat- convex; free part of the wall with brownish silky tomentum. Distr. Malesia: NE. Sumatra (Langkat at 4° N), also Riouw and Lingga Arch., Malay Peninsula (scattered; also Singapore). Ecol. Forests, to 1000 m. Fi. Nov.-Dec., fr. April-Dec. Note. The species was included in C. javanica by J. D. HOOKER and by KING. 23. Castanopsis microphylla SoEPADMO, Rein- wardtia 7 (1968) 395. Tree, 20-30 m by 40-60 cm g. Branchlets ini- tially with a tomentum of fulvous stellate hairs of various Size, later glabrescent, mostly slender, dark purplish brown, smooth, sparsely lenticel- late; terminal bud ellipsoid, 5-7 by 2-3 mm, scales linear-acute, 5—6 by 1-2 mm, hairy. Stipules linear- acute, 5—7 by 1-2 mm, hairy. Leaves chartaceous, 5-9 by 14%4-3 cm (index 2.5-4.6), widest at the middle to slightly below; base attenuate-acute to 1972] seldom nearly rounded, top gradually acuminate with a sharp to blunt tip 14-1 cm; surfaces + dis- colorous, above glabrous and rather dull, be- neath densely to very sparsely set with stellate scales (magnification to 60!), their nucleus often orangish, and on the nerves, particularly when young, long simple hairs; midrib + prominent on both surfaces; nerves 8-10 pairs at an angle of 45-60°, ascending, parallel, arcuating towards the margin, thin; reticulation + scalariform, fine, dense, obscure on both surfaces; petiole 3-10 mm, adaxially flat. Male rachis 5-10 cm by | mm g; bracts and bracteoles ovate-acute; ¢ flowers in clusters of 3—7, perianth lobes ovate-acute, 1 by 1 mm, densely tomentose outside, stamens 12, filaments 1-144 mm, anthers 0.2 mm long, pistil- lode 1 mm g. Female rachis 3—5 cm long, 1 mm g, bracts and bracteoles ovate-acute; 2 flowers in clusters of 3, perianth lobes 14 mm long, densely stellate-tomentose outside, staminodes 12, rudi- mentary, styles 3, conical, 1 mm. Young infruc- tescence: rachis 5-7 cm by 2 mm g, carrying 1-8 young cupules ovoid-globose, sessile, 5 by 5 mm, outside with 4 distinct clusters of short spines alternating with the scaly later sutures. Fruits 3, immature, completely enclosed by the cupule. Distr. Malesia: Borneo (Sarawak, Sabah, Kalimantan: Sanggau in the western part, and western Kutei). Ecol. Lowland dipterocarp forest to submon- tane forest up to 1600 m, often on slopes; one rec- ord from basalt derived soil. F/. Aug.—Oct. Notes. Vegetatively distinct; somewhat sug- gestive of C. acuminatissima, but the latter has leaves with a different indumentum, often remote- ly serrulate, and a single fruit in each cupule, whereas in C. microphylla there must be 3. The mature cupule and fruit are unknown. 24. Castanopsis catappaefolia KING ex Hook. f. Fl. Br. Ind. 5 (1888) 621 (‘catalpaefolia’); KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 100, t. 92; A. Camus, Chat. (1930) 341, t. 38; SoEpPADMo, Rein- wardtia 7 (1968) 388. Tree, 20-27 m by 35-50 cm g. Branchlets terete, blackish brown with purple tinge, with minute lenticels; tomentum consisting of small stellate hairs, glabrescent; terminal bud ovoid, 1-114 by 0.8 cm, scales ovate-acute, 1 by 0.4 cm, with stellate tomentum outside. Leaves thick-coriaceous, rigid, lanceolate-oblong, 35-50 by 14-19 cm; base atten- uate-rounded, top rounded-emarginate or shortly acute; above olive-greenish, glossy, glabrous, be- neath dull, with dense cover of appressed stellate scales (magnification 30!), interspersed with a few bundle-hairs, like the midrib above choco- late-brownish; midrib prominent on both surfaces; nerves 23-27 pairs at an angle of + 80°, parallel, arcuating and anastomosing near the margin, flat to impressed above, prominent beneath; reticulation lax, scalariform except near the mid- rib, distinct beneath; petiole 1-144 cm by 5-7 mm, distinctly swollen at the base. Female rachis 25-30 cm; bracts ovate-acute, 2 by 2 mm; @ flow- ers solitary or in clusters of 3, perianth lobes FAGACEAE (Soepadmo) a11 rounded-acute, 5 by 5 mm, with dense stellate tomentum outside; staminodes 12, rudimentary, ovary -+ rounded-trigonous, | by 1 mm, styles 3-4, conical, 1 mm. Cupule (nearly ripe) enclosing the fruits but for a small opening at the top, + ovoid-globose, sometimes distinctly 2-lobed, flat and smooth adaxially, 314-4 by 3 cm, base at- tenuating towards the short peduncle, top roun- ded-acute; wall thin, inside with red-brown silky hairs, outside with some fulvous felt, rather den- sely set with sturdy simple or little-branched spi- nes 6-8 mm long, dull brown and subglabrous (dehiscence not observed). Fruits 1 or 3 per cupule, + ovoid, 114-3 by 3 cm; the scar compa- ratively small; free part of the wall reddish, dense- ly covered with short simple silky fulvous hairs. Distr. Malesia: Malay Peninsula (Goping in Perak), one collection known. Ecol. In open jungle on low hill, 100-170 m. Fr. Sept. Notes. The spelling of the name was corrected by KING in the year after its first publication. The male inflorescence is as yet unknown. 25. Castanopsis argentea (BL.) A.DC. J. Bot. 1 (1863) 182; A. Camus, Chat. (1930) 308, t. 31: 4-5; BACKER & BAKH. f. FI. Java 2 (1965) 4; SOEPADMO, Reinwardtia 7 (1968) 387. — Fagus argentea BL. Flora 7 (1824) 291. — Castanea argentea Bui. Bijdr. (1826) 525; Fl. Jav. Cupul. (1829) 40, t. 21; Koorp. Teysmannia 11 (1900) 134; K. & V. Bijdr. 10 (1904) 5; Koorp. Atlas 1 (1913) t. 37. — Castanea argentea var. rigida Bi. Mus. Bot. 1 (1850) 283. Tree, 15-30 m by 14-1 m g; bark dark grey, fissured and lenticellate. Branchlets initially dense- ly to sparsely fulvous stellate hairy, later dark- coloured, glabrescent, densely lenticellate; ter- minal bud 6-10 by 3—5 mm, scales linear, acute, 5-10 by 1 mm, glabrescent. Stipules linear, acute, 10-15 by 2-3 mm, subglabrous. Leaves thin-coria- ceous, (9—)13-16(—20) by (3—)5—7(—12) cm (index 2-3.5), widest at the middle, seldom above or below; surfaces discolorous, above glabrous, glossy in various shades, beneath silvery-grey with dense, stellate scales (magnification 30!), subglabres- cent; base rounded, attenuate-acute, top acute to 1-2 cm sharply acuminate; midrib and nerves strongly prominent beneath, slightly so above; nerves (9—)11—13(—15) pairs at an angle of 60—70°, subparallel, arcuating, not confluent; reticulation fine, rather lax, more or less scalariform, distinct on both surfaces; petiole (44-)1-14(-2%) cm, adaxially flat. Male rachis 15-25 cm by 2-3 mm @, sometimes with short branches near the base; bracts and bracteoles ovate, rounded to acute, coriaceous, 144-3 by 1-14% mm; ¢ flowers in clusters of 3—7, perianth lobes ovate-acute, 11-2 by 1-144 mm, hairy outside, stamens 12, ifila- ments 2-3 mm, anthers 0.2—-0.25 mm long, pis- tillode 1-2 mm @. Female rachis 5-15 cm by 24mm 9, bracts and bracteoles thick-coriaceous, broadly Ovate-acute, 3-5 by 3-4 mm; @ flowers in clusters of 3, perianth lobes ovate-acute, 1-144 by 1 mm, hairy on both sides, staminodes 12, well-developed, 212 FLORA MALESIANA [ser. I, vol. 7? styles 3, conical, 14,-2 mm, slightly recurved. Young infructescence 15-30 cm long; young cupule subglobose, the spines in 4 sectors with scaly parts in between. Ripe cupule 44-1 cm stalk- ed, ovoid-globose, 3-4 cm through; wall 1-2 mm thick, outside with dense very short fulvous in- dumentum, the spines densely set in bundles of sturdy and slender ones, 1-144 cm long, un- branched, mostly straight; dehiscence into 4 seg- ments or irregular. Fruits 3 in a cupule, ovoid- conical with acute top, 2-244 by 1-1% cm, scar comparatively small, wall 1 mm thick, the free part densely reddish-brown tomentose; cotyle- dons flat-convex. Distr. Malesia: Sumatra (scattered), Java (common, eastwards to Mt Ungaran, 110° 20’ E, and on Mt Wilis at Ngebel). Ecol. Forests, c. 150-1400 m. F/. Aug.—Oct., fr. Nov.—Febr. Use. The fruits are edible. Notes. The leaves underneath are more dis- tinctly silvery than in the other scaly species. Records for Burma have generally been based on specimens of C. Aystrix. 26. Castanopsis scortechinii GAMBLE, Kew Bull. (1914) 178; J. As. Soc. Beng. 75, ii (1915) 455; A. Camus, Chat. (1930) 334, t. 34: 9-11; SoEPAD- MO, Reinwardtia 7 (1968) 402. Small tree. Branchlets initially densely set with fulvous tomentum, later glabrescent, greyish black; terminal bud ovoid, c. 5 by 3 mm, scales linear-acute. Stipules linear-acute, c. 10 by 3 mm, densely fulvous tomentose, soon caducous. Leaves thin-coriaceous, elliptic-oblong or oblong- lanceolate, 10-1214 by 3-44% cm; discolorous, above glabrous, glossy, dark chocolate-brown, underneath with a dense cover of fulvous ad- pressed, minute stellate hairs and erect long simple hairs; base, acute or rounded, margin entire, re- curved, apex abruptly acute or 4-1 cm sharp acuminate; midrib and nerves prominent beneath, flattish above; nerves 10-15 pairs, parallel, ascend- ing, at an angle of c. 70°, arcuating but not ana- stomosing towards the margin; reticulation fine, dense, subscalariform, distinct beneath; petiole c. 5 mm, 2 mm g, terete, densely fulvous tomentose. Female rachis 15-20 cm; @ flowers in clusters of 3. Cupule subglobose, 2-3 cm through; spines much-branched, tips sharp, 1-114 cm long, dense- ly fulvous tomentose, densely but irregularly set. Fruits ovoid-conical, asymmetrical, 1-144 cm through, adaxially flat; wall thin, free part densely fulvous tomentose; scar small, convex or flat. Distr. Malesia: Malay Peninsula (Perak and Cameron Highlands). Ecol. Forests, 1000-1200 m. Notes. Quite near to C. fulva, but different by its leaf with a shorter petiole, and different in- dumentum, and by its cupule set with long, dense- ly fulvous-tomentose spines. The male inflorescence is not yet known. 27. Castanopsis fulva GAMBLE, Kew Bull. (1914) 179: A. CAMus, Chat. (1930) 327, t. 33: 6-9; SOEPADMO, Reinwardtia 7 (1968) 391. Tree, 18-24 m by 25-50 cm go. Branchlets initially angular and covered with dense fulvous tomentum, later glabrescent, rather sturdy, sparse- ly lenticellate; terminal bud ovoid-globose, 3 by 3 mm, scales ovate-acute 3 by 1-2 mm. Stipules 10 by 4 mm, elliptic with acute top. Leaves 9-17 by 3-614 cm (index 2.1—3.2), widest at the middle to sometimes above; base rounded and abruptly acute, top sharply acute to acuminate; discolo- rous, above glossy (greenish) brown, glabrous, beneath dull brown, sparsely to densely covered with closely adpressed, stellate scales (magnifica- tion 30!) and, particularly on the nerves, with some longer simple or 2-3-fid hairs, sometimes late glabrescent; midrib strongly prominent be- neath, slightly so above; nerves 10-15 pairs at an angle of 60—70°, ascending, parallel, arcuating and disappearing near the margin, prominent beneath, flattish above; reticulation fine, scalariform, ob- scure on both surfaces; petiole 1-3 cm, shallowly furrowed adaxially, with distinct basal joint. Male rachis 10-20 cm, bracts ovate-acute, thick-coria- ceous, 1144-2 by 1-144 mm, bracteoles membra- nous, smaller; 3 flowers in clusters of 3—7, perianth lobes membranous, ovate-acute, 1 by 1 mm, densely hairy, stamens 12, filaments 144-2 mm, anthers 0.2-0.25 mm long, pistillode 144-2 mm @. Female rachis 10-15 cm, 2-3 mm g, bracts and bracteoles thick-coriaceous, broadly ovate-acute, 1-14% by 144-2 mm; 2 flowers in clusters of 3, perianth lobes membranous, 0.7 by 0.7 mm, dense- ly hairy on both sides, staminodes 12, rudimen- tary, styles 3, conical, slightly recurved, 1 mm. Young infructescence: rachis 15-20 cm, 3-5 mm 9, young cupule sessile, ovoid-globose, 4-1 by % cm, the 4 spiny and 4 scaly sectors + equal. Cupule (nearly ripe) on a stalk 4% cm long and wide, or subsessile with tapering base, ovoid-globose, 2-244 cm through; wall 1 mm thick, inside hairy, out- side fulvous pubescent, densely set with sturdy, recurved spines 14-1 cm long, in bundles and/or branched themselves, arranged in 4-6 concentric rows, except the top densely puberulous; dehis- cence (always?) irregular. Fruit (only one of the 3 well-developed!) ovoid-conical with acute top, 114-2 cm through, scar comparatively small, wall 1 mm thick, free part densely fulvous-pubescent; cotyledons flat-convex. Distr. Malesia: Central Sumatra (SW. of Pakanbaru), Malay Peninsula (Perak, Selangor, Pahang, Johore), Borneo (W. Kalimantan: G. Kenepai; Sarawak; Brunei; Sabah). Ecol. Primary lowland dipterocarp forest, in the hills up to 200 m, on poor sandy soil. Fi. Sept.-May, fr. July-Jan. 28. Castanopsis costata (BL.) A.DC. J. Bot. 1 (1863) 182; Prod. 16, 2 (1864) 110; HANcE, J. Bot. 13 (1875) 367; GamBLE, J. As. Soc. Beng. 75, ii (1915) 456; A. Camus, Chat. (1930) 328, t. 33: 10-11; SozpADMO, Reinwardtia 7 (1968) 388. — Castanea costata BL. Mus. Bot. 1 (1850) 284. — Castanea brevicuspis Mia. Fl. Ind. Bat. 1, 1 (1858) 1972] FAGACEAE (Soepadmo) 313 Fig. 14. Castanopsis costata (BL.) A. DC. a. Habit, x 24, b. 3 flower, c. 9 flowers, d. longitudinal section of a 2 flower, e. ripe cupule, abaxial side, f. ripe cupule, adaxial side, g. fruit. — C. foxworthyi SCHOTTKY. h. Ripe cupule, abaxial side, i. ripe cupule, side view. All flowers x 4, cupules and fruit nat. size (a RASTI- NI 203, b DELMAAR 1899, c-d KURNIASIH 29, e SAR 15116, f-g SAN 16747, h-i SAR 4416). 866. — Castanea spectabilis Mia. l.c. — C. brevi- cuspis (MiqQ.) A.DC. J. Bot. 1 (1863) 182. — C. spectabilis (Miq.) A.DC. lc. — C. trisperma ScHEFF. Nat. Tijd. N. I. 31 (1870) 362. — C. costata B bancana ScuerrF. |.c. — Fig. 14 a-g. Tree, 10-36 m by 15-80 cm @9; bark yellowish to reddish brown. smooth and peeling off profuse- ly. Branchlets when young densely set with ful- vous bundle-hairs, later glabrescent, slender, grey- ish or blackish brown, with many minute lenticels; terminal bud ovoid, 3—5 by 2-3 mm, scales ovate- acute, 3-4 by 2-3 mm. Stipules linear-triangular, 13 by 14% mm. Leaves sometimes thickly coria- ceous, (9-)14-17(-25) by (4-)5-8(-10) cm (index (1.7-)2.5(-3.2)), widest at or below the middle to sometimes above; base attenuate-acute or rare- ly rounded and abruptly acute, apex bluntly acute or sharply 144-1 cm acuminate; above glabrous or sometimes on and near the midrib with a few hairs, glossy olive-green, underneath 314 dull brown with a dense cover of stellate scales (magnification 60!), sometimes also roughish with 2-3-fid bundle-hairs; midrib and nerves strongly prominent beneath, slightly so above; nerves (14—)16—-17(—20) pairs, ascending, at an angle of 50—70°, parallel, arcuating and disappear- ing towards the margin; reticulation fine, dense, scalariform, obscure on both surfaces; petiole (44-)1-1%(-2) cm, adaxially flat. Male rachis 5-15 cm, 1-2 mm 9; bracts and bracteoles ovate- acute, 1-144 by 1 mm; ¢ flowers solitary or in clusters of 3-7, perianth lobes rounded or acute, membranous, 0.7—1 mm long, densely tomentose; filaments 2—3 mm, anthers 0.2-0.25 mm long; pistillode *4-14 mm @. Female rachis 10-20 cm, 2-3 mm 9g, bracts and bracteoles broadly ovate- acute, thick-coriaceous, 1-144 by 1 mm; & flowers in clusters of 3; perianth lobes acute, 34-1 mm long, densely hairy outside; staminodes 10-12, exceeding the perianth; styles 3, conical-cylindri- cal, recurved, 1-144 mm. Young infructescence up to 20 cm; young cupules numerous, subsessile, obovoid-globose; the segments between the 4 sutures covered with spines. Ripe (?) cupule sub- sessile to 7 mm stalked, ovoid-globose, 24-3 by 3—4 cm; wall 1-2 mm thick, outside densely fulvous hairy, and more or less densely set with bundles of spines 4-14 mm long, fulvous-puberulous ex- cept the top, sturdy and more or less recurved or sometimes the inner ones of a bundle slender and straight; inside of wall with dense yellowish brown, silky tomentum. Fruits 1-3 per cupule, more or less conical, 144-2 by 1-114 cm, the scar covering Y, part, convex; the free part with dense yellowish brown silky tomentum; wall thin. Distr. Malesia: Sumatra (scattered), also Ban- ka and Billiton Is., Malay Peninsula (Perak), Borneo (scattered all over), also Laut and Nunu- kan Is. Ecol. Lowland to submontane forests, up to c. 1500 m. FI. March-June, fr. Sept.-May. Notes. The cupule resembles those of C. fulva and C. foxworthyi, but the leaf is different. In SAN 27360 from North Borneo, in young fruit, the leaves are completely glabrous, and glaucous underneath. In SAN 16445, SAN 28260, CLEMENS 10791 and 32169, all from Mt Kinabalu, the leaves, although with the characteristic close, parallel nerves, have underneath the long hairs much more densely set than in the other specimens; the young twigs being also very densely hairy. In SAN 16445, the fruit is 514 cm in largest @, the spines being very sturdy and 1-2 cm long. 29. Castanopsis motleyana KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 96, t. 86 (‘mottleyana’); A. Camus, Chat. (1930) 339, t. 37: 6-10; SoEPADMO Reinwardtia 7 (1968) 396. — C. javanica (non A.DC. ) Evmer, Leaf]. Philip. Bot. 3 (1910) 936. — C. pearsonii MERR. Philip. J. Sc. 30 (1926) 79. ‘Tree, 10-40 m by 20-70 cm g; buttresses up to 3-4 m tall, 1-2 m out, 5-10 cm thick, spreading; bark rough, fissured or flaky, chocolate- or reddish brown. branchlets when young with dense fulvous FLORA MALESIANA [ser. I,@vol: <7? indumentum of longer and mostly also shorter bundle-hairs, glabrescent, slender to sturdy; ter- minal bud ovoid-ellipsoid, 5-10 by 3-5 mm, scales ovate- or linear-acute, 4-5 by 1-2 mm. Stipules Ovate-acute, 7-10 by 3 mm. Leaves thick-coria- ceous, (7—)15—23(—32) by (244—)7-10(-16) cm (in- dex 2-2.8(—3.5)), widest at the middle to some- times above; base attenuate-acute, top acute with a small sharp acumen; above variable in colour and shine, when young sometimes near the nerves sparsely set with simple or 2—3-fid bundle-hairs which leave a pit when disappeared, underneath dull brown, glabrous to densely set with stellate scales (magnification 30-60!), and/or simple or 2—3-fid bundle-hairs; midrib and nerves strongly prominent beneath, slightly so above; nerves (11—)14-18(-21) pairs at an angle of (S0—)70-80°, mostly parallel, ascending, arcuating and anasto- mosing near the margin; reticulation coarse, dense, subscalariform, distinct beneath; petiole (44—-)1- 144(2\%) cm, hairy as the twig, adaxially flat. Male rachis 10-20 cm, 2—3 mm g; bracts and bract- eoles membranous, ovate-acute, 10 by 7 mm; 3S flowers in clusters of 3-7, perianth 4—6—lobed, the lobes acute, 1-144 by 0.7-1 mm, densely yellowish brown stellate-tomentose; stamens 10— 12, filaments 2-24 mm, anthers 0.2-0.25 mm long, pistillode rounded-triangular, 144 mm o. Female rachis 15-25 cm, 3—5 mm @; bracts and bracteoles thick-coriaceous, ovate-acute, 1 by 0.7 mm; @ flowers in clusters of 3, perianth 6—lobed, the lobes 0.7—1 by 0.3—0.6 mm, with dense stellate tomentum on both surfaces; staminodes 12, rudi- mentary; styles 3, conical, recurved, 1-144 mm. Young infructescence 10-20 cm, rachis 3-5 mm @, carrying 10-25 young fruits. Young cupule sessile, depressed-globose, with short, sharp, slender and/ or flat sturdy spines pointing to all directions. Ripe cupule ellipsoid-globose, 3-4 cm through, sometimes flattened adaxially and asymmetrical; wall densely hairy both outside and inside, 3-4 mm thick, densely set with spines 34-11% cm long and densely fulvous or golden puberulous (magnifica- tion 10!), their top glabrous, of a slender, straight kind and in bundles, and/or of a sturdy kind like trees with recurved branches; dehiscence into 4 segments or irregular. Fruit ovoid-conical, 2-244 by 1144-2 cm, the scar covering % part, flat or convex, the free part densely yellowish brown tomentose, top acute; wall 1 mm thick; cotyledons flat-convex. Distr. Malesia: Borneo (throughout), also Nunukan I., Philippines (Mindanao). Ecol. Primary, rarely secondary or disturbed forest in hilly country up to 500 m, the soil record- ed sandy clayey. Fi. Sept.-May, fr. May-Dec. Notes. In D. D. Woop (coll. EVANGELISTA) 1151 from North Borneo, the type of C. pearsonii, the cupule (detached from the leaves) is 5 by 64% cm, the sutures being free of spines. In the few Philippine specimens partly the leaves resemble those of the type of C. pearsonii, i.e. lanceolate with comparatively few arcuating veins under a sharp angle, and scaly beneath, partly they resemble other materials from Borneo, 1972] with wider leaves, more and parallel veins under a wider angle, and glabrous beneath. Leaves longer than 20 cm were never found to be scaly beneath, but no further correlations could be detected in the considerable diversity as contained in the above description, neither in structure nor in geographical or ecological distribution, and the extremes are all connected by intergrades. 30. Castanopsis pedunculata SopPpADMO, Reinward- tia 7 (1968) 399. Tree, 18 m. Branchlets greyish brown, glabrous, densely lenticellate; terminal bud ovoid-globose, 2-3 by 2 mm, with dense stellate tomentum, glabrescent. Leaves concolorous, elliptic-oblong, 8-16 by 2-21 cm; base rounded and abruptly acute, apex acute to 44-1 cm acuminate; glabrous on both surfaces; midrib prominent beneath, flat above; nerves 10-13 pairs, ascending, subparallel, arcuating towards the margin, slightly prominent on both surfaces; reticulation distinct on both sides; petiole 1-2 cm, adaxially shallowly furrow- ed. Rachis of the infructescence rigid, up to 18 cm, % cm thick, densely lenticellate, with adpres- sed stellate tomentum, glabrescent. Ripe cupule compressed obovoid-globose, more or less sym- metrically 4-lobed, thin-walled, completely enclos- ing the fruits; surface with regular undulating ridges of flattened minute tubercles, fulvous-hoary, 2-3 by 3-314 cm; dehiscence into 4 + equal seg- ments; peduncle 1-144 by 1% cm, rugose and usually twisted. Fruits 3 in each cupule, ovoid- conical, 144-2 by 1.5-1.8 cm, the adjoining sides of both central and lateral fruits found concave, the scar covering 1/3—2/5 part, the free part glossy with a thin short indumentum; top abruptly point- ed; cotyledons flat-convex. Distr. Malesia: North Borneo (one collection). Ecol. Forest at 300 m. Fr. Sept. Notes. Distinct by its almost smooth cupule on the long peduncle. Inflorescences are unknown. 31. Castanopsis psilophylla So—EPADMO, Reinward- tia 7 (1968) 401. — C. inermis (non B. & H.) MERR. Pl. Elm. Born. (1929) 42. — Fig. 15 e-f. Tree, 10-27 m by 20-45 cm g; buttresses up to 2 m tall, 34 m out; bark greyish brown, smooth or scaly. Branchlets glabrous almost from the be- ginning, smooth, dark purplish brown, lenticels none or very few; terminal bud ovoid-ellipsoid, 2-3 by 1-2 mm, scales linear-acute, sparsely pube- rulous. Stipules linear, 3-4 mm, very early cadu- cous. Leaves somewhat discolorous, mostly glossy above, glabrous on both sides, (6—)10(-16) by (1-)314(5) cm (index (2.3—)2.8-4.2), widest about the middle, sometimes below; base rounded to acute, slightly decurrent, top acute to sharply acuminate with tip 44-2 cm; midrib and nerves more or less prominent on both surfaces; nerves 9-12 pairs at an angle of 60—70°, parallel, arcuating and disappearing towards the margin; reticulation fine, obscure on both surfaces; petiole 44-114 cm, adaxially flat or shallowly sulcate, glabrous. /nflo- rescence male, female, or mixed. Male rachis 5-15 FAGACEAE (Soepadmo) 315 cm, 1—2 mm @, with dense stellate tomentum; bracts ovate-acute, | by 5mm, also hairy; 9 flowers in clusters of 3—7, perianth deeply 6—-lobed, the lobes 1 mm long, hairy outside; stamens 12, fila- ments 144-2 mm, glabrous, anthers 0.2-0.25 mm long, pistillode rounded-triangular, 1 mm @, den- sely woolly pubescent. Female or mixed rachis 6 cm, with dense stellate tomentum; @ flowers in clusters of 3-7, perianth deeply 6—lobed, lobes 0.5-0.75 mm long, acute, hairy outside; stamino- des 12, well-developed and producing good pollen or rudimentary; styles 3, conical, recurved, 1 mm. Young infructescence to 20 cm, carrying 4-12 young fruits. Young cupule subsessile, turbinate, beset with concentric bands of short tubercles. Ripe cupule asymmetrically pear-shaped or depres- sed-obovoid, with 2 or more lobes, completely enclosing the 3—7 fruits, green-brownish velvety with stellate or simple hairs, 114-24 cm through; wall 4% mm thick, covered with several symmetri- cal ridges of short tubercles sometimes recurved; dehiscence into 2—4 segments, more or less regular. Fruits 3 or more, ovoid-complanate, 144-2 cm through, scar covering 2/5—4/5 part of the surface, the remainder densely brownish tomentose. Distr. Malesia: Borneo (Sabah, common; Sarawak, rare, in the northern parts; Kalimantan, scattered in the eastern half), also Banguey I., Philippines (Palawan, EBALO 408). Ecol. In forests on sandy or basaltic soil, in the lowlands up to 1000 m. Fil. April and July, /r. June and Nov. Notes. The nearly ripe fruit recorded green. Nearest to C. inermis and C. philipensis; see the key. Bornean records of C. inermis belong here. C. brevispina ScHoTTKY, Bot. Jahrb. 49 (1913) 358, non HAYATA 1911, might belong here, but the description of the cupule does not agree in all points, and the type specimen, Foxwortuy 156 from Sarawak, seems to be lost. 32. Castanopsis inermis (LINDL. ex WALL.) B. & H. Gen. Pl. 3 (1880) 409; A. Camus, Chat. (1930) 447, t. 63, p.p.; CORNER, Ways. Trees (1940) 292, f. 93, pl. 219; SoppADMo, Reinwardtia 7 (1968) 392. — Castanea inermis LINDL. ex WALL. Pl. As. Rar. 2 (1830) 6; A.DC. Prod. 16, 2 (1864) 116. — Castanea glomerata [non (ROxB.) WALL. ex BL.] BL. Mus. Bot. 1 (1850) 283, quoad specimina, excl. syn. — Calleocarpus sumatrana Mia. Pl. Jungh. (1851) 14; Fl. Ind. Bat. 1, 1 (1858) 868. — C. suma- trana (Miq.) A.DC. J. Bot. 1 (1863) 182; Prod. 16, 2 (1864) 113; Hook. f. Fl. Br. Ind. 5 (1888) 623, p.p.; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 103, t. 97; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 460. — Castanea sumatrana (MiQ.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 378. — C. mitifica HANCE, J. Bot. 16 (1878) 200. Tree, 10-30 m by 30-90 cm g; bark grey, rough and lenticellate. Young branchlets greyish black, tomentose, older ones glabrous, lenticellate; ter- minal bud ovoid-ellipsoid, 5 by 2 mm, scales ovate- acute, 5 by 2 mm, more or less hairy. Stipules, if distinct, triangular, 1 mm, caducous. Leaves (6-)10-15(-20) by 4-714 cm (index 2.1—3), widest 316 FLORA MALESIANA [ser. I, vol. 7? slightly below to slightly above the middle; discol- orous, upper surface glabrous, dull, lower surface with greyish brown stellate tomentum (magnifi- cation 35!); base rounded to acute, somewhat decurrent, apex more or less acute; midrib and nerves -++ prominent beneath, slightly so above; nerves 10-15 pairs at an angle of 60—80°, parallel, arcuating towards the margin; reticulation fine, parallel, obscure on both surfaces; petiole glab- rous, 1-2 cm, adaxially flat or shallowly furrow- ed. Male rachis 10-15 cm, 1 mm 9g; bracts ovate- acute, 14 by 4% mm, with dense tomentum; ¢ flow- ers solitary or in clusters of 3-7; perianth deeply 6—lobed, lobes rounded, 4% mm long, hairy; sta- mens 12, filaments 2-3 mm, glabrous, anthers 0.1-0.2 mm long; pistillode rounded-triangular, 1 mm g, densely woolly pubescent. Female and mixed androgynous rachis 10-15 cm, 1-2 mm g; 2 flowers solitary or in cluster of 3-4; perianth deeply 6—lobed, lobes rounded, 44 mm long hairy, staminodes 12, styles 3, conical-cylindrical, 4% mm, pubescent at base. Young cupule on stalk 2-5 mm, subglobose, densely covered with the future spines pointing to the top. Ripe cupule 2-4 by 2-3 cm, hoary with minute fulvous hairs, depressed obo- void-globose, regularly or irregularly 3—4—lobed, sometimes constricted at both ends; surface mar- ked with 4-5 more or less parallel, curving bands bearing short, more or less distinct thick tubercles; wall 1-2 mm thick, completely enclosing the 1-4 fruits. Dehiscence badly known; by 4 valves? Fruits 1-4, with dense reddish brown tomentum, 2-3 by 14-2 cm, completely free from the cupule except for the convex, glabrous, rugose base; central fruit in cross-section asymmetrically round- ed-triangular, ovoid-conical, flattened laterally and rounded adaxially; lateral fruits smaller, ovoid-globose, flattened adaxially and rounded on the free sides. Distr. Malesia: Sumatra (mostly in the south- ern part; twice found in Tapanuli), also Banka and Lingga Is., and Malay Peninsula (common; also Singapore). Ecol. Forests, mostly in the hills, also found in bamboo forest, and in groves on a river-bank; up to 600 m. Fi. Oct.-June, fr. July-Dec. Vern. Berangan, with variants. Notes. Branches spreading. Leaves darkish green. Flowers cream-coloured or the lobes brown- ish-tipped; stamens white. Fruit when unripe pale bluish green or greyish. Materials from Burma formerly reckoned here, generally belong under C. /anceaefolia, which differs in the cupule having concentrical rings out- side and a single fruit inside. C. inermis, C. psilophylla from Borneo, and C. philipensis from the Philippines are closely related, C. lucida being more remote. They can best be distinguished by comparison and then will be found to differ in a number of correlating characters, summarized in the key. Records of this species from Borneo relate to C. psilovhylla. 33. Castanopsis philipensis (BLANCO) VIDAL, Rev. Pl. Vasc. Filip. (1886) 265 (‘philippinensis’); MERR. Philip. J. Sc. 3 (1908) Bot. 319 (‘philippensis’); Brown, Min. Prod. Philip. For. 2 (1921) 260, f. 11; Merr. En. Philip. 2 (1923) 25; SoEPADMo, Reinwardtia 7 (1968) 399. — Fagus philipensis BLANCO, FI. Filip. ed. 2 (1845) 503; ed. 3, 3 (1879) 132. — C. sumatrana (non A.DC.) F.-VILL. Nov. App. (1880) 210; VIDAL, Sinopsis Atlas (1883) 41, t. 92: 4; Evmer, Leaf]. Philip. Bot. 3 (1910) 936. — C. javanica (non A.DC.) VIDAL, Sinopsis Atlas (1883) 41, t. 92: 1; Etmer, Leaf]. Philip. Bot. 3 (1910) 936; Merr. En. Philip. 2 (1923) 24. — C. glabra MERR. Philip. J. Sc. 9 (1914) Bot. 354. — C. inermis (non B. & H.) MerR. En. Philip. 2 (1923) 24. — Fig. 15 a-d. Tree, 6-28 m by 0.2—1 m g; bark yellowish or greyish brown, smooth, densely lenticellate. Inno- vations with dense stellate tomentum. Branchlets soon glabrous, dark purplish coloured, with sparse minute lenticels; terminal bud ovoid or ellipsoid, 2-3 by 1 mm, scales ovate- or linear- acute, 2-3 by 1 mm. Stipules linear-acute, 3-4 by 1 mm, very soon caducous. Leaves 6-16 by 2—5 cm (index 2.3-4.2), widest below the middle to rarely above, discolorous, above green-brownish and often glossy, glabrous, beneath dull brownish with adpressed scale-like stellate tomentum (magnifi- cation 35!); base acute to rarely rounded, slightly decurrent ,+ asymmetrical; apex mostly gradually tapering and acuminate with a sharp or blunt tip 4-2 cm; midrib and nerves on both surfaces brownish and more or less prominent; nerves 7-10 pairs, ascending, subparallel, arcuating and dis- appearing towards the margin, at an angle of 50— 70°; reticulation obscure on both surfaces; petiole (3—)5-10(-20) mm, adaxially flat. Male rachis 10-25 cm rather densely tomentose; bracts and bracteoles ovate-acute, 2 by 1 mm, densely hairy; 3 flowers in clusters of 3-7, perianth lobes 1 mm long, rounded or acute, densely hairy outside; stamens 12, filaments 2-214 mm, glabrous, anthers 0.2-0.25 mm long; pistillode rounded-triangular, 1-144 mm g, densely woolly pubescent. Female rachis 10-20 cm, rather densely tomentose; bracts and bracteoles ovate-acute, 1144-2 by 1 mm; @ flo- wers in clusters of 3 or sometimes solitary, perianth lobes 144-1 mm long, rounded or acute, densely hairy outside; staminodes 12, rudimentary; styles 3, conical, recurved, 1-2 mm. Young cupule sub- sessile to 7 mm stalked, obovoid-globose, covered all over with thick, scale-like appendages. Ripe cupule pear-shaped or depressed-subglobose, 2-3 by 3-4 cm, with two rounded lobes more or less symmetrical; surface with undulate ridges of short, pointed tubercles, hoary to velvety with fulvous to brownish hairs; peduncle 5-7 mm long, 3-4 mm thick. Fruits 3 (rarely 1?) per cupule, the scar covering about half the surface, the remainder free and red-brown, with golden hairs; central fruit 14, by 2 cm, lateral ones as large to slightly smaller. Distr. Malesia: Philippines (Luzon, Mindanao, Leyte, Basilan, Samar, Mindoro). Ecol. Forested slopes, c. 400-1800 m. Dec.- May, fr. June-Oct. 1972] FAGACEAE (Soepadmo) 317 Fig. 15. Castanopsis philipensis (BLANCO) VIDAL. a. Habit, x 24, b. young cupule, c. ripe cupule, abaxial view, d. ripe cupule, seen from above, all nat. size. — C. psilophylla SOEPADMO. e. ripe cupule, aba- xial view, f. ripe cupule, side (lateral) view, both nat. size. — C. rhamnifolia (Miq.) A. DC., g. ripe cupule, adaxial view, h. ripe cupule, abaxial view, i. ripe fruit, side view, all x 2 (a VIDAL 611, b AHERN’S Coll. 8100, c-d ELMER 11315, e-f SAN 25953, g MEIJER 7660, h-i GRASHOFF 362). 318 Uses. Timber tree. Fruits edible; see BRown. Notes. Flowers are recorded as being white. MERRILL (1908), assuming a typographical er- ror, proposed to change the name into C. philip- pensis. In our opinion, there is not enough evidence to adopt this. Very close to C. inermis and C. psilophylla; see the key. 34. Castanopsis lucida (NEES) SOEPADMO, Rein- wardtia 7 (1968) 394. — Alseodaphne lucida NEES in Wall. Pl. As. Rar. 2 (1831) 72. — Laurus lucida WALL. Cat. (1830) 2590, nomen. — C. hullettii KING ex Hook. f. FI. Br. Ind. 5 (1888) 623; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 104, t. 98; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 459; A. Camus, Chat. (1930) 453. Tree, 12-20 m by 40-60 cm 9; buttresses up to 1.6 m tall, 1.2 m out, straight; bark pale-brown, fissured. Innovations with dense stellate tomen- tum. Branchlets sturdy, initially dull dark-col- oured, later greyish, with many lenticels; ter- minal bud ovoid-globose, 3-6 by 2-4 mm, scales Ovate-acute, 3-5 by 2-3 mm. Stipules deltoid, minute. Leaves thick-coriaceous, (10—)16—-20 by (3144-) 8-10 cm (index 1.7—3), widest about the middle, on sterile basal shoots to 38 by 18 cm; above glabrous, glossy sometimes dull, beneath with dense, brownish, stellate scale-like tomentum (magnification 50!); base rounded to attenuate- acute, apex rounded to acutish, sometimes 14-1 cm acuminate; midrib prominent on both sides; FLORA MALESIANA (ser. Live 7 nerves 15-20 pairs, parallel, ascending, arcuating towards the margin, at an angle of (45-)70-80 (-90)°, prominent beneath, flat to subprominent above; reticulation fine, obscure on both surfaces; petiole 144-314 cm, adaxially flat. Male rachis 10-20 cm, bracts ovate-acute; ¢ flowers in clusters of 3-7, perianth deeply 6-lobed, lobes rounded- acute, 1 by 1 mm, hairy outside; stamens 12, fila- ments 344-4 mm, glabrous, anthers 0.25 mm long; pistillode roundish, 1-114 mm g, woolly pubescent. Female rachis 10 cm; @ fiowers in clusters of 3(-4-5); perianth deeply 6-lobed, lobes acute, 4 cm long, hairy outside; staminodes 12, rudi- mentary; styles 3, conical, %-1 mm. Young cu- pule sessile, subglobose, the future spines scale- like, densely set all over the surface, acroscopical. Ripe cupule subsessile, depressed-subglobose or sometimes constricted at base and top, 3—4-lobed, 3144-4 cm through, thick, woody, covered with 3-4 prominent, transverse, curved ridges bearing sharp short spines; dehiscence into 3-4 equal seg- ments. Fruits 2-4 in each cupule, complanate- subglobose, 114-2 cm through, shining, smooth, with sparse, adpressed, stellate tomentum; base convex, rugose, glabrous. Distr. Malesia: Malay Peninsula (Penang, Perak, Negri Sembilan, Malacca), also Singapore; Borneo (SW. and NE. Kalimantan). Ecol. Lowland forests, to c. 500 m. Fi. Aug.— May, fr. April-May. Note. KinG’s record for Riouw and Billiton is not confirmed; he may have had C. inermis. 3. LITHOCARPUS BLUME, Bijdr. (1826) 526; Fl. Jav. Cupul. (1829) 34; OupEM. Natuurk. Verh. Kon. Akad. 11 (1865) 19; ReHp. & Wits. in Sargent. Pl. Wils. 3 (1917) 205; BARN. Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 332; A. Camus, Chénes 3 (1954) 511; SOEPADMO, Reinwardtia 8 (1970) 197. — Synaedrys LINDL. Intr. Nat. Syst. ed. 3 (1836) 441; HANcE in Hook. J. Bot. 1 (1849) 175; Kompz. Bot. Mag. Tokyo 30 (1916) 186, p.p., excl. sect. Chlamydobalanus. — Arcaula RaFin. Alsog. Am. (1838) 30. — Balanaulax RAFIN. Alsog. Am. (1838) 28. — Cyclobalanus (ENDL.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80; ScHwarz, Notizbl. Berl.-Dahl. 13 (1936) 6. — Pasania (MIQ.) OrrstT. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81, p.p., excl. subg. Chlamydobalanus; PRANTL in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 55, p.p., excl. ditto; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 411; HickEL & A. CAMuUus, Ann. Sc. Nat. Bot. 3 (1921) 386, p.p., excl. sect. Pseudocastanopsis; Fl. Gén. I.-C. 5 (1930) 962, p.p., excl. ditto; SCHWARZ, Notizbl. Berl.-Dahl. 13 (1936) 6. — Fig. 18-29. Trees or shrubs; buttresses or stilt-roots occasionally present. Innovations dense- ly yellowish brown to rufous tomentose by simple or stellate hairs. Terminal bud ovoid to ellipsoid; scales spirally imbricate. Stipules extrapetiolar, mostly caducous. Leaves spirally arranged, entire, thin- to thick-coriaceous, margin sometimes strongly recurved, glabrous or variously hairy at least beneath. Petiole terete or adaxially flat or furrowed. Inflorescence male, female, androgynous, or mixed, always erect, including bracts and bracteoles variously densely hairy. Male rachis solitary in the axil of a lower leaf or in a dense paniculate cluster 1972] FAGACEAE (Soepadmo) 319 on lateral or subterminal shoots, simple or much-branched; ¢ flowers solitary or in dichasial clusters of 3—7(—30) along the rachis, or in an androgynous inflores- cence along the upper part of the rachis; perianth campanulate, (4-)6(—7)-lobed; stamens (8—-)12(-15); filaments filiform, glabrous, anthers reniform, dorsifixed, 0.2-0.3(-0.5) mm long; pistillode always present, globose or subglobose, some- times compressed, hairy. Female, androgynous or mixed rachis solitary in the axil of a higher leaf or on the upper part of the paniculate cluster; ° flowers solitary or in dichasial clusters of 3—7(-15) along the rachis, or in an androgynous inflores- cence along the basal part of the rachis; perianth regularly 6-lobed, staminodes 10-12, rudimentary or well-developed and exceeding the perianth, sometimes polliniferous; style 3-6(-15), conical or terete, more or less connate or recurved, densely tomentose at base, stigma punctiform, terminal. Cupule solitary or in dichasial clusters of 3-7(—15) along the rachis, sessile or 1-2 cm stalked, enclosing a solitary fruit, cup-shaped, saucer-shaped to almost globular, variously lamellate, squamose, tuberculate or muricate, never truly spiny, never with vertical sutures; dehiscence none or irregular. Fruit in cross-section rounded, sometimes falsely multi-septate by intrusions of the endocarp; pericarp woody or bony, for the great- er part free to adnate to the cupule, scar small to very large, glabrous or hairy. Cotyledons flat-convex. Germination hypogeal. Distr. About 300 spp., in the NE. parts of India (Nepal, Bhutan, Assam), Burma, China (except in the NW. and W. parts), Korea, Japan, Formosa, Hainan, Indochina, Siam, Malesia, and 1 species in the SW. part of the United States. The most primitive species (L. elegans) occurs in most areas except in Korea, Japan, Hainan, and the United States. In Malesia c. 110 spp., distributed throughout the region eastwards to Sudest and Rossel Is. The genus is rare in the E. part of Java, absent in Madura, the Lesser Sunda Islands, and the SE. Moluccas. Fig. 16-17. Fossil records. Very few fossils have been recorded, viz those enumerated by LA MotTE (Mem. Am. Geol. Soc. 51, 1952, 205-206) from various Miocene-Pliocene deposits in several localities in the United States, and probably also some pollen grains mentioned by MULLER (Proc. 10th Int. Bot. Congr. Edinb. 1964, 371) from the Tertiary of Borneo. Ecol. In Malesia the genus occurs mainly in the everwet lowland to montane forest, between 0-3000 m, more commonly below 1500 m, on various types of soil including limestone. In Central Java a few species have been recorded from pockets of everwet forest on Mts Diéng, Merapi, Ungaran, Muria, Lawu, and in E. Java on Mts Wilis, Kawi, Ardjuno, Tengger, and Idjen. oT, \Yr-~ \y i | gat Fig. 16. Distribution of the genus Lithocarpus, the fossil records indicated by triangles. 320 FLORA MALESIANA [ser. ILavolre? KEY TO THE SPECIES! 1. Fruit for the greater part adnate to the cupule, i.e. the scar occupying the greater part of its surface. 2. Cupule variously lamellate. 3. Leaves with an areolate reticulation; nerves obscure on both sides . . ...... 41. L. javensis 3. Leaves with scalariform or subscalariform reticulation; nerves prominent beneath. 4. Cupule and fruit as broad as or broader than long. 5. Leaves (15—)18—22(—30) by (6—)8-10 (-13) cm; nerves 10-16 pairs. 6. Cupule 5—7 cm long and wide, smooth but set with spiral or rarely concentric rings. 2. L. hallieri 6. Cupule 2—3 cm long, 3-44% cm 9; surface see ridged with the mses recurved or folded to- wards the base . . ... .. J3.L. porecatus 5. Leaves (6—)9-12(—15) by (2- )4-6(— 8) c cm; nerves a fewer than 10 pairs. 7. Cupule 3-4% cm long, 5-6 cm g, apical opening up to 3 cm g; fruit wall woody, %-1% cm thick, ruminate: ss) e .... . 4. L. ruminatus 7. Cupule 2-3% cm long and wide, completely enclosing ‘the fruit or with an apical opening of 3mm g. 8. Cupule sessile; lamellae 7-10, denticulate. Fruit wall thinner than 1 mm. Leaves thick-coria- ceous; petiole 1-2 cm .. . . 5.1L. hendersonianus 8. Cupule Y%—1 cm stalked; lamellae cle 4, entire. Fruit wall woody, 2- 5 mm thick. Leaf thin- coriaceous; ‘petiole’ 24=1 tem) s4 2°50 Fae NS we ee 22 SG lebomialtn 4. Cupule and fruit longer than wide. 9. Cupule with a truncate top. Leaf thick-coriaceous, rigid . . ....... . 7.4L. turbinatus 9. Cupule umbonate at the top. Leaf coriaceous but not rigid. 10. Cupule lengthwise ridged, ape when young. Leaves (7—)10-12(-16) by 3-6 cm; petiole 1-2 cm; nerves 5-9 pairs . . . . 8. L. beccarianus 10. Cupule not lengthwise ridged, even “when young. Leaves s (13 )17-20(-2) by (5—)6—8(-10) cm; petiole 2-3 cm; nerves (13—)16—-20(-22) pairs. . . . . . 9. L. maingayi 2. Cupule tuberculate or muricate. . Cupule obovoid, pear- -shaped, the lower 73 part smooth or eee ridged, the remainder tuber- culate. Leaf thin-coriaceous; nerves 9— 11 PalnSmaee: ... . 10. L. rotundatus 11. Cupule obconical, surface covered all over with short thick fuibercles or with recurved, hook-shaped thick scales. Leaf thick-coriaceous, rigid; nerves (9—)11—18(—22) pairs. 12. Leaves densely greyish tomentose beneath; midrib and nerves thin on both sides; nerves 9-13 pairs; reticulation obscure on both sides. Cupule set with hook-shaped, thick scales 2-5 mm long. 11. L. echinifer 12. Leaves densely yellowish to rufous tomentose; midrib and nerves thick, strongly prominent beneath; nerves 12-22 pairs; reticulation distinct beneath. Cupule tuberculate. . 12. L. pulcher 1. Fruit for the greater part free from the cupule, though sometimes enclosed by it, i.e. scar relatively small. 13. Greater part of the fruit enclosed by the cupule, although not adnate to its wall. 14. Cupule densely set with soft or stiff, spine-like appendages of 2-15 mm long. 15. Branchlets glabrous. Appendages of the cupule soft, 1-1% cm long. Styles 2-2¥% mm long, 4-5 times as long as the perianth . . . . 13. L. longispinus 15. Branchlets densely pubescent. Appendages of the cupule rigid, shorter ‘than 5 mm. Styles 1-14 mm long, 2-3 times as long as the perianth. 16. Branchlets and undersurface of the leaves densely pubescent by stiff simple hairs. Nerves more than 15 pairs. Fruit broader than long . . . . « » « 4ST wrayi 16. Branchlets and undersurface of the leaves densely pubescent by woolly, stellate hairs. Nerves less than 15 pairs. Fruit as long as orlonger than broad. . ..... . . . 15. L. coopertus 14. Cupule lamellate, squamose, tuberculate, to almost smooth. 17. Cupule tuberculate or squamose with the scales adpressed and irregularly set; in L. kostermansii there are in addition 3—4 thin lamellae in the basal part. 18. Cupule tuberculate; wall up to2 mm thick. ............. 16. L. confragosus 18. Cupule squamose; wall thinner than 1 mm. 19. Fruit as long as or longer than broad. Leaves with more than 12 nerves; reticulation dense, scalariform, distinct beneath ..... . . . . 17. L. neorobinsonii 19. Fruit broader than long. Leaves with less than 12 pairs of nerves; reticulation lax, subscalari- form, obscure on both sides .........2.2.2.2. 2.4... + 18. L. kostermansii 1 The term ‘tomentose’ is used for any indument which is continuous interwoven and does not show the parenchyma beneath, irrespective of its thickness and the size of the hairs; the latter may be of mi- croscopical dimension and the tomentum very thin. 1972] FAGACEAE (Soepadmo) 321 17. Cupule lamellate, or squamose with the scales set in concentric lines to almost smooth. 20. Fruit densely fulvous to silvery tomentose; top not elongate-acuminate. 21. Leaves thick-coriaceous, rigid; petiole 2-4 mm thick. eo sessile. Fruit densely rufous tomentose os . .. . 19. L. pattaniensis 21. Leaves coriaceous but not rigid; petiole thinner than 2 mm. Cupule 1%4,—2 % cm stalked. Fruit densely yellowish to silvery tomentose. 22. Leaves 8-15 by 4-6 cm; nerves less than 12 pairs. Cupule 1-2 % cm stalked. 23. Cupule with a rounded top, outside with distinct longitudinal streaks. Leaves dark greyish brown; petiole c. 1 cm; nerves ket ieee near the margin; reticulation lax, subscalariform toitresular ~~. 575 = . . 20. L. encleisacarpus 23. Cupule with an acute top, ‘smooth or with obscure longitudinal streaks. Leaves greenish to chocolate-brown; petiole 114-2 fa cm; nerves not anastomosing near the margin; reticula- tion dense, scalariform . . . . os 3, 6 2, ena ae 22. Leaves 15—22 by 6—8 cm; nerves more ‘than 12 pairs. “Cupule subsessile . . 22. L. macphailii 20yEruteclabrous, topielongate-acuminate .....:..=...+..... 2d A. Dinmeanns 13. Greater part of the fruit exserted from the cupule. 24. Cupule and fruit larger than 314% cm o. 25. Cupule variously lamellate. 26. Lamellae prominent. 27. Cupule %-2'% cm stalked. 28. Reticulation areolate. Fruit glabrous. 29. Branchlet yellowish grey. Reticulation obscure on both sides.Lamellae strongly prominent. Fruit with a depressed-umbonate top. . . . . . 24.L. platycarpus 29. Branchlet blackish brown. Reticulation distinct ‘beneath. ‘Lamellae not strongly prominent. Fruit witharoundedtop. ... bt See teeth oi 2S 1L. perakensis 28. Reticulation scalariform. Fruit densely tomentose eee ae Pes Oe? 26516. sericobalanus 27. Cupule sessile. 30. Reticulation obscure, areolate or irregular. 31. Leaves glabrous; top not acuminate; nerves thin, obscure on both sides; petiole 2-5 mm long. Cupule woody, lamellae strongly prominent. Fruit glabrous, wall 3-5 m thick. 27. L. lucidus 31. Leaves tomentose beneath; top 1-144 cm acuminate; nerves prominent beneath; petiole 10-15 mm long. Cupule and its lamellae thin. Fruit densely tomentose, wall c. 1 mm thick. 28. L. eichleri 30. Reticulation scalariform, distinct beneath ............ . 29. L. korthalsii 26. Lamellae obscure. 32. Scar of the fruit deeply concave, conical in longitudinal section. Rim of the cupule recurved. 30. L. urceolaris 32. Scar of the fruit flat to convex. Rim of the cupule not recurved. 33. Fruit longer than broad. Rim of the cupule thinner than 44mm... . . . 31. L. indutus 33. Fruit broader than long. Rim of the cupule thicker than 1 mm. 34. Leaves (18—)20—25(—30) by (6—)7—9(—12) cm; nerves 14-20 pairs, strongly prominent beneath. Fruit with a depressed-umbonate top; scar strongly convex. . . . . 32. L. cyclophorus 34. Leaves (7—-)9-12(-15) by (2%4-)3 een cm; nerves 9-12 pairs, thin on both sides. Fruit with an acute or rounded top; scar flat. . ; en Ooh ie. lates 25. Cupule variously muricate or squamose. 35. Cupule obconical, 3-3 4% cm high, 44-54% cma, Corea 2, part of the fruit; rim thin, incurved over the rounded top of the fruit. . .. oo ae se ged 35. Cupule broadly saucer-shaped or cup- -shaped, 1-24, cm “high, 4-5 Vy ¢ cm g, covering 144-4 part of the fruit. 36. Leaves thick-coriaceous, rigid, concolorous. Scar of the fruit concave to slightly convex at the centre. 37. Petiole 6-10 mm, 2—2'% mm g. Leaf-margin not strongly recurved; top acute to 4-1 cm acuminate. Cupule with a thick rounded rim; scales concentrically set. Scar of the fruit 242-4 cm @, concave but convex at the centre . . . . ... . 35. L. megacarpus 37. Petiole 5-6 mm, 3-4 mm 9. Leaf margin strongly recurved; top rounded-emarginate or bluntly acute. Cupule with a thin rim; scales imbricate. Scar of the fruit 2-214 cm g, concave. 36. L. revolutus 36. Leaves thin-coriaceous, not rigid, discolorous. Scar of the fruit strongly convex to conical. 38. Cupule 14-24% cm high, 44% cm 9g; scales adpressed. Fruit 244-3 cm long, 34cm g; wall 5—7 mm thick. Nerves 7-10 pairs. 39. Cupule with a thick rounded rim; scales geen obscure. Leaves with an acute top; re- ticulation obscure SVS eS 2 S71 brassii 39. Cupule with a thin rim; scree weedy: fhick) dauaee ree V1 cm acuminate; reticula- Lien GIStinee- OCNcath s 2. 4 a0 = ok ee eae ak Se i es 2 °SS. dee terbacn 322 FLORA MALESIANA [ser. I, vol. 7? 38. Cupule 44-1 cm high, 344-5 cm g; scales patent, woody, conical, 2-3 mm long. Fruit 1-2 cm long, 3-4'4% cm 9; wall thinner than 3 mm. Nerves 11-16 pairs ... . . 39. L. pallidus 24. Cupule and fruit smaller than 314 cm @. 40. Cupule variously lamellate. 41. Branchlet and leaf densely tomentose by adpressed stellate hairs and erect simple or tuft-hairs. 42. Nerves anastomosing near the margin; reticulation coarse, distinct beneath. 43. Fruit densely greyish tomentose by adpressed simple hairs, top long-acuminate. Leaf-base acute; indumentum yellowish brown torufous ... . . . . . 40. L. conocarpus 43. Fruit densely fulvous tomentose by simple and stellate hairs, without a long-acuminate top. Leaves with a rounded to subcordate base; indumentum greyish brown . 41. L. atjehensis 42. Nerves not anastomosing near the margin; reticulation fine, obscure on both sides. 44. Leaf thick-coriaceous, rigid; nerves 12-14 pairs. Scar of the fruit deeply concave. 42. L. dasystachyus 44. Leaf papery to thin-coriaceous; nerves 8-10 pairs. Scar of the fruit flat to convex. 43. L. caudatifolius 41. Branchlet and leaf densely tomentose by adpressed stellate hairs to glabrous. 45. Rim of the cupule thinner than % mm. 46. Leaves with regular, scalariform or subscalariform reticulation. 47. Fruit variously tomentose. 48. Scar of the fruit deeply concave. Nerves (11—)12—15(-16) pairs. 49. Leaves thin-coriaceous; nerves and reticulation dense. . ...... . 44. L. ewyckii 49. Leaves thick-coriaceous, rigid; nerves and reticulation lax. 50. Cupule 4% cm stalked. Nerves (12—)14-15(—16) pairs; reticulation distinct beneath. 45. L. cantleyanus 50. Cupule sessile. Nerves 11-13 pairs; reticulation obscure. . . ... . 46. L. meijeri 48. Scar of the fruit flat or convex. Nerves (7—)8—10(-12) pairs. 51. Leaves 7-13 by 3-5 cm, index (174—)2-2 %4(—3 14), broadest at or below the middle. 52. Cupule obconical; lamellae 8-10, entire. Nerves not anastomosing near the margin. 47. L. daphnoideus 52. Cupule cup-shaped; lamellae 5-8, denticulate. Nerves anastomosing near the margin. 48. L. philippinensis 51. Leaves 12-17 by 5-8 cm, index (123—)2(-3), broadest about the middle. 53. Leaf thick-coriaceous, rigid; midrib prominent on both sides; nerves flattish above. Cupule 4% cm stalked, all over with distinct lamellae. Fruit as long as or longer than broads .... . 49. L. apoensis 53. Leaves thin-coriaceous, not rigid: aides flattish above: nerves impressed above. Cupule sessile, lower part ridged or tuberculate, upper part with 5—7 obscure lamellae. Fruit broadenthanvone ee see er rene a n ss ete 8 eee pe , SOR oIOTMOSHS 47. Fruit glabrous. 54. Leaf thin-coriaceous. 55. Cupule 44-1 cm high, 2-214 cm @. Scar of the fruit flat. 56. Indumentum of the branchlet and leaf greyish brown. Cupule cup-shaped, %-—1 cm high, covering 4-1, part of the fruit. ... . : . . 51. L. solerianus 56. Indumentum yellowish brown. Cupule saucer-shaped, Yo Ye cm ‘high, covering the basal part of the fruit .. : . . 52. L. mindanaensis 55. Cupule 4% cm high, 1-1 Vy ¢ cm @. “Scar of the fruit concave. 57. Reticulation obscure. Fruit longer than broad, top rounded-acute . . 53. L. bennettii 57. Reticulation distinct beneath. Fruit broader than long, top abruptly acuminate. 54. L. confertus 54. Leaf thick-coriaceous, rigid. 58. Leaf strongly bullate; petiole shorter than5mm ......... . 55.L. bullatus 58. Leaf not bullate; petiole 7-15 mm long. 59. Leaf sparsely pubescent; base long decurrent down to the base of the petiole; margin strongly recurved; petioleS5-6 mmg... . .... . 56. L. rigidus 59. Leaf densely tomentose beneath; base short decurrent: margin not strongly recurved; petiole thinner than 3 mm. 60. Fruit broader than long. Leaf index 1.6—2, blade broadest at or below the middle. 61. Reticulation obscure. Cupule and fruit usually smaller than 2 cm 9; lamellae 4-6. 57. L. nodosus . Reticulation distinct beneath. Cupule and fruit usually ee than 2 cm g; lamellae A—VO0r 50s 5 .. . 58. L. woodii 60. Fruit longer than broad. Leaf index 2- 2, blade broadest at or above the middle. 62. Leaf with a thin cover of pale yellowish brown tomentum beneath; nerves not anas- tomosing near the margin. Cupule 44 cm high, 1.2—1.7 cm g. Scar of. the fruit concave. 59. L. hatusimae 1972] FAGACEAE (Soepadmo) 323 62. Leaf with a thick cover of fulvous to rufous tomentum beneath; nerves anastomosing near the margin. Cupule 1 cm high, 2.2 cm g. Scar of the fruit flat. . 60. L. vidalii 46. Leaf with an areolate or irregular reticulation. 63. Fruit tomentose. 64. Nerves with shorter ones not reaching the margin in between . . . . 61. L. reinwardtii 64. Leaves without extra shorter nerves. 65. Leaf thin- eS ain ee )9-11(-13) by (24%—)4—5(-6) cm, index 2-21%, broadest about the middle . . eh.) 202.1, Snkitrs 65. Leaves thick-coriaceous, (9- )12- 20(- 23) by (4-)5- 8(- 10), index 12 2p. broadest about the middle. 66. Leaf discolorous; petiole 5-12 mm long; nerves anastomosing near the margin. 63. L. bancanus 66. Leaf concolorous; petiole 15-25 mm long; nerves not anastomosing near the margin. 64. L. ovalis 63. Fruit glabrous. 67.) Leaf orbicular; petiole 3)mm long, 2immigi 2. 2 ce) G0...) 65. LE. orbicularis 67. Leaf not orbicular; petiole 5-20 mm long, 1-144 mm @. GserCupule: icmihigh; 2, cmon fie ses fi Genesee Ges Gi 20. (66: 1: pusillus 68. Cupule %-1 cm high, 144-274 cm g@. 69 saLeat c: 1S—20'by 5—8:cms nerves: (14—15' pairs’. . 5... = 5 « = 2 67. 1. eracilis 69. Leaf 5—13 by 2-4 cm; nerves less than 14 pairs. 70. Cupule with 6-10, prominent lamellae. Fruit broader than long. Petiole up to 2 cm. 68. L. rassa 70. Cupule with 5-6 thin lamellae. Fruit as long as or longer than broad. Petiole shorter than 1 cm. 71. Midrib strongly prominent on both sides; nerves Pe aus near the margin. Fruit dark purplish brown ... . .. . . 69. L. andersonii 71. Midrib thin on both sides; nerves ‘not anastomosing near the margin. Fruit pale cho- colate-brown , alah SUL ee ei ea Boris © Po! OAK. Vink 45. Rim of the cupule thicker than 1 mm. 72. Leaf elliptic-lanceolate. Cupule sessile, sometimes in clusters of ra 1-1 4 cm high, 2-244 cm @. Fruit broader than long, top rounded . . . . . . 71. L. clementianus 72. Leaf ovate-elliptic. Cupule 44—'% cm stalked, hd cm n high, oe 2Y, 56 cm 9g, always solitary. Fruit as long as or longer than broad, top acute .. . . .. . 72. L. suffruticosus 40. Cupule squamose to muricate. 73. Cupule variously squamose. 74. Cupules, even when ripe, in clusters of 3-10. 75. Leaf and fruit glabrous. 76. Leaf pale brown, (9—)12-17(-20) by (3—)4-6(-8) cm, index (1 44—)2-3(—5), base acute or rounded. Cupules in clusters of (3—)5—7(—10), scales distinct .. Hiya Paso: elegans 76. Leaf greyish green, (18—)2440(—56) by (5—)9-11(— ee cm, index 354: base cordate to auri- culate. Cupules in clusters of 3, scales obscure .. . : hokey 74. Jacopsil 75. Leaf and fruit tomentose. 77. Fruit broader than long, densely yellowish brown tomentose. Leaf with a distinct, scalari- form reticulation. 78. Leaf densely glaucous to greyish brown tomentose by adpressed stellate hairs beneath. Branchlet and petiole sturdy. Cupule larger than 2 cm g. Scar of the fruit up to 2 cm g. 75. L. lampadarius 78. Leaf densely yellowish brown tomentose by adpressed stellate hairs and erect tuft-hairs beneath. Branchlet and petiole slender. Cupule smaller than 1144 cm g. Scar of the fruit smallerthanicmg... . . . 76. L. wallichianus 77. Fruit longer than broad, densely rufous tomentose. Teat with | an irregular, obscure reticu- lation? yates meee ls Waele? 2aat Ils bela hoe ea ery throcarpus 74. Cupules solitary, rarely i in clusters of a 79. Leaf with a rounded or obtuse top. 80. Leaf 2%-5 by 144-4 cm, index (1—)1%(-2); base rounded to subcordate; reticulation areo- late; petiole 2-3 mm long, 1-14 mmg. Cupulec.24%cmg@. .... . 78. L. oreophilus 80. Leaf 6-13 by 3-7 cm, index 114-2 %; base acute; reticulation scalariform or subscalariform; petiole S—13 mm long, 1-5 mm g. Cupule smaller than 1% cm g. 81. Leaf without glandular papillae, thick-coriaceous, rigid, margin strongly recurved; nerves flattish on both sides, anastomosing near the ur Indumentum consisting of adpressed and erect stellate hairs sens ... . . 79. L. obtusifolius 81. Leaf with glandular papillae above, not rigid coriaceous; - margin not strongly recurved ; nerves prominent beneath, impressed above, not anastomosing near the margin. Indumen- tum consisting exclusively of adpressed stellate hairs ...... . . 80. L. papillifer 324 FLORA MALESIANA [ser. I, vol. 7? 79. Leaf with an acute or acuminate top. 82. Leaf concolorous, glabrous, rarely sparsely pubescent. 83. Fruit densely reddish brown tomentose, as broad as or broader than long. 81. L. ferrugineus 83. Fruit glabrous, longer than broad. 84. Petiole (8—)10-15(-20) mm long. 85. Bud-scales densely silvery tomentose by simple hairs. Nerves (12—)14-17(-20) pairs, anastomosing near the margin . . . . . . 82. L. falconeri 85. Bud-scales densely fulvous tomentose by stellate hairs. Nerves (6-)9-10(-12) pairs, not anastomosing near the margin .............. . 93.1. crassimeryius 84. Petiole 2-6 mm long. 86. Leaf sparsely pubescent by stellate hairs on both sides; base rounded to subcordate; nerves 10-13 pairs, anastomosing near the margin. ees . . . 84. L. kunstleri 86. Leaf glabrous; base attenuate-acute; nerves 7—8 pairs, not anastomosing near the margin. 85. L. pseudokunstleri 82. Leaf discolorous, densely tomentose at least beneath. 87. Fruit glabrous. 88. Branchlet and undersurface of the leaf with a thick cover of yellowish brown to rufous, adpressed, stellate hairs and erect, glandular tuft-hairs with bulbous base. 86. L. havilandii 88. Branchlet and undersurface of the leaf with a thin cover of greyish to fulvous adpressed and erect, non-glandular, stellate hairs without bulbous base. 89. Scar of the fruit conical. Cupule with obscure lamellae on the lower half. 87. L. sogerensis 89. Scar of the fruit flat to concave. Cupule all-over with distinct scales set in concentric lines or imbricate. 90. Fruit longer than broad; scar deeply concave . . . . .. . . .88. L. celebicus 90. Fruit broader than long; scar flat to shallowly concave. 91. Indumentum on the branchlet and leaf consisting of adpressed, stellate hairs and curled or erect tuft-hairs. Nerves (10—)12—14(-16) pairs; reticulation scalariform. 89. L. sundaicus 91. Indumentum of the branchlet and leaf consisting exclusively of adpressed, stellate hairs. Nerves 10-12 pairs; reticulation irregular to subscalariform . 90. L. pseudomoluccus 87. Fruit variously tomentose. 92. Leaf smaller than 10 by 4 cm; nerves 6-8 pairs. . . . .. . . 91. L. submonticolus 92. Leaf larger than 10 by 4 cm; nerves more than 9 pairs. 93. Cupule 3-10 mm stalked; rim never undulate. 94. Fruit longer than broad; scar Beg convex. Petiole e )10(-11) mm long. Older branchlet glabrous. . : . . 92. L. castellarnauianus 94. Fruit as broad as or broader ‘than long; s scar ar deeply « concave, rarely flat. Petiole 3-6 mm long. Branchlet densely tomentose. . . , Bees de eee SAE Meptogyne 93. Cupule sessile, rarely subsessile, rim often undulate. 95. Indumentum of the branchlet and leaf consisting of adpressed, stellate hairs and erect long simple hairs. Nerves strongly prominent beneath, anastomosing near the margin. 94. L. nieuwenhuisii 95. Indumentum of the branchlet and leaf consisting exclusively of adpressed stellate hairs. Nerves not strongly prominent beneath and not anastomosing near the margin. 95. L. curtisii 73. Cupule variously muricate. 96. Leaf glabrous. 97. Leaf thick-coriaceous, rigid; nerves more than 8 pairs; petiole 10-13 mm long. Fruit longer than broad, glabrous, scar deeply concave .. . .. . . 96. L. scortechinii 97. Leaf coriaceous but not rigid; nerves less than 8 pairs, ‘usually 6-7; petiole 4-8 mm long. Fruit as broad as or broader than long, densely fulvous tomentose, scarflat. . . 97. L. kingianus 96. Leaf variously tomentose or pubescent. 98. Leaf concolorous, greenish grey, beneath with a thin cover of stellate tomentum. 99. Leaf (3 4—)5-6(-8) by (1 %-)2%-3(-3%) cm, index 2-2.7; petiole 4-6 mm long. Cupule cup-shaped, scales concentrically set. Fruit longer than broad, scar c. 6 mm 9. 98. L. luzoniensis 99. Leaf (9—)13-18(-21) by (3—)4-6(-7) cm, index 3-3%; petiole 8-15 mm long. Cupule flat saucer-shaped, scales irregularly set. Fruit broader than long, scar up to 1144 cm @. 99. L. echinulatus 98. Leaf discolorous, not greenish grey, indumentum thick, consisting of adpressed, stellate hairs and erect or curled, simple or tuft-hairs. 100. Leaf 6-10 by 4-5 cm, index 1.3—2; indumentum yellowish brown. 1972] FAGACEAE (Soepadmo) 325 101. Leaf thick-coriaceous, rigid, margin strongly recurved; reticulation fine, dense, scalariform; indumentum velvety. Fruit longer than broad. 100. L. jordanae 101. Leaf coriaceous but not rigid, margin not strongly recurved; ‘reticulation coarse, lax sub- scalariform, arched. Fruit as broad as or broader than long . 101. L. aspericupula 100. Leaf 10-20 by 31%4—7 cm, index 2—3.3; indumentum fulvous to rufous. 102. Fruit and cupule smaller than 2 cm g. Nerves less than 9 pairs 102. L. buddii 102. Fruit and cupule 2-3 1% cm g. Nerves more than 9 pairs, usually 10-14. 103. Indumentum consists of adpressed stellate hairs and curled stellate or tuft-hairs. Nerves less than 12 pairs, usually about 10; reticulation lax. Fruit chocolate-brown, top rounded. 103. L. rufovillosus 103. Indumentum consists of adpressed stellate hairs and erect stellate or tuft-hairs. Nerves 12-16 pairs; reticulation dense. Fruit dark purplish brown, top with a long acuminate end, especially when young 1. Lithocarpus javensis BL. Bijdr. (1826) 527; FI. Jav. Cupul. (1829) 35, t. 20; OuDEM. Natuurk. Verh. Kon. Akad. 11 (1865) 19, t. 11; A. Camus, Chénes 3 (1954) 572, t. 351: 1-20; SoEPADMo, Reinwardtia 8 (1970) 249. — Quercus costata BL. Bijdr. (1826) 522; Fl. Jav. Cupul. (1829) 25, t. 13, incl. var. B, l.c. 26, t. 14; KorTH. Kruidk. (1844) 212; Bt. Mus. Bot. 1 (1850) 301, incl. Fig. 17. Species density of Lithocarpus in Malesia, above the hyphen the number of non-endemic species, below the hyphen the number of endemic species in the island or island group. var. convexa et var. subrecurvata BL. l.c. 302; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 116; A.DC. Prod. 16, 2 (1864) 93, incl. var. convexa BL. /.c. 94; WENzIG, Jahrb. Bot. Gart. Berl. 4 (1886) 239; K. & V. Bijdr. 10 (1904) 60; Koorpb. Atlas 1 (1913) t. 46; BACKER & BAKH. f. Fl. Java 2 (1965) 6. — Quercus javensis (BL.) Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 117; A.DC. Prod. 16, 2 (1864) 104; Wenzic, Jahrb. Bot. Gart. Berl. 4 (1886) 238; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 83, t. 76 B; K. & V. Bijdr. 10 (1904) 58; Koorp. Atlas 1 (1913) t. 48; BACKER & Baku. f. Fl. Java 2 (1965) 6. — L. scutigera OupDEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 207; Natuurk. Verh. Kon. Akad. 11 (1865) 20, t. 12. — Cyclobalanus javensis (BL.) Okrrst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Cyclobalanus costata (BL.) OERST. l.c. — Pasania javensis (M1Q.) PRANTL in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 55. — Synaedrys javensis (BL.) Korpz. Bot. Mag. Tokyo 30 (1916) 189. — Synaedrys costata (BL.) Korpz. lc. 188. — L. costata (BL.) REHD. J. Arn. Arb. 1 (1919) 124; 104. L. hystrix A. Camus, Chénes 3 (1954) 580, t., incl. var. convexa (BL.) A. CAMuS, l.c. 582, t. 353: 1-9. — L. costata var. scutigera (OUDEM.) A. CAMUS, Bull. Soc. Bot. Fr. 92 (1945) 254; Chénes 3 (1954) 582, t. 354: 1-5. — L. costata var. typica A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 254; Chénes 3 (1954) 582, t. 353: 10-15, t. 354: 6-8. — Fig. 18. Tree 20—50 m, 30-100 cm g; buttresses up to 114 m tall; bark grey, shallowly fissured. Branch- lets initially with a dense reddish brown indu- ment, later glabrous, pale to greyish brown, sparsely lenticellate; terminal bud ovoid-—ellip- soid, 3-5 by 2-3 mm, scales narrowly ovate to linear. Stipules narrowly ovate or subulate, 5-9 by 1-2 mm, rather long persistent. Leaves thick- coriaceous, (5—)8-12(-16) by (2-)3-4(-6) cm (index 2-3), broadest at or slightly below the mid- dle; surfaces more or Jess concolorous, above glab- rous, dull, beneath densely greyish tomentose by adpressed, stellate hairs, glabrescent; base acute to cuneate, rarely rounded, margin entire, re- curved, top bluntly acute to %-1 cm acumi- nate; midrib prominent beneath, slightly so above; nerves 10-14 pairs, obscure on both sides, subparallel, at an angle of 45—60°, arcuating and anastomosing towards the margin; reticulation fine, areolate, distinct beneath; petiole 14-114 cm, 1-2 mm g, adaxially flat. Inflorescence male, androgynous, or mixed, densely fulvous-tomen- tose by stellate hairs; bracts linear-acute, 1-3 by 14-1 mm, bracteoles ovate-acute, 1-114 by 75-1 mm. Male rachis 10-20 cm, 1-2 mm g; ¢ flowers in clusters of 3—7, perianth 6-lobed, stamens 12-16, filaments 3-4 mm, anthers 0.3—-0.4 mm long, pistil- lode subglobose, 1-144 mm g. Androgynous or mixed rachis 5-10 cm, i14%-2 mm g; female flowers solitary, sometimes in clusters of 2- ae perianth 6-lobed, staminodes 12, rudimentary, styles 3, conical, 1-144 mm, connate. Young cupule solitary or in clusters of 2-3, obovoid to ellipsoid, 1-244 cm stalked; outside with 5-8 more or less concentric, prominent lamellae, densely greyish brown stellate-tomentose, apical opening 4-114 cm o. Ripe cupule sessile or 1-1% cm stalked, ellipsoid-globose or obovoid, 3—5 cm long, 344-5144 cm @, covering the fruit almost completely or with an apical opening of 44-3 cm 0; wall woody, 2—3 mm thick, outside with 5-10 thin to strongly prominent lamellae, densely greyish tomentose by simple and stellate hairs. Ripe fruit 326 FLORA MALESIANA [ser. I, vol. 7? Mia 20 Fig. 18. Lithocarpus javensis BL. (pasang batu) in the West Javanese mountain forest of Priangan Prov. (J. VAN ROOSENDAAL, Dec. 1919). 1972] FAGACEAE (Soepadmo) 327 depressed ovoid-globose, 3-4 cm long, 314-5 cm g; wall woody, 14-1 cm thick, for the greater part adnate to the cupule, upper free part flat to convex, %-3'% cm g, densely greyish brown to- mentose, glabrescent; scar rounded, rugose; cotyledons flat-convex. Distr. Malesia: Sumatra (various localities), also Singkep I., Malay Peninsula (rare), Java (rather common in the western parts, eastwards to Mt Ungaran in Central Java). Ecol. Forests up to 1800 m. Fl. July-Sept., fr. Oct.-Aug. Notes. In the previous works L. costata was distinguished from L. javensis by its cupule with apical aperture wider than 44 cm. In JUNGHUHN .n. from Mt Malabar (described by OUDEMANS as L. scutigera) and JAcoss 4564 (from Mt Ker- intji, Central W. Sumatra), however, the cupule is intermediate between that of L. javensis and L. costata. Furthermore the leaves of these three species are exactly the same. In 1826, and also in 1829, BLUME recognized two varieties under Quercus costata, differing from each other by the fruit. In the typical variety the upper free part of the fruit is flat to concave, while in the other it is convex. In 1850, BLUME named the latter variety var. convexa. The type specimens at Leyden show that no sharp boundary between these two vari- eties can be drawn. In 1888, Hooker f. attributed several specimens from Malaya to Quercus costata, apparently without examining the original speci- men of BLUME. KING (1889) followed Hooker, and on account of the convex fruit included the Malayan specimens in var. convexa. GAMBLE (1914) working on the Malayan specimens only, included Quercus costata in the genus Pasania, and placed them in the subgenus Cyclobalanus. In 1954, A. Camus disagreed with HOoKER, KING, and GAMBLE, and transferred Quercus costata to the genus Lithocarpus, recognizing four distinct varieties, viz. var. typica, convexa, scutigera (all from Java) and var. kingii for the Malayan spe- cimens. I consider L. costata (BL.) REHD. var. kingii A. CAMUS as a separate species, differing from L. javensis (incl. L. costata and its varieties, and L. scutigera) by its cupule enclosing the lower part of the fruit only (see 25. L. perakensis). 2. Lithocarpus hallieri (VON SEEMEN) A. CAMUS, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 579, t. 352; SOEPADMo, Reinwardtia 8 (1970) 244. — Quercus hallierii VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 11; Merr. Univ. Calif. Publ. Bot. 15 (1929) 44. — Synaedrys hallierii (von SEEMEN) Korpz. Bot. Mag. Tokyo 30 (1916) 189. Tree 12-48 m, 10-70 cm g; bark smooth to flaky, lenticellate, greyish brown. Branchlets ini- tially densely reddish brown adpressed-stellate- hairy, later glabrescent, terete, dark coloured, smooth to shallowly fissured; terminal buds ovoid- ellipsoid, c. 8 by 3 mm, scales ovate to lanceolate. Leaves thick-coriaceous, (15—)16—-20(-30) by (6-) 8-10(-11) cm (index 2-3), broadest above or at the middle; surfaces more or less concolorous, above glabrous, dull to glossy, underneath spar- sely to densely greyish tomentose by stellate hairs; base rounded to cuneate, decurrent, top bluntly acute to 1 cm acuminate; midrib prominent beneath, slightly so above; nerves (8—)10—11(—13) pairs, parallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin, prom- inent beneath, impressed above; reticulation subscalariform, obscure on both sides; petiole 1-3 cm, 2-3 mm @g, adaxially flat. Male inflores- cence not known. Female rachis c. 15 cm, 2 mm 6; bracts and bracteoles ovate, c. 1-2 by 1 mm; 2 flowers solitary, perianth 6-lobed, staminodes 12, rudimentary, styles 3, conical, c. 2 mm, more or less connate. Young infructescence 15-20 cm, 4-5 mm a, densely tomentose by stellate hairs, carrying 15-20 young cupules. Ripe cupule ses- sile to 44-1 cm stalked, obovoid-globose, 5-7 cm through; wall woody, 2-3 mm thick, outside den- sely greyish tomentose, with 5-7 spiral or rarely concentric lamellae; base rounded-attenuate, top rounded with a small opening of 5-7 mm g. Ripe fruit subglobose, 3-414 cm long, 4-6 cm g; wall woody, 44—1% cm thick, for the greater part adnate to the cupule; free part (top) truncate- umbonate, densely yellowish brown tomentose, base rounded. Distr. Malesia: Borneo (scattered in the western part of Kalimantan, Sarawak, and Sabah). Ecol. Forests up to 1350 m, on yellowish sandy soil. Fr. Oct.—July. 3. Lithocarpus porcatus SOEPADMO, Reinwardtia 8 (1970) 268. Tree 10-45 m, 20-50 cm g; bark smooth to flaky. Branchlets initially densely fulvous, ad- pressed stellate-hairy, later subglabrous, warty lenticellate; terminal buds ovoid, 3 by 2 mm, scales ovate-acute. Leaves thick-coriaceous, (15—)20-22 (—26) by (7-)9-10(—13) cm (index 1.8-2.5), broad- est at or above the middle; surfaces more or less discolorous, above glabrous, dark choco- late-brown, glossy, underneath densely greyish tomentose by adpressed stellate hairs, subglabres- cent; base abruptly rounded-acute to cuneate, margin undulate, top bluntly acute; midrib and nerves strongly prominent beneath, flat above; nerves (12—)13-14(-16) pairs, parallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, dense, sca- lariform, obscure on both sides; petiole %-1%4 cm, 2 mm g, rugose and thickened at base, adax- ially flat. Inflorescence male or female, densely yellowish brown tomentose by adpressed stellate hairs; bracts and bracteoles ovate to linear, 23-114 by 4-24 mm. Male rachis 10-30 cm, 1-2 mm 9; 3 flowers in clusters of 3, rarely solitary, perianth 6-lobed, stamens 12, filaments 2-314 mm, anthers 0.3—-0.4 mm long, pistillode subglobose, c. 1 mm 9. Female rachis 10-15 cm, 3 mm 9g; @ flowers soli- tary, perianth 6-lobed, staminodes 12, rudimen- tary, styles 3-4, terete, 24-1 mm. Infructescence 10-15 cm, 44-1 cm 9, densely warty lenticellate, carrying many young cupules. Cupule (not yet fully developed) 14-1 cm stalked, depressed-ovoid- globose, 2-3 cm long, 3-4 cm g, top flat with a 328 small opening of 1-5 mm g, base rounded; out- side with 5-10 concentric or rarely spiral promi- nent lamellae which are sometimes folded towards the base, densely yellowish brown tomentose. Fruit depressed-subglobose, 1-144 cm long, 144-2 cm 9; wall woody 1-2 mm thick, for the greater part adnate to the cupule; top flat-umbonate, densely yellowish brown tomentose, base rounded. Distr. Malesia: Borneo (SW. Sarawak: Mt Penrissen; Sabah). Ecol. Forests up to 1500 m, usually by river- banks. Fi. Sept., fr. Oct._June. 4. Lithocarpus ruminatus SOEPADMO, Reinwardtia 8 (1970) 277. Tree 10-20 m, 20-30 cm g; bark greyish brown, smooth to flaky. Branchlets initially densely ful- vous-tomentose by adpressed stellate hairs, later subglabrous, greyish, sparsely lenticellate; ter- minal buds ovoid, 2-3 by 2 mm, scales ovate. Sti- pules linear-acute, 2-3 by 14 mm, caducous. Leaves thick-coriaceous, (6—)9-12(-16) by (3—)4-S(-8) cm (index 2-3), broadest at or below the middle; surfaces more or less discolorous, above glabrous, more or less glossy, greyish green, underneath densely greyish tomentose with adpressed, stellate hairs; base rounded and abruptly acute to cuneate, margin recurved, top bluntly to sharply acute; midrib prominent beneath, slightly so above; nerves 6-8 pairs, subparallel, at an angle of 45—50°, he Fig. 19. Characteristic stilt-rooted stem-base of young Lithocarpus sp. in mountain forest at 2000 m on Mt Gedeh, West Java (W. MEER, Aug. 1956). FLORA MALESIANA [ser:. I wolh?? arcuating but not anastomosing towards the mar- gin; reticulation fine, lax, subscalariform, ob- scure on both sides; petiole 44-1 cm, 2 mm 4g, thick- ened and rugose at base, adaxially flat. Inflo- rescence male, androgynous or mixed, densely yel- lowish brown to fulvous tomentose by stellate hairs; bracts and bracteoles ovate acute, 4-1 by 1, mm. Male flowers (seen on the upper part of an androgynous rachis) solitary, perianth 6-lobed, stamens 12, filaments 244-4 mm long, anthers 0.35 mm long, pistillode subglobose, c. 2 mm g. Androgynous rachis 15-20 cm, 2 mm g; female flowers solitary, perianth 6-lobed, staminodes 12, well developed and exceeding the perianth, styles 3, conical, 73-1 mm. Ripe cupule subsessile, sub- hemispherical to depressed obovoid-globose, 3-414 cm long, 5—6 cm g, base attenuate-rounded, top truncate with an opening of 2-3 cm g; wall woody, c. 4% cm thick, outside with 8-10 concen- tric lamellae, 5-6 of which on the truncate top, densely greyish brown tomentose by short stel- late hairs, subglabrescent. Ripe fruit subhemisphe- rical to depressed-obovoid-globose, 3—4 cm long, 4-5 cm g; wall woody, ruminate, %-14%4 cm thick, for the greater part adnate to the cupule; top convex, depressed-umbonate at the centre, densely reddish brown tomentose, base rounded. Distr. Malesia: Borneo (Sabah). Ecol. Forests, from the lowland up to 2000 m. Fl. Sept., fr. March-Aug. 5. Lithocarpus hendersonianus A. CAMus, Bull. Mus. Hist. Nat. Paris II, 6 (1934) 92; Chénes 3 (1954) 589, t. 358: 9-14; SoEPADMO, Reinwardtia 8 (1970) 246. Tree 20-30 m, c. 60 cm o; bark smooth, grey. Branchlets initially densely yellowish brown to- mentose by stellate hairs, soon glabrescent, greyish black, with dense minute lenticels; terminal buds ovoid-globose, 1—2 mm, scales ovate-acute. Leaves thick-coriaceous, (8—)10—15(—18) by (24%4-)4-6(-7) cm (index 2144-3 44), broadest at or below the mid- dle; surfaces more or less concolorous, above glab- rous, glossy, underneath sparsely yellowish grey tomentose by stellate hairs; base rounded and abruptly acute to cuneate, margin recurved, top abruptly acute to sharply 1-2 cm acuminate; midrib and nerves prominent beneath, obscure and flattish above; nerves 7-10 pairs, subparallel, at an angle of 45-60°, arcuating but hardly anas- tomosing near the margin; reticulation fine, lax, subscalariform, obscure on both sides; peti- ole 1-2 cm, 1-2 mm g, adaxially flat. Inflorescence male or androgynous, densely yellowish brown tomentose by short stellate hairs; bract and bract- eoles ovate-acute 1-144 by 0.7 mm. Male rachis 5-10 cm, 1 mm g; ¢ flowers in clusters of 3-7, perianth 6-lobed, stamens 12, filaments 1-144 mm, anthers 0.2-0.25 mm long, pistillode subglobose, c. 1 mm @. Androgynous rachis 5-10 cm, 1 mm 9; female flowers in clusters of 2—5, perianth deeply 6-lobed, staminodes 12, rudimentary, styles 3, conical, c. 14% mm, recurved. Cupule (not fully developed) sessile, ellipsoid-globose, 2-3 cm through; wall thinner than 1 mm, outside densely 1972] yellowish brown tomentose, with 7-10 more or less concentric, undulating, denticulate lamellae; base rounded, top truncate, with a small opening of less than 3 mm @. Fruit (not fully developed) globose, 2-244 cm through; wall thinner than 1 mm, for the greater part adnate to the cupule; free part (top) c. 1 cm, rotundate, densely tomentose, base rounded. Distr. Peninsular Siam (Ranawang at 10° N), in Malesia: Malay Peninsula (Cameron High- lands). Ecol. Forests at 1300-1500 m. F/. July-April, fr. April.—Nov. 6. Lithocarpus burkillii A. Camus, Bull. Mus. Hist. Nat. Paris II, 5 (1933) 88; Chénes 3 (1954) 578, t.; SOEPADMO, Reinwardtia 8 (1970) 225. Tree c. 24 m. Branchlets glabrous, blackish grey, densely rusty lenticellate. Leaves thin-coria- ceous, 6-10 by 2-4 cm (index 214-3), broadest at or below the middle; surfaces discolorous, above glabrous, dark greyish brown, dull, beneath densely glaucous-tomentose by adpressed stellate hairs; base rounded-acute, margin undulate, top sharply acute to acuminate; midrib more or less prominent on both sides; nerves 8-10 pairs, par- allel, at an angle of c. 60°, arcuating but not anas- tomosing towards the margin, obscure on both sides; reticulation fine, subscalariform, obscure on both sides; petiole 44-1 cm, 1-2 mm 9, adaxial- ly flat. Inflorescence not known. Cupule Y-1 cm stalked, obovoid-globose, 244-314 cm through; wall woody, 1-2 mm thick, outside with 3-4 more or less concentric lamellae, densely greyish brown tomentose by short stellate hairs; base rounded- attenuate, top rotundate, covering the fruit almost completely except for a very small opening at the tip. Fruit depressed-subglobose, 1144-2 cm long, 2-3 cm 9; wall woody, 2-5 mm thick, for the great- er part adnate to the cupule; top %4-1 cm a, depressed-umbonate, base rounded. Distr. Malesia: Malay Peninsula (Fraser Hill). Ecol. Forests at c. 1200 m. Fr. Sept.—Nov. 7. Lithocarpus turbinatus (STAPF) FORMAN, Kew Bull. 18 (1966) 423; Sogpapmo, Reinwardtia 8 (1970) 284. — Castanopsis turbinata STAPF, Trans. Linn. Soc. Lond. II, Bot. 4 (1894) 232. — Fig. 20. Tree 5-12 m, c. 30 cm 9, usually crooked; bark peeling off profusely. Branchlets initially densely rufous-tomentose by stellate hairs, later subglab rous, sturdy with thick nodes, yellowish grey, smooth to finely fissured; terminal buds ovoid- ellipsoid, 3-6 by 3-4 mm, scales ovate-acute. Stipules linear-acute, 144-2 by 14-23 mm, soon caducous. Leaves thick-coriaceous, rigid, (6-) 8-12(-15) by (2-) 3-6-8) cm (index 1.6-2.8), broadest below or at the middle; surfaces more or less discolorous, above dull, chocolate-brown, glabrous except the midrib, beneath densely yel- lowish brown tomentose by adpressed stellate hairs; base rounded and abruptly acute to acute, margin recurved, top abruptly acute to 1-144 cm acuminate, tip sharp, usually oblique; midrib and FAGACEAE (Soepadmo) 329 nerves strongly prominent beneath, impressed above; nerves (6-)7-9(-12) pairs, subparallel, at an angle of 60-80°, arcuating and anastomosing near the margin; reticulation fine, lax, scalari- form, distinct beneath; petiole 14-14% cm, 1-4 mm 9, adaxially flat. Inflorescence male, female, androgynous or mixed, densely fulvous to rufous tomentose by stellate hairs; bracts and bracteoles ovate-acute to linear, thick-coriaceous, 114-3 by *3-1 mm. Male rachis c. 10 cm, 11%4-2 mm o; 3 flowers solitary or in clusters of 3-7, perianth 6— lobed, stamens usually 12, filaments 4-5 mm, an- thers 0.4-0.5 mm long, pistillode subglobose, 1144-2 mm @. Female, androgynous or mixed rachis 4-6 cm, 3-5 mm 9; 9 flowers solitary or in clusters of 3-7, perianth thick-coriaceous, 6-lobed, sta- minodes 12, sometimes well-developed and ex- ceeding the perianth, styles 3-5, conical, 2-3 mm, recurved. Rachis of the infructescence sturdy. Young cupule subsessile, in clusters of 3-7, out- side with acroscopical scales arranged in concen- tric lines, greyish black, densely tomentose by short stellate hairs. Ripe cupule sessile, obovoid- ellipsoid, 6-7 cm long, 5-6 cm @, base rounded, top rounded, depressed at the centre, opening less than 14 cm 9g; wall woody, greyish black, 2-5 mm thick, outside lenticellate, with 8-10 concentric, rarely spiral, entire or denticulate lamellae, mainly on the rotundate top. Fruit ellipsoid-globose, 5—6 cm long, 4-5 cm 9; wall woody, 1-2 cm thick, for the greater part adnate to the cupule; top c. 2 cm 9, densely tomentose, base rounded. Distr. Malesia: Borneo (N. Sarawak, and se- veral localities in Sabah, especially on Mt Kina- balu). Ecol. In mossy forest at 1200-3000 m, usually on steep ridges. F/. Febr.—Oct.. fr. Noy.—July. Note. Though the young cupules are in clus- ters of 3-7, at maturity only one or two are fully developed, the others remain abortive and some- times may be seen at the base of the well-devel- oped cupule(s). 8. Lithocarpus beccarianus (BENTH.) A. CAMUS, Riviera Scient. 18 (1932) 39 (‘beccariana’); Chénes 3 (1954) 575, t. 355: 1-7; So—EPADMO, Reinwardtia 8 (1970) 220. — Quercus beccariana BENTH. in Hook. f. Ic. Pl. (1880) t. 1315; Hook. f. Fl. Br. Ind. 5 (1888) 618; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 83.— Pasania beccariana (BENTH.) PRANTL in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 55. — Synae- drys beccariana (BENTH.) Ko1pz. Bot. Mag. Tokyo 30 (1916) 188. Tree 15-30 m, 20-70 cm @g; bark greyish brown to dark brown, smooth to scaly, lenticellate. Branchlets initially densely yellowish brown to- mentose by adpressed stellate hairs, late glabres- cent, slender, fissured to flaky, sparsely lenticellate; terminal buds ovoid-ellipsoid, 3-4 by 2-3 mm, scales ovate to linear-acute. Stipules linear-acute, 3-4 by 1/3-4 mm, soon caducous. Leaves more or less coriaceous, (7—)10—12(—16) by 3-6 cm (index 2-3.7), broadest at or slightly above the middle; surfaces discolorous, above glabrous, glossy, grey- ish brown, underneath densely yellowish tomen- 330 FLORA MALESIANA [ser. I, vo?" SSSE"Z SA Fig. 20. Lithocarpus turbinatus (STAPF) FORMAN. a. Habit with 3 rachis, b. 2 rachis, c. terminal bud, d. 3 flower, e. 2 flowers, f. longitudinal section of a 2 flower, g. young cupules, A. ripe cupule, i. ripe cupule seen from above, j. ripe fruit, k. ripe fruit seen from above, /. longitudinal section of cupule and fruit; a-b, g-l x %, c-f ¥4 (a & d SAN 29053, b & eSAN A 4472, c & g CLEMENS 33654, f & / Jacoss 5777, h-i HOoLtTtTuM s.n., j-k SAR 20209). 1972] tose by adpressed stellate hairs; base acute to cuneate, rarely rounded, margin recurved, top abruptly acute to 44-1 cm acuminate, with a blunt tip; midrib strongly prominent beneath, impressed above; nerves (5—)7—8(-9) pairs, more or less prominent beneath, impresssed above, parallel, at an angle of 45—70°, arcuating but not anastomos- ing towards the margin; reticulation dense, fine, scalariform, distinct beneath; petiole 1-2 cm, 1-24% mm 9, adaxially flat. Inflorescence male or androgynous, densely yellowish brown tomentose by short stellate hairs; bracts and bracteoles ovate- acute, 1-144 by 1/3—4%4 mm. Male rachis 5-10 cm, 1 mm @; ¢ flowers in clusters of 3 or solitary, perianth 6-lobed, stamens 12, filaments 11%4-144 mm, anthers 0.25 mm long, pistillode subglobose, c. 1 mm @. Androgynous rachis c. 10 cm, 2 mm @; female flowers solitary, perianth 6—lobed, stami- nodes 12, rudimentary, styles 3, conical, 1-144 mm, recurved. Ripe cupule 1-114 cm stalked, ellipsoid, up to 10 by 6 cm, covering the fruit completely; wall woody, 3—5 mm thick, outside densely yellow- ish brown tomentose by short stellate hairs, longitudinally ridged especially when young, with 5-8 undulating lamellae; base attenuate-rounded, top rotundate-umbonate. Ripe fruit ellipsoid, 5—7 cm long, 4-5 cm 9; wall woody, %-11% cm thick, for the greater part adnate to the cupule, scar rounded-attenuate, free part convex, umbonate, densely yellowish brown tomentose by simple hairs; cotyledons flat-convex. Distr. Malesia: Borneo (Sarawak, Sabah, Kali- mantan, scattered). Ecol. Forests up to 1500 m, on dark brown sandstone derived soil. F/. March-May, fr. June- Nov. Note. Authors who recorded this species from Penang and Singapore are supposed to have had specimens of L. burkillii, BANGHAM 1118 from Sumatra, referred by MERRILL to L. beccariana, belongs to L. javensis. 9. Lithocarpus maingayi (BENTH.) REHD. J. Arn. Arb. 1 (1919) 128; A. Camus, Chénes 3 (1954) 577, t. 356; SOEPADMO, Reinwardtia 8 (1970) 257. — Quercus maingayi BENTH. in Hook. f. Ic. Pl. (1880) t. 1314; Hook. f. Fl. Br. Ind. 5 (1888) 617 (‘main- gayii’); KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 82, t. 77; CORNER, Ways. Trees (1940) 304, f. 98. — Pasania maingayi (BENTH.) SCHOTTKY, Bot. Jahrb. 47 (1912) 627; GamBLE, J. As. Soc. Beng. 75, i (1915) 452. — Synaedrys maingayii (BENTH.) Koiwz. Bot. Mag. Tokyo 30 (1916) 189.— L. subnucifera A. CAMUS, Bull. Mus. Hist. Nat. Paris Il, 4 (1932) 123; Chénes 3 (1954) 583, t. 354: 9-16. Tree 10-15 m, 30-40 cm 9. Branchlets initially densely rufous-tomentose by short stellate hairs, later subglabrescent, sturdy, sparsely lenticellate; terminal buds ovoid, 3—5 by 2-3 mm, scales ovate- acute. Stipules ovate-acute, 4-1 by 1/3—% cm, soon caducous. Leaves thick-coriaceous, (13—)17— 20(—22) by (5—)6—8(—10) cm (index 2-2.7), broad- est at or below the middle; surfaces more or less concolorous, above subglabrous, dull to glossy, beneath densely yellowish brown, tomentose by FAGACEAE (Soepadmo) 331 adpressed, stellate hairs; base acute to cuneate, rarely rounded, margin recurved, top abruptly acute to 1 cm acuminate, tip sharp, oblique; midrib strongly prominent beneath, slightly so above; nerves (13—)16—20(-22) pairs, strongly prominent beneath, impressed above, parallel at an angle of 50-70°, arcuating and anastomosing near the margin; reticulation fine, lax, subscalariform, dis- tinct beneath; petiole 2-3 cm, 144-3 mm 9, adaxi- ally flat. Inflorescence male or female, densely velvety rufous stellate hairy; bracts linear-acute, 1144-31, by 0.7-1 mm, bracteoles broadly ovate- acute, 0.7 by 1 mm. Male rachis 10-15 cm, 14%-2 mm 9; flowers in clusters of 3-7, perianth deeply 6—lobed, stamens 12, filaments 114-2 mm, anthers 0.35 mm long, pistillode subglobose, 1-114 mm @. Female rachis 5-10 cm, 3 mm 9; @ flowers solitary, perianth 6—lobed, staminodes 12, rudimentary, styles 3, conical, 1-144 mm, recurved. Young cupule 1 cm stalked, obovoid-ellipscid, outside with 6-8 more or less concentric lamellae, base attenuate-acute, top rotundate-umbonate with an opening of 3-5 mm @g; densely rufous tomentose. Ripe cupule subsessile, 4-5 cm long, 3-314 cm @; wall 3-5 mm thick, covering the fruit almost com- pletely except for an opening of 2-3 mm @; base rounded-acute, top rotundate-umbonate. Ripe fruit ovoid-globose, 344-4 cm long, 244-314 cm 9; wall woody, c. 4% cm thick, for the greater part adnate to the cupule, base rounded-attenuate, top (free part) rotundate-umbonate, 2-3 cm g. Distr. Malesia: Malay Peninsula (Perak, Selan- gor, Negri Sembilan; Fraser Hill, Pahang; Mt Ophir, Malacca; also Penang). Ecol. Hill forest, 600-1000 m. Fi. Oct., fr. Dec.-July. 10. Lithocarpus rotundatus (BL.) A. Camus, Ri- viera Scient. 18 (1932) 41 (‘rotundata’); Chénes 3 (1954) 624, t.; SoEPADMO, Reinwardtia 8 (1970) 275. — Quercus rotundata BL. Verh. Bat. Gen. K. & W. 9 (1823) 219; Bijdr. (1826) 521; Fl. Jav. Cupul. (1829) 22, t. 11; A.DC. Prod. 16, 2 (1864) 88; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 85, t. 100 A; K. & V. Bijdr. 10 (1904) 64; BACKER & Baku. f. Fl. Java 2 (1965) 6. — Pasania rotundata (BL.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83. — Quercus sp. VIDAL, Sinopsis Atlas (1883) 41, t. 92: G. — Quercus clathrata von SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 15; K. & V. Bijdr. 10 (1904) 49; Koorp. Atlas 1 (1913) t. 62. — Quercus pyriformis VON SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 17; K. & V. Bijdr. 10 (1904) 62; Koorp. Atlas 1 (1913) t. 54 (‘piriformis’). — Quercus curranii MERR. Philip. J. Sc. 3 (1908) Bot. 329; En. Philip. 2 (1923) 27. — Synaedrys clathrata (VON SEEMEN) Korpz. Bot. Mag. Tokyo 30 (1916) 190. — Synaedrys curranii (MERR.) Korpz. /.c. 189. — Synaedrys pyriformis (VON SEEMEN) Kotpz. /.c. 189. — Synaedrys ‘rotunda’ (BL.) Korpz. l.c. 189. — L. clathrata (VON SEEMEN) REHD. J. Arn. Arb. 1 (1919) 123; A. Camus, Chénes 3 (1954) 603, t. 363: 1-7. — L. curranii (MERR.) REHD. J. Arn. Arb. 1 (1919) 124; A. Camus, Chénes 3 (1954) 628, t. — L. pyriformis (VON SEEMEN) REHD. J. FLORA MALESIANA 332 Ny 2 ed Bae es ASC Denog eer Laegis- See S55 = pO ea Ses Fig. 21. Lithocarpus echinifer (MERR.) A. CAMUS. 4. of © flower with young cupule, x2, d. young cupule, x 4, g. ripe cupule and fruit in longitudinal section, ripe cupule, d-e SAN 24792, f-g SAN 49504). [ser. I vole? KP Ke ay Yb fd q Oh ay ol ay} Ox oy! ef te AY ny DP 161) Of bi g fs as ps a) Bi fs ip p ga pe ate! Of My , fe) dy OP, AN LD in iy “] HY TH vy, vs 4 4 ; ae] Sa Habit with 3 rachis, x 24, b. 3 flower, <4, ¢. section x %, e. young cupule seen from above, x %, f. x 4 (a-b ELMER 20640, c SAN A 3456, 1972] Arn. Arb. 1 (1919) 130; A. Camus, Chénes 3 (1954) 584. Tree 20—40 m, up to 100 cm @. Branchlets initi- ally densely rufous tomentose by simple and stel- late hairs, later subglabrescent, greyish brown, finely fissured, sparsely to densely warty lenticel- late; terminal buds ovoid-ellipsoid, 3-4 by 14-24% mm, scales ovate to linear-acute. Stipules linear- acute, 4-5 by 0.7-1 mm, with longitudinal ribs, soon caducous. Leaves thin-coriaceous, (10—)13-— 15(—20) by (4-)5-6(-9) cm (index 214-3), broadest at or slightly above the middle; surfaces more or less concolorous, above glabrous, dull, under- neath densely greyish tomentose by adpressed stellate hairs; base rounded to acute, top bluntly acute to sharply 44-1 cm acuminate; midrib and nerves thin, impressed above; nerves 9-11 pairs, subparallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; re- ticulation fine, dense, subscalariform, obscure on both sides; petiole 44-114 cm, c. 1 mm 9, terete or adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous tomentose by short stellate hairs; bracts and bracteoles ovate-acute, 23-1 by 24 mm. Male rachis 10-15 cm, 1-2 mm @; 3 flowers in clusters of 3-7, perianth 6-lobed, stamens 12, filaments 3-4 mm, anthers 0.3-0.5 mm long, pistillode subglobose, i44-2 mm ag. Androgynous or mixed rachis 10-15 cm, 1-2 mm @; female flowers solitary, perianth 6-lobed, sta- minodes 12, rather well-developed but not exceed- ing the perianth, styles 3-4, conical, 1-114 mm, recurved. Ripe cupule %-2 cm stalked, obovoid to pear-shaped, 114-2 cm long, 3-4 cm 9g, wall 2-3 mm thick, densely pale grey stellate-hairy, lower part smooth or ridged, attenuate-rounded, top truncate set with imbricate scales or woody tubercles; covering the greater part of the fruit except for an opening of c. 1 cm 9. Ripe fruit sub- hemispherical, 2 cm long, 3-344 cm 9; wall 2-3 mm thick, for the greater part adnate to the cupule, top flat-convex. Distr. Malesia: Java (mountainous country in W. Java), N. Borneo (SAN/For. Dept. 4943, from Sandakan), Philippines (Luzon, several localities). Ecol. Forest, from sea-level (Borneo) up to 1500 m (Java). Fl. July-Oct., fr. Oct.-Febr. 11. Lithocarpus echinifer (MERR.) A. CAMus, Bull. Soc. Bot. Fr. 80 (1934) 818 (‘echinifera’); Chénes 3 (1954) 632, t. 372; SoppADMOo, Reinwardtia 8 (1970) 235. — Quercus echinifera MeRR. Pl. Elm. Born. (1929) 43. — Fig. 21. Tree 20-35 m, 20-80 cm 9; buttresses up to 3 m tall, 2 m out, 5 cm thick; bark greyish brown, ir- regularly fissured to flaky. Branchlets initially densely yellowish brown tomentose by woolly, simple or stellate hairs, later subglabrous, densely or sparsely warty lenticellate; terminal buds ovoid-ellipsoid, %-1 by 14-14 cm, scales ovate to linear-acute, recurved. Stipules linear-acute, 4-1 by %4—-% cm, recurved, rather long-persistent, and sometimes crowded near the tip of a young twig. Leaves thick-coriaceous, (15—)18-20(-30) by (S—)7-10(-15) cm (index 2144-3), broadest at or FAGACEAE (Soepadmo) 333 rarely above the middle; surfaces more or less con- colorous, above glabrous, dull to glossy, under- neath densely greyish tomentose with adpressed stellate hairs; base rounded to cuneate, margin entire, sometimes undulate, top abruptly acute to 144-1 cm acuminate; midrib prominent on both sides; nerves (9—)11-12(-13) pairs, thinly prom- inent beneath, subparallel, at an angle of 45-60°, arcuating but not anastomosing towards the mar- gin; reticulation fine, dense, subscalariform, ob- scure on both sides; petiole 144-244 cm, 24 mm ®, adaxially flat. Inflorescence densely fulyous stellate-hairy; bracts linear-acute, recurved, 5-6 by 4% mm, bracteoles ovate-acute, 1-114 by %4 mm. Male rachis 10-25 cm, 14%-2 mm @; 3 flowers in clusters of 3-7, perianth 6—7-lobed, stamens 12, filaments 344-5 mm, anthers 0.3-0.4 mm long, pistillode subglobose, 1-144 mm 9. Female rachis not known. Young infructescence 15-20 cm, 3 mm 9; 2 flowers (seen on a young fruit): perianth 6-lobed, staminodes 12, rudimentary, styles 3, conical, 144-2 mm, recurved; young cupule ob- ovoid, densely fulvous-tomentose, scales recurved, hook-shaped. Ripe cupule subsessile to 1-2 cm stalked, obconical, 2-4 cm long, 3-5 cm @; wall up to 4% cm thick, outside densely brownish tomentose by short stellate hairs; scales spirally set, recurved, 2-5 mm long, rather woody; enclos- ing the greater part of the fruit except for an open- ing of %-1 cm o. Ripe fruit subhemispherical, 2-3 cm long, 3-4 cm 9; wall woody, 4-8 mm thick, for the greater part adnate to the cupule, base rounded, top rotundate-umbonate, densely greyish brown stellate-hairy. Distr. Malesia: Borneo (Sarawak, Brunei, Sa- bah, scattered). Ecol. Forests up to 1800 m, on sandy clayey soil, usually on river-banks. F/. Aug.—Nov., /r. Dec.—Sept. 12. Lithocarpus pulcher (KING) MArRKGrR. Bot. Jahrb. 49 (1925) 67 (‘pulchra’); A. CAMus, Chénes 3 (1954) 626, t. 370: 1-4; SoOEPADMoO, Reinwardtia 8 (1970) 269. — Quercus pulchra KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 85, t. 81, p.p., excl. BEc- CARI PB 3269. — Synaedrys pulchra (KING) Korpz. Bot. Mag. Tokyo 30 (1916) 189. Tree 24-36 m, 40-60 cm @g; buttresses up to 2% m tall, 10 cm thick; bark irregularly fissured to scaly, chocolate-brown. Branchlets initially densely rufous pubescent by woolly, simple or stellate hairs, later subglabrous, ridged or terete, sparsely lenticellate; terminal buds ovoid-glob- ose, 3-5 by 2-3 mm, scales ovate to linear-acute. Stipules narrowly ovate to linear, 10 by 2-5 mm, soon caducous. Leaves thick-coriaceous, rigid, (10—)15-20(-22) by (4-)6-8(C-12) cm (index 2-3.2), broadest at or rarely above the middle; sur- faces discolorous, above glabrous except the mid- rib and nerves, dull to glossy, dark, chocolate- brown, beneath densely yellowish to reddish brown tomentose by adpressed stellate hairs; base acute to cuneate, rarely rounded, margin recurved, top acute to 144-1 cm acuminate; mid- rib and nerves strongly prominent beneath, slight- 334 FLORA MALESIANA fser. [yak v7? ere nee ern, =D D- 2. ly so to impressed above; nerves (12-)15—-18(-22) pairs, parallel, at an angle of 60-70°, arcuating but not anastomosing towards the margin; re- ticulation dense, fine, scalariform, distinct be- neath; petiole 1-2 cm, 2-3 mm 9, adaxially flat to shallowly furrowed. Inflorescence male or female, densely rufous stellate-hairy; bracts and bracteo- les ovate-acute, 1-2 by 0.7-1 mm. Male rachis 10-15 cm, 1-144 mm 9; flowers in clusters of 3, perianth 6-lobed, stamens 10-12, filaments 2-3 mm, anthers 0.3-0.35 mm long, pistillode sub- globose, c. 1 mm 9. Female rachis c. 10 cm, 3 mm 9; @ flowers solitary or rarely in clusters of 2-3, perianth 6~—7-lobed, staminodes 12, rather well- developed but not exceeding the perianth, styles 3-4, conical, 1-144 mm, recurved. Ripe cupule sessile, obconical, 3-4 cm long, 4-6 cm @; wall woody up to 8 mm thick; outside densely greyish brown stellate-hairy, upper half with short, thick, pointed tubercles, spirally set, lower half with irregularly set tubercles; covering the greater part of the fruit except for an opening of 144-3 cm 9; base acute, top truncate or rotundate. Ripe fruit subhemispherical, 2-3 cm long, 4-5 cm 9; wall woody, 3-7 mm thick, for the greater part adnate to the cupule, base rounded, top (free part) depressed-umbonate, densely greyish brown to- mentose. Distr. Malesia: Borneo (Sarawak, Sabah, and Kalimantan, scattered). Ecol. Forests, up to 1000 m. Fr. March-Nov. Note. BECCARI PB 3269, on which KING based his description of the inflorescence, belongs to 1bp beccarianus. 13. Lithocarpus longispinus BARNETT, Kew Bull. (1938) 100; A. Camus, Chénes 3 (1954) 887, t.; SoEPADMO, Reinwardtia 8 (1970) 254. — Fig. 22 g-i. Tree 8-30 m; except the buds and inflorescen- ces all parts glabrous. Branchlets greyish brown, sparsely lenticellate; terminal buds ellipsoid, 5-8 by 1-2 mm, scales narrowly ovate to linear-acute. Leaves chartaceous, (12-)16-20(-27) by (3-) 5 ~7(-8) cm (index 3-3.8), broadest at or slightly above the middle; surfaces more or less concol- orous, greenish; base cuneate, top 1-114 cm acu- minate, tip oblique; midrib and nerves thinly prominent on both surfaces; nerves 12-16 pairs, subparallel, at an angle of 60—70°, arcuating but not anastomosing towards the margin; reticula- tion fine, subscalariform, obscure on both sides; petiole 4-1 cm, 2 mm 9, adaxially fiat. Inflores- cence male or androgynous, sparsely simple- hairy; bracts narrowly ovate, c. 2 by 43 mm. Male- rachis 14-20 cm, 1-2 mm @; ¢ flowers in clusters of 3, perianth deeply incised, lobes 5—6, stamens 10-12, filaments 4-5 mm, anthers 0.35 mm long, pistillode subglobose, 1-144 mm @. Androgynous rachis 10-15 cm, 1-2 mm 9; female flower soli- tary, perianth 5—6—lobed, staminodes 10-12, rudi- mentary, styles 3, terete, 144-5 mm, recurved. Ripe cupule sessile, more or less globose, 2-24 cm through, covering the fruit almost completely except for an opening of 3-7 mm 9; wall thinner than 1 mm; scales densely but irregularly set, spine-like but soft and recurved, 1-114 cm long. Ripe fruit ovoid, 1 ¥3-2 cm through; free part dense- ly fulvous-tomentose; scar concave, 1 cm 9; wall c. 1 mm thick, for the greater part free from the cupule. Distr. Siam (common in the Peninsula, scat- tered in the northern parts), in Malesia: Malay Peninsula (Selangor). Ecol. Evergreen forests, up to 600 m, often on river-banks. Fl. Nov.—March, fr. April—-Sept. 14. Lithocarpus wrayi (KiNG) A. Camus, Riviera Scient. 18 (1932) 42; Chénes 3 (1954) 891, t.; SoEPADMO, Reinwardtia 8 (1970) 288. — Quercus lappacea (non Roxs.) Hook. f. Fl. Br. Ind. 5(1888) 607, quoad Malaya; KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 41, t. 33; CoRNER, Gard. Bull. S. S. 10 (1939) 279; Ways. Trees (1940) 303, f. 97. — Quercus wrayii KING, Ann. R. Bot. Gard. Calc. 2 (1889) 77, t. 104. — Pasania wrayii (KING) Gams_e, J. As. Soc. Beng. 75, ii (1915) 446. — Pasania lappacea (non OERST.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 446, quoad Malaya. — Syn- aedrys wrayii (KING) Ko1nz. Bot. Mag. Tokyo 30 (1916) 187. —L. lappacea var. perakensis A. Ca- mus, Bull. Soc. Bot. Fr. 92 (1945) 254; Chénes 3 (1954) 856, t. — Fig. 22 c-d. Tree 12-30 m, 20-50 cm 9. Branchlets initially densely yellowish brown to fulvous-pubescent by erect simple hairs, later glabrescent, whitish grey, densely or sparsely lenticellate; terminal buds ellipsoid, 4-8 by 2 mm, scales linear-acute. Stipules linear-acute or broadly spathulate, 7-10 by 2-3 mm, soon caducous. Leaves thin-coriaceous (10-)15-20(-24) by (3-)5-7(-8) cm (index 2V%,-31%4), broadest at or rarely above the middle; surfaces more or less concolorous, greyish green, sparsely pubescent on both surfaces, densely on the midrib and nerves; base acute to rounded, a- symmetrical, margin recurved, sometimes un- dulate, top bluntly to sharply 1-2 cm acuminate; midrib prominent on both sides; nerves (14-)16-— 18(-19) pairs, parallel, at an angle of 45-70°, prominent beneath, arcuating but not anastomos- ing near the margin; reticulation fine, dense, sca- lariform, obscure on both surfaces; petiole (2-) 3-5(-10) mm, 1-2 mm @, densely simple-hairy, adaxially flat. Inflorescence male or androgynous, densely yellowish brown pubescent with erect sim- ple hairs; bracts narrowly ovate-acute, 2-3 by 0.7-1 mm, bracteoles 0.7—1 by 0.3-0.5 mm. Male rachis 10-20 cm, 1-2 mm 9; ¢ flowers solitary or in clusters of 3, perianth (5—)6(—7)-lobed, stamens 10-12, filaments 3-4 mm, anthers 0.25-0.30 mm long, pistillode subglobose, 1-144 mm 9. Andro- gynous rachis 10-15 cm, 2 mm 9; female flowers solitary, perianth 6-lobed, staminodes 12, rather well-developed but not exceeding the perianth, styles 3, conical, 144-2 mm. Young cupule de- pressed-subglobose to discoid, completely covered by imbricate, acroscopical subulate scales. Ripe cupule sessile, ovoid-globose to depressed-ovoid, 144-2 cm long, 24-3 cm 9; wall 14-1 mm thick, outside densely tomentose; scales subulate, re- 1972] FAGACEAE (Soepadmo) B39 curved; covering the fruit completely; dehiscence irregular. Ripe fruit ovoid- to depressed ovoid- globose, 1-114 cm long, 2-2% cm 9; wall c. %-1 mm thick, for the greater part free from the cupule; free part densely pale brown to rufous tomentose; scar flat to convex. Distr. Peninsular Siam (Trang, BKF 2131), in Malesia: Sumatra (East Coast and near Palem- bang), Malay Peninsula (Kedah, Perak, Pahang, Trengganu). Ecol. Forests, up to 1350 m. Fl. Jan.—April, fr. May-Nov. 15. Lithocarpus coopertus (BLANCO) ReEuHD. J. Arn. Arb. 1 (1919) 124 (‘cooperta’); A. CAMus, Chénes 3 (1954) 890, t.; So—EPADMO, Reinwardtia 8 (1970) 231. — Quercus cooperta BLANCO, FI. Filip. ed. 2 (1845) 503; A.DC. Prod. 16, 2 (1864) 105; Merr. Philip. J. Sc. 3 (1908) Bot. 329; Sp. Blanc. (1918) 121; En. Philip. 2 (1923) 27. — Castanea cooperta (BLANCO) OERsT. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. — Castanopsis costata (non A.DC.) F.-ViLL. Noy. App. (1880) 209. — Quercus fernandesii VIDAL, Sinopsis Atlas (1883) 41, t. 92: E. — Quercus reflexa KING, Ann. R. Bot. Gard. Calc. 2 (1889) 78, t. 72; MERR. Philip. J. Sc. 3 (1908) Bot. 329; Etmer, Leafl. Philip. Bot. 3 (1910) 940; ibid. 6 (1913) 1981; Merr. En. Philip. 2 (1923) 30. — Quercus prui- nosa (non BL.) ELMER, Leafl. Philip. Bot. 3 (1910) 940. — Synaedrys cooperta (BLANCO) KolIpz. Bot. Mag. Tokyo 30 (1916) 186. — Synaedrys refle- xa (KING) Korpz. /.c. 187. — Castanopsis re- flexa (KING) ReHD. J. Arn. Arb. 1 (1919) 122. — Quercus boholensis MeErRR. Philip. J. Sc. 29 (1926) 476. — L. boholensis (MERR.) REHD. J. Arn. Arb. 10 (1929) 132; A. Camus, Chénes 3 (1954) 905, t. — L. reflexa (KING) A. Camus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 867, t. — Fig. 22 e-f. Tree 10-45 m, 15-70 cm @g; buttresses up to 2 m tall, 14% m out, 10 cm thick; bark greyish brown, smooth to flaky, lenticellate. Branchlets initially densely yellowish brown to fulvous to- mentose by stellate hairs, later subglabrescent, terete, greyish brown, sparsely lenticellate; ter- minal buds ovoid, 2-4 by 2-3 mm, scales ovate to linear-acute. Stipules deltoid, 4-8 by 1-3 mm, soon caducous. Leaves thin-coriaceous, (5—)10—14(-17) by (2-)4-6(-7) cm (index 2-4), broadest about the middle; surfaces discolorous, above glabrous, dull, greyish brown, underneath densely yellow- ish brown to fulvous tomentose by adpressed, stellate hairs; base rounded-acute, top acute to 1-2 cm acuminate; midrib strongly prominent on both surfaces; nerves (10—)12—-14(-16) pairs, more or less prominent on both sides, subparallel, at an angle of 45-70°, arcuating but not anastomos- ing near the margin; reticulation fine, arched, sca- lariform, obscure on both surfaces; petiole 4—6 mm, 1-2 mm @, densely stellate-hairy, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely yellowish brown stellate-hairy; bracts narrowly ovate-acute, 1-144 by 4-24 mm, bracteoles linear, 4%-1 by 4% mm. Male rachis 5-15 cm, 1-144 mm 9; ¢ flowers solitary, rarely in clusters of 2—3, perianth (5—)6(—7)—lobed, stamens 10-12, filaments 3-4 mm, anthers 0.25-0.30 mm long, pistillode subglobose, 1-144 mm @. Andro- gynous or mixed rachis 7-10 cm, 1-2 mm 9; female flower solitary, perianth 6-lobed, staminodes 12, rudimentary, styles 3-4, conical, 14%4,-2 mm, recurved. Ripe cupule subsessile, ovoid-conical, 114-2 cm long, 1144-2¥% cm 9; wall thinner than 1 mm, covering the fruit completely; scales ir- regularly set, densely tomentose, subulate, recury- ed; dehiscence irregular. Ripe fruit ovoid-coni- cal, c. 144-2 cm through, densely yellowish brown tomentose; top acute, scar deeply concave to flat; wall thinner than 1 mm, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (rare), Bor- neo (Sarawak, Brunei, Sabah, fairly common), Philippines (common in Mindanao, rare and scat- tered in Luzon, Leyte, Samar, and Surigao). Ecol. Forests, up to 1800 m, occasionally in peat swamp and heath forest, usually on yellow- ish sandy soil. F/. Febr.—Oct., fr. Nov.—May. Uses. In Borneo the wood is used by the local people for housing construction, but serves mainly for fire-wood. Notes. The Bornean specimens usually have smaller leaves than those from the Philippines, but the fruit and cupule are similar. ELMER 14012 from the Philippines is chosen as the neotype of BLANCO’s species. 16. Lithocarpus confragosus (KING ex Hook. /f.) A. Camus, Riviera Scient. 18 (1932) 40 (con- fragosa’); Chénes 3 (1954) 832, t.; SOEPADMO, Reinwardtia 8 (1970) 230. — Quercus confragosa KING ex Hook. f. Fl. Br. Ind. 5 (1888) 616; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 77, t. 71. — Pasania confragosa (KING ex Hook. f.) SCHOTTKY, Bot. Jahrb. 47 (1912) 662; GAmBLE, J. As. Soc. Beng. 75, ii (1915) 446. Tree 12-29 m, 25-100 cm 9; bark greyish brown, smooth to scaly, lenticellate. Branchlets initially densely greyish brown tomentose by adpressed, stellate hairs, glabrescent, densely warty lenticel- late; terminal buds ovoid-globose, 3—4 by 2-3 mm, scales ovate. Stipules linear to lanceolate, 10-15 by 2-3 mm. Leaves thick-coriaceous, rigid, (10-) 12-18(—27) by (3%4—)5-7(-10) cm (index 2.2-4.5), broadest about the middle; surfaces more or less concolorous, above glabrous, glossy, underneath densely greyish brown, tomentose by adpressed, stellate hairs, glabrescent; base acute to cuneate, rarely rounded, margin recurved, top acute to 1-114 cm acuminate; midrib prominent beneath, slightly so above; nerves (5—)6-8(-10) pairs, prominent beneath, obscure above, subparallel, at an angle of 45-60°, arcuating but not anasto- mosing towards the margin; reticulation lax, sub- scalariform, obscure on both sides; petiole 1-2 cm, 2-4 mm 9, glabrous, adaxially flat. Ripe cupule sessile to 1 cm stalked, depressed-ovoid-globose, 24%-3 cm long, 3-5% cm 9; wall 2-10 mm thick, brittle, covering the fruit almost completely ex- cept for an opening of smaller than 14 cm 9, out- 336 FLORA MALESIANA [ser. I, vol. 7? side irregularly set with rounded to pointed short tubercles; dehiscence irregular. Ripe fruit depressed ovoid-globose, 1144-21% cm long, 2-4 cm @, base truncate, scar flat to concave, 1144-21 cm 9, top rounded and depressed umbonate at the centre; wall for the greater part free from the cupule, 2-8 mm thick, densely yellowish brown to ful- vous tomentose; cotyledons flat-convex. Distr. Malesia: N. Sumatra (Gajo Lancs), Malay Peninsula (Perak), Borneo (Sarawak: Nyabau F. R.; Sabah, scattered; Nunukan I.). Ecol. Forests, up to 2000 m, on yellowish brown sandy soil. Fr. Aug.—Febr. Note. Inflorescence not known. 17. Lithocarpus neorobinsonii A. CAMus, Chénes, Atlas 3 (1949) 77, t. 410: 1-6; Chénes 3 (1954) 780; SoOEPADMO, Reinwardtia 8 (1970) 261. — Quercus robinsonii RipL. J. Fed. Mal. St. Mus. 5 (1914) 46, non MerRR. 1915. — Pasania robinsonii (RIDL.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 450. — Fig. 22 a-b. Tree c. 15 m, 40—50 cm 9; bark smooth. Branch- lets initially densely fulvous tomentose by stel- late hairs, later glabrous, sparsely lenticellate; terminal buds ovoid-globose, c. 3 by 2 mm, scales broadly ovate. Stipules linear to lanceolate, 8-10 by 2-3 mm. Leaves thick-coriaceous, (10—)12-14 (-1514) by 4-6 cm (index 2-3), broadest at or ra- rely above the middle; surfaces discolorous, above glabrous, glossy, dark brown, beneath densely silvery grey-tomentose by adpressed, stellate hairs; base acute, margin sometimes undulate, top acute to %-144 cm acuminate; midrib more or less prominent on both sides, dark brown; nerves 13-15 pairs, prominent beneath, obscure above, parallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation dense, scalariform, distinct beneath; petiole 4-144 cm long, 2 mm 9g, adaxially flat. Inflorescence male or androgynous, densely fulvous stellate-hairy; bracts and bracteoles ovate, 1-3 by 1 mm. Male rachis c. 10 cm, 1-14% mm 9; g flowers solitary or in clusters of 3, perianth 6-lobed, stamens 12, filaments 2-3 mm, anthers 0.3 mm long; pistillode subglobose, c. 1 mm @. Androgynous rachis c. 10 cm, 1-2 mm 9; female flowers solitary, pe- rianth 6-lobed, staminodes 12, rudimentary, styles 3-4, conical, 2-3 mm, recurved. Ripe cupule ses- sile to 4% cm stalked, ovoid, 2-244 cm through; base truncate to concave, top acute; wall thinner than 14 mm, covering the fruit almost completely, densely yellowish brown to greyish tomentose; scales minute, adpressed, irregularly set; dehis- cence irregular. Ripe fruit ovoid-conical, 144 cm long, 2-244 cm 9; wall bony, 1-114 mm thick, densely yellowish brown tomentose, for the great- er part free from the cupule; top acute, base rounded, scar flat to concave, c. 1 cm @; cotyle- dons flat-convex. Distr. Malesia: Malay Peninsula (Selangor, rare; Fraser Hill, Pahang). Ecol. Forests, at 1500-1700 m. Fi., fr. Oct.— May. 18. Lithocarpus kostermansii SOEPADMO, Rein- wardtia 8 (1970) 251. — Quercus blumeana (non Kortn.) K. & V. Bijdr. 10 (1904) 57; Koorp. Atlas 1 (1913) t. 53; BACKER & BAKH. f. Fl. Java, 2 (1965) 6. — L. blumeana (non REHD.) A. CAMUS Chénes 3 (1954) 773, t. 408: 1-11, quoad Java. Tree 18-30 m, 30-60 cm 9; buttresses 1-4 m tall, 44-14 m out; bark grey. Branchlets initially densely fulvous tomentose by adpressed, stellate hairs, later subglabrous, greyish brown, sparsely lenticellate; terminal buds ovoid-globose, 2-4 by 1-2 mm, scales narrowly ovate to linear. Stipules linear, 3-6 by 1%4-1 mm. Leaves thick-coriaceous, (12—)16-22(-30) by (4-)6-8(--10) cm (index (1.6—)3-314(-4)), broadest about the middle; sur- faces more or less discolorous, above glabrous, dull, chocolate-brown, beneath densely greyish tomentose by adpressed, stellate hairs; base acute to cuneate, rarely rounded, top acute to 1-3 cm acuminate; midrib prominent beneath, slightly so above; nerves (8—)10—11(—13) pairs, prominent beneath, obscure above, parallel at an angle of 45-60° with the midrib, arcuating but not ana- stomosing towards the margin; reticulation lax, subscalariform, obscure on both sides; petiole 1-2 cm, 2-344 mm @, glabrous, adaxially flat. Inflorescence male, androgynous or mixed, den- sely brownish stellate-hairy; bracts linear acute, 1-2 by 144 mm, bracteoles narrowly ovate, 4-24 by %-'% mm. Male rachis 10-20 cm, 2 mm 9; g flowers in clusters of 3, perianth 6-lobed, stamens 12, filaments 4-5 mm, anthers 0.3-0.4 mm long, pistillode subglobose, 1-14 mm @. Androgynous or mixed rachis 15-35 cm, 2-3 mm @; female flowers solitary, rarely in clusters of 2-3, perianth 6-lobed, staminodes 12, rudimentary, styles 3-4, conical, 144-2 mm, recurved. Ripe cupule sub- sessile, depressed ovoid-globose, 2-24 cm long, 234-3 cm 9; wall thinner than 44 mm, covering the fruit completely, outside at the basal half with 3-4 thin, concentric lamellae, at the upper half set with obscure adpressed scales to almost smooth densely fulvous stellate-hairy; dehiscence irre- gular. Ripe fruit depressed ovoid-globose, c. 2 cm long, 244-3 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule, densely fulvous tomentose; base rotundate, scar concave, top abruptly acute. Distr. Malesia: W. Java. Ecol. Forests, up to 1000 m. Fi. July—Sept., fr. Oct.—Jan. 19. Lithocarpus pattaniensis BARNETT, Kew Bull. (1938) 104; A. Camus, Chénes 3 (1954) 770, t.; SOEPADMO, Reinwardtia 8 (1970) 266. Small tree. Branchlets initially densely greyish tomentose by adpressed, stellate hairs, later gla- brous, dark greyish brown, sparsely lenticellate; ter- minal buds ovoid, c. 3 by 2 mm, scales ovate- acute. Stipules deltoid, c. 1 by 4% mm, soon ca- ducous. Leaves thick-coriaceous, (8—)14-16(-18) by (3-)6-8 (-11) cm (index 1.8-2.2), broadest at or rarely above the middle; surfaces discolor- ous, above glabrous, glossy, chocolate-brown, underneath densely greyish brown tomentose by 1972] FAGACEAE (Soepadmo) 337 Fig. 22. Lithocarpus neorobinsonii A. CAMUS. a. Habit, b. longitudinal section of cupule and fruit, both x 24. — L. wrayi (KiNG) A. Camus. c. Cupule, side view, d. cupule seen from above, both nat. size. — L. coopertus (BLANCO) REHD. e. Cupule, side view, f. cupule seen from above, both nat. size. — L. long- ispinus BARNETT. g. Cupule, side view, h. cupule seen from above, i. section of cupule showing the fruit, all nat. size (a-b KEP 32856, c-d HENDERSON s.n., e—f BECCARI PB 4057, g-i RAJAB 523). adpressed, stellate hairs; base rounded, sometimes asymmetrical, margin recurved, top bluntly acute to 1 cm acuminate; midrib and nerves strongly prominent beneath, flat to impressed above; nerves 11-14 pairs, subparallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation subscalariform, distinct beneath; petiole 1-2 cm, 2-3 mm 2, adaxially flat. Inflorescence male or female, densely pale yellowish brown stellate hairy; bract and brac- 338 FLORA MALESIANA [ser. I, vol. 7? eoles ovate-acute, 1144-3 by 1-2 mm. Male rachis 10-15 cm, 2 mm @; ¢ flowers in clusters of 3, perianth 6-lobed, stamens 12, filaments 2-3 mm, anthers 0.3-0.35 mm @, pistillode subglobose, 144-2 mm o@. Female rachis 10-15 cm, 3-4 mm 0; 2 flowers solitary or in clusters of 2-3, perianth 6-lobed, or occasionally irregularly lobed, sta- minodes 12, rudimentary, styles 4-6, conical, 144-2 mm, connate. Ripe cupule solitary or in clusters of 2-3, sessile, ovoid-globose, c. 2 cm long, 24% cm 9; wall thinner than 1 mm, covering the fruit almost completely except for an opening smaller than 1 cm 9, outside densely fulvous to greyish tomentose by stellate, adpressed hairs; lamellae 8-10, concentric or spiral, thin; dehiscen- ce irregular. Ripe fruit depressed ovoid-globose, 1-11 cm long, 2-2% cm 9; top rounded, base ro- tundate, scar concave; wall 1-2 mm thick, woody, for the greater part free from the cupule, outside densely fulvous to rufous tomentose. Distr. Siam (two localities, in the northern part and in the Peninsula), in Malesia: Malay Peninsu- la (Perak, Pahang). Ecol. Forests, in N. Siam at 700 m, in Malaya at 1500-1700 m. Fi. in Malaya Sept.—Nov., /r. March-Sept. 20. Lithocarpus encteisacarpus (KORTH.) A. CAMUS Riviera Scient. 18 (1932) 40 (‘encleisocarpa’); Chénes 3 (1954) 767, t. 406: 1-5, incl. var.; SOEPAD- MO, Reinwardtia 8 (1970) 239. — Quercus encleisacarpa KorTH. Kruidk. (1844) 209, t. 45; BL. Mus. Bot. 1 (1850) 288, incl. var. divergens; A.DC. Prod. 16, 2 (1864) 103; WENziGc, Jahrb. Bot. Gart. Berl. 4 (1886) 238; Hook. f. Fl. Br. Ind. 5 (1888) 617, incl. var. aperta KING ex HooK. f.; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 80, t. 75: 1-4, incl. var. aperta; CORNER, Ways. Trees (1940) 302, f. 95, 98. — Cyclobalanus encleisa- carpa (KORTH.) OgersT. Vid. Medd. Naturh. For. Kjo6bn. 8 (1867) 81. — Pasania encleisacarpa (KORTH.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 449, incl. var. aperta. — Synaedrys encleisacarpa (KorTH.) Komz. Bot. Mag. Tokyo 30 (1916) 186. — Castanopsis encleisacarpa (KORTH.) REHD. J. Arn. Arb. 1 (1919) 122. — L. encleisacarpa var. typica A. Camus, Bull. Soc. Bot. Fr. 93 (1945) 84. Tree 10-40 m, 25-75 cm 9; buttresses up to 1 m tall; bark grey, smooth to scaly, lenticellate. Branch- lets initially densely fulvous to rufous tomen- tose by adpressed, stellate hairs, later subglabrous, dark greyish brown, sparsely lenticellate; termi- nal buds subglobose, 2-3 by 23-1 mm, scales broadly ovate. Leaves thin-coriaceous, 12-15 by 4—6 cm (index 2-3), broadest about the middle; surfaces more or less discolorous, above dark greyish brown, dull, subglabrous, beneath densely glaucous adpressed stellate-hairy; base rounded or acute, margin occasionally undulate, top acute to bluntly 1-2 cm acuminate; midrib prominent beneath, slightly so to impressed above; nerves (7—)8-10(—12) pairs, parallel, at an angle of 45—70°, more or less prominent beneath, arcuating and anastomosing near the margin; reticulation lax, subscalariform to irregular, more or less distinct beneath; petiole 4-144 cm, 1-2 mm 4@, terete or adaxially flat. Inflorescence male, female or an- drogynous, densely fulvous stellate-hairy; bract and bracteoles ovate-acute, 1-144 by 1 mm. Male rachis 10-15 cm, 1 mm @; ¢ flowers in clusters of 3 or solitary, perianth 6-lobed, stamens 12, filaments 3-4 mm, anthers 0.30-0.35 mm long, pistillode subglobose, 1-144 mm go. Female or androgynous rachis 10-20 cm, 2 mm @; @ flowers solitary, rarely in clusters of 2-3, perianth 6- lobed, staminodes 12, rudimentary, styles 3-4, conical, c. 1 mm. Ripe cupule 1-1% cm stalked, ovoid-globose, 144-2 cm long, 2-3 cm 9; top rounded, base truncate; wall thinner than | mm, covering the fruit completely, outside densely fulvous tomentose by stellate hairs, longitudinally, thinly ribbed; lamellae thin, 5-7, more or less con- centric; dehiscence irregular. Ripe fruit depressed ovoid-globose, 1144 cm long, 2-2% cm 9; wall thinner than 1 mm, for the greater part free from the cupule, outside densely silvery tomentose; scar concave. Distr. Peninsular Siam (Songkla, KERR 15883), in Malesia: Sumatra (scattered), Malay Penin- sula (common; also Singapore), Borneo (scat- tered in Sarawak and Sabah). Ecol. Forests, up to 1300 m. Fi. Febr.—Aug., fr. Sept.—April. Note. Specimens from above 900 m, usually have a smaller cupule and fruit, and the cupule is cup-shaped, covering c. ¥ of the fruit; these have sometimes been distinguished as var. aperta. 21. Lithocarpus mariae SOEPADMO, Reinwardtia 8 (1970) 258. Tree 15-33 m, 50-60 cm 9, buttresses c. 23 m tall; bark smooth, lenticellate, yellowish grey. Branchlets initially densely fulvous to rufous tomentose by stellate hairs, soon glabrescent, dark greyish brown, smooth or sparsely lenti- cellate; terminal buds ovoid, 144-2 by 1-2 mm, scales ovate. Leaves thin-coriaceous, (8—)10—-12 (-13) by 4-5 cm (index 2-3), broadest at or below the middle; surfaces slightly discolorous, above glabrous, glossy, greenish to chocolate-brown, beneath densely glaucous tomentose by adpressed minute stellate hairs; base rounded acute, top abruptly acute to 1-14% cm acuminate; midrib prominent beneath, flat above; nerves 10-12 pairs, thin, prominent beneath, flat to impressed above, parallel, at an angle of 50-60°, arcuating but not anastomosing towards the margin; re- ticulation fine, dense, scalariform, distinct be- neath; petiole 114-21 cm, 1-114 mm 9, adaxially flat or shallowly furrowed. Inflorescence male or androgynous, simple, densely fulvous tomentose by adpressed stellate hairs; bracts and bracteoles ovate-acute, 23-1 by 4-23; mm. Male rachis c. 10 cm, 2 mm @; g flowers solitary or in clusters of 2-3, perianth 6-lobed, stamens 12, filaments 3-4 mm, anther 0.35 mm long, pistillode globose, 1-144 mm 9g, densely whitish tomentose, Andro- gynous rachis c. 10 cm, 2 mm @; female flowers solitary, perianth 6-lobed, staminodes 12, rudi- mentary, styles 3-5, conical, 1-144 mm, connate. Young cupule 1-11 cm stalked, globose, densely rufous-tomentose by adpressed stellate hairs. Ripe cupule on a peduncle 1—2 cm long, c. 1 cm @, ovoid, rounded-acute at both ends, 2-214 cm long, 2%-3 cm @, enclosing the fruit except for an ope- ning of 3 mm; wall thinner than 1 mm; inside densely fulvous-tomentose with adpressed, simple hairs, outside smooth or with obscure longitudi- nal streaks, densely glaucous adpressed stellate- tomentose. Ripe fruit ovoid, 2—-21% cm long, 2%-23/4, cm @, densely fulvous-tomentose by ad- pressed simple hairs, top rounded-acute, base rounded, scar deeply concave, c. 2 cm @, wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: rare). Ecol. In forest on ridges at 300-600 m. FI. fr. Oct.—Nov. Notes. Leaves collected from a basal shoot measure 22 by 10 cm, with 14 pairs of nerves. Near L. encleisacarpus, but differering in its leaves with more nerves, dense scalariform re- ticulation, and in its cupule without concentric rings and rounded-acute at both ends. Borneo (Sarawak, Sabah, 22. Lithocarpus macphailii (HENDERS.) BARNETT, Trans. & Proc. Bot. Soc. Edinb. 34 (1944) 178; A. Camus, Chénes 3 (1954) 773, t.; SOEPADMO, Reinwardtia 8 (1970) 257. — Pasania macphai- lii HENDERS. Gard. Bull. S. S. 5 (1930) 76, f. Tree 10-45 m, up to 50 cm 9; bark smooth to fissured, greyish brown. Branchlets initially den- sely fulvous to rufous adpressed stellate-hairy, later glabrous, dark, greyish brown, sparsely to densely lenticellate; terminal buds ovoid-ellip- soid, 2-5 by 2-3 mm, scales narrowly to broadly ovate-acute. Stipules narrowly ovate-acute, c. 5 by 2mm, soon caducous. Leaves thin-coriaceous, 15-22 by 6-8 cm (index 2-3), broadest at or below the middle; surfaces discolorous, above glabrous, greyish to dark brown, dull to glossy, underneath densely glaucous tomentose with adpressed, stellate hairs; base rounded or acute, top abruptly, bluntly 1-144 cm acuminate; mid- rib and nerves prominent beneath, almost so above; nerves (10—)12-16(-18) pairs, subparal- lel, at an angle of 50—70°, arcuating but not anas- tomosing towards the margin; reticulation dense, fine, subscalariform, distinct beneath; petiole 1-1.7 cm, 1-2 mm 9, shallowly furrowed. In- florescence male, female or androgynous, densely fulvous stellate hairy; bracts and bracteoles nar- rowly ovate-acute, 0.7-144 by %-0.7 mm. Male rachis 10-15 cm, 2 mm @; ¢ flowers in clusters of 3, filaments 3-4 mm, anthers 0.3—0.35 mm long, pistillode subglobose, 1-14 mm o. Female or androgynous rachis 10-25 cm, 2 mm @; @ flowers solitary or in clusters of 2-3, staminodes rudi- mentary, styles 3-5, conical, 1144-2 mm, connate. Ripe cupule sessile to 4% cm stalked, deeply cup- shaped to almost completely covering the fruit, 25-114 cm long, 2-3 cm 9, wall thin, outside den- sely fulvous-tomentose, with 5—8 thin, more or FAGACEAE (Soepadmo) ne EEE UEnEIEN EEE EEE 339 less concentric, lamellae. Ripe fruit depressed ovoid-globose, 1-14% cm long, 2-244 cm @, top rounded-acute, base rounded, scar flat-concave; wall woody, 1-2 mm thick, densely greyish brown to silvery tomentose by simple adpressed hairs, for the greater part free from the cupule. Distr. Peninsular Siam, in Malesia: rather common in Sumatra and Malay Peninsula. Ecol. Forests, up to 900 m, usually on river- banks. Fi/. June—Oct., fr. May. Note. Records of Quercus and Pasania blu- meana from the Malay Peninsula may belong here. 23. Lithocarpus blumeanus (KorTH.) REHD. J. Arn. Arb. 10 (1929) 132; A. CAmus, Chénes 3 (1954) 774, excl. fig., et specim. e Java et Ma- laya; SOEPADMO, Reinwardtia 8 (1970) 221. — Quercus blumeana Kortu. Kruidk. (1844) 208, t. 44; A.DC. Prod. 16, 2 (1864) 103. — Cyclo- balanus blumeana (KorTH.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Synaedrys blumeana (KortH.) Korpz. Bot. Mag. Tokyo 30 (1916) 186. — Castanopsis blumeana (KORTH.) Renp. J. Arn. Arb. 1 (1919) 122. Tree 5-15 m, 10-25 cm @; bark scaly, greyish brown. Branchlets initially densely yellowish to reddish brown tomentose by adpressed, stellate hairs, later subglabrous, greyish brown, sparsely lenticellate; terminal buds ovoid, 1-1% by 1 mm, scales ovate-acute. Stipules deltoid, c. 1 by 14 mm soon caducous. Leaves thin-coriaceous (8-)10-15(-17) by (31%4-) 4-5 (-6) cm (index 2-2.4), broadest at or slightly below the middle; surfaces more or less discolorous, above glabrous, dull to glossy, dark brown, beneath densely glau- cous tomentose by adpressed stellate hairs; base acute, shortly decurrent, margin undulate, top 1-114 cm acuminate; midrib prominent on both surfaces, stronger beneath; nerves (11—)12—14(-16) pairs, more or less prominent beneath, parallel, at an angle of 45-60°, arcuating but not anasto- mosing towards the margin; reticulation fine, den- se, scalariform, obscure; petiole 14-1 cm long, 1 mm 9, adaxially flat or shallowly furrowed. In- florescence male, female or androgynous, densely pale greyish brown stellate hairy; bract and bract- eoles ovate to linear-acute, 1-2 by 14-4 mm. Male rachis 15-20 cm, 1 mm @; ¢ flowers solitary or in clusters of 3, filaments 3-4 mm, anthers 0.3-0.35 mm long, pistillode subglobose, c. 1 mm @. Female or androgynous rachis 10-15 cm, Imm 9; 2 flowers solitary, staminodes rudimentary, styles 3, conical, 1-144 mm, recurved. Ripe cupule Y%-1 cm stalked, ovoid or deeply cup- shaped, 1-114 cm long, 244-3 cm 9, covering the fruit for 24-34 or almost completely; wall thinner than 14% mm, densely greyish brown tomentose on both surfaces; lamellae thin, concentric, den- ticulate; dehiscence irregular. Ripe fruit de- pressed-ovoid, c. 144 cm long, 244-3 cm 9, gla- brous except for a small part at the long, acumi- nate top, base cordate, scar concave; wall bony, c. 1mm thick, dark purplish brown, for the greater part free from the cupule. 340 FLORA MALESIANA [ser. I, vol. 7? Distr. Malesia: Borneo (scattered in Sarawak, E. Kalimantan, and Sabah). Ecol. Forests, up to 1650 m, usually on river- banks, on sandstone or basalt derived soil. Fi. June—Aug., fr. Aug.—May. Notes. Records from Malaya, under Quercus, may belong to L. macphailii, and those from Java to L. kostermansii. In SAN 16199, 16448, and 41760 from 1200— 1650 m, the leaves and cupules are larger than those from the lowland forest. Several interme- diates exist, however, so that they may be regarded as montane form of the species. 24. Lithocarpus platycarpus (BL.) REHD. J. Arn. Arb. 1 (1919) 130; A. Camus, Chénes 3 (1954) 698, t.; SOEPADMO, Reinwardtia 8 (1970) 268. — Quercus platycarpa Bu. Fl. Jav. Cupul. (1829) 27, t. 15; A.DC. Prod. 16, 2 (1864) 92; WeEnzic, Jahrb. Bot. Gart. Berl. 4 (1886) 234; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 70, t. 65; K. & V. Bijdr. 10 (1904) 53; BACKER & BAKH. f. Fl. Java 2 (1965) 7. — Cyclobalanus platycarpa (BL.) OersT. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80. — Synaedrys platycarpa (BL.) Koz. Bot. Mag. Tokyo 30 (1916) 192. Tree 15-35 m, 40—50 cm 9; bark fissured, whitish grey. Branchlets glabrous, whitish grey, smooth to superficially fissured; terminal buds ovoid- ellipsoid, 4-5 by 2-3 mm, scales linear-acute. Sti- pules linear-acute, 5-7 by 1-2 mm. Leaves thick- coriaceous, (8—) 12-15(-20) by (4~-)5-8(-9) cm (index 2-2.5), broadest at or above the middle; surfaces more or less concolorous, above gla- brous, glossy, underneath densely whitish grey tomentose by adpressed stellate hairs, subgla- brescent; base rounded-acute, margin recurved, top acute to bluntly 1 cm acuminate; midrib prominent on both surfaces; nerves 11-14 pairs, subparallel, at an angle of 45-60°, arcuating and anastomosing near the margin; reticulation fine, irregular to areolate, obscure; petiole Y%-114 cm, 2-3 mm @, adaxially flat. Inflorescence male or female, densely stellate hairy; bracts and bracteoles linear acute, 2-4 by 44 mm. Male rachis 10-15 cm, 1-2 mm 9; ¢ flowers in clusters of 3—4, rarely sol- itary, filaments 3-4 mm, anthers 0.3 mm long, pistillode subglobose, c. 1 mm @. Female rachis c. 10 cm, 2-3 mm 9; @ flowers solitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3, conical, 1-144 mm. Ripe cupule Y-1% cm stalk- ed, saucer-shaped, 44-1 cm long, 3-4 cm 9; wall woody, covering c. 144 part of the fruit, outside with 5-7 strongly prominent lamellae. Ripe fruit depressed ovoid-globose, 1144-2 cm long, 3-4 cm @, chocolate-brown, glabrous, glossy, top de- pressed-umbonate, base rounded, scar flat-convex; wall woody, 2-5 mm thick, for the greater part free from the cupule. Distr. Malesia: W. and S. Central Java (once collected in SW. Bantam, several times in Nusa Kambangan). Ecol. Forests, at low alutude. F/. April, fr. Aug. —Sept. 25. Lithocarpus perakensis SOEPADMO, Reinward- tia 8 (1970) 266. — Quercus costata (non BL.) Hook. f. Fl. Br. Ind. 5 (1888) 617. — Quercus costata var. convexa (non BL.) KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 82, t. 76 A. — Pasania costata (BL.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 443, quoad Malaya. — L. costata var. kingii A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 254; Chénes 3 (1954) 583, t. 353: 16-19. Tree 15-30 m, 30-90 cm 9. Branchlets initially densely fulvous stellate hairy, later glabrous, black- ish brown, fissured; terminal buds ovoid-ellip- soid, 5-7 by 3—4 mm, scales ovate to linear-acute. Stipules linear, 5—7 by 2 mm. Leaves thick-coria- ceous, (10—)12-16(—17) by (4-)5—6(-7) cm (index 2-21), broadest at or above the middle; upper surface glabrous, dull or glossy, underneath spar- sely greyish tomentose by adpressed stellate hairs, glabrescent; base rounded-acute to cuneate, top bluntly 44-2 cm acuminate; midrib strongly prominent beneath, flat above; nerves 11-14 pairs, thin and obscure on both sides, subparallel, at an angle of 45-60°, arcuating and anastomosing near the margin; reticulation fine, areolate, dis- tinct beneath; petiole 1-2 cm, 1-2 mm 9, adaxially flat to shallowly furrowed. Inflorescence male or female, densely fulvous stellate hairy; bracts and bracteoles linear, 2-3 by % mm. Male rachis 10-20 cm, 2-3 mm 9; ¢ flowers in clusters of 3, filaments 3-4 mm, anthers 0.3-0.35 mm long, pistillode subglobose, 1-144 mm 9. Female rachis 15-20 cm, 2-3 mm 9; @ flowers solitary, perianth thick-coriaceous, staminodes rudimentary, styles 3, conical, 1-114 mm, recurved. Ripe cupule 1-2 cm stalked, cup-shaped, 1-144 cm long, 3-4 cm 9; wall woody, thick, rim incurved, covering 14-14 part of the fruit, outside with 5-6 more or less concentric, flat or prominent lamellae. Ripe fruit broadly depressed ovoid-globose, 1-144 cm long, 3-4 cm @, glabrous, dark brown, glossy, top round- ed-umbonate, base rounded, scar convex; wall woody, 3—5 mm thick, for the greater part free from the cupule. Distr. Malesia: Johore, Trengganu). Ecol. Forests, 90-1200 m. Fi. fr. Sept--March Malay Peninsula (Perak, 26. Lithocarpus sericobalanus E. F. WARB. Kew Bull. (1936) 20; A. Camus, Chénes 3 (1954) 716, t.; SOEPADMO, Reinwardtia 8 (1970) 279. Tree 20-50 m, 30-100 cm @; buttresses up to 1.8 m tall, 2 m out, 10 cm thick; bark deeply fissur- ed to scaly, pale to dark brown. Branchlets initial- ly densely greyish brown tomentose by adpressed stellate hairs, later subglabrous, finely fissured; terminal buds ovoid-ellipsoid, 244-4 by 2 mm, scales narrowly ovate. Leaves coriaceous, (9—)12— 15(—20) by (3-)4—6 (—8) cm (index 2-3), broadest at the middle; surfaces discolorous, above gla- brous, chocolate-brown, dull to glossy, under- neath densely fulvous to glaucous tomentose by adpressed stellate hairs; base rounded-acute, top bluntly %-1 cm acuminate; midrib and nerves prominent beneath; nerves 10-12 pairs, parallel, at an angle of 45-60°, arcuating but not anasto- 1972] FAGACEAE (Soepadmo) 341 mosing towards the margin; reticulation fine, den- se, scalariform, obscure; petiole 1-2 cm, 2 mm 9, terete or adaxially flat. Inflorescence male or an- drogynous, densely greyish brown tomentose by adpressed stellate hairs; bracts and bracteoles ovate, 3-4 by 2-3 mm. Male rachis 10-15 cm, 2 mm 9; ¢ flowers in clusters of 3, filaments 3-344 mm, anthers 0.2-0.3 mm long, pistillode subglo- bose, 1-14%4 mm 9. Androgynous rachis 10-15 cm, 1-2 mm 9; female flowers solitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3-5, con- ical, 1-2 mm, recurved. Ripe cupule %-1 cm stalked, broadly cup-shaped, 44-2 cm long, 3-444 cm 9; wall woody, rim thick, covering Y,_2% part of the fruit, outside with 6-10 concen- tric or spiral lamellae. Ripe fruit broadly de- pressed ovoid-globose to discoid, 1-2 cm long, 3-4 cm @, top rounded-umbonate, occasionally depressed at the centre, base rounded, scar strong- ly convex; wall woody, 2-3 mm thick, outside densely greyish tomentose by adpressed simple hairs, for the greater part free from the cupule. Distr. Malesia: Borneo (all parts except in Brunei; also Nunukan I.). Ecol. In primary, also in secondary and heath forests, on hills or low ridges, up to 1200 m, on sandy soil. F/. Aug.—Dec., fr. Jan.—_May. 27. Lithocarpus lucidus (RoOxsB.) REHD. J. Arn. Arb. 1 (1919) 128; A. Camus, Chénes 3 (1954) 690, t. 386: 1-15; SoEPADMO, Reinwardtia 8 (1970) 254. — Quercus lucida Roxs. FI. Ind. ed. Carey 3 (1832) 635; Hook. f. Fl. Br. Ind. 5 (1888) 614; KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 69, t. 64; CORNER, Ways. Trees (1940) 304, f. 95. — Quercus omalokos KortH. Kruidk. (1844) 214; A.DC. Prod. 16, 2 (1864) 92; Hook. f. FI. Br. Ind. 5 (1888) 614 (omalkos’); KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 70, t. 23 B. — Quercus cu- neata RoxsB. ex A.DC. Prod. 16, 2 (1864) 108. — Cyclobalanus omalokos (KoOrRTH.) OERsT. Vid. Medd. Naturh. For. Kj6bn. 8 (1867) 80. — Pasa- nia omalokos (KORTH.) SCHOTTKY, Bot. Jahrb. 47 (1912) 676; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 439. — Pasania lucida (ROxB.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 440. — Synaedrys omalkos (KORTH.) Koipz. Bot. Mag. Tokyo 30 (1916) 192. — L. omalokos (KorTH.) REHD. J. Arn. Arb. 1 (1919) 129; A. CAMus, Chénes 3 (1954) 695, t. 387: 9-20. Tree 15-40 m, 20-120 cm 9; buttresses up to 1 m tall; bark greyish brown, shallowly fissured. Branchlets initially densely fulvous stellate hairy, soon glabrescent, with 3-5 sharp ribs decurrent under the leaf-insertion, pale to dark brown, spar- sely lenticellate; terminal buds ovoid-ellipsoid, 4-5 by 1-2 mm, scales ovate to linear. Stipules linear-acute to subulate, 5 by 2mm, soon caducous. Leaves chartaceous to coriaceous, (4-)8—15(-18) by (2-)3-5(-7) cm (index 2-314), broadest at or more commonly above the middle; surfaces concolorous, pale to dark brown, glabrous, above glossy; base cuneate and decurrent into the pet- iole, top rounded, obtuse-emarginate, acute to abruptly cuspidate; midrib prominent on both sides; nerves (10—)14—16(—20) pairs, thin and ob- scure on both surfaces, parallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, areolate, obscure; petiole 2-5 mm long, 2-3 mm 9g, glabrous, terete or adaxially flat. Inflorescence male, female or an- drogynous, densely fulvous stellate hairy; bracts and bracteoles ovate, c. 1 by 4% mm. Male rachis 10-20 cm, 2-3 mm 9; 3 flowers in clusters of 3, perianth with 4-6 lobes, stamens 8~—12, filaments 2-3 mm, anthers 0.2— 0.3 mm long, pistillode sub- globose, c. 1 mm 9. Female or androgynous rachis 10-15 cm, 2-3 mm @; flowers soiitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3, conical, 1-114 mm. Ripe cupule sessile, cup- or saucer-shaped, 14-1 cm long, 3-3% cm 9; wall woody, rim thick, incurved, covering the basal part of the fruit; lamellae 8-10, concentric, thick, densely greyish stellate hairy outside. Ripe fruit depressed ovoid, 2-24% cm long, 3-344 cm 9, glabrous, glossy, chocolate-brown, top rounded- acute, base truncate, scar concave, 1144-2 cm 9; wall woody, 3-5 mm thick, for the greater part free from the cupule; cotyledons flat-convex, ir- regularly divided by the endocarp intrusions. Distr. Malesia: Sumatra (Asahan, Langkat, Indragiri, and Riouw), Malay Peninsula (common, also Singapore and Penang), Borneo (Sarawak, NE. Kalimantan, and Nunukan I.). Ecol. Forests up to 1600 m. Fi., fr. June-May. Notes. Specimens from the lowland forest, attributed by most previous authors to L. oma- lokos, usually have a thin, pale greyish brown leaf with an acute or abruptly cuspidate top, and those from above 1000 m, usually credited to L. lucidus, have the leaves thick-coriaceous, dark brown, with rounded to obtuse-emarginate top. Recent col- lections from the Malay Peninsula and Borneo show that the lowland and montane forms can not be considered as separate species, as there are many intermediates. Among the loan of Fagaceae received from Brussels, I found the MSS. name Quercus cuneata, a name originally proposed by ROxBURGH, but which he in the Calcutta Herbarium deliberately changed into Quercus lucida Roxs. A. DE CAN- DOLLE, unaware of this, validated Quercus cuneata Roxs. which is consequently a later, superfluous homonym of Quercus lucida. 28. Lithocarpus eichleri (WENzIG) A. CAMus, Ri- viera Scient. 18 (1932) 40; Chénes 3 (1954) 718, t. 395; SoEPADMO, Reinwardtia 8 (1970) 236. — Quercus eichleri WENzI1G, Jahrb. Bot. Gart. Berl. 4 (1886) 236; Hook. f. Fl. Br. Ind. 5 (1888) 615; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 72, t. 68. — Pasania eichleri (WENZIG) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 438. — Synaedrys eichleri (WENZIG) Koipz. Bot. Mag. Tokyo 30 (1916) 191. Tree 10-15 m, 25-60 cm 9. Branchlets initially densely fulvous to rufous tomentose by adpressed stellate hairs, later glabrous, whitish grey, dense- ly warty lenticellate; terminal buds ovoid-ellip- soid, 2-3 by 1-2 mm, scales linear to narrowly 342 FLORA MALESIANA [ser. I, yor 7? ovate-acute. Stipules narrowly ovate to linear, 3-4 by 1-144 mm, soon caducous. Leaves thin- coriaceous, (14-)18—26(-30) by (4-)6-8(-12) cm (index 214-5), broadest at or above the middle; surfaces more or less concolorous, above glabrous, dull to glossy, underneath densely greyish tomen- tose by adpressed stellate hairs, subglabrescent; base acute, rarely rounded, margin recurved, top bluntly to sharply 1-144 cm acuminate; midrib prominent on both sides; nerves (7—)8-11(-15) pairs, prominent beneath, impressed above, sub- parallel, at an angle of 45—-60°, arcuating but not anastomosing towards the margin; reticulation fine, lax, subscalariform, obscure; petiole 1-1% cm, 144-4 mm @, adaxially flat. Inflorescences male, androgynous or mixed, densely pale grey stellate hairy; bracts and bracteoles ovate, c. 1 by % mm. Male rachis 10-15 cm, 1 mm @; 3 flowers solitary or in clusters of 3, perianth (5—) 6(-8)-lobed, stamens (10—)12(-13), filaments 3-4 mm, anthers 0.3-0.35 mm long, pistillode subglobose, longitudinally compressed, 1-2 mm @. Androgynous or mixed rachis 10-20 cm, 2-4 mm @; female flowers solitary, staminodes well- developed and exceeding the perianth, sometimes polliniferous; styles 3, conical, c. 2 mm, recurved. Ripe cupule sessile, saucer-shaped, 4%-1 cm long, 2-314 cm @, covering the basal part of the fruit; rim thin, entire or undulate; outside densely grey- ish brown stellate hairy; lamellae 5—8, concentric, thin. Ripe fruit depressed-ovoid, 1-144 cm long, 214-3 cm @, densely fulvous short stellate-tomen- tose; top rounded-umbonate, base rounded, scar deeply concave, 1-2 mm 9; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: S. Sumatra (Palembang and vicinity), Malay Peninsula (Perak, Selangor, Johore). Fig. 23. Lithocarpus korthalsii (ENDL.) SOEPADMO. a. Habit with f and 9 rachis, x 24, b. part of g rachis, «4, c. 2 flower, «4, d. young cupules, x 24, e. ripe cupule and fruit, « 2 (a-c & e Jacoss 4565, d KoORTHALS 5S.71.). 1972] FAGACEAE (Soepadmo) 343 Ecol. Forests, up to 240 m, usually in swampy places. Fl. June-Sept., fr. Oct.—Febr. 29. Lithocarpus korthalsii (ENDL.) SOEPADMO, Reinwardtia 8 (1970) 251. — Quercus korthalsii ENDL. Gen. Pl. Suppl. 4, 2 (1847) 28, non BL. 1850; Mia. Ann. Mus. Bot. Lugd. -Bat. 1 (1863) 113. — Quercus annulata Kortu. Kruidk. (1844) 213, t. 46: 21-22, non SmiTH 1819. — Quercus pseudoannulata BL. Mus. Bot. 1 (1850) 299. — Quercus tysmannii BL. Mus. Bot. 1 (1850) 300; Mia. FI. Ind. Bat. 1, 1 (1856) 860 (‘teysmannii’); A. DC. Prod. 16, 2 (1864) 92; OUDEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 205; Natuurk. Verh. Kon. Akad. 11 (1865) 14, t. 8; WENziIG, Jahrb. Bot. Gart. Berl. 4 (1886) 235; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 71, t. 66; K. & V. Bijdr. 10 (1904) 52; Koorp. Atlas 1 (1913) t. 47; BACKER & BAKH. f. FI. Java 2 (1965) 7. — Quercus pseudomolucca var. incrassata BL. Mus. Bot. 1 (1850) 291; A.DC. Prod. 16, 2 (1864) 86; OupeM. Natuurk. Verh. Kon. Akad. 11 (1865) 9. — Quercus laurifolia Mia. Pl. Jungh. 1 (1851) 11, non Micwx. 1801. — Quercus hypoleuca Mia. Fl. Ind. Bat. 1, 1 (1858) 869. — Cyclobalanus tysmanni (BL.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80; Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 375 (tysmanninii’). — Pasania teysmanni (BL.) PRANTL in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 55. — Quercus heliciformis VON SEE- MEN, Bot. Jahrb. 27, Beibl. 64 (1900) 15; K. & V. Bijdr. 10 (1904) 47; Koorp. Atlas 1 (1913) t. 49. — Synaedrys heliciformis (VON SEEMEN) Koipz. Bot. Mag. Tokyo 30 (1916) 191. — Synae- drys teysmanni (BL.) Korpz. /.c. 193. — L. heli- ciformis (VON SEEMEN) REHD. J. Arn. Arb. 1 (1919) 127; A. Camus, Chénes 3 (1954) 700, t. 389: 1-14. — L. teysmannii (BL.) REHD. J. Arn. Arb. 1 (1919) 131; A. Camus, Chénes 3 (1954) 692, t. 387: 1-8. — Fig. 23-24. Tree 25-45 m, 100-150 cm @; bark dark grey, lenticellate. Branchlets initially densely fulvous to rufous-tomentose with simple or stellate hairs, angular, later glabrous, terete, densely or sparsely lenticellate; terminal buds ovoid-ellipsoid, 5—12 by 2—3 mm, scales linear-lanceolate. Stipules linear to subulate, 4-144 by \% cm, long persistent. Leaves thick-coriaceous, (11—)13—16(—23) by (3-) 4-7(-9) cm (index 2.3—3.3), broadest at the middle; above glabrous, glossy, pale to dark chocolate- brown, beneath densely glaucous tomentose by adpressed stellate hairs; base rounded-acute, mar- gin recurved, top bluntly or sharply 1-144 cm acuminate; midrib and nerves prominent beneath, flat above; nerves (13—)15—20(—25) pairs, subparal- lel, at an angle of 50-60°, arcuating but not anas- tomosing towards the margin; reticulation fine, dense, subscalariform, obscure to distinct be- neath; petiole 144-2 cm, 2-4 mm 4g, glabrous, ad- axially flat. Inflorescence male or androgynous, densely yellowish to greyish brown stellate hairy; bracts and bracteoles narrowly ovate to linear-acute, 144-5 by 34-1144 mm. Male rachis 10-20 cm, 144-3 mm 9; ¢ flowers in clusters of 3, stamens 10-12, filaments 3-4 mm, anthers 0.35 mm long, pistillode globose, 1-144 mm @. Androgynous rachis 10-20 cm, 2-3 mm @; female flowers soli- tary or in clusters of 2—3, staminodes rudimen- tary, styles 3-4, conical, 1-3 mm. Ripe cupule subsessile, saucer-shaped, 7-2 cm long, 344-414 cm 9; rim thick, entire or undulate, covering 44-4 part of the fruit; lamellae 6-9, entire or undulate, Fig. 24. Lithocarpus korthalsii (ENDL.) SOEPADMO. Fruiting rachis, x 2/3 (JAcoBs 4565). 344 FLORA MALESIANA [ser. I, vol. 72 strongly prominent, outside densely fulvous to- mentose. Ripe fruit depressed ovoid-globose to subhemispherical, 144-224 cm long, 3-414 cm @, glabrous, dark chocolate-brown, top depressed or rounded-umbonate, base truncate or rounded, scar deeply concave, rarely convex, 244-3 cm 9; wall woody, 2-4 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Sumatra (scattered), Java (western part, eastwards to Mts Wilis and Idjen). Ecol. Forests at 150-1900 m. F/. June-Oct., fr. Oct.-Aug. 30. Lithocarpus urceolaris (JACK) MERR. J. Arn. Arb. 33 (1952) 241, p.p., excl. syn. Quercus eichle- ri; SOEPADMO, Reinwardtia 8 (1970) 285. — Quercus urceolaris JACK, Mal. Misc. 2, 7 (1822) 86; Hoox. Comp. Bot. Mag. 1 (1836) 256; A.DC. Prod. 16, 2 (1864) 89; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 92; A. Camus, Chénes 3 (1954) 1172. — Quercus oligoneura KortH. Kruidk. (1844) 203; BL. Mus. Bot. 1 (1850) 294; A.DC. Prod. 16, 2 (1864) 88; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 91; A. Camus, Chénes 3 (1954) 1165. — Pasania urceolaris (JACK) OxErRsT. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83. — Pasania oligoneura (KortTH.) Ogrst. I. c. 84. — Quercus bancana (non ScCHEFF. 1870) SCHEFF. Nat. Tijd. N. I. 32 (1871) 416; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 62, t. 56 B. — Pasania cra- terophora FISCHER, Kew Bull. (1932) 319, f. 1-6. — UL. ‘cratephora’ (FiscHER) A. CAmus, Not. Syst. 5 (1935) 75; Chénes 3 (1954) 952, t. 458, 459: 1-8. — L. bancana (non REHD.) A. CAMus, Chénes 3 (1954) 694, t., excl. Nova Guinea. Tree 15-33, 20-130 cm 9; buttresses up to 2 m tall, spreading; bark greyish brown, smooth to deeply fissured or scaly. Branchlets initially dense- ly fulvous to rufous tomentose by simple and stellate hairs, later subglabrous, sparsely to densely lenticellate; terminal buds ovoid-ellipsoid, 4—5 by 2-3 mm, scales linear. Leaves thick-coriaceous, (10—)18-25(-35) by (5-)8-10(-17) cm (index 1.7—3.4), broadest about the middle; above gla- brous, or sometimes with some rufous tomentum on midrib and nerves, dull to glossy, underneath glaucous-tomentose by adpressed stellate hairs; base rounded to acute, margin recurved, top bluntly to sharply acute to 1-2 cm acuminate; midrib strongly prominent on both surfaces; nerves (7—)9-10(-12) pairs, flat on both sides, parallel, at an angle of 45-60°, arcuating but not anasto- mosing towards the margin; reticulation dense, fine, scalariform, obscure to distinct beneath; petiole (%4-)1-144(-2%) cm, 2-3 mm g, subglabrous, adaxially flat. Inflorescence male or androgynous, densely yellowish brown stellate hairy; bract and bracteoles narrowly ovate, 1-114 by 4% mm. Male rachis 15-20 cm, 2-3 mm 9; ¢ flowers in clusters of 3, filaments 3-5 mm, anthers 0.20- 0.35 mm long, pistillode globose, c. 1% mm @. Androgynous rachis 10-25 cm, 2-3 mm 9; female flowers solitary, rarely in clusters of 3, staminodes rudimentary, styles 3-4, conical, 1-2 mm, recur- ved. Ripe cupule sessile to 1 cm stalked, deeply cup-shaped, 114-2 cm long, 4-5 cm ag, rim re- curved (sometimes very strongly), entire or un- dulate, covering 14—' part of the fruit; wall woody, densely glaucous to fulvous adpressed stellate- tomentose; lamellae thin, obscure, denticulate, the scales sometimes rather distinct. Ripe fruit de- pressed subglobose or globular-cylindrical, 3-4 cm in size, top rounded to depressed-umbonate, base truncate, scar deeply concave, conical, c. 144-2 cm 9; wall woody, 2-4 mm thick, outside densely greyish tomentose by adpressed simple hairs, greater part free from the cupule; cotyle- dons flat-convex. Distr. Malesia: Sumatra (scattered), Banka, Malay Peninsula (Pahang, Johore), Borneo (com- mon, especially in Sabah). Ecol. In primary, also in secondary and swamp forests, up to 1800 m, on granitic sandy or loamy soil. Fertility seems to be throughout the year, with the fruit ripe between Aug. and Febr. 31. Lithocarpus indutus (BL.) REHD. J. Arn. Arb. 1 (1919) 127; A. Camus, Chénes 3 (1954) 702, t. 390: 1-12; SoEPADMo, Reinwardtia 8 (1970) 247. — Quercus induta BL. Verh. Bat. Gen. K. & W. 9 (1823) 220; Bijdr. (1826) 522; Fl. Jav. Cupul. (1829) 23, t. 12: 1=2, incl. var.)B; Ve: 25a, A.DC. Prod. 16, 2 (1864) 96, incl. var. microcarpa, I.c. 97; OUDEM. Natuurk. Verh. Kon. Akad. 11 (1865) 13, t. 7; WENzIG, Jahrb. Bot. Gart. Berl. 4 (1886) 228; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 55, t. 51; K. & V. Bijdr. 10 (1904) 37; Koorp. Atlas 1 (1913) t. 60; BACKER & BAKH. f. Fl. Java 2 (1965) 7. — Cyclobalanus induta (BL.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80. — Synaedrys induta (BL.) Korbz. Bot. Mag. Tokyo 30 (1916) 195. — Pasania induta (BL.) S. Moore, J. Bot. 63 (1925) Suppl. 144. Tree 20-45 m, 50-150 cm g; buttresses numer- ous; bark dark grey, fissured, lenticellate. Branch- lets initially densely greyish brown tomentose by adpressed simple and stellate hairs, later glabrous, smooth or sparsely warty lenticellate; terminal bud ovoid, c. 4 by 3 mm, scales linear to lance- olate. Leaves thick-coriaceous, rigid, (15—)18—20 (-26) by (5—)7-9(-11) cm (index 2144-3), broadest at or slightly below the middle; surfaces more or less discolorous, above glabrous, dull to glossy, greyish brown, underneath with a thin cover of glaucous tomentum, hairs adpressed, minute, stellate or simple; base rounded-acute, margin re- curved, top bluntly acute to sharply 1—2 cm acumi- nate; midrib strongly prominent on both sides; nerves (11—)12-14(-15) pairs, flat on both sur- faces, parallel, at an angle of 50—-70°, arcuating but not anastomosing towards the margin; reti- culation fine, dense, subscalariform, distinct beneath; petiole 1-2 cm, 2-4 mm @g, glabrous, terete. In florescence male, androgynous or mixed, densely greyish adpressed stellate hairy; bract and bracteoles linear, 1-24% by ¥% mm. Male rachis 10-20 cm, 1144-2 mm 9g; ¢ flowers solitary or in clusters of 2-3, filaments 3—5 mm, anthers 0.2-0.3 mm long, pistillode subglobose, 1-14 mm @. Androgynous or mixed rachis 10-25 cm, 1-2 mm 1972] FAGACEAE (Soepadmo) 345 g; female flowers solitary or in clusters of 2-3, staminodes rudimentary, styles 3, terete, 2-3 mm, recurved. Ripe cupule 1-14% cm stalked, broadly cup-shaped, 1-2 cm long, 3-4 cm g, rim thin, covering %4-24 part of the fruit; outside ob- scurely tuberculate, the tubercles in concentric rows or irregularly. Ripe fruit subhemispherical, 2-3 cm long, 3—4 cm g, densely greyish tomentose, top flat-umbonate, base truncate, scar flat, c. 144-2 cm g; wall woody, 3-5 mm thick, for the greater part free from the cupule. Distr. Malesia: Java (W. Java, rather common, eastwards to Mt Slamet, 109° 15’ BE). Ecol. In submontane forest up to 1800 m. Fi. mainly in March, fr. Aug.—Oct. DocTers VAN LEEUWEN (Ned. Kruidk. Arch. 51, 1941, 134) described a leaf-gall caused by a gall-midge, and a stem-gall by a coccid. 32. Lithocarpus cyclophorus (ENDL.) A. CAMuUs, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 714, t. 393; BARN. Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 334; ibid. 34 (1944) 177; SopEPADMo, Rein- wardtia 8 (1970) 233. — Quercus depressa ROXB. Fl. Ind. ed. Carey 3 (1832) 640, non Hums. & BonpL. 1809, nec BL. 1826. — Quercus cyclophora ENDL. Gen. Pl. Suppl. 4, 2 (1847) 28; A.DC. Prod. 16, 2 (1864) 102; Hook. f. Fl. Br. Ind. 5 (1888) 615; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 71, t. 67; CORNER, Ways. Trees (1940) 302, f. 97. — Quercus penangensis Mia. FI. Ind. Bat. 1, 1 (1856) 859. — Quercus umbonata HANCE, J. Bot. 12 (1874) 241; ibid. 13 (1875) 364. — Pasania cyclophora (ENDL.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 442. — Synaedrys cyclophora (ENDL.) Korpz. Bot. Mag. Tokyo 30 (1916) 191. — L. pseudoplatycarpus A. CAMUS, Bull. Mus. Hist. Nat. Paris II, 4 (1932) 914; Chénes 3 (1954) 716, t. 394: 1-6. Tree 18-40 m, 20-120 cm g; buttresses up to 2 m tall and out; bark deeply fissured or scaly. Branchlets initially densely yellowish brown to ful- vous tomentose by adpressed stellate hairs, later glabrous, lenticellate; terminal buds ovoid-ellip- soid, 3-6 by 2-3 mm, scales ovate to linear. Stip- ules narrowly lanceolate to linear, 10 by 3 mm. Leaves thick-coriaceous, rigid, (18—)20—25(—30) by (6—-)7-9(—12) cm (index 2.5-3.5), broadest at or slightly below the middle; surfaces discolorous, above glabrous, dark chocolate-brown, glossy, underneath densely yellowish brown to fulvous adpressed stellate hairy; base rounded-acute, top bluntly 1-144 cm acuminate; midrib and nerves strongly prominent beneath, slightly so to flat above; nerves (14-)15—17(—20) pairs, parallel, at an angle of 45—-60°, arcuating but not anastomo- sing towards the margin; reticulation dense, scala- riform, distinct beneath; petiole 1-2 cm, 2-3 cm g, adaxially flat. Inflorescence male or androgynous, densely yellowish brown adpressed stellate hairy; bract and bracteoles thick-coriaceous, linear, 24%4— 3¥% by % mm. Male rachis 10-15 cm, 2%-3 mm @; 3 flowers in clusters of 3, filaments 3-4 mm, anthers 0.25-0.3 mm long, pistillode globose, c. 1 mm g. Androgynous rachis c. 10 cm, 3-4 mm g; female flowers solitary or in clusters of 2-3, sta- minodes rudimentary, styles 3, conical, c. 1 mm. Ripe cupule sessile, broadly obconical to cup- shaped or saucer-shaped, 114-214 cm long, 4-6 cm g, rim %-1 cm thick, incurved, covering 14- Y, part of the fruit; wall woody, lamellae 8-10, obscure, rounded, concentric or irregular, densely fulvous stellate hairy. Ripe fruit broadly depressed subglobose, 1144-2% cm long, 344-41% cm g, top depressed-umbonate, base rounded, scar strongly convex; wall woody, 3—5 mm thick, outside dense- ly yellowish brown to fulvous tomentose with adpressed simple hairs, for the greater part free from the cupule. Distr. Peninsular Siam (Betong at 5° 45’ N), in Malesia: Sumatra (scattered), Malay Peninsula (Perak, Selangor, Malacca; also Penang and Singa- pore). Ecol. In primary, rarely also in secondary forest, at 150-1500 m. F/. Nov.-April, fr. Aug.-Nov. 33. Lithocarpus luteus SOEPADMO, Reinwardtia 8 (1970) 255. Tree 18-36 m, 30-100 cm g; buttresses up to 2 m tall and out; bark deeply fissured to scaly, reddish brown. Branchlets initially densely yellow- ish brown to fulvous tomentose by adpressed stel- late hairs, later glabrous, dark greyish brown, densely warty lenticellate; terminal buds ovoid, 3 by 2 mm, scales narrowly ovate-acute. Stipules linear-acute, 4-6 by 1 mm. Leaves thick-coriaceous, (7-)9-12(-15) by (2%4-)3%4-5(-6) cm (index 24% —3), broadest at or slightly below the middle; sur- faces more or less concolorous, yellowish brown, above glabrous, glossy, underneath with a thin cover of adpressed stellate hairs; base acute to cuneate, margin recurved, top acute to 1 cm acumi- nate; midrib and nerves yellowish, thin, prominent beneath, flat above; nerves 9-12 pairs, dense, parallel, at an angle of 45-50°, arcuating but not anastomosing towards the margin; reticulation fine, dense, scalariform, distinct beneath; petiole 8-13 mm, 2 mm g, adaxially flat. Inflorescence male or androgynous, densely yellowish brown stellate hairy; bracts and bracteoles narrowly ovate to linear, 2-244 by 1 mm. Male rachis 10- 15 cm, 1% mm g; ¢ flowers in clusters of 3-7, stamens 10-12, filaments 2144-314 mm, anthers 0.35 mm long, pistillode subglobose, c. 1 mmg. Androgynous rachis 5-10 cm, 2-3 mm 9; female flowers in clusters of 2-3 or solitary, staminodes well-developed and exceeding the perianth, styles 3, conical, c. 1 mm, recurved. Cupule sessile, soli- tary or more commonly in clusters of 2-3, cup- shaped, 1-114 cm long, 244-3 cm 9g; rim thick, sometimes incurved, covering + 14 part of the fruit; wall woody, dark chocolate-brown, lamellae obscure, undulate, densely stellate hairy. Fruit ovoid to subhemispherical, 1-114 cm long, 2-24% cm g, densely fulvous to greyish tomentose, top acute to rounded-umbonate, base truncate, scar flat, c. 2 cm 9; wall woody, 2-3 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Mt Mulu, Mt Murut, 346 FLORA MALESIANA [ser. I, vol. 7 Kapit, Sarawak; Mt Kemul, Kalimantan; Mt Ki- nabalu, Sabah, very common). Ecol. Forests, at 1100-1800 m, on brownish sandstone derived soil. F/. July—Jan., fr. Dec.—June. 34. Lithocarpus schlechteri MARKGR. Bot. Jahrb. 59 (1924) 69, f. 2; A. CAMus, Chénes 3 (1954) 633, t. 373: 14-17; SoEPADMO, Reinwardtia 8 (1970) 278. — L. perclusa MARKGR. Bot. Jahrb. 59 (1924) 68; A. CAMus, Chénes 3 (1954) 608. Tree 10-26 m, 10-50 cm g; bark light grey- brown, lenticellate. Branchlets initially densely ful- vous to rufous tomentose by adpressed stellate hairs, later glabrous, dark greyish brown, finely fissured or sparsely lenticellate; terminal buds ovoid, 1-2 by 1 mm, scales ovate. Stipules ovate, 1-114 by 1 mm. Leaves thin-coriaceous, (7—)8—12 (-16) by (3-)314-5(-6) cm (index 2.2-2.7), broad- est at or rarely below the middle; surfaces discolor- ous, above glabrous, dark chocolate-brown, dull to glossy, underneath with a thin cover of pale greyish brown tomentum, hairs adpressed, mi- nute, stellate; base rounded-acute to acute, top bluntly acute to 44-1 cm acuminate; midrib and nerves thinly prominent beneath, flat to impressed above; nerves (7—)8—9(—10) pairs, subparallel, at an angle of c. 45°, arcuating but not anastomosing towards the margin; reticulation fine, subscalari- form, distinct beneath; petiole glabrous, 5-8 mm, 1-2 mm g, adaxially flat to furrowed, dark colour- ed. Inflorescence male or androgynous, densely greyish stellate hairy; bracts and bracteoles ovate, 1-14% by %-1 mm. Male rachis 7-10 cm, 1-1% mm g; ¢ flowers in clusters of 2—3, filaments 2-3 mm, anthers 0.30 mm long, pistillode subglobose, c. 14% mm @. Androgynous rachis c. 5 cm, 2-3 mm 9; female flowers solitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3-6, conical, 1-144 mm. Ripe cupule sessile, obconical, 3-314 cm long, 414-51, cm g; rim thin, incurved over the rounded top of the fruit, covering + 2% part of the fruit; wall woody, 3—5 mm thick; scales woody, thick, adpressed, with rounded edges or keeled, imbri- cate but set on concentric rows. Ripe fruit de- pressed-subglobose, 2144-31 cm long, 344-414 cm 9, glabrous, dark chocolate-brown, top depressed- umbonate at the centre, base rounded, scar strong- ly convex, c. 2-3 cm g; wall woody, 44-1 cm thick, for the greater part free from the cupule; cotyle- dons flat-convex. Distr. Malesia: New Guinea (from the Vogel- kop Peninsula to the eastern parts, rather com- mon). Ecol. Often a common and gregarious consti- tuent of the Fagaceous forest at 800-1900 m, sporadic in the lower limit of Nothofagus forest at 2200 m, on sandy clayey soil. Fl. March—Oct., fr. Nov.—July. Notes. Quercus imperialis VON SEEMEN in K. Sch. & Laut. Fl. Schutzgeb. (1901) 263, t. 4 F. — Synaedrys imperialis (VON SEEMEN) KoIpz. Bot. Mag. Tokyo 30 (1916) 195. — L. imperiulis (von SEEMEN) MARKGR. Bot. Jahrb. 59 (1924) 69; A. Camus, Chénes 3 (1954) 632, t., may belong here, but as the author did not indicate any particular specimen, and described the species based on a single cupule (lost), and the description cannot b2 recognized with certainty, the later name schlechteri is here accepted. L. perclusa was based on a specimen with very young cupules, while L. schlechteri was based on a specimen with ripe cupules. Recent collections from New Guinea show that they are conspecific. The cupule of L. schlechteri is so far the largest among known species in New Guinea. 35. Lithocarpus megacarpus SOEPADMO, Reinward- tia 8 (1970) 259. Tree 9-30 m, 10-50 cm g. Branchlets glabrous, sturdy, dark greyish brown, with sparse or dense lenticels; terminal buds ovoid, 2-3 by 2 mm, scales narrowly ovate-acute, rufous tomentose. Stipules narrowly ovate, 1-144 cm by 4%-1 mm. Leaves thick-coriaceous, rigid, (9—)11-14(-16) by (3%) 5-6(—8) cm (index (2—)2.2-2.6(—3.2)), broadest at or rarely below the middle; surfaces concolorous, greyish brown, above glabrous, dull to glossy, un- derneath with a thin cover of minute, adpressed stellate hairs, soon glabrescent; base rounded-acute to attenuate-acute, margin recurved, top acute to 14-1] cm acuminate; midrib and nerves prominent beneath, flat above; nerves (7—)8-10(—11) pairs, subparallel, at an angle of 45—50°, arcuating but not anastomosing towards the margin; reticulation coarse, lax, subscalariform, distinct beneath; peti- ole glabrous, 6-10 mm, 2-214 mm g, shallowly furrowed. Inflorescence male or androgynous, spar- sely greyish brown stellate-tomentose; bracts and bracteoles ovate-acute, 0.7-1 by 0.7 mm. Male rachis c. 10 cm, 2 mm g; ¢ flowers in clusters of 3, filaments 3-4 mm, anthers 0.30 mm long, pis- tillode globose, 1-114 mm g. Androgynous rachis c. 5 cm, 2 mm @; female flowers solitary or in clusters of 2-3, staminodes rudimentary, styles 3, conical, c. 1 mm. Ripe cupule subsessile, broadly saucer-shaped, 1-144 cm long, 4-54% cm g; rim thick, rounded, covering the basal part of the fruit; wall woody, densely greyish brown tomentose by adpressed stellate hairs; scales thick, adpressed, imbricate but concentrically set. Ripe fruit de- pressed-subglobose, rarely ovoid, glabrous, choco- late-brown, 2-34 cm long, 4-5 cm g, top rounded- apiculate to depressed-umbonate, base truncate, scar concave but convex at the centre, 2-4 cm g; wall woody, 5-8 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: New Guinea (rather rare in the western part, more common in the eastern part). Ecol. Forests, 1200-1900 m, common on steep slopes. Fi. Jan._June, fr. July—Nov. 36. Lithocarpus revolutus HATUS. ex SOEPADMO, Reinwardtia 8 (1970) 273, f. 12. — Fig. 25. Tree 12 m, 30 cm g; bark soft, corky, creamy in colour. Branchlets initially densely yellowish brown, stiff, stellate-pubescent, soon glabrescent, greyish brown, sparsely lenticellate; terminal buds ovoid, 4-5 by 3-4 mm, scales narrowly ovate or linear. Stipules narrowly ovate-acute, 4-5 by 2- 2% mm. Leaves thick-coriaceous, rigid, 844-14 by 1972] 3-5 cm (index 2-3), broadest at or above the middle; surfaces concolorous, greenish brown, sparsely stiff, stellate-pubescent on both sides, glabrescent; base acute, margin strongly revolute, top rounded to bluntly acute, tip emarginate; midrib and nerves strongly prominent beneath, flat to impressed above; nerves 6-8 pairs, sub- parallel, at an angle of 45—60°, arcuating but not anastomosing near the margin; reticulation sub- N ']G FAGACEAE (Soepadmo) 347 scalariform, fine, obscure to rather distinct be- neath; petiole 5-6 mm, 3-4 mm g, adaxially flat. Male rachis 10-15 cm, 3 mm g, densely yellowish brown stiff stellate hairy; bracts narrowly ovate, 2-21% by 1-114 mm, bracteoles ovate, 1 by 1 mm; 3 flowers in clusters of 3, filaments 3144-5 mm, anthers 0.3-0.35 mm long, pistillode subglobose, 1144-2 mm g. Female flowers (seen as young fruit) solitary or in clusters of 2-3, staminodes well Fig. 25. Lithocarpus revolutus HATUS. ex. SOEPADMO. a. Habit with 3 rachis, x 2, b. details of 3 flower, x4, c. 2 flowers, <4, d. ripe cupule and fruit, x 24, e. longitudinal section of cupule and fruit, x 75 (RSNB 4171). 348 FLORA MALESIANA [ser. I, vol. 72 developed and exceeding the perianth, styles 3, terete, 3-4 mm long, recurved. Ripe cupule sessile, woody, cup-shaped, 114-2 cm long, 334-444 cm 9; rim thin but rigid, covering 44-14 part of the fruit; inside densely silvery brown simple sericeous, outside densely fulvous stellate tomentose; scales ovate, adpressed, imbricate. Ripe fruit depressed ovoid-conical, 2-2 14 cm long, 3-4 cm g, glabrous, dark chocolate-brown; top rounded-acute, base rotundate, scar concave, 2-244 cm g; wall woody, up to 1 cm thick, for the greater part free from the cupule. Distr. Malesia: North Borneo (Mt Kinabalu; Sarawak, rare). Ecol. In forests at 1500 m. Fi. fr. Jan._May. Note. The female inflorescence is not yet known. 37. Lithocarpus brassii SOEPADMO, Reinwardtia 8 (1970) 221. Tree 15-35 m, 30-45 cm g; bark greyish brown, lenticellate. Branchlets initially with a dense cover of fulvous stellate hairs, soon glabrous, greyish black, sparsely lenticellate; terminal buds ovoid, 3 by 2 mm, scales ovate-acute. Stipules ovate- acute, 1-2 by 1 mm. Leaves coriaceous, 10-14 by 4-6 cm (index 2.2-2.6), broadest at or slightly below the middle; surfaces discolorous, above glabrous, greenish grey to dark chocolate-brown, dull, underneath with a thin cover of yellowish brown to fulvous adpressed, minute stellate hairs; base acute, top bluntly acute; midrib and nerves thin, flat on both sides; nerves 8-10 pairs, sub- parallel, at an angle of 45-50°, arcuating but not anastomosing towards the margin; reticulation fine, subscalariform to almost irregular, obscure; petiole glabrous, 6-10 mm, 1-2 mm g, adaxially flat to furrowed. Inflorescence male or androgy- nous, densely fulvous woolly pubescent; bracts and bracteoles narrowly ovate-acute, 2-3 by 1 mm. Male rachis c. 5 cm, 14% mm g; 3 flowers mostly solitary, filaments 3-4 mm, anthers 0.30-0.35 mm long, pistillode subglobose, c. 1 mm g. Androgy- nous rachis 5-6 cm, 1-14 mm g; female flowers solitary or in clusters of 2-3, staminodes rudimen- tary, styles 3-4, terete, c. 144 mm. Ripe cupule sessile to 1-14 cm stalked, broadly cup-shaped, 14%-2 cm long, 4-4% cm g; rim thick, slightly incurved, covering 14-14 part of the fruit; wall woody, 3-5 mm thick, outside densely greyish brown tomentose by adpressed stellate hairs; scales obscure, ovate-acute, adpressed, imbricate but concentrically set. Ripe fruit depressed ovoid- globose, 24-3 cm long, 3-4 cm g, glabrous except the depressed-umbonate top, dark chocolate- brown, base rounded to conical, scar 21-3 cm @; wall woody, 5-7 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: East New Guinea (Highlands Districts). Ecol. Forests at 1000-2500 m, on badly drained clayey soil. Fl. May-Sept., fr. Sept._May. 38. Lithocarpus lauterbachii (VON SEEMEN) MARK- GR. Bot. Jahrb. 59 (1924) 69; A. Camus, Chénes 3 (1954) 672, t.; SOEPADMO, Reinwardtia 8 (1970) 253. — Quercus lauterbachii VON SEEMEN, Bot. Jahrb. 23, Beibl. 57 (1897) 54; in K. Sch. & Laut. Fl. Schutzgeb. (1901) 264, t. 4: A-E. — Synaedrys lauterbachii (VON SEEMEN) Ko1pz. Bot. Mag. Tokyo 30 (1916) 196. — L. solanicarpa MARKGR. Bot. Jahrb. 59 (1924) 67; A. Camus, Chénes 3 (1954) 681, t. Tree 12-36 m, 35-80 cm g; bark grey-brown. Branchlets initially densely fulvous-tomentose by adpressed stellate hairs, later glabrous, dark grey- ish brown, smooth to sparsely lenticellate; terminal bud ovoid, 2-3 by 2 mm, scales ovate. Stipules not seen. Leaves thin-coriaceous, (8—)10—-13(-16) by (4-)5-—6(-8) cm (index 1.8-2.8), broadest at or below the middle; surfaces discolorous, above glabrous, pale to dark chocolate-brown, under- neath densely glaucous to pale grey tomentose by adpressed stellate hairs; base acute, margin re- curved, top bluntly and abruptly 14-1 cm acumi- nate; midrib and nerves thin, prominent beneath, flat above; nerves (7—)8—-9(—10) pairs, subparallel, at an angle of 45-50°, arcuating but not anasto- mosing towards the margin; reticulation lax, fine, subscalariform, distinct beneath; petiole 44-1 cm, 1144-2 mm g, glabrous, adaxially flat to furrowed. Inflorescence male or androgynous, densely ful- vous stellate hairy; bracts and bracteoles ovate, 1-114 by %4-1 mm. Male rachis 6-15 cm, 2 mm 2; 6 flowers in clusters of 3, filaments 3-4 mm, anthers 0.35 mm long, pistillode subglobose, longi- tudinally compressed, c. 1 mm g. Androgynous rachis 5-7 cm, 144-2 mm g; female flowers soli- tary, staminodes rudimentary, styles 3, conical, 1-114 mm, connate at the base. Ripe cupule %- 1144 cm stalked, deeply and broadly cup-shaped, 1144-2144 cm long, 3-4 cm g; rim thin, dentate, covering %4-'4 part of the fruit; wall woody, inside densely silky tomentose, outside densely fulvous stellate hairy; scales thick, adpressed, im- bricate but concentrically set. Ripe fruit depressed subglobose, 214-234 cm long, 3-344 cm g, gla- brous, chocolate-brown, top rounded and depres- sed-umbonate at the centre, base rounded, scar strongly convex, c. 24% cm g; wall woody, 5—7 mm thick, for the greater part free from the cupule. Distr. Malesia: New Guinea (locally found in NW. parts, rather common in the eastern parts). Ecol. In primary, occasionally also in secon- dary forests, at 300-2400 m. Fi. July—Febr., fr. March—Oct. Uses. Wood locally used for building construc- tion and fencing. 39. Lithocarpus pallidus (BL.) REHD. J. Arn. Arb. 1 (1919) 129; A. Camus, Chénes 3 (1954) 679, t. 382: 11-21; SoppADMO, Reinwardtia 8 (1970) 265. — Quercus pallida Bu. Bijdr. (1826) 524; Fl. Jav. Cupul. (1829) 12, t. 4, 5; Mia. Fl. Ind. Bat. 1, 1 (4856) 851; A.DC. Prod. 16, 2 (1864) 84; OUDEM. Natuurk. Verh. Kon. Akad. 11 (1865) 11, t. 6; WENziIG, Jahrb. Bot. Gart. Berl. 4 (1886) 225; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 57, t. 53 A; K. & V. Bijdr. 10 (1904) 35; Koorp. Atlas 1972] FAGACEAE (Soepadmo) 349 1 (1913) t. 52; Backer & BAKH. f. FI. Java 2 (1965) 7. — Quercus pseudomolucca var. rostrata Bi. Mus. Bot. 1 (1850) 291. — Quercus pseudo- molucca var. pallida (BL.) Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 108. — Pasania pallida (BL.) Orrst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83; S. Moore, J. Bot. 63 (1925) Suppl. 114. — Synaedrys pallida (BL.) Koibz. Bot. Mag. Tokyo 30 (1916) 197. Tree 10-20 m tall. Branchlets initially densely fulvous woolly pubescent by stellate hairs, later subglabrous, greyish brown, sparsely lenticellate; terminal buds ovoid-ellipsoid, 5-8 by 2-3 mm, scales narrowly ovate to linear, 3-5 by 1-2 mm. Stipules linear to subulate, 8-11 by 1-144 mm, rather long persistent. Leaves coriaceous, (10—)12- 16(-19) by (3-)4-5(-7) cm (index 24-31%), broadest at the middle; surfaces discolorous, above pale chocolate-brown, glabrous, dull to glossy, underneath densely yellowish tomentose by ad- pressed stellate hairs; base rounded-acute, top abruptly acute to 1-2 cm acuminate; midrib prom- inent on both sides, sometimes pinkish coloured; nerves (11—)12-14(—16) pairs, prominent beneath, impressed above, subparallel, at an angle of 45- 50°, arcuating but not anastomosing towards the margin; reticulation dense, fine, scalariform, dis- tinct beneath; petiole 6-10 mm, 1—2 mm g, adaxi- ally flat. Male rachis 5-10 cm, 1-114 mm g, usually much branched; bracts and bracteoles narrowly ovate-acute to linear, 3-5 by 44-1 mm; ¢ flowers solitary or in clusters of 2-3, filaments 2-3 mm, anthers 0.3 mm long, pistillode subglobose, c. 1 mm @. Young fruit: staminodes rudimentary, styles 3, conical, 1-144 mm, connate. Ripe cupule sessile, saucer-shaped, 44-1 cm long, 3144-5 cm g; rim thick, covering the basal part of the fruit; wall woody, inside densely fulvous tomentose by simple hairs, outside densely fulvous stellate hairy; scales woody, conical, 2-3 mm long, free, imbricate. Ripe fruit depressed-subglobose, 1-2 cm long, 3%-4144 cm g, glabrous, dark chocolate-brown, top rounded and depressed-umbonate at the cen- tre, base rotundate, scar flat, 3-4 cm 9; wall woody, c. 3 mm thick, for the greater part free from the cupule. Distr. Malesia: S. Sumatra (ForRBEs, fr., 1881, from Mt Dempo), Java (rather common in the western parts, eastwards to Mts Slamet and Wilis). Ecol. Forests, 1300-2100 m. Fi. May-June, /r. Aug.—Jan. Note. Female inflorescence not known. 40. Lithocarpus conocarpus (OUDEM.) REHD. J. Arn. Arb. 1 (1919) 123; A. Camus, Chénes 3 (1954) 754, t. 403: 1-9; So—EPADMO, Reinwardtia 8 (1970) 231. — Quercus conocarpa OUDEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 206; A.DC. Prod. 16, 2 (1864) 93; OupEM. Natuurk. Verh. Kon. Akad. 11 (1865) 18, t. 10; WENziIG, Jahrb. Bot. Berl. 4 (1886) 230; Hook. f. Fl. Br. Ind. 5 (1888) 612; K1nc, Ann. R. Bot. Gard. Calc. 2 (1889) 61, t. 56 A; K. & V. Bijdr. 10 (1904) 50; CORNER, Ways. Trees (1940) 302, f. 96, pl. 49; BACKER & BAKH. f. Fl. Java 2 (1965) 8. — Cyclo- balanus conocarpa (OUDEM.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Pasania cono- carpa (OUDEM.) SCHOTTKY, Bot. Jahrb. 49 (1913) 357, pro auct. Oerst. citato; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 430. — Synaedrys conocarpa (OUDEM.) Korpz. Bot. Mag. Tokyo 30 (1916) 191. — L. conocarpa ssp. malaccensis A. CAMUS, Bull. Soc. Bot. Fr. 90 (1943) 200; Chénes 3 (1954) 757, t. 403: 10-19, atque ssp. euconocarpa. Tree 10-45 m, 10-90 cm g; bark greyish brown, lenticellate. Branchlets initially densely greyish brown tomentose with stellate hairs, later subgla- brous with a few or many lenticels; terminal buds ovoid, 3-4 by 2-3 mm, scales narrowly ovate. Stipules ovate, 2-3 by 144 mm. Leaves coriaceous, rigid, (6—)8-12(-14) by (2—)3-4(-5\4) cm (index (2.2—)2.5—3 (-4)), broadest at or above the middle; surfaces discolorous, above dark greyish to choco- late-brown, by sparse erect stellate hairs, dull to glossy, underneath with a dense cover of yellowish brown to rufous adpressed and erect stellate hairs; base acute, margin recurved, top bluntly acute to 1-114 cm acuminate; midrib strongly prominent on both sides, densely pubescent with erect stellate hairs; nerves (9—)10-12(—15) pairs, prominent be- neath, impressed above, subparallel, at an angle of 45-70°, arcuating and anastomosing near the mar- gin, densely pubescent by erect stellate hairs; reticu- lation lax, coarse, subscalariform, distinct be- neath; petiole densely stellate hairy, 14-1 cm long, 1-2 mm g, terete or adaxially flat. Inflorescence male, androgynous or mixed, densely yellowish brown to fulvous stellate hairy; bracts and brac- teoles narrowly ovate, 1144-24% by %4-144 mm. Male rachis 10-17 cm, 1-14% mm g; 3 flowers in clusters of 3, filaments 3-4 mm, anthers 0.35 mm long, pistillode globose, c. 1 mm g. Androgynous or mixed rachis 7-18 cm, 1144 mm g; female flowers solitary, rarely in clusters of 2-3, staminodes rudi- mentary, styles 3, conical, 1-144 mm. Ripe cupule 3-8 mm stalked, cup-shaped, 2-6 mm long, 1.6—2 cm g; rim thin, covering the basal part of the fruit; lamellae 6-7, concentric, denticulate, outside densely yellowish brown to rufous stellate hairy. Ripe fruit ovoid-conical, 1.6-2.4 cm long, 1.7-2 cm g, densely greyish tomentose, subglabrescent, top long-acuminate, base rounded, scar concave, 1-1.2 cm g; wall woody, c. 1 mm thick, greater part free from the cupule. Distr. Malesia: Sumatra (rare, scattered), Ma- lay Peninsula (rare, scattered), Java (W. parts, rare), Borneo (common, especially in Sabah). Ecol. Forests, from sea-level up to 1800 m. Fi. July—March, fr. April—Oct. DocTERS VAN LEEUWEN (Ned. Kruidk. Arch. 51, 1941, 134) recorded a leaf gall caused by a gall-midge from Java. 41. Lithocarpus atjehensis HATUS. ex SOEPADMO, Reinwardtia 8 (1970) 217. Shrub. Branchlets densely fulvous tomentose by adpressed stellate and erect woolly tuft-hairs, blackish brown, brittle; terminal buds ovoid- globose, 3 by 2 mm, scales ovate. Stipules nar- rowly ovate-acute, 2 by 1 mm. Leaves thick- 350 coriaceous, rigid, brittle, (6-)8—10(-12) by (3—)4~ 5(—7) cm (index 1.7—2), broadest below the middle; surfaces discolorous, above dark chocolate-brown, more or less glossy, sparsely pubescent by woolly tuft-hairs especially on midrib and nerves, beneath pale greyish brown, densely tomentose by ad- pressed stellate hairs interspersed with woolly tuft- hairs; base rounded to subcordate, top sharply 1-144 cm acuminate; midrib prominent on both sides; nerves 12-14 pairs, prominent beneath, im- pressed above, subparallel, at an angle of 60-70°, arcuating and anastomosing near the margin; reticulation coarse, subscalariform, distinct be- neath; petiole 7-10 mm, 2 mm g, adaxially flat, densely stellate hairy. Male rachis 2-4 cm, 2 mm @, densely tomentose by erect stellate hairs; bracts and bracteoles ovate, 1-144 by %4-1 mm; ¢ flowers solitary, stamens 10-12, filaments 2-2% mm, anthers 0.30 mm long, pistillode subglobose, c. 1 mm g. Young infructescence 3-5 cm, 3 mm g, carrying 5-10 obconical young cupules. Female flowers (seen in a young fruit) solitary or in clusters of 2-3, staminodes rudimentary, styles 3, conical, 2-24% mm, recurved. Young cupule sessile, cup- shaped, c. 1 cm long, 2 cm g; rim thin, covering + 1/3 part of the fruit; wall thin, woody, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous tomentose by stel- late hairs; lamellae 5—6, thin, concentric, denticu- late. Young fruit ovoid-conical, c. 114% cm long and g, densely fulvous tomentose by adpressed simple and stellate hairs, top acute, base rotundate, scar concave, c. 0.7 cm 9; wall woody, c. 4% mm thick, for the greater part free from the cupule. Distr. Malesia: N. Sumatra (Atjeh: Gajo Lands: VAN STEENIS 8419, 8563, 9143). Ecol. Montane forest at 2500-3000 m. Fi. fr. Jan.—Febr. 42. Lithocarpus dasystachyus (MiQ.) REHD. J. Arn. Arb. 1 (1919) 124; A. Camus, Chénes 3 (1954) 798, t.; SOEPADMO, Reinwardtia 8 (1970) 234. — Quer- cus dasystachya Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 221; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 53, t. 45 B. — Pasania dasystachya (Mia.) ScHoTtTky, Bot. Jahrb. 49 (1913) 356. — Pasania winkleriana SCHOTTKY, I.c. 357. — Synaedrys dasy- stachya (M1Q.) Koipz. Bot. Mag. Tokyo 30 (1916) 194. — Quercus winkleriana (SCHOTTKY) MERR. En. Born. (1921) 216. — L. winkleriana (SCHOTTKY) A. Camus, Riviera Scient. 18 (1931) 42; Chénes 3 (1954) 800. Tree 5-18 m, 10-25 cm 9g; stilt-roots occasion- ally present; bark smooth to flaky, lenticellate, greyish brown. Branchlets densely velvety tomen- tose by greyish to reddish brown adpressed stel- late hairs and erect simple or stellate hairs; ter- minal bud ovoid-globose, 2-3 by 2 mm, scales ovate. Stipules ovate-acute to deltoid, 1-2 by 1% mm. Leaves thick-coriaceous, rigid, (12—)15-20(— 25) by (44%-)6-8(-10) cm (index 2-3), broadest about the middle; surfaces more or less discolor- ous, above subglabrous, dull to glossy, pale greyish brown, underneath densely yellowish to reddish brown, rarely greyish, tomentose by adpressed FLORA MALESIANA [ser. I, vol. 7? stellate and erect simple or stellate hairs; base rounded to acute, decurrent, margin recurved, top sharply 1-244 cmacuminate; midrib strongly prom- inent on both sides, densely pubescent by erect simple hairs, glabrescent; nerves 12-14 pairs, lax, prominent on both sides or impressed above, sub- parallel, at an angle of 60—70°, arcuating but not anastomosing towards the margin, densely pubes- cent by erect simple hairs, glabrescent above; retic- ulation lax, subscalariform, obscure; petiole dense- ly yellowish brown pubescent, 8-15 mm, 2-3 mm g, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely yellowish brown to rufous tomentose by erect simple and adpressed stellate hairs; bracts and bracteoles ovate or deltoid, 0.7-1 by 0.5-0.7 mm. Male rachis 10-25 cm, 1-2 mm g; ¢ flowers in clusters of 3, rarely solitary, perianth 5—6-lobed, stamens 10-12, filaments 2-244 mm, anthers 0.25 mm long, pistil- lode subglobose, c. 1 mm @. Androgynous or mixed rachis 10-30 cm, 2 mm g; female flowers solitary or rarely in clusters of 2-3, staminodes rudimen- tary, styles 3-6, conical, 1-2 mm. Ripe cupule 2-5 mm stalked, saucer- to cup-shaped, 5-7 mm long, 13-18 mm 9g; rim thin, covering c. % part of the fruit; wall thin, inside densely yellowish brown to rufous-tomentose by adpressed simple hairs, out- side densely yellowish brown to fulvous-tomentose by stellate hairs and erect simple hairs; lamellae 7-8, concentric, denticulate. Ripe fruit ovoid- conical, 13-15 mm long, 10-13 mm g, densely yellowish brown to fulvous tomentose by adpressed simple hairs, subglabrescent, pale chocolate-brown, top abruptly acuminate, base rotundate, scar deep- ly concave, c. 4% cm g; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (scattered in all parts). Ecol. Forests up to 750 m, common in secon- dary peat-swamp and heath forest, or in primary lowland forest on poor sandy, badly drained pod- zolized soil. F/. Sept._Febr., fr. March—July. Notes. Three more or less distinct forms may be recognized, viz one which occurs exclusively in peat-swamp forest, characterized by its large leaf with greyish brown indumentum, a form which is apparently confined to heath forest with a smaller leaf and less tomentum on the leaf and fruit, and one which occurs in forest on a better drained soil having a large leaf densely set with yellowish to reddish brown, erect simple hairs, and with strong- ly prominent venation. Several intermediates have, however, been collected. Records of L. dasystachyus from the Philippines are referred to L. mindanaensis. 43. Lithocarpus caudatifolius (MERR.) REHD. J. Arn. Arb. 1 (1919) 123; A. Camus, Chénes 3 (1954) 796, t.; SOEPADMO, Reinwardtia 8 (1970) 226. — Quercus caudatifolia MERR. Philip. J. Sc. 3 (1908) Bot. 324; EL_mer, Leafl. Philip. Bot. 3 (1910) 939; Merr. En. Philip. 2 (1923) 26. — Quercus minahassae KoorD. ex ELMER, Leaf. Philip. Bot. 3 (1910) 941; Koorp. in Koord.- Schum. Syst. Verz. 3 (1914) 28. — Synaedrys caudatifolia (MERR.) Koipz. Bot. Mag. Tokyo 30 1972] (1916) 190. — L. minahassae (KOORD. ex ELMER) Reup. J. Arn. Arb. 10 (1929) 133; A. CAmus, Chénes 3 (1954) 1162. — Quercus bulusanensis Eimer, Leafl. Philip. Bot. 10 (1939) 3736. — L. bulusanensis (ELMER) A. CAMus, Not. Syst. 13 (1948) 265; Chénes 3 (1954) 764, t. Tree 5—30 m, 10-80 cm g; buttresses 0.3 m tall, 0.6 m out, 5 cm thick; bark scaly, greyish brown. Branchlets densely yellowish brown to fulvous vel- vety tomentose by soft erect simple and stellate hairs, sparsely lenticellate; terminal bud ovoid, 2-3 by 2 mm, scales ovate. Stipules ovate to linear, 2-6 by 1 mm. Leaves thin-coriaceous to papery, (6—)10-14(-17) by (2—)3-6(-9) cm (index (2—)2.5 —3(—4)), broadest about the middle; surfaces more or less discolorous, above glabrous except the mid- rib and nerves, dull to glossy, pale chocolate- brown to greyish brown, underneath densely ful- vous velvety tomentose by adpressed stellate and erect simple and stellate hairs; base acute to cune- ate, rarely rounded-acute, occasionally asymmet- rical, top bluntly to sharply 1-244 cm caudate- acuminate, rarely acute; midrib and nerves prom- inent beneath slightly so above; nerves (7—)8— 10(-11) pairs, lax to dense, parallel, at an angle of 40-60°, arcuating but not anastomosing towards the margin; reticulation lax, fine, subscalariform, obscure; petiole 4-144 cm, 1-2 mm 4g, terete or adaxially flat. Inflorescence male, androgynous or mixed, densely greyish brown to fulvous tomentose by soft adpressed and erect simple and stellate hairs; bracts and bracteoles ovate-acute, 0.7—1 by 0.7 mm. Male rachis 10-30 cm, 1-2 mm g, simple and axillary or much-branched and subterminal; 3 flowers in clusters of 3—5, rarely solitary, peri- anth 6-8-lobed, stamens 10-14, filaments 3-4 mm, anthers 0.25-0.35 mm long, pistillode globose, c. 1 mm g. Androgynous or mixed rachis 6-25 cm, 1-2 mm g; female flowers solitary, staminodes rudi- mentary, styles 3, conical, 0.7-1 mm, recurved. Ripe cupule sessile to 3-5 mm stalked, cup- to saucer-shaped, 3—6 mm long, 1.2—2 cm g; rim thin, covering the basal part of the fruit; lamellae 5-6, concentric, denticulate; densely fulvous tomentose by simple and stellate hairs. Ripe fruit ovoid-conical to depressed ovoid-globose, 9-17 mm long, 12-19 mm g, densely greyish brown fulvous tomentose by adpressed stellate hairs, top rounded-acute, base truncate, scar concave to convex, 8-10 mm @; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (common in Sabah, rare in SE. Kalimantan), Philippines (Luzon, Min- danao, and several other islands). Ecol. Primary and secondary forest, up to 1350 m, on ridges or in swampy places near rivers, on yellowish sandy clayey soil or basalt-derived soil. Fl. Sept._June, fr. May—Jan. Note. Lithocarpus menadoensis (KOORD.) SOE- PADMO may belong here: see under the doubtful species. 44. Lithocarpus ewyckii (KORTH.) REHD. J. Arn. Arb. 10 (1929) 132; A. Camus, Chénes 3 (1954) 759, t.404: 11-19; SopEPADMo, Reinwardtia 8 (1970) FAGACEAE (Soepadmo) 351 240. — Quercus ewyckii KorTH. Kruidk. (1844) 212,t.46: 1-20; A.DC. Prod. 16, 2 (1864) 94; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 68, t. 62 A: 1-2. — Cyclobalanus ewyckii (KORTH.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80. — Quercus ewyckii var. latifolia KING ex HOOK. f. FI. Br. Ind. 5 (1888) 614; KinGc, Ann. R. Bot. Gard. Calc. 2 (1889) 68, t. 62 A: 3-6. — Pasania ewyckii (KorTH.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 431, incl. var. latifolia, l.c. 432. — Pasania lam- ponga var. ewyckioides GAMBLE, I.c. 425. — Synae- drys ewyckii (KORTH.) Korpz. Bot. Mag. Tokyo 30 (1916) 191. — L. pseudolamponga A. CAMUs, Chénes, Atlas 3 (1949) 110, t. 504; Chénes 3 (1954) ile Tree 10-30 m, 20-90 cm g; buttresses 0.6—1.2 m tall, 0.9-2.4 m out, 7-10 cm thick; bark smooth to scaly, greyish brown to reddish brown. Branchlets initially densely fulvous tomentose by adpressed stellate hairs, later subglabrous, greyish to dark brown, sparsely lenticellate; terminal buds ovoid- ellipsoid, 3-5 by 1-2 mm, scales narrowly ovate to linear. Stipules narrowly ovate to linear, 5 by 1-2 mm, soon caducous. Leaves thin-coriaceous, (6—)12-15(-18) by (2-)4-6(-7) cm (index 2-3), broadest about the middle; surfaces discolorous, above glabrous, glossy, pale to dark chocolate- brown, underneath densely yellowish to greyish brown tomentose by adpressed stellate hairs; base acute, rarely rounded, top bluntly to sharply 44-2 cm acuminate; midrib and nerves prominent be- neath, slightly so above; nerves (12—)13—15(-16) pairs, dense, parallel, at an angle of 45-60°, ar- cuating but not anastomosing towards the margin; reticulation dense, fine, scalariform, distinct be- neath; petiole 4-114 cm, 1-144 mm g, adaxially flat to furrowed. Inflorescence male, androgynous or mixed, densely brownish stellate hairy; bracts and bracteoles narrowly ovate-acute, 1-2 by %-1 mm. Male rachis 10-15 cm, 1-2 mm 9g; ¢ flowers in clusters of 3 or solitary, stamens 10-12, fila- ments 3—5 mm, anthers 0.30-0.50 mm long, pistil- lode subglobose, 1-144 mm g. Androgynous or mixed rachis 7-15 cm, 1-2 mm 9; female flowers solitary, rarely in clusters of 2-3, staminodes rudi- mentary, styles 3-4, conical, 1-144 mm long, con- nate. Ripe cupule 4%4—1 cm stalked, cup- to saucer- shaped, 14-1 cm long, 2-3 cm 9; rim thin, covering the basal part of the fruit; lamellae thin, 6-8, con- centric, densely pale brown stellate hairy. Ripe fruit ovoid, 2-2% cm through, densely greyish to- mentose, glabrescent, pale brown, glossy, top ab- ruptly acuminate, base rounded, scar deeply con- cave, 1-144 cm g; wall bony, 1-1 cm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (most parts), Malay Peninsula (Perak, Pahang, Selangor, Malacca, Jo- hore; Singapore), Borneo (common in all parts, also Nunukan I.). Ecol. In primary, also secondary and swamp forest up to 1800 m, on ridges or in rocky places. Fertility seems to be irregular. Note. The flowers have been recorded as having a musty fragrance. 352 45. Lithocarpus cantleyanus (KING ex Hook. /f.) Reup. J. Arn. Arb. 10 (1929) 132; A. Camus, Chénes 3 (1954) 709, t.; SoEPADMO, Reinwardtia 8 (1970) 225. — Quercus cantleyana KING ex HOOK. f. Fl. Br. Ind. 5 (1888) 613; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 64, t. 59. — Pasania cant- leyana (KING ex Hook. f.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 434. — Synaedrys cantleyana (KING ex Hook. f. )Kombz. Bot. Mag. Tokyo 30 (1916) 190. Tree 10-36 m, 10-120 cm g; bark greyish brown, fissured to scaly, lenticellate. Branchlets initially strongly ribbed under the leaf-insertion, densely yellowish brown tomentose by adpressed stellate hairs, later terete, subglabrous, sparsely lenticel- late; terminal buds ovoid, 2—4 by 2—3 mm, scales narrowly ovate-acute. Stipules ovate to linear- acute, 3-5 by 1-3 mm, soon caducous. Leaves thick-coriaceous, rigid, (10—)14-18(-24) by (4-) 5—6(—8) cm (index 2-3), broadest at or above the middle; above glabrous, glossy chocolate-brown to olive-green, beneath densely yellowish brown to pale glaucous tomentose by adpressed stellate hairs; base acute to cuneate, decurrent, margin recurved, top abruptly acute to %-1% cm acu- minate; midrib strongly prominent on both sides; nerves (12—)14-15(-16) pairs, prominent beneath, impressed above, parallel, at an angle of c. 60°, arcuating but not anastomosing towards the mar- gin; reticulation lax, fine, scalariform, distinct be- neath; petiole 1-144 cm, 1-244 mm g, shallowly furrowed. Inflorescence male, female or androgy- nous, densely yellowish brown stellate hairy; bracts and bracteoles narrowly ovate to linear, 2-214 by 0.7-1 mm. Male rachis 10-20 cm, 1144-2 mm 9g; 6 flowers in clusters of 3, perianth 5—7-lobed, stamens 10-12, filaments 3-4 mm, anthers 0.30- 0.35 mm long, pistillode subglobose, 1-114 mm g. Female or androgynous rachis 10-15 cm long, 2 mm g; 2 flowers solitary or in clusters of 3—5, staminodes rudimentary, styles 3, conical, 1-14% mm, connate. Ripe cupule c. Y cm stalked, cup- shaped, 14-1 cm long, 144-214 cm g; rim thin, covering ¥4—1,4 part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely stellate hairy; lamel- lae 5-8, concentric, entire or denticulate. Ripe fruit depressed-ovoid, 1-14% cm long, 144-2 cm 9, densely pale greyish brown simple-tomentose, gla- brescent, top abruptly acute, base cordate, scar concave, c. 1 cm g; wall bony, thinner than 1 mm, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (Perak, com- mon, Dindings, Johore; Singapore), Borneo (Sara- wak, Sabah, rare in E. Kalimantan). Ecol. In primary forest up to 850 m, occasion- ally on river-bank on sandy clayey soil or ultra- basic soil. Fl. May—Dec., fr. Febr.—Nov. 46. Lithocarpus meijeri SOEPADMO, Reinwardtia 8 (1970) 260. Tree 12—42 m, 20-100 cm g; buttresses spreading, rounded; bark smooth to deeply fissured or scaly, greyish to reddish brown. Branchlets initially dense- ly fulvous stellate hairy, later subglabrous, grey- FLORA MALESIANA [ser. I, yale” ish to blackish brown, sparsely lenticellate; ter- minal bud ovoid, 3 by 2 mm, scales narrowly ovate-acute. Stipules linear-acute to subulate, 3-4 by 4-1 mm. Leaves coriaceous, (11—)13-16(—25) by (4-)5—7(-11) cm (index 2-3), broadest at or above the middle; surfaces discolorous, above glabrous, chocolate-brown, glossy, underneath densely greyish to fulvous tomentose by adpressed stellate hairs; base cuneate, decurrent, margin re- curved, top bluntly acute to abruptly %-1 cm acuminate; midrib and nerves prominent beneath, flat above, often pinkish beneath; nerves 11-13 pairs, subparallel, at an angle of c. 45°, arcuating but not anastomosing towards the margin; reticu- lation dense, fine, subscalariform, obscure; petiole subglabrous, 1-114 cm, 1144-21 mm g, thickened and rugose at the base, shallowly furrowed. In- florescence male, female, androgynous or mixed, densely yellowish brown to fulvous stellate hairy; bracts and bracteoles narrowly ovate-acute, 0.7— 1.5 by 0.3-0.5 mm. Male rachis 10-25 cm, 2 mm @; 3 flowers in clusters of 3-7, rarely solitary, perianth 6-8-lobed, recurved, stamens 12-15, fila- ments 3—5 mm, anthers 0.30 mm long, pistillode globose, 1-14 mm g. Female, androgynous or mixed rachis 10-15 cm, 2—3 mm @g; @ flowers soli- tary, rarely in clusters of 2-3, staminodes rudimen- tary, styles 3, conical, 0.7-1 mm. Ripe cupule ses- sile, cup-shaped, 1-114 cm long, 2-2% cm g; rim thin, covering the basal part of the fruit; wall woody, inside densely greyish brown tomentose by adpressed simple hairs, outside densely fulvous stellate hairy, lamellae 6-8, concentric, entire or undulate, sometimes obscure. Ripe fruit depressed ovoid, 0.6-1.5 cm long, 2—3 cm @, densely silvery grey tomentose by adpressed simple hairs, top abruptly acuminate or depressed-apiculate, base truncate, scar deeply concave, 144-2 cm g; wall woody, 2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Borneo (Sarawak, Brunei, Sa- bah). Ecol. Primary, mixed Dipterocarp forest on low ridges up to 1000 m, occasionally also in secondary forest; on yellowish brown, sandy loamy soil or blackish basalt-derived soil. F/. Aug.—Dec., fr. March—Oct. 47. Lithocarpus daphnoideus (BL.) A. CAMus, Ri- viera Scient. 18 (1932) 40; Chénes 3 (1954) 711, t. 396: 1-9; SopPADMO, Reinwardtia 8 (1970) 234. — Quercus daphnoidea BL. Fl. Jav. Cupul. (1829) 28, t. 16; A.DC. Prod. 16, 2 (1864) 96; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 60, t. 54 A; K. & V. Bijdr. 10 (1904) 45; Koorp. Atlas 1 (1913) t. 56; BACKER & BAKH. f. Fl. Java 2 (1965) 8. — Quercus nitida BL. Mus. Bot. 1 (1850) 294, non RAFIN. 1838, nec MART. & GAL. 1843, incl. var. grisea; A.DC. Prod. 16, 2 (1864) 95; WENZIG, Jahrb. Bot. Gart. Berl. 4 (1886) 235; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 91; VON SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 11. — Cyclobalanus daphnoidea (Bu.) OERsT. Vid. Medd. Naturh. For. Kjobn. 8 (1867) 81. — Cyclobalanus nitida (BL.) OrrstT. /.c. 81. — Quercus poculiformis VON SEEMEN, 1972] FAGACEAE (Soepadmo) 353 Bot. Jahrb. 27, Beibl. 64 (1900) 13; K. & V. Bijdr. 10 (1904) 46; Koorp. Atlas 1 (1913) t. 61. — Synaedrys daphnoidea (BL.) Korbz. Bot. Mag. To- kyo 30 (1916) 191. — Synaedrys poculiformis (VON SEEMEN) Korpz. /.c. 192. — Pasania daphnoidea (BL.) S. Moore, J. Bot. 63 (1925) Suppl. 114. — L. poculiformis (VON SEEMEN) A. CAMus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 763. — L. nitida (BL.) A. CAMus, Bull. Soc. Bot. Fr. 92 (1945) 255; Chénes 3 (1954) 1116, t. — L. sara- wakensis E. F. WARB. Kew Bull. (1936) 19; A. Camus, Chénes 3 (1954) 697, t. Tree 10-24 m, 25-65 cm g; bark smooth, grey, lenticellate. Branchlets initially densely fulvous- tomentose with adpressed stellate hairs, later sub- glabrous, greyish to blackish brown, sparsely to densely lenticellate; terminal buds ovoid, 3-4 by 2-24% mm, scales narrowly ovate-acute to linear, 2%-3 by 0.7-1 mm. Stipules linear-acute, 3-4 by 1 mm. Leaves thin-coriaceous, (8—)10—13(—16) by 4-5 cm (index 2.43.4), broadest at the middle; surfaces discolorous, above glabrous, pale greyish green, beneath densely greyish to yellowish brown tomentose by adpressed stellate hairs; base acute, margin recurved, top bluntly acute to 1-2 cm acum- inate; midrib prominent on both sides; nerves (7—)8-10(—12) pairs, more or less prominent be- neath, subparallel, at an angle of 45—60°, arcuating but not anastomosing towards the margin; retic- ulation lax, fine, subscalariform, distinct beneath; petiole glabrous, 1-1.2 cm, 1 mm g, furrowed. Inflorescence male or androgynous, densely greyish brown stellate hairy; bracts and bracteoles nar- rowly ovate to linear, 1-2 by 4% mm. Male rachis 10-15 cm, 1-114 mm g; ¢ flowers in clusters of 3, rarely solitary, perianth 5—6-lobed, stamens 10-12, filaments 214-3 mm, anthers 0.25-0.30 mm long, pistillode subglobose, c. 1 mm g. Androgynous rachis 10-15 cm, 1-2 mm g; female flowers soli- tary, rarely in clusters of 3, staminodes rudimen- tary, styles 3-4, conical, c. 1 mm, connate. Ripe cupule sessile, obconical, cup-shaped, 1-14 cm long, 2-24% cm g; rim thin, covering the basal part of the fruit; wall woody, inside densely silvery grey tomentose by adpressed simple hairs, outside densely fulvous to greyish stellate hairy; lamellae 8-10, rounded, prominent, concentric. Ripe fruit ovoid-conical, 2-3 cm long, 1.8—2.5 cm g, densely fulvous tomentose by simple adpressed hairs, top abruptly acute, base rotundate, scar flat to convex, 1-144 cm g; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (scattered), Malay Peninsula (Perak, Kelantan, Pahang), W. Java (several collections from Bantam and Priangan Prov.), Borneo (scattered in Sarawak and W. Kali- mantan). Ecol. Forests, up to 1350 m. Fl. Dec.—April, fr. May-Nov. Note. Quercus dolichocarpa VON SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 14. — Synaedrys doli- chocarpa (VON SEEMEN) Koipz. Bot. Mag. Tokyo 30 (1916) 191. — L. dolichocarpa (VON SEEMEN) REuD. J. Arn. Arb. 1 (1919) 125, may belong here. The type and the other collections mentioned by VON SEEMEN are, however, very inadequate for a definite conclusion. The leaves and cupules are smaller and the fruit is longer than those of L. daphnoideus, otherwise they are very similar. 48. Lithocarpus philippinensis (A. DC.) Reup. J. Arn. Arb. 1 (1919) 129; A. Camus, Chénes 3 (1954) 723, t.; SOEPADMO, Reinwardtia 8 (1970) 267. — Quercus philippinensis A. DC. Prod. 16, 2 (1864) 97; WeENziIG, Jahrb. Bot. Gart. Berl. 4 (1886) 230, excl. syn.; MERR. Philip. J. Sc. 3 (1908) Bot. 328; En. Philip. 2 (1923) 29. — Cyclobalanus ‘philippensis’ (A. DC.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Quercus caraballoana F.-VILL. Nov. App. (1880) 209. — Quercus wen- zelii MERR. Philip. J. Sc. 10 (1915) Bot. 267; En. Philip. 2 (1923) 31. — Synaedrys philippinensis (A. DC.) Korpz. Bot. Mag. Tokyo 30 (1916) 192. — Synaedrys wenzelii (MERR.) Koipz. lc. 193. — Quercus rizalensis MERR. Philip. J.Sc.13 (1918) Bot. 272; En. Philip. 2 (1923) 30. — L. wenzelii (MerRR.) REHD. J. Arn. Arb. 1 (1919) 132; A. Camus, Chénes 3 (1954) 1111. — L. rizalensis (Merr.) ReHD. J. Arn. Arb. 10 (1929) 133; A. Camus, Chénes 3 (1954) 812, t. 420: 1-9. — L. oligocarpa A. Camus, Bull. Soc. Bot. Fr. 81 (1934) 816; Chénes 3 (1954) 762, t. 405: 13-16. Shrub or tree 4-20 m, 18-30 cm g. Branchlets initially densely rufous tomentose by scurfy stel- late hairs, later glabrous, whitish grey to blackish brown, sparsely to densely lenticellate; terminal buds ovoid, 3 by 1 mm, scales ovate. Stipules narrowly ovate-acute or deltoid, 14-1 by 4% mm, soon caducous. Leaves thin-coriaceous, rigid, (5-)7-9(-1014) by (214—-)3-4(-6) cm (index 1.6- 2.5), broadest about the middle; surfaces discolor- ous, above glabrous, glossy, pale to dark choco- late-brown, rarely greyish green, underneath dense- ly glaucous tomentose by adpressed stellate hairs; base acute to cuneate, decurrent, margin recurved, top bluntly 14-214 cm acuminate-caudate; midrib prominent on both sides; nerves (7—)8—10(—12) pairs, prominent beneath, flat to impressed above, subparallel, at an angle of 45—50°, arcuating and anastomosing near the margin; reticulation dense, fine, scalariform, distinct beneath; petiole glabrous, 14-114 cm, 1-2 mm 9g, adaxially flat or furrowed. Inflorescence male or mixed, densely fulvous to rufous stellate hairy, simple and axillary or much- branched and subterminal; bracts and bracteoles ovate, 14-1 by 14 mm. Male rachis 5-15 cm, 144 mm 9g; ¢ flowers in clusters of 3, rarely solitary, stamens 10-12, filaments 2-3 mm, anthers 0.20- 0.30 mm long, pistillode globose, c. 1 mm @. Mixed rachis 5-10 cm, 114—mm 9; female flowers solitary, staminodes rudimentary, styles 3, conical, c. 1 mm. Ripe cupule sessile to 3 mm stalked, deeply cup-shaped, 8-13 mm long, 15-25 mm 9g; rim thin, covering 14—% part of the fruit; wall woody, thin, inside densely whitish grey simple tomentose, outside densely greyish to fulvous stellate hairy; lamellae 5-6, denticulate, concen- tric, thin to prominent. Ripe fruit ovoid, 1144-2 cm in size, densely greyish tomentose by adpressed stellate hairs; top rounded-acuminate, base round- 354 ed, scar strongly convex to flat, 9-15 mm 9g; wall woody, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, Mindanao, Samar, Leyte). Ecol. Forests at 1000-2000 m. Fi. Dec.-July, fr. Oct.-Nov. 49. Lithocarpus apoensis (ELMER) REHD. J. Arn. Arb. 1 (1919) 123; A. CAMus, Chénes 3 (1954) 708, t.; SOEPADMO, Reinwardtia 8 (1970) 217. — Quer- cus apoensis ELMER, Leaf]. Philip. Bot. 3 (1910) 945, incl. var. ulayan, I.c. 946; MERR. En. Philip. 2 (1923) 25. Tree 10-25 m, 30-130 cm g; bark pale grey, finely fissured. Branchlets initially densely yellow- ish brown stellate hairy, later glabrous, greyish brown, densely lenticellate; terminal buds ovoid- globose, 3-4 by 2-3 mm, scale broadly ovate. Stipules ovate, 214 by 1144 mm. Leaves thick- coriaceous, rigid, (12—)15—17(—22) by (5—)6—8(-10) (index 1.6—2.8), broadest at or below the middle; surfaces discolorous, above glabrous, pale to dark greyish brown, glossy, beneath densely glaucous- tomentose by adpressed-stellate hairs; base round- ed-acute to cuneate, margin recurved, top bluntly acute to 1-114 cm acuminate; midrib prominent on both sides; nerves (7—)8-9(-10) pairs, promi- nent beneath, flat above, subparallel, at an angle of 45-60°, arcuating but not anastomosing to- wards the margin; reticulation fine, subscalariform, obscure to distinct beneath; petiole glabrous, 1—2 cm, 3-4 mm 9, terete or adaxially flat. Inflorescence male or mixed, densely yellowish brown stellate hairy; bracts and bracteoles ovate-acute, 1-114 by 2/3 mm. Male rachis 10-25 cm, 2-3 mm @g; 6 flowers solitary or in clusters of 2-3, filament 214-3 mm, anthers 0.35—0.50 mm long, pistillode subglobose, 114-2 mm g. Mixed rachis 5-20 cm, 2-3 mm @; female flowers solitary or in clusters of 3, staminodes rudimentary, styles 3-4, conical, c. 1 mm long. Ripe cupule subsessile, cup-shaped, 1.2— 1.5 cm long, 2-214 cm g; rim thin, covering c. 4 part of the fruit; wall woody, rather thick, outside densely greyish brown stellate hairy; lamellae 7-9, thin, entire, concentric. Ripe fruit ovoid, 2-2 cm through, densely greyish tomentose by adpressed, simple hairs, top gradually rounded-apiculate, base truncate, scar flat to slightly convex, 1-2 cm @g; wall woody, 2-3 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, Mindanao, Catanduanes, Panay). Ecol. Forests, up to 1700 m. Fl. March-Dec., fr. April-Sept. 50. Lithocarpus glutinosus (BL.) SOEPADMo, Rein- wardtia 8 (1970) 243. — Quercus glutinosa BL. Mus. Bot. 1 (1850) 304; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 88; A. Camus, Chénes 3 (1954) 1158. — Pasania glutinosa (BL.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. — Quercus koordersii VON SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 16. — Quercus zschokkei ELMER, Leaf]. Philip. Bot. 3 (1910) 944; MeErr. En. Philip. FLORA MALESIANA [ser. I, vols ?? 2 (1923) 31. — Quercus copelandii Elmer, Leafl. Philip. Bot. 6 (1913) 1984; MeErr. En. Philip. 2 (1923) 27. — L. copelandii (ELMER) REHD. J. Arn. Arb. 1 (1919) 124; A. Camus, Chénes 3 (1954) 705, t. 390: 13-25. — L. zschokkei (ELMER) REHD. J. Arn. Arb. 1 (1919) 132; A. Camus, Chénes 3 (1954) 702, t. — L. koordersii (VON SEEMEN) MarkGr. Bot. Jahrb. 59 (1924) 66; A. CAmus, Chénes 3 (1954) 713, t. 392: 11-19. Tree 20 m, 60-70 cm g; bark greyish brown. Branchlets initially densely yellowish brown stel- late hairy, later glabrous, greyish, sparsely to densely lenticellate; terminal buds ovoid, 0.7—1 by 114-2 mm, scale narrowly ovate. Leaves thin- coriaceous, (8—)12—-17(-20) by (4-)5-6(-8) cm (index 2-3), broadest at or slightly above the middle; above glabrous, dull to glossy, greyish green to chocolate-brown, beneath densely greyish tomentose by adpressed stellate hairs; base acute, margin recurved, top abruptly acute to 1-3 cm acuminate; midrib strongly prominent beneath, flat above; nerves (7—)8—-9(—10) pairs, prominent beneath, impressed above, subparallel, at an angle of 40-50°, arcuating but not anastomosing to- wards the margin; reticulation fine, subscalari- form, obscure; petiole glabrous, 8-15 mm, 1-3 mm @g, adaxially flat. Inflorescence male or andro- gynous, densely yellowish brown stellate hairy; bracts and bracteoles narrowly ovate to linear, 1-2 by %4-'4 mm. Male rachis 10-15 cm, 1-144 mm 9; ¢ flowers in clusters of 3—5, filaments 3-5 mm, anthers 0.35 mm long, pistillode globose, 0.7 —1 mm g. Androgynous rachis 5-10 cm long, 1 mm g; female flowers solitary, staminodes rudimen- tary, styles 3-4, conical, c. 1 mm, recurved. Ripe cupule sessile, deeply cup-shaped, 114-2 cm long, 214-3 cm g; rim thin, covering 4%—14 part of the fruit; wall woody, densely fulvous tomentose; basal part ridged or tuberculate, upper part with 5-7 obscure lamellae. Ripe fruit subhemispherical, 114-134 cm long, 2-214 cm g, densely fulvous tomentose by adpressed stellate hairs, top rounded -umbonate, base truncate, scar flat, c. 2 cm 9; wall woody, c. 2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Mindanao), Cele- bes (Northern Peninsula and Central part). Ecol. Forests, up to 900 m. Fertility mainly in the second half of the year. Notes. Quercus molucca RuMPH. ex L. 1753 may belong here, but the original description and figure are not recognizable, the later name gluti- nosa is here accepted. In his original description of Quercus glutinosa, BLUME mentioned Java as the type locality. In the Rijksherbarium (Leyden) there are 3 specimens which seem to be duplicates of one collection, bear- ing 2 different labels and localities. Two of these specimens (H.L.B. 901.310-309 and 310) were inscribed by BLUME as Quercus glutinosa and said to be from Java, and the third (901.310—380) bears REINWARDT’S original green label with Tondano (N. Celebes) as the locality and is dated October 1821. As BLUME never was in Celebes, and no other 1972] FAGACEAE (Soepadmo) 355 collections confirm his record of Quercus glutinosa from Java, it is concluded that two of REINWARDT’S specimens were mislabelled and that all came from Celebes, where L. glutinosus has repeatedly been collected by KooRDERS, EyMA, and others. 51. Lithocarpus solerianus (VIDAL) REHD. J. Arn. Arb. 1 (1919) 131; A. Camus, Chénes 3 (1954) 979, t. 467: 1-15; SopPADMO, Reinwardtia 8 (1970) 279. — Quercus molucca (non RuMmpH. ex L.) BLANCO, FI. Filip. (1837) 726. — Quercus concen- trica (non Lour.) BLANCO, FI. Filip. ed. 2 (1845) 502. — Quercus reinwardtii (non KORTH.) F.-VILL. Nov. App. (1880) 207. — Quercus costata var. convexa (non BL.) NAVES in Blanco, FI. Filip. ed. 3 (1880-83) t. 441. — Quercus soleriana VIDAL, Rey. Pl. Vasc. Filip. (1886) 261; Merr. Philip. J. Sc. 3 (1908) Bot. 327; E_mer, Leaf]. Philip. Bot. 6 (1913) 1983; Merr. Sp. Blanc. (1918) 121; En. Philip. 2 (1923) 30. — Quercus clementiana (non KinG ex Hook. f.) MERR. Philip. J. Sc. 1 (1906) Suppl. 41. — Synaedrys soleriana (VIDAL) Ko1pDz. Bot. Mag. Tokyo 30 (1916) 193. Tree 10-15 m, 20-30 cm g; bark brown. Branch- lets initially densely greyish brown adpressed stellate hairy, later glabrous, dark blackish brown, sparsely to densely lenticellate; terminal buds ovoid, 14-1 by 14 mm, scales ovate. Stipules ovate to deltoid, 14-1 by 14 mm. Leaves thin-coriaceous, (5—)8-12(-16) by (3-)4-6(-714) cm (index 1!4- 214), broadest about the middle; above glabrous, dull to glossy, pale to dark chocolate-brown, underneath densely greyish to glaucous tomentose by adpressed stellate hairs; base rounded and abruptly acute to attenuate-acute, top abruptly, bluntly acuminate to sharply 1-114 cm caudate- acuminate; midrib and nerves thin, prominent beneath, flat above; nerves (9—)10(—12) pairs, parallel, at an angle of 45—50°, arcuating but not anastomosing towards the margin; reticulation dense, fine, scalariform, distinct beneath; petiole glabrous, blackish coloured, 7-15 mm, 1-114 mm @, shallowly furrowed. Inflorescence male, andro- gynous, or mixed, densely greyish brown stellate hairy; bracts and bracteoles ovate-acute, 0.7—1 by 14 mm. Male rachis 10-15 cm, 1 mm g, simple and axillary or more commonly much-branched and subterminal; ¢ flowers in clusters of 3, stamens 10-12, filaments 2-3 mm, anthers 0.35 mm long, pistillode globose, c. 1 mm g. Androgynous or mixed rachis c. 10 cm, 1-2 mm 9; female flowers solitary, staminodes rudimentary, styles 3, conical, more or less recurved, 0.7-1 mm. Ripe cupule subsessile, cup-shaped, 14-1 cm long, 2-2!4 cm g; rim thin, entire, erect, covering 14-14 part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous to glaucous stellate hairy; lamellae 5-7, thin, concentric or spiral at the base, minutely denticulate. Ripe fruit ovoid, 1.5—2.2 cm long, 1.8-2 cm g, glabrous, dark chocolate-brown, top gradually rounded acute, base truncate, scar flat, c. 144 cm g; wall bony, thinner than 1 mm, for the greater part free from the cupule; cotyledons flat- convex. Distr. Malesia: Philippines (Luzon, Mindanao, perhaps also Mindoro). Ecol. Forests, at 700-1200 m. Fl. Aug.-May, fr. April-Oct. 52. Lithocarpus mindanaensis (ELMER) REHD. J. Arn. Arb. 1 (1919) 128; A. Camus, Chénes 3 (1954) 1110, t.; SogpADMO, Reinwardtia 8 (1970) 261. — Quercus philippinensis (non A.DC.) MERR. Bull. Bur. For. Philip. 1 (1903) 16. — Quercus celebica (non MiQ.) VON SEEMEN in Perkins, Fragm. Fl. Philip. (1904) 41. — Quercus acumina- tissima MERR. Philip. J. Sc. 3 (1908) Bot. 326, non A. DC. 1864. — Quercus ‘tasystachya’ (non Mig.) Evmer, Leaf]. Philip. Bot. 3 (1910) 938; Merr. En. Philip. 2 (1923) 27. — Quercus minda- naensis ELMER, Leaf]. Philip. Bot. 3 (1910) 942, quoad nom.; MeErRR. En. Philip. 2 (1923) 28. — Synaedrys acuminatissima (MERR.) KoIDZz. Bot. Mag. Tokyo 30 (1916) 190. Small tree, c. 10 cm g@; bark rough, light grey. Branchlets initially densely yellowish to fulvous tomentose by stellate hairs, later glabrous, greyish to dark brown, with sparse minute lenticels; terminal buds ovoid, 1 by 1 mm, scales ovate- acute. (Stipules not seen.) Leaves chartaceous, (714-)10-12(-1414) by (214—-)3-5(-6) cm (index (2-)2'4(-3)), broadest at the middle; surfaces discolorous, above dark chocolate-brown, sparsely stellate-pubescent, glabrescent, dull or glossy, beneath densely yellowish tomentose by adpressed stellate hairs; base attenuate-acute or rounded- acute, sometimes decurrent and asymmetrical, margin sometimes undulate, top bluntly acute to sharply 1-2 cm acuminate, tip usually oblique; midrib thin but prominent on both sides; nerves (8-)9-10(-12) pairs, thin, prominent beneath, flat above, subparallel, at an angle of 50-60°, arcuating but not anastomosing towards the margin; reticulation fine, dense, scalariform, obscure, or sometimes rather distinct beneath; petiole 7-13 mm, 1-2 mm @g, sparsely pubescent, shallowly furrowed. Inflorescence male or andro- gynous, densely yellowish to fulvous stellate-hairy; bracts and bracteoles ovate-acute, c. 1 by 2/3 mm. Male rachis 10-20 cm, 1-114 mm 9g; ¢ flowers in clusters of 3, filaments 3-314 mm, anthers 0.3 mm long, pistillode subglobose, c. 1 mm @. Androgy- nous rachis 10-20 cm, 2 mm g; female flowers solitary, staminodes rudimentary, styles 3, conical, c. 1 mm, recurved. Ripe cupule sessile, saucer- shaped, %-'4 cm long, 2-244 cm 9g; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely adpresssed pubescent by sim- ple hairs, outside densely fulvous tomentose by adpressed stellate hairs; lamellae thin, denticulate, concentric or somewhat spiral. Ripe fruit ovoid- conical, 2-214 cm through, glabrous, pale to dark chocolate-brown, top rounded-acute to acuminate, base rotundate, scar flat, c. 114 cm 9; wall woody, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Mindanao, com- mon, Leyte, scattered). 356 Ecol. In forests, at 300-1200 m, common below 600 m. F/. Oct.—March, fr. May-Sept. Note. ELMER 11922, the only specimen cited by ELMER (1910) when he proposed the new name Quercus mindanaensis for Quercus acuminatissima MerR., non A. DC., belongs to L. caudatifolius. 53. Lithocarpus bennettii (M1Q.) REHD. J. Arn. Arb. 1 (1919) 123; A. Camus, Chénes 3 (1954) 742, t.; SOEPADMO, Reinwardtia 8 (1970) 220. — Quer- cus bennettii Mriq. Fl. Ind. Bat. 1, 1 (1856) 857; A. DC. Prod. 16, 2 (1864) 94; Hook. f. Fl. Br. Ind. 5 (1888) 613; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 64, t. 58 A; CorNER, Ways. Trees (1940) 301. — Quercus miqueliana SCHEFF. Nat. Tijd. N. I. 31 (1870) 360; ibid. 32 (1871) 416. — Cycloba- lanus bennettii (M1Q.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 375. — Pasania bennettii (Miq.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 433. — Synaedrys bennettii (M1Q.) Kor1pz. Bot. Mag. Tokyo 30 (1916) 190. Tree 10-35 m, 20-80 cm g; buttresses low, up to 3 m out, stilt-roots occasionally present; bark grey-brown, smooth to fissured, lenticellate. Branch- lets initially densely yellowish brown to rufous- tomentose by adpressed stellate hairs, later gla- brous, greyish to blackish brown, sparsely lenti- cellate; terminal buds ovoid-ellipsoid, 114-3 by 1-114 mm, scales ovate to linear-acute. Stipules linear-acute, 214-314 by 2/3-1 mm. Leaves thin -coriaceous, (7—)9-12(-15) by (214—-)3-5(-6) cm (index (2—)214-3(-4)), broadest at or rarely above the middle; surfaces discolorous, above glabrous, dull to glossy, usually dark chocolate-brown, underneath densely greyish to glaucous tomentose by adpressed stellate hairs; base attenuate-acute, top bluntly or sharply 14-2 cm acuminate; midrib prominent on both sides; nerves (9—)10—12(—13) pairs, thin but prominent beneath, parallel, at an angle of 45-60°, arcuating and anastomosing towards the margin; reticulation fine, subscalari- form, obscure; petiole glabrous, 14-114 cm, 1-3 mm 9g, furrowed. Inflorescence male or androgy- nous, densely greyish to yellowish brown stellate hairy; bracts and bracteoles ovate, 1-114 by 1%4- 1%, mm. Male rachis 10-20 cm, 1-144 mm g, simple and axillary or much-branched and subter- minal; 3 flowers solitary or in clusters of 2-3, stamens 10-12, filaments 2-3 mm, anthers 0.35 mm, pistillode globose, c. 1 mm 9g. Androgynous rachis 10-15 cm, 1-114 mm g; female flowers solitary, staminodes rudimentary, styles 3, conical, c. 1mm. Ripe cupule 14 cm stalked, saucer-shaped, Y%—'y cm long, 1-2 cm g; rim thin, covering the basal part of the fruit; wall thin, inside densely greyish brown tomentose by adpressed simple hairs, outside densely greyish adpressed stellate hairy; lamellae 5—7, thin, concentric, minutely denticulate. Ripe fruit ovoid-conical, 1-114 cm through, usually slightly longer than wide, gla- brous, dark purplish brown, top gradually round- ed-acute, base rotundate, scar deeply concave, c. 1-114 cm g; wall bony, thinner than 1 mm, for the greater part free from the cupule; cotyledons flat-convex. FLORA MALESIANA [ser. I, vol. 7? Distr. Malesia: Sumatra (scattered), Banka (common), Malay Peninsula (Kelantan, Negri Sembilan, frequent in Johore; Singapore), Borneo (scattered, mainly N. of the equator). Ecol.Primary, swampy, heath forest or mixed Dipterocarp forest, up to 1000 m, on various types of soil. F/. March—Nov., fr. July—Febr. Note. Philippine records of L. bennettii were based on specimens of L. solerianus or L. sulitii. 54. Lithocarpus confertus SOEPADMO, Reinwardtia 8 (1970) 229. Tree 12-30 m, 20-100 cm g; buttresses up to 0.9 m tall, 144 m out, 10 cm thick; bark greyish brown, densely lenticellate, occasionally scaly. Branchlets initially densely yellowish brown to fulvous stellate hairy, later subglabrous, dark greyish brown, densely or sparsely lenticellate; terminal buds ovoid-ellipsoid, 2-3 by 1-2 mm, scales narrowly ovate. Stipules narrowly ovate- acute to lanceolate, 2—3 by 1 mm. Leaves thin-coria- ceous, (6—)9-10(-13) by (3—-)4-5(-7) cm (index (114-)2-214 (-3)), broadest at or below the middle; surfaces more or less discolorous, above glabrous, dull to glossy, greyish green to chocolate-brown, beneath densely set with greyish to yellowish brown (rarely rufous) adpressed stellate hairs; base rounded and abruptly acute, margin recurved, top bluntly and abruptly acute to 1-114 cm acu- minate; midrib prominent on both sides; nerves (8-)9-10(-11) pairs, prominent beneath, impres- sed above, subparallel, at an angle of 40—50°, arcuating and anastomosing towards the margin; reticulation dense, scalariform, distinct beneath; petiole subglabrous, 14-1 cm, 1-144 mm g, fur- rowed. Inflorescence male, androgynous or mixed, densely greyish to yellowish brown stellate hairy; bracts and bracteoles narrowly ovate to linear, 114-5 by 14-1 mm. Male rachis 10-15 cm, 144 mm @, in dense paniculate clusters, subterminal; g flowers in clusters of 3, filaments 2-244 mm, anthers 0.25—0.30 mm long, pistillode subglobose, c. 1 mm g. Androgynous or mixed rachis 5-15 cm, 114 mm 9; female flowers solitary or in clusters of 2-3, staminodes rudimentary, styles 3, terete, 1- 214 mm, strongly recurved. Ripe cupule subsessile, saucer-shaped, 2-5 mm long, 114-134 cm 9g; rim thin, covering the basal part of the fruit; wall thin, inside densely fulvous-tomentose by adpressed simple hairs, outside densely greyish brown to fulvous stellate hairy; lamellae 5—6, thin, concen- tric, minutely denticulate. Ripe fruit depressed- ovoid, 1-114 cm long, 114-2 cm g, glabrous, chocolate-brown to purplish brown, top abruptly acuminate, base rounded, scar concave, 1-114 cm @; wall bony, thinner than 1 mm, for the greater part free from the cupule . Distr. Malesia: Borneo (common on Mt Kina- balu). Ecol. Forests, at 900-1800 m, usually on ultra- basic soil. F/. Sept.—April, fr. July—April. 55. Lithocarpus bullatus HATus. ex SOEPADMO, Reinwardtia 8 (1970) 223, f. 11. Tree 9-25 m, 15-40 cm 9g; bark fissured to scaly, 1972] FAGACEAE (Soepadmo) ad | dark greyish brown to reddish brown. Branchlets initially densely yellowish brown to rufous-tomen- tose by adpressed, stellate hairs, later subglabrous, pale yellowish brown to dark greyish brown, fis- sured, sparsely lenticellate; terminal buds ovoid, 1-114 by 1 mm, scales narrowly ovate-acute to lanceolate. Stipules lanceolate, 1-144 by 4%-'4 mm. Leaves thick-coriaceous, rigid, strongly bul- late, (5—)7-9(-11) by (214-)3-4(-5) cm (index 1.3-2.8), broadest at or slightly below the middle; above glabrous, glossy, olive-green, beneath dense- ly pale grey-brown tomentose by adpressed stel- late hairs; base acute, margin strongly recurved, top bluntly acute to 14-1 cm acuminate, rarely rounded-emarginate; midrib strongly prominent on both sides; nerves 9-10 pairs, strongly promi- nent beneath, impressed above, parallel, at an angle of 60—-80°, arcuating but not anastomosing towards the margin; reticulation lax, subscalari- form, obscure; petiole glabrous, 3-5 mm, 1-14 mm 9, adaxially flat. Inflorescence male, female or mixed, densely yellowish brown stellate hairy; bracts and bracteoles ovate-acute, 1 by 14-24 mm. Male rachis 5-10 cm, 1-144 mm g; ¢ flowers solitary or in clusters of 3, filaments 2!4-3!4 mm, anthers 0.5—0.7 mm long, pistillode subglobose, c. 1 mm g. Female or mixed rachis 5-10 cm, 2 mm g; © flowers solitary, staminodes rudimentary, styles 3-4, conical, 2-214 mm, more or less connate. Ripe cupule subsessile, cup-shaped, 14-1 cm long, 114-1%4 cm 9g; rim thin, covering 4-14 part of the fruit; wall woody, inside densely yellowish brown to silvery tomentose by adpressed simple hairs, outside densely greyish brown stellate hairy; lamellae 5-8, concentric, minutely denticulate. Ripe fruit ovoid-conical, c. 114 cm through, top abruptly acute, base rotundate, scar flat, 8-10 mm @, glabrous, dark chocolate-brown; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak, rare, Mt Kinabalu, common). Ecol. Forests, at 800-4000 m, on sandy acid or ultra-basic soil. Fertility throughout the year. 56. Lithocarpus rigidus SOEPADMO, Reinwardtia 8 (1970) 275. Tree 6-18 m, 30 cm g. Branchlets sturdy, with short internodes and thick nodes, glabrous, dark- brown, sparsely lenticellate; terminal buds ovoid- ellipsoid, 7-10 by 3—5 mm, scales narrowly ovate, 5-7 by 2-3 mm, initially with sparse minute stellate hairs, soon glabrescent. Stipules ovate- acute to lanceolate, 6-8 by 3-4 mm. Leaves thick- coriaceous, rigid, (8—)14-16(-18) by (5—)6—8(—10) cm (index 114-2), broadest at or above the middle; surfaces concolorous, pale yellowish brown, above glabrous, glossy, underneath sparsely tomentose by minute, adpressed, stellate hairs; base acute, decurrent to the base of the petiole, margin strongly recurved, top bluntly rounded-acute or rounded; midrib broad, prominent on both sur- faces, nerves 8-11 pairs, prominent beneath, flat to impressed above, subparallel, at an angle of 50-80°, arcuating but not anastomosing towards the margin; reticulation fine, subscalariform to irregular, obscure on both sides; petiole 7-10 mm, 5-6 mm @, adaxially flat, glabrous. Male rachis (not fully developed) c. 5 cm, 3-4 mm g, sparsely stellate-tomentose; bracts linear, 5-7 by 1-2 mm, bracteoles ovate, 3-4 by 2 mm; ¢ flowers in clusters of 3. Female flowers (on young cupule) with peri- anth 6—lobed, staminodes 12, rudimentary, styles 3, conical, 1-2 mm, recurved and glabrous near the top. Fruiting rachis (not fully developed) 15-20 cm, 14-1 cm g, sparsely fulvous stellate-tomentose; young cupule sessile, mostly in clusters of 3, densely fulvous stellate-tomentose; scales minute, adpressed, concentrically arranged. Young fruit ovoid, glabrous, chocolate-brown, top rounded- acuminate. Distr. Malesia: Borneo (Sabah: Mts Kinabalu and Tambuyokan). Ecol. Everwet forests, at 1500-2500 m. FI. March—April, fr. July. Notes. Fully developed inflorescences, cupules, and fruits not known. Vegetatively distinct by its extremely rigid thick- coriaceous leaves, with long-decurrent base, and by its sturdy branchlets, inflorescence and infruc- tescence. 57. Lithocarpus nodosus SOEPADMO, Reinwardtia 8 (1970) 262. Tree 10-20 m, 20 cm g; bark brownish, smooth. Branchlets glabrous, with thick nodes, dark grey- ish brown, sparsely lenticellate; terminal bud ovoid, 1-3 by 14-1 mm, scales narrowly ovate, initially densely rufous lepidote, glabrescent. Stip- ules linear-acute, c. 2 by 14 mm. Leaves thick- coriaceous, rigid, (6—)8-10(-12) by (3—)4-5(-6) cm (index (1.6—)2(-3)), broadest at the middle; surfaces slightly discolorous, above glabrous, glossy, chocolate-brown, underneath densely yel- lowish brown tomentose by adpressed, minute, stellate hairs; base acute, top abruptly acute or 1-114 cm acuminate; midrib broad, prominent above, flattish beneath, nerves 8-10 pairs, thinly prominent beneath, flat above, subparallel, at an angle of 30—50°, arcuating but not anastomosing towards the margin; reticulation fine, subscalari- form, obscure; petiole glabrous, 8-12 mm, 2-3 mm 9, adaxially flat. Inflorescence male or andro- gynous, solitary or in dense paniculate clusters, densely or sparsely tomentose by stellate hairs; bracts narrowly ovate, 1-2 by 14-24 mm, bracte- oles ovate, c. 14 by 14 mm. Male rachis 3-10 cm, 1-2 mm @; ¢ flowers in clusters of 3, filaments 2-3 mm, anthers 0.25 mm long, pistillode subglo- bose, 24-1 mm @. Androgynous rachis c. 5 cm, 2 mm @; female flowers solitary or rarely in clusters of 2-3, staminodes rudimentary, styles 3, conical, 1-2 mm, connate. Young cupule solitary or in clusters of 2-3; scales adpressed, minute, concen- trically arranged. Ripe cupule sessile, solitary, cup-shaped, 24-1 cm long, 2-214 cm g; rim thin, covering 14-1, part of the fruit; wall woody, thin, inside densely greyish tomentose by adpressed simple hairs, outside densely fulyvous-tomentose by short stellate hairs; lamellae thin, 4-6, ob- 358 scure, minutely denticulate. Ripe fruit ovoid-glo- bose, 1-114 cm long, 134-2!4 cm 9, glabrous, pale chocolate-brown, top rounded-acute, base round- ed, scar deeply concave, c. 1 cm @; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak: Mts Mulu and Poi; Sabah: Mt Kinabalu). Ecol. Everwet forests, at 900-2400 m; S. 4706 was recorded from a limestone hill. F/. June-— March, fr. April—Oct. Note. Though the young cupules are sometimes in clusters of 2-3, only one is well-developed, the others being abortive. 58. Lithocarpus woodii (HANCE) A. CAMus, Ri- viera Scient. 18 (1932) 42; Chénes 3 (1954) 1173; SOEPADMO, Reinwardtia 8 (1970) 287. — Quercus woodii HANCE, J. Bot. 12 (1874) 240; Merr. Philip. J. Sc. 3 (1908) Bot. 326; En. Philip. 2 (1923) 31. — Synaedrys woodii (HANCE) Korpz. Bot. Mag. Tokyo 30 (1916) 193. — L. loheri A. Camus, Chénes, Atlas 3 (1949) 111, t.; Chénes 3 (1954) iW 2s Tree 5-30 m, 20-90 cm g. Branchlets initially densely fulvous to rufous stellate hairy, later gla- brous, greyish black, sparsely to densely lenticel- late; terminal buds ovoid, 1-114 by 0.7-1 mm, scales ovate. Stipules ovate to linear, 1-3 by 14 mm. Leaves thick-coriaceous, (6—)9—12(—17) by (4~)5—7 (-10) cm (index (1.3—)1.6—2(—2.2)), broadest at or below the middle; surfaces more or less discolor- ous, above glabrous, dull to glossy, pale to dark chocolate-brown, underneath densely glaucous tomentose by adpressed stellate hairs; base round- ed-acute to cuneate, margin recurved, occasion- ally undulate, top bluntly acute to bluntly or sharply 1-114 cm acuminate; midrib and nerves strongly prominent beneath, flat above; nerves 8-11 pairs, subparallel, at an angle of 45—50°, arcuating but not anastomosing towards the mar- gin; reticulation fine, lax, subscalariform, distinct beneath; petiole glabrous, 7-15 mm, 1-3 mm @g, adaxially flat. Inflorescence male or androgynous, densely yellowish to greyish brown stellate hairy; bract and bracteoles ovate to linear, 1-2 by 4 mm. Male rachis 10-15 cm, 1-2 mm @, much-branched, axillary or subterminal; ¢ flowers in clusters of 3 or solitary, perianth 5—6—-lobed, stamens 10-12, filaments 2-3 mm, anthers 0.35 mm long, pistillode subglobose, c. 1 mm @. Androgynous rachis c. 10 cm, 1-144 mm @g; female flowers solitary or rarely in clusters of 2-3, perianth 5—6—lobed, staminodes 10-12, rudimentary, styles 3, terete, 114-2 mm, recurved. Ripe cupule subsessile, cup- shaped, 24-1 cm long, 214-314 cm @; rim thin, covering c. 14 part of the fruit; wall woody, thin, inside densely pale yellowish brown tomentose by adpressed simple hairs, outside densely greyish to fulvous stellate hairy; lamellae 7-10, obscure or distinct, concentric, minutely denticulate. Ripe fruit subhemispherical, 1.5—2.2 cm long, 2.5—3.2 cm @, glabrous except near the umbo, pale yellowish brown to dark chocolate-brown, top rounded- umbonate, base rotundate, scar deeply concave FLORA MALESIANA [ser. [avGhky? to flat, 144-2 cm g; wall woody, c. 2 mm thick, for the greater part free from the cupule. Distr. Malesia: Philippines (Luzon, Mindanao, Samar). Ecol. Forests, at 1200-2300 m. F/. Aug.—June, fr. April-Nov. 59. Lithocarpus hatusimae SOEPADMO, Reinwardtia 8 (1970) 244. Tree 13-18 m, 30 cm g; bark smooth. Branchlets initially densely yellowish brown tomentose by adpressed stellate hairs, later subglabrous, dark greyish brown, sparsely lenticellate; terminal buds ovoid, 2-3 by 2 mm, scales ovate-acute. Stipules thick-coriaceous, ovate, 2-3 by 1-2 mm. Leaves thick-coriaceous, rigid, (8—)11-14(-16) by (3—)4-6 (-7) cm (index 2-3), broadest at or slightly below the middle; above glabrous, glossy, pale chocolate- brown, underneath with a thin cover of pale yellowish brown, adpressed, stellate hairs; base acute, rarely rounded, margin recurved, top acute to sharply 1-2 cm caudate; midrib strongly promi- nent on both sides; nerves prominent beneath, impressed above, (9—)11—12(—13) pairs, parallel, at an angle of 60—70°, arcuating but not anasto- mosing towards the margin; reticulation lax, sub- scalariform, distinct beneath; petiole glabrous, 1-114 cm, 1-3 mm g, furrowed. Inflorescence male, female, androgynous or mixed, densely yellowish brown stellate hairy; bract and bracteo- les ovate, 1-114 by 24 mm. Male rachis c. 10 cm, 1-114 mm g; ¢ flowers solitary or in clusters of 3, perianth 5—6—lobed, stamens 10-12, filaments 3-4 mm, anthers 0.30-0.35 mm long, pistillode subglobose, 1-114 mm g, densely yellowish brown pubescent. Female, androgynous or mixed rachis 5-10 cm long, 1-2 mm 9; female flowers solitary or in clusters of 3, perianth 6—lobed, staminodes 12, rudimentary, styles 3, conical, 144-2 mm, connate. Ripe cupule subsessile, cup-shaped, c. 14 cm long, 1.2-1.7 cm g; rim thin, covering c. 14 part of the fruit; wall woody, thin, inside densely pale greyish brown tomentose by adpres- sed simple hairs, outside densely greyish brown stellate hairy; lamellae 5-8, concentric, minutely denticulate. Ripe fruit ovoid-conical, 1-144 cm through, top abruptly acute, base cordate, scar concave, c. 1 cm 9g; wall bony, thinner than 1 mm, outside glabrous, glossy, chocolate-brown, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak: Mt Mulu; Sabah: Mt Kinabalu, common). Ecol. Primary forest, at 1200-2000 m, on ultra-basic soil. F/. June—March, fr. March—July. 60. Lithocarpus vidalii (F.—ViLL.) ReHpD. J. Arn. Arb. 10 (1929) 134; SoppADMO, Reinwardtia 8 (1970) 285. — Quercus vidalii F.-V1LL. Nov. App. (1880) 209; VIDAL, Sinopsis Atlas (1883) 41, t. 92: B; Evmer, Leaf]. Philip. Bot. 6 (1913) 1981; MERR. En. Philip. 2 (1923) 31. Tree 10-25 m, 30-40 cm g; bark greyish brown, scaly. Branchlets initially densely fulvous to rufous scurfy tomentose by adpressed stellate hairs, later subglabrous, greyish to chocolate-brown, sparsely 1972] to densely lenticellate; terminal buds ovoid-ellip- soid, 2-6 by 2-3 mm, scales ovate to lanceolate. Stipules ovate to lanceolate, 3-6 by 2-4 mm. Leaves coriaceous, rigid, (6—)9-12(-14) by (214-) 3—4(—S) cm (index (1.8—)2—3(—3.6)), broadest at or above the middle; above glabrous, glossy, pale to dark chocolate-brown, underneath with a thick cover of fulvous to rufous adpressed stellate hairs; base acute to cuneate, decurrent, top rounded to abruptly acute; midrib strongly prominent be- neath slightly so above; nerves (9—)10-11(—13) pairs, prominent beneath, impressed above, sut parallel, at an angle of 40—-45°, arcuating and anastomosing near the margin; reticulation fine, subscalariform, lax, distinct beneath; petiole sub- glabrous, 14-1 cm, 1!4-3 mm g, adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous stellate hairy; bract and bracteoles ovate, 1-114 by 24mm. Male rachis 7-15 cm, 1-2 mm g, usually much-branched; ¢ flowers in clusters of 3-7, filaments 2-3 mm, anthers 0.35 mm long, pistillode globose, c. 1 mm @. Androgynous or mixed rachis 5-10 cm, 1-2 mm @g; female flowers solitary or in clusters of 2-3, staminodes rudimen- tary, styles 3-6, terete, 1-114 mm, recurved. Ripe cupule subsessile, cup-shaped, 1 cm long, 2—2.2 cm 2; rim thin, covering 14-14 part of the fruit; wall woody, thin, inside densely dark chocolate-brown tomentose by adpressed stellate hairs, outside densely fulvous stellate hairy; lamellae 6-8, con- centric, denticulate. Ripe fruit ovoid-conical, 114— 2 cm long, 1!4-1.8 cm g, glabrous, chocolate- brown, top abruptly acute, base truncate, scar flat, c. 1.1 cm 9; wall woody, thinner than 1 mm, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, Mindanao, Palawan). Ecol. Forests, at 1100-1800 m. F/. Febr.—May, fr. June-Nov. 61. Lithocarpus reinwardtii (KoRTH.) A. CAMus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 726, t. 397: 1-8; SOEPADMO, Reinwardtia 8 (1970) 272. — Quercus reinwardtii KoRTH. Kruidk. (1844) 211; OupEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 205; A. DC. Prod. 16, 2 (1864) 92; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 63, t. 57 A. — Cyclobalanus reinwardtii (KORTH.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Pasa- nia reinwardtii (KORTH.) PRANTL in E. & P. Nat. Pfi. Fam. 3, 1 (1889) 55. — Synaedrys reinwardtii (KortH.) Korpz. Bot. Mag. Tokyo 30 (1916) 192. Tree of medium size; bark grey. Branchlets initially densely fulvous tomentose by adpressed stellate hairs, later subglabrous, dark grey, densely lenticellate; terminal buds ovoid, 114-2 by 1-114 mm, scales ovate-acute. Stipules ovate-acute, 2-3 by 1 mm. Leaves thin-coriaceous, rigid, (8—)12—14 (-16) by (214—)3-4(-5) cm (index 2.3-3.3), broad- est at or rarely slightly below the middle; sur- faces discolorous, above glabrous, glossy, dark- brown, underneath densely glaucous tomentose by adpressed stellate hairs; base acute to cuneate, margin recurved, top bluntly acute to 1-114 cm FAGACEAE (Soepadmo) 359 acuminate; midrib thin, prominent on both sides; nerves (10—)12—-15(—17) pairs, in between with shorter ones not reaching the margin, obscure, subparallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, areolate, obscure; petiole subglabrous, 1-2 cm, 1 mm @g, adaxially flat or furrowed. /nflores- cence male or androgynous, densely fulvous stel- late hairy; bract and bracteoles ovate, 1-114 by 4-24, mm. Male rachis c. 10 cm, 1 mm @;¢ flow- ers in clusters of 3, stamens 10-12, filaments 1— 1144 mm, anthers 0.2-0.25 mm long, pistillode subglobose, c. 1 mm g. Androgynous rachis 10-20 cm, 2-214 mm 9; female flowers solitary, perianth (5—)6(—7)-lobed, staminodes rudimentary, styles 4-5, conical, c. 1 mm, connate. Ripe cupule 14-1 cm stalked, cup-shaped, 3-5 mm long, 114-2 cmg; rim thin, covering 4—1% part of the fruit; wall woody, thin, outside densely fulvous to glaucous tomentose by adpressed stellate hairs; lamellae 6-8, thin, concentric. Ripe fruit ovoid-conical, 114-2 cm long, 1-114 cm g, densely fulvous to glaucous tomentose by adpressed simple hairs, top abruptly acute, base rotundate, scar deeply concave, c. 1 cm 9; wall woody, thinner than 1 mm, for the greater part free from the cupule. Distr. Malesia: Sumatra (Central part: Mt Melintang, near Pajakumbuh). Ecol. Forests, at 600-1200 m. F/. June—Sept., fr. Nov.—June. Note. Records from Indo-China, Malay Penin- sula and Borneo are not confirmed. 62. Lithocarpus sulitii SoEPADMO, Reinwardtia 8 (1970) 280. — Quercus wenzigiana (non KiNG ex Hook. f.) MERR. Philip. J. Sc. 1 (1906) Suppl. 41. — Quercus bennettii (non MiQ.) MERR. Philip. J. Sc. 3 (1908) Bot. 328, quoad FB 759, 781, WuIT- FORD 365 et WILLIAMS 705; En. Philip. 2 (1923) 25, quoad ditto et BS 32958. Tree 15-20 m, 25-80 cm g. Branchlets initially sparsely tomentose by fulvous, adpressed stellate hairs, later glabrous, dark greyish brown, sparsely lenticellate; terminal buds ovoid, 2 by 2 mm, scales ovate. Leaves thin-chartaceous, (6—)9—11(— 13) by (214-)4-5(-6) cm (index (2-)24%4(-2'4)), broadest about the middle; surfaces discolorous, above glabrous, dull, dark greyish brown, beneath pale chocolate-brown, densely tomentose by ad- pressed stellate hairs, subglabrescent; base round- ed-acute, top acute or more commonly bluntly 14-1 cm acuminate; midrib and nerves thin on both sides ; nerves (8—)9—10(—13) pairs, obscure, subparallel, at an angle of 45-70°, arcuating but not anastomosing towards the margin; reticulation fine, areolate, obscure; petiole 7-12 mm, 1-114 mm 9g, glabrous, adaxially flat. Inflorescence male, androgynous or mixed, sparsely greyish tomentose by adpressed stellate hairs; bracts and bracteoles ovate-acute, 24-1 by 14 mm. Male rachis 8-20 cm, 114 mm 9g, simple and axillary or much-branched and subterminal; flowers in clusters of 3, filaments 2-214 mm, anthers 0.25 —0.30 mm long, pistillode globose, 24-1 mm 9g. Androgynous or mixed rachis 10-15 cm, 2 mm 9g; female flowers solitary, 360 FLORA MALESIANA [ser. I, vol. 7? staminodes rudimentary, styles 3, conical, 1-114 mm long, more or less connate. Ripe cupule sessile, saucer-shaped, 5-6 mm long, 2-244 cm 9; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely glaucous stellate hairy; lamellae 4-5, thin, spiral. Ripe fruit ovoid, 2.3 cm long, 2—2.2 cm @, densely greyish tomentose by adpressed simple hairs, top rounded- acute, base rotundate, scar deeply concave, c. 1 cm @; wall woody, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, common, Mindoro, rare). Ecol. Forests, up to 600 m. F/. Sept.—Jan., fr. Febr.—June. Note. Other records of Quercus bennettii from the Philippines, partly belong to L. solerianus, partly to an undescribed species. 63. Lithocarpus bancanus (SCHEFF.) REHD. J. Arn. Arb. 10 (1929) 132; SoepADMO, Reinwardtia 8 (1970) 218. — Quercus bancana ScuEFF. Nat. Tijd. N. I. 31 (1870) 361; Kinc, Ann. R. Bot. Gard. Fig. 26. Lithocarpus bancanus (SCHEFF.) REHD. a. Habit, x 2%, b. ¢ flowers, x4, c. 2 flowers, x4, d. fruit and cupule, e. basal view of the cupule, f. basal view of the fruit showing the deeply concave, small scar, all nat. size (a—-b BURKILL 2509, c RASTINI 207, d~f RACHMAT Si Toroes 3357). 1972] FAGACEAE (Soepadmo) 361 Calc. 2 (1889) 67. — Quercus rajah HANCE, J. Bot. 16 (1878) 198; Kinc, Ann. R. Bot. Gard. Calc 2 (1889) 67, t. 61 B. — Quercus scyphigera var. riedelii KING, Ann. R. Bot. Gard. Calc. 2 (1889) 39. — Synaedrys bancana (SCHEFF.) KoIbDZz. Bot. Mag. Tokyo 30 (1916) 190. — Synaedrys rajah (HANCE) Korpz. l/.c. 192. — Pasania bancana (SCHEFF.) MARKGR. Bot. Jahrb. 59 (1924) 79, quoad nomen. — Pasania rajah (HANCE) S. Moore, J. Bot. 63 (1925) Suppl. 115. — L. rajah (HANCE) A. CAMus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 737, t. — L. scyphigera var. riedelii (KING) A. Camus, Chénes 3 (1954) 962. — Fig. 26. Tree 10-25 m, 20-60 cm go; bark smooth to finely scaly, yellowish to greyish brown. Branchlets initially densely fulvous to rufous stellate hairy, later glabrous, smooth, greyish to dark chocolate- brown; terminal buds ovoid, 14-1 by 1 mm, scales ovate-acute. Stipules deltoid, 44-1 by 4% mm. Leaves thick-coriaceous, (9—)12-14(-17) by (4-) 5—6(-7) cm (index 2-2.7), broadest about the middle; surfaces more or less discolorous, above glabrous, pale to dark chocolate-brown, glossy, beneath yellowish to greyish brown tomentose by adpressed, minute stellate hairs; base acute to cuneate, top bluntly to sharply 14-114 cm acumi- nate, rarely acute; midrib strongly prominent on both sides; nerves (9—)10-12(—16) pairs, obscure on both surfaces, subparallel, at an angle of 45- 60°, arcuating and anastomosing towards the margin; reticulation fine, irregular, obscure; peti- ole subglabrous, (144-)1(-1144) cm, 1-3 mm g, adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous to rufous adpressed stel- late hairy; bracts and bracteoles ovate, 14-1 by 14 mm. Male rachis 5-20 cm, 1 mm g; ¢ flowers solitary, filaments 244-314 mm, anthers 0.3-0.4 mm long, pistillode globose, c. 1 mm g. Androgy- nous or mixed rachis 10-16 cm, 114-2 mm g; female flowers solitary, staminodes rudimentary, styles 3, conical, 1-114 mm, recurved. Ripe cupule sessile, saucer-shaped, 2-5 mm long, 15-21 mm g; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely silvery grey tomentose by adpressed stellate hairs, outside densely yellow- ish to greyish brown adpressed stellate hairy; lamellae 4-8, concentric, entire. Ripe fruit ovoid, 16-18 mm long, 20-22 mm g, densely whitish grey tomentose by adpressed simple hairs, glabrescent, top abruptly rounded-acute, base rotundate, scar deeply concave, 8-10 mm g; wall woody, thinner than 1 mm, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Sumatra (scattered in the main- land), Banka and Belitung Is., Malay Peninsula (Pahang, Johore), Borneo (scattered in Sarawak, Brunei, Kalimantan). Ecol. Forests up to 250 m. Fi. April—Nov., /r. Dec.—March. Notes. When ScHEFFER described Quercus bancana he had specimens with flowers and young cupules, and apparently also some detached, ripe cupules. A year later (1871) he stated that the detached, ripe cupules on which he based his description of Quercus bancana, did not belong to the species, and he replaced it with another one, based on a specimen collected by TEYSMANN (HB 7639) from Pangkal Pinang in Banka, which, however, belongs to L. urceolaris. HANCE, who never saw the type of Quercus bancana SCHEFF. (1870), described Quercus rajah based on speci- mens collected from a tree cultivated in the Bogor Botanic Gardens, the seedlings (or seeds?) of which had been brought back by TEYSMANN from Banka. These specimens are exactly the same with SCHEFFER’S Quercus bancana (1870). Believing that SCHEFFER’S identification (1871) was correct, KING (1889) and most later authors, who never checked the type of Quercus bancana, referred all specimens of the true bancana to rajah, and the true urceo- laris to bancana. These two species are, however, perfectly different from one another by the charac- ters of the leaf and cupule (see the key). The material from New Guinea cited by MARK- GRAF under Pasania bancana, belongs to L. mega- carpus. 64. Lithocarpus ovalis (BLANCO) REHD. J. Arn. Arb. 1 (1919) 129; A. Camus, Chénes 3 (1954) 750, t.; SOEPADMO, Reinwardtia 8 (1970) 264. — Quercus glabra BLANCO, FI. Filip. (1837) 727, non THUNB. 1784. — Quercus ovalis BLANCO, FI. Filip. ed. 2 (1845) 502; A. DC. Prod. 16, 2 (1864) 97; F.- VILL. Noy. App. (1880) 208; Merr. Philip. J. Sc. 3 (1908) Bot. 325; Sp. Blanc. (1918) 120; En. Philip. 2 (1923) 29. — Quercus blancoi A. DC. Prod. 16, 2 (1864) 97; VIDAL, Sinopsis Atlas (1883) 41, t. 92: C. — Cyclobalanus ovalis (BLANCO) Orrst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Cyclobalanus blancoi (A.DC.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 376. — Synaedrys ovalis (BLANCO) Korpz. Bot. Mag. To- kyo 30 (1916) 192. Tree 15-25 m, 30-50 cm g. Branchlets glabrous, dark greyish brown, sparsely lenticellate; terminal buds globose, c. 1 by 1 mm, scales ovate-acute. Leaves thick-coriaceous, (9—)15—20(—23) by (414-) 7-8(-10) cm (index (1.6-)2.5(-3)), broadest about the middle; surfaces glabrous, above pale grey-green, glossy, beneath pale yellowish brown to glaucous; base acute to cuneate, top bluntly or sharply 1-2 cm acuminate; midrib prominent on both sides; nerves (11—)12—14(—16) pairs, obscure on both surfaces, subparallel, at an angle of 45—60°, arcuating but not anastomosing towards the margin; reticulation irregular, obscure; petiole 114-214 cm, 2-3 mm @, glabrous, adaxially flat. Inflorescence male or female, sparsely stellate hairy; bracts and bracteoles ovate-acute, 24-1 by %3 mm. Male rachis 15-30 cm, 1-2 mm g; ¢ flowers in clusters of 2-3, filaments 3-4 mm, anthers 0.3 mm long, pistillode subglobose, 1-114 mm @. Female rachis 15-23 cm, 114 mm g; @ flowers solitary, stamino- des rudimentary, styles 3-4, conical, 1-144 mm long. Ripe cupule sessile, cup-shaped, 5-6 mm long, 2-214 cm g; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely greyish tomentose by adpressed simple hairs, out- side densely greyish to fulvous adpressed stellate 362 hairy; lamellae 5S—6, thin, concentric, entire. Ripe fruit ovoid-conical, 2-2.4 cm long, 1.9-2.1 cm 9, densely greyish to brownish tomentose by ad- pressed simple hairs, top rounded-acute, base truncate, scar concave, 0.9-1 cm @; wall woody, thinner than 1 mm, for the greater part free from the cupule. Distr. Malesia: Philippines (Luzon, in various Provinces). Ecol. Lowland forest. F/. Nov.—April, fr. June— Aug. 65. Lithocarpus orbicularis SoEpADMO, Reinwardtia 8 (1970) 264. — Fig. 27. Shrub 1-3 m. Branchlets initially sparsely to- mentose by fulvous, adpressed stellate hairs, later glabrous, blackish brown; terminal buds ovoid, c. 1 by 24 mm, scales ovate. Stipules linear, c. 2 by 14 mm. Leaves thick-coriaceous, rigid, (2—)3-3'4 (414) by (2-)214(-3) cm (index 1-114), broadest about the middle; surfaces more or less concolor- ous, chocolate-brown, above glabrous, glossy, with some minute glands, underneath with a thin cover of minute, adpressed stellate hairs, subgla- brescent; base rounded to subcordate, top round- ed-emarginate; midrib thin, prominent on both sides; nerves 8-10 pairs, obscure, parallel, at an angle of c. 60°, arcuating but not anastomosing towards the margin; reticulation fine, areolate, obscure; petiole c. 3 mm, 2 mm g, glabrous, adax- ially flat. (Inflorescence not known.) Fruiting rachis (young) c. 3 cm, 5mm 9, carrying more than 10 subglobose young cupules. Cupule (not yet ripe) sessile, cup-shaped, c. 8 mm long, 2 cm g; rim thin, covering c. 1% part of the fruit; wall thin, woody, inside densely tomentose by adpressed simple hairs, outside densely fulvous stellate hairy, hairs minute, adpressed; lamellae thin, 5—6, con- centric. Fruit (not yet ripe) depressed-ovoid, 1 cm long, 124 cm g, glabrous, dark chocolate-brown; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: N. Sumatra (once collected on top of Goh Lembuh, Gajo Lands, Atjeh: VAN STEENIS 8977). Ecol. Montane forest, at 2800 m. Fr. 19-II-— 1937. Notes. Vegetatively distinct, recognizable by its thick-coriaceous, rigid, orbicular leaf with areolate reticulation and rounded-emarginate top. Inflorescences and ripe fruits are unknown. 66. Lithocarpus pusillus SoEPADMO, Reinwardtia 8 (1970) 270. Tree 7-25 m, 10-80 cm g; stilt-roots occasionally present, up to 1.2 m tall; bark finely scaly, greyish brown. Branchlets initially densely greyish brown adpressed stellate hairy, later subglabrous, greyish brown to greyish black, sparsely lenticellate or finely fissured; terminal buds ovoid, c. 1 by 1 mm, scales ovate. (Stipules not seen.) Leaves thin- coriaceous, rigid, (4-)6-10(—1214) by (1.7—)244-4 (—-5) cm (index (1.6—)2-3(—3.6)), broadest about the middle; above glabrous, dull to glossy, pale FLORA MALESIANA [ser. I, vali @? greyish green to chocolate-brown, underneath (except midrib and nerves) densely greyish brown tomentose by adpressed, minute, stellate hairs; base rounded-acute, top bluntly 1-3 cm acuminate -caudate; midrib thin, prominent on both sides; nerves (6—)8—10(—11) pairs, thin, obscure to dis- tinct beneath, subparallel, at an angle of 50-60°, arcuating and faintly anastomosing near the mar- gin; reticulation fine, irregular or areolate, ob- scure; petiole 5-10 mm, 1-114 mm g, glabrous, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely greyish brown stellate hairy, subglabrescent; bracts and bracte- oles ovate, %-2%4 by %-1 mm. Male rachis 7-20 cm, 1-114 mm g, mostly simple and axillary; 3 flowers solitary, perianth 5—6-lobed, stamens 10-12, filaments 214-314 mm, anthers 0.25-0.3 mm long, pistillode globose, c. 1 mm @. Andro- gynous or mixed rachis 8-15 cm, 1144 mm g; female flowers solitary, staminodes rudimentary, styles 3, conical, 1-2 mm, recurved. Ripe cupule sessile to 2-3 mm stalked, c. 3 mm long, 8-12 mm g; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely set with adpressed simple hairs, outside densely greyish stellate hairy, hairs minute, adpressed; lamellae 4—5, thin, con- centric, entire or denticulate. Ripe fruit ovoid- conical, 8-13 mm through, glabrous, light choco- late-brown, top rounded-acute, base rotundate, scar concave, 3-7 mm g; wall bony, 44-1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak, Kalimantan, Sabah). Ecol. In forests, from sea-level up to 1800 m; rather common in heath forest. F/. Aug.—May, /r. Febr.—July. 67. Lithocarpus gracilis (KORTH.) SOEPADMO, Rein- wardtia 8 (1970) 243. — Quercus gracilis KORTH. Kruidk. (1844) 207; A. DC. Prod. 16, 2 (1864) 93; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 88; A. Camus, Chénes 3 (1954) 1158. — Quercus cyrtorhyncha MiqQ. Sumatra (1861) 350; Hook. f. Fl. Br. Ind. 5 (1888) 613; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 66, t. 60 B. — Quercus diepen- horstii Mia. Sumatra (1861) 349; A. DC. Prod. 16, 2 (1864) 95; WENziG, Jahrb. Bot. Gart. Berl. 4 (1886) 231; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 67, t. 61 A. — Cyclobalanus diepenhorstii (Miq.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 80. — Cyclobalanus gracilis (KORTH.) Oerst. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 376. — Pasania cyrtorhyncha (MiQ.) GAM- BLE, J. As. Soc. Beng. 75, ii (1915) 432. — Synaedrys cyrtorhyncha (MiqQ.) Koz. Bot. Mag. Tokyo 30 (1916) 191. — Synaedrys diepenhorstii (MiIQ.) Korpz. /.c. 191. — L. cyrtorhyncha (MiqQ.) REHD. J. Arn. Arb. 1 (1919) 124; A. Camus, Chénes 3 (1954) 734, t. — L. diepenhorstii (M1Q.) BARNETT, Trans. & Proc. Bot. Soc. Edinb. 34 (1944) 177; A. Camus, Chénes 3 (1954) 752, t. — L. cyathifor- mis A. CAMus, Bull. Soc. Bot. Fr. 94 (1947) 4; Chénes 3 (1954) 802, t. 417: 1-8. Tree 10-40 m, 20-90 cm g; buttresses up to 114 m tall, 1 m out, 10 cm thick; bark smooth to 1972] Fig. 27. Lithocarpus orbicularis SoppADMO. Habit of fruiting shrub, Gajolands, N. Sumatra, c. 2800 m FAGACEAE (Soepadmo) : i pan ts alt. (VAN STEENIS 8977; Febr. 1937). fissured, rarely scaly, greyish brown. Branchlets initially angular, densely or sparsely set by silvery to greyish brown, short simple or stellate hairs, later terete, glabrous, greyish to dark brown, sparsely lenticellate; terminal buds subglobose, c. 1-2 by 14%4 mm, scales ovate. Stipules ovate to linear-acute, 114-4 by 0.7-1 mm. Leaves thin- coriaceous, (10—)15—20(—33) by (4~-)5-8(-13) cm (index (2—)214-3(—3.6)), broadest at the middle; above glabrous, dull to glossy, pale to dark brown, underneath with a thin cover of pale brown to glaucous, adpressed, minute stellate hairs; base 364 FLORA MALESIANA [ser. I, vol. 7? abruptly acute to cuneate, occasionally asymmetri- cal, top bluntly acute to 1-114 cm acuminate- cuspidate; midrib strongly prominent on both sides; nerves (11—)14-16(-17) pairs, thin, promi- nent on both surfaces, subparaliel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, subscalariform to irregular, distinct beneath; petiole 7-10 mm, 2 mm @, adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous stellate hairy; bract and bracteoles ovate, c. 1 by 1/3 mm. Male rachis 10- 20 cm, 1-2 mm g, simple and axillary or much- branched and subterminal; 3 flowers solitary or in clusters of 3, filaments 3-4 mm, anthers 0.35-0.4 mm long, pistillode subglobose, c. 1 mm @. Andro- gynous or mixed rachis 10-15 cm, 1-2 mm 9g; female flowers solitary or rarely in clusters of 2-3, staminodes rudimentary, styles 3, conical, c. 114 mm, connate. Ripe cupule sessile to 14 cm stalked, saucer- to cup-shaped, 0.3-1 cm long, 2-2.7 cm @; rim thin, covering the basal part of the fruit to c. ¥% part of the fruit; wall thin, inside densely fulvous to silvery tomentose by adpressed simple hairs, outside densely fulvous to glaucous tomen- tose by minute, adpressed, stellate hairs; lamellae 6-8, thin, distantly denticulate, more or less con- centric. Ripe fruit ovoid-conical, 1.5-1.8 cm long, 1.5-2.6 cm g, glabrous, dark chocolate-brown to purplish black, top abruptly acuminate, base ro- tundate, scar flat to shallowly concave, 1.4-1.7 cm @; wall bony, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: S. Sumatra (Palembang and vicinity, Simalur I.), Malay Peninsula (Perak, Pahang, Selangor, Negri Sembilan; Singapore), Borneo (common in all parts; Nunukan and Laut Is.). Ecol. Primary, sometimes also in secondary and heath forests, on ridges or hills or flat swampy lands, on sandy or rarely limestone derived soil, often on river-banks, up to 1500 m, more common- ly below 800 m. Fertility throughout the year. 68. Lithocarpus rassa (MIQ.) REHD. J. Arn. Arb. 1 (1919) 130; A. Camus, Chénes 3 (1954) 739, t.; SOEPADMO, Reinwardtia 8 (1970) 271. — Quercus rassa MiQ. Sumatra (1861) 350; A. DC. Prod. 16, 2 (1864) 95; Hook. f. Fl. Br. Ind. 5 (1888) 613, incl. var. montana KiNG ex Hook. f.; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 66, t. 60 A; CoRNER, Ways. Trees (1940) 304, f. 96. — Cyclobalanus rassa (MiQ.) OERsT. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 376. — Quercus wenzigiana KING ex Hook. f. Fl. Br. Ind. 5 (1888) 613; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 65, t. 58 B; CORNER, Ways. Trees (1940) 305, f. 96. — Quercus rassa var. lanuginosa RIDL. J. Str. Br. R. As. Soc. n. 61 (1912) 37. — Pasania rassa (MIQ.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 436, incl. var. lanu- ginosa, l.c. 437. — Pasania wenzigiana (KING ex Hook. f.) GAMBLE, /.c. 435. — Synaedrys rassa (MiQ.) Korpz. Bot. Mag. Tokyo 30 (1916) 192. — Synaedrys wenzigiana (KING ex Hook. f.) Kotpz. I.c. 193. — L. rangeriana A. Camus, Bull. Mus. Hist. Nat. Paris II, 4 (1932) 912; Chénes 3 (1954) 791, t. 415: 9-11. — L. ridleyana A. Camus, Bull. Mus. Hist. Nat. Paris II, 4 (1932) 913; Chénes 3 (1954) 752, t. 404: 20-27. — L. symingtoniana A. Camus, Bull. Mus. Hist. Nat. Paris IT, 4 (1932) 913; Chénes 3 (1954) 736, t. 399: 14-15. — L. wenzigiana (KING ex Hook. f.) A. CAMus, Riviera Scient. 18 (1932) 42; Chénes 3 (1954) 744, t. Shrub or tree, 3-24 m, 15-90 cm g; buttresses rounded, up to 0.6 m tall, 114 m out, 5 cm thick; bark fissured to scaly, lenticellate, pale grey- brown. Branchlets initially densely set with yellow- ish brown, short, simple or stellate hairs, later glabrous, greyish to blackish brown, finely fissured or sparsely lenticellate; terminal buds ovoid-ellip- soid, 3-5 by 2-3 mm, scales ovate to linear. Stip- ules linear-acute to subulate, 3-4 by 1-2 mm. Leaves thin-coriaceous, rigid, (314—)5-13(-17) by (1.2-)2-4(-614) cm (index (2-)2144-314%4(4)4)), broadest about the middle; above glabrous, glossy, chocolate-brown, underneath densely set with glaucous to fulvous, adpressed, minute stellate hairs, subglabrescent; base acute to cuneate, rarely rounded, margin recurved, top bluntly acute to bluntly or sharply 14-114 cm acuminate-caudate; midrib strongly prominent on both sides; nerves thin, more or less prominent on both surfaces, (8-)10-14(-16) pairs, subparallel at an angle of 40-50°, arcuating and anastomosing near the margin; reticulation fine, areolate, obscure; petiole 14-2 cm, 1-114 mm 9g, glabrous, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely yellowish brown, stellate tomen- tose; bracts thick-coriaceous, narrowly ovate- acute, 0.7—-by 0.3 mm, bracteoles broadly ovate, 0.3 by 0.7 mm. Male rachis 7-20 cm, 1-3 mm 9g; 3 flowers in clusters of 3, filaments 2-3 mm, anthers 0.2-0.3 mm long, pistillode globose, 0.7—1 mm g. Androgynous or mixed rachis 6-20 cm, 1-3 mm 9; female flowers solitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3-5, coni- cal, 0.7-1 mm. Ripe cupule sessile to 3-4 mm stalk- ed, cup- to saucer-shaped, 5-12 mm long, 14-22 mm 9; rim thin, covering 14-14 part of the fruit; wall woody, rather thick, inside densely silvery grey tomentose by adpressed simple hairs, outside densely fulvous stellate hairy, hairs minute, ad- pressed; lamellae 6-10, prominent, entire or denti- culate. Ripe fruit subhemispherical, 12-15 mm long, 15-20 mm g, pale chocolate-brown, glabrous except around the umbo, top rounded-apiculate, base rotundate, scar concave, c. 1 cm @; wall bony, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (scattered), Malay Peninsula (Perak, Pahang, Trengganu, Malacca, Johore), Borneo (Sarawak, Sabah). Ecol. In primary forest, from sea-level up to 1800 m. Fi. July—June, fr. May—March. Notes. The typical form of this species grows in the submontane forest at 600-1500 m, and has small, narrowly ovate to linear-lanceolate leaves, small fruits and cupules with 5—6 lamellae. Speci- mens from the lowland forest, recognized by KING and others as L. (Quercus) wenzigiana, 1972] FAGACEAE (Soepadmo) 365 nn CEU UE EEE SESE Sa usually have larger leaves, cupules and fruits. In KEP 94394, SFN 10775, from Malay Peninsula, Honinc s.n. from Sumatra, and SAN 26163, S. 5385, 18030, from Borneo, the leaves, cupules and fruits are intermediate. Specimens from 1500-1800 m, formerly recognized as L. symingtoniana, have much smaller leaves, cupules and fruits than those of L. rassa, but several intermediates also occur. 69. Lithocarpus andersonii SOEPADMO, Reinwardtia 8 (1970) 215, f. 10. Tree 7-27 m, 20-60 cm @g; stilt-roots occasion- ally present, up to 1.8 m tall; bark smooth, light grey. Branchlets initially sparsely greyish stellate hairy, later glabrous, greyish to blackish brown, sparsely to densely lenticellate; terminal buds ovoid-ellipsoid, 3-4 by 2 mm, scales narrowly ovate-acute. Stipules narrowly ovate-acute to subulate, 3-4 by 1-2 mm. Leaves thin-coriaceous, rigid, (6-)7-10(--1114) by (2-)214-3144(-5) cm (index 2-3), broadest about the middle; above glabrous, glossy, pale greyish green to yellowish brown, underneath sparsely tomentose by greyish to chocolate-brown, adpressed, minute, stellate hairs, glabrescent; base acute or rounded, margin recurved, top bluntly or sharply 1-114 cm acumi- nate; midrib strongly prominent on both sides; nerves thin, obscure to rather prominent beneath, 9-11 pairs, subparallel, at an angle of 45-60°, arcuating and anastomosing near the margin; reticulation fine, areolate, obscure; petiole 5-8 mm, 1 mm g, glabrous, furrowed. Inflorescence male or androgynous, densely greyish brown, stellate hairy, hairs adpressed, minute; bracts and bracteoles broadly or narrowly ovate, 4-114 by 2,mm. Male rachis 5-15 cm, 1-2 mm 9; ¢ flowers in clusters of 3, perianth 5—6—-lobed, stamens 10— 12, filaments 214-314 mm, anthers 0.25 mm long, pistillode subglobose, c. 24-1 mm 9. Androgynous rachis 5-10 cm, 1-114 mm 9g; female flowers soli- tary, staminodes rudimentary, styles 3, conical, c. 1 mm, recurved. Ripe cupule sessile to 3-4 mm stalked, saucer-shaped, 2-6 mm long, 15-22 mm g; rim thin, undulate, recurved, covering the basal part of the fruit; wall woody, thin, inside densely tomentose by adpressed, simple hairs, outside densely greyish stellate hairy; lamellae 5—6, thin, concentric, denticulate. Ripe fruit ovoid-conical, 1.2-2.2 cm long, 1.6-2.3 cm g, glabrous, dark purplish brown, top rounded-acuminate, base rotundate, scar deeply concave, 9-15 mm g; wall bony, c. 1.2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Borneo (Sarawak; Sampit in S. Central Kalimantan). Ecol. Peat swamp forest at low altitude. Fi. April-June, fr. Dec._May. 70. Lithocarpus vinkii SoEPADMO, Reinwardtia 8 (1970) 286. Tree 15-30 m, 15-50 cm 9; buttresses sometimes present, up to 2!4 m tall and out, 10 cm thick; bark grey-brown, smooth to shallowly fissured. Branchlets initially densely set with a fulvous ad- pressed stellate tomentum, soon glabrescent, grey- ish brown, finely fissured, with many minute lenticels; terminal buds ovoid, c. 3 by 2 mm, scales ovate. Stipules ovate, 1-2 by 1 mm. Leaves coriaceous, (7—)8—-14(-18) by (3-)4-5(-7) cm (index (1.7-)2-214(-314)), broadest at or rarely below the middle; surfaces more or less discolor- ous, above glabrous, dull or glossy, pale to dark greyish brown, beneath yellowish grey to glaucous tomentose by adpressed minute stellate hairs; base acute, rarely rounded, top abruptly acute to 14-1 cm acuminate, tip blunt or pointed, sometimes oblique; midrib and nerves thin, flattish on both sides to impressed above; nerves (8—)10-12 pairs, subparallel, at an angle of 45-S0°, arcuating but not anastomosing towards the margin; reticulation fine, irregular, obscure on both sides; petiole 7-10 mm, 114-2 mm g, adaxially flat. Inflorescence male or androgynous, mostly simple and axillary, densely fulvous tomentose by short stellate hairs; bracts and bracteoles narrowly ovate, 74-1 by 4-24 mm. Male rachis 10-20 cm, 14% mm 9; 3 flowers in clusters of 3, filaments c. 2 mm, an- thers 0.25 mm long, pistillode subglobose, 73-1 mm 9. Androgynous rachis 10-15 cm, 114-2 mm 9; female flowers solitary or rarely in clusters of 2, staminodes rather well-developed but not exceed- ing the perianth, styles 3(-4), conical, c. 1 mm, recurved. Ripe cupule solitary or rarely in clusters of 2, subsessile, saucer-shaped, 14-14 cm long, 114-2 cm 9g, inside densely greyish tomentose by simple hairs, outside densely fulvous to glacous tomentose by adpressed stellate hairs; rim thin, incurved, covering the basal part of the fruit; lamellae 4-6, concentric, thin, denticulate. Ripe fruit conical, glabrous, pale chocolate-brown, 1344- 214 cm long, 114 cm g, top acute, base truncate or rounded, scar concave, 34-1 cm 9; wall woody, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: New Guinea (common in the Vogelkop Peninsula, scattered in the eastern part), also Japen I., eastward to Normanby, Misima, Sudest and Rossel Is. Ecol. Forest, from sea-level up to 1800 m, more commonly below 700 m, on sandy to clayey soil overlying limestone. Both in the Vogelkop Penin- sula and in the islands east of New Guinea, this species has been recorded as forming pure stands on low ridges. Fl. March—Dec., fr. Sept—March. 71. Lithocarpus clementianus (KING ex Hook. /-) A. Camus, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 707, t. 391: 1-8; SOEPADMO, Reinwardtia 8 (1970) 229. — Quercus clementiana KING ex Hook. f. Fl. Br. Ind. 5 (1888) 614; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 69, t. 63 A; CORNER, Ways. Trees (1940) 301, f. 96. — Pasania clemen- tiana (KiNG ex Hook. f.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 439. — Synaedrys clementiana (KinG ex Hook. f.) Kornz. Bot. Mag. Tokyo 30 (1916) 191. — Quercus teysmannii (non BL.) HEINE in Fedde, Rep. 54 (1951) 225. Tree 10-36 m, 30-70 cm g; bark deeply fissured, brownish. Branchlets initially angular, sparsely stellate hairy, later terete, glabrous, blackish 366 FLORA MALESIANA [ser. I, vol. 7? brown, finely fissured; terminal buds ellipsoid, 3-5 by 1-2 mm, scales linear. Leaves coriaceous, (10—)12—16(—20) by (3-)4—S(-7) cm (index 3-314), broadest at or slightly below the middle; above glossy, greyish green to pale chocolate-brown, underneath densely fulvous to pale grey tomentose by adpressed, minute, stellate hairs, subglabrescent; base acute, margin recurved, top 1-114 cm acumi- nate; midrib prominent on both sides; nerves 10— 14 pairs, thin, parallel at an angle of c. 45°, arcuat- ing but not anastomosing towards the margin; reticulation fine, dense, subscalariform, obscure; petiole 14-1 cm, 2 mm @g, glabrous, adaxially flat. Inflorescence male, androgynous or mixed, densely yellowish brown, adpressed stellate hairy, sub- glabrescent; bracts and bracteoles ovate to linear, 14-2 by 4%-2, mm. Male rachis 10-15 cm, 2-3 mm @; 4 flowers in clusters of 3-7, filaments 214— 314 mm, anthers 0.2-0.25 mm long, pistillode subglobose, 1-114 mm g. Androgynous or mixed rachis 5-10 cm, 2-5 mm 9g; female flowers solitary or in clusters of 2-3, staminodes rudimentary, styles 3, conical, 2-3 mm, recurved. Ripe cupule sessile, cup-shaped, 1-114 cm long, 214-3 cm @g; rim 1-2 mm thick, covering 4-14 part of the fruit; wall woody, 3-4 mm thick, inside densely fulvous tomentose by adpressed simple hairs, outside sparsely stellate hairy by adpressed stellate hairs, glabrescent; lamellae 6-8, rather prominent, concentric. Ripe fruit depressed subglobose, 1—11/4 cm long, 2-3 cm g, glabrous, top rounded-umbo- nate, base truncate, scar concave, c. 1 cm 9; wall woody, c. 2 mm thick, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (Pahang; Penang), Borneo (Sarawak, scattered; Sabah: Mt Kinabalu, rather common). Ecol. Forests at 600-1900 m. F/. April-June, fr. July—Febr. Note. Specimens from Borneo usually have thicker leaves than those from Malay Peninsula. The cupule and fruit are, however, very similar. 72. Lithocarpus suffruticosus (RIDL.) SOEPADMO, Reinwardtia 8 (1970) 280, f. 13. — Pasania rassa var. suffruticosa R1IDL. Fl. Mal. Pen. 3 (1924) 382. — L. rassa var. suffruticosa (RIDL.) A. CAMUS, Chénes 3 (1954) 741. Shrub or tree, 3-15 m tall. Branchlets initially densely set with rufous, adpressed stellate hairs, later glabrous, greyish to blackish brown, sparsely lenticellate; terminal buds ovoid, 1-2 by 1 mm, scales ovate-acute. Stipules broadly ovate to del- toid, c. 4% by 144 mm. Leaves thick-coriaceous, rigid, (6—)8—10(—12) by 3—5 cm (index (1.6—)2(-3)), broadest at or below the middle; above glabrous, glossy, dark chocolate-brown, underneath densely glaucous tomentose by adpressed, minute, stellate hairs; base acute to cuneate, margin recurved, top 1-2 cm acuminate-caudate, tip blunt or sharp; midrib strongly prominent on both sides; nerves (7—)9-10(-14) pairs, obscure, subparallel at an angle of c. 45°, arcuating and anastomosing near the margin; reticulation fine, subscalariform, ob- scure; petiole (S—)10-13(-16) mm, 1-2 mm ag, glabrous, adaxially flat. Inflorescence male, andro- gynous or mixed, densely tomentose by greyish to rufous, adpressed stellate hairs; bracts and brac- teoles ovate, 14-24 by 4 mm. Male rachis 10 cm, 114 mm @g, simple and axillary or much-branched and subterminal; 3 flowers solitary or in clusters of 2-3, perianth 5—6-lobed, stamens 10~-12, fila- ments 2 mm, anthers 0.35 mm long, pistillode globose, c. 1 mm @. Androgynous or mixed rachis 6cm, 1-2mmg; female flowers solitary, staminodes rudimentary, styles 3-4, conical, 1 mm. Fruiting rachis up to 20 cm, with numerous cupules. Ripe cupule 3—5 mm stalked, cup-shaped, 0.6-1 cm long, 2-214 cm @; rim 114-2 mm thick, covering 14—, part of the fruit; wall woody, 2-3 mm thick, inside densely greyish brown adpressed-tomentose by simple hairs, outside densely greyish tomentose by adpressed, minute stellate hairs; lamellae 8-10, concentric, entire or distantly denticulate. Ripe fruit ovoid-conical, 17-25 mm long, 17-20 mm g, glabrous, dark chocolate-brown, top acute, base rotundate, scar deeply concave, 1-114 cm @; wall bony, 1—2 mm thick, for the greater part free from the cupule. Distr. Malesia: N. Sumatra (Mts Kemiri and Talamau), Malay Peninsula (Perak, Selangor, Pahang, Trengganu). Ecol. Submontane to mossy montane forest at 600-3150 m. Fertility seems to be throughout the year. Note. Though the leaves are suggestive of L. rassa, the cupule and fruits are different. 73. Lithocarpus elegans (BL.) HATUS. ex SOEPADMO Reinwardtia 8 (1970) 236. — Quercus spicata SM. in Rees, Cyclop. (1814) Quercus n. 12, non HUMB. & BonpL. 1809; D. Don, Prod. Fl. Nepal. (1825) 56; WALL. Pl. As. Rar. 1 (1830) 40, t. 46; Mia. Fl. Ind. Bat. 1, 1 (1856) 848; Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 106; A. DC. Prod. 16, 2 (1864) 85; Kurz, For. Fl. Burma 2 (1877) 486; WENZzIG, Jahrb. Bot. Gart. Berl. 4 (1886) 224; Hook. f. Fl. Br. Ind. 5 (1888) 609; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 47, t. 41: 1-2; K. & V. Bijdr. 10 (1904) 39; BranpIs, Indian Trees (1906) 629, f. 194; Cras, Kew Bull. (1911) 473; CorNeErR, Ways. Trees (1940) 304, f. 98. — Quercus racemosa Jack, Mal. Misc. 2 (1822) 86, non LAMK, 1783; in Hook. Comp. Bot. Mag. 1 (1836) 255; KorTH. Kruidk. (1844) 205; Merr. J. Arn. Arb. 33 (1952) 241. — Quercus elegans BL. Verh. Bat. Gen. K. & W. 9 (1823) 208; Bijdr. (1826) 518; Fl. Jav. Cupul. (1829) 21, t. 10; OupEM. Natuurk. Verh. Kon. Akad. 11 (1865) 5, t. 2, 3; BACKER & BAKH. f. FI. Java 2 (1965) 7. — Quercus depressa BL. Verh. Bat. Gen. K. & W. 9 (1823) 209, t. 1, non Hums. & BonpL. 1809. — Quercus glaberrima Bu. Verh. Bat. Gen. K. & W. 9 (1823) 210, t. 2; Bijdr. (1826) 519; Fl. Jav. Cupul. (1829) 17, t. 8. — Quercus placentaria BL. Bijdr. (1826) 518; Fl. Jav. Cupul. (1829) 19, t. 9; A. DC. Prod. 16, 2 (1864) 87; WENZzIG, Jahrb. Bot. Gart. Berl. 4 (1886) 226; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 72. — Quercus arcaula BUCH.-HAM. ex SPRENG. Syst. Veg. 3 (1826) 857; BL. Mus. Bot. 1 (1850) 290, 1972] incl. var. racemosa (JACK) BL. et var. microcalyx (KorTH.) BL. — Quercus glomerata Roxs. FI. Ind. ed. Carey 3 (1832) 640. — Arcaula spicata (SM.) RAFIN. Alsog. Am. (1838) 28. — Quercus anceps KortH. Kruidk. (1844) 204. — Quercus microcalyx KorTH. Kruidk. (1844) 206; OuUDEM. Versl. Med. Kon. Ak. Wet. Natuurk. 12 (1861) 204; Natuurk. Verh. Kon. Akad. 11 (1865) 6, t. 4: 3. — Quercus pyrifolia BL. Mus. Bot. 1 (1850) 304. — Quercus sphacelata BL. I.c. 304. — Quercus spicata var. glaberrima (BL.) Miq. FI. Ind. Bat. 1, 1 (1856) 848; A. DC. Prod. 16, 2 (1864) 86; WENZIG, Jahrb. Bot. Gart. Berl. 4 (1886) 225. — Quercus spicata var. microcalyx (KORTH.) MIQ. Fl. Ind. Bat. 1, 1 (1856) 848; A.DC. Prod. 16, 2 (1864) 86; Hook. f. Fl. Br. Ind. 5 (1888) 610; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 48, t. 43: 9-11. — Quercus spicata var. placentaria (BL.) Mia. FI. Ind. Bat. 1, 1 (1856) 849; Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 106. — Quercus gracilipes Mia. Sumatra (1861) 347. — Quercus spicata var. gracilipes (MiQ.) Mig. Ann. Mus. Bot. Lugd.— Bat. 1 (1863) 106; A. DC. Prod. 16, 2 (1864) 86; Hook. f. Fl. Br. Ind. 5 (1888) 610; K. & V. Bijdr. 10 (1904) 42; Koorp. Atlas 1 (1913) t. 42. — Quercus spicata var. racemosa (JACK) Miq. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 106; A. DC. Prod. 16, 2 (1864) 85; ScHEFF. Nat. Tijd. N. I. 31 (1870) 359. — Pasania spicata (‘WALL.’) OrERsST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83; ScHottT- KY, Bot. Jahrb. 47 (1912) 664; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 419; HickeL & A. Camus, FI. Gen. I.-C. 5 (1930) 983. — Pasania placentaria (BL.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83. — Pasania glomerata (ROXB.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. — Quercus spicata var. latifolia SCHEFF. Nat. Tijd. N. I. 31 (1870) 359. — Quercus rhioensis HANCE, J. Bot. 16 (1878) 198. —- Quercus leucocarpa Hoox. f. & THomMs. ex WENZIG, Jahrb. Bot. Gart. Berl. 4 (1886) 225. — Quercus spicata var. depressa (BL.) KING, Ann. R. Bot. Gard. Calc. 2 (1889) 48, t. 43: 8; K. & V. Bijdr. 10 (1904) 42. — Quercus spic- ata var. genuina Koorp. Exk. Fl. Java 2 (1912) 66. — Pasania spicata var. gracilipes (MI1Q.) SCHOTT- KY, Bot. Jahrb. 47 (1912) 664; GAmBLE, J. As. Soc. Beng. 75, ii (1915) 421; RipL. Fl. Mal. Pen. 3 (1924) 377, f. 159. — Pasania spicata var. placen- taria (BL.) ScHoTTKy, Bot. Jahrb. 47 (1912) 664. — Pasania spicata var. microcalyx (KORTH.) GAM- BLE, J. As. Soc. Beng. 75, ii (1915) 421. — Synae- drys spicata (SM.) Korpz. Bot. Mag. Tokyo 30 (1916) 198. — L. spicata (SM.) REHD. & WILS. in Sargent, Pl. Wils. 3 (1917) 207. — L. spicata var. gracilipes (M1Q.) REHD. J. Arn. Arb. 1 (1919) 131. — L. spicata var. placentaria (BL.) REHD. ibid. 10 (1929) 133; A. Camus, Chénes 3 (1954) 1035, t. 481: 8-13. — L. rhioensis (HANCE) A. CAMus, Riv- iera Scient. 18 (1932) 40; Chénes 3 (1954) 1026, t. 478: 8-15. — L. microcalyx (KoRTH.) A. CAMUS, Bull. Soc. Bot. Fr. 92 (1945) 83; Chénes 3 (1954) 1055, t.; SMITINAND, Nat. Hist. Bull. Siam Soc. 20 (1962) 126. — L. spicata var. elegans (BL.) A. Camus, Chénes, Atlas 3 (1949) 102, t. 481: 1-7; FAGACEAE (Soepadmo) 367 Chénes 3 (1954) 1034. — L. spicata var. glaberrima (BL.) A. CAmus, Chénes, Atlas 3 (1949) 103, t. 482: 7-15; Chénes 3 (1954) 1035. Tree 5—30 m, 20-70 cm g; bark greyish brown, deeply fissured, lenticellate. Branchlets glabrous, greyish brown, sparsely to densely lenticellate; terminal buds ovoid, 2-5 by 2—3 mm, scales nar- rowly ovate to linear, densely yellowish brown stellate hairy, glabrescent. Stipules narrowly ovate to linear, 3-6 by 1-2 mm. Leaves thick-coriaceous, rigid, (9—)12-17(—20) by (3—)4-6(-8) cm (index (1 14-)2-3(—-5)), broadest at or above the middle; surfaces glabrous, concolorous, glossy, pale to dark chocolate-brown; base acute, rounded to (extra-Mal.) auriculate, margin recurved, top bluntly acute to 1-2 cm acuminate; midrib strong- ly prominent beneath, slightly so to flat above; nerves (10—)12—15(—18) pairs, prominent beneath, obscure above, subparallel at an angle of 45-60°, arcuating and anastomosing near the margin; reticulation subscalariform to irregular, fine, ob- scure to distinct beneath; petiole 44-214 cm, 1-2 mm @g, adaxially flat, glabrous, occasionally thick- ened at the base. Inflorescence sturdy, male, andro- gynous or rarely mixed, densely fulvous simple and stellate hairy; bracts and bracteoles ovate- acute, 1-2 by 1 mm. Male rachis 10-30 cm, 2-3 mm @, sometimes much-branched; ¢ flowers in clusters of (3-)7-15(-24), filaments 214-3 mm, anthers 0.3 mm long, pistillode subglobose, 1-114 mm g. Androgynous or mixed rachis 15-30 cm, 2-34 mm @; female flowers in clusters of (3—)5—7(-10), staminodes rudimentary, styles 3, terete, 24-1 mm. Fruiting rachis up to 30 cm, carrying numerous clustered cupules. Ripe cupule sessile to 14 cm stalked, cup- to saucer-shaped, (14—)14-1(-2) cm long, (1—)2—-3(—314) cm 9; rim thin to rather thick, covering 14-'% part of the fruit; wall woody, inside densely yellowish brown tomentose by simple adpressed hairs, outside densely fulvous adpressed stellate hairy; scales adpressed, some- times rather woody, distinct but not free, imbricate to more or less concentrically set. Ripe fruit ovoid- conical to depressed ovoid-globose, (1—)114-2(- 24%) cm long, (1-)2-21%4(-3%) cm g, glabrous, chocolate-brown, top abruptly rounded acute to rounded and depressed at the centre, base rotun- date, scar flat to concave, 34-114 cm 9; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. India (Bhutan, Nepal), E. Pakistan (Chittagong), Burma, Indo-China; in Malesia: Sumatra, Malay Peninsula, Java, Borneo (com- mon), Celebes (rare). Ecol. Forests, from sea-level up to 2400 m, more commonly at 1000-1600 m, on various types of soil. North of the equator f/. Oct.—June, fr. Aug.—April; south of the equator fertile through- out the year. Notes. L. elegans is the most widely distributed, and narurally the most variable species, and pro- bably the most primitive in the genus. In the past seven or more varieties have been distinguished, and some of them even had been recognized as 368 FLORA MALESIANA [ser. I, vol. 7? distinct species. After examining a vast number of collections, I came to the conclusion that none of these varieties deserves to be regarded as sepa- rate species. There are many intergrades, and the variations are mainly due to different habitats. The species as accepted here is recognizable by the glabrous, pale to dark chocolate-brown leaves, glossy on both sides, with somewhat irregular reticulation, and by the cupules which are always squamose and set in dichasial clusters of 3-10. In the flowering stage, it may be distinguished by the rigid and sturdy inflorescence with the flowers in clusters of 3-15. Records from New Guinea relate to L. megacar- pus; and those from China probably to L. henryi. The above synonymy covers Malesia only, the complete one is given in Reinwardtia /.c. 74. Lithocarpus jacobsii SoEPADMO, Reinwardtia 8 (1970) 248. Tree 9-36 m, 10-35 cm; stilt-roots occasion- ally present; bark pale brown, smooth, lenticel- late. Branchlets initially with a sparse adpressed stellate tomentum, the hairs sometimes intersper- sed by minute, reddish brown tinged (scales?), later glabrous, dull greyish brown, sparsely lenti- cellate; terminal buds ovoid, 5-6 by 3-4 mm, scales narrowly ovate, densely rufous stellate hairy. Stipules linear, 5-10 by 1-2 mm, rather long persis- tent. Leaves thick-coriaceous, rigid, (18—)24—40 (-56) by (5—)9-11(-16) cm (index 3-4), broadest slightly above the middle; surfaces more or less discolorous, glabrous, greyish green, paler be- neath; base cordate to auriculate, margin recurved, top bluntly acute to abruptly, sharply 1 cm acumi- nate; midrib strongly prominent beneath, flat above, nerves (13-)14-17(-19) pairs, prominent beneath, impressed above, subparallel, at an angle of c. 60°, arcuating and anastomosing near the margin; reticulation lax, subscalariform to irregu- lar, distinct beneath; petiole 14-114 cm, 3-6 mm @, terete or adaxially flat. Inflorescence male or female, densely fulvous stellate hairy; bracts and bracteoles linear-acute, 2-3 by 14-1 mm. Male rachis (not fully developed) c. 5 cm, 2 mm 9; ¢ flowers in clusters of 3. Female rachis c. 25 cm, 214 mm g; @ flowers in clusters of 3, staminodes well-developed and exceeding the perianth, styles 3, conical, 2-3 mm. Ripe cupule 1-1 cm stalked, saucer-shaped, 144-14 cm long, 2-214 cm @; rim thin, covering the basal part of the fruit; wall woody, thin, dull grey-brown stellate hairy on both sides; scales obscure, set in inconspicuous con- centric rows. Ripe fruit ovoid-globose, 114-2 cm long, 2-214 cm g, glabrous, pale chocolate-brown to dark brown, top abruptly acute, base truncate, scar deeply concave, c. 1144 cm @; wall bony, c. 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Borneo (Sarawak, Sabah, rare). Ecol. Forests, below 500 m, usually in swampy places. Fi. Febr., fr. Sept.-Nov. 75. Lithocarpus lampadarius (GAMBLE) A. CAMUS, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 717, t. 398: 7-10; SopPADMO, Reinwardtia 8 (1970) 253. — Pasania lampadaria GAMBLE, Kew Bull. (1914) 177; J. As. Soc. Beng. 75, ii (1915) 444. — Synaedrys lampadaria (GAMBLE) KoIbZz. Bot. Mag. Tokyo 30 (1916) 192. — Quercus lampadaria (GAMBLE) BURKILL, Kew Bull. (1935) 318; Dict. 2 (1935) 1857; CorNER, Ways. Trees (1940) 303, f. 96. Tree 15-36 m, 30-100 cm g; buttresses up to 1.2 m tall, 1144 m out, 10 cm thick; bark dark brown, scaly. Branchlets initially densely fulvous to rufous stellate hairy, later glabrous, sturdy, greyish to blackish brown, sparsely lenticellate, the lenticels large; terminal buds ellipsoid, 5-10 by 2-3 mm, scales linear. Stipules linear-acute, 6-10 by 1-3 mm. Leaves thick-coriaceous, rigid, (15—) 20-25(-30) by (6-)8-10(-15) cm (index 1.7-3), broadest about the middle; above glabrous, dull to glossy, pale greyish green, underneath densely greyish brown to glaucous tomentose by adpressed, minute stellate hairs; base acute to cuneate, rarely rounded, margin recurved, top bluntly or sharply acute to 114-214 cm acuminate; midrib strongly prominent on both sides; nerves (10—)12-15(-18) pairs, prominent beneath, impressed above, paral- lel, at an angle of c. 60°, arcuating but not anasto- mosing towards the margin; reticulation scalari- form, distinct beneath; petiole 1-3 cm, 2-3 mm g, glabrous, terete or adaxially flat. Inflorescence male or androgynous, densely fulvous stellate hairy; bracts linear acute, 6-8 by 144-2 mm, bracteoles broadly ovate, 2-3 by 114-2 mm. Male rachis 10-30 cm, 3-4 mm g, much-branched; 3 flowers in clusters of 3-7 (or more), filaments 3-4 mm, anthers 0.35 mm long, pistillode subglo- bose, 1-2 mm @. Androgynous rachis 10-15 cm, 3-4 mm g; female flowers in clusters of 3—7, very rarely solitary, staminodes rudimentary, styles 3-5, coni- cal, 114-2 mm, recurved. Fruiting rachis sturdy, densely warty lenticellate. Ripe cupule subsessile, cup-shaped, 1-1.2 cm long, 2-3 cm 9; rim thick, covering the basal part of the fruit; wall woody, rather thick, inside densely yellowish brown, sil- very tomentose by adpressed simple hairs, outside densely fulvous adpressed stellate hairy; scales obscure, set in 5-6 more or less concentric rows. Ripe fruit depressed ovoid-globose, 2-24%4 cm long, 3-314 cm g, densely yellowish brown tomen- tose by adpressed, simple hairs, top rotundate, depressed-umbonate at the centre, base rotundate, scar deeply concave, up to 2 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (Perak, Pa- hang, Trengganu), Borneo (Sabah: Mt Kinabalu, common). Ecol. Forests, at 900-2000 m. F/. Sept.—April, fr. May—Aug. Uses. Twigs are used by the local people as torches. 76. Lithocarpus wallichianus (LINDL. ex HANCE) Reup. J. Arn. Arb. 1 (1919) 132; A. CAmus, Chénes 3 (1954) 1102, t. 503: 1-8; SOEPADMO, Reinwardtia 8 (1970) 287. — Quercus wallichiana LINDL. ex HANCE, J. Bot. 8 (1870) 4; Hook. f. Fl. 1972] FAGACEAE (Soepadmo) 369 Br. Ind. 5 (1888) 610; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 51, t. 46; CorNER, Ways. Trees (1940) 305, f. 96. — Pasania wallichiana (LINDL. ex HANCE) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 425. — Quercus hystrix var. longispica GAMBLE, lc. 429. — Synaedrys wallichiana (‘LINDL. in WALL.’) Korpz. Bot. Mag. Tokyo 30 (1916) 199. Tree 10-30 m, 10-90 cm g. Branchlets initially densely set with fulvous erect stellate hairs, angu- lar, later subglabrous, terete, slender, dark greyish brown, sparsely to densely lenticellate; terminal buds ovoid-globose, 2-5 by 2-3 mm, scales ovate to linear. Stipules lanceolate to linear, 214-5 by 1-114 mm. Leaves rigid, coriaceous, (10—)14—-18 (-7) cm (index (214—)3-314(-S5)), broadest at or above the middle; surfaces discolorous, above pale chocolate-brown, with erect stellate hairs especially on the midrib and nerves, underneath densely set with pale yellowish brown, adpressed and erect stellate hairs; base acute, margin recur- ved, top acute to 1-2 cm acuminate; midrib strong- ly prominent on both sides; nerves (10—)12-16 (-18) pairs, prominent beneath, impressed above, subparallel at an angle of 50-60°, arcuating and faintly anastomosing near the margin; reticulation dense, scalariform, distinct beneath; petiole (5—)10 —12(-17) mm, 1-2 mm @g, densely stellate hairy, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely yellowish brown to fulvous stellate hairy; bracts and bracteoles ovate, 1-114 by 14-24 mm. Male rachis 10-15 cm, 2 mm 9g; ¢ flowers in clusters of 3, filaments 3-5 mm, anthers 0.35 mm long, pistillode subglobose, c. 1 mm g. Androgynous or mixed rachis 10-20 cm, 114-2 mm 9; female flowers in clusters of 3, stami- nodes rudimentary, styles 3, conical, 1-3 mm, recurved. Ripe cupule subsessile, saucer- to cup- shaped, 3-5 mm long, 1.5-1.7 cm g; rim thin, covering the basal part of the fruit; wall woody, 1-2 mm thick, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous to rufous stellate hairy; scales set in concentric rows, adpressed. Ripe fruit depressed ovoid to subhemispherical, 1-144 cm long, 1.8-2 cm 29, densely pale yellowish brown tomentose by ad- pressed, minute simple hairs, top depressed-umbo- nate, base rotundate, scar concave, 8-10 mm @g; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Peninsular Siam (at 6° 30’ N: Kerr 15875); in Malesia: Sumatra (scattered in the northern half, also Singkep I.), Malay Peninsula (common in all parts; also Penang, Tioman, and Singapore Is.). Ecol. Primary forests, from sea-level up to 1600 m, usually on poor soil. F/. Dec.—May, /r. July—Febr. Note. The leaves are suggestive of L. cyclopho- rus, but the cupule and fruit are much smaller and of completely different shape. 77. Lithocarpus erythrocarpus (RIDL.) A. CAMUS, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 962, t. 461: 6-12; SoppADMo, Reinwardtia 8 (1970) 240. — Pasania erythrocarpa RipL. J. Bot. 42 (1924) 301. Tree 9-24 m, c. 40 cm g; buttresses concave, 2/3 m tall; bark whitish grey, scaly, lenticellate. Branchlets initially densely fulvous-tomentose by adpressed, minute stellate hairs, later glabrous, sturdy, greyish brown, with numerous large lenti- cels; terminal buds ovoid, 2—3 by 2 mm, scales ovate. Leaves coriaceous, 20—26 by 6-9 cm (index 2.6-3.3), broadest at or below the middle; above glabrous greyish green, dull, underneath with a thin cover of pale green, adpressed, stellate hairs; base acute, top acute, margin recurved; midrib broad, slightly prominent on both sides; nerves 12-15 pairs, flat on both surfaces, parallel, at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, irregular, obscure; petiole 114-2 cm, 3-5 mm 4g, terete or adaxially flat, corky. Fruiting rachis sturdy, c. 10 cm, 14-1 cm g, densely lenticellate, carrying nu- merous, clustered cupules. Young cupule sessile, in clusters of 2-3, or rarely solitary, blackish brown. Young fruit: staminodes rudimentary, styles 3, terete, 2 mm, recurved. Ripe cupule sessile, cup-shaped, 1-114 cm long, 2 cm 9; rim thin, covering 14 part of the fruit; wall woody, densely dark reddish brown tomentose on both sides; scales minute, adpressed, whitish, set in obscure concentric rows. Ripe fruit cylindrical, 244 cm long, 114-134 cm g, densely dark reddish brown tomentose by adpressed simple hairs, top rounded- apiculate, base truncate, scar deeply concave, c. 1 cm g; wall woody, c. 2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Malay Peninsula (Selangor, Negri Sembilan, Pahang). Ecol. Primary forests, on ridges below 900 m. Fr. May—Dec. Notes. Though the young cupules are in clus- ters of 2-3, at maturity only one is fully developed, the others rudimentary. Inflorescences are unknown. 78. Lithocarpus oreophilus SOEPADMO, Reinwardtia 8 (1970) 264. — Quercus monticola KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 44, t. 37, non PETZOLD & KircHner, 1864; RipL. J. Fed. Mal. St. Mus. 8 (1917) 86. — Synaedrys monticola (KING) KoIbDz. Bot. Mag. Tokyo 30 (1916) 197. — L. monticola (KING) ReuD. J. Arn. Arb. 1 (1919) 129; A. Camus, Chénes 3 (1954) 1130, t. 510: 6-11. Tree 9-20 m, c. 30 cm @. Branchlets initially densely set with fulvous adpressed, minute stellate hairs, later glabrous, greyish to blackish brown, sparsely lenticellate; terminal buds ovoid, 1-2 by 1 mm, scales ovate. Stipules linear-acute, 3-4 by 1 mm, rather long persistent. Leaves thick-coria- ceous, rigid, 214-5 by 114-4 cm (index 1-2), broad- est at the middle; surfaces more or less discolorous, above glabrous, glossy, chocolate-brown to grey- ish green, beneath with a thick cover of greyish, adpressed, minute stellate hairs; base rounded to subcordate, margin recurved, top rounded; midrib thin, prominent on both sides; nerves 5-6 pairs, flat on both surfaces, subparallel at an angle of 370 FLORA MALESIANA [ser. I, vol. 72 45-60°, arcuating but not anastomosing towards the margin; reticulation fine, areolate, obscure; petiole 2-3 mm, 1-1'4 mm 9, adaxially flat. In- florescence male or female, densely stellate hairy; bract and bracteoles narrowly ovate, 1-2 by 14-1 mm. Male rachis 3—5 cm, 1-114 mm 9g; ¢ flowers in clusters of 3, filaments 4-5 mm, anthers 0.3-0.4 mm long, pistillode subglobose, 1-114 mm g. Female rachis 3-4 cm, 2-3 mm g; 2 flowers solitary or rarely in clusters of 2—3, staminodes rather well-developed but not exceeding the perianth, styles 3, conical, 2-214 mm, recurved. Ripe cupule subsessile, saucer-shaped, 2—3 mm, 214-3 cm g; rim thin, covering the basal part of the fruit; wall woody, inside densely yellowish brown tomentose by adpressed, simple hairs, outside densely fulvous to greyish stellate hairy; scales imbricate, distinct and free. Ripe fruit depressed-subglobose, 114-2 cm long, 2-214 cm g, glabrous, dark chocolate- brown, glossy, top rounded-umbonate and de- pressed at the centre, base cordate, scar deeply concave, 114-124 cm @; wall woody, 1-2 mm thick, for the greater part free from the cupule; cotyledons flat-convex, ruminate. Distr. Malesia: Sumatra (rare). Ecol. Montane mossy forest, at 2400-3300 m. Fl. fr. May-July. Note. Records from the Philippines (ELMER, Leafl. Philip. Bot. 3, 1910, 942) and from Borneo (KING, /.c.) are not confirmed. ELMER had speci- mens of L. submonticolus, and according to STAPF, Trans. Linn. Soc. Lond. II, Bot. 4 (1894) 231, KinG had a specimen of Myrica. 79. Lithocarpus obtusifolius SoEPADMO, Reinward- tia 8 (1970) 263. — Quercus rufa VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 4, non VuKorT. 1889; in Fedde, Rep. 3 (1906) 173; MerRR. En. Born. (1921) 215. — Synaedrys rufa (VON SEEMEN) Ko1pz. Bot. Mag. Tokyo 30 (1916) 198. — L. rufa (VON SEEMEN) A. CAMUS, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 968, t. 463: 9-18. Tree. Branchlets, petiole, undersurface of leaves, fruiting rachis, and cupules densely velvety rufous hairy by erect and adpressed stellate hairs. Leaves thick-coriaceous, rigid, 9-13 by 5-714 cm (index 114-2), broadest slightly above the middle; sur- faces discolorous, above subglabrous, dark choco- late-brown; base acute, margin strongly recurved, top rounded to obtuse; midrib strongly prominent beneath, slightly so above; nerves 6-7 pairs, thin, prominent beneath, flat to impressed above, paral- lel, at an angle of c. 45°, arcuating and anastomos- ing near the margin; reticulation lax, subscalari- form, obscure; petiole 1-144 cm, %4-'4 cm g, adaxially flat. Ripe cupule sessile, solitary or in clusters of 2-3, 4-5 mm long, 15-16 mm 9g; rim thin, covering the basal part of the fruit, incurved; scales minute, adpressed, in more or less concentric rows. Ripe fruit ovoid-globose, c. 114 cm in size, glabrous, dull to glossy, chocolate-brown, top abruptly rounded-acute, base truncate, scar con- cave, c. 8 mm 9g; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Mt Klam in SW. Kalimantan, HALLIER 2481, /r.). Note. The inflorescences are still unknown. 80. Lithocarpus papillifer HATus. ex SoEPADMO, Reinwardtia 8 (1970) 265. Tree 15-25 m, 30-40 cm g; bark smooth, grey- brown. Branchlets initially densely set with fulvous to rufous, adpressed stellate hairs, later glabrous, greyish black, densely lenticellate; terminal buds ovoid, 2 by 2 mm, scales narrowly ovate. Stipules ovate or deltoid, 1-2 by 14-1 mm. Leaves thick- coriaceous, (4—)6—-10(—12) by (2-)3-4(-514) cm (index 114-214), broadest at or below the middle; surfaces more or less discolorous, above dark choc- olate-brown, dull to glossy, with some adpressed stellate hairs especially on the midrib and nerves, and blackish glandular papillae, underneath den- sely yellowish brown to rufous tomentose by ad- pressed, minute, stellate hairs; base acute, top acute or rounded-emarginate; midrib prominent beneath, slightly so above; nerves (5—)6—7(-8) pairs, prominent beneath, impressed above, paral- lel, at an angle of 45—50°, arcuating but not anasto- mosing towards the margin; reticulation fine, dense, scalariform, obscure to rather distinct be- neath; petiole 144-1 cm, 1-3 mm g, shallowly furrowed. Inflorescence male or androgynous, in densely subterminal, paniculate clusters, densely yellowish brown to rufous stellate hairy; bracts and bracteoles narrowly ovate, 1-114 by 14-1 mm. Male rachis c. 5 cm, 1-114 mm g; ¢ flowers in clusters of 2-3, rarely solitary, filaments 114-2 mm, anthers 0.25 mm long, pistillode globose, c. 1 mm @. Androgynous rachis 4-5 cm, 2 mm @; female flowers solitary, rarely in clusters of 2, staminodes rudimentary, styles 3, conical, 1-114 mm, recurved. Cupule (not fully ripe) sessile, saucer-shaped, 6 mm long, 10-15 mm g; rim thin, covering the basal part of the fruit; wall thin, densely fulvous tomentose on both sides; scales minute, adpressed, imbricate but concentrically set. Fruit (not yet ripe) ovoid-conical, c. 1 by 114 cm, glabrous, chocolate-brown, glossy, top acuminate, base ro- tundate, scar concave, c. 8 mm g; wall bony, thinner than '4 mm, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak, rare: Mt Mulu, Mt Kalulong 3° 14’ N, 114° 41’ E; W. Kutei in Central E. Kalimantan; Sabah: Mt Kina- balu, common). Ecol. Submontane forest, at 600-1800 m. Fi. June—Jan., fr. Febr.—June. Note. In SAN 29006 (with female flowers) the staminodes are absent, and the styles are up to 15. 81. Lithocarpus ferrugineus SOEPADMO, Reinward- tia 8 (1970) 242. — Quercus kunstleri (non KING ex Hook. f. ) VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 4; Merr. En. Born. (1921) 214. Tree 16-24 m, 20-30 cm g; buttresses c. 0.7 m tall, 1-2 m out, 10 cm thick; bark smooth, grey. Branchlets initially densely yellowish brown to- mentose by simple and stellate, adpressed hairs, later glabrous, greyish green, finely fissured, sparse - 1972] ly lenticellate; terminal buds ovoid-globose, 2-4 by 2-3 mm, scale ovate. Leaves thin-coriaceous, (13-)17—24(-26) by (3—)4~-5(-6) cm (index 3-5), broadest at or above the middle; surfaces con- colorous, glabrous, pale greenish brown, above dull, beneath glossy; base acute, top abruptly acute to 1-114 cm acuminate; midrib strongly prominent on both sides; nerves (7—)9—10 (—12) pairs, prominent beneath, impressed above, sub- parallel, at an angle of 30—50°, arcuating but not anastomosing towards the margin; reticulation lax, fine, subscalariform to irregular, distinct be- neath; petiole (14—)1—114(-134) cm, 114-2 mm g, adaxially flat. Inflorescence male, androgynous or mixed, sparsely to densely fulvous adpressed, stellate hairy; bracts and bracteoles ovate, 24-2 by %-1 mm. Male rachis c. 10 cm, 1 mm @g; 3 flowers solitary, filaments 3-4 mm, anthers 0.30 mm long, pistillode subglobose, 114-2 mm g. Androgynous or mixed rachis 10-15 cm, 2 mm g; female flowers solitary, staminodes well-developed and exceeding the perianth, styles 3, terete, 2-3 mm, connate at the base. Ripe cupule subsessile, cup-shaped, 14-1 cm long, 114-2 cm g; rim thin, covering 14-1, part of the fruit; wall woody, thin, inside densely silvery to fulvyous tomentose by adpressed, minute simple hairs, outside densely fulvous adpressed, stellate hairy; scales adpressed, imbricate. Ripe fruit ovoid, 114-2 cm in size, densely covered by rufous, adpressed, simple hairs, top rounded-acute, base rotundate, scar deeply concave, c. 1 cm 9; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak; SW. Kali- mantan; Sabah: Beaufort Hill, rare). Ecol. Forests, below 500 m, usually on ridges. FI. April-July, fr. Aug.—Dec. 82. Lithocarpus falconeri (KURZ) ReuHD. J. Arn. Arb. 10 (1929) 133; BARNETT, Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 333; A. Camus, Chénes 3 (1954) 955, t. 460: 1-10; SoppADMo, Reinwardtia 8 (1970) 241.— Quercus falconeri Kurz, J. As. Soc. Beng. 44, ii (1875) 197; For. Fl. Burma 2 (1877) 485; Hook. f. Fl. Br. Ind. 5 (1888) 608; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 42, t. 34; Corner, Ways. Trees (1940) 303. f. 96. — Quercus minor RIDL. J. Str. Br. R. As. Soc. n. 57 (1910) 95, non SARGENT, 1889, nec GANDOGER, 1890. — Pasa- nia falconeri (KURZ) SCHOTTKY, Bot. Jahrb. 47 (1912) 679; GamBLg, J. As. Soc. Beng. 75, ii (1915) 423. — Synaedrys falconeri (KURZ) Koz. Bot. Mag. Tokyo 30 (1916) 195. Tree 6-10 m. Branchlets initially densely set with fulvous erect simple hairs, later subglabrous, greyish to blackish brown, sparsely to densely lenticellate; terminal buds ovoid-ellipsoid, 1-2 by Y-*/, mm, scales ovate-acute, densely silvery pubescent by simple adpressed hairs. Stipules linear-acute, 15-20 by 1-114 mm. Leaves thin- coriaceous, (12—)20-30(—34) by (4~)5-8(-10) cm (index (214-)3-4(-414)), broadest at or rarely above the middle; surfaces more or less concolor- ous, glabrous, pale greyish green, above glossy, FAGACEAE (Soepadmo) 371 beneath dull; base acute, rarely rounded, top abruptly acute; midrib strongly prominent on both sides; nerves (12—)14-17(-20) pairs, strongly prominent beneath, impressed above, subparallel, at an angle of 45-60°, arcuating and anastomosing near the margin; reticulation lax, fine, scalariform, rather distinct beneath; petiole 34-2 cm, 2-4mm ag, adaxially flat. Inflorescence male or female, sparse- ly to densely fulvous stellate hairy; bracts and bracteoles ovate-acute, 24-3 by 74-1 mm. Male rachis 15-35 cm, 114-2 mm g; 3 flowers solitary or in clusters of 2—3, filaments 3-4 mm, anthers 0.3 mm long, pistillode subglobose, compressed, 2-2 mm g. Female rachis 10-15 cm, 3 mm 9; 2 flowers solitary, rarely in clusters of 2-3, stami- nodes rudimentary, styles 3-4, conical, 1-114 mm, recurved. Ripe cupule sessile, solitary, rarely in clusters of 2-3, saucer- to cup-shaped, 4%-1 cm long, 2-224 cm g; rim thin, undulate and recurved, covering 14—1% part of the fruit; wall woody, thin, inside densely fulyous tomentose by adpressed simple hairs, outside densely fulvous stellate hairy; scales adpressed, imbricate. Ripe fruit ovoid-ellip- soid, 2-214 cm long, 124-2 cm g, glabrous, pale chocolate-brown, glossy, top gradually rounded- acute, base rotundate, scar concave, 74-1 cm g; wall woody, c. 1 mm thick, for the greater part free from the cupule. Distr. Burma, Peninsular Siam, in Malesia: Malay Peninsula (Perlis, Kedah, Perak, rather common; also Langkawi Is.). Ecol. Primary forests, below 200 m. In Burma and Siam confined to the evergreen forest. Fi. Sept.—Nov., fr. Febr.—July. 83. Lithocarpus crassinervius (BL.) REHD. J. Arn. Arb. 1 (1919) 124; A. Camus, Chénes 3 (1954) 950, t. 457: 3-19; SoEPADMO, Reinwardtia 8 (1970) 232. — Quercus crassinervia BL. Mus. Bot. 1 (1850) 292; A. DC. Prod. 16, 2 (1864) 87 (dassinervia’); KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 87; von SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 13; K. & V. Bijdr. 10 (1904) 28; Koorp. Atlas 1 (1913) t. 63; BACKER & BAKH. f. FI. Java 2 (1965) 8. — Quercus pseudomolucca var. crassinervia (BL.) MIQ. FI. Ind. Bat. 1, 1 (1856) 849; WENziG, Jahrb. Bot. Gart. Berl. 4 (1886) 227. — Pasania crassinervia (BL.) Oerst. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 84. — Synaedrys crassinervia (BL.) KOIDZ. Bot. Mag. Tokyo 30 (1916) 194. Tree 15-20 m, 50 cm go. Branchlets initially densely set with fulvous, adpressed, stellate hairs, later glabrous, greyish, sparsely lenticellate; ter- minal buds ovoid-ellipsoid, 2-3 by 1-144 mm, scales narrowly ovate to linear, densely fulvous tomentose by adpressed, stellate hairs. Stipules linear-acute, 5—10 by 1-1!4 mm, rather long persis- tent. Leaves thin-coriaceous, (9—)15—20(—28) by (4-)5—7(—10) cm (index (1.7—)2-3(—3.7)), broadest at or above the middle; surfaces concolorous, glabrous, greyish green, dull; base acute, rarely rounded, margin recurved, top acute; midrib and nerves strongly prominent beneath, flat to im- pressed above; nerves (6—)9-10(—12) pairs, paral- lel, at an angle of 50—-60°, arcuating but not anasto- 372 FLORA MALESIANA [ser. 1, vou W7? mosing towards the margin; reticulation fine, dense subscalariform to irregular, rather distinct beneath; petiole 8-15 mm, 2-4 mm 9g, corky, adaxially flat. Male rachis 6-10 cm, 1-2 mm g, densely fulvous stellate hairy; bracts and bracteoles narrowly ovate 114-2 by 44-24 mm; ¢ flowers solitary, perianth strongly recurved, filaments 3-4 mm, anthers 0.3- 0.4 mm long, pistillode subglobose, 1-114 mm @. Fruiting rachis 15-35 cm, 5 mm @g, carrying a few solitary cupules. Young fruit: perianth strongly recurved, staminodes well-developed and exceed- ing the perianth, styles 3-4, terete, 2-214 mm, strongly recurved. Ripe cupule sessile, cup-shaped, *,-1 cm long, 2144 cm g; rim thin, occasionally recurved, covering the basal part of the fruit; wall woody, thin, inside densely silky, fulvous tomen- tose by minute, adpressed simple hairs, outside densely fulvous stellate hairy by adpressed hairs; scales thick, ovate, adpressed, imbricate. Ripe fruit ovoid-ellipsoid, 3-3'4 cm long, 2-214 cm @, glabrous, chocolate-brown, top abruptly rounded- acute, base truncate, scar concave to flat, 1-114 cm @; wall woody, 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Java (W. and Central parts, rare). Ecol. Primary forests, at 200-2000 m. Fr. Nov. —April. Vern. Pasang balung, p. djambé, wrakar, J, tangogo, S. 84. Lithocarpus kunstleri (KING ex Hook. f.) A. Camus, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 974, t. 465: 1-9; SoppADMo, Reinwardtia 8 (1970) 252. — Quercus kunstleri KING ex Hook. f. Fl. Br. Ind. 5 (1888) 606; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 40, t. 31; CorNER, Ways. Trees (1940) 303, f. 96. — Pasania kunstleri (KING ex Hook. f.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 419. — Synaedrys kunstleri (KING ex Hook. f.) KoipZ. Bot. Mag. Tokyo 30 (1916) 196. Tree 6-15 m, 15-35 cm g. Branchlets initially densely set with yellowish brown to fulvous erect, stellate hairs, later subglabrous, whitish grey to greyish brown, finely fissured or sparsely lenticel- late; terminal buds ovoid-ellipsoid, 3—5 by 2-3 mm, scales narrowly ovate-acute. Stipules narrowly ovate-acute to linear, 5 by 1 mm. Leaves thin- coriaceous, (714—)13-16(—20) by (3-)5-6(-7) cm (index (2-)214(-3)), broadest at or above the middle; surfaces concolorous, sparsely stellate pubescent on both sides, dark greyish green; base rounded to subcordate, top abruptly acute to 1 cm acuminate; midrib strongly prominent on both surfaces, densely fulvous stellate hairy; nerves 10- 13 pairs, strongly prominent beneath, impressed above, densely fulvous stellate hairy, subparallel, at an angle of 50—-60°, arcuating and anastomosing near the margin; reticulation fine, lax, subscalari- form, distinct beneath; petiole 2-5 mm, 1 14-2 mm @, terete or adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous stellate hairy; bract and bracteoles narrowly ovate-acute, Y,-1¥y by % mm. Male rachis 20-30 cm, 1-2 mm 9; 3 flowers solitary, filaments 3-4 mm, anthers 0.3—0.35 mm long, pistillode subglobose, 1 mm g. Androgynous or mixed rachis 10-15 cm, 1 mm @g; female flowers solitary, staminodes well-developed and exceeding the perianth, styles 3, terete, 114-2 mm, strongly recurved. Ripe cupule 4%—-Y% cm stalked, cup-shaped, 5-7 mm long, 13-15 mm 9; rim thin, entire or denticulate, covering the basal part of the fruit; wall woody, thin, inside dense- ly fulvous-tomentose by simple hairs, outside densely fulvous-tomentose by adpressed, stellate hairs; scales adpressed, distinct but not free, imbri- cate. Ripe fruit cylindrical, 244-3 cm long, 1-1!4 cm, glabrous, dull chocolate-brown, top rounded- acute, base truncate, scar flat, 74-34 cm 9; wall woody, 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Malay Peninsula (Langkawi, Perak, Selangor, Pahang, Johore). Ecol. Primary forests, below 300 m, sometimes also in secondary forest, in swampy places or on hill-sides. FJ. Sept._Nov., fr. Jan.—June. 85. Lithocarpus pseudokunstleri A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 10; Chénes 3 (1954) 976, t. 465: 10-17; SoppaApMmo, Reinwardtia 8 (1970) 268. Tree 10-25 m, 20-60 cm g; buttresses 14-1 m tall, 44-14 m out, 6 cm thick; bark greyish brown, scaly. Branchlets initially densely fulvous tomen- tose by stellate hairs, soon glabrous, dark greyish brown or yellowish grey, sparsely to densely lenti- cellate; terminal buds ovoid-ellipsoid, 2-4 by 1- 144 mm, scales narrowly ovate-acute. Stipules lanceolate to subulate, 2-5 by 1-2 mm. Leaves thin-coriaceous, (8—)11-15(-16) by (2'4-)4-6(- 614) cm (index (2—-)214-3(-34)), broadest at or above the middle; surfaces concolorous, glabrous, glossy, pale greyish green; base attenuate-acute, top abruptly acute to 1 cm acuminate, tip blunt or sharp; midrib strongly prominent on both sides; nerves 7-8 pairs, prominent beneath, flat above, subparallel, at an angle of 45—S0°, strongly arcuat- ing but mostly not anastomosing towards the margin; reticulation fine, dense, scalariform, more or less distinct on both surfaces; petiole 4-6 mm, 114-2 mm g, terete or adaxially flat, glabrous. Inflorescence male or androgynous, densely ful- vous stellate hairy, subglabrescent; bracts and bracteoles ovate-acute, 144-114 by %-% mm. Male rachis 5—25 cm, 1 mm 9; ¢ flowers in clusters of 2-3, stamens 8-12, filaments 2-214 mm, anthers 0.25-0.3 mm long, pistillode subglobose, 1 mm 9g. Androgynous rachis 5-10 cm, 1 mm @g; female flowers solitary, staminodes rather well-developed but not exceeding the perianth, styles 3, terete, 214-3 mm, recurved. Ripe cupule sessile, cup- shaped, 1-114 cm long, 124-214 cm g; rim thin, entire, covering the basal part of the fruit; wall woody, thin, inside densely greyish brown tomen- tose by adpressed, simple hairs, outside densely fulvous stellate hairy: scales adpressed, imbricate, rather woody and corky, occasionally fused at the base to form obscure lamellae. Ripe fruit cylin- drical, 3-414 cm long, 114-2 cm 9g, initially densely greyish tomentose by adpressed, minute simple hairs, soon glabrous, dull chocolate-brown, top 1972] abruptly rounded-acute, base truncate, scar flat, 1 cm g; wall woody, 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Borneo (Sarawak, rather com- mon; Kalimantan, rare; Sabah, rare). Ecol. Primary forests, from sea-level up to 1500 m, occasionally also in peat-swamp forest. Fl. April-May, fr. June—March. 86. Lithocarpus havilandii (STAPF) BARNETT, Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 176; A. CAMus, Chénes 3 (1954) 677; So—EPADMO, Reinwardtia 8 (1970) 245. — Quercus havilandii Starr, Trans. Linn. Soc. Lond. II, Bot. 4 (1894) 231, t. 18: A 1-4, p.p., excl. VIDAL 1864. — Quercus abendanonii VALETON, Icon. Bogor. 4 (1913) 179, t. 354. — L. abendanonii (VALETON) A. CAMus, Bull. Soc. FAGACEAE (Soepadmo) DIS Bot. Fr. 92 (1945) 84; Chénes 3 (1954) 908, t. 446: 15-21. t. 447. — Fig. 28 a-c. Tree or shrub, 2-25 m, 20-100 cm g; bark grey— brown, smooth to scaly, lenticellate. Branchlets densely set with yellowish brown to rufous short stellate hairs and erect glandular tuft—hairs with bulbous base, later subglabrous, greyish brown, sparsely lenticellate; terminal buds ovoid, 3 by 2 mm, scales narrowly ovate-acute. Stipules linear, 3-6 by 1-2 mm. Leaves coriaceous, rigid, (4—)5-8 (-9) by (3-)4(-S) cm (index (1.2—)1.3-1.8(-2)), broadest at or below the middle; surfaces discolor- ous, above pale to dark chocolate-brown, dull to glossy, sparsely pubescent, glabrescent, beneath with a thick cover of yellowish brown to rufous, adpressed stellate hairs and erect, glandular tuft- hairs; base acute, rounded or subcordate, margin Fig. 28. Lithocarpus havilandii (STAPF) BARNETT. a. Habit with fruiting rachis and a few ¢ rachis, x 24, b. erect glandular tuft-hairs on the underside of leaf, x 12, c. ripe fruit and cupule, x 24. — L. scortechi- nii (KING ex Hook. f.) A. Camus. d. Ripe cupule and fruit, < 24. — L. kingianus (GAMBLE) A. CAMUS. e. Young cupule and fruit, f. ripe cupule and fruit, both x 23 (a-b RSNB 5932, c Toxopgeus 15, d KING’s Coll. 2188, e Poore 1282, f MAHMUD 606). 374 FLORA MALESIANA [ser. I, vol) 7? recurved, top bluntly acute to 14-1 cm acuminate; midrib strongly prominent beneath, slightly so above; nerves (7—)8-10(—12) pairs, prominent be- neath, impressed above, subparallel, at an angle of 40-50°, arcuating but not anastomosing towards the margin; reticulation lax, subscalariform, ob- scure tor ather distinct beneath; petiole 6-12 mm, 1-2 mm 9g, terete or adaxially flat. Inflorescence male or androgynous, in dense subterminal or axil- lary paniculate clusters, densely fulvous to rufous stellate hairy, hairs glandular; bracts and bracte- oles ovate-acute, 1 by 24 mm. Male rachis 3-5 cm, 114-2 mm 9g; ¢ flowers solitary or in clusters of 2-3, filaments 214-3 mm, anthers 0.25 mm long, pistillode globose, 24-1 mm @. Androgynous rachis 3-4 cm, 2 mm 9g; female flowers solitary, staminodes rudimentary, styles 3, conical, 1-114 mm, connate at the base. Ripe cupule sessile, cup-shaped, 1-1!4 cm long, 2-214 cm 9; rim thin, covering c. 14 part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous to rufous stellate hairy; scales minute, adpressed or free, ovate, in concentric rows. Ripe fruit conical, 214- 3 cm long, 114-2 cm g, glabrous, dark chocolate- brown, top acute, base rotundate, scar flat to slightly convex, 1 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak: twice col- lected from Lawas area, 5th Division; Sabah: common on Mt Kinabalu), S. and Central Celebes (scattered on Mts Lompobatang, Rante Mario, and Lumut). Ecol. In montane mossy forests, at 1300-3000 m. Fl. fr. throughout the year, fruits ripe in June- July. 87. Lithocarpus sogerensis (S. MOoRE) MARKGR. ex A. Camus, Chénes 3 (1954) 795, t. 415: 12-14; SOEPADMO, Reinwardtia 8 (1970) 279. — Pasania sogerensis S. Moore, J. Bot. 61 (1923) Suppl. 54. Tree 36 m, 80 cm 9; bark pale brown, finely fissured. Branchlets initially densely set with ful- vous adpressed stellate hairs, later glabrous, grey- ish brown, sparsely lenticellate; terminal buds ovoid, 1-114 by 1 mm, scales ovate-acute. Stipules ovate-acute, 1 by 24 mm. Leaves thin-coriaceous, (6—-)8-10(-12) by (3-)4(—5) cm (index (2-)2.2-2.5 (—2.7)), broadest at or slightly below the middle; surfaces discolorous, above glabrous, chocolate- brown or greyish green, dull to glossy, beneath sparsely tomentose by pale greyish brown adpres- sed stellate hairs; base rounded-acute, top bluntly 4-1 cm acuminate; midrib thin, prominent be- neath, flat above; nerves 8-10 pairs, thin, promi- nent beneath, flat to impressed above, subparallel, at an angle of 45-5S0°, arcuating but not anasto- mosing towards the margin; reticulation fine, lax, subscalariform, distinct beneath; petiole 22-1 cm, 1 mm 9g, adaxially flat or furrowed. Male rachis 6-10 cm, 1-114 mm g, in dense paniculate clusters, densely fulvous tomentose by short stellate hairs; bracts and bracteoles ovate, 114-2 by 1 mm; 3 flowers in clusters of 3, perianth 5—7-lobed, recurved, stamens 10-12, filaments 2-214 mm, anthers 0.35 mm long, pistillode subglobose, 1 mm @. Ripe cupule solitary, rarely in clusters of 2-3, 2-1 cm stalked, cup-shaped 2 cm long, 2-3 cm 9, base gradually narrowed; rim thin, covering 4-% part of the fruit; scales mainly distinct on the upper half of the cupule, adpressed, set in concentric rows. Ripe fruit conical, 3 cm long, 24% cm g, glabrous, chocolate-brown, top acute, base rotun- date, scar conical, 2 cm 9; wall woody, 3 mm thick, for the greater part free from the cupule; cotyle- dons flat-convex. Distr. Malesia: New Guinea (scattered in the NE. parts). Ecol. Forests, at 900-1600 m. Fi/. March, fr. July-Oct. 88. Lithocarpus celebicus (M1Q.) REHD. J. Arn. Arb. 1 (1919) 123; A. Camus, Chénes 3 (1954) 903, t.; SOEPADMO, Reinwardtia 8 (1970) 227. — Quercus celebica Mi1Q. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 110; A. DC. Prod. 16, 2 (1864) 95; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 50, t. 45 A; Evmer, Leaf]. Philip. Bot. 3 (1910) 938; MERR. En. Philip. 2 (1923) 26. — Quercus llanosii A. DC. Prod. 16, 2 (1864) 97; Merr. Philip. J. Sc. 3 (1908) Bot. 323; En. Philip. 2 (1923) 28. — Cyclobalanus celebica (M1Q.) OERsT. Vid. Medd. Naturh. For. Kjobn. 8 (1867) 81. — Cyclobalanus llanosii (A. DC.) Oerst. /.c. 80; Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 375 (lanosii’). — Quercus com- Panoana VIDAL, Sinopsis Atlas (1883) 41, t. 92: D. — Quercus dalbertisii F. vy. M. Vict. Natural. 1 (1884) 124, ibid. 1 (1885) 167. — Quercus gulli- veri F. v. M. lc. 167. — Quercus pseudomolucca var. papuana O. Wars. Bot. Jahrb. 13 (1891) 286; K. Sco. & Laut. FI. Schutzgeb. (1901) 263. — Quercus brachyclada VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 7. — Quercus lipacon ELMER, Leafl. Philip. Bot. 6 (1913) 1983; Merr. En. Philip. 2 (1923) 27. — Synaedrys brachyclada (VON SEEMEN) Korpz. Bot. Mag. Tokyo 30 (1916) 194. — Synaedrys celebica (Mi1Q.) Koinz. l.c. 194. — Synaedrys d’albertisii (F. v. M.) Koz. Le. 194. — Synaedrys gulliveri (F. vy. M.) Kotz. L.c. 189. — Synaedrys llanosii (A. DC.) Korpz. lL.c. 196. — Quercus mabesae MeERR. Philip. J. Sc. 3 (1908) Bot. 7; En. Philip. 2 (1923) 28. — L. lipacon (ELMER) Reup. J. Arn. Arb. 1 (1919) 128; A. Ca- Mus, Chénes 3 (1954) 1114, t. — L. llanosii (A. DC.) Reup. J. Arn. Arb. 1 (1919) 128; A. CAmus, Chénes 3 (1954) 757, t. — Pasania aculeata MARK- GR. Bot. Jahrb. 59 (1924) 73, excl. SCHLECHTER 19296. — Pasania d’albertisii (F. v. M.) MARKGR. l.c. 74. — Pasania companoana (VIDAL) MARKGR. l.c. 78. — Pasania papuana (O. WARB.) MARKGR. Lc. 74. — L. aculeata (MARKGR.) REHD. J. Arn. Arb. 10 )(1929) 132; A. Camus) ‘@Gheness3 (1954) 914, t. — L. d’albertisii (F. v. M.) REHD. J. Arn. Arb. 10 (1929) 132; A. Camus, Chénes 3 (1954) 1154. — L. papuana (O. WARB.) REHD. J. Arn. Arb. 10 (1929) 133; A. Camus, Chénes 3 (1954) 1115, t. — L. brachyclada (VON SEEMEN) A. Camus, Bull. Soc. Bot. Fr. 90 (1943) 201; Chénes 3 (1954) 792, t. 413: 1-8. — L. mabesae 1972] FAGACEAE (Soepadmo) 375 (MerR.) A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 84; Chénes 3 (1954) 904, t. Tree 10—33 m, 30-100 cm 9; buttresses up to 1 m tall and out; bark grey-brown, fissured to scaly. Branchlets initially densely yellowish brown to rufous short stellate hairy, later glabrous, grey- brown, finely fissured, sparsely lenticellate; termi- nal buds ovoid, 3—5 by 2-3 mm, scales narrowly ovate-acute, densely tomentose by adpressed stel- late hairs. Stipules linear-acute, 3-4 by 14-1 mm. Leaves thin-coriaceous, (8—)12—16(—20) by (3—)4— 6(-8) cm (index (2—)214-3(—314)), broadest at or below the middle; surfaces discolorous, above greyish brown, dull to glossy, sparsely pubescent, glabrescent, beneath with a thin cover of yellowish brown to greyish adpressed, minute, stellate hairs, on the midrib and nerves sometimes also with some erect stellate hairs; base rounded-acute to cuneate, decurrent, occasionally asymmetrical, top abruptly or gradually 14-2 cm acuminate, tip blunt or sharp; midrib thin, prominent on both sides; nerves (6—)8-10(—12) pairs, flat on both surfaces, subparallel, at an angle of 40—50°, arcu- ating but not anastomosing towards the margin; reticulation fine, subscalariform, lax, obscure; petiole subglabrous, 14-1 cm, 2 mm @g, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, densely fulvous stellate hairy; bracts and bracteoles ovate-acute, 1-114 by 24-1 mm. Male rachis 10-20 cm, 1-114 mm 9; ¢ flowers in clusters of 3, filaments 3-4 mm, anthers 0.3—-0.35 mm long, pistillode globose, 1 mm g. Androgynous or mixed rachis 10-15 cm, 1-2 mm g; female flowers solitary, very rarely in clusters of 2-3, staminodes rudimen- tary, styles 3, terete, 114-2 mm, recurved. Ripe cupule sessile to 14 cm stalked, cup-shaped, 74-1 cm long, 2-3 cm g; rim thin, covering c. 14 part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely yellowish brown stellate hairy; scales ovate, adpressed, rather distinct but not free, in concentric rows or imbricate. Ripe fruit ovoid- conical, 2-2'4 cm long, 124-214 cm g, glabrous, chocolate-brown, top rounded-acute, rarely ro- tundate-apiculate, base rotundate, scar deeply concave, 1-114 cm g; wall woody, 2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines, Celebes (scattered), Moluccas, New Guinea (also including Goode- nough and Fergusson Is.). Ecol. Forests, at 20-1200 m, commonly below 800 m, on clayey soil. F/. Sept.-May, fr. June— April. In New Guinea usually as co-dominant forming an association with species of Anisoptera (Dipt.) and Eugenia (Myrt.). 89. Lithocarpus sundaicus (BL.) REHD. J. Arn. Arb. 1 (1919) 131; A. Camus, Chénes 3 (1954) 910, t. 448: 1-24; SoEpADMO, Reinwardtia 8 (1970) 282. — Quercus sundaica BL. Verh. Bat. Gen. K. & W. 9 (1823) 216; Bijdr. (1826) 520; Fl. Jav. Cupul. (1829) 11, t. 2, 3; Mig. FI. Ind. Bat. 1, 1 (1856) 850; A. DC. Prod. 16,2 (1864) 89; OupEM. Natuurk. Verh. Kon. Akad. 11 (1865) 11; Hook. f. FI. Br. Ind. 5 (1888) 611; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 51, t.47,48; K. & V. Bijdr. 10 (1904) 31; E_mer, Leaf]. Philip. Bot. 3 (1910) 941; Koorp. Atlas 1 (1913) t. 51; Merr. En. Philip. 2 (1923) 31; CorRNER, Ways. Trees (1940) 305; BACKER & BAKH. f. Fl. Java 2 (1965) 8. — Quercus pruinosa Bu. Verh. Bat. Gen. K. & W. 9 (1823) 217, t. 5; Bijdr. (1826) 521; Fl. Jav. Cupul. (1829) 9, t. 1, incl. var. B, l.c. 10; ENDL. Gen. Pl. Suppl. 4, 2 (1847) 28; Mia. FI. Ind. Bat. 1, 1 (1856) 850; Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 107; A. DC. Prod. 16, 2 (1864) 87; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 56, t. 53B; K. & V. Bijdr. 10 (1904) 33. — Castanea latifolia BL. Bijdr. (1826) 526. — Quercus muricata Rox. FI. Ind. ed. Carey 3 (1832) 635. — Quercus mappacea KorTH. Kruidk. (1844) 202; Mia. FI. Ind. Bat. 1, 1 (1856) 850. — Quercus korthalsii BL. Mus. Bot. 1 (1850) 292, non ENDL. 1847; A. DC. Prod. 16, 2 (1864) 89; OUDEM. Natuurk. Verh. Kon. Akad. 11 (1865) il. — Quer- cus korthalsii BL. var. kajan BL. Mus. Bot. 1 (1850) 293; A. DC. Prod. 16, 2 (1864) 89. — Quercus korthalsii BL. var. mappacea (KORTH.) BL. Mus. Bot. 1 (1850) 293; A. DC. Prod. 16, 2 (1864) 90. — Quercus korthalsii BL. var. pachyphylla BL. Mus. Bot. 1 (1850) 293; Mia. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 107; A. DC. Prod. 16, 2 (1864) 90. — Quercus pruinosa var. alpina JUNGH. Java ed. 2, 1 (1853) 496. — Quercus lamponga Mia. Sumatra (1861) 348; A. DC. Prod. 16, 2 (1864) 95; WeENzic, Jahrb. Bot. Gart. Berl. 4 (1886) 229; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 53, t. 49; CoRNER, Ways. Trees (1940) 303, f. 96. — Cyclobalanus lamponga (MIQ.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Pasania pruinosa (Bv.) OERST. I.c. 83; S. Moore, J. Bot. 63 (1925) Suppl. 114. — Pasania sundaica (BL.) Oerst. Vid. Medd. Naturh. For. Kj6bn. 8 (1867) 83; GAMBLE, J. As. Soc. Beng. 75, ii (1915) 426; S. Moore, J. Bot. 63 (1925) Suppl. 114. — Pasania korthalsii (BL.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83. — Cyclobalanopsis muricata (ROXxB.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. — Quercus pseudo- molucca var. korthalsii (BL.) WENZ1iG, Jahrb. Bot. Gart. Berl. 4 (1886) 227. — Quercus pseudomo- lucca var. pruinosa (BL.) WENZIG, lI.c. 227. — Quercus pseudomolucca var. sundaica (BL.) WEN- z1G, l.c. 227. — Quercus grandifrons KING ex HOOK. f. Fl. Br. Ind. 5 (1888) 610; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 49, t. 35 B; CoRNER, Ways. Trees (1940) 303. — Pasania lamponga (MIQ.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 423, excl. var. — Pasania grandifrons (KING ex Hook. /.) GAMBLE, /.c. 421. — Synaedrys grandifrons (KING ex Hook. f.) Kompz. Bot. Mag. Tokyo 30 (1916) 195. — Synaedrys lamponga (MiqQ.) Kotpz. l.c. 196. — Synaedrys pruinosa (BL.) Korpz. I.c. 197. — Synaedrys sundaica (BL.) Ko1pz. l.c. 198. — L. lamponga (Miq.) ReEuD. J. Arn. Arb. 1 (1919) 128; A. Camus, Chénes 3 (1954) 1107, t. 504: 9-20. — L. pruinosa (BL.) REHD. J. Arn. Arb. 1 (1919) 130; A. Camus, Chénes 3 (1954) 673, t. 383, 384: 1-16. 376 — Quercus hystrix var. mappacea (KORTH.) MERR. En. Born. (1921) 213. — L. grandifrons (KING ex Hook. f.) A. CAMus, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 1122, t. 507: 7-18. Tree 10-36 m, 20-90 cm g; bark grey-brown, fissured to scaly. Branchlets initially densely yellow- ish brown to fulvous tomentose by adpressed stellate hairs and woolly or erect tuft-hairs, later subglabrous, dark greyish brown, with numerous minute lenticels; terminal buds ovoid, 3—5 by 2-3 mm, scales narrowly ovate. Stipules narrowly ovate to linear, 5—10 by 2-3 mm. Leaves coriaceous (10—)12-16(-24) by (4-)5-6(—10) cm (index (2-) 2'4-(-3)), broadest at or below the middle; surfaces discolorous, above greyish brown to dark choco- late-brown, initially with some woolly or erect tuft-hairs especially on the midrib and nerves, soon glabrescent, beneath densely yellowish brown to fulvous tomentose by adpressed stellate hairs interspersed with woolly or erect tuft-hairs; base rounded-acute, rarely acute, top acute to 14-2 cm acuminate, tip blunt or sharp; midrib strongly prominent on both sides; nerves (10-)12-14(-16) pairs, prominent beneath, flat above, subparallel, at an angle of 45-60°, arcuating but not anasto- mosing towards the margin; reticulation fine, lax, scalariform, obscure to distinct beneath; petiole 5-12 mm, 114-3 mm 4g, initially densely stellate hairy, glabrescent, adaxially flat or furrowed. Inflorescence male, androgynous or mixed, in dense, subterminal paniculate clusters, densely yellowish brown to fulvous stellate hairy; bracts and bracteoles narrowly ovate-acute, 1-114 by Y%-2, mm. Male rachis 10-15 cm, 1-114 mm 9; 6 flowers in clusters of 3, stamens 10-12, filaments 214-3 mm, anthers 0.25-0.3 mm long, pistillode globose, 1 mm g. Androgynous or mixed rachis 10-15 cm, 114-2 mm 9g; female flowers solitary or rarely in clusters of 2-3, staminodes rudimentary, styles 3-4, terete, 1-2 mm, recurved. Ripe cupule sessile to 14-1 cm stalked, saucer-shaped, 3-6 mm long, (2—)244(-3) cm 9; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely fulvous-tomentose by adpressed, simple hairs, outside densely fulvous stellate hairy; scales adpressed, ovate, imbricate but concentrically set. Ripe fruit depressed-ovoid, 114-2 cm long, (2-)214 —214(-3) cm g, glabrous, dark chocolate-brown to purplish brown, top rounded-acute, base ro- tundate, scar concave to flat, 114-2 cm g; wall woody, 1—2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Peninsular Siam (rare), in Malesia: Sumatra (scattered), Malay Peninsula (common; also in Penang and Singapore), Java (common in the western part, scattered in the central and eastern parts, eastwards to Mts Ardjuno, Kawi, Tengger and Idjen), Borneo (scattered in Sarawak, Kalimantan, and Sabah), Philippines (Luzon, Mindoro, rare). Ecol. Primary forests, from sea-level up to 2600 m, more commonly between 500-1500 m. In Central and E. Java confined to pockets of FLORA MALESIANA [ser. I, walt w# everwet forest at higher altitude. F/. Sept.—July, fr. Aug.—March. Notes. The typical form of this species occurs at 1000-1500 m, and is characterized by the rela- tively thin leaves with a thin indumentum. Speci- mens from higher localities (Quercus pruinosa) usually have a somewhat smaller but thicker leaf with a thicker indumentum, and larger cupule and fruit than those of the typical form. Specimens from below 500 m, formerly included in Quercus mappacea and Quercus grandifrons, have larger leaves with less indumentum, smaller cupule and fruit than those of L. sundaicus. The nervation, reticulation, the type of indumentum, fruit and cupule are, however, very similar in the whole series of specimens. The Sumatran specimens, formerly recognized as Quercus lamponga, have smaller, almost gla- brous leaves, but again the type of the cupule and fruit is the same. 90. Lithocarpus pseudomoluccus (BL.) REHD. J. Arn. Arb. 1 (1919) 130; A. Camus, Chénes 3 (1954) 947, t. 456: 8-24; SoppADMO, Reinwardtia 8 (1970) 269. — Quercus pseudomolucca BL. Verh. Bat. Gen. K. & W. 9 (1823) 214, t. 4; Bijdr. (1826) 519; Fl. Jav. Cupul. (1829) 14, t. 6; OuDEM. Na- tuurk. Verh. Kon. Akad. 11 (1865) 8, t. 5; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 43, t. 36; K. & V. Bijdr. 10 (1904) 26; Koorp. Atlas 1 (1913) t. 57; BACKER & BAKH. f. Fl. Java 2 (1965) 8. — Quercus angustata BL. Verh. Bat. Gen. K. & W. 9 (1823) 212, t. 3; Bijdr. (1826) 520; Fl. Jav. Cupul. (1829) 15, t. 7. — Quercus pseudomolucca var. angustata (BL.) BL. Mus. Bot. 1 (1850) 292; Mia. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 108; A. DC. Prod. 16, 2 (1864) 86; OupEM. Natuurk. Verh. Kon. Akad. 11 (1865) 9. — Quercus thelecarpa Mia. Pl. Jungh. (1851) 9; Fl. Ind. Bat. 1, 1 (1856) 851. — Quercus thelecarpa var. angustata (BL.) Mia. PI. Jungh. (1851) 9; Mra. Fl. Ind. Bat. 1, 1 (1856) 852. — Quercus neurophylla Mia. Sumatra (1861) 351. — Pasania pseudomolucca (BL.) OERST. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 83. — Pasania neurophylla (M1Q.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. — Synaedrys pseudomolucca (BL.) Korpz. Bot. Mag. Tokyo 30 (1916) 197. Tree 15-25 m, 60 cm g; bark grey, fissured; stilt-roots and aerial roots occasionally pre- sent. Branchlets initially densely fulvous stellate hairy, later glabrous, greyish to blackish brown, with many large lenticels; terminal buds ovoid, 3-5 by 3-4 mm, scales ovate to linear. Stipu- les linear-acute, 4-8 by 1-114 mm. Leaves coriaceous, (8—)13-17(-22) by (2!4-) 5-6 (-8) cm (index 2-3), broadest about the middle; surfaces more or less discolorous, above glabrous, pale greyish green, glossy, underneath densely greyish to glaucous tomentose by adpressed stel- late hairs; base rounded-acute to acute, margin recurved, top bluntly acute to sharply 1-2 cm acuminate; midrib strongly prominent on both sides; nerves 10-12 pairs, prominent beneath, flat above, subparallel, at an angle of 45—-60°, arcuating 1972] FAGACEAE (Soepadmo) eM faintly anastomosing near the margin; reticulation subscalariform to irregular, distinct beneath; pe- tiole 24-2 cm, 1-2 mm 9, adaxially flat or furrow- ed. Inflorescence male, female or mixed, densely fulvous stellate hairy; bracts and bracteoles nar- rowly ovate, 1-4 by 24-1 mm. Male rachis 10-20 cm, 114-2 mm g; ¢ flowers in clusters of 3, or solitary, filaments 3-4 mm, anthers 0.3-0.35 mm long, pistillode subglobose, 1-114 mm 9. Female or mixed rachis 10-20 cm, 2 mm g; & flowers solitary or rarely in clusters of 3, staminodes rudi- mentary, styles 3, conical, 114-2 mm. Ripe cupule sessile, saucer-shaped, 3-8 mm long, (2—)214-3 (-314) cm g; rim thin, covering the basal part of fruit; wall woody, thin, inside densely greyish brown tomentose by adpressed, simple hairs, out- side densely fulvous stellate hairy; scales distinct, imbricate or occasionally in more or less concen- tric rows. Ripe fruit depressed-subglobose or ovoid, 114-2 cm long, 2-3 cm g, glabrous except around the umbo, top acute or rounded and de- pressed at the centre, base cordate, scar flat, 114- 2\%4 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (rare), Java (mainly in the western part, eastwards to Mt Jang at I} SiO 12). Ecol. Forests, at 600-1700 m. F/. April-June, fr. July—Dec. Note. Records from the Malay Peninsula re- late to L. rassa, and from New Guinea to L. mega- carpus. 91. Lithocarpus submonticolus (ELMER) REHD. J. Arn. Arb. 1 (1919) 131; A. Camus, Chénes 3 (1954) 972; SopPADMO, Reinwardtia 8 (1970) 280. — Quercus submonticola ELMER, Leaf]. Philip. Bot. 3 (1910) 943; Merr. En. Philip. 2 (1923) 31. — Quercus monticola (non KING, nec PETZ. & KIRCHN.) ELmer, Leaf]. Philip. Bot. 3 (1910) 942; ibid. 6 (1913) 1982; MERR. En. Philip. 2 (1923) 29. Tree 7-13 m, 10-15 cm 9; bark grey, smooth. Branchlets initially densely tomentose by adpressed stellate hairs, later glabrous, greyish to blackish brown, densely lenticellate; terminal buds ovoid- ellipsoid, 2-4 by 1-2 mm, scales narrowly ovate- acute. Stipules linear-acute, 4-5 by 1 mm. Leaves thin-coriaceous, (3—)6-8(-10) by (2—)3—314(-4) cm (index (2-)2!4(-3)), broadest at or below the middle; above glabrous, greyish green to choco- late-brown, dull to glossy, beneath with a thin cover of adpressed, minute stellate hairs, some- times interspersed with some erect stellate hairs especially on the midrib and nerves, dull pale brown to greyish brown; base rounded-acute to acute, top rounded-acute to 1 cm acuminate, tip usually oblique; midrib prominent on both sides; nerves 6-8 pairs, thin, flat on both surfaces to impressed above, subparallel, at an angle of 45— 60°, arcuating but not anastomosing towards the margin; reticulation subscalariform to irregular, obscure; petiole 5-8 mm, 1-114 mm g, adaxially flat or furrowed. Inflorescence male or androgynous, simple and axillary or much-branched and sub- terminal, densely fulvous stellate hairy; bract and bracteoles narrowly ovate-acute, 2-3 by 24-1 mm. Male rachis 5-10 cm, 1-114 mm g; ¢ flowers in clusters of 3, filaments 3-314 mm, anthers 0.3 mm long, pistillode globose, 24-1 mm g@. Androgynous rachis 3-8 cm, 114 mm @; female flowers solitary, staminodes rather well-developed but not exceed- ing the perianth, styles 3, conical, 1-144 mm, recurved. Ripe cupule \4—%, cm stalked, cup- shaped, 24-1 cm long, 114-2 cm g; rim thick, covering 14-1, part of the fruit; wall woody, 1-2 mm thick, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous tomentose to adpressed, minute, rather distinct on the upper half of the cupule, imbricate but set in concentric rows. Ripe fruit cylindrical to depressed subglobose, 114-214 cm long, 114-134 cmg, dense- ly fulvous tomentose by adpressed simple hairs, top acute or rounded-umbonate, base truncate, scar flat, 1-114 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Philippines (Luzon, common; Mindoro, rare). Ecol. Forests, at 1000-1300 m. Fi/. Sept._May, fr. June—Dec. 92. Lithocarpus castellarnauianus (VIDAL) A. CA- Mus, Bull. Soc. Bot. Fr. 92 (1945) 84; Rev. Bot. Appl. Trop. 25 (1945) 32; Chénes 3 (1954) 817; SOEPADMO, Reinwardtia 8 (1970) 226. — Quercus castellarnauiana VIDAL, Rev. Pl. Vasc. Filip. (1886) 264; MeErRR. Philip. J. Sc. 3 (1908) Bot. 326; En. Philip. 2 (1923) 26. — Quercus merrittii MERR. Philip. J. Sc. 3 (1908) Bot. 325; En. Philip. 2 (1923) 28. — Quercus obliquinervia MERR. Philip. J. Sc. 4 (1909) Bot. 250; En. Philip. 2 (1923) 29. — Synaedrys castellarnauiana (VIDAL) Koipz. Bot. Mag. Tokyo 30 (1916) 190. — Synaedrys merrittii (MErRR.) Korpz. /.c. 192. — Quercus cagayanensis Merk. Philip. J. Sc. 13 (1918) Bot. 6; En. Philip. 2 (1923) 26. — L. merrittii (MERR.) REHD. J. Arn. Arb. 1 (1919) 128; A. Camus, Chénes 3 (1954) 818. — L. obliquinervia (MERR.) A. CAMus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 818. — Quercus pinatubensis ELMER, Leaf]. Philip. Bot. 9 (1934) 3188. — L. cagayanensis (MERR.) A. CA- Mus, Bull. Soc. Bot. Fr. 92 (1945) 256; Chénes 3 (1954) 819. — L. pinatubensis (ELMER) A. CAMUS, Chénes, Atlas 3 (1949) 117, t. 518: 15-18; Chénes 3 (1954) 804. Tree 10-20 m, 20-30 cm g. Branchlets initially densely fulvous to rufous tomentose by adpressed stellate hairs, later glabrous, greyish, sparsely to densely lenticellate; terminal buds ovoid, 3-4 by 2-3 mm, scales narrowly ovate-acute. Stipules ovate to lanceolate, 3-5 by 1-3 mm. Leaves thin- coriaceous, (914-)13-17(-21) by (31%4-)414-6 (714) cm (index 2.3-3), broadest at or rarely above the middle; above glabrous, dull to glossy, greyish green to chocolate-brown, beneath with a thin cover of greyish to glaucous adpressed, minute stellate hairs, subglabrescent; base rounded-acute to attenuate-acute, top 1-214 cm acuminate, tip sharp, oblique; midrib prominent beneath, slightly so above, rather broad especially near the base; nerves (9—)10(—11) pairs, thin, prominent beneath, FLORA MALESIANA (ser. Te yvalry? Fig. 29. Lithocarpus leptogyne (KORTH.) SOEPADMoO. A stilt-rooted tree locally common in Johore, S. Malaya (CORNER SF 28959; 1935). 1972] FAGACEAE (Soepadmo) 379 flat above, subparallel, at an angle of 45-60°, arcu- ating but not anastomosing towards the margin; reticulation dense, subscalariform, obscure to dis- tinct beneath; petiole 7-11 mm, 1—3 mm 9, adaxi- ally flat. Inflorescence male or androgynous, densely yellowish brown to fulvous stellate hairy; bracts and bracteoles ovate-acute, 14-1 by 14 mm. Male rachis 10-20 cm, 2 mm 9; 3 flowers in clusters of 3, filaments 3-4 mm, anthers 0.3 mm long, pistillode globose, 1 mm @. Androgynous rachis 10 cm, 114-2 mm 9; female flowers solitary, rarely in clusters of 2, staminodes well-developed but not exceeding the perianth, styles 3, conical, 1 mm, strongly recurved. Ripe cupule 1 cm stalked, cup- shaped, 24-1 cm long, 2-214 cm 9; rim thin, covering 4-1 part of the fruit; scales ovate, minute, adpressed, in more or less concentric rows, distinct on the upper half of the cupule only. Ripe fruit cylindrical or ovoid, 2-274 cm long, 174-2 cm @, densely fulvous tomentose by adpressed, simple hairs, top rounded-acute, base rounded, scar strongly convex, 1-114 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, common; Mindoro, Marinduque, Samar, Leyte, rare). Ecol. Forests, at 200-2200 m. Fl. Jan.—May, fr. June—Dec. 93. Lithocarpus leptogyne (KoRTH.) SOEPADMO, Reinwardtia 8 (1970) 254. — Quercus leptogyne Kortu. Kruidk. (1844) 206; A. DC. Prod. 16, 2 (1864) 93; A. Camus, Chénes 3 (1954) 1160. — Cyclobalanus leptogyne (KORTH.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 376. — Quercus lamponga (non Miq.) Hook. f. Fl. Br. Ind. 5 (1888) 611. — Quercus conocarpa (non OUDEM.) Merkr. PI. Elm. Born. (1929) 43. — Fig. 29. Tree 10—40 m, 10—90 cm g; buttresses up to 114 m tall, 114 m out, 10 cm thick; bark pale grey- brown, lenticellate. Branchlets initially densely fulvous tomentose by stellate hairs, later subgla- brous, sparsely lenticellate; terminal buds ovoid- ellipsoid, 3-5 by 2-3 mm, scales narrowly ovate- acute. Stipules linear, 4-5 by 24-1 mm. Leaves thin-coriaceous, (8—)12-15(-18) by (3-)314-5 (—514) cm (index (2!4—)2.7-3.3 (-3.6)), broadest at or above the middle; surfaces more or less discolor- ous, above} dull, greyish green to pale chocolate- brown, sparsely stellate hairy especially on the midrib and nerves, underneath with a thin cover of pale yellowish brown to pale greyish green, adpressed, minute stellate hairs; base acute to rarely rounded, usually asymmetrical, top abrupt- ly 1-114 cm acuminate; midrib prominent on both sides; nerves 11-14 pairs, flat beneath, impressed above, subparallel at an angle of 45-60°, arcuating but not anastomosing towards the margin; reticu- lation fine, lax, subscalariform, obscure to rather distinct beneath; petiole 3-6 mm, 1—2 mm g, dense- ly tomentose by stellate hairs, shallowly furrow- ed. Inflorescence male, androgynous or mixed, densely greyish brown stellate hairy; bracts and bracteoles ovate, 1-114 by 24-1 mm. Male rachis 10-15 cm, 1 mm g, usually much-branched; 3 flow- ers in clusters of 3, filaments 3-4 mm, anthers 0.35 mm long, pistillode subglobose, 1-114 mm g@. Androgynous or mixed rachis 5-10 cm, 1 mm @; female flowers solitary, staminodes rudimentary or well-developed and exceeding the perianth, styles 3, terete, 1-114 mm, recurved. Ripe cupule ¥, cm stalked, saucer-shaped, 3-6 mm long, 1.3— 2 cm @; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely fulvous- tomentose by adpressed simple hairs, outside densely fulvous stellate hairy; scales adpressed, in 6-10 concentric rows. Ripe fruit depressed ovoid, 1.2-1.7 cm long, 114-2 cm @, densely fulvous to- mentose by adpressed simple hairs, soon glabres- cent, top rounded-acute, base rounded, scar deeply concave to rarely flat, 1 cm @; wall woody, 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (rare), Malay Penin- sula (scattered), Borneo (rather common in Sara- wak and Sabah). Ecol. Forests, from sea-level up to 1500 m. Fl. Aug.—July, fr. Sept.-May. 94. Lithocarpus nieuwenhuisii (VON SEEMEN) A. Camus, Bull. Soc. Bot. Fr. 92 (1945) 255; Chénes 3 (1954) 732, t.; SoEPADMO, Reinwardtia 8 (1970) 262. — Quercus nieuwenhuisii VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 6. — Quercus clementis MERR. Philip. J. Sc. 3 (1908) Bot. 321; En. Philip. 2 (1923) 26. — Pasania clementis (MerRR.) SCHOTTKY, Bot. Jahrb. 49 (1913) 358. — Pasania ochracea SCHOTTKY, l.c. 357. — Synaedrys ‘clementi’ (MERR.) Koipz. Bot. Mag. Tokyo 30 (1916) 194. — Synaedrys ‘nieawenhuisii’ (VON SEEMEN) Korpz. /.c. 197. — Quercus ochracea (SCHOTTKY) MerR. En. Born. (1921) 214. — Quer- cus borneensis MERR. Philip. J. Sc. 21 (1922) 516; Pl. Elm. Born. (1929) 42. — L. borneensis (MERR.) Reup. J. Arn. Arb. 10 (1929) 132; A. Camus, Chénes 3 (1954) 956, t. 460: 11-19. — L. clementis (MerR.) A. Camus, Riviera Scient. 18 (1932) 39; Chénes 3 (1954) 983, t. 467: 16-22. — L. ochracea (ScHoTTKy) A. Camus, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 985. — L. crateriformis MERR. ex A. CAMus, Chénes, Atlas 3 (1949) 97, t.; Chénes 3 (1954) 967. Tree 5-25 m, 15-70 cm 9; buttresses and stilt- roots sometimes present; bark greyish brown, scaly, lenticellate. Branchlets initially densely yel- lowish brown to fulvous or rufous tomentose by long, erect simple hairs and adpressed stellate hairs, later subglabrous, greyish to blackish brown, sparsely lenticellate; terminal buds ovoid, 3-5 by 2-3 mm, scales narrowly ovate to linear. Stipules ovate to lanceolate, 5-6 by 2-3 mm. Leaves char- taceous to coriaceous, (9—)14—-20(-23) by (3-)4-6 (-8) cm (index (2.6-)3-3!4(-4)), broadest about the middle; surfaces discolorous, above glabrous, except the midrib and nerves, dull greyish green, rarely glossy, underneath dull chocolate-brown, rarely glossy, densely yellowish brown to rufous hairy by erect simple hairs and adpressed stellate hairs, glabrescent except the midrib and nerves; base attenuate-acute to rounded-acute, occasion- ally asymmetrical, top bluntly acute with emargin- 380 ate tip to sharply or bluntly 1-2 cm acuminate; midrib strongly prominent beneath, slightly so above; nerves (9—)10(—14) pairs, strongly promi- nent beneath, impressed above, subparallel, at an angle of 45—60°, arcuating and anastomosing near the margin; reticulation lax, subscalariform, dis- tinct beneath; petiole densely fulvous to rufous stellate hairy, (3—)5—10(—12) mm, 1—3 mm 9g, terete or adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous to rufous stellate hairy; bracts and bracteoles ovate, 1-2 by 14-1 mm. Male rachis 10-15 cm, 1-114 mm g; ¢ flowers solitary, filaments 3-4 mm, anthers 0.3-0.35 mm long, pistillode subglobose, 1-114 mm g. Andro- gynous or mixed rachis 10-20 cm, 114-2 mm g; female flowers solitary, staminodes rather well- developed but not exceeding the perianth, styles 3, terete, 2-3 mm, strongly recurved. Ripe cupule subsessile, cup-shaped, 114-2 cm long, 2-224 cm @;rim thin, erect and entire or undulate and recury- ed, covering 14—1% part of the fruit; wall woody, thin, inside densely fulvous to rufous tomentose by adpressed stellate hairs, outside densely fulvous to rufous-tomentose by adpressed stellate hairs; scales minute, adpressed, imbricate or occasionally in more or less concentric rows. Ripe fruit ovoid- globose to cylindrical, 2-3 cm long, 114-2 cm @, densely greyish brown tomentose by adpressed simple hairs, subglabrescent, top rounded-acute to rotundate-apiculate, base truncate, scar deeply concave to flat, 144-1 cm g; wall woody, 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (Sarawak, Kalimantan, Sabah, Brunei), also Nunukan Is.; Philippines (Mindanao, Basilan, rare). Ecol. Forests, at low altitude, sometimes also in peat-swamp or heath forest, on sandy clayey to ultra-basic soil; usually along stream. Fi. fr. throughout the year. 95. Lithocarpus curtisii (KING ex Hook. f.) A. Camus, Riviera Scient. 18 (1932) 40; Chénes 3 (1954) 1104, t. 503: 9-12; BARN. Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 333; SoppapMo, Reinwardtia 8 (1970) 233. — Quercus curtisii KING ex Hook. f. Fl. Br. Ind. 5 (1888) 612; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 56, t. 52. — Pasania curtisii (KING ex Hook. f.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 429. — Synaedrys curtisii (KING ex Hook. J.) Kotpz. Bot. Mag. Tokyo 30 (1916) 194. Tree 6-27 m, 25-60 cm 9; bark pale grey, lenti- cellate. Branchlets initially densely fulvous to rufous tomentose by adpressed stellate hairs, later glabrous, greyish brown, sparsely lenticellate; terminal buds ovoid, 1-114 by 24 mm, scales nar- rowly ovate. Leaves thick-coriaceous, (12—)15-22 (24) by (4-)5-7(-10) cm (index (214-)3-3 14 (-4)), broadest at or below the middle; surfaces more or less concolorous, above glabrous, dull to glossy, pale greyish brown, underneath densely tomentose by adpressed minute, stellate hairs, glabrescent; base acute, rarely rounded, top bluntly or sharply 1-2 cm acuminate; midrib strongly prominent on both sides; nerves 9-11 pairs, prominent beneath, impressed above, subparallel, at an angle of 45-— FLORA MALESIANA [ser. 1, wolt 7? 60°, arcuating but not anastomosing towards the margin; reticulation fine, subscalariform to irregular, obscure; petiole /,-1 cm, 114-214 mma, adaxially flat. Inflorescence male, androgynous or mixed, densely fulvous stellate hairy; bracts and bracteoles ovate-acute, 24-1 by 14 mm. Male rachis 5-15 cm, 1-114 mm 9g; ¢ flowers solitary, or rarely in clusters of 2-3, perianth 5—6—lobed, stamens 10-12, filaments 214-3 mm, anthers 0.3 mm long, pistillode subglobose, 1 mm @. Andro- gynous or mixed rachis 15-25 cm, 1144-2 mm 9; female flowers solitary, staminodes well-developed and exceeding the perianth, occasionally pollini- ferous, styles 3, conical, 1-114 mm, recurved. Ripe cupule sessile, saucer-shaped, 3-5 mm long, 2-274 cm g; rim thin, undulate and recurved, covering the basal part of the fruit; wall woody, thin, inside densely greyish tomentose by adpres- sed simple hairs, outside densely fulvous stellate hairy; scales adpressed, obscure. Ripe fruit ovoid- globose, 124-214 cm long, 114-2 cm g, densely greyish tomentose by adpressed simple hairs, subglabrescent, top rounded-acute, base cordate, scar deeply concave, 1 cm g; wall bony, thinner than 1 mm, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (Perak, Se- langor, Pahang, Trengganu, Kelantan, also Pe- nang I.). Ecol. Forests, below 300 m, on clayey soil. Fl. fr. June—Nov. 96. Lithocarpus scortechinii (KING ex HOooK. /.) A. Camus, Riviera Scient. 18 (1932) 42; Chénes 3 (1954) 894, t. 442: 5-10; BARN. Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 333; SOEPADMO, Rein- wardtia 8 (1970) 278. — Quercus scortechinii KING ex Hook. f. Fl. Br. Ind. 5 (1888) 608; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 43, t. 35A; CORNER, Ways. Trees (1940) 304, f. 96. — Pasania scortechi- nii (KING ex Hook. f.) ScHoTTKy, Bot. Jahrb. 47 (1912) 676; GamBte, J. As. Soc. Beng. 75, ii (1915) 418.— Synaedrys scortechinii (KING ex Hook. f.) Korpz. Bot. Mag. Tokyo 30 (1916) 198. — L. eriolepis A. CAmus, Bull. Soc. Bot. Fr. 84 (1937) 176; Chénes 3 (1954) 888, t. 448: 25-27. — L. smitinandiana A. CAMus, Not. Syst. 14 (1952) 257; Chénes 3 (1954) 1270, f. 28: 5-8. — Fig. 28d. Tree 15-20 m, 30-90 cm 9g. Branchlets initially densely yellowish brown to fulvous tomentose by woolly stellate hairs, later glabrous, greyish brown, sparsely to densely lenticellate; terminal buds ellipsoid, 5-8 by 2-3 mm, scales linear-acute to lanceolate, densely yellowish brown erect stellate hairy. Stipules linear-acute, 5—6 by 1-2 mm. Leaves thick-coriaceous, rigid, (10-)14-16(-20) by 3-6 cm (index (214-)3(—5)), broadest at or rarely below the middle; surfaces concolorous, glabrous, pale greyish green, dull beneath, glossy above; base acute, top bluntly acute to rounded, rarely 1-2 cm acuminate; midrib strongly prominent on both sides; nerves 8-10 pairs, thin, prominent beneath, impressed above, subparallel, at an angle of 45-50°, arcuating but not anastomosing to- 1972] FAGACEAE (Soepadmo) 381 wards the margin; reticulation fine, subscalariform | to irregular, obscure; petiole 10-13 mm, 2 mm 9, adaxially flat, towards the base gradually thicken- ed and corky. Inflorescence male, androgynous or mixed, densely yellowish brown tomentose by adpressed stellate hairs; bracts and bracteoles ovate, 2-3 by 1 mm. Male rachis 10-25 cm, 1-114 mm @; 6 flowers in clusters of 2-3, filaments 3—5 mm, anthers 0.3 mm long, pistillode subglobose, occasionally compressed, c. 1 mm 9g. Androgynous or mixed rachis 10-20 cm, 2 mm 9; female flowers solitary or rarely in clusters of 2, staminodes rudimentary, styles 3, conical, 2 mm, recurved. Ripe cupule sessile, saucer-shaped, 14-1 cm long, 214-3 cm g; rim thin, incurved, covering the basal part of the fruit; wall woody, 1-2 mm thick, inside densely greyish brown tomentose by ad- pressed simple hairs, outside densely fulvous stel- late hairy; scales subulate, 3-4 mm long, free, recurved or patent, densely but irregular set. Ripe fruit subglobose to cylindrical, 2!44-4 cm long, 2-234 cm g, glabrous, dark chocolate-brown, top rounded umbonate or rounded-acute, base rotun- date or cordate, scar deeply concave, 1-114 cm 9; wall woody, 2 mm thick, for the greater part free from the cupule. Distr. Indo-China (Annam), Peninsular Siam (Pattani), in Malesia: Malay Peninsula (Perak, Kelantan, Pahang). Ecol. Evergreen forests, at 700-1200 m. FI. Dec., fr. April-Nov. 97. Lithocarpus kingianus (GAMBLE) A. CAMUS, Riviera Scient. 18 (1932) 41; Chénes 3 (1954) 678, t.; SOEPADMO, Reinwardtia 8 (1970) 250. — Pasa- nia kingiana GAMBLE, Kew Bull. (1914) 117; J. As. Soc. Beng. 75, ii (1915) 417. — Synaedrys kingiana (GAMBLE) KoIpz. Bot. Mag. Tokyo 30 (1916) 196. — Fig. 28 e-f. Tree 12-20 m, 20-30 cm g. Branchlets initially densely fulvous to rufous tomentose by woolly stellate hairs, later glabrous, greyish brown, sparse- ly lenticellate; terminal buds ovoid-globose, 2—4 by 1-2 mm, scales broadly ovate. Leaves coria- ceous, 7-13 by 4~7 cm (index 1.7—214), broadest at the middle; surfaces more or less concolorous, glabrous, dark greyish green, above glossy, be- neath dull; base rounded-acute, top abruptly acute; midrib thin, prominent on both sides; nerves (4-)6—7(—8) pairs, prominent beneath, flat above, subparallel, at an angle of c. 45°, arcuating but usually not anastomosing towards the margin; reticulation fine, subscalariform to irregular, rather distinct beneath; petiole 4-8 mm, 114-2 mn, gla- brous, adaxially flat. Inflorescence male or andro- gynous, densely fulvous stellate hairy; bracts and bracteoles ovate-acute, 74-2 by 24-1 mm. Male rachis 10-20 cm, 2 mm 9g; ¢ flowers solitary or in clusters of 2-3, stamens 8-12, filaments 214-4 mm, anthers 0.3-0.4 mm long, pistillode subglobose, 1-114 mm @. Androgynous rachis 5-15 cm, 11-2 mm 9; female flowers solitary, staminodes well- developed and exceeding the perianth, styles 3, terete, 214-314 mm, recurved. Ripe cupule sessile, cup-shaped, 2 cm long, 214 cm g; rim thin, cover- ing 14-% part of the fruit; wall woody, thin, inside and outside densely fulvous-tomentose; scales subulate, 4~5 mm long, free, recurved, spiral- ly set. Ripe fruit ovoid, 244 cm long, 2 cm @, densely fulvous-tomentose by adpressed simple hairs, top rounded-acute, base truncate, scar flat, 1144 cm g; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Malay Peninsula (Pahang). Ecol. Forests, at 1350-2100 m. Fl. March— April, fr. May. 98. Lithocarpus luzoniensis (MERR.) REHD. J. Arn. Arb. 10 (1929) 133; A. Camus, Chénes 3 (1954) 1111, t.; SoppADMO, Reinwardtia 8 (1970) 256. — Quercus luzoniensis MERR. Philip. J. Sc. 3 (1908) Bot. 323; En. Philip. 2 (1923) 28. — Synaedrys ‘Juzonensis’ (MERR.) Ko1pz. Bot. Mag. Tokyo 30 (1916) 197. Tree 5-12 m, 20-30 cm g. Branchlets initially angular, densely stellate hairy, later terete, gla- brous, greyish black, densely lenticellate; terminal buds ovoid-ellipsoid, 3-5 by 2-3 mm, scales nar- rowly ovate to linear. Stipules linear-acute, 6-8 by 1 mm. Leaves thin-coriaceous, (3!4—)5—-6(-8) by (114-)214-3(-3'4) cm (index 2-2.7), broadest at or below the middle; surfaces concolorous, greenish, above glabrous, dull to glossy, beneath with a thin cover of adpressed, minute, stellate hairs; base acute to rarely rounded-acute, margin recurved, top bluntly acute to 1-2 cm acuminate; midrib prominent beneath, slightly so above; nerves (6-)7-8(-10) pairs, thin, flat beneath, im- pressed above, parallel, at an angle of 40-50°, arcuating but not anastomosing towards the mar- gin; reticulation fine, subscaliform to irregular, obscure; petiole 4-6 mm, 1 mm 9g, adaxially flat. Inflorescence male or androgynous, simple and axillary or much-branched and subterminal, dense- ly fulvous stellate hairy; bracts and bracteoles linear, 2-3 by 14-1 mm. Male rachis 3-5 cm, 1-14 mm 9g; 3 flowers solitary or in clusters of 3, fila- ments 2-214 mm, anthers 0.25-0.3 mm long, pistil- lode subglobose, 1 mm 9. Androgynous rachis 3-5 cm, 1-114 mm g; female flowers solitary, stami- nodes rather well-developed but not exceeding the perianth, styles 3, conical, 1 mm. Cupule (not fully developed) sessile, cup-shaped, 44-1 cm long, 114-114 cm g; rim thin, covering c. ¥, part of the fruit; wall woody, thin, inside densely silky tomentose by adpressed simple hairs, outside densely fulvous stellate hairy; scales distinct, nar- rowly ovate, 1 mm long, imbricate but concentri- cally set. Ripe fruit ovoid, 144 cm long, 1 cm 9, glabrous, chocolate-brown, top acute, base rotun- date, scar flat to concave, 7 mm 9; wall woody 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Philippines (Luzon, rather common on Mts Pulog, Pauai, Data, and Nanga- oto). Laat Forests, at 1500-2700 m. Fi. Jan.—June, fr. July—March. 382 FLORA MALESIANA [ser. I, vol. 7? 99. Lithocarpus echinulatus So—EPADMO, Reinward- tia 8 (1970) 235. Tree 25-40 m, 50-80 cm 9g; buttresses 1—2 m tall and out, concave; bark smooth, pale greyish brown. Branchlets initially densely tomentose by adpressed stellate hairs, later subglabrous, dull grey-brown, sparsely lenticellate; terminal buds ovoid, 3-4 by 2-3 mm, scales narrowly ovate. Leaves thin-coriaceous, (9Q—)13—18(—21) by (3-)4— 6(-7) cm (index 3-314), broadest at the middle; surfaces concolorous, dull greyish green, above glabrous, beneath densely tomentose by adpressed, minute stellate hairs; base attenuate-acute, top sharply 1-114 cm acuminate; midrib prominent beneath, flat above; nerves 7—9 pairs, prominent beneath, impressed above, subparallel, at an angle of 30-45°, arcuating but not anastomosing to- wards the margin; reticulation fine, dense, sub- scalariform, obscure; petiole 34-114 cm, 1-2 mm @, adaxially flat. Ripe cupule sessile, saucer-shaped, 4-4 cm long, 2-214 cm g; rim thin, undulate, covering the basal part of the fruit; wall woody, thin, insides densely silky tomentose by adpressed simple hairs, outside densely fulyous tomentose by adpressed stellate hairs; scales subulate, 1-2 mm long, free, patent, densely but irregular set. Ripe fruit depressed ovoid-globose, 1-114 cm long, 114-214 cm @, densely greyish tomentose by adpressed simple hairs, glabrescent, top abruptly rounded-apiculate, base rounded, scar flat, 114 cm 9; wall woody, 1 mm thick, for the greater part free from the cupule. Distr. Malesia: Borneo (scattered in SE. Kali- mantan and Sabah). Ecol. Forests, below 300 m, on sandy loamy soil. Fr. April—-Aug. Note. Inflorescences as yet unknown. 100. Lithocarpus jordanae (LAGUNA) REHD. J. Arn. Arb. 1 (1919) 127; A. Camus, Chénes 3 (1954) 981, t.; SOEPADMO, Reinwardtia 8 (1970) 250. — Quer- cus jordanae LAGUNA, Apuntes Roble Filip. (1875) 7, t.; VIDAL, Rev. Pl. Vasc. Filip. (1886) 265; Me_rkR. Philip. J. Sc. 3 (1908) Bot. 322, excl. syn.; En. Philip. 2 (1923) 27, excl. syn. — Quercus sundaica (non BL.) MeErRR. Philip. J. Sc. 1 (1906) Suppl. 41. — Synaedrys jordanae (LAGUNA) Koipz. Bot. Mag. Tokyo 30 (1916) 196. Tree 5-12 m, 20-30 cm g; bark greyish brown, smooth, lenticellate. Branchlets initially densely set with yellowish brown to fulvous velvety, sim- ple and adpressed stellate hairs, later glabrous, dark greyish brown, finely fissured and densely lenticellate; terminal buds ovoid, 3—5 by 2-3 mm, scales broadly ovate. Stipules ovate-acute, 4-5 by 2-3 mm, longitudinally ribbed and rather long persistent. Leaves thick-coriaceous, rigid, (4-)6- 8(-10) by (3-)4-S(-6) cm (index (1.3-)1.7(-2)), broadest at or below the middle; surfaces disco- lorous, above dark chocolate-brown, dull to glossy, sparsely stellate pubescent especially on the mid- rib and nerves, beneath densely set with yellowish brown, adpressed stellate and erect simple or stellate hairs; base acute to rounded-acute, some- times asymmetrical, margin strongly recurved, sometimes undulate, top bluntly acute to abruptly 4-1 cm acuminate; midrib and nerves strongly prominent beneath, flat to impressed above; nerves 8-10 pairs, dense, parallel, at an angle of 40-60°, arcuating but not anastomosing towards the mar- gin; reticulation fine, dense, scalariform, obscure to distinct beneath; petiole S-7 mm, 2-3 mm g, densely stellate pubescent, glabrescent, adaxially flat. Inflorescence male or androgynous, densely yellowish to fulvous stellate hairy; bracts and bracteoles narrowly ovate-acute, 1-4 by 14-1 mm. Male rachis 5 cm, 1-114 mm 9g; 3 flowers in clusters of 3, filaments 214-3 mm, anthers 0.3-0.35 mm long, pistillode globose, 1 mm g. Androgynous rachis 3-4 cm, 1-114 mm 9; female flowers solitary, staminodes rudimentary, styles 3, terete, 1-114 mm, recurved. Ripe cupule subsessile, cup-shaped, 14-1 cm long, 2-214 cm g; rim thin, covering c. Y part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely fulvous adpressed stellate hairy; scales linear, 5 mm long, free, irregularly set or in obscure concentric rows. Ripe fruit ovoid- conical, 2-214 cm long, 114-2 cm @g, glabrous, chocolate-brown, top rounded-acute, base rotun- date, scar flat, 1 cm @; wall woody, 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, rather common). Ecol. Forests, at 1200-2400 m. Fi. Jan.—July, fr. Aug.—April. Note. VIDAL 1814, placed by StTaprF (1894) under his Quercus havilandii, belongs here. 101. Lithocarpus aspericupula (MARKGR.) REHD. J. Arn. Arb. 10 (1929) 123; A. Camus, Chénes 3 (1954) 903, t.; So—EPADMO, Reinwardtia 8 (1970) 217. — Pasania aspericupula MARKGR. Bot. Jahrb. 59 (1924) 78, f. 1 E. Tree, c. 10 m tall. Branchlets initially densely fulvous tomentose by adpressed stellate hairs, later subglabrous, dark brown, densely lenticellate; terminal buds ovoid, 3 by 3 mm, scales ovate. Stipules ovate, 3 by 2 mm. Leaves coriaceous, elliptic-oblong, 8-14 by 4-6 cm; surfaces disco- lorous, above glabrous except the midrib, glossy, chocolate-brown, beneath densely yellowish to- mentose by adpressed stellate hairs interspersed by erect, brownish tuft-hairs; base rounded-acute, top rounded and abruptly, sharply 14-1 cm acu- minate; midrib prominent on both sides; nerves 8-10 pairs, prominent beneath, impressed above, subparallel, at an angle of c. 50°, arcuating but not anastomosing towards the margin; reticulation lax, arched, subscalariform, distinct beneath; petiole 7-10 mm, 2 mm @g, densely stellate pubes- cent, glabrescent, adaxially flat. Cupule (? ripe) 4 cm stalked, saucer-shaped, 14 cm long, 114-1% cm @, rim thin, covering the basal part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside densely yellow- ish tomentose by adpressed stellate hairs; scales ovate to subulate with pointed tips, recurved, 2-4 mm long, irregular set. Fruit ovoid, 144-174 cm 1972] FAGACEAE (Soepadmo) 383 long, 114 cm @, glabrous, chocolate-brown, top rounded-acute, base rounded, scar concave, 7 mm @; wall woody, 1 mm thick, for the greater part free from the cupule. Distr. Malesia: New Guinea (Vogelkop Penin- sula and Sepik area). Ecol. Forests, up to 250 m, on sandy clayey soil overlying limestone. F/. Sept., fr. March. Notes. The male inflorescence in LEDERMANN’S specimens is too badly preserved for description. The fruiting specimen BW 15331 agrees very well with MARKGRAr’s drawing of the cupule and fruit of LEDERMANN 11523 (+) but for the smaller size, which was cited by MARKGRAF (nuts up to 3 by 214 cm); the leaves perfectly agree with those of LEDERMANN 8729 (WRSL). 102. Lithocarpus buddii (Merr.) A. CAMus, Bull. Soc. Bot. Fr. 90 (1943) 85; Chénes 3 (1954) 846, t. 428; SoppADMO, Reinwardtia 8 (1970) 222. — Quercus robinsonii MERR. Philip. J. Sc. 10 (1915) Bot. 287; En. Philip. 2 (1923) 30, non RIDL. 1914. — Synaedrys robinsonii (MERR.) KOIDZ. Bot. Mag. Tokyo 30 (1916) 192. — L. robinsonii MerR. J. Arn. Arb. 1 (1919) 130. — Quercus buddii MERR. Philip. J. Sc. 35 (1928) 6. Tree 10-25 m, 25-40 cm g. Branchlets initially densely rufous pubescent with adpressed and erect, stellate and simple hairs, later glabrous, dark greyish brown, finely fissured, sparsely lenti- cellate; terminal buds ovoid, 3—5 by 2-4 mm, scales linear-acute. Stipules linear-acute, 5-10 by 1-114 mm. Leaves thin-coriaceous, (6—)10—14(—17) by (214-)3'4-5C7) cm (index (2—)2.3-2.8(-3)), broadest at or below the middle; surfaces disco- lorous, above dull to glossy, greyish brown, initi- ally densely rufous pubescent by stellate hairs, soon glabrescent, beneath densely rufous tomen- tose by adpressed and erect stellate hairs, later subglabrous; base acute, top bluntly 1-114 cm acuminate; midrib strongly prominent beneath, slightly so to flat above; nerves (6—)7—8(—9) pairs, thin, prominent beneath, flat above, subparallel, at an angle of 45—50°, arcuating but not anasto- mosing towards the margin; reticulation fine, dense, subscalariform, obscure; petiole !4—1 cm, 1- 144 mm 9g, adaxially flat. Inflorescence male or androgynous, simple and axillary or much-bran- ched and subterminal, densely rufous stellate hairy; bracts and bracteoles linear-acute, 2-5 by Y%-1 mm. Male rachis 8-15 cm, 1-114 mm @g; 6 flowers solitary or in clusters of 2-3, stamens 10- 12, filaments 2-3 mm, anthers 0.25-0.3 mm long, pistillode globose, 1 mm 9. Androgynous rachis 6-17 cm, 1144 mm @; female flowers solitary or very rarely in clusters of 2-3, staminodes rudi- mentary, styles 3-4, terete, 2-3 mm, recurved. Ripe cupule subsessile, cup-shaped, 14-1 cm long, 114-2 cm 9g; rim thin, covering the basal part of the fruit; wall woody, thin, inside densely fulvous tomentose by adpressed simple hairs, outside den- sely rufous tomentose by adpressed stellate hairs; scales subulate, recurved, 2-4 mm long, free, in 4-6 concentric rows. Ripe fruit depressed ovoid, 114-144 cm long, 114-2 cm g, glabrous, chocolate- brown, top rounded-acute, base rotundate, scar flat to concave, 1 cm 9; wall woody, 1 mm thick, for the greater part free from the cupule; cotyledons flat-convex. Distr. Malesia: Philippines (Luzon, common on Mt Maquiling; Samar, rare; Leyte, rare). Ecol. Forests, at 200-900 m. Fi/. Oct.—June, fr. June—Dec. 103. Lithocarpus rufovillosus (MARKGR.) REHD. J. Arn. Arb. 10 (1929) 133; A. Camus, Chénes 3 (1954) 807, t.; SOEPADMO, Reinwardtia 8 (1970) 276. — Pasania rufovillosa MARKGR. Bot. Jahrb. 59 (1924) 74, f. 3, 4: B-C. Tree 15-40 m, 20-60 cm g; bark pale to dark grey-brown, fissured to scaly. Branchlets initially densely fulvous to rufous-tomentose by adpressed stellate hairs and woolly tuft-hairs, later subgla- brous, dark grey-brown, sparsely lenticellate; terminal buds ovoid, 3—5 by 2-3 mm, scales ovate. Stipules ovate to lanceolate, 3-5 by 2 mm. Leaves coriaceous, (6—)10—-15(-18) by (3-)4-6(-8) cm (index (2—)214(-314)), broadest at or below the middle; surfaces more or less discolorous, above greyish brown to dark chocolate-brown, dull to rarely glossy, initially sparsely pubescent by erect, woolly tuft-hairs especially on the midrib and nerves, soon glabrescent, beneath densely set with a thick cover of fulvous to rufous, adpressed stellate hairs and woolly tuft-hairs, the latter subglabrescent; base rounded-acute, very often asymmetrical, margin occasionally strongly re- curved, top rounded and abruptly acute to more commonly 14-114 cm acuminate, tips sharp, often oblique; midrib strongly prominent on both sides; nerves 9-11 pairs, prominent beneath, fiat to impressed above, subparallel, at an angle of (45—) 60-70°, arcuating but not anastomosing towards the margin; reticulation lax, subscalariform, rather distinct beneath; petiole 14-1 cm, 114-3 mm @, terete, densely rufous pubescent. Jnflorescence male or androgynous, in densely subterminal pani- culate clusters, densely rufous stellate hairy; bracts and bracteoles ovate-acute, 1-244 by 24-1 mm. Male rachis 7-15 cm, 2 mm 9; 3 flowers in clusters of 3, stamens 10-12, filaments 214-3 mm, anthers 0.3-0.4 mm long, pistillode subglobose, 1 mm 9. Androgynous rachis 5-10 cm, 2 mm 9; female flowers solitary, very rarely in clusters of 2-3, staminodes rather well-developed but not exceed- ing the perianth, styles 3-4, terete, 114-2 mm, recurved. Ripe cupule sessile, cup-shaped, 1-214 cm long, 2-3(—314) cm @; rim 1 mm thick, covering the basal part of the fruit; wall woody, 1-2 mm thick, inside densely fulvous tomentose by ad- pressed simple hairs, outside densely fulvous to rufous tomentose by adpressed stellate hairs; scales ovate with pointed, subulate tips, 1-3 mm long, alternate but concentrically set. Ripe fruit ovoid-conical, 2-3 cm long, 2-2! cm 9, glabrous, chocolate-brown, top rounded, base rotundate, scar concave, 114 cm 9; wall bony, 2-3 mm thick, for the greater part free from the cupule; cotyle- dons flat-convex. 384 Distr. Malesia: New Guinea (common in all parts), also Japen and Misool Is. Ecol. Forests, from sea-level up to 2300 m, more commonly above 700 m, on sandy or clayey soils. F/. Sept.—June, fr. July—April. Note. Specimens from above 1000 m usually have a thicker and darker indumentum and a larger cupule and fruit than those from the low- lands. 104. Lithocarpus hystrix (KORTH.) REHD. J. Arn. Arb. 1 (1919) 127; A. Camus, Chénes 3 (1954) 856, t. 432: 5-22; So—EPADMO, Reinwardtia 8 (1970) 246. — Quercus hystrix KortTH. Kruidk. (1844) 201, t. 43; Mia. Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 108; Wenzic, Jahrb. Bot. Gart. Berl. 4 (1886) 223; Hoox. f. Fl. Br. Ind. 5 (1888) 611; KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 54, t. 50; CORNER, Ways. Trees (1940) 303, f. 96. — Quercus korthalsii BL. var. hystrix (KORTH.) BL: Mus. Bot. 1 (1850) 293; A. DC. Prod. 16, 2 (1864) 90. — Quercus mappacea var. hystrix (KORTH.) Mia. FI. Ind. Bat. 1, 1 (1856) 851. — Quercus cyrtopoda Mia. ibid. 1, 1 (1858) 869; A. DC. Prod. 16, 2 (1864) 97; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 86. — Castanea furfurella Mig. Sumatra (1861) 352. — Cyclobalanus hystrix (KORTH.) OrrstT. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. — Cyclobalanus cyrtopoda (Mi1qQ.) OrrstT. l.c. 80. — Quercus brevipetiolata ScHEFF. Nat. Tijd. N. I. 31 (1870) 359. — Pasania hystrix (KORTH.) GAMBLE, J. As. Soc. Beng. 75, ii (1915) 428. — Synaedrys hystrix (KORTH.) Korpbz. Bot. Mag. Tokyo 30 (1916) 195. — L. cyrtopoda (Miaq.) A. Camus, Chénes, Atlas 3 (1949) 74, t.; Chénes 3 (1954) 753. Tree 10-30 m, 20-90 cm g; bark pale-brown. Branchlets densely set with fulvous to rufous erect tuft-hairs with 2-many arms, later subgla- brous, dark greyish brown, finely fissured, sparsely to densely lenticellate; terminal buds ovoid, 3-5 by 2-3 mm, scales ovate. Stipules broadly ovate to lanceolate, 3-10 by 2-5 mm, rather long persistent. Leaves thin-coriaceous, (12—)14—18(—20) by (5—)6— 7(-8) cm (index 2-3), broadest at the middle; surfaces discolorous, above dark brown, densely pubescent especially on the midrib and nerves, subglabrescent, dull to glossy, beneath densely set with fulvous to rufous, adpressed stellate hairs and erect tuft-hairs; base rounded, margin occa- sionally strongly recurved, top rounded and ab- ruptly acute to 14-1 cm acuminate; midrib strong- ly prominent on both sides; nerves 12-16 pairs, strongly prominent beneath, flat to impressed above, parallel, at an angle of 50-60°, arcuating and anastomosing near the margin; reticulation dense, scalariform, distinct beneath; petiole 14-1 cm, 2 mm 9, terete or adaxially flat, densely ful- vous to rufous pubescent. Inflorescence male, androgynous or mixed, in densely subterminal paniculate clusters, densely fulvous to rufous pubescent; bracts and bracteoles narrowly ovate, 1-114 by \%4-% mm. Male rachis 5-15 cm, 1-2 mm 9; 3 flowers solitary or in clusters of 2-3, filaments 2-3 mm, anthers 0.25-0.3 mm long, FLORA MALESIANA [ser. I, voln?? pistillode globose, 1 mm 9. Androgynous or mixed rachis 10-20 cm, 2 mm @; female flowers solitary, rarely in clusters of 2-3, staminodes rudimentary, styles 3-4, conical, 1-144 mm, recurved. Ripe cupule sessile to 14 cm stalked, saucer- to cup-sha- ped, 14-1 cm long, 2-3 cm g; rim thin, covering the basal part of the fruit; wall woody, 2-3 mm thick, inside densely fulvous tomentose by ad- pressed, simple hairs, outside densely fulvous to rufous tomentose by adpressed stellate hairs; scales subulate, 3-5 mm long, patent or recurved, more or less concentrically set. Young fruit with a long acuminate top. Ripe fruit depressed-ovoid, 114-2 cm long, 2-3 cm g, glabrous, dark purplish brown, top rounded-acute to rounded-acuminate, base rotundate, scar flat to concave, 114-2 cm 9; wall woody, 1-2 mm thick, for the greater part free from the cupule. Distr. Malesia: Sumatra (common; also in Simalur, Riouw, and Karimun Is.), Malay Penin- sula (Perak, Selangor, Negri Sembilan, Pahang, Malacca, Johore; also in Penang and Singapore), Borneo (scattered in SE. Kalimantan). Ecol. Forests, from sea-level up to 1800 m, mostly below 800 m. Fi. Sept.—July, fr. Aug.—June. Doubtful Lithocarpus argyrocarpa A. CAMUS, Bull. Soc. Bot. Fr. 94 (1947) 4; Chénes 3 (1954) 806, t. 417: 9-15. KEP 21712 (type, K!) with young cupules and fruits, was collected in the border area between Kedah (Malay Peninsula) and Siam. Except for the smaller size, the cupules, fruits and leaves, it very closely resembles specimens of L. sootepensis (CRAB) A. CAMuUS from Doi Angka in Siam. More material of both species is, however, needed to draw a definite conclusion. Lithocarpus debaryana (O. WARB.) MARKGR. Bot. Jahrb. 59 (1924) 69; A. Camus, Chénes 3 (1954) 608. — Quercus debaryana O. WARB. Bot. Jahrb. 13 (1891) 286; VON SEEMEN in K. Sch. & Laut. Fl. Schutzgeb. (1901) 263. — Synaedrys debaryana (O. WARB.) KoIpz. Bot. Mag. Tokyo 30 (1916) 189. When WARBURG described this species from E. New Guinea, he did not indicate any particular specimen on which he based his new species. He described the leaves as 18 by 9 cm, and the cupule 4 cm long, 314 cm g, narrowed at the base and covering the fruit almost completely. MARKGRAF mentioned WARBURG 20584 (consisting of one leaf and one fruit) from Sattelberg in the NE. part of New Guinea as the type, and added HELLWIG 515, also from Sattelberg. Both speci- mens were lost during the World War IJ. Among the later collections from the same area, CLEMENS 11360 (Eot-Wonimbu, Morobe Distr.) is the only specimen of which the cupules and fruits agree with WARBURG’S description. The cupule and nut show some resemblance in size to those of 38. L. lauterbachii (VON SEEMEN) MARKGR. but 1972] are clearly different. It seems a good species but more material is needed for its proper definition of the inflorescence and foliage. The leaves (collect- ed from a basal shoot) are, however, much smaller than those described by WARBURG, the largest being 12 by 5 cm. Lithocarpus dolichocarpa (VON SEEMEN) REHD. J. Arn. Arb. 1 (1919) 125; A. Camus, Chénes 3 (1954) 801. — Quercus dolichocarpa VON SEEMEN, Bot. Jahrb. 27, Beibl. 64 (1900) 14; K. & V. Bijdr. 10 (1904) 44; Koorp. Atlas 1 (1913) t. 50; BACKER & BAKH. f. Fl. Java 2 (1965) 8. — Synaedrys dolichocarpa (VON SEEMEN) Koripz. Bot. Mag. Tokyo 30 (1916) 191. Ecxept for the smaller size of the cupule, fruit and leaf, the species which is based on Javanese specimens is suggestive of 47. L. daphnoideus (BL.) A. Camus. The specimens which I have seen are, however, far too incomplete for a final decision. Lithocarpus menadoensis (KOORD.) SOEPADMO, Reinwardtia 8 (1970) 290. — Quercus menadoensis Koorp. in Koord.—Schum. Syst. Verz. 3 (1914) 28, nomen. KoorvDerRS 16632 from Minahassa (NE. Cele- bes), the type, now preserved in the Rijksherba- rium, Leyden, has glabrous leaves, 10-12 by 3-414 cm, with 6—7 pairs of nerves, irregular reticulation, and a cupule 1 cm long and 1.7 cm 9, with 5-6 obscure denticulate lamellae. The fruit is ovoid, c. 144 cm through, glabrous and chocolate-brown. There are, at Leyden, several other collections mainly from the vicinity of Malili, Central Celebes, which resemble KoorDERS 16632, but unfortunate- ly most of them are sterile. The species is sugges- tive of 43. L. caudatifolius (MERR.) REHD. from Borneo and the Philippines, but in the latter the leaves and branchlets are densely velvety tomen- tose. Lithocarpus moluccus (RUMPH. ex L.) SOEPADMO, Reinwardtia 8 (1970) 290. — Quercus molucca [RumpPH. Herb. Amb. 3 (1743) 85, t. 56 ex] LINNE, Sp. Pl. (1753) 1199; WILLD. Sp. PI. 4, 1 (1809) 427; BL. Mus. Bot. 1 (1850) 291; Mra. FI. Ind. Bat. 1, 1 (1856) 849; A. DC. Prod. 16, 2 FAGACEAE (Soepadmo) 385 (1864) 86; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 90; Merr. Int. Rumph. (1917) 186; A. Camus, Chénes 3 (1954) 1163. — Balanaulax molucca (RUMPH. ex L.) RAFIN. Alsog. Am. (1838) 25. — Pasania molucca (RUMPH. ex L.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 373. See notes under 50. L. glutinosus (BL.) SOEPAD- MO. Lithocarpus ‘nymanniana’ (MARKGR.) REHD. J. Arn. Arb. 10 (1929) 133; A. Camus, Chénes 3 (1954) 1164. — Pasania nymaniana MARKGR. Bot. Jahrb. 59 (1924) 77. The species was described from a specimen (NYMAN 692, New Guinea, .v.) with a very young cupule and fruit. As the type seems to be lost, and the description is not recognizable, the species is treated as doubtful. Lithocarpus oogyne (MiqQ.) A. CAMUS, Bull. Soc. Bot. Fr. 92 (1945) 255; Chénes 3 (1954) 781. — Quercus oogyne MiQ. Sumatra (1861) 351; A. DC. Prod. 16, 2 (1864) 95; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 92. — Cyclobalanus oogyne (MiqQ.) OerrsT. Vid. Medd. Naturh. For. Kjébn. 8 (1867) 81. DIEPENHORST HB 1315 (type, U!) from Priaman, NW. Sumatra, has very young cupules and fruits which are suggestive of 89. L. sundaicus REHD., though the leaves (badly preserved) are much smaller and less pubescent. MIQUEL compared his species with Quercus blumeana KorTH. and Quer- cus encleisacarpa KORTH., which are, however, two completely different species. Lithocarpus plumbea (BL.) SOEPADMO, Rein- wardtia 8 (1970) 291. — Quercus plumbea BL. Mus. Bot. 1 (1850) 293; A. DC. Prod. 16, 2 (1864) 88; A. Camus, Chénes 3 (1954) 1166. — Pasania plumbea (BL.) OrErst. Vid. Medd. Naturh. For. Kjoébn. 8 (1867) 84. The type (? KORTHALS s.n., Sumatra) now pre- served at Leyden, has very young cupules and fruits which resemble those of 89. L. sundaicus REHD. The leaves are, however, smaller and less pubescent than those of L. sundaicus. 4. QUERCUS! LINNE, Gen. Pl. ed. 5 (1754) 431; A.DC. Prod. 16, 2 (1864) 2, p.p., excl. sect. Andro- gyne, Chlamydobalanus, Cyclobalanus, Lithocarpus et Pasania; B. & H. Gen. Pl. 3 (1880) 407, p.p., excl. ditto et sect. Synaedrys; Hook. f. Fl. Br. Ind. 5 (1888) 600, p.p., excl. ditto; PRANTL in E. & P. Nat. Pfl. Fam. 3, 1 (1889) 55; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 19, p.p., excl. ditto; A. CAMUS, Chénes 1 (1938) 7; BARNETT, Trans. & Proc. Bot. Soc. Edinb. 33 (1942) 329; HUTCHINSON, Gen. FI. Pl. 2 (1967) 13; SoppADMo, Gard. Bull. Sing. 22 (1968) 355. — Cyclobalanopsis OrrsT. Vid. Medd. Nat. For. Kjébn. 8 (1867) 77. — Fig. 32-34. (1) This generic description covers only the Malesian species. 386 FLORA MALESIANA [ser. I, volt7? Trees, sometimes buttressed, rarely with small stilt-roots. Branchlets initially densely tomentose by simple or stellate hairs, or densely brownish, stiff pubescent, glabrescent; terminal buds ovoid-globose or ovoid-conical, rarely ovoid-ellipsoid, usually conferted, scales densely fulvous-tomentose, with a tendency to ortho- stichy. Stipules extrapetiolar, linear-acute, densely tomentose or woolly pubescent, caducous. Leaves spirally arranged, crowded near the top of the branchlets, or rarely pseudo-whorled; midrib and nerves flattened or impressed or slightly raised above, prominent beneath; margin entire or remotely minutely serrate in the api- cal half; glabrous to densely pubescent or tomentose (sometimes by stellate hairs) at least on the lower surface; petiole thickened at base. Inflorescence male or female. Male rachis solitary in the axil of a lower leaf or in paniculate clusters on the lateral or subterminal new shoots, flexuous, pendent, compound or simple; bracts ovate- linear, acute, densely tomentose, caducous; 3 flowers in clusters of 3-4; perianth (4—)6-lobed, the lobes connate at base, densely tomentose; stamens (4-)6(-9), filaments slender, filiform, glabrous or tomentose at base, anthers 14-1 mm long, basifixed; pistillode normally absent, sometimes replaced by a tuft of stiff simple hairs. Female rachis solitary in the axil of a higher leaf, erect, densely woolly pubes- cent, few- to many-flowered; bracts linear-acute, densely pubescent, caducous; 2 flowers always solitary; perianth (4—)6(—9)-lobed; staminodes 0 or 5-7; styles 3—4(-6), cylindrical, free and recurved or connate at base; stigmas broadly capitate, glabrous; ovary cells as many as styles. Cupule cup- or saucer-shaped, obconical or obovoid-globose, lamellate, hairy both inside and outside; lamellae c. 5—12, denticulate and free at the rim or more or less smooth and connate, thin or thick. Fruit ovoid-conical, ovoid-globose or ovoid-cylindrical; apex rounded, attenuate- acute or abruptly depressed, umbonate; perianthodium (umbo) provided with many rings, well-developed; glabrous and shining or densely tomentose. Cotyledons flat-convex; radicle vertical. Germination hypogeal. yd oe eR Fig. 30. Distribution of the genus Quercus, the fossil records indicated by dots. Distr. Quercus has most of its c. 600 spp. in the northern temperate zone, with an extension into the subtropics and tropics into W. Malesia (to 8° S) and the NE. corner of S. America (see map in Gard. Bull. Sing. 22, 1968, 419). In Malesia occurs only Quercus subg. Cyclobalanopsis, which is confined to 1972] FAGACEAE (Soepadmo) : 387 E. and SE. Asia, viz Japan (Kanto Prov., Honshu), Korea, China, Hainan, Formosa, Indo-China (with a centre of speciation), NE. India, Burma, Thailand; in Malesia it is found in Sumatra, the Malay Penin- sula, Banka, Billiton, Anambas Is., W. and Central Java, Borneo, and Palawan. The wider spread species are rather equally spread over the Malay Peninsula, Sumatra, and Borneo, but few in Java. In Borneo oc- cur 9 endemic spp., in N. Sumatra 1. Fig. 30-31. Fossil records. LA Motte (Mem. Am. Geol. Soc. 51, 1952, 285-301) listed not less than 200 fossil “species” recorded from various localities in America and Greenland of different geological ages (Upper Cretaceous to Pleistocene). Most fossil records which have been discovered in Europe refer to fossils younger than Late Tertiary. According to TRALAU (Kongl. Svensk. Vet. Akad. Handl. IV, 9, 1963, 18-21), all these records might belong to one species. From the Tertiary of Asia, FLORIN (Kongl. Svensk. Vet. Akad. Handl. 63, 1920, 1-71), OGuRA (Jap. J. Bot. 6, 1932, 173-181), and WATARI (Jap. J. Bot. 11, 1938, 385-438) reported several fossils from Japan, Korea and Manchuria, and RAo (Paleaeobot, 4, 1955, 37-89) from India. In Australia and Tasmania, ETTINGSHAUSEN (Mem. Geol. Surv. N.S.W., Palaeont. 2, 1888) and JoHN- STON (Pap. & Proc. R. Soc. Tasmania 1885, 322-325) described 21 species from various Tertiary depo- sits. According to DUIGAN (Proc. R. Soc. Victoria n.s. 63, 1951, 46-47) and Couper (Proc. R. Soc. Lon- don, Sect. B, 152, 1960, 491—500) the identification of most of these Australasian records is rather doubt- ful. Ecol. In Malesia, Quercus is confined to everwet conditions. Therefore it is very scarce in Central and East Java. The genus occurs from sea-level to 3350 m, with a preference for the region between 600 and 1500 m, in various sorts of primary forest, viz lowland mixed Dipterocarp forest, swamp forest (with fluctuating water-level), low or hill kerangas or heath forest, ridge forest. Types of soil recorded are sandy clay, sandy loam, and ultrabasic soil overlying sandstone or granite. Notes. The distribution of Malesian species of Quercus outside Malesia has but partially been investi- gated. O. SCHWARZ (Notizbl. Berl. Dahl. 13, 1936, 7-8) distinguished besides Cyclobalanopsis also Erythro- balanus and Macrobalanus as distinct genera; they are both New World taxa which, in my opinion, should not be segregated from Quercus. KEY TO THE SPECIES?! (based on flowering and fruiting specimens) 1. Rim of the cupule thinner than 144 mm, lamellae with more or less free margin. 9 Inflorescences usually many-flowered. 2. Cupule deeply cup-shaped, covering 14—%,, part of the fruit, (1—-)2-3(-3 %) cm high, 2-4 cm o. 3. Cupule 2-3 4% cm high, 244-4 cm g. Fruit 3—S by 2-3 cm. Leaves pseudo-whorled, base cordate to auriculate; petiole 4-1 cm .. Sula) soe eee ee ol. OU pseudoverticillata 3. Cupule (1-)14%4-2(-2%) cm high, d- )1%-20-2% 6) cm @. Fruit 2-3(—3 14) by 1 42-2(-3) cm. Leaves not pseudo-whorled, base not cordate nor auriculate; petiole 1 4% —5 cm. 4. Cupule tapering towards the base. Leaves densely stellate-tomentellous, glaucous or silvery be- neath, entire. 5. Fruit elongate conical or ovoid-globose. Styles free and recurved. Staminodes 0. Leaves silvery beneath; petiole not sulcate on the adaxial side. . . . . . 2. Q. argentata 5. Fruit obovoid-globose. Styles connate, not recurved. Staminodes ee 1. Leaves glaucous beneath; petiole deeply sulcate on the adaxial side .. . we ies. Of nived 4. Cupule rounded or truncate at base. Leaves glabrous ¢ or densely brownish stiff-pubescent or sparse- ly stellate-tomentose, neither glaucous nor silvery beneath; margin remotely minutely serrate in the apical half. 6. Cupule 2-2 by 2-2% cm; lamellae 6-8. Fruit ovoid-conical, 2-3 by c. 2 cm; base convex. 4. Q. gaharuensis 6. Cupule 3-3 % by 3-3% cm; lamellae 9-11. Fruit Saeko tae or ovoid-cylindrical, 2 4%4-3 4 by 2-3 cm; base flat or convex .... a) os Os idbiirps 2. Cupule shallowly cup-shaped or saucer- shaped, ‘covering 1 ay part of the fruit, %4-1.2 cm high, (0.7—)1-1.8(-24%) cm @. 7. Leaves elliptic-lanceolate or ovate-lanceolate, (3—)10-15(—24) by (1-)3—5(-9) cm; petiole 1-3 cm. 8. Leaves elliptic-lanceolate, chartaceous, base attenuate-acute. 9. Cupule covering 4%—- vs eae of the fruit; base of the fruit concave. Petiole terete or flattened on the adaxial side ... 5 RT ee | ae ee eee en ee Same Orit gies (1) The term ‘tomentose’ is used for any indument which is continuous interwoven and does not show the parenchyma beneath, irrespective of its thickness and the size of the hairs; the latter may be of mi- croscopical dimension and the tomentum very thin. 388 FLORA MALESIANA [ser. 1, vale@e 9. Cupule covering te part of the fruit; base of the fruit convex. Petiole sulcate on the adaxial sidehy’ bay A ae eS at 4. Q. gaharuensis 12. Leaves sparsely stellate- tomentose; petiole and midrib deeply sulcate on the adaxial side. 5. Q. oidocarpa 10. Leaves elliptic-lanceolate, (1 4—)5-13(—24) by 2%-5 cm. 13. Leaves glaucous beneath, reticulation fine, obscure... ........-. 6. Q. sumatrana 13. Leaves not glaucous beneath; reticulation coarse, distinct beneath. 14. Leaves densely glaucous, adpressed-pubescent beneath; nerves 10-20 pairs, dense; reticulation dense, distinct beneath ep efat i ie, Weel sist ata em etre Dra wee Leereh oils Se P eee el Ve) tan ee 16. Q. lineata 14. Leaves densely rufous, woolly-pubescent beneath; nerves 5-12 pairs, lax; reticulation obscure, lax. SAME ee ales © “es” ce) \ 60 e* aereae eal) le 19. Q. elmeri on day merrell se, 5) a)! wy “epee, et 4) ert. a he Rie 5. Leaves linear-lanceolate or ovate-elliptic, 5-10 by 2-4 cm. 15. Leaves linear-lanceolate, with a dense cover of simple pubescence beneath . . 18. Q. treubiana 15. Leaves ovate-elliptic, densely or sparsely stellate-tomentose beneath, or glabrous. 16. Leaves with attenuate-acute or attenuate-rounded base, glabrous or sparsely stellate-tomentose beneath. 17. Leaves glabrous, glaucous beneath; petiole 0.8-1.2 cm : 17. Leaves sparsely stellate-tomentose beneath, not glaucous; petiole 1-2 % cm. 12. Q. kerangasensis 7. Q. subsericea 16. Leaves with rounded or cordate base, with densely stellate-tomentose by simple hairs beneath. 18. Leaves remotely serrulate in the apical half, beneath densely stellate-tomentose or glabrous; reticulation more or less areolate 8. Q. lowii ot Meme ee 6). cms! Se Neu OM Ree Beek On ee seh (© aoe 18. Leaves entire, beneath densely yellowish pubescent; reticulation scalariform. 1. Quercus pseudoverticillata SOEPADMO, Gard. Bull. Sing. 21 (1966) 380, f. 1; ibid. 22 (1968) 384, map 3. Tree, c. 30 m, c. 90 cm g; buttresses up to 2 m tall; bark scaly, peeling off profusely into small, rectangular pieces. Branchlets glabrous, lenticel- late; terminal buds ovoid-globose, 2-3 by 2 mm. Leaves pseudo-whorled, thick-coriaceous, elliptic- lanceolate or oblanceolate-oblong, 7-17 by 3-614 cm, entire, base cordate to auriculate, apex round- ed or bluntly acute; above glabrous, beneath sparsely minute-pubescent; midrib and nerves strongly prominent beneath, flattened to slightly impressed above, especially in the apical half; nerves 8-15 pairs, at an angle of 30-45° with the midrib, parallel arcuating; reticulation obscure, scalariform, dense; petiole 14-1 cm by 0.2 cm, tomentose, glabrescent, adaxially flat. Cupule cup- shaped, obconical-globose, pointed at base, cover- ing 144-¥, part of the fruit, 2-314 cm high, 214-4 cm @; outside brownish sericeous, glabrescent; inside densely brownish stiff-pubescent; lamellae 10-12, thin, rims free, dentate, especially the lower ones. Fruit cylindrical-globose, 3-5 by 2-3 cm, sericeous, apex rounded or depressed, base convex or flat; umbo conical. Distr. Malesia: North Borneo (Mt Kinabalu). Ecol. Rain-forest, c. 1650 m. Fr. Oct.—Febr. Note. Stipules and inflorescences are as yet unknown. 2. Quercus argentata KorTu. Kruidk. (1844) 215, t. 47: 1-17; A. DC. Prod. 16, 2 (1864) 91; SCHEFF. Nat. Tijd. N. I. 32 (1871) 417; WeENziIG, Jahrb. Bot. Gart. Berl. 4 (1886) 230; Kina, Ann. R. Bot. Gard. Calc. 2 (1889) 30, t. 24A; A. Camus, Chénes 13. Q. kinabaluensis Fig. 31. Species density of Quercus in Malesia, above the hyphen the number of non-endemic species, below the hyphen the number of endemic species in the island or island group. 1 (1938) 311, t. 23: 1-9; So—PADMo, Gard. Bull. Sing. 22 (1968) 384, f. 6, map 4. — Q. argentata var. concolor BL. Mus. Bot. 1 (1850) 299. — Q. pinanga BL. I.c. 303; A. DC. Prod. 16, 2 (1864) 107; Mia. FI. Ind. Bat. 1, 1 (1856) 864; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 92; K. & V. Bijdr. 10 (1904) 65. — Cyclobalanopsis argentata (KORTH.) Orrst. Vid. Medd. Nat. For. Kjébn. 8 (1867) 79, t. 1-2 f. 5-8. — Q. wilhelminae VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 9, excl. fr. — Synae- drys wilhelmianae (VON SEEMEN) Kot1pZ. Bot. Mag. Tokyo 30 (1916) 193. — Lithocarpus argentata (KortH.) MerR. Contr. Arn. Arb. 8 (1934) 42, excl. BANGHAM 1117 which is L. ewyckii. — Fig. 32; FLORA MALESIANA fser. Rival t7* 390 Sabah, c. 1500 m alt. Bembangan R., Mt Kinabalu, ’ Tree base (RSNB 4427; CorRNER, 1964). Quercus argentata KORTH PZ. Fig 1972] FAGACEAE (Soepadmo) 391 Tree, 7-37 m, 20-100 cm 9; buttresses up to 1.2 m tali; bark smooth, lenticellate, pale grey, some- times with horizontal cracks; inner bark c. 2 cm thick, pale brown, brittle. Branchlets glabrous, densely lenticellate, greyish brown; terminal buds ovoid-globose, 2-3 by 2-214 mm, tomentose by stellate or simple hairs, glabrescent. Stipules linear- acute, densely stiff-pubescent, 3-5 by 1-114 mm. Leaves coriaceous, elliptic- or lanceolate-oblong, 8-22 by 3-7 cm; beneath with dense, silvery, stel- late tomentum, above glossy, glabrous; base atten- uate-rounded or attenuate-acute, slightly asym- metrical, top 14—1 cm acuminate, entire and slight- ly undulate: midrib and nerves strongly prominent beneath, impressed above, glabrous; nerves 10-17 pairs at an angle of 60-75”, parallel, arcuating near the margin; reticulation fine, obscure; petiole 114-3 cm by 1-1!4 mm, adaxially flat, glabrous. Male rachis 5-10 cm, in a paniculate cluster of 3 or 4 0n a lateral new shoot; bracts ovate-acute, densely tomentose by simple hairs, 1-114 by 4 cm; ¢ flowers: perianth lobes connate at base; stamens normally 6, filaments hairy at base. Female rachis many-flowered, slender, 2-3 cm, densely tomentose by simple hairs; bracts linear- acute, 14-1 mm; @ perianth densely tomentose outside; staminodes absent; styles hairy at base, free and slightly recurved. Young cupule turbinate or ovoid-conical, flattened or rounded at the top, attenuate at base; lamellae 8-10, thin, dentate, with dense brownish tomentum, covering the fruit except for the umbo. Mature cupule cup-shaped, obconical or obovoid, 114-2 cm high, 1-114 cm @, densely pubescent; lamellae 8-10, free and denticulate at the rim. Fruit elongated conical or ovoid-globose, attenuate towards the acute top and rounded base, 3-314 by 114 cm. Distr. Malesia: Sumatra (scattered), also Ban- ka, Malay Peninsula (Malacca, Selangor, Johore, Trengganu, and Singapore), Anambas Is., Borneo (all parts except Brunei; also P. Laut), West Java. Ecol. Forests up to 2700 m. Fi. July—Sept., fr. Oct.—May. 3. Quercus nivea KING, Ann. R. Bot. Gard. Calc. 2 (1889) 31,t. 24B; A. Camus, Chénes 1 (1938) 313, t. 23: 10-16; SoEPADMO, Gard. Bull. Sing. 22 (1968) 387, f. 7, map 3. Tree, c. 25 m, c. 40 cm g; buttresses c. 2 m tall, 10-15 cm thick; bark pale grey, rough; inner bark chocolate-brown, fibrous. Branchlets with a dense brownish, stellate tomentum, glabrescent; lenticels sparse, splitting longitudinally into shallow fur- rows; terminal bud ovoid-ellipsoid, 3-4 by 2-3 mm, scales ovate, glabrous. Stipules linear-acute, stel- late-tomentose, 2-3 by 4-1 mm. Leaves thick- coriaceous, elliptic-oblong or lanceolate-oblong, 6-15 by 214-514 cm; above glossy, glabrous, be- neath glaucous, with dense stellate tomentum; entire, slightly undulate in upper part; base round- ed or abruptly acute, sometimes asymmetrical, top 14-1 cm acuminate or caudate; midrib and nerves prominent beneath, impressed or flattened above; nerves 6-12 pairs, parallel, arcuating near the margin, at an angle of 60—70°, petiole densely stellate-tomentose when young, soon glabrous, 2-414 cm, deeply sulcate especially near the top. Male rachis in a paniculate cluster on the lateral new shoot, c. 5—20 cm, densely stellate-tomentose; 3 flowers: perianth lobes rather thick-coriaceous; filaments slender, hairy at base. Female rachis solitary in the axil of a higher leaf; bracts ovate- acute; 2 flowers: perianth densely stellate-tomen- tose, staminodes well-developed, styles short, con- nate at base. Young cupule obconical-turbinate, base attenuate, lamellae thin, densely stellate-to- mentose. Ripe cupule cup-shaped, obconical, taper- ing towards the base, 1—-1.7 cm high and 114-2 cm @; lamellae c. 7. Fruit obovoid-globose, c. 2 by 2 cm, top depressed, base convex. Distr. Malesia: Malay Peninsula (Pahang, Trengganu), Borneo (Sarawak, Mts Gaharu and Pueh). Ecol. In Malaya it is found in hill Dipterocarp forest at c. 1000 m, and in Borneo in heath forest, also at c. 1000 m. Fi. Jan.—Febr., fr. June—July. 4. Quercus gaharuensis SOEPADMO, Gard. Bull. Sing. 21 (1966) 384, f. 3; ibid. 22 (1968) 389, map 5. Tree, 10-30 m, 0.3-1 m 9; buttresses short, spreading, up to 0.7 m tall; bark smooth, mottled hooped, lenticels scattered in longitudinal rows. Branchlets densely stellate-tomentose, glabrescent, lenticellate; terminal bud ovoid-globose, c. 2 by 2 mm. Stipules linear-acute, c. 5 by 1 mm. Leaves thin-coriaceous, elliptic-lanceolate or elliptic-ob- long, 5-23 by 2-9 cm, entire or remotely serrulate in the apical part, base attenuate-acute or rounded, slightly asymmetrical, top rounded or bluntly acute; above glabrous, beneath densely stellate- pubescent, glabrescent; midrib and nerves pro- minent beneath, slightly so above; nerves 8-15 pairs, parallel, arcuating towards the margin, at an angle of 45-60°, reticulation distinct beneath; petiole 1-314 cm, slender, terete or adaxially flat, densely stellate-tomentose, glabrescent. Male rachis 14-314 cm; bracts ovate-acute, 1-114 by 1 mm; ¢ flowers: filaments hairy at base, 1-2 mm; anthers c. 14 by 14 mm; pistillode replaced by a tuft of stiff hairs. Female rachis 1-1 cm; bracts ovate-linear, 1-2 by 14-1 mm; 2 flower: stamino- des absent, styles slender, cylindrical, hairy at base, 1-2 mm. Young cupule ovoid-globose, 11-2 cm through, top rounded or truncate, base atten- uate-rounded, densely sericeous outside and densely brownish stiff-pubescent inside. Mature cupule deeply cup-shaped, 2-24 cm through; base rounded, enclosing 14-24 part of the fruit; lamellae 6-8, rims thin, slightly recurved and denticulate. Fruit ovoid-conical, 2-3 by 2 cm, top acute or rounded, convex at base. Distr. Malesia: Sumatra (East Coast Res. and SE. of Padang), Malay Peninsula (Perak, P. Pe- nang), Borneo (Sarawak and adjoining Sipitang in Sabah). Ecol. Forest, 100-1400 m. Fr. Sept.—Febr. Note. Specimens from Malaya reckoned by Hook. f. Fl. Br. Ind. 5 (1888) 603 and by later authors to Q. oidocarpa, belong here. 392 FLORA MALESIANA [ser. I, vol. 7? 5. Quercus oidocarpa Kortu. Kruidk. (1844) 216, t. 47: 18; BL. Mus. Bot. 1 (1850) 302; Mra. FI. Ind. Bat. 1, 1 (1856) 856; Ann. Mus. Bot. Lugd.—Bat. 1 (1863) 115; A. DC. Prod. 16, 2 (1864) 99; WeEN- ziG, Jahrb. Bot. Gart. Berl. 4 (1886) 233; Hook. f. Fl. Br. Ind. 5 (1888) 603; KinG, Ann. R. Bot. Gard. Calc. 2 (1889) 28; HickeL & A. CAmus, FI. Gén. I.-C. 5 (1930) 952; A. Camus, Chénes 1 (1938) 213, t. 7: 10-15; SoEPADMo, Gard. Bull. Sing. 22 (1968) 390, map 6. — Cyclobalanopsis oidocarpa (KORTH.) OERST. Vid. Medd. Nat. For. Kjobn. 8 (1867) 78. — Q. brevistyla A. CAMus, Bull. Soc. Bot. Fr. 80 (1933) 353; Chénes 1 (1938) 276, t. 17. — Fig. 33. Tree, 25-30 m, c. 50 cm g; bark grey, scaly and peeling off profusely into rectangular pieces. Branchlets initially with dense stiff pubescence, gla- brescent, lenticellate; terminal bud ovoid-globose, 3—5 by 2-3 mm. Stipules linear-acute, 3-4 by 14-1 mm, densely fulvous-tomentose by simple hairs. Leaves oblong-lanceolate or elliptic-oblong, 7-17 by 3-7 cm, remotely serrulate in the apical half; base acute or rounded or subcordate, top acute or 14-114 cm acuminate; above glabrous, beneath sparsely stellate-tomentose; midrib and nerves prominent beneath, impressed above; nerves c. 9-13 pairs, parallel, at an angle of c. 45°; reticula- tion distinct beneath, scalariform, dense; petiole Ja Fig. 33. Quercus oidocarpa Kortu. a. Habit, b. young cupules, c. ripe cupule and fruit, d. ripe fruit, e. Jongitudinal section of cupule and fruit, all x 24 (a BACKER 25956, b Ja. 4784, c—e IcuLas 115). 1972] FAGACEAE (Soepadmo) 393 114-314 cm, shallowly furrowed, glabrous. Male rachis 5—7 cm, densely stiff pubescent; bracts ovate -acute, 114-2 by 1-1!14 mm, glabrous; ¢ flowers: perianth membranous, glabrous; filaments slender, glabrous, c. 2 mm; anthers c. 0.7 mm long, Female rachis 3-7 cm, subglabrous, carrying c. 3-7 flowers, bracts ovate-acute; 2 flowers: staminodes absent, styles free, recurved, c.2 mm, hairy at base. Cupule deeply cup-shaped, 3-314 cm through; base atten- uate-rounded; lamellae thin, 9-11, lower ones denticulate, the others entire, yellowish brown tomentose. Fruit ovoid-globose or ovoid-cylin- drical, 2144-314 by 2-3 cm, top rounded or abrupt- ly depressed, base rounded or flat. Distr. Malesia: Sumatra (western central part), Malay Peninsula (Pahang, Selangor), Java (east- wards to Banjumas, Pringombo). Ecol. Forests, 150-1500 m. F/. March-April, fr. April-Febr. 6. Quercus sumatrana SoEPADMO, Gard. Bull. Sing. 21 (1966) 387, f. 4; ibid. 22 (1968) 392, map7. Tree, c. 35 m, 130 cm g; buttresses 1-214 m tall; bark rough, grey. Branchlets initially densely brownish, stiff-pubescent, late glabrescent, smooth lenticellate; terminal buds ovoid-globose, 2-3 by 114-2 mm. Stipules linear-acute membranous, 7 by 1 mm, densely fulvous-tomentose by simple hairs. Leaves chartaceous, elliptic-lanceolate, 8-— 24 by 2-8 cm, entire and undulate or remotely serrulate in the upper part, base attenuate-acute, top sharply 14-1!4 cm acuminate; above glabrous, underneath sparsely set with stiff, simple hairs, glaucous; midrib and nerves prominent beneath, slightly so above; nerves 8-13 pairs, rarely oppo- site, at an angle of 45—60°, parallel, arcuating near the margin; reticulation scalariform, obscure on both surfaces; petiole slender, 1-2'4 cm, terete or adaxially flat, sparsely pubescent. Ripe cupule shallowly cup-shaped, 0.7—1 cm high, 2-214 cm 9, covering 4%—1/, part of the fruit, base attenuate- rounded, densely tomentose on both faces; lamel- lae free at the rim. Ripe fruit ovoid-conical, 1.8—2 by 114-2 cm, densely sericeous, top acute, base concave. Distr. Malesia: Sumatra (Bencoolen, also Simalur), Borneo (northern Sarawak; western Sabah; in Kalimantan near Balikpapan and Sama- rinda). Ecol. Forests, up to 1300 m, on sandy loam or basalt-derived soil. Fr. April—Dec. Note. Inflorescences are as yet unknown. 7. Quercus subsericea A. Camus, Bull. Soc. Bot. Fr. 80 (1933) 354; Chénes 1 (1938) 367, t. 33: 21-26; SoEPADMO, Gard. Bull. Sing. 22 (1968) 392, f. 9, map 8. — Q. sericea SCHEFF. Nat. Tijd. N. I. 31 (1870) 361; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 63, t. 57B, non WILLD. (1805). — Cyclobalanopsis sericea (SCHEFF.) SCHOTTKY, Bot. Jahrb. 47 (1912) 656. — Synaedrys sericea (SCHEFF.) Korpz. Bot. Mag. Tokyo 30 (1916) 192. — Q. lineata (non Bui.) S. Moore, J. Bot. 63 (1925) Suppl. 114, p.p., quoad specim. FoRBES 572. — Q. oidocarpa (non KortTuH.) Merr. En. Born. (1921) 214, p.p., quoad specim. BECCARI 2919. Tree, 6-15 m, 10-40 cm @g; bark finely fissured or scaly, thin, greyish brown; inner bark fibrous, brownish, c. 2!4 cm thick. Branchlets initially densely set with stiff simple hairs or adpressed stellate hairs, glabrescent, later lenticellate; termi- nal buds ovoid-globose, 3—4 by 2—3 mm, scales narrowly ovate, densely simple sericeous. Stipules linear-acute, 7 by 1-2 mm. Leaves thin-coriaceous, elliptic-lanceolate, rarely ovate, 5-16 by 1-5 cm, base acute or roundish, margin entire or remotely serrulate near the sharply acute or 14-114 cm acuminate apex; beneath with a sparse stellate tomentum, above glossy, glabrous; midrib and nerves prominent beneath, impressed above; ner- ves 6-12 pairs, at an angle of 45-60°, parallel, arcuating towards the margin; reticulation obscure; petiole slender, 1-244 cm, adaxially furrowed. Male rachis 2—5 cm; bracts ovate-acute, tomentose outside, 1-2 by 1 mm; ¢ flowers 1-3; perianth tomentose, filaments slender, 1-2 mm, anthers c. 1 by 1 mm. Female rachis \4-1% cm, carrying 2-5 flowers; bracts ovate-acute, 0.5—0.7 by 0.3-0.5 mm; & flowers: ovary ovoid-cylindrical, c. 4 mm long, rounded-triangular, staminodes 0, styles c. 1 mm, tomentose at base. Young cupule ovoid- globose or obconical, lamellae 4-5, the 2-3 lower ones denticulate, the others entire. Ripe cupule cup-shaped, obconical, covering 4—! part of the fruit, 0.5—1.2 cm high, 0.7—-1 cm @, base rounded or attenuate, lamellae densely tomentose. Fruit ovoid-conical or ovoid-globose, 1-214 by 1-14 cm, densely sericeous, base obtuse or convex, apex acute or abruptly depressed-umbonate. Distr. Malesia: Sumatra (scattered), Malay Peninsula (Kelantan, Negri Sembilan, Pahang, Malacca), Banka (common), Borneo (scattered), W. Java (Pasir Orai: FORBES 572). Ecol. Forests, mostly at low altitude, in Java found at 850 m, on Mt Kinabalu at 1500 m; on sandy clay, sandy loam or granitic soil. Fertility seems irregular. J 8. Quercus lowii KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 28, t. 21B; A. Camus, Chénes 1 (1938) 366, t. 33: 27-29; SoppADMO, Gard. Bull. Sing. 22 (1968) 394, f. 10, map 9. — Cyclobalanopsis lowii (KING) ScHoTTKY, Bot. Jahrb. 47 (1912) 653. Tree, 10-20 m, 30-40 cm g. Branchlets initially densely set with brownish stellate hairs, late gla- brescent, dark grey, shallowly fissured; older branchlets lenticellate, glabrous; terminal buds ovoid-globose, densely tomentose. Stipules linear- acute, 2-4 by 14-14 mm. Leaves thick-coriaceous, ovate-elliptic or ovate-lanceolate, (3—)5—10(-14) by 2-514 cm; base rounded or cordate, slightly asym- metrical, top sharply acute or 14-2 cm acuminate; margin entire or remotely serrulate in the apical half; beneath densely brownish stellate tomentose or almost glabrous, above glabrous and glossy or dull and with sparse stellate tomentum (especially on the midrib); midrib and nerves impressed above, prominent or flattened beneath; nerves 5-8 pairs, at an angle of 50—-60°, parallel, arcuating towards 394 FLORA MALESIANA [ser. I, valk (7? the margin; reticulation fine, more or less areolate, distinct beneath; petiole 1-214 cm, sulcate, densely tomentose by simple or stellate hairs. Male rachis slender, 5-10 cm, simple or rarely much-branched; bracts ovate-acute, 1-2 by 14 mm, densely stellate- tomentose, glabrescent; § flowers 1-3; perianth densely tomentose by simple or stellate hairs, gla- brescent, anthers | by 14-1 mm, filaments 1 mm, pistillode replaced by a cluster of woolly, simple hairs. Female rachis 1-2 cm, with 2-5 2 flowers, staminodes none, styles c. 1 mm, recurved. Young cupule obconical, 0.8—1 cm high, 0.7—1.2 cm 9; lam- ellae thin, 5—7, the lower ones denticulate, the rest entire, densely stellate-tomentose. Ripe cupule cup- shaped, 0.8—1 cm high, 1.3—1.8 cm 9, base rounded, covering !4—1% part of the fruit. Fruit ovoid-cylin- drical, 114-2 by 1.3—1.5 cm, top rounded or acute, base convex; densely stellate-tomentose, glabres- cent. Distr. Malesia: Borneo (Sabah, various places, common on Mt Kinabalu; HALLIER 2950 from Mt Liang Gagang in western Kalimantan). Ecol. Forests, up to 2500 m, mostly in the mont- ane zone on blackish ultra-basic soil. Fr. July— March. Note. In the lowland specimens the leaves are glabrous, in mountain specimens they are mostly tomentose beneath; intermediates exist, see SOE- PADMO (1968). 9. Quercus percoriacea SOEPADMO, Gard. Bull. Bull. Sing. 21 (1966) 382, f. 2; ibid. 22 (1968) 396, map 6. Tree, trunk c. 50 cm 9; buttresses up to 1 m tall; bark smooth, grey, hoop-marked. Branchlets grey, glabrous, lenticellate; terminal buds ovoid- globose, c. 3 by 2 mm, densely tomentose. Leaves thick-coriaceous, ovate-elliptic, (10—)15-18(—20) by 5-9 cm; base rounded, top acute or 1-114 cm acuminate; margin incurved, remotely serrulate in the apical half, sparsely set with stellate hairs, glaucous, above glossy, glabrous; midrib and nerves prominent beneath, impressed and obscure above; nerves 8-10 pairs, at an angle of 45-60°, parallel, arcuating towards the margin; reticula- tion subscalariform, distinct beneath; petiole 2— 344 cm, glabrous, terete or adaxially + flat. Young infructescence 2-3 cm, rachis lenticellate, with 1-4 solitary young fruits. Young cupule obo- void, sericeous, 0.7—1 cm high, 0.8-1 cm @, base attenuate; lamellae 4-8, the lower ones denticulate, the others entire; young fruits ovoid-globose, attenuate towards the conical, ringed umbo, seri- ceous; styles recurved, sericeous. Ripe cupule flattened cup-shaped, 0.7-1 cm high, 2 cm a9, covering 14%—1/; part of the fruit, base rounded; lamellae 6-8, thin, more or less free at the rim, densely tomentose. Fruit ovoid-globose, c. 2 by 2 cm, densely tomentose, top rounded, umbonate, base convex. Distr. Malesia: Borneo (northern Sarawak: Ulu Baram). Ecol. Primary heath forest on terrace sand, with Agathis, 1100-1200 m. Fr. June-July. Note. The inflorescence is as yet unknown. 10. Quercus chrysotricha A. Camus, Chénes, Atlas 3 (1949) 50, t. 347: 7-14; Chénes 3 (1954) 1212; SOEPADMO, Gard. Bull. Sing. 22 (1968) 397, f. 11: la—c, map 10. Tree, c. 19 m, c. 30 cm 9; bark smooth or finely fissured. Branchlets initially densely brownish pubescent, late glabrescent, lenticellate; terminal buds ovoid-globose, c. 3 by 2 mm, scales ovate, glabrous. Stipules linear-acute, 8-10 by 1 mm, densely woolly rufous-tomentose by simple hairs. Leaves thin-coriaceous, obovate-elliptic, 1-514 by 114-34 cm; base attenuate-rounded or attenuate- acute, slightly asymmetrical, margin entire, apex rounded or truncate-emarginate; above glabrous, underneath sparsely pubescent, glabrescent; mid- rib and nerves slightly prominent beneath, flatten- ed or impressed above; nerves 4-5 pairs, at an angle of 60—-70°, parallel, arcuating towards the margin; reticulation obscure on both surfaces; petiole 1-5 mm, adaxially flat. Cupule cup-shaped, obconical, 1-114 cm through, base attenuate- acute, covering 14—14 part of the fruit; lamellae thin, 6-7, densely tomentose, denticulate. Fruit ovoid-conical, 114-2 by 1-114 cm, densely tomen- tose, glabrescent; top attenuate-acute, base con- vex. Distr. Malesia: Borneo (northern Sarawak, Dulit Ridge). Ecol. Mossy forest, 1200-1300 m. Fr. Sept. Notes. RICHARDS 1885, the type, was referred to Q. arbutifolia HickeL & A. Camus by E. F. WARBURG (Kew Bull. 1936, 19). The inflorescence is as yet unknown. 11. Quercus merrillii VON SEEMEN in Fedde, Rep. 5 (1908) 21; Merr. Philip. J. Sc. 3 (1908) Bot. 329; En. Philip. 2 (1923) 28; A. Camus, Chénes 1 (1938) 210, t. 7: 6-9; SoEPADMO, Gard. Bull. Sing. 22 (1968) 397, f. 11: 2a—c, map 10. — Cyclo- balanopsis merrillii (VON SEEMEN) SCHOTTKY, Bot. Jahrb. 47 (1912) 649. Small tree. Branchlets initially densely set with brownish, stiff hairs, glabrescent, lenticellate; terminal buds ovoid-conical, c. 3 by 2 mm, scales ovate, glabrous. Stipules linear-acute, densely pubescent outside, 5-10 by 1 mm. Leaves thin- coriaceous, obovate or elliptic-ovate, 114-5 by 0.7-214 cm, base attenuate-acute, top bluntly acute; margin remotely serrulate in the apical half; densely pubescent on both surfaces, late glabres- cent; midrib and nerves prominent beneath, flat- tened or impressed above; nerves 5-8 pairs, paral- lel, at an angle of 50-60°, arcuating towards the margin; reticulation obscure on both surfaces; petiole 3-6 mm, densely pubescent, glabrescent, adaxially flat. Young infructescence 1-2 cm, rachis densely tomentose, glabrescent, lenticellate, with 1-2 solitary young fruits. Young cupule obovoid or obconical, c. 14 cm through, densely tomentose; styles 1-2 mm. Ripe cupule cup-shaped, obconical, covering 14-14 part of the fruit; 1 cm high, 1-114 cm @; lamellae 7-8, thin, denticulate, densely tomentose. Fruit ovoid or ovoid-cylindrical, 2- 214 by 1-114 cm, glabrous, glossy, top acute, base convex. 1972] FAGACEAE (Soepadmo) 395 Distr. Malesia: Borneo (scattered in Sarawak and Sabah, also Nunukan I.), Philippines (Pala- wan only). Ecol. Forest, 100-500 m. Fr. April—Aug. Note. Inflorescences are as yet unknown. 12. Quercus kerangasensis SOEPADMO, Gard. Bull. Sing. 22 (1968) 399, f. 12, map 11. Tree, 20-30 m, 25-50 cm @; buttresses small, fluted; bark smooth or rough, lenticellate, grey. Branchlets initially densely stiff-pubescent, late glabrescent, greyish brown, densely warty lenti- cellate; terminal buds ovoid-globose, 3-6 by 3-4 mm. Stipules linear-acute, 2-3 by 14-1 mm. Leaves elliptic-lanceolate, coriaceous, 4-11 by 2-4 cm, above glossy, glabrous, underneath pale glaucous, glabrous except on the midrib, entire; base atte- nuate-acute, top acute or 14-1 cm acuminate; . midrib and nerves prominent beneath, slightly so above; nerves 7-10 pairs, ascending, subparallel, arcuating towards the margin, at an angle of 60- 70°; reticulation obscure on both surfaces; petiole 8-12 by 14-1 mm, densely pubescent, glabrescent, adaxially flat or shallowly furrowed. Male rachis 1-2 cm; bracts membranous, ovate-acute, densely stiff-pubescent outside, 1 by 1 mm; ¢ flowers with membranous perianth, filaments 1-2 mm. Ripe cupule cup-shaped, obconical, 8-12 mm high, 114-2 cm @Q, base attenuate, rim 1-114 mm thick, densely brownish tomentose on both surfaces; lamellae 5—7, entire or denticulate. Ripe fruit ovoid-conical or ovoid-cylindrical, 2-3 cm by 12-15 mm, densely sericeous, glabrescent, top rounded or acute, base convex. Distr. Malesia: Borneo (Sarawak; Brunei; in Kalimantan: Central Kutei near Long Bleh). Ecol. Primary heath (‘kerangas’) forests, to 100 m. Fi. June, fr. Sept. Note. The female rachis is still unknown. 13. Quercus kinabaluensis SOEPADMo, Gard. Bull. Sing. 22 (1968) 401, f. 13, map 11. Tree, 10—40 m, 20-30 cm 9; bark smooth, brown- ish with white patches. Branchlets initially den- sely brownish, stiff-pubescent, glabrescent, later with sparse lenticels; terminal bud ovoid-globose or ovoid-conical, 4-10 by 3-4 mm. Stipules linear- acute, 5-10 by 14-1 mm. Leaves thick-coriaceous, ovate-elliptic, 5-10 by 2-5 cm; surfaces densely yellowish brown pubescent, glabrescent; base asymmetrical, rounded or acute, top acute or %,-1\4 cm acuminate; margin entire; midrib and nerves slightly prominent beneath, flattened or impressed above; nerves 6-8 pairs, ascending, arcuating towards the margin at an angle of 45-60°; reticulation obscure on both surfaces; petiole 1-114 cm by 1-114 mm, adaxially flat. Male rachis 5-11 cm, 1 mm 9g; bracts ovate-acute, membranous, densely pubescent outside, 2-3 by 1-2 mm; ¢ flowers 3, filaments 2 mm, anthers c. 1 by 1 mm. Female rachis 1-3 cm by 2-3 mm, carrying 1-3 flowers; perianth thick-coriaceous, staminodes none, styles recurved, 2-3 by ',-1 mm. Young cupule cup-shaped, obconical, 0.7—1 cm high, 1-114 cm g, rim thick, base attenuate; lamellae 6-8, denticulate or entire, with some brownish pubescence on both surfaces. Fruit ovoid-conical, densely brownish tomentose, glab- rescent, 1-1.2 by 0.8-1 cm; top rounded or attenuate-acute, base convex. Distr. Malesia: Borneo (Sabah: Mt Kinabalu and Bk. Ampuon). Ecol. Forests, 500-2600 m, on ultra-basic soil. Fl. Febr., fr. March—May. 14. Quercus valdinervosa SOEPADMO, Gard. Bull. Sing. 22 (1968) 404, f. 14, map 12. — Q. mespili- folia var. borneensis HEINE in Fedde, Rep. 54 (1951) 225. Tree, 20-35 m, 30-60 cm @; bark smooth, grey. Branchlets initially densely brownish pubescent, hairs stiff, simple or stellate, glabrous, lenticellate; terminal buds ovoid-globose, 12-1 by 14 cm, the scales in four vertical rows. Stipules linear-acute, tomentose, 10-15 by 1-2 mm. Leaves thick-coria- ceous, elliptic or obovate-oblong, 8-15 by 3-6 cm; above glabrous, beneath densely pubescent, soon glabrescent; base acute or attenuate-rounded, top 14-1 cm acuminate; margin remotely serrulate in the apical half; midrib and nerves prominent beneath, flattened or impressed above; nerves 10— 15 pairs, dense, parallel and straight, arcuating near the margin at an angle of c. 60°; reticulation dense, scalariform, prominent beneath; petiole 1— 2'4 cm, glabrous, adaxially flat. Male rachis 5-10 cm; bracts lanceolate or linear-acute, membranous, 3-5 by 2-3 mm; ¢ flowers: perianth densely tomen- tose, filaments 3-4 mm, anthers c. 1 by 1 mm. Female rachis 1-11 cm, carrying 4-6 flowers; bracts ovate or linear, 2-3 by 1 mm; staminodes none, styles 1-2 mm. Ripe cupule cup-shaped, obconical, base attenuate, 1.5-1.7 cm high, 1-214 cm 9g, covering 144—1% part of the fruit; lamellae 7-8, densely tomentose rim crenate and thick. Fruit cylindrical-conical or ellipsoid, 3—3.7 by 1.3—1.7 cm, tomentose, top acute, base convex. Distr. Malesia: Borneo (Sarawak: Kalabit Highland and Mt Mulu; Brunei: Mt Pagon Periuk; Sabah: Mt Kinabalu; Kalimantan: Mt Kemul in W. Kutei). Ecol. Forests, 1200-2300 m, on brownish soil. Fl. Jan.—_Febr. and June, fr. Febr.—April and July. 15. Quercus gemelliflora BL. Verh. Bat. Gen. K. & W. 9 (1823) 222, t. 6, excl. fr.; Bijdr. (1826) 523; Fl. Jav. Cupul. (1829) 30, t. 17; A. DC. Prod. 16, 2 (1864) 88; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 88; K. & V. Bijdr. 10 (1904) 24; A. Camus, Chénes 1 (1938) 363, t. 23: 15-25; BACKER & BAKH. f. Fl. Java 2 (1965) 5; SoepADMo, Gard. Bull. Sing. 22 (1968) 406, f. 15, map 13. — Q. turbinata BL. Bijdr. (1826) 523, non Roxs. (1832); Fl. Jav. Cupul. (1829) 31, t. 18; K. & V. Bijdr. 10 (1904) 22; Ripu. Fl. Mal. Pen. 5 (1925) 336; HICKEL & A. Camus, FI. Gén. I.-C. 5 (1930) 949; A. Camus, Chénes 1 (1938) 204, t. 6: 9-16. — Q. merkusii ENDL. Gen. Pl. Suppl. 4, 2 (1847) 28; A. DC. Prod. 16, 2 (1864) 98. — Q. horsfieldii Mia. Fl. Ind. 396 FLORA MALESIANA [ser. I, vol. 7? Bat. 1, 1 (1856) 856; A. DC. Prod. 16, 2 (1864) 99. — Q. horsfieldii var. longifolia Mia. Fl. Ind. Bat. 1, 1 (1858) 869. — Cyclobalanopsis horsfieldii (MiQ.) Ogrst. Vid. Medd. Nat. For. Kjébn. 8 (1867) 78. — Cyclobalanopsis merkusii (ENDL.) OkrsT. /.c. 79. — Q. lineata var. merkusii (ENDL.) WENZIG, Jahrb. Bot. Gart. Berl. 4 (1886) 232; KING, Ann. R. Bot. Gard. Calc. 2 (1889) 33, t. 26: 2. — Q. semiserrata (non Rox.) Hook. f. Fl. Br. Ind. 5 (1888) 604; Kinc, Ann. R. Bot. Gard. Calc. 2 (1889) 28, quoad specim. e Sumatra et Banka; K. & V. Bijdr. 10 (1904) 25, p.p. — Cyclobalanop- sis turbinata (BL.) SCHOTTKY, Bot. Jahrb. 47 (1912) 648. — Q. turbinata var. crassilamellata GAMBLE, J. As. Soc. Beng. 75, ii (1915) 410. — Q. crassilamellata (GAMBLE) A. CAMUS, Bull. Mus. Hist. Nat. Paris II, 3 (1931) 689; Chénes 1 (1938) 226.t. 10> 1—7. Tree, 20-30 m, 20-60 cm g; buttresses up to 1 m tall; bark smooth or finely fissured, greyish brown. Branchlets initially densely brownish hairy, late glabrescent, lenticellate; terminal buds ovoid, 3-5 by 2-3 mm, scales broadly ovate, densely fulvous tomentose, by simple hairs, glabrescent. Stipules linear-acute, 5-10 by 1-2 mm. Leaves thin-coriace- ous, elliptic-lanceolate or elliptic-oblong, 5-15 by 2-5'%4 cm; base attenuate-acute, asymmetrical, top shortly acuminate; margin remotely serrulate in the apical half; above glabrous, beneath densely pubescent, soon glabrescent; midrib and nerves prominent beneath, flattened or slightly raised above, especially the midrib; nerves 8-10 pairs, parallel, straight, arcuating towards the margin, at an angle of c. 60°; reticulation obscure on both surfaces; petiole 1-3 cm by 1-2 mm. Male rachis c. 6 cm; J flowers: filaments c. 2 mm, anthers 14-] mm long. Female rachis carrying 2-7 flowers: bracts ovate-acute, 1-2 by 1 mm; staminodes 0-6, styles recurved, 1-2 mm. Young cupule turbinate- obconical or cylindrical-globose, covering the greater part of the fruit, base attenuate; lamellae 5-7, rim thick, entire or denticulate, densely to- mentose. Ripe cupule cup-shaped, 114-2 cm high, 114-244 cm @, covering 14-14 part of the fruit, rim c.2 mm thick; lamellae 7-8, densely tomentose. Ripe fruit conical cylindrical, 2-514 by 1-2 cm; top acute, base rounded; densely tomentose, gla- brescent. Distr. Malesia: Sumatra (common), Malay Peninsula (most provinces, also P. Penang), Banka, Borneo (scattered, in Sarawak, Sabah and eastern Kalimantan), Java (West Java, eastwards only on Mts Lawu, Wilis and Ardjuno). Ecol. Forests in lowland and montane zone, in E. Java up to 2160 m, usually near streams on red sandy clay or ultra-basic soil overlying sand- stone or granitic formations. Fertility seems to be irregular. DOcTERS VAN LEEUWEN, Zoocecidia (1926) 105 and 107, described some galls from Java and Sumatra, and BACKER & BAKHUIZEN f. hold it, that the large, globose, brown-pilose galls often borne by the twigs of this species, are unknown from other Quercus species in Java. 16. Quercus lineata BL. Bijdr. (1826) 523; Fl. Jav. Cupul. (1829) 32, t. 19; Mig. Fl. Ind. Bat. 1, 1 (1856) 855; A. DC. Prod. 16, 2 (1864) 98; WeNziG, Jahrb. Bot. Gart. Berl. 4 (1886) 232; K. & V. Bijdr. 10 (1904) 20, et Koorp. Atlas 1 (1913) t. 58, quoad Java; A. Camus, Chénes 1 (1938) 207, t. 6: 1-5; BACKER & BAKuH. f. Fl. Java 2 (1965) 5; SOEPADMO, Gard. Bull. Sing. 22 (1968) 408, f. 16, map 14. — Q. polyneura Mia. PI. Jungh. (1851) 11. — Q. lineata var. heterochroa Miq. Fl. Ind. Bat. 1, 1 (1856) 855. — Q. oxyrhyncha Miq. Sumatra (1861) 347. — Cyclobalanopsis lineata (BL.) OERST. Vid. Medd. Nat. For. Kjébn. 8 (1867) 78. — Q. lineata var. ‘typical lineata’ KING, Ann. R. Bot. Gard. Calc. 2 (1889) 33, t. 26: 1. — Q. lineata var. oxyrhyncha (MI1Q.) VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 4. — Q. hendersoniana A. Camus, Bull. Mus. Hist. Nat. Paris II, 4 (1932) 123; Chénes 1 (1938) 210, t. 6: 6-8. — Fig. 34. Tree, 20-30 m, 20-60 cm g; bark smooth grey; inner bark fibrous, reddish brown. Branchlets ini- tially densely pubescent, late glabrescent, dark grey, lenticellate; terminal buds globose or ovoid-conical, densely puberulous, glabrescent. Stipules linear- acute, 10-15 by 1-2 mm. Leaves thin-coriaceous, ovate-elliptic or ovate-lanceolate, 5-16 by 2-6 cm; base attenuate-acute, sometimes asymmetrical, top acute to 14-114 cm acuminate-caudate; margin remotely serrulate in the apical half, above with some pubescence, especially on midrib and nerves, beneath densely adpressed-pubescent; midrib and nerves prominent: beneath, slightly so above; ner- ves 10-20 pairs, straight, parallel, dense, arcuating near the margin but not anastomosing, at an angle of 45-60°; petiole 1-2 cm, shallowly furrowed. Male rachis 5-10 cm; bracts ovate-acute, 2-3 by 2mm, membranous; J flowers: filaments 14-1 mm, anthers c. 14 by 14 mm, pistillode replaced by a tuft of stiff, simple hairs. Female rachis 114-2 cm, carrying 5—6 flowers; bracts ovate-acute, 1-2 mm; staminodes none, styles recurved, 1-3 mm. Young cupule ovoid-globose, densely sericeous, lamellae 3-4, thick, the 2 lower ones denticulate, the others entire. Ripe cupule cup-shaped, obconical, 1-1.2 cm high, 2-214 cm @g, attenuating towards the base, densely tomentose, rim thick; lamellae 8-10, free. Ripe fruit conical-cylindrical, 2-3 by 1—2 cm, densely tomentose; apex attenuate-round- ed, base convex. Distr. Malesia: Sumatra (scattered in the N. and central part), Malay Peninsula (Pahang, Trengganu), Java (scattered in W., in Central Java: Mt Telemojo and Mt Wilis), Borneo (Sara- wak: Mts Pueh and Berumput; Sabah: Mt Kina- balu). Ecol. Forests, 1000-2000 m, on yellow sandy or ultra-basic soil. F/. April-May, fr. Aug. —April. Note. Non-Malesian records of Q. lineata relate to different species. 17. Quercus steenisii SoEPADMO, Gard. Bull. Sing. 21 (1966) 389, f. 4; ibid. 22 (1968) 411, map 3. Tree, c. 15 m, c. 60 cm g. Branchlets initially densely brownish stiff-pubescent, late glabrescent, lenticellate; terminal buds ovoid-ellipsoid, 1-114 1972] FAGACEAE (Soepadmo) 397 TW DCE Fig. 34. Quercus lineata BL. a. Habit with 3 rachis, x 25, b. 9 flower, <4, c. 9 flower seen from above, <4, d.stamen, x 8, e. habit with 2 rachis, x 24, f.Q flower, x4, g. style, <8, A. longitudinal section of a ° flower, <4, i. ripe cupule and fruit, x 24 (a HoLTTUM SF 31253, b-d RSNB 4927, e-h MEIER SAN 38069, i RAJAB 607). by 14 cm. Stipules linear, 5-10 by 1-2 mm. Leaves thick-coriaceous, ovate-elliptic or ovate-orbicular, 3-8 by 2-5 cm; base rounded, obtuse, or cordate, top rounded, emarginate or bluntly acute; margin entire or remotely serrulate in the apical half; above with some brownish, stiff simple hairs, glabrescent, beneath brownish, woolly tomentose, soon glabrescent; midrib and nerves prominent beneath, keeled or flattened above; nerves 6-10 pairs, straight, parallel, arcuating towards the margin, at an angle of 45—60°; reticulation scalari- form, prominent beneath, dense; petiole 2-5 mm long, adaxially flat. Male rachis 5 cm; bracts ovate- obtuse, c. 2 by 1 mm, densely pubescent; 3 flowers: filaments 1—-1.3 mm long, anthers c. 1 by 14 mm, pistillode replaced by a tuft of stiff, simple hairs. Young infructescence 114-3 cm, sparsely lenticel- late, carrying 2—7 young fruits; bracts ovate-acute; perianth thick-coriaceous, staminodes 0-6, rudi- mentary, styles recurved, 1-2 mm long. Young cupule ovoid-globose, densely pubescent; lamellae thick, 3-4, the 2 lower ones denticulate, the others 398 entire. Ripe cupule cup-shaped, obconical, or patelliform, 0.7-1 cm high, 1.2—-214 cm g, enclos- ing 44-4 part of the fruit; lamellae 7-8, + re- curved, densely brownish pubescent. Fruit globose or ovoid, densely brownish tomentose, glabrescent, 1-2 cm through; top rounded- or depressed-umbo- nate, base convex or fiat. Distr. Malesia: N. Sumatra (Gajo Lands). Ecol. Mossy forest on ridges, 2000-3350 m. Pilani Renn. Note. The female rachis and flowers are as yet unknown. 18. Quercus treubiana VON SEEMEN, Bull. Dép. Agr. Ind. Néerl. 1 (1906) 3; A. Camus, Chénes 1 (1938) 368, t. 33: 15-17; SoEPADMO, Gard. Bull. Sing. 22 (1968) 412, f. 17, map 15. — Cyclobalanopsis treubiana (VON SEEMEN) SCHOTTKY, Bot. Jahrb. 47 (1912) 648. Tree, c. 30 m, c. 60 cm @, bole irregular; but- tresses up to 1 m tall; bark rough, peeling off profusely into rectangular pieces, rusty; inner bark c. 114 cm thick, ridged. Branchlets initially densely brownish pubescent, later glabrous, lenti- cellate; terminal buds ovoid-ellipsoid, scales lin- ear-acute. Stipules linear-acute, 5-10 mm, dense- ly pubescent. Leaves thin-chartaceous, linear- lanceolate or elliptic-lanceolate, 3-10 by 1-3 cm; base attenuate-acute, sometimes asymmetrical, top sharply acute or 14-1 cm acuminate; margin re-. motely serrulate in the apical half; beneath densely pubescent, glabrescent, above glabrous except the midrib and nerves; midrib and nerves more or less prominent on both surfaces; nerves c. 5—10 pairs, straight, parallel, arcuating towards the margin, at an angle of 45—60°; reticulation obscure on both surfaces; petiole 44-114 cm, adaxially flat. Male flowers: perianth densely sericeous outside, filament c. 2 mm, anther 14-1 mm through, pistil- lode reduced to a tuft of stiff, simple hairs. Young infructescence: rachis 1-2 cm by 1—2 mm, densely woolly pubescent, glabrescent, carrying 2-5 young fruits; bracts ovate-acute, 1-114 by 14-1 mm, densely tomentose; perianth (in young fruit): styles recurved, c.2mm. Young cupule ovoid-glob- ose or obconical, lamellae c. 6, densely tomentose. Ripe cupule cup-shaped, base attenuate-rounded, 114-2 cm high, 1-2 cm @, covering %4,—!4 part of the fruit; lamellae 10-12, thick, densely seri- ceous. Ripe fruit cylindrical-globose, 2-3 by 114-2 cm; top depressed-umbonate, base convex. Distr. Malesia: Sumatra (near Palembang), Borneo (Sabah: Mts Kinabalu and Tambuyokan; Kalimantan: scattered). Ecol. Forests, 600-2100 m, on sandy waterlog- ged soil. Fr. July—Aug. FLORA MALESIANA (ser. I, vole a? Note. The female rachis is not yet known. 19. Quercus elmeri MerR. PI. Elm. Born. (1929) 43; A. Camus, Chénes 1 (1938) 194, t. 4: 9-12; Sog- PADMO, Gard. Bull. Sing. 22 (1968) 414, f. 18, map 16. Tree, 18-40 m, 25-60 cm @; bark greyish brown, cankered with longitudinal rows of lenticels or scaly, inner bark 14-114 cm thick, fibrous, reddish brown; buttresses narrow, up to 1.3-3 m tall. Branchlets initially densely rufous-tomentose, later glabrous, lenticellate; terminal buds subglo- bose, 3—5 by 4-5 mm. Stipules linear-acute, 5—7 by 1 mm. Leaves thin-coriaceous, elliptic-lanceolate or elliptic-oblong, 5-14 by 1-5 cm; above glabrous, glossy, beneath densely rufous-tomentose, gla- brescent; base attenuate-acute, usually asymmet- rical, margin remotely serrulate near the acute or sharply acuminate apex; midrib and nerves strong- ly prominent beneath, flattened or slightly raised above; nerves 5-12 pairs, parallel, arcuating to- wards the margin, at an angle of 60—70°; reticula- tion fine, subscalariform, obscure on both sur- faces; petiole 1-3 cm, terete or adaxially flat. Young infructescence 1-2 cm, carrying 1-5 young fruits; bracts linear-acute, 1-2 mm long, densely tomentose; perianth of the 2 flower (in young fruit): staminodes 0, styles 2-3 mm, recurved. Young cupule ovoid-conical; lamellae 3-4. Ripe cupule shallowly cup-shaped, or patelliform, 5-7 mm high, 2-214 cm g, covering 4%4—-'4 part of the fruit; lamellae 5-7, thick; rim denticulate; base truncate or rounded. Rive fruit ovoid-conical or conical-cylindrical, 2-3 by 1144-2 cm, densely sericeous, glabrescent; base truncate or convex, top rounded- or depressed-umbonate. Distr. Malesia: Central Sumatra, Malay Penin- sula (Pahang), Borneo (N. Sarawak; Sabah; W. of Samarinda in Kalimantan). Ecol. Forests, up to 1400 m, on sandy loam or ultra-basic soil. Fr. Sept. —April. Note. Inflorescences not known. Excluded Quercus litoralis BL. Mus. Bot. 1 (1850) 303; Mio. FI. Ind. Bat. 1, 1 (1856) 864; A. DC. Prod. 16, 2 (1864) 106; KiNG, Ann. R. Bot. Gard. Calc. 2 (1889) 90; A. Camus, Chénes 3 (1954) 1161. — Pasania litoralis (BL.) OERST. Kong. Danske Vid. Selsk. Skrift. V, 9 (1871) 379. The type (BLUME s.n.) now preserved in Leyden, has been identified by Dr. A. J. G. H. KOSTERMANS as Cyclandrophora scabra Hassk. (Rosaceae). 5. TRIGONOBALANUS ForMAN, Taxon 11 (1962) 140; Kew Bull. 17 (1964) 381; ibid. 21 (1967) 331.— Fig. 35-38. Trees. Innovations densely set with fulvous to rufous adpressed and erect, simple 1972] FAGACEAE (Soepadmo) 399 Fig. 35. Trigonobalanus yerticillata FORMAN. Male. a. Habit, x 23, b. node, showing stipules, x5, c. g rachis, <6, d. 5 flower, x12, e. stamen, <8, f. bract and bracteoles, < 8 (a PooRE 1337, b—c CORNER RSNB 2732, d—f WHITMORE FRI 8631, e from Kew Bull. 17, 1964, 382, f. 1, 5). and stellate hairs. Terminal buds ovoid, scales imbricate. Stipules extra- or inter- petiolar. Leaves spirally arranged or in whorls of 3. Inflorescence catkin-like, male, female, androgynous or mixed. Male rachis slender, erect or pendent, simple or with some lateral branches, arising from the axil of a lower, normal leaf or in the axil of reduced leaves and crowded together in a lateral or subterminal paniculate cluster. Male flowers in dichasial clusters of (1—)3—7(—12), subtended by a bract and bracteoles; perianth membranous, campanulate, 6—lobed, the lobes imbricate; stamens 6, filaments free, exserted, anthers ovoid, 14-1 mm long, cordate at the base, basifixed, 2—loculed, longitudinally dehiscent; pistillode replaced by a cluster of erect simple hairs. Female, androgynous or mixed rachis slender, erect, not branch- ed, arising from the axil of al higher leaf. Female flowers in dichasial clusters of (1—)3-7(-15), subtended by a bract and bracteoles; perianth with 6 imbricate lobes, the lower parts adnate to the ovary; staminodes 6, well-developed and ex- ceeding the perianth or remaining shorter than the perianth, sometimes pollinifer- ous; ovary 3-celled, ovules 2 per cell, axillary and apical; styles 3,! recurved or connate at the base, stigmas capitate. Cupule sessile or %—-14 cm stalked, spirally 400 FLORA MALESIANA (ser. I vorr/* Vi BAL Q7, =; Sh NY HNN \ ait Nw Sit NAN \ i} Nyy \\ \\ \\ \ il | Pee f ve 7s) 2@ Fig. 36. Trigonobalanus verticillata FORMAN. Female. a. Habit, in fruit, 74, b. flowering rachis, x 6, c. longitudinal section of 2 flower, and cupule, x 12, d. fruiting cupule, x6, e. cupules showing scars of nuts, <6, f. nut, x6 (a, d—f ASHTON 19624, b—-c ILIAS Pate S 26547). set or in whorls of 2-3 along the rachis, open, 3—12—lobed, the adaxial lobe usually reduced and rounded; outside covered with transverse dentate lamellae or imbri- cate scales; containing 1—7(-12) fruits. Fruit trigonous in cross-section, ovoid-tri- angular in outline; wall thinner than 14 mm, for the greater part free from the cupule. Germination epigeal. Distr. Two species, one in N. Siam (Mts Doi Chang and Doi Nang Ka, in Chieng Mai Prov., +19° N, 99°E) and one in Malesia: Malay Peninsula, Borneo, and Central Celebes. Fig. 39. Ecol. Confined to everwet, montane forest at 850-1765 m; in Malesia fl. Aug.—June, fr. April-Noy., in Siam fl. Nov. - Jan., fr. March. 1. Trigobalanus verticillata FORMAN, Taxon 11 (1962) 140; Kew Bull. 17 (1964) 383, f. 1 & 2. — Fig. 35-38. Tree 10-36 m, 20-70 cm @, sometimes fluted, producing many sucker-shoots at the base; bark smooth, shallowly fissured to scaly, pale grey- brown. Branchlets initially rather densely fulvous to rufous stellate-hairy, trigonous, later glabrous, terete, dark greyish brown, densely lenticellate or verrucose, lenticels minute; terminal buds ovoid, 3-5 by 2-4 mm, scales narrowly ovate. Stipules interpetiolar, ovate-lanceolate, 4-S by 2-3 mm, sparsely stellate-hairy, soon caducous, leaving prominent scars. Leaves in whorls of 3, coriaceous, rigid, (4-)6—-8(-12) by (2-)3-4(-514) cm (index 1.8—-2.5), broadest at or slightly above the middle; above glabrous, dull to glossy, beneath sparsely puberulous by stellate hairs, especially on midrib and nerves; base acute to cuneate, decurrent, mar- gin crenate in the apical half, or entire and re- curved, top bluntly acute to rounded emarginate; midrib prominent beneath, slightly so above; nerves (6—)7—8(-10) pairs, prominent beneath, slightly so above, subparallel, at an angle of 50- 70°, arcuating but not anastomosing towards the margin; reticulation irregular to areolate, obscure to rather distinct beneath; petiole (3—)5—8(—10) mm, 1-2 mm @, adaxially flat, thickened and rugose at the base. Inflorescence male, androgy- nous or mixed, sparsely pubescent by stellate hairs; bracts and bracteoles membranous, broadly to narrowly ovate-acute, ciliate, 14-1 by 1-144 mm. Male rachis 5-10 cm, 1 mm 9, erect, simple or much-branched, arising from the axils of normal, lower leaves or in the axils of reduced leaves and crowded together in lateral or subterminal pani- culate clusters; ¢ flowers in clusters of 3-12, perianth usually 6-lobed, filaments c. 1 mm, an- 401 1972] FAGACEAE (Soepadmo) Fig. 37. Trigonobalanus verticillata FORMAN. Tree habit showing the basal suckers, Mt. Kinabalu, Sabah (CORNER, 1964). 402 FLORA MALESIANA [ser. Ty vols 7? Fe: 4 Fig. 38. Trigonobalanus verticillata FORMAN. Branchlets with two 2 rachis, Mt Kinabalu, Sabah (CORNER, 1964). 1972] FAGACEAE (Soepadmo) 403 Fig. 39. Distribution of the genus Trigonobalanus, viz T. verticillata FORMAN (dots) and T. doichan- gensis (A. CAMUS) FORMAN (triangle). thers 14-1 mm long. Androgynous or mixed rachis 5-10 cm, 114-2 mm 9; female flowers in clusters of 3—7(-15), rarely solitary, staminodes 6, rather well-developed but not exceeding the perianth, styles terete, c. 1 mm, connate but at the distal part. Infructescence 5-10 cm, 2-3 mm 9, carrying numerous cupules arranged in whorls of 2-3, rachis more or less triangular. Ripe cupule sessile, enclosing (1—)3—7(—10) fruits, 4-6 mm long, width 8-15 by 5-10 mm, open, 3—12—lobed, the lobes acute, 3-4 mm long, inside densely fulvous tomen- tose by adpressed simple hairs, outside densely yellowish brown to fulvous stellate hairy and with 4-7 transverse rows of acutely denticulate lamel- lae. Ripe fruit sharply trigonous but not winged, 5-7 mm long, near the base 3—5 mm wide, outside sparsely stellate hairy; base truncate, scar flat, triangular, c. 2 mm Q, top acute. Distr. Malesia: Malay Peninsula (common on Fraser Hill), Borneo (Sarawak: Mt Hose, Carapa Pila, Balleh, Ulu Baram; Sabah, common on Mt Kinabalu), Central Celebes (twice collected from Pena, above Binuang, Mamasa, at 3° S, 119° 3’ EB). Fig. 39. Ecol. Forests at 850-1500 m, usually on ridges, on blackish ultra-basic or loamy soil overlying Tertiary sandstones. F/. Sept.—Febr., fr. April—Nov. Both in Malaya and Borneo, the species grows gregariously in association with Dacrydium elatum, Podocarpus imbricatus, and Agathis alba. Vern. Borneo: bérangan béluyan, Iban, salad rettan Kelabit; Celebes: mérang, Toradja. Note. Leaves collected from a basal shoot and also from the seedling measured 12-18 by 4-6 cm, with a long acuminate top, and are either alternate or decussate; the stipules are linear, up to 6 by 2 mm. Both in Malaya and in Borneo, the main trunk has been reported after dying off to be replaced by several subsidiary boles arising from the sucker-shoots. Fig. 37. Dr. D. H. Mat (Jb. Geol. 3 fiir 1967, 1970, 381-— 409, t. 1-4) has attributed fossil remains of Dryo- phyllum from Europe to Trigonobalanus, which would enormously extend its former range. Mr. FORMAN rejects this reduction. € 2 i PASSIFLORACEAE (W.J.J.O. de Wilde, Leyden) Mostly climbing herbs or lianas with axillary tendrils, rarely erect herbs, shrubs or small trees, glabrous or hairy, in Mal. not spiny. Branching usually by a supra- axillary serial bud. Leaves (mostly) spirally arranged, simple or compound, pinni- nerved or palminerved, entire or lobed; petiole or blade-base often with 1—-many glands, and often glands on margin and lower surface of the blade. Stipules present. Inflorescences essentially axillary, cymose, sessile or peduncled, 1—many-flowered, ending in (a) tendril(s) or not. Bracts and bracteoles mostly small. Flowers often stiped, articulate to the pedicel, actinomorphic, bisexual or functionally unisexual (either with staminodes or a vestigial ovary, and then plants mostly dioecious) or polygamous. Perianth mostly 2-seriate, mostly persistent, the segments free or partially connate (Adenia p.p.), inserted on the rim of the saucer- or cup-shaped or tubiform hypanthium. Sepals (4-)5(—6), imbricate. Petals (4-)5(—6), mostly imbri- cate. Corona inserted on the hypanthium, mostly a complicated structure, com- posed either of filaments, hairs, or appendages, or membranous, annular, or compo- sed of scales (disk), or in addition with ‘septa’ (Adenia p.p.), rarely corona absent (Adenia p.p.). Stamens 4-10, inserted mostly at the base of the hypanthium, or on an androgynophore (mostly hypogynous), (mostly) opposite the sepals; filaments free or partially connate into a tube; anthers 2-celled, longitudinally dehiscent, sometimes apiculate. Ovary superior, subsessile or on a gynophore or androgyno- phore, |-celled, 3(—5)-carpellate; placentas 3(—5), parietal; ovules many, anatrop- ous; integuments 2; styles 1 or 3(—5), very short to distinct, sometimes partially connate; stigmas + globose, or capitate, or papillate, or much divided. Fruit a loculicidally 3(—5)—valved capsule, or berry-like. Seeds mostly numerous, mostly compressed, often beaked, enveloped by a (membranous or juicy) aril; funicles often distinct; testa crustaceous (coriaceous), mostly striate, reticulate or pitted; endosperm (copious) horny; embryo straight; cotyledons foliaceous. Cf. HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 470-507. Distribution. About 10 genera and 500 spp., almost entirely confined to the tropics: in America c. 350 spp. (mainly Passiflora, a few species in Dilkea, Mitostemma, Tetrastylis), in Africa (incl. Madagas- car) c. 110 spp. (mainly Adenia c. 80 spp., Tryphostemma c. 20 spp., Deidamia, incl. Efulensia, c. 6 spp., Crossostemma, incl. Schlechterina, 2 spp.), in Asia and Australia c. 40 spp. (Passiflora c. 20 spp., Adenia 14 spp., Hollrungia | sp., Tetrapathaea | sp. in New Zealand). The two largest genera are each distributed in two continents, viz Passiflora in America and Austral- asia (S. & SE. Asia, Malesia, NE. & E. Australia, W. Pacific islands), and Adenia in Africa, Madagascar, SE. Asia, and Queensland. In addition the tribe Pariopsieae is now arbitrarily reckoned to the Passifloraceae and excluded from the Flacourtiaceae. They are all woody, erect plants and occur in the palaeotropics, only one species being represented in Malesia. See SLEUMER, Bull. Jard. Bot. Brux. 40 (1970) 49 and De wiLpbeE, Blumea 19 (1971) 99. The characters by which this tribe differs from Passifloraceae in the strict sense are here not incorporated in the family circumscription. See further below under Taxonomy. Ecology. Mostly heliophilous climbers at low and medium altitude, Adenia kinabaluensis at c. 2000 m on Mt Kinabalu; in Andean Peru Passiflora mixta up to 3500 m. Usually scattered, only the introduced Passiflora foetida sometimes very common in thickets. Pollination. Most genera are monoecious with bisexual flowers, Passiflora having a marked protandry; Adenia is monoecious or dioecious, or polygamous. Functionally unisexual flowers occur in Adenia, Hollrungia, and in the New Zealandian Tetrapathaea. Pollination by bumble-bees and kolibris is known from American Passifloras; for a number of narrow- flowered species of Adenia (Mal. spp.!) pollination by (small) insects is likely. Some African Adenias have fragrant flowers. Morphology. The axillary tendrils in sterile shoots replace the axillary inflorescences; inflorescences are essentially cymose and the first flower or first 3 flowers of the lowest triad may be replaced by (a) 406 FLORA MALESIANA [ser. Evel, tendril(s). Sometimes the cyme has become monochasial or is deformed by partial concaulescence, or the cymes reduced to 1-3 flowers are contracted into raceme- or panicle-like inflorescences (Passiflora race- mosa). Ramification of the plant takes place through the serial bud. Studies dealing with the inflorescences and tendrils of Passifloraceae have been made by HARMS (Bot. Jahrb. 24, 1897, 163-178), Cusser (Bull. Soc. Bot. Fr. 115, 1968, 45-61), and myself (Thesis, 1971, 16-17). With hypanthium in the flower descriptions is meant the usually cup-shaped basal part, which bears on its margin the tepals (mostly free; sepals in Adenia are often partially united into a calyx tube) and the co- rona (mostly filamentous). Lower down in the hypanthium various types of a disk may be found, mostly annular, in Adenia mostly consisting of 5 scale-like or strap-shaped appendages. The leaves mostly bear (often large) nectarial glands on petiole and blade. Seedlings of extra-Malesian species (Passiflora, Adenia) are depicted by LusBock (Seedlings 1, 1892, 582-593) and DE WILDE (Thesis, 1971, 24, fig. 4). Phytoc hemistry. Many members of the family are toxic. The toxic constituents are still incompletely “known. Most species of Adenia and Passiflora, and Deidamia clematoides and Barteria fistulosa (of the Paropsieae) release appreciable amounts of prussic acid on wounding. They contain the gynocardin-like glucosides deidaclin (first named deidamin) and barterioside and, most probably, also gynocardin itself (TANTISEWIE c.s. Pharm. Weekblad 104, 1969, 1341; Paris c.s. C. R. Ac. Sc. Paris 268 D, 1969, 2804; CLAPP c.s. J. Am. Chem. Soc. 92, 1970, 6378). The occurrence of this highly characteristic type of cyanogenic glucosides points to a rather intimate relationship between Passifloraceae and Flacourtiaceae- Pangieae. Besides cyanogenic glucosides other toxic principles seem to be present in some members of the family; a toxalbumin, modeccin, was reported as a constituent of Adenia (= Modecca) digitata. Notwithstanding their often toxic nature several species of Passiflora produce edible ‘Passion Fruits’ (see PurTHI, Advances in Food Research 12, 1963, 203-282; MARTIN & NAKASONE, Econ. Bot. 24, 1970, 333-343). Leaves and stems of the non-cyanogenic, temperate American species Passiflora incarnata are used in medicine as a sedative drug; Herba Passiflorae contains harman and related simple indolic alkaloids. Such alkaloids are also present in other species of Passiflora. The phenolic constituents of Passifloraceous plants are still very incompletely known. The so-called tannin cells of plant anatomists seem to contain catechins and leucoanthocyanins rather than true tannins. True tannins are lacking in the family or at the most are present in small amounts. Leaf flavonoids are represented by glucosides of kaempferol and quercetin (but not of myricetin) and espe- cially by C-glykoflavones like saponaretin (= isovitexin), vitexin and orientin. On the whole our knowledge of chemical characters of Passifloraceae is still very restricted. However, the common occurrence and the very peculiar nature of the gynocardin-like cyanogenic glucosides ac- centuate a Flacourtiaceous relationship. For a summary of phytochemical literature and references see HEGNAUER, Chemotaxonomie der Pflanzen 5 (1969) 293-298. — R. HEGNAUER. Anatomy. See HArs, Bot. Jahrb. 15 (1893) 548-633 and METCALFE & CHALK, Anat. Dicot. 1 (1950) 674. Uses. Various Passifloras are ornamental or have edible fruit. Adenias are sometimes used as fish poison. Because of the showy flowers or edible fruits many species are cultivated in the tropics and subtropics and frequently run wild. Most species are nitrophilous or ruderal and are found in secondary vegetation. Many species are easily propagated either by seeds or cuttings. See HEYNE, Nutt. Pl. (1927) and BuRKILL, Dict. (1935). Taxonomy. Related to Flacourtiaceae to which the tribe Paropsieae (shrubs or trees) forms a tran- sitional group. Recent anatomical evidence (AYENSU & STERN, Contr. U.S. Nat. Herb. 34, 1964, 45-73) and palynological studies (PRESTING, Pollen et Spores 7, 1965, 194-247; SpiRLET, ibid. 7, 1965, 249-301; PACQUE, ined.) point to a closer relationship of Paropsieae with Passifloraceae than with Flacourtiaceae. Bibliographical note. In my monograph of the genus Adenia published in the Med. Landbouw- hogeschool Wageningen 71-18 (1971) 1-281 I have also alluded to many general aspects of the family. This study is referred to as my ‘Thesis’. KEY TO THE GENERA 1. Climbing plants with tendrils. 2. Androgynophore long; flowers bisexual; styles long, stigmas -+ globular, smooth; anthers versatile. 1. Passiflora 2. Androgynophore 0 or short; flowers unisexual or polygamous; styles rather short, stigmas finely lobed or divided. 3. Corona 0 or consisting of fine hars; disk glands 5, era or strap- ignisick anthers narrow, erect, basifixed; flowers unisexual ... . ashy egpespAdenta 3. Corona double: outer corona consisting ‘of oe 3 rows ‘of filaments, inner corona (‘operculum’) laciniate; disk 0 or as a faint annulus at bottom of hypanthium; anthers broad, deeply sagittate, versatile; flowers (functionally) polygamous. srislbiecce cig |'> veg Se) RES . Erect shrubs or small trees. No tendrils. See Fl. Mal. I, 5 (1954) 13; eid. leone sceneu se Aa aronsia 1972] PASSIFLORACEAE (de Wilde) 407 1. PASSIFLORA Linne’, Gen. Pl. ed. 5 (1754) 410; Sp. Pl. 1 (1753) 955; DC. Mém. Soc. Phys. Genéve 1, 2 (1822) 434; Prod. 3 (1828) 322; BL. Rumpia | (1837) 169; Roem. Syn. Mon. 2, Pepon. (1846) 131, 165; BENTH. FI. Austr. 3 (1866) 311; BENTH. & Hook. f. Gen. Pl. 1 (1867) 810; MAst. Trans. Linn. Soc. 27 (1871) 593; Fl. Bras. 13 (1872) 531; in Hook. f. Fl. Br. Ind. 2 (1879) 599; HARMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 86; F. M. BAILEy, Queens]. Fl. 2 (1900) 686; GAGN. FI. Gen. I.-C. 2 (1921) 1016; HALL. f Med. Rijksherb. 42 (1922) 5; HARMs in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 495; Crap, FI. Siam. En. 1 (1931) 742; KiLLtp, Field Mus. Publ. Bot. 19, 1 (1938) 1-613 (American spp.); BACK. & Baku. f. Fl. Java 1 (1963) 289;° CussET, Fl. Camb., Laos & Vietn. 5 (1967) 106; HutcuH. Gen. FI. Pl. 2 (1967) 370. Climbers (in Mal.). Leaves (mostly) spirally arranged, very rarely (sub)opposite, simple or (extra-Mal.) compound, entire or lobed, pinni- or palminerved; petiole with or without glands. Stipules (in Mal.) minute. Jnflorescences sessile or pedun- cled, |—many -flowered, with or without a simple tendril; flowers rarely collected into pseudoracemes. Bracts (in Mal.) small. Flowers bisexual, 5-merous; hypan- thium saucer-shaped to cylindrical. Sepals and petals mostly free, often highly coloured; petals often resembling the sepals, membranous, (extra-Mal.) sometimes absent. Corona composed of a mostly complicated outer corona, and mostly a flat or plicate inner corona; nectary ring annular, within the operculum, or absent; disk at the base of, or on the androgynophore, or absent (in Australian spp.). Sexual organs on a distinct androgynophore. Stamens 5(-8), free filaments at first erect, later on mostly reflexed; anthers dorsifixed, versatile, elliptic to linear. Ovary glo- bose to fusiform, sessile or stalked; styles 3(-4), (in Mal.) free, long; stigmas capitate. Fruit mostly indehiscent, -—- baccate, often with coriaceous exocarp, globose to (rarely) fusiform. Distr. About 370 spp., of which c. 350 in the New World and c. 20 spp. in Indo-Australia and the West Pacific. The genus is absent in Africa. The species described from Madagascar (P. calcarata Mast.) and the Mascarene Is. (P. mauritiana THOUARS and P. mascarensis PRESL) pertain to early introduced species from America, the first most likely P. subpeltata ORTEGA, the latter two being P. alata DRYAND. Ecol. Rather low climbers, in primary and secondary forests, scrub and savannahs, below 1800 m, rather rare, under everwet climatic conditions except P. moluccana which prefers a seasonal climate. Uses. A number of species is introduced as ornamentals or for the edible fruit with delicate flavour. Edible are e.g. P. edulis Sims (Purple granadilla), P. /aurifolia L. and P. quadrangularis L. (Marquesa, Grenadilla). Uses and vernaculars of various Passifloras are given by HEYNE, Nutt. Pl. 2 (1927) 1142- 1144; Ocuse (& BAKH.) Fruit & Fruitculture (1931) 99-103; Ind. Groenten (1931) 575-581; BURKILL, Dict. 2 (1935) 1704-1706; ALLEN, Mal. Fruits (1967) 134-142. Taxon. Harms (1925) accepted 21 sections, K1LL1p (1938) 22 subgenera and many sections and series for the New World. The indigenous Old World species all belong to sect. Decaloba in the sense of HARMS, a section also represented in America. These species do not fit, however, into the section as conceived by KILLIP for the American species. Within sect. Decaloba three groups can be distinguished. Group | is characterized by creamy or white flowers less than 5 cm 9. To this group belong all continen- tal SE. Asian species and spp. 1-3 in this work. A. P. de CANDOLLE (1828) knew of this group only P. (1) The synonymy of the group of indigenous Old World species is given under sect. Decaloba. Other synonyms of Passiflora are: Cieca MEDIK., Murucuja [TOURN.] MEDIK., Granadilla [TOURN.] MEDIK., Tacsonia [TOURN.] Juss., Anthactinia Bory, Asephananthes Bory, Monactineirma Bory, Decaloba (DC.) ROEM., Astrophea (DC.) RoEM., Dysosmia (DC.) RoEM., Rathea KARSTEN. For further informa- tion one is referred to e.g. HARMS (1925), KILLIP (1938) and HUTCHINSON (1967). 408 FLORA MALESIANA [ser. I, voli\7= Ww Fig. 1. Passiflora hollrungii K.Scu. a. Habit, x ¥%, b. flower in longitudinal section, nat. size, c. fruits, x Y%, d. seed, X 6. — P. perakensis HALL. f.e. Flower in longitudinal section, <2. — P. aurantia Forst. f. var. aurantia. f. Flower in longitudinal section, nat. size (a BRASS 32290, b CARR 14428, c-d SAYERS 21268, e RAHMAT SI ToroEs 51, f BRASS 27591). 1972] PASSIFLORACEAE (de Wilde) 409 moluccana and P. leschenaultii which he referred to sect. Polyanthea, reduced to a subsection of sect. Decaloba by HArms (1925) and CusseT (1967). HARMS created two other new sections among the continental SE. Asian species, viz sect. Octandranthus to fit P. octandra GAGN. with 6-8 stamens and sect. Anomopathanthus for P. cochinchinensis SPRENG. with (sub)opposite leaves, both species from Indo-China. These two species are, however, very related to P. perakensis and P. moluccana. Group II is characterized by large, pinkish, orange or red flowers 5—10 cm g; the outer corona filamen- tous, the inner corona tubiform. To this group belong 4 species confined to New Guinea, Australia and the SW. Pacific. This group was formerly accommodated in a genus Disemma; HARMS reduced this to sect. Decaloba subsect. Eudecaloba. Group III is allied to group II but differs in having blue flowers in which the outer corona is tubiform and the inner corona small and incised. This comprises only one species, P. hollrungii, from New Guinea. Though these groups reflect affinity within the Old World species of sect. Decaloba, it appears to me that they should not be given separate taxonomic rank. Chromosomes. Of some tens of American species chromosomes have been counted; this yielded the numbers 2n = 12, 18 and 20, the most common number being 18. BEAL (Austr. Pl. 6, 1970, 13-14) reveal- ed that the three native Australian species P. aurantia, P. herbertiana, and P. cinnabarina, are all 2n = 12, adding that this would point to their primitiveness. Notes. I am confronted with the situation that of Passiflora in Malesia there are few indigenous and many introduced species, quite a few of which have run wild, sometimes profusely so, which causes con- fusion to collectors and their field notes. The five commonest among these are P. foetida L., P. suberosa L., P. edulis Sims, P. mollissima (H.B.K.) BAILEY, and P. mixta L. f. Under these circumstances it appeared out of proportion to study critically all these introduced species at the same level as the indigenous ones. Therefore, two keys are presented, one for the native species and one for the introduced ones. It is not possible to separate these by a single character; therefore several forks of the first key refer to the second key containing the introduced species. As a native New Guinean species occurs far into the Pacific (P. aurantia) I have included in the key also the two other Australasian species. KEY TO THE MALESIAN & AUSTRALASIAN-PACIFIC SPECIES 1. Bract and bracteoles distinct, (sub)foliaceous, forming an involucre. See Key to introduced species 1. Bracts and bracteoles minute, setaceous or linear, mostly scattered, not involucre-like. 2. Hypanthium long, mostly bowl-shaped or infundibuliform or tubiform. See Key to introduced species 2. Hypanthium rather shallow, -- saucer-shaped. 3. Operculum (inner corona) not plicate. .. . . . . See Key to introduced species 3. Operculum (inner corona) plicate, mostly with crenulatey or crispy or laciniate edge. ems TAUSCOE: =. tee sr nes ov alls Ou) DS) wtiea male SUM le | Seer Imey te imeroduced species 4. Petals present. 5. Inflorescences mostly shortly peduncled, (1—)3—30-flowered; flowers 2-414 cm @ (sepals 10-20 mm), tepals (creamy—) white, outer corona filaments in 2 rows. 6. Petiole without glands. . » 2 Wo ote 0. eeiSee iKey. tolimtroduced species 6. Petiole provided with glands (Group I). 7. Leaves shallowly 3-lobed, each lobe distinctly mucronate, leaf-base also with 2 mucros. See Key to introduced species 7. Leaves without distinct mucros. 8. Petiolar glands at apex or in upper 44 of petiole. Siar fone les) pel SE moluccana 8. Petiolar glands situated in the lower 2% of the petiole. 9. Filament connate for the lower 14- in enveloping the ovary; ovary hairy; leaf top acute. 2. P. perakensis 9. Filaments above androgynophore + free; ovary glabrous; leaf truncate or + 2-3-lobed. 3. P. sumatrana 5. Inflorescences sessile, 1—2-flowered; flowers 5-10 cm © (sepals (20—)25—50 mm), tepals pink to red, or bluish; outer corona filaments in 1 row or connate into a tube. 10. Outer corona filamentous, inner corona (operculum) tubiform, 5—15(-20) mm, reddish or whitish; flowers pinkish to orange or red (Group II). 11. Blade-base or petiole with glands. 12. Plant pubescent, rarely glabrescent; glands often + stiped; tops of leaf lobes + acute; co- rona filaments whitish, inner corona distinctly plicate with shallowly lobulate-crenulate edge. Australia: Queensland, New South Wales. ......... .. P. herbertiana 410 FLORA MALESIANA (ser. I, vol. 7? 12. Glabrous; glands sessile, flat; tops of leaf lobes acute, rounded, or emarginate; corona fila- ments purplish red, inner corona + wrinkled in the upper half, with shallowly lobulate- crenulate edge. Australia, Norfolk I. . ... Joe seat S040. ow) bea Parenti Blade-base or petiole without glands (extra- -Malesian). a Gynophore long; outer corona filaments whitish, longer than the whitish inner corona which has a densely fine-fimbriate edge. Australia: Victoria... .. . . . DP. cinnabarina 13. Gynophore short; outer corona filaments purplish red, about as long as or shorter than the inner corona, which has a shallowly lobed edge. Australia, Norfolk I. . . . 4. P. aurantia 10. Outer corona broadly tubiform, 20-30 mm, purple-blue, inner corona small, incised, 1-2 mm; flowers: bluish (Group TM) . 2°. «08 ud, hs, @ oe KEY TO INTRODUCED SPECIES 1. Bracts and bracteoles conspicuous, forming an involucre. 2. Involucral bracts finely and deeply divided; plant densely glandular hairy; flowers 2144-5 cm 9; petals white; corona filaments + purplish; fruit subglobose, c. 2 cm @, yellow to orange; variable, a ‘common, weed; 0-1500.m. . 2. . oc a SO FURS, OR? [So OP foci 2. Involucral bracts not divided. 3. Involucral bracts partially connate in a tube; flowers + pinkish, with hypanthium 6-9 cm long. 4. Bracts to 4cm, connate up to halfway; fruit ellipsoid, 7-12 cm, edible; ornamental and cultivated for the fruit; 500-2500 m... . . . . P. mollissima (H.B.K.) BAILeEy 4. Bracts to 5 cm, connate for + %; fruit ellipsoid: to 6 cm; occasionally cultivated (closely related to, and hybridizing with P. mollissima) ; 500-2000 m. . . . . P. mixta LINN. f/. 3. Involucral bracts free or shortly connate at base; hypanthium less than 3(—4) cm. 5. Flowers pink or red, pending on up to 20 cm long pedicels; hypanthium 2—4 cm; fruit + fusiform, longitudinally ribbed, to c. 12 cm, edible; also cultivated as ornamental; New Guinea, locally running wild; c. 2000 m. oe es ee we ew eee ee eee os ES Ps an tloquicnsisskeansn 5. Pedicels much shorter. 6. Leaves pinnately nerved, not lobed. 7. Stems 4-angular or -winged. 8. Petioles 6-glandular; stipules (lanceolate-) ovate, more than 1 cm wide; flowers 7-10 cm 9, purple-red; corona filaments banded, purple-white; fruit 12-30 cm long, edible; cultivated, sometimes escaped; to 1000 m.. . . . . PP. quadrangularis L. 8. Petioles 2—4-glandular; stipules lanceolate- linear: less than Ie cm . wide: flowers 10-12 cm g; co- rona filaments variegated with red, purple and white; fruit 8-10 cm long; occasionally cul- tivated as an ornamental. See also P. X alato-coerulea LINDL., below; 0-1000 m. P. alata DRYAND. 7. Stems not winged. 9. Stipules thread-like, c. 4% cm; leaves ellipsoid to oblong, coriaceous; flowers c. 8 cm 9, flushed with purple or purple-dotted; corona filaments purple with white cross-bands; fruit ovoid, 5—8 cm long, edible; frequently cultivated, 0-1000m.. ....... . .. P. laurifolia L. 9. Stipules foliaceous; leaves herbaceous. 10. Glands on petiole thread-like or long-cavate; leaves deeply cordate; flowers 6-9 cm 9, white or pale pinkish; corona filaments banded; fruits ovoid, 6-8 cm long, edible; locally cultivated and escaped; 0-1000 m ... . . . P. ligularis Juss. 10. Glands on petiole wart-like, sessile; leaf base 4 cordate, rounded or acute. 11. Involucral bracts very large, extending beyond the flower, acute-acuminate; stipules lanceo- late, longly acuminate, 1-114 cm; flowers c. 10 cm 9, mottled purple-red; corona filaments banded; fruit c. 4 cm 9; ornamental, sometimes escaped; 0-1000 m . . P. maliformis L. 11. Involucral bracts not extending beyond the flower, obtuse; stipules linear-subulate, 5—6 mm; flowers c. 10 cm 9, white and pinkish; corona filaments purplish banded; fruit globose, 3-6 cm 9; sometimes cultivated;0-1000m..... . wills its) Sates te aepemitidarhicbens: 6. Leaves palmately nerved, mostly lobed or partite. 12. Stipules lanceolate or filiform; involucral bracts serrate-denticulate. 13. Petiole with wart-like glands at apex; flowers 4-6 cm 9. 14. Plant glabrous; involucral bracts 14-2 cm; petals white, corona filaments white with purple base; fruit ellipsoid, 4-6 cm long, purplish, sometimes yellow; cultivated and profusely escaped: “0-2000 smite). Gresdeete). Menieeeed cee } auceeeirel cnet P. edulis Sims! (1) According to MERRILL, Sp. Blanc. (1918) 276 and En. Philip. 3 (1923) 118 this introduced species has twice been described by BLANCO from the Philippines, viz as P. minima Buco, FI. Filip. (1837) 647, oe L. 1753 and P. serrulata Buco, ibid. ed. 2 (1845) 452; ibid. ed. 3, 3 (1879) 50, t. 414, non JAcQ. 67. 1972] PASSIFLORACEAE (de Wilde) 411 14. Plant pilosulous; involucral bracts c. 1 cm; petals white or lavender, corona filaments mostly purplish or pinkish; fruit up to 5cm 9; sometimes cultivated; c.1500m. . P. incaranata L. 13. Glands at base of petiole; flowers c. 10 cm 9, red; plant ferruginous- re ene fruit ovoid, c.5cm long; 0-1000 m .. . > fmesieos, Sra. PR ERR vitioliathise kk. 12. Stipules foliaceous; involucral bracts (sub)entire. 15. Stipules 1-2 cm long, falcate, mostly dentate. 16. Glands on petiole wart-like, subsessile, c. 1 mm long; flowers white, lilac or purplish, 6-8 cm @. 17. Leaves (3—)5(—9)-lobed, incisions nearly to the base; fruit ovoid to subglobose, c. 6 cm long; mostly ornamental; 0-2000m...... edits con ted een > Doe RACERS Sls. 17. Leaves 3—5—lobed, incisions to about 34, of the blade. 18. Stem 4-angular; leaf 3-lobed, the lobes broad; niall ‘Impératrice Eugénie’; ornamen- tal; 0-2000m .. : . . P. x alato-caerulea LINDL. 18. Stem terete; leaves 3- i= lobed, the lobes narrower; ‘ornamental and escaped; 0—2000 m. P. caeruleo-racemosa SABINE 16. Glands on petiole filiform, 1-2 mm; flowers red, c. 6.8 cm 9; fruit not known; ornamental. P. kermesina LINK & OTTO 15. Stipulesiarge, 114-4 cm, straight, entire; flowers 4-5 cm wide, white; sepals 4-1 cm horned; fruit 244-4 cm o, with thick pericarp; ornamental and locally profusely escaped (India, Philippines, Queensland); 0-1000 m ... , . . P. subpeltata ORTEGA 1. Bracts and bracteoles inconspicuous (filiform or linear), « or “caducous before anthesis; no involucre. 19. Flowers red, arranged in pending racemes; involucral bracts caducous before anthesis; flowers red, c. 7cm g; fruit narrowly ovoid, S—7 cm; ornamental; 0-2000m..... . P. racemosa BROT. 19. Flowers 1 or mo in the axils of normal leaves; flowers greenish yellow or white, not red. 20. Flowers small, 1-114 cm g, apetalous, pale egies fruit a purple-black berry, 1-144 cm 9; variable, eee established; 0-1500 m. . . Di tarreh cia. ee . . . P. suberosa L. 20. Flowers more than 1% cm g, provided with petals. 21. Leaves herbaceous, 2— or 3—lobed, not truncate. 22. Plant finely hairy. 23. Flowers 1-2 per inflorescence; leaves 2-lobed; flowers 2-6 cm 9; fruit + fusiform, hexagonal, 5-6 cm; locally escaped in W. Java; 100-1000 m. ........ Se b capsolaris 5. 23. Flowers 4-8 per inflorescence; leaves 3—lobed. flowers 3-4 cm @; fruit globose, c. 1% cm © (not yet observed in Old World specimens); occasionally cultivated; 0-1000 m. P. holosericea L. 22. Plant glabrous; leaves 3-lobed. 24. Leaves variegated; lower surface with 2 large glands near the base of the midnerve; petiole without glands; flowers 3-4 cm g; fruit c. 2 cm 9, glaucous; ornamental; 0-500 m. P. trifasciata LEM. 24. Leaves not variegated, without glands at base of midrib; petiole with glands; flowers c. 2% cm @; fruit subglobose, purple, c. 2% cm long; occasionally cultivated; 0-1500 m. P. gracilis Jaca. 21. Leaves + coriaceous, 2-lobed or hemi-orbicular or obreniform, top truncate, lower surface with 2 rows of glands; flowers c. 3 cm 9g; fruit globose, 1-2 cm 9; locally cultivated and escaped; OF 1000.) Ay soie) ws eS ees SE stip.) co SEES See eats tate ernie ener ee ae P. biflora LAMK Section Decaloba DC. Mém. Soc. Phys. Genéve 1, 2 (1822) 435; Prod. 3 (1828) 325; G. Don, Gen. Syst. 3 (1834) 49. — Sect. Decaloba subsect. Eudecaloba Mast. Trans. Linn. Soc. 27 (1871) 632 (under subg. Plectostemma); HARMS in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 88; ibid. ed. 2, 21 (1925) 499. - Sect. Polyanthea DC. Mém. Soc. Phys. Genéve 1, 2 (1822) 435; Prod. 3 (1828) 322. — Sect. Decaloba subsect. Polyanthea (DC.) ENDL. Gen. PI. (1839) 926; MAsrT. Trans. Linn. Soc. 27 (1871) 630, 642 (under subg. Plectostemma); HARMS in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 88; ibid. ed. 2, 21 (1925) 498; CusseT, Fl. Camb., Laos & Vietn. 5 (1967) 107. Disemma LABILL. Sert. Austr. Caled. (1824) 78, t. 79; DC. Prod. 3 (1828) 332; G. Don, Gen. Syst. 3 (1834) 56; Spacu, Hist. Nat. Vég. Phan. (1838) 276; VAN Houtte, Hort. 2 (1847?) t. 11 (Distemma’); Roem. Syn. Mon. 2, Pepon. (1846) FLORA MALESIANA [ser. I, vol. 7? 131, 188; Mast. Trans. Linn. Soc. 27 (1871) 626, 630; Mia. Fl. Ind. Bat. 1, 1 (1855) 699; SEEM. FI. Vit. (1865) 96. Blephistelma Rar. Fl. Tellur. 4 (1836) 103. Anthactinia (non BORY) ROEM. Syn. Mon. 2, Pepon. (1846) 190. Sect. Octandranthus HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 506. — Sect. Decaloba subsect. Octodranthus (HARMS) CUSSET, Fl. Camb., Laos & Vietn. 5 (1967) 108. Sect. Anomopathanthus HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 506. — Sect. Decaloba subsect. Anomopathanthus (HARMS) CussET, Fl. Camb., Laos & Vietn. 5 (1967) 108. Sect. Hollrungiella HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 501. Distr. About 70 spp. in the tropical and subtropical Americas according to KILLIP, some 20 spp. in Indo-Australia and SW. Polynesia. Notes. I have refrained from drawing a sectional description, as this would have necessitated close examination of diagnostic characters of both the American and Asian species, which would necessarily lead too far in considering a re-evaluation of the infrageneric systematy of the genus, leading me beyond the scope of this regional revision. 1. Passiflora moluccana REINW. ex BL. Bijdr. (1826) 938. See for references and synonymy under the varieties. Climber to 6 m, sparsely to densely short-hairy. Leaves spirally arranged or distichous, rarely sub- opposite (in extra-Mal. mostly subopposite), en- tire (very rarely faintly 3-lobed), mostly (sub) coriaceous, glossy above, dull pale green, densely velutinous to (sub)glabrous beneath, lanceolate (in juvenile shoots) to elliptic or ovate, top acute to obtuse or rounded, sometimes retuse or up to 1 cm acuminate, base obtuse to (sub)cordate, 5-16 by (1-)1!4-10 cm, pinninerved; nerves 3-8 pairs; petiole 34-3 cm. Glands on lamina (in Mal.) 2-5 pairs, 4-2 mm 9g; petiolar glands 2, 1-3 mmg, always in the upper half of the petiole. Stipules c. 1 mm. Inflorescences sessile, (2—)4—10(—15)-flow- ered, with a central tendril, the two lateral partial inflorescences up to 1 cm stalked; pedicels 14-214 cm; bracts and bracteoles linear, acute, mostly hairy, 2-6 mm. Tendrils 5-20 (—25) cm. Flowers pubescent to subglabrous outside, 3-414 cm 9; stipe 5—20(-25) mm; hypanthium saucer-shaped, 2-3 by 6-9 mm. Sepals lanceolate(-triangular), subacute to obtuse, 10-20 by 4-6 mm. Petals lan- ceolate, obtuse, (sub)entire, 10-18 by 2-5 mm. Corona double, outer corona consisting of a rather dense row of filaments 10-18 mm and a somewhat lower inserted row of short filaments 1-3 mm, inner corona a rather stiff, densely plicate ‘collar’ 114-2 by 114-214 mm, with finely serrate-laciniate edge, curving inward towards a low fleshy rim-like disk 44—'4 mm high. Androgynophore 5-10 mm. Filaments + subulate, (5—)6—10 mm; anthers ellip- tic(-oblong), obtuse, 3—414(-5) by 114-2 mm. Ovary (sub) sessile, globose to ellipsoid, glabrous to densely hairy, 2-4 by 114-3 mm; styles 3, 5-8 mm; stigmas (14-)1 mm @. Fruits 1-3 per inflores- cence, globose to ellipsoid, 2-3 by 114-214 cm, glabrous or hairy; pericarp coriaceous, c. 1 mm @. Seeds c. 30-50, suborbicular to obovoid, 314-414 by 3-4 by 114 mm, 7-9 small pits per g. Distr. Indo-China, S. China, in Malesia: Phi- lippines (Luzon), Java, Lesser Sunda Is., S. Cele- bes, Moluccas. Fig. 2: la—b. Ecol. Prefers apparently a seasonal climate; 0-900 m. Notes. After comparison of the Malesian mate- rial of P. horsfieldii with a large amount of speci- mens of P. cochinchinensis from the Asian conti- nent and Hainan, there appeared to be only one differentiating character, viz the mostly suboppo- site leaves in the latter species; this is of course insufficient for distinction. Field notes. \n fresh flowers the sepals are green- ish white, the petals white, corona filaments yellowish with purple-brown base, inner corona purplish brown. Fresh fruits are greenish. KEY TO THE VARIETIES 1. Petiolar glands inserted at (2—)3-8 mm from the blade-base. Leaves + coriaceous or herba- ceous, velutinous to (sub)glabrous beneath. Ovary (sub)glabrous, rarely pubescent. Fruit (sub)glabrous. . : a. var. moluccana 1. Petiolar glands at the apex of the petiole at 0-3 mm from the blade. Leaves mostly coria- ceous, velutinous beneath. Ovary densely pubes- cent. Fruit thinly hairy. b. var. teysmanniana a. var. moluccana. — P. moluccana REINW. ex BL. Bijdr. (1826) 938; Rumphia 1 (1837) 169, t. 51; DC. Prod. 3 (1828) 323; G. Don, Gen. Syst. 3 (1834) 47; WALP. Rep. 2 (1843) 221; Mast. Trans. Linn. Soc. 27 (1871) 631; BRITTEN in Forbes, Wand., App. 6 (1885) 506; HARMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 88; ibid. ed. 2, 21 (1925) 498; Merk. Philip. J. Sc. 11 (1916) Bot. 294; HALL. f. Med. Rijksherb. 42 (1922) 6. — Anthactinia moluccana ROEM. Syn. Mon. 2, Pepon. (1846) 190. PASSIFLORACEAE (de Wilde) 413 1a A Passiflora moluccana var. moluccana 1b © Passiflora moluccana var. teysmanniana 2 @ Passiflora perakensis 3 * Passiflora sumatrana 5 @ Passiflora hollrungii Fig. 2. Distribution of various Passiflora species, extra-Malesian localities of P. moluccana var. teysman- niana omitted. — Disemma moluccana Mia. Fl. Ind. Bat. 1, 1 (1855) 699. P. moluccana var. timorensis BL. Rumphia 1 (1837) 169, t. 51 A; Mast. Trans. Linn. Soc. 27 (1871) 631. — P. timoriana SPAN. Prod. Fl. Timor., Icon. ined., Linnaea 15 (1841) 207, Icon. ined. t. 76, nom. nud. — Anthactinia timorensis ROEM. Syn. Mon. 2, Pepon. (1846) 191. — Disemma timoriana Miq. Fl. Ind. Bat. 1, 1 (1855) 700; BRITTEN in Forbes, Wand. etc., App. 6 (1885) 506. Leaves spirally arranged, ovate to oblong, apex obtuse to acute-acuminate, subcoriaceous to her- baceous (membranous when dry), pale green or -+ glaucous, glabrous or subglabrous, or hairy only near the nerves beneath. Glands on the petiole at 3-8 mm from the blade. Petioles 144-3 cm. Bracts and bracteoles, flower stipe, hypanthium and outer side of sepals more or less hairy to (sub)glabrous. Flower stipe 5-11 mm. Anthers 3-4 mm. Ovary (sub)glabrous, sometimes hairy. Fruits (sub)-glabrous. Distr. Malesia: Lesser Sunda Is. (W. & E. Timor), Moluccas (Buru, Ceram, Ambon, Haruku 1a) - big. 2: 1a: Ecol. Fl. mostly July(-Noyv.). b. var. teysmanniana (MIQ.) DE WILDE, comb. nov. P. pallida (non L.) Lour. Fl. Coch. 2 (1790) 527. — P. cochinchinensis SPRENG. Syst. Veg. 4, Curae post. (1827) 346. P. horsfieldii BL. Rumphia 1 (1837) 170, t. 52, 1-4; WaLpP. Rep. 2 (1843) 221; TEysm. Nat. Tijd. N. I. 11 (1856) 178; Mast. Trans. Linn. Soc. 27 (1871) 631; HARMS in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 88; ibid. ed. 2, 21 (1925) 498; HALL. f. Med. Rijksherb. 42 (1922) 6; Back. & Baku. f. FI. Java 1 (1963) 291; CusseT, Adansonia 7 (1967) 375, 381. — Anthactinia horsfieldii ROEM. Syn. Mon. 2, Pepon. (1846) 191. Disemma horsfieldii Mig. Fl. Ind. Bat. 1, 1 (1855) 700. — Disemma horsfieldii var. teysman- niana Mia. l.c. P. horsfieldii var. elbertiana HALL. f. Med. Rijks- herb. 42 (1922) 6. P. ligulifolia Mast. Trans. Linn. Soc. 27 (1871) 632. P. hainanensis HANCE, J. Bot. 16 (1878) 227. P. philippinensis ELM. Leaf]. Philip. Bot. 1 (1908) 326; MerR. En. Philip. 3 (1923) 118. Leaves spirally arranged, distichous, sometimes partly subopposite, coriaceous, ovate or elliptical to lanceolate, apex mostly obtuse, often retuse, dark green above, pale green to glaucous green, velutinous beneath. Glands on the petiole at 0-3 mm from the blade, rarely partially on the blade. Petiole 14—-2(—2!4) cm. Bracts and bracteoles, and flower stipe, hypanthium and outer side of sepals more or less densely hairy. Flower stipe 7-20 mm. Anthers 3-5 mm. Ovary densely hairy. Fruits thinly hairy. Distr. Indo-China, S. China (Hainan), in Malesia: Java (W. & E.; Madura I.), Lesser Sunda Is. (Bali, Sumbawa, Flores), SE. Celebes (Kabaéna I.), Philippines (Luzon). Fig. 2: 1b. Ecol. Stony slopes, scrub, open forest, teak forest, 0-900 m. F/. March—June, fr. Sept._Nov. 414 Vern. Java: rambuset, Md., prabuset, p. moto, J; Luzon: quachal, Ig. Note. In one of the two known collections from the Philippines (RAMos & EDANO BS 38098) the basal blade glands are situated for a large part on the blade. 2. Passiflora perakensis HALL. f. Med. Rijksherb. 42 (1922) 5; CusseT, Adansonia 7 (1967) 375, 381. — Fig. le. P. horsfieldii (non BL.) Kina, J. As. Soc. Beng. 71, ii (1902) 50; RipL. Fl. Mal. Pen. 1 (1922) 839; Burk. & HEND. Gard. Bull. S. S. 3 (1925) 378. P. horsfieldii var. distans CRAIB, Fl. Siam. En. 1 (1931) 743. Climber 2—-10(-15) m; sparsely, rather stiff- hairy on stem and inflorescential branches, often glabrescent. Leaves spirally arranged, rarely sub- opposite, herbaceous to thinly coriaceous, above (sub)glabrous, green, beneath (sub)glabrous to sparsely hairy, ovate-elliptic to oblong, top acute, sometimes 1 cm acuminate, (0O-)1—-3 mm mucro- nate, base rounded or shallowly cordate, 5-18 by 214-9 cm, pinninerved, nerves 4-8 pairs, reticula- tion distinct; petiole 1-4(-5) cm. Glands on the lamina in 2—7(-8) pairs, in two rows about halfway margin and midrib, 14-114 mm g; petiolar glands 2, opposite, 1-3 mm g, + halfway to 14 from the base of the petiole. Stipules minute, linear, c. 1 mm. Inflorescences sessile, 4-20-flowered, with a cen- tral tendril, the two lateral main branches up to 15 mm stalked; pedicels 1—-2(—3) cm; bracts and bracteoles linear, acute, with finely hairy margin, 1-2'4 mm. Tendrils 8-20 cm. Flowers thinly hairy to glabrescent, 2—3(-314) cm 9; stipe 5—15(—22) mm; hypanthium saucer-shaped, c. 2 by 7-10 mm. Sepals lanceolate, obtuse, 10-15(—20) by (3—)4—5 mm. Petals lanceolate, obtuse, 8-15 by 3-4 mm. Corona double; outer corona filaments 6-10 mm, often strongly sinuate, inner filaments (1—)2—3 mm; inner corona 2-3 by 2—3 mm, as a densely plicate ‘collar’, with finely serrate edge, -+_ curving in- wards over the c. 1 mm high disk. Androgyno- phore 5—6(—7) mm. Filaments connate into a + inflated or cup-shaped tube entirely enveloping the ovary, (3—)5-10(—11) by 2-3 mm, free part of filaments 3-5 mm; anthers oblong, obtuse, 214-4 (—5) by 1-114 mm. Ovary on gynophore up to 114 mm, ellipsoid to obovate, hirsute, 2—4(-5) by 114-2(-2'4) mm; styles 3, free or up to 3 mm con- nate; style arms 3-7 mm; stigmas 1-114 mm @. Fruits 2-7 per inflorescence, globose to ellipsoid, (1144-)2-244 by 114-2 cm, thinly hispid; pericarp coriaceous, c. 1 mm @. Seeds c. 40-70, obovate, c. 4-414 by 314-4 by 114 mm, 6-9 pits per go. Distr. S. Thailand, in Malesia: Sumatra (East Coast Res.), Malay Peninsula (Kedah, Perak, Pahang), and North Borneo (Keningau Distr. SAN 65383). Fig. 2: 2. Ecol. Rain-forests, 0-1000 m. Fl. Sept.—April, fr. mainly May. Notes. Closely related to P. siamica CRAIB (P. octandra GAGN.), which has also partly connate filaments, but in which the leaves are distinctly FLORA MALESIANA [ser. I, vo.1 7? hairy also on the upper surface and which has (5—)6—8 stamens. Field notes. Leaves bluish green or glaucous underneath. Sepals greenish, petals white, corona filaments white or greenish yellow, + mottled at base, inner corona purplish; ovary purplish or pale green, androgynophore pale green, filaments and styles and stigmas green, anthers yellow. Fruits (bluish-)green, pericarp fleshy. Seeds black- ish, arils juicy, + colourless, of a flat taste. 3. Passiflora sumatrana BL. Rumphia 1 (1837) 170; Mast. Trans. Linn. Soc. 27 (1871) 631. — Anthac- tinia sumatrana ROEM. Syn. Mon. 2, Pepon. (1846) 191. — Disemma sumatrana Mia. FI. Ind. Bat. 1, 1 (1855) 700. Climber 5-15 (?) m, glabrous to glabrescent. Leaves spirally arranged, thinly coriaceous, gla- brous, ovate-elliptic, mostly -- semi-orbicular, apex truncate to 3-lobed, mostly 1-2 mm mucro- nate, rarely acutish, base rounded to broadly cordate, 6-12 by 5-13 cm, sub—5S—plinerved and with 1-2 pairs of smaller nerves higher up, ascend- ing or straight, the upper two main nerves ending in the lobes, reticulation distinct; petiole 2-614 cm. Glands on lamina absent or 1—2, small, 44-1 mm g, submarginal in the sinusses between the lobes; petiolar glands 2, 1-2 mm g, at 1/5—1/3 from the base. Stipules linear, 14(-1) mm. Inflorescences sessile with a central tendril, the two lateral main branches up to 8 mm stalked, 2-14-flowered; pedicels 5-15 mm; bracts and bracteoles linear, acute, 14-114 mm. Tendrils 10-20 cm. Flowers glabrous, 3-4 cm g; stipe c. 8-12 mm; hypanthium saucer-shaped, c. 214 by 7 mm. Sepals lanceolate, 13-15 by 4-6 mm, with a minute subapical horn or wart c. 14 mm. Petals lanceolate, 10-13 by 3 mm. Corona double, outer corona filaments 5-8 mm, inner filaments 2-3 mm; inner corona 2 by 2-214 mm, consisting of a densely plicate, + in- ward curving ‘collar’ with subentire edge; disk c. Y4 mm. Androgynophore 7-8 mm. Filaments free, c. 7 mm; anthers oblong, obtuse, c. 4 by 114 mm. Ovary sessile, ellipsoid, glabrous, c. 4 by 114 mm; styles free, c. 7 mm; stigmas 1-144 mm g. Fruits c. 1-4 per inflorescence, subglobose to ellip- soid, (2-)214 by 2 cm, glabrous; pericarp coria- ceous, 14-1 mm 9. Seeds c. 60, obovate, c. 414 by 3 by 114 mm, 6-8 pits per @. Distr. Malesia: West Central Sumatra (Tapa- nuli, West Coast Res.). Fig. 2: 3. Ecol. Forests, 1600-1800 m. Notes. The species is closely related to P. assa- mica CHAKRAVARTY, P. burmanica CHAKRAVARTY, P. wilsoni HEMSLEY, P. jugorum W. W. SMITH and P. celata Cusset from continental SE. Asia. Field notes. According to an unpublished water- colour drawing of a KorTHALs specimen (in L) the sepals are pale greenish, the petals white, an- drogynophore, filaments and styles greenish, an- thers yellow, ovary dirty-green, corona filaments pale yellowish with purplish base, inner corona purplish. 1972] 4. Passiflora aurantia Forst. f. Fl. Ins. Austr. Prod. (1786) 621. a. var. aurantia. — P. aurantia Forst. f. Fl. Ins. Austr. Prod. (1786) 621 (n. 336); CAv. Diss. 10 (1790) 457; WILLD. Sp. Pl. 3, 1 (1800) 620; SPRENG. Syst. Veg. 4, Curae post. (1827) 250; ANDREws, Bot. Rep. 5 (1803) t. 295; Masr. Trans. Linn. Soc. 27 (1871) 634; F. v. M. Fragm. Phyt. Austr. 9 (1875) 68; K. Scu. Bot. Jahrb. 9 (1888) 211; K. Scu. & HOoLir. FI. Kais. Wilh. Land. (1889) 82; K. Scu. & Laut. Fl. Schutzgeb. (1901) 456; WARB. Bot. Jahrb. 13 (1891) 384; HARMs in E. & P. Nat. Pfi. Fam. 3, 6a (1893) 89; ibid. ed. 2, 21 (1925) 500; Bot. Jahrb. 15 (1893) 581 (anat.); F. M. BAILeEy, Queens]. Fl. 2 (1900) 689; HECKEL, Ann. Mus. Col. Marseille II, 10 (1912) 272, t. 30; GUILLAUM. J. Arn. Arb. 12 (1931) 262; FI. Nouv. Caléd. (1948) 224; Ann. Mus. Col. Marseille 55/56 (1948) 37; Mém. Mus. Hist. Nat. Paris. sér. B, Bot. 8 (1959) 148; ibid. 8 (1962) 270; RECHINGER, Denk- schr. Wien. Akad. Wiss. 85 (1910) 314; LLoyp & AIKEN, Bull. Lloyd Libr. n. 33 (1934) 75; BEAL, Austr. Pl. 6 (1970) 13, photogr. — Murucuja aurantia PERS. Syn. Pl. 2 (1807) 222; SpRENG. Syst. Veg. 3 (1826) 43. — Disemma aurantia LABILL. Sert. Austr. Caled. (1824) 78, t. 79; DC. Prod. 3 (1828) 332; G. Don, Gen. Syst. 3 (1834) 56; SPACH, Hist. Nat. Vég. Phan. (1838) 276; Hook. in Curtis, Bot. Mag. (1845) t. 4140; Roem. Syn. Mon. 2, Pepon. (1846) 188. — Blephistelma aurantia RAF. Fl. Tellur. 4 (1836) 103. — Distem- ma aurantiacum LEMAIRE, I11. Hortic. 14 (1867) Misc. 57. — Fig. If. P. glabra WENDL. Coll. Plant. 1 (1805) 55, t. 17, non MILL. 1768; Mast. Trans. Linn. Soc. 27 (1871) 634; F. v. M. Fragm. Phyt. Austr. 9 (1875) 69. — P. adiantum WILLD. En. Hort. Berol. 2 (1809) 698; SPRENG. Syst. Veg. 3 (1826) 42. — P. adiantifolia KER-GAWL, Bot. Reg. 3 (1817) t. 233; LAWRENCE, Passionfi. (1802) t. 11; HARMS in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 89; Bot. Jahrb. 15 (1893) 581 (anat.). — Murucuja adiantifolia Sweet, Hort. Brit. ed. 1, pt 2 (1826) 355. — Disemma adianthi- folia DC. Prod. 3 (1828) 333; G. Don, Gen. Syst. 3 (1834) 56; Spacu, Hist. Nat. Vég. Phan. (1838) 277; Roem. Syn. Mon. 2, Pepon. (1846) 188; LEMAIRE, I11. Hortic. 14 (1867) Misc. 57 (‘Dis- temma adiantifolium ). Murucuja baueri LinpL. Coll. Bot. (1821) t. 36. — Disemma baueriana ENDL. Prod. Fl. Norfolk (1833) 66; G. Don, Gen. Syst. 3 (1834) 56 (‘bau- eri’); WALP. Rep. 2 (1843) 221; Roem. Syn. Mon. 2, Pepon. (1846) 188-189; Lemarrg, I11. Hortic. 14 (1867) Misc. 57. — P. baueriana MAST. Trans. Linn. Soc. 27 (1871) 634; Harms, Bot. Jahrb. 15 (1893) 581 (anat.). Disemma coccinea DC. Prod. 3 (1828) 333, non P. coccinea AuUBL. 1775; G. Don, Gen. Syst. 3 (1834) 56; Spacu, Hist. Nat. Vég. Phan. (1838) 277; LEMAIRE, 111. Hortic. 14 (1867) Misc. 57. — P. banksii BENTH. Fl. Austr. 3 (1866) 312; MAsT. Trans. Linn. Soc. 27 (1871) 634; HARMS in E. & P. Nat. Pfil. Fam. 3, 6a (1893) 89; ibid. ed. 2, 21 (1925) 483, 500; Bot. Jahrb. 15 (1893) 581 (anat.); PASSIFLORACEAE (de Wilde) 415 BRITTEN, I11. Austr. Pl. etc. 2 (1901) 42, t. 129; Domin, Bibl. Bot. Heft. 89 (1928) 986-987. — P. aurantia var. banksii F. M. BAILEY, Queensl. Agric. J. 26 (1911) 315; ibid. 27 (1911) 66; Compr. Cat. Queens]. (1913) 220, f. 191. Disemma brachystephanea F. vy. M. Fragm. Phyt. Austr. 1 (1858) 56. — P. brachystephanea BENTH. FI. Austr. 3 (1866) 312; MaAsr. Trans. Linn. Soc. 27 (1871) 634; F. v. M. First Syst. Census (1882) 76; Second Syst. Census (1889) 128; F. M. BAILEy, Syn. Queensl. Fl. (1883) 199; Cat. Pl. Queens]. (1890) 20; Queens]. Fl. 2 (1900) 689; Compr. Cat. Queensl. (1913) 220; Moore, Handb. Fl. N.S. W. (1893) 254. — P. banksii var. brachystephanea Doin, Bibl. Bot. Heft. 89 (1928) 987. Disemma_ caerulescens SEEM. (1862) 366. Disemma barclayi SEEM. Fl. Vit. (1865) 96. — P. barclayi Mast. Trans. Linn. Soc. 27 (1871) 634; DRAKE DEL CASTILLO, I11. Fl. Ins. Mar. Pacif. (1890) 175; Harms in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 89; PARHAM, PI. Fiji Is. (1964) 111. Disemma storckii SEEM. Fl. Vit. (1865) 96. — P. storkii DRAKE DEL CASTILLO, J11. Fl. Ins. Mar. Pacif. (1890) 175; PARHAM, PI. Fiji Is. (1964) 111. Disemma vitiensis SEEM. Fl. Vit. (1865) 96. — P. yitiensis Mast. Trans. Linn. Soc. 27 (1871) 634; DRAKE DEL CASTILLO, I11. Fl. Ins. Mar. Pacif. (1890) 175; Harms in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 89; GiBpss, J. Linn. Soc. Bot. 39 (1909) 148, excl. specim. Samoensis; TURRILL, J. Linn. Soc. Bot. 43 (1915) 23; PARHAM, PI. Fiji Is. (1964) 2 P. samoénsis (non EXELL) YUNCKER, Bern. P. Bish. Mus. Bull. 220 (1959) 193. Climber to 6 m, glabrous or glabrescent, rarely finely pubescent (distinctly pubescent in var. pubes- cens). Serial buds sometimes developed into mi- nute shoots. Leaves spirally arranged, herbaceous, rarely coriaceous, broadly ovate to suborbicular in outline, 3(—5, especially in some New Cale- donian specimens)—lobed in the upper part, some- times 3(—5)—dissected, rarely entire, top acute or mostly obtuse or rounded, sometimes retuse, often up to 1 mm mucronate, base subacute to shortly cordate, (144-)214-10 by (2-)4-13 cm, 3(-5)- plinerved with 2-6 pairs of smaller nerves higher up, main nerves straight, ending in the lobe tips; reticulation distinct; lobes 14-5 cm, narrow to broad; petiole 1-4 cm. Glands on lamina 44-4 mm 9, 2-35 scattered between the main nerves; petiolar glands 2 (outside Mal. rarely 0 and absent in var. pubescens), flat to + crateriform, 1,-2 mm 9, in upper half of petiole, usually close to the blade, rarely (var. samoénsis) in the lower half. Stipules linear, c. 144 mm. Inflorescences sessile, 1(—2?)- flowered; pedicels 14—2 cm, bracteoles (2—)3, linear acute, 1-3 mm. Tendrils 7-20 cm. Flowers gla- brous, 414-8(-10) cm g; stipe 144-5(-7) mm; hypanthium saucer-shaped, 3-4 by (8—)10-17 mm. Sepals lanceolate, (sub)acute, keeled, (2-)21,-414 by 14-2 cm. Petals oblong to lanceolate, acute to obtuse, (0O—-)S-15 by 3-5 mm. Corona double, Bonplandia 10 416 outer corona filaments slender, rather spaced, (5—)8—-12(—18) mm (+ as long as the inner corona; inner corona tubular, membranous, 7—15(—20) mm long, gradually narrowed and + wrinkled to- wards the 7-12 mm wide throat, with a shallowly lobed undulate paler edge; disk 0. Androgyno- phore 20-35 mm. Filaments 7-11 mm, free or up to 2 mm connate at base; anthers lanceolate, obtuse, 5-10 by 114-3 mm. Ovary stalked, 1-214 mm, (ob)ovate-ellipsoid, glabrous or glabrescent, 214— 6 by 114-3(—314) mm; styles free, (5—)7-10 mm; stigmas (1-)114-3 mm @. Fruit subglobose to ellipsoid, 214-5 by 2'4-4'4 cm; pericarp coria- ceous, c. '4 mm @ thick. Seeds c. 100-200, obovate, 214-3 by 2-214 by 1-114 mm, 4-6 pits per 9; embryo c. 114-2 mm, cotyledons ellipsoid, broadly obtuse, c. 1144 by 1 mm. Distr. Australia, SW. Pacific, and Malesia: East New Guinea, Thursday I., Louisiades, Queens- land, Lord Howe I., Norfolk I., New Caledonia, New Hebrides, Fiji, Tonga, Niue I. Fig. 3: 4a. . ‘a . ©. PC e. Ss 42a @ Passiflora aurantia var aurantia 4b a Passiflora aurantia var Samoensis 4c APassiflora aurantia var pubescens %& Passiflora cinnabarina @Passiflora herbertiana Fig. 3. Distribution of various Passiflora species. Ecol. (Coastal) scrub, monsoon scrub, (rain-) forest edges and regrowths, forest clearings, lime- stone cliffs, on sandy and loamy soils, 0-500(— 1250) m. Fi. Jan._Dec. (New Guinea), ditto, but mainly May—June & Oct. (Australia); var. samoén- sis fl. May—Aug. Notes. Some specimens from New Caledonia with (-+ deviating) 5—palmate(-lobed) and rather coriaceous leaves, and with rather small flowers, are included in the type variety. Field notes. Fresh flowers are described as follows. Sepals, corona, androgynophore: orange, salmon, pink, flesh-coloured or pale red, turning deep (bright) red with age; petals: whitish, creamy, to reddish; corona-filaments deep red; fresh fruits are reported as (pale) greenish. KEY TO THE VARIETIES 1. Petiole provided with glands. 2. Glands in the upper half of the petiole; lami- nal glands not (only) approximate to the nerve bases. . Sie >, eel esha ear aurantia 2. Glands in the lower 14 of the petiole; laminal glands rather approximate to the nerve bases. Samoa. Be, b. var. samoénsis FLORA MALESIANA [ser. I, vol. 7? 1. Petiole without glands. 3. Plant glabrous; sometimes with minute petio- lar glands. Norfolk I. a. var. aurantia 3. Plant (incl. ovary) finely pubescent. Australia. c. var. pubescens b. var. samoénsis (EXELL) DE WILDE, comb. nov. — P. samoénsis EXELL, J. Bot. 63 (1925) 203; Curis- TOPHERSEN, Bern. P. Bish. Mus. Bull. 128 (1935) 153. — Fig. 3: 4b. c. var. pubescens F. M. BAILEY, Queensl. Agric. J. 26 (1911) 315, t. 31 f. 2; Compr. Cat. Queensl. Pl. (1913) 220, f. 191—bis. — P. baileyana Domin, Bibl. Bot. Heft 89 (1928) 433. — Fig. 3: 4c. Passiflora cinnabarina LINDL. Gard. Chron. (1855) 724; OLIv. in Curtis, Bot. Mag. 97 (1871) t. 5911; Mast. Trans. Linn. Soc. 27 (1871) 634; F. v. M. Fragm. Phyt. Austr. 9 (1875) 68; LINDL. J. R. Hort. Soc. Lond. n.s. 4 (1877) 134; Domin, Bibl. Bot. Heft 89 (1928) 433; YouNG, Rec. Auckl. Inst. Mus. 7 (1970) 148, f. 9-11. ? Disemma muelleriana REGEL, Index Sem. Hort. Petrop. (1866) 101. —? P. muelleriana MAst. J. R. Hort. Soc. Lond. n.s. 4 (1877) 133. ? Distemma eglandulosum LEMAIRE, I11. Hortic. 14 (1867) Misc. 56. Distr. Australia (Victoria). Fig. 3. Passiflora herbertiana KER-GAWL, Bot. Reg. 9 (1823) t. 737; SPRENG. Syst. Veg. 3 (1826) 41; LoppicEs, Bot. Cab. 14 (1828) t. 1364; BENTH. FI. Austr. 3 (1866) 311; Mast. Trans. Linn. Soc. 27 (1871) 634; F. v. M. Fragm. Phyt. Austr. 9 (1875) 68; Harms, Bot. Jahrb. 15 (1893) 581 (anat.); in E. & P. Nat. Pfil. Fam. 3, 6a (1893) 89; F. M. BAILEY, Queensl. Fl. 2 (1900) 688; Compr. Cat. Queensl. Pl. (1913) 220. — Murucuia herbertiana SweEET, Hort. Brit. ed. 1, pt 2 (1826) 355. — Di- semma herbertiana DC. Prod. 3 (1828) 332; G. Don, Gen. Syst. 3 (1834) 56; DECNE, Herb. Timor. Descr. (1835) 123 (= Nouv. Ann. Mus. 3: 451); SPACH, Hist. Nat. Vég. Phan. (1838) 276; SPAN. Linnaea 15 (1841) 207; Roem. Syn. Mon. 2, Pepon. (1846) 189; Mra. FI. Ind. Bat. 1, 1 (1855) 701; LEMAIRE, I11. Hortic. 14 (1867) 57; BRITTEN in Forbes, Wand., App. 6 (1885) 506. P. verruculosa WEINMANN, Syll. Plant. Ratisb. 1 (1824) 228; SteuD. Nom. ed. 2, 2 (1841) 276 (‘verrucosula’); ROEM. Syn. Mon. 2, Pepon. (1846) 177; Mast. Trans. Linn. Soc. 27 (1871) 639. P. biglandulosa CALEY, in Hb. Lambert, nom. nud. — Disemma herbertiana var. caleyana DC. Prod. 3 (1828) 332; G. Don, Gen. Syst. 3 (1834) 56. — Disemma caleyana Roem. Syn. Mon. 2, Pepon. (1846) 189. — P. herbertiana var. caleyana MAST. Trans. Linn. Soc. 27 (1871) 634. P. distephana F. v. M. ex HArMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 89, nomen. Distr. Australia (Queensland and New South Wales). Fig. 3. Note. The record of this species from Timor by DECAISNE, SPANOGHE, MIQUEL, and BRITTEN, /l.cc. is an erroneous localisation of an Australian col- lection, as Dr. H. Hetne kindly informed me that 1972] in the Decaisne Herbarium at Paris there are 3 identical specimens, 2 apparently wrongly labelled with the provenance ‘Timor’, the 3rd one marked to come from ‘N. holl. port jackson’. 5. Passiflora hollrungii K. Scu. Bot. Jahrb. 9 (1888) 211; K. Scu. & Hoxtr. FI. Kais. Wilh. Land (1889) 82; Harms, Bot. Jahrb. 15 (1893) 581; K. Sco. & Laur. FI. Schutzgeb. (1901) 456; HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 501; JERMY & SAYERS, J. R. Hort. Soc. 92, 3 (1967) 121. — Fig. la-—d. Climber to c. 10 m, sparsely to densely greyish pubescent. Leaves spirally arranged, herbaceous, hairy especially on and near the nerves beneath, broadly ovate to elliptic(-oblong), entire, top acute or up to 1 cm acuminate, up to 2 mm mucronate, base cordate to rounded, up to 2 mm peltate, 5-14 by 3-914 cm, 3(—5)—-plinerved, the nerves arching towards the apex, larger veins often -+ trabeculate; petiole 2-4 cm. Glands on lamina 1—5 submarginal on either side, 14—14(-1) mm 9, petiolar glands 0. Stipules linear, c. % mm. Inflorescences sessile, 1—2-flowered, either with a tendril or not; pedicels 34-114 cm; bracteoles 3, linear, 1!,-4 mm, rather near the articulation. Tendrils 6-25 cm. Flowers (sub)glabrous; stipe 5-12 mm; hypanthium bowl- shaped, 3-4 by 9-12 mm. Sepals lanceolate, sub- acute to obtuse, + hairy at the top, 4-5 by 4-34 (- 1) cm. Petals lanceolate, acute, 314-444 cm by 4-6 mm, membranous, glabrous. Corona double, outer corona erect, tubular to + skirt-shaped, PASSIFLORACEAE (de Wilde) 417 + fleshy, 2—3 cm, with up to 14 mm deep irregular- ly undulate-lobed edge; inner corona 1—2 mm high, curved inward, stiff, up to halfway laciniate into acute teeth; both coronas inserted on the rim of the hypanthium; disk fleshy, annular sharp-edged, c. 1 mm high near the base of the androgynophore, connected by 5 low septa to the hypanthium. Androgynophore 214-4 cm. Filaments 7-12 mm; anthers lanceolate, bluntly c. 14 mm apiculate, shortly sagittate at base, 8-9 by 2-3 mm. Ovary 2 mm stalked, densely pubescent, obovate-ellip- soid(-oblong), 5—7 by 3-4 mm; styles free, 5-714 mm; stigmas c. 2 mm @. Fruit subglobose to ellip- soid, sometimes + fusiform, excluding stipe 214— 314 by 2-214 cm; pericarp coriaceous, c. 1 mm g, sparsely to densely pubescent. Seeds c. 100-200, obovate-oblong, 2-214 by 1-114 by 2/3 mm, 3-5 pits per 9; embryo 114-2 mm, cotyledons obovate, broadly obtuse, c. 1 by 2/3 mm. Distr. Malesia: East New Guinea. Fig. 2: 5. Ecol. Secondary forest, scrub, 1000-1700 m. Fl. fr. mainly Oct., once in May and July. Vern. Gamoogaka mamerga, Finisterre Mts. Notes. In a single leaf I have found one minute petiolar gland at 2 mm below the blade. Field notes. Sepals and petals of fresh specimens are reported as lavender or pale bluish green or greyish purple, the corona (tube) as purple-blue to deep purple or blackish; filaments, styles and stigmas (pale) green. Ripe fruit green with a fine white pubescence. 2. ADENIA Forsk. Fl. Aegypt.-Arab. (1775) 77; ENGL. Bot. Jahrb. 14 (1891) 374; HARMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 83; ibid. ed. 2, 21 (1925) 488; HALL. f/ Med. Rijksherb. 42 (1922) 8; CHAKRAVARTY, Bull. Bot. Soc. Beng. 3 (1951) 68; HUTCH. Gen. Fl. Pl. 2 (1967) 373; DE WILDE, Thesis, Med. Landb. Hogeschool Wagen. 71-18 (1971) 1-281. — Modecca [RHEEDE, Hort. Mal. 8 (1688) t. 20-23] LAMK, Encycl. Méth. Bot. 4 (1797) 208; DC. Prod. 3 (1828) 336; BENTH. & HooK. f. Gen. Pl. 1 (1867) 813. — Microblepharis (W. & A.) RoEM. Syn. Mon. 2, Pepon. (1846) 133, 200. — Erythrocarpus Roem. I.c. 133, 204. Unarmed climbers (in Mal.), often with tubers. Leaves (in Mal.) simple, entire or lobed, pinni- or palminerved; apex of petiole or blade-base with 1-2 glands, sessile or on auricles. Stipules (in Mal.) minute. Inflorescences mostly stalked, few- to many-flowered, often with 1(—3) tendrils, rarely collected into raceme-like short-shoots. Bracts small. Flowers unisexual (plants mostly dioecious), in ¢ with vestigial ovary, in 9 the stamens reduced to + subulate staminodes (rarely bisexual in Afr.), mostly greenish to yellowish; hypanthium saucer- to cup-shaped, or tubi- form, sometimes S—saccate. Sepals (4-)5(-6), free or partially connate into a calyx tube. Petals (4-)5(-6), free or partially connate with the calyx tube, greenish, creamy, or white, mostly smaller than the sepals. Corona mostly a simple laciniate membrane or composed of hairs, sometimes fleshy, situated at the transition of hypanthium and sepals (or calyx tube), or absent. Disk mostly composed of 5 strap- shaped or clavate often outward curved appendages, opposite the sepals, inserted near mostly the bottom of the hypanthium. Androgynophore 0, rarely short. 418 FLORA MALESIANA [ser. Lo wale? Stamens 5, (in Mal.) inserted at the base of the hypanthium, free or partially connate into a filamental tube, the tube often connected with the hypanthium by septa opposite the petals; anthers narrow, mostly acute or acuminate, erect, basi- fixed. Ovary subsessile, globose to fusiform; styles 3(—5), free or connate, distinct or not; stigmas finely lobed to papillate or ramified. Capsule 3—valved, (in Mal.) coriaceous or woody. Distr. About 92 spp. in 6 sections in tropical and subtropical Africa (c. 60 spp.), Madagascar (c. 20 spp.) and SE. Asia, Malesia, and N. & NE. Australia, in the Pacific east to the Solomon Is. (A. heterophyl- la). Especially in Africa and Madagascar quite a number of local endemics. Ecol. The genus occurs in a wide range of habitats, varying from rain-forest to savannahs and almost desert conditions where most species have more os less succulent stems. In Africa and Madagascar a number of species have striking, swollen main stems or tubers; some are thorny or spiny. In Malesia up to 1500(—2000) m, in everwet forest, except A. heterophylla which prefers seasonal climat- ic conditions. Malesian species are medium-sized to large lianas, but A. penangiana is usually a small climber of but a few metres. Morph. & Taxon. In most species the filaments are in the basal part united into a (short) tube, where- as this tube is connected by septa with the hypanthium, opposite the petals. In some members of sect. Microblepharis and sect. Blepharanthes the thus formed apartments are + bulging, forming a 5-saccate hypanthium in which the 5 disk glands are situated. As pointed out in my monograph (DE WILDE, Thesis 1971, 27-36), 6 sections are recognized, mainly based on the flower structure, and sustained by characters found in the position of the glands on the leaves. In many species the sepals (and often also the petals) are partially connate into a tube; and this condition is considered as an advanced state against free sepals. The Indo-Malesian species belong to 3 sections: (i) sect. Microblepharis (Africa, Madagascar, SE. Asia; A. penangiana in Malesia), (ii) sect. Blepharanthes (Africa, SE. Asia; not in Malesia), and (iii) sect. Erythrocarpus (most Malesian spp.). Sect. Microblepharis, with free sepals and petals, is considered as primitive; sect. Erythrocarpus as relatively derived because of the tubiform flowers with partially connate sepals and petals, the absence of the corona, and the basal blade-glands situated on auricles at the top of the petiole. As to the leaf glands 3 types are distinguished: (i) glands at blade-base 2, or by connation (reduction) 1, sessile at the very blade-base, or on auricles at the transition of blade and petiole or at the top of the pe- tiole; (ii) blade-glands on the lower surface of the blade scattered or in + fixed places; in lobed leaves of- ten corresponding with the sinusses between the lobes; (iii) marginal glands, mostly very small, at the end of a small vein, in some species on teeth. In all Indo-Malesian species the fruits are red, and the funicles of the seeds are longer than in all other African or Madagascan species. Heteroblasty and heterophylly. As in many lianas the leaves of most Adenias are very variable, and in many species lobed as well as entire leaves are found. Also in juvenile forms the leaves may be quite dif- ferent in shape and in presence or position of the glands, as compared with the adult stage. In adult A. cordifolia, for instance, the basal glands of the cordate, ovate leaves are situated in two distinct, separate, hollowed auricles lateral at the top of the petiole, whereas in the juvenile stage these auricles are absent, and the blade of a peculiar lunate- or 3-lobed shape with mostly peltate base, with or without 1-2 very small glands. KEY TO THE SPECIES 1. Sepals free (flowers + campanulate). Corona present. Leaves with 1-2 mm wide peltate base; basal glands 2, free or contiguous. Sect. Microblepharis. . . . . . 1. A. penangiana 1. Sepals largely connate (flowers -+ tubiform). Corona absent. Leaves. not peltate: basal glands 2, on 2 auricles at the transition of petiole and blade. Sect. Erythrocarpus'. 2: Calyx lobes (1-)144-3 mm, reflexed in anthesis. Leaves entire or lobed, suborbicular to lanceolate in outline, with sails to acute base, 5-25 cm long. Gland-bearing auricles shallowly concave, more or less adnate to the blade, sometimes + peltately connate. 3. Stipe of 3 flowers 4-15 mm; ¢ flowers including stipe (10—)15-25(—30) by 114-5(-7 4) mm. Flower buds oval to obovate. Fruits ellipsoid to oblong; dry pericarp coriaceous. Leaves entire or lobed. 2. A. heterophylla (1) Two species of this section, viz. A. cardiophylla (MAST.) ENGL. and A. viridiflora CRA1B which oc- cur outside Malesia in the Himalayan-Indochinese region, are not entered in the key. They are closely allied to A. heterophylla, but distinguished by the thick pericarp, which is 5-20 mm @g at the valve sutures. 419 1972] PASSIFLORACEAE (de Wilde) 3. Stipe of ¢ flowers 1-4(-8) mm. Flower buds ovate. Fruits subglobular or + fusiform; dry pericarp woody. Leaves entire. 4. g Flowers including the 1 %4—4(—8) mm long stipe 9-15 by 2-3 4% mm. Anthers (4—)4 4-7 by *4-1% mm. Leaves suborbicular, ovate or obovate to oblong-lanceolate, palmati- to pinninerved, with acute-acuminate to rounded base and obtuse to acute, up to c. 1 cm acuminate apex; when dry green to dark brown above, pale brownish or greenish to whitish beneath. Glands at blade-base restricted to the auricles. Fruits globular or + fusiform; the valves 4%—3 mm thick. 3. A. macrophylla 4. g Flowers including the 1-114 mm long stipe 7-9 by 4(-5) mm. Anthers 3—4 by | mm. Leaves ovate to ovate-elliptic, sub-3—5-plinerved, with cordate to broadly rounded base and %—1 4% cm acumi- nate apex; when dry dark brown at both surfaces. Glands at blade-base large, extending beyond the auricles on the blade. Fruits globular, with c. 3 mm thick valves. 4. A. kinabaluensis 2. Calyx lobes 1-2 mm, erect in anthesis. Leaves entire, ovate to oblong, with deeply cordate to sub- truncate base, 244-10(-17) cm long. Gland-bearing auricles hemiglobular, deeply concave, + sep- arate from the blade, not peltately connate. 5. Fruits fusiform, -- 3(—6)-angular; valves when dry hard-coriaceous, 1-144 mm thick. 3 Flowers including the 10-20 mm long stipe 18-35 by 14%-3(-4) mm. . . oot COA “cordifolia 5. Fruits globular, not ribbed; dry valves -- Wg oe y mia 6 mm 1 thick. 3 Flowers including the 9-10 mm long stipe 16-18 by 242-3 mm Fer se aeae : 6. A. crassa 1. Section Microblepharis (W. & A.) ENGL. Bot. Jahrb. 14 (1891) 376; HARMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 84; ibid. ed. 2, 21 (1925) 492; HALL. f. Med. Rijksherb. 42 (1922) 8, p.p. — Modecca subg. Microblepharis W. & A. Prod. 1 (1834) 353. — Microblepharis Roem. Syn. Mon. 2, Pepon. (1846) 133, 200. — Modecca sect. Microblepharis ENDL. Gen. Pl. (1839) 928; Mia. Fl. Ind. Bat. 1, 1 (1856) 702, quoad bas.; BENTH. & Hook. f. Gen. Pl. 1 (1867) 813; Mast. in Hook. f. Fl. Br. Ind. 2 (1879) 601. Sepals free. Petals free. Hypanthium about as long as wide. Corona present. Distr. Africa (9 spp.), Madagascar (1 sp.), SE. Asia and Malesia (5 spp.). 1. Adenia penangiana (WALL. ex G. DON) DE WILDE, Blumea 15 (1967) 266. See for references and synonymy under the varie- ties. Climber or creeper to 6 m, with tuberous root- stock. Leaves herbaceous to coriaceous, entire, ovate-elliptic to linear, top acute, up to 2 cm acu- minate, base rounded, rarely subacute, (1 1 —)2—16 by (4-)%-7% cm, pinninerved; nerves 5-12 pairs; petiole (44-)'4-344 cm. Glands at blade- base 2, free or contiguous, 14-114 mm 9, situated on the mostly slightly 2-lobed, up to 5 mm broad peltate base; blade-glands up to 14 mm 9, 0-9 at either side, situated marginal or rarely submarginal Inflorescences up to 10 cm peduncled, rarely (sub)- sessile in short-shoots up to 3 cm; ing up to 30- flowered, often cincinnate, in 2 1—3-flowered; tendrils 0-3, 14-1 cm. Sterile tendrils simple or 3-fid, 1-10 cm, sometimes ending in adhesive disks. Bracts and bracteoles triangular to oblong, acute, 14-114 mm. Plants dioecious or monoecious. — 9 Flowers including the 1-7 mm long stipe 8-17 by 1-4 mm, the sepals spreading in anthesis up to c. 10 mm wide. Hypanthium cup-shaped, 1—2(-3) mm; calyx tube 0(-2) mm. Sepals free or nearly so, oblong to lanceolate-linear, acute to obtuse, 414-15 mm. Petals elliptic to oblong, ++ unguicu- late, obtuse to subcute, 4-10 by 114-3 mm, finely serrulate. Filamentsa 214-314 mm, connate for 1-214 mm. Anthers 2-314 by 14-34 mm, obtuse to subacute, up to 1 mm apiculate. Septa 1-2 mm high. Corona composed of fine hairs or a finely laciniate membrane 0.1—-'4 mm. Disk glands 14-1 mm. — & Flowers including the 1—3(—5) mm long stipe 6-16 by 1-2!4 mm. Hypanthium cup- shaped !4-11!4 mm. Calyx tube 0. Sepals oblong to lanceolate-linear, acute to obtuse, 4-10 mm. Petals elliptic to oblong, obtuse, 2-5 by *4-144 mm, + finely serrulate. Staminodes 1-114 mm, connate for 144-14 mm. Septa 4-4 mm high. Corona hairs 14-14 mm. Disk glands c. 4 mm. Pistil 314- 6 mm; gynophore c. 14 mm; ovary ellipsoid to oblong, 2-3 by 114-114 mm; styles connate for up to 14 mm, style-arms 1-2 mm; stigmas finely papillate, each c. 1 mm @. Fruit 1, ellipsoid to oblong, top obtuse to subacute, excluding the 2-25 mm long gynophore (134—-)2-5'2(-6) by 114-3 cm; pericarp (dry) coriaceous, 4-4 mm. Seeds (3—)5-15 subglobular or flattened or sub- triangular, 5-11 by 5-11 by 3-4 mm, smooth or grooved or variously pitted; embryo 5-9 mm; cotyledons suborbicular, sometimes obliquely trun- cate, 5-9 by 5-8 mm. Distr. Nicobar Is., Peninsular Thailand, in Malesia: Malay Peninsula and Sumatra. Fig. 7. Notes. Beside inflorescences in the axils of 420 FLORA MALESIANA [ser. 1 walle? Fig. 4. Adenia cordifolia (BL.) ENGL. a. Branch with infructescence, x 24, b. ditto with 3 inflorescence, « 2%, c. $ flower, <2, d. 3 flower in longitudinal section, x4, e. fruit, x 24, f. seed with aril, x2, g. seed, x2 (a ALVINS 2288, b-d RIDLEY 10197, e—f BLUME 2030, in spirit, g BECCARI 2155). 1972] PASSIFLORACEAE (de Wilde) 421 | Mf %, Bail 4 il HH Poona ae 2 soe @ shy iP o & 4968 ie ) Ry Fig. 5. Adenia penangiana (WALL. ex G. DON) DE WILDE var. penangiana. a. Habit, b—-c. leaves from above. — Ditto var. parvifolia (PIERRE ex GAGN.) DE WILDE. d. Leaf from beneath, e. tuber; all x 24 (@ RAHMAT SI TOROES 2329, b MARADJO 158, c BECCARI 4409, d HANIFF & NuR 7497, e HENDERSON S.n.). 422 FLORA MALESIANA [ser. I, vol. 7? Fig. 6. Adenia penangiana (WALL. ex G. DON) DE WILDE var. penangiana. a—b. 3 Flowers in longitudinal section, x6, c. petal, x6, e. 9 flower in longitudinal section, x6, f. fruit, x 24, h. seed, x 2, j. detail of inflorescence with tendrils, «4, k. detail of inflorescence, x 4. — Ditto var. parvifolia (PIERRE ex GAGN.) DE WILDE. d. g Flower in longitudinal section, x6, g. fruit, x 73, i. seed, x2 (a LORZING 6832, b-c, j RAHMAT SI ToROES 2329, d HANIFF & NUR 7497, e MEIJER 3196, f JELINEK 148, g, i CORNER SFN 37889, h HAnNiFF 3909, k CurTIS 3504). 1972] normals leaves sometimes (sub)sessile inflorescen- ces arranged in short-shoots are found. These short-shoots develop from the serial bud in the axils of the sterile tendrils, and later on often grow through into normal shoots. Sometimes monoecious specimens with 3 and 2 flowers in different inflorescences are found in var. parvifolia. A variable species in which arbitrarily two varie- ties are recognized; most of the specimens from limestone in Peninsular Thailand belong to var. parvifolia. KEY TO THE VARIETIES 1. Fruit 3-6 cm, gynophore 4-25 mm. Seeds flat- tened, 714-11 mm 9g, shallowly pitted. Hypan- thium 114-3!14 (-4) mm wide. Anthers (2!4-) 3-314 mm, up to 1 mm apiculate. Leaves never iDeenir ot ne eee eee cee a. var. penangiana 1. Fruit 134-4 cm, gynophore 2-5 mm. Seeds globular to flattened, 4-7 mm 9, smooth or shallowly pitted. Hypanthium 1-2 mm wide. Anthers 2—3(—3!4) mm, up to 0.2 mm apiculate. Leaves sometimes lanceolate-linear. b. var. parvifolia a. var. penangiana. — A. penangiana (WALL. ex G. Don) DE WILDE, Blumea 15 (1967) 266; Thesis (1971) 84, 88, f. 9-10. — Passiflora penangiana WALL. [Cat. (1829) n. 1233, nom. nud.] ex G. Don, Gen. Syst. 3 (1834) 55; Masr. Trans. Linn. Soc. 27 (1871) 631. — Anthactinia penangiana ROEM. Syn. Mon. 2, Pepon. (1846) 192. — Disemma penangiana Mia. FI. Ind. Bat. 1, 1 (1855) 700. — Modecca nicobarica KURZ ex TRIM. J. Bot. 13 (1875) 326; J. As. Soc. Beng. 45, ii (1876) 132; Mast. in Hook. f. Fl. Br. Ind. 2 (1879) 603. — A. nicobarica KiNG, J. As. Soc. Beng. 71, ii (1903) 52; Ripi. Fl. Mal. Pen. 1 (1922) 840, pro maj. parte; HALL. f. Med. Rijksherb. 42 (1922) 9; HEND. Gard. Bull. S. S. 4 (1928) 264, p.p.; J. Str. Br. R. As. Soc. 17 (1939) 47, p.p.; CRAIB, Fl. Siam. En. 1 (1931) 747, incl. var. obliqua CRAIB; MERR. Contr. Arn. Arb. 8 (1934) 110; CHAKRAVARTY, Bull. Bot. Soc. Beng. 3 (1951) 65. — A. catharinae Merr. Contr. Arn. Arb. 8 (1934) 110, t. 7. — Fig. 5 a—c, 6 a—c, e—f, h, j-k. Climber to c. 6 m. Leaves broadly ovate-elliptic to (ob)lanceolate, top mostly distinctly acuminate, base rounded, rarely subacute, 314-16 by 1-714 cm. (Sub)marginal glands present or not. Inflores- cences peduncled for 4-10 cm. — ¢ Flowers including the 1-5 mm long stipe 8-17 by 114-4 mm. Hypanthium 114-2 mm. Calyx tube 0. Sepals ob- long to lanceolate, 5-13 mm. Petals 4-10 mm. Filaments 214-314 mm, connate for 1-2 mm. Anthers (2!4-)3-3144 mm, up to 1 mm apiculate. Corona-filaments (0.1-)44-’%4 mm. — 2 Flowers including the 1-5 mm long stipe 6-16 by 114-214 mm. Hypanthium 1-114 mm. Calyx tube 0. Sepals 5-9 mm. Fruit excluding the 4-25 mm long gyno- phore 3-6 by 1!4-3 cm. Seeds flattened, nearly smooth or shallowly or pitted, 714-11 mm @. Distr. Nicobar Is., Peninsular Thailand, in PASSIFLORACEAE (de Wilde) 423 r 7 af inna f r % o a 1a @ Adenia penangiana var. penangiana 1b A Adenia penangiana var parvifolia Fig. 7. Distribution of Adenia penangiana (WALL. ex G. DON) DE WILDE. Malesia: Malay Peninsula and N. & Central Sumatra. Fig. 7: la. Ecol. Forest and scrub, sometimes on limestone, 0-1200 m. Fl. fr. Jan.—Dec. Notes. The fruits of most specimens have a long gynophore, of at least 1 cm; specimens from the Taiping-area (Mal. Pen.), however, often have rather short-stiped fruits. Fresh flowers are green or greenish yellow, fruits yellowish red to brilliant cinnamon. b. var. parvifolia (PIERRE ex GAGN.) DE WILDE, Thesis (1971) 84, 89, f. 9-10. — A. parvifolia PIERRE ex GAGN. Bull. Mus. Hist. Nat. Paris 25 (1920) 127; Bull. Soc. Bot. Fr. 65 (1918) 76-77 (fl. morph.); Fl. Gén. I.-C. 2 (1921) 1028, f. 113, 8-13; CraiB, FI. Siam. En. 1 (1931) 748, incl. var. insularis et var. nervosa; CUSSET, Fl. Camb., Laos & Vietn. 5 (1967) 149, t. 2 f. 7, t. 5 f. 13-16; Adan- sonia 7 (1967) 373, 383. — A. nicobarica (KURZ) KING ex RIDL. J. Str. Br. R. As. Soc. 59 (1911) 106; BurK. & HEND. Gard. Bull. S. S. 3 (1925) 378. — A. angustisepala CrRAtB, Kew Bull. (1930) 406; Fl. Siam. En. 1 (1931) 745; Cusset, Adansonia 7 (1967) 372, 383. — A. linearis CRAB, Kew Bull. (1930) 407; Fl. Siam. En. 1 (1931) 747; CussET, Adansonia 7 (1967) 373, 383. — Fig. 5 d-e, 6 d, g, i. Climber or creeper up to 2 m, growing from a tuber. Leaves ovate-oblong to lanceolate-linear, top longly acute or acuminate, base rounded, (114-)2-13 by 4-414 cm. Marginal glands absent. Inflorescenses either in short-shoots or in the axils of normal leaves, up to 4 cm peduncled. — 3 Flo- wers including the 3-7 mm long stipe 8-15 by 1-2 mm. Hypanthium 114-3144 mm. Calyx tube 0-2 mm. Sepals lanceolate to linear, 4-8 mm. Petals 4-5 mm. Filaments 214-314 mm, connate for 114-214 mm. Anthers 2—3(-314) mm, up to 0.2 mm apiculate. Corona-hairs 0.1-0.3 mm. — 424 2 Flowers including the 1-—3(—5) mm long stipe 8-12 by 1-2 mm. Hypanthium 1-114 mm. Calyx tube 0. Sepals 4-7 mm. Fruit excluding the 2-5 mm long gynophore 134-4 by 1-2 cm. Seeds subglobu- lar to + flattened, smooth or shallowly grooved or pitted, 4-7 mm g. Distr. Peninsular Thailand, in Malesia: NW. FLORA MALESIANA [ser. I, wale@? Malay Peninsula (Perlis, Kedah, Langkawi Is.). Big: 7/2 lb: Ecol. Limestone hills, ridges, 0-600 m. Fi. fr. mostly Sept.—Jan. Note. Fresh flowers recorded as greenish or greenish yellow, fruits as greenish turning bright red. 2. Section Erythrocarpus (ROEM.) DE WILDE, Thesis (1971) 209. — Erythrocarpus Roem. Syn. Mon. 2, Pepon. (1846) 133, 204. — Modecca subg. Erythrocarpus Mia. Fl. Ind. Bat. 1, 1 (1856) 703. — Modecca subg. *Modeccae verae’ (incl. sect. Microblepharis et sect. Blepharanthes pro maj. parte, typo excl.) Miq. Fl. Ind. Bat. 1, 1 (1856) 702. — Adenia sect. Microblepharis HALL. f. Med. Rijksherb. 42 (1922) 8, pro maj. parte, typo excl.). Sepals largely connate into a tube. Petals largely connate with the calyx tube; hypanthium not differentiated. Corona absent. Distr. SE. Asia and Malesia (7 spp.). 2. Adenia heterophylla (BL.) Koorp. Exk. FI. Java 2 (1912) 637; DE WILDE, Thesis (1971) 212. See for references and synonymy under the sub- species and varieties. Climber to 30 m. Leaves membranous (herba- ceous) to coriaceous, entire to 5—partite, orbicular to ovate to lanceolate, top rounded to acute, up to 3 cm acuminate, base acute to cordate, (314—)5-25 by (114-)214-19 cm, 3—5(—7)-plinerved to pinni- nerved by 4-10 pairs of nerves, margin entire or up to 4 cm dentate; lobes triangular to lanceolate, up to 15 cm; petiole 1-10 cm. Glands at blade-base 2, 1-4 mm g, on two auricles 2-6 mm @ at the apex of the petiole, -- adnate with the blade, either -+ connate over the apex of the petiole or not; blade-glands 0-2 pairs, 14-2 mm g, submarginal; marginal glands minute, 0-25 at either side. Inflo- rescences peduncled up to 20 cm, rarely in short- shoots, ing up to 40-flowered, in 2 (1-)2-4(-8)- flowered; tendrils 1(—3), 1-5 cm. Sterile tendrils simple, rarely 3-fid, up to 25 cm. Plants sometimes monoecious with g and 2 flowers mixed in one inflorescence. Bracts and bracteoles narrowly triangular, acute, 14-114 mm. — ¢ Flowers tubi- form to urceolate, including the 3-15 mm long stipe (10—)15—25(—30) by 1'4-5(-714) mm. Hypanthium including calyx tube 5—12(-14) mm, fleshy. Calyx lobes triangular, acute to subobtuse, 1-3 mm, re- flexed, inserted near the throat of the calyx tube. Petals narrowly triangular to lanceolate, subacute, 2-4 mm, reflexed. Filaments 1-4 mm, connate for 14-3 mm, inserted at the base of the hypanthium, or on an androgynophore up to 4 mm. Anthers 3-5 mm, subacute, up to 14 mm apiculate. Septa 1-3 mm high. Corona 0. Disk glands 1-3 mm. — 2 Flowers tubiform, including the 1-6(-10) mm long stipe (6—)7—18(—22) by 3-5(-6) mm. Hypan- thium including calyx tube (4-)6-13 mm. Calyx lobes elongate triangular, subacute, 1-214 mm. Petals oblong to lanceolate, (sub)acute, 2-4 mm, inserted near the throat of the calyx tube. Stami- nodes 1-3 mm, connate for up to 1144 mm. Septa 14-2 mm high. Corona 0. Disk glands 14-214 mm. Androgynophore up to 244 mm. Ovary 1-314 mm stiped, subglobose to oblong, 3-5 by 2-3 mm, 3(—S)—carpellate; styles 3(—5), 44-1 m up to halfway connate; stigmas papillate, each c. 114 mm 9g. Fruits 1—3(-4), ellipsoid to oblong-lan- ceolate, sometimes ++ 3-ribbed, base and top obtuse or acute, excluding the (14—)1—3(—4) cm long gynophore 2-13 by 134-414 cm; pericarp coria- ceous, 1-3 mm g, when fresh + fleshy, yellowish to bright red. Seeds 10-60, orbicular to obliquely triangular, (4-)5-10 by 4!4-10 by 214-314 mm, pitted, sometimes muricate; embryo 4-814 mm; cotyledons orbicular to ovate, emarginate or truncate at one side, or shallowly 3—lobed, 4-71, by 47 mm. Distr. From the Andaman Is., Indo-China and S. China through Malesia to N. Australia, east to the Solomon Is., absent in Sumatra, Malaya, and Borneo. Fig. 8. Ecol.Ina variety of habitats in forest and scrub. The species shows as a whole a distinct preference for seasonal climatic conditions and is absent in Sumatra, the Malay Peninsula and Borneo. Fi. fr. Jan.—Dec., but mostly in the rainy season. The species is usually dioecious, but not rarely monoecious specimens occur with 3 and 2 flowers in one inflorescence. The tubular, narrow-throated flowers suggest pollination by insects. Uses. The plant as a whole, and especially the fruit, is reported as poisonous, and used as poison for hunting; the juicy aril is sometimes mentioned as sweet and edible, whereas POILANE reported for Indo-China that the leaves are eaten by the Mois. PASSIFLORACEAE (de Wilde) 425 2,1a @ Adenia heterophylla ssp. heterophylla var. heterophylla 2,1b *& Adenia heterophylla ssp. heterophylla var. celebica 2,2 A Adenia heterophylla ssp. australis © Adenia kinabaluensis Fig. 8. Distribution of Adenia heterophylla (BL.) Koorp. and A. kinabaluensis DE WILDE. In the Philippines a decoction of the root is a remedy for stomach trouble. Vern. (Areuj) patok manok,S, kabelo, Kangean, sasariwu, tatanaru warawo, Talaud; Philippines: binoyok-béyok, meléng-uadk, salapong, Tag., saka- sdka, s.-ti-udk, Ik., tababayung-udk, Sulu, nomad- nomad, Palawan, tabungau, Mindoro, tambal baya (s), Pint.-Sbl.; daun bobok, Tanimbar, malasibi, NE. New Guinea. Notes. Modecca heterophylla Bu. and the synonym M. acuminata BL., the oldest two names available, are both of 1826. The epithet hetero- phylla is chosen because of the current misinter- pretation of specimens from the Malay Peninsula which belong to a different species, A. macro- phylla, under the name Adenia acuminata (non BL.) KING. A variable species, in which rather arbitrarily four largely allopatric subspecies — one of which with two varieties — are recognized. Two of the subspecies do not occur in the Malesian region, but these have been entered in the key. KEY TO THE SUBSPECIES AND VARIETIES 1. Stipe of 2 flowers (1—)2—6 mm, in fruit (1—)3—13 mm. Gynophore in fruit 8-30 mm. Fruits (4-)6-13 cm, mostly with acute apex. Stipe of 6 flowers 5-15 mm, as long as to longer than the remainder of the flower. Leaves orbicular to lanceolate-linear in outline with cordate to acute base, palmately to pinninerved, vena- tion distinct or not . .1. ssp. heterophylla 2. Leaves subherbaceous to coriaceous, orbicular to lanceolate(-linear), entire to deeply 5— lobed, base cordate to acute, nerves palmate to pinnate, margin entire or dentate. Gland- bearing auricles well marked off from the blade, glands limited to the auricles. a. var. heterophylla 2. Leaves strongly coriaceous, ovate-oblong to oblong, entire, base rounded to subacute, nerves pinnate, margin entire. Gland-bearing auricles broadly adnate with the blade, the glands extended on the blade. b. var. celebica 1. Stipe of 2 flowers c. 1 mm, in fruit 1-2 mm. Gynophore in fruit 3-13 mm. Fruits 2-7 cm. Stipe of ¢ flowers 3-8 mm, as long as to shorter than the remainder of the flower. Leaves orbi- cular to ovate in outline, with cordate to trun- cate base, mostly palmately nerved, venation distinct beneath. 3. Leaves mostly membranous, margin entire. Marginal glands mostly absent. Gland-bearing auricles + peltately connate, sometimes free. Stipe of 3 flowers about as long as the remain- der of the flower. Filaments connate about halfway. Fruits 4-7 cm, apex obtuse. 2. ssp. australis 426 3. Leaves + coriaceous, margin entire or dentate. Marginal glands present. Gland-bearing auri- cles free. Stipe of g flowers shorter than the remainder of the flower. Filaments more than halfway connate. 4. Leaves (sub)orbicular, up to 1 cm acuminate, distinctly reticulate at both sides. Fruits 2-314 (-4) cm, apex obtuse. Thailand, Laos, Cam- bodia, southern S. Vietnam (Cochinchina). ssp. arcta (CRAIB) DE WILDE 4. Leaves suborbicular to ovate, 1-2 cm acumi- nate, reticulate only beneath. Fruits 414-7 cm, apex acute. Great Coco I., Andamans & Nicobars. . .ssp. andamanica DE WILDE la. ssp. heterophylla var. heterophylla. — A. heter- ophylla (BL.) Koorp. Exk. Fl. Java 2 (1912) 637; HALL. f. Med. Rijksherb. 42 (1922) 8; BAck. & Baku. f. Fl. Java 1 (1963) 289; Cusser, Adansonia 7 (1967) 373, 382; DE WILDE, Thesis (1971) 216, f. 35. — Modecca heterophylla Bu. Bijdr. (1826) 940; DC. Prod. 3 (1828) 336; G. Don, Gen. Syst. 3 (1834) 59; Hassk. Cat. Hort. Bog. (1844) 187; Mia. Fi. Ind. Bat. 1, 1 (1855) 702; F.—VILL. Nov. App. (1880) 95. — Microblepharis heterophylla Roem. Syn. Mon. 2, Pepon. (1846) 133, 200. Modecca acuminata BL. Bijdr. (1826) 940; DC. Prod. 3 (1828) 336; G. Don, Gen. Syst. 3 (1834) 59; HAssk. Cat. Hort. Bot. (1844) 187; Mra. FI. Ind. Bat. 1, 1 (1855) 702. — Microblepharis acuminata Roem. Syn. Mon. 2, Pepon. (1846) 133, 200. — A. acuminata KING, J. As. Soc. Beng. 71, ii (1903) 55, quoad basionym; Koorp. Exk. FI. Java 2 (1912) 437; Faiz. f. Med. Rijksherb. 42 (1922) 11; Back. & BAKH. f. Fl. Java 1 (1963) 289; Cusser, Adan- sonia 7 (1967) 372, 383. Passiflora parviflora BLco, FI. Filip. ed. 1 (1837) 647, non SwArRtTZ, 1788. — Modecca parviflora Bico, Fl. Filip. ed. 2 (1845) 453, non G. Don, 1834; ibid. ed. 3, 3 (1879) 52; MerR. Philip. J. Sc. 10 (1915) Bot. 331. — A. parviflora Cusset, FI. Camb., Laos & Vietn. 5 (1967) 145, t.2 f. 1, t. 5 f. 3-12, t. 7 f. 5-6, quoad basionym, spec. p.p., nom. illeg.; Adansonia 7 (1967) 373, 383. Passifiora zucca Buco, FI. Filip. ed. 1 (1837) 648. — A. zucca MERR. Sp. Blanc. (1918) 276; En. Philip. 3 (1923) 117; HARMs in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 472, 492. Passiflora coccinea Buco, FI. Filip. ed. 1 (1837) 650, non AUBL. 1775, nec BANKS & SOLAND. ex BENTH. 1867. — Modecca coccinea BLco, FI. Filip. ed. 2 (1845) 453; ibid. ed. 3, 3 (1879) 53; MERR. Philip. J. Sc. 1 (1906) Suppl. 100. — A. coccinea Merr. Philip. J. Sc. 3 (1908) Bot. 421; Fl. Manila (1912) 337; Philip. J. Sc. 10 (1915) Bot. 331. Modecca lobata (non JAcg.) Hassk. Cat. Hort. Bog. (1844) 187; Mia. FI. Ind. Bat. 1, 1 (1855) 703. Modecca kardiocarpa Hassk. Cat. Hort. Bog. (1844) 187; WALP. Rep. 5 (1846) 774; Roem. Syn. Mon. 2, Pepon. (1846) 203; Mra. Fl. Ind. Bat. 1, 1 (1855) 703. — A. cardiocarpa Koorp. Exk. FI. Java 2 (1912) 637. Modecca oblonga Hassk. Cat. Hort. Bog. (1844) 187; WALP. Rep. 5 (1846) 774; Roem. Syn. Mon. 2, Pepon. (1846) 203; Mra. FI. Ind. Bat. 1, 1 (1855) FLORA MALESIANA [ser. I, vol. 7? 703. — A. oblonga Koorp. Exk. FI. Java 2 (1912) 637. Modecca trilobata (non Roxs.) Btco, FI. Filip. ed. 2 (1845) 452; ibid. ed. 3, 3 (1879) 52; F.-VILL. Nov. App. (1880) 95; Merr. Philip. J. Sc. 1 (1906) Suppl. 100 (‘triloba’); ibid. 10 (1915) Bot. 331. Modecca cardiophylla (non Mast.) F.—VILL. Nov App. (1880) 95. Modecca palmata (non LAMK) F.-VILL. L.c. Modecca populifolia (non BL.) K. Scu. & HOLLR. Fl. Kais. Wilh. Land (1889) 83. — A. populifolia (non BL.) K. Sco. & Laut. Fl. Schutzgeb. (1900) 456; PULLE, Nova Guinea 8 (1912) 673 (aff. populi- folia); Wuite, J. Arn. Arb. 10 (1929) 244. Momordica sp., PULLE, Nova Guinea 8 (1910) 405. Modecca formosana HAYATA, Ic. Pl. Form. 4 (1914) 8, f. 1-2; Ivo, Ill. Formos. Pl. (Taiwan Shokubutu Dzusetu) (1927) t. 11; SAsAK1, Cat. Govt. Herb. Formos. Dept. For. (1930) 362. — A. formosana HAYATA, Ic. Pl. Form. 4 (1914) 8, f. 1-2; HARMs in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 492; CusseT, Adansonia 7 (1967) 373, 384. A. longifolia MERR. Philip. J. Sc. 10 (1915) Bot. 330; En. Philip. 3 (1923) 117. A. palmatifolia MERR. Philip. J. Sc. 10 (1915) Bot. 330; En. Philip. 3 (1923) 117. A. chevalieri GAGN. Bull. Mus. Hist. Nat. Paris 25 (1920) 126; Bull. Soc. Bot. Fr. 65 (1918) 76-77 (fl. morph.); Fl. Gén. I.-C. 2 (1921) 1030, f. 114, 1-5; HArMs in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 490; Crais, Fl. Siam. En. 1 (1931) 746; Merr. & CHUN, Sunyatsenia 1 (1934) 73; CHUN, ibid. 1 (1934) 276; MASAM. FI. Kaitan. (1943) 216; Wana, Acta Phytotax. Sin. 6 (1957) 237; CHUN, CHANG & CHEN, FI. Hainan. 1 (1964) 467, f. 258; CusseET, Fl. Camb., Laos & Vietn. 5 (1967) 140, f. 2, t. 5 f. 20-23, t. 7 f. 2-3; Adansonia 7 (1967) 372, 383. A. cordifolia (non BL.) GAGN. FI. Gén. I.-C. 2 (1921) 1025; Bull. Soc. Bot. Fr. 65 (1918) 76-77 (fl. morph.). A. nicobarica (non KURZ) GAGN. FI. Gén. I.-C. 2 (1921) 1028; CusseT, Fl. Camb., Laos & Vietn. 5 (1967) 148, t. 2 f. 3. A. maclurei MERR. Philip. J. Sc. 21 (1922) 349; men. Scads 5 (192 7)N33. A. diversifolia HALL. f. Med. Rijksherb. 42 (1922) 10. A. sumbawana HALL. f. l.c. A. pandurata HALL. f. I.c. 12; HoLttH. & H. J. Lam, Blumea 5 (1942) 215. A. pinnatisecta (non CRAIB) PHAM-HOANG-HO, Fl. Vietn. (1960) 148, f. D. Leaves up to 3 cm acuminate, 314-25 by 114-19 cm; petiole 1-10 cm. Gland-bearing auricles pel- tately connate over the top of the petiole or not. — 3$ Flowers including the 7-15 mm long stipe 15-30 by 2-5(-714) mm. — 2 Flowers including the (114-)2-6 mm long stipe (10—)12-18 by 2-6 mm. Fruit excluding the 8—-30(-40) mm long gynophore (4-)6-13 by 2!4-414 cm; flower stipe below the 1972] PASSIFLORACEAE (de Wilde) 427 withered perianth (1 !4—)3-13 mm. Seeds (5—)6—10 mm 9g. Distr. Tropical SE. Asia to the Solomons and Queensland, in Malesia: Central Sumatra (once), Java, Lesser Sunda Is. (once, Sumbawa), Philip- pines, Celebes, Moluccas, New Guinea (also Bis marcks). Fig. 8: 2—la. Ecol. Forest and scrub, often in secondary veg- etation, growing on a variety of soils, sand, clay, silt, rocks, etc., 0-1000 m, in New Guinea to 2000 m. Distinctly preferring a seasonal climate, hence absent from Peninsular Thailand, the Malay Pen- insula, Borneo, and Sumatra. In the latter island there is a single, old collection (KORTHALS 682b), which might have been mislocalized and could have come from Java. In islands with a mosaic- climate also found in everwet parts, e.g. in West Java, the Philippines, Celebes and New Guinea, possibly facilitated through devastation. Notes. Var. heterophylla is a variable entity in which a number of intergrading local forms (para- morphs) have been described as species. For a discussion of the synonymy, with argumentation, see my Thesis (1971) 219. Flowers are sometimes 4—merous, with 4 calyx lobes. HAyATA (1914) mentioned for the type specimen of A. formosana 4-S-carpellate ovaries; 4 or 5 carpels are also occasionally found in A. macrophylla. In specimens from the Philippines several times galled. club-shaped flowers were found. Field notes. Leaves are often reported as very glossy; the flowers as greenish, creamy or yellow- ish; ripe fruits as often + 3(—6)—angular, yellow to bright red; seeds blackish covered by a whitish aril. 1b. ssp. heterophylla var. celebica (KOCRD.) DE WILDE, Thesis (1971) 220, f. 35. — Modecca celebica KoorD. Minah. (1898) 638, 478. — A. celebica KoorD. in Koord.—Schum. Syst. Verz. 3 (1914) 90. Leaves up to 114 cm acuminate, 5—16 by 3—7(-9) cm; petiole (44—-)1-S5 cm. Gland-bearing auricles not peltately connate, broadly adnate with the auricles, the glands partly extending on the blade. — ¢ Flowers including the 5-11 mm long stipe 12-22 by 114-3 mm. — °& Flowers including the (1144-)2-5 mm long stipe 10-13 by 2-314 mm. Fruit fusiform, excluding the 10-20 mm long gy- nophore 5-8 by 2-3 cm; flower stipe below the withered perianth (114—)2-6 mm. Seeds 5-6 mm @. Distr. Malesia: N. Borneo (E. Sabah: Elphin- stone Bay), Celebes (North Peninsula), Moluccas (Ceram). Fig. 8: 2—1b. Ecol. Up to c. 200 m. Fi. Febr., March, Sept., fr. March, Sept. 2. ssp. australis (R. BR. ex DC.) DE WILDE, Thesis (1971) 220, f. 35. — Modecca australis R. BR. ex DC. Prod. 3 (1828) 337; G. Don, Gen. Syst. 3 (1834) 59; ENDL. Ic. Gen. Pl. (1838) t. 114-115; Roem. Syn. Mon. 2, Pepon. (1846) 203; SCHNITZL. Ic. 3 (1851) t. 197; BENTH. FI. Austr. 3 (1866) 312; F. v. M. Fragm. Phyt. Austr. 9 (1875) 69; First Syst. Census (1882) 76; Second Syst. Census (1889) 128; F. M. BAILEY, Syn. Queensl. Fl. (1883) 200; Cat. Pl. Queens]. (1890) 20; Queensl. Fl. 2 (1900) 689; Compr. Cat. Queens]. PI. (1913) 220, f. 192; Domin, Bibl. Bot. Heft 89 (1928) 987. — A. australis ENGL. Bot. Jahrb. 14 (1891) 376; Harms, ibid. 15 (1893) 572-573; in E. & P. Nat. Pfi. Fam. 3, 6a (1893) 85; ibid. ed. 2,21 (1925) 492; Ewart & Davies, Fl. North. Terr. Austr. (1917) 196; SpecHT, Rec. Am.—Austr. Exp. Arnhem Land 3 (1958) 262. Modecca populifolia Zipp. ex BL. Rumphia | (1837) 168, t. 50; SPAN. Linnaea 15 (1841) 207; WALP. Rep. 2 (1843) 222; Mia. Fl. Ind. Bat. 1, 1 (1855) 703; Mast. in Hook. f. Fl. Br. Ind. 2 (1879) 603; BRITTEN in Forbes, Wand. efc., App. 6 (1885) 506; F. M. BAILEY, Queens]. Agr. J. 1, 3 (1897) 228; Queensl. Fl. 2 (1900) 690; Compr. Cat. Queensl. (1913) 220, f. 193. — Erythrocarpus populifolius RoEM. Syn. Mon. 2, Pepon. (1846) 204. — A. populifolia ENGL. Bot. Jahrb. 14 (1891) 376; HARMS, ibid. 15 (1893) 573, 553 (anat.); in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 85; ibid. ed. 2, 21 (1925) 492: Hatt. f. Med. Rijksherb. 42 (1922) 9; RIDL. Disp. (1930) fig. frontisp. Leaves up to 1 cm acuminate, entire or lobed, 5-18 by 3-15 cm; petiole 1-5'4 cm. Gland-bearing auricles mostly peltately connate over the top of the petiole. — g Flowers including the 5-8 mm long stipe 11-18 by 2-3!4 mm. — 9 Flowers includ- ing the 1-114(—2) mm long stipe 6-8 by 2-3 mm. Fruit with obtuse apex, excluding the 3-13 mm long gynophore 4—7 by 2-3 cm; flower stipe below the withered perianth 1-2 mm. Seeds 4-7 mm 9. Distr. N. Australia, in Malesia: E. Java, Lesser Sunda Is. (incl. also Tanimbar Is.), Fig. 8: 2-2. Ecol. Scrub vegetation in a seasonal climate, monsoon forest, 0-150 m, mostly on sandy or calcareous soils near the coast. Fi. fr. (Java) mainly Febr._May, in Lesser Sunda Is. & Australia March—Dec. 3. Adenia macrophylla (BL.) Koorpb. Exk. Fl. Java 2 (1912) 637; DE WILDE, Thesis (1971) 226, f. 34. See for references and synonymy under the varie- ties. Liana to 25 m, at base up to 15 cm 9. Leaves (sub)coriaceous, entire or up to 14 cm lobed in upper half, suborbicular to oblong-lanceolate or (ob)ovate, top (sub)obtuse to acute, up to 114 cm acuminate, base acute-acuminate, or rounded or subcordate, (4—)5-21 by (114-)214-12 cm, 3—5 subplinerved or + pinninerved; nerves 3-5(-10) pairs, arching towards the top; petiole 14-714 cm. Glands at blade-base 2, 1-4 mm g, entirely or largely situated on two semi-orbicular auricles 114-5 mm @ at the apex of the petiole; blade-glands 0-4(—6), 14-1 mm 9, submarginal in the upper half of the blade; marginal glands 0-25 at either side. Inflorescences peduncled for up to 14 cm, sometimes subsessile in short-shoots up to 25 cm, lax or condensed, in f up to 150—flowered, in 2 2-10-flowered, tendrils 0-3, up to 4 cm long. Sterile tendrils simple or 3—fid, up to 20 cm. Bracts 428 and bracteoles narrowly triangular, acute, 14-1 mm. Flower buds ovate, not ellipsoid. —¢ Flowers narrowly tubiform-urceolate, including the 114-4 (-8) mm long stipe 9-15 by 2-314 mm. Hypan- thium including calyx tube tubiform-urceolate, fleshy, 514-9 mm. Calyx lobes triangular to oblong, subobtuse, 2—214(—3) mm, reflexed. Petals oblong- lanceolate, obtuse to subacute, subentire, 214-4 by 1-1'%4(-2'4) mn, inserted at or near the throat of the calyx tube. Filaments 1-114 mm, connate up to 14 mm, inserted at the base of the hypanthium. Anthers 4-7 mm, -+ tapering to above, subacute, 14-1 mm apiculate. Septa 0-44 mm high. Corona 0. Disk glands 14-1 mm. — @ Flowers tubiform- campanulate, including the 14-1 mm long stipe 5—7 by 214-3 mm. Hypanthium including calyx tube 3-4 mm. Calyx lobes 2-4 mm. Petals 2-3 mm, inserted near the throat of the calyx tube. Stami- nodes c. 1 mm. Septa 0. Corona 0. Disk glands c. 1 mm. Pistil 4-6 mm; gynophore 1—2 mm. Ovary subglobose to ovoid, 134-2144 by 114-2144 mm; styles 14-34 mm, + free; stigmas irregularly lobed, each c. 144 mm @g. Fruit 1-2(—3?), globose to broadly ovoid, base rounded, top rounded to subacute, or fruit + fusiform, excluding the (7—)10-30 mm long gynophore 2-6(—614) by 114-5 cm; pericarp coriaceous to woody, 14-3 mm thick. Seeds 15-40, + orbicular, 5-10 by 5-10 by 214-4 mm, + muricate and pitted; embryo 5-8 mm; cotyledons ovate to elliptic, often emarginate at one side, 414-714 by 4-6 mm. Distr. Malesia: Sumatra, Malay Peninsula, W. & Central Java, Borneo. Fig. 9. Ecol. Rain-forest; see further under the varie- ties. Uses. Fruits several times recorded as poison- ous. According to various authors the roots and leaves are medicinal. According to HEYNE (Nutt. Pl. 1927, 1142) the bark is used for spinning threads for fishing tackle in Sumatra’s West Coast. JACoBsS (n. 5003) mentioned: ‘The wood smells a bit of HCN; vessels contain potable water’. Vern. Akar gélumpong, a. lémpudu gajah, a. lupok, a. mérapoh, a. saut, a. sianun dundang, a. timon dandang, dérik-dérik, méntimun gajah mérah, m. pajah, pedendang, Mal. Pen.; akar djala, a. talun tungang, andor sidari, lakom gadjah, sautan, Sumatra; olor bauwo dotan, o. lawis, Simalur I.; areuj guntang, patok manok, J; lantiong, Anambas Is.; buah émpérah, Iban; gurtah, Sarawak; akar pétjah tutuban, Brunei; barabah, takup, Dusan, Sabah. Notes. The fruits are sometimes 4—carpellate. Often larva feeding on the pollen are found. De- formed galled flowers occur regularly. Flowers are once reported as odorless; the urceolate flowers enclosing the anthers, however, suggest entomo- gamy. Field notes. Fresh flowers are yellow or lemon, sometimes ‘waxy’, sometimes reddish spotted inside, or orange at the base; pollen bright yellow to orange-yellow. Fresh fruits are red, when dry orange-brown or purplish; the funicles in fresh fruits are pinkish, the arils whitish. FLORA MALESIANA [ser. I, voly 7? 3a @ Adenia macrophylla var. macrophylla 3b Adenia macrophylla var smilacina 3c x Adenia macrophylla var singaporeana Fig. 9. Distribution of Adenia macrophylla (BL.) KOorRD. KEY TO THE VARIETIES 1. Fruit globular to ovoid. Leaves mostly greenish beneath. 2. Leaves suborbicular to lanceolate, palmately to pinninerved. Fruit (2—)3-6(-6!4) cm; gy- nophore (10—)15—30 mm; valves (14—)1—3 mm thick. . . ... . . .a. var. macrophylla 2. Leaves suborbicular to ovate, 3—5—(sub)pli- nerved. Fruit 2-3 cm; gynophore c. 10 mm; valves 14-1 mm thick.. .b. var. smilacina 1. Fruit (ovate-)oblong, + fusiform; valves c. 1 mm thick. Leaves oblong to lanceolate, + pinninerved, mostly whitish green beneath. c. var. Singaporeana a. var. macrophylla; DE WILDE, Thesis (1971) 229, f. 34. — A. macrophylla (BL.) Koorp. Exk. FI. Java 2 (1912) 637; HALL. f. Med. Rijksherb. 42 (1922) 12; BAcK. & BAKH. f. Fl. Java 1 (1963) 289; CusseET, Adansonia 7 (1967) 373, 384. — Modecca macrophylla Bu. Bijdr. (1826) 939; DC. Prod. 3 (1828) 337; G. Don, Gen. Syst. 3 (1834) 59; HAssk. Cat. Hort. Bog. (1844) 187; Mia. Fl. Ind. Bat. 1, 1 (1855) 702. — Microblepharis macrophylla ROEM. Syn. Mon. 2, Pepon. (1846) 202. 2? Modecca dubia Roxs. [Hort. Beng. (1814) 49, nom. nud.| Fl. Ind. ed. Carey 3 (1832) 135. Modecca quintuplinervia Miq. Fl. Ind. Bat. 1, 1 (1855) 1093; Sum. (1860) 132, 333. — A. quintu- plinervia HALL. f. Med. Rijksherb. 42 (1922) 16. Modecca palmata (non LAMK) Kurz, Nat. Tijd. N. I. 27 (1864) 168. Modecca sp. RiDL. Trans. Linn. Soc. Bot. II, 39 (1893) 304. A. acuminata (non BL.) KiNG, J. As. Soc. Beng. 71, ii (1903) 55; RIDL. Fl. Mal. Pen. 1 (1922) 841; RENDLE, J. Bot. (1924) Suppl. 43; Burk. & HEND. 1972] PASSIFLORACEAE (de Wilde) 429 Gard. Bull. S. S. 3 (1925) 378; BARTLETT, Pap. Mich. Ac. Sc. 6 (1926) 31; HEND. Gard. Bull. S. S. 4 (1928) 264; Burk. Dict. 1 (1935) 48; RipL. Kew Bull. (1938) 112; MAsAm. En. Phan. Born. (1942) 506; HEND. Mal. Nat. J. 4 (1949) 153, f. 147; Cus- seET, Adansonia 7 (1967) 372, 383. A. clementis MERR. Philip. J. Sc. 13 (1918) Bot. 95; En. Born. (1921) 413; Univ. Cal. Publ. Bot. 15 (1929) 210; Rip. Kew Bull. (1938) 112; HEINE in Fedde, Rep. 54 (1951) 242; Pfl. Clemens Kina- balu (Thesis) (1953) 68. A. longipedunculata MeErRR. Philip. J. Sc. 13 (1918) Bot. 96; En. Born. (1921) 413. A. grandifolia RIDL. J. Fed. Mal. St. Mus. 10 (1920) 136; Fl. Mal. Pen. 1 (1922) 842. A. borneensis HALL. f. Med. Rijksherb. 42 (1922) 13; HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 492; RipL. Kew Bull. (1938) 112. A. borneensis var. microcarpa HALL. f. Med. Rijksherb. 42 (1922) 16. A. palmata (non LAMK) STEEN. Bull. Jard. Bot. Btzg. III, 12 (1932) 165. Leaves thinly to thickly coriaceous, pale green to grey-green beneath, entire or shallowly lobed, suborbicular or ovate to lanceolate, top rounded to acute, up to 1 cm acuminate, base acute-acumi- nate to broadly rounded, 4-21 by 114-12 cm, 3(—5)—plinerved and 1-2 pairs of lesser nerves from the midrib, or + pinninerved; petiole 14—-6(—7!/4) cm. Glands at blade-base confined to the auricles; submarginal glands (O—)1—2(—-3) pairs; marginal glands 5-25 at either side. Inflorescences peduncled up to 14 cm. Fruit globular to ovoid, excluding the (10—)15—30 mm long gynophore (2—)3-6(-614) by (1144-)2-5 cm; pericarp thickly coriaceous to woody, (J4-)1-3 mm thick. Seeds 7-10 mm g. Distr. Malesia: Sumatra, Malay Peninsula, W. & Central Java, Borneo. Fig. 9: 3a. Ecol. Primary and secondary forest, peat swamp forest, swamp edges; recorded from sand, loam, ‘red’ soil, sandstone, sandy loam soil with lime, rich yellow soil, podsolized sands, peat; 0—1000 (-1500) m. FI. fr. mostly April—Dec. Note. Some small-fruited, pale-leaved speci- mens from Java resemble var. singaporeana. b. var. smilacina (HALL. f.) DE WILDE, Thesis (1971) 230, f. 34. — A. smilacina HALL. f. Med. Rijksherb. 42 (1922) 17; RrpL. Kew Bull. (1938) 125 Leaves thinly coriaceous, pale green beneath, entire, suborbicular to ovate, top acute, 14-1 cm acuminate, base broadly rounded, mostly shortly acuminate, 6-12 by 3-9 cm, 3—5—(sub)plinerved; petiole 2-7 cm. Glands at blade-base + extending beyond the auricles on the blade; submarginal glands 0; marginal glands c. 5 at either side of the blade. Inflorescences peduncled for (1—)6—-13 cm. Fruit globular to ovoid, excluding the c. 10 cm long gynophore c. 2-3 by 114-2 cm; pericarp coriaceous, 14-1 mm thick. Seeds 5-7 mm @. Distr. Malesia: Sarawak and NE. Borneo. Fig. 9: 3b. Ecol. Forests and forest-edges, secondary for- est; rich yellow soil; low altitude. Fr. July—Oct. Notes. The leaves resemble those of A. kina- baluensis. The fruits are reported as sea-green turning bright red. c. var. singaporeana (WALL. ex G. DON) DE WILDE, Thesis (1971) 231, f. 34. — Passiflora singaporeana WALL. [ Cat. (1829) n. 1232, nomen] ex G. Don, Gen. Syst. 3 (1834) 55; StEuD. Nom. ed. 2, 2 (1841) 276 (‘sengaporeana’); Mast. Trans. Linn. Soc. 27 (1871) 631. — Anthactinia singaporeana Roem. Syn. Mon. 2, Pepon. (1846) 192. — Mo- decca singaporeana MaAstv. in Hook. f. Fl. Br. Ind. 2 (1879), 601: Rip. Jo. Sti. Br. ik. AS, Soc: 33 (1900) 87. — A. singaporeana ENGL. Bot. Jahrb. 14 (1892) 376; KiNG, J. As. Soc. Beng. 71, ii (1903) 55; RIDL. Fl. Mal. Pen. 1 (1922) 841; HEyNneg, Nutt. Pl.’ (1927) 1142; Burk. Dict. 1 (1935) 48; KENG, Ord. & Fam. Mal. Seed Pl. (1969) 76, f. 41. Leaves thickly coriaceous, grey-green to whitish green beneath, entire, (obovate-)oblong to lanceo- late, top acute, up to 14 cm acuminate, base acute to rounded, 5-15 by 134-7(-8) cm, faintly 3—pli- nerved and (1—)2-4 pairs of nerves from the midrib; petiole 14-214 cm. Glands at blade-base confined to the auricles; submarginal glands 0—1(—2) pairs; marginal glands 3-10 at either side of the blade. Inflorescences peduncled for up to 6 cm. Fruit (ovate-)oblong, + fusiform, excluding the (7—)10- 25 mm long gynophore 214-6 by 114-214 cm; pericarp thickly coriaceous or woody, c. 1 mm thick. Seeds 6-9 mm @. Distr. Malesia: southern Malay Peninsula (Johore), Singapore. Fig. 9: 3c. Ecol. Forest edges; low altitudes. F/. Sept.— March, fr. Jan., July—Oct. 4. Adenia kinabaluensis DE WILDE, Thesis (1971) 225s Soe Liana up to 20 m. Leaves thinly-coriaceous, brownish when dry, entire, ovate to ovate-elliptic, top acute, 1-114 cm acuminate, base cordate to broadly rounded, 6-14 by 3144-11 cm, 3(—5)-pli- nerved and with 1 (—2) pair(s) of strong nerves from near the base of the midrib, arching towards the top; petiole (1!4—)2-6 cm. Glands at blade-base 2, elliptic to reniform, 2-4 mm long, mainly on two auricles at the apex of the petiole but extending on the blade to or beyond the insertion of the basal nerves; blade-glands 0; marginal glands mi- nute, 0—S5 at either side of the blade. Inflorescences peduncled for 2-12 cm, in d up to 30—flowered, in 2 2-5-flowered; tendrils 1(—3), 14-3 cm. Sterile tendrils up to 15 cm. Bracts and bracteoles elon- gate triangular, acute, 14-1 mm. — ¢ Flowers urceolate, including the 1-114 mm long stipe 7-9 by 4(-5) mm. Hypanthium including calyx tube urceolate, fleshy, 4-5 mm. Calyx lobes elongate triangular, subobtuse, 2-214 mm, reflexed in an- thesis. Petals oblong-lanceolate, subobtuse, c. 3 mm, inserted near the throat of the calyx tube. Filaments 1-114 mm, free, inserted at the base of the hypanthium. Anthers 3-4 mm, acute, 0.1—0.3 mm apiculate. Septa 0. Corona 0. Disk glands c. 1 mm. — & Flowers not known. Fruit 1-2, globose, 430 FLORA MALESIANA [ser. I, vol. 7? excluding the 20-30 mm long gynophore 314-4 by 314-4 cm; pericarp woody, (2—)3 mm. Seeds c. 15, suborbicular, 8-10 by 8-9 by 2-2!4 mm, pitted; embryo 7-8 mm; cotyledons ovate, deeply emargi- nate at one side, c. 714 by 7 mm. Distr. Malesia: N. Borneo (Sabah: Mt Kina- balu region). Fig. 8: 4. Ecol. Montane forest, 1500-1800 m. Note. The leaves resemble those of A. macro- phylla var. smilacina and certain broad-leaved forms of var. macrophylla, but differ by the usually cor- date base, the basal glands which extent onto the blade, and by the brown colour and different tex- ture when dry. The 3 flowers are smaller. The fruits resemble thick-valved forms of A. macrophylla var. macrophylla. 5. Adenia cordifolia (BL.) ENGL. Bot. Jahrb. 14 (1891) 376; HARMs in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 84; ibid. ed. 2, 21 (1925) 490; Koorb. Exk. Fl. Java 2 (1912) 637; HALL. f. Med. Rijks- herb. 42 (1922) 11; Heyne, Nutt. Pl. (1927) 1142; Merr. Un. Cal. Publ. Bot. 15 (1929) 210; RIDL. Kew Bull. (1938) 112; MASAM. En. Phan. Born. (1942) 506; BAck. & BAKH. f. Fl. Java 1 (1963) 289; CusseT, Adansonia 7 (1967) 372, 383; DE WILDE, Thesis (1971) 232, f. 34, 36. — Modecca cordifolia Bu. Bijdr. (1826) 939; DC. Prod. 3 (1828) 336; G. Don, Gen. Syst. 3 (1834) 59; BL. Rumphia 1 (1837) 167, t. 49 f. 1-7; Hassx. Cat. Hort. Bog. (1844) 187; Mia. Fl. Ind. Bat. 1, 1 (1855) 702. — Microblepharis cordifolia ROEM. Syn. Mon. 2, Pepon. (1846) 202. — Fig. 4. Modecca obtusa BL. Bijdr. (1826) 939; DC. Prod. 3 (1828) 336; G. Don, Gen. Syst. 3 (1834) 59; BL. Rumphia 1 (1837) 166, t. 48 f. 1-10; HAssk. Cat. Hort. Bog. (1844) 187; Mig. Fl. Ind. Bat. 1, 1 (1855) 702. — Microblepharis obtusa ROEM. Syn. Mon. 2, Pepon. (1846) 200. — A. obtusa ENGL. Bot. Jahrb. 14 (1891) 376; Harms in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 84, f. 30 A-E; ibid. ed. 2, 21 (1925) 489, 492, f. 223: Koorp: Exk: FI Java 2 (1912) 637; HALL. f. Med. Rijksherb. 42 (1922) 11; STEEN. Acta Bot. Neerl. 15 (1966) 41. A. populifolia var. pentamera KiNG, J. As. Soc. Beng. 71, ii (1903) 54; RIpDL. Fl. Mal. Pen. 1 (1922) 841. A. quadrifida MERR. Philip. J. Sc. 13 (1918) Bot. 94; En. Born. (1921) 413; HARMS in E. & P. Nat. Pfl. Fam. ed. 2, 21 (1925) 492; MAsam. En. Phan. Born. (1942) 506. A. vespertilio HALL. f. Med. Rijksherb. 42 (1922) 8; MASsAM. En. Phan..Born. (1942) 506. A. sp. BARTLETT, Pap. Mich. Ac. Sc. 6 (1926) 31. A. populifolia (non BL.) RipL. Kew Bull. (1926) 66; ibid. (1938) 112; HEND. Gard. Bull. S. S. 4 (1928) 264; Burk. Dict. 1 (1935) 48; MASAM. En. Phan. Born. (1942) 506. Liana to 20(—50?) m. Leaves herbaceous to subcoriaceous, pale to glaucous green beneath, entire, broadly ovate to oblong, top obtuse to acute, up to | cm acuminate, base rounded to deeply cordate, 214-10(-17) by 114-6(-9) cm, 3—5—plinerved and with 2-10 pairs of lesser nerves from the midrib, arching towards the top; petiole 14—3(—414) cm. Glands at blade-base 2, 1-2'4 mm @, in two deeply hollowed hemispherical auricles 214-414 mm @ at the apex of the petiole; blade- glands 0-6, 14-1 mm @, submarginal; marginal glands minute, 0-8 at either side. Inflorescences peduncled up to 514 cm, in 3 up to 60-flowered, in 2 3—S—flowered, tendrils 0-3, 14-214 cm. Sterile tendrils 3—5(—7)-fid, up to 10 cm, sometimes ending in adhesive disks. Bracts and bracteoles narrowly triangular, acute, 14-1 mm. — 39 Flowers narrowly tubiform-urceolate, including the 10-20 mm long stipe 18-35 by 1!4-3(—4) mm. Hypanthium includ- ing calyx tube tubiform-urceolate, + narrowed at the throat, fleshy leathery, 8—12(—14) mm. Calyx lobes elliptic-oblong, obtuse to subacute, 1-2(-214) mm, (sub)erect in anthesis. Petals oblong-lanceo- late, obtuse to subacute, 1-2 mm, inserted in the throat of the calyx tube. Filaments (1!4—)2—5 mm, connate for 1-2 mm, inserted at the base of the hypanthium. Anthers 3—4 mm, acute, up to 144 mm apiculate. Septa 1-2 mm high. Corona 0. Disk glands 1-2 mm. — 2 Flowers tubiform(-urceolate), including the 4-10 mm long stipe 12-18 by 214-3 mm. Hypanthium including calyx tube c. 7 mm. Calyx lobes c. 1144 mm. Petals c. 1'4 mm, inserted at or near the throat of the calyx tube. Staminodes 1-114 mm, + connate at base. Septa c. 44 mm high. Corona 0. Disk glands c. 1 mm. Pistil 5-6 mm; gynophore c. 1 mm; ovary ellipsoid c. 4-414 by 2'4 mm; styles c. 14 mm, free; stigmas + papil- late, each c. 1 mm 9. Fruit 1-2, ellipsoid-oblong, fusiform, -- 3-angular, top acute, up to 1 cm acuminate, excluding the 5—10(-15) mm long gynophore (4!4-)5—8(-9) by 114-314 cm; pericarp woody-coriaceous 1-1!4 mm. Seeds 10-30, orbi- cular to reniform, 7-814 by 8-10 by 4-414 mm, pitted; embryo 7-9 mm; cotyledons ovate-elliptic, 5 @ Adenia cordifolia 6 A Adenia crassa Fig. 10. Distribution of Adenia cordifolia (BL.) ENGL. and A. crassa MERR. 1972) PASSIFLORACEAE (de Wilde) 431 broadly emarginate at one side towards the top, 7-9 by 514-614 mm. Distr. Malesia: Sumatra, Malay Peninsula, W. Java, Borneo, Philippines (Palawan: MERRILL 806, sterile). Fig. 10: 5. Ecol. Rain-forest, thickets, forest clearings, peat swamp forest; 0-1200 m. Fi. fr. Jan.—Dec., mostly fl. Sept.—March, most fr. Nov. Because of the narrow-urceolate flowers, entire- ly enclosing either the stamens or the pistil, pollination is likely effected by small insects (moths?). Uses. The roots and fruits are known as poison- ous; leaves and stems have medicinal properties. According to HEYNE the branches are resistant to humidity and therefore used as binding material under water. Vern. Akar kail, M, layang-layang (Kedayan), Mal. Pen.; andor loting, Asahan; areuj babaling- bingan, a. patok manok, a. tjalingtjing, S. Notes. Most records from the Philippines concern the related A. crassa. Juvenile forms are often found creeping on open places, on tree trunks and rocks, and are provided with typical lobed or lunate leaves with more or less peltate blade-base; A. vespertilio is a juvenile from with lunate leaves. Occasionally 4—merous 3 flowers are found; once a flower with a 4—carpellate pistil; once a ° flower with 8 staminodes. A. quadrifida is based on a specimen in which part of the flowers have 4 calyx lobes. Modecca cordifolia and M. obtusa date both from 1826. M. obtusa was listed first in the synony- my of A. cordifolia by KOORDERS, 1912. Often galled flowers or slightly deformed flowers with an insect larva within are found. Field notes. Fresh flowers are reported as pale greenish to yellow, when dry they are often inside reddish brown spotted; fresh ripe fruits are bright glossy red. 6. Adenia crassa MerR. Philip. J. Sc. 10 (1915) Bot. 331; En. Philip. 3 (1923) 117; DE WILDE, Thesis (1971) 235, f. 34. — A. quadrifida (non MERR.) Merk. En. Philip. 3 (1923) 117. Climber to 10 m. Leaves herbaceous, entire or shallowly toothed in the lower half, ovate-elliptic to oblong, top longly acute or faintly acuminate up to 3(—5) cm, base cordate, 3-15 by 114-914 cm, 3—5—plinerved and 1-3 pairs of lesser nerves from the midrib; nerves often -- reddish tinged; petiole ¥%-3'\%4 cm. Glands at blade-base 2, 1-214 mm @, in two deeply hollowed auricles 214-6 mm @ at the apex of the petiole; blade-glands 0-2, c. 14 mm 9, submarginal; marginal glands minute, 0-5 at either side. Inflorescences peduncled for 114-6 cm, in 3 up to 30-flowered, in 2 2—5-flowered; tendrils 0, or 1 or 3, '4-2'4 cm. Sterile tendrils up to 10 cm, in juvenile forms 3—fid, ending in small adhesive disks. Bracts and bracteoles narrowly triangular to oblong, acute, 14-1 mm. — 9 Flowers narrowly tubiform-urceolate, including the 9-10 mm long stipe 16-18 by 214-3 mm. Hypanthium including calyx tube narrowed to the throat, rather fleshy (not leathery), 6-7 mm. Calyx lobes elongate triangular, subacute, 1-114 mm, suberect in anthe- sis (not reflexed). Petals elongate triangular, acute, 1-1!4 mn, inserted at the throat of the calyx tube. Filaments c. 3 mm, connate for 1 !4—2 mm, inserted at the base of the hypanthium. Anthers 3-314 mm, obtuse. Septa c. 1'4 mm high. Corona 0. Disk glands 1-114 mm. — 2 Flowers + urceolate, in- cluding the 2!4-3 mm long stipe 8-10 by 3 mm. Hypanthium including calyx tube 4414 mm. Calyx lobes triangular, subobtuse, c. 1 mm. Petals oblong, c. 144 mm, inserted at the throat of the calyx tube. Staminodes c. 1 mm, connate at base for c. 4% mm. Septa c. 14 mm high. Corona 0. Disk glands c. 14 mm. Pistil c. 5% mm; gynophore c. 1 mm; ovary subglobose, c. 3 by 244 mm; styles connate for c. 14 mm, style-arms c. 14 mm. Stig- mas papillate-laciniate, each c. 1 mm @. Fruit 1, subglobose, excluding the 10-15 mm long gyno- phore 5-6 by 414-5 cm; pericarp woody-coria- ceous, inside + spongy, 1!4-2!4 mm thick. Seeds 30-40, suborbicular, c. 714 by 714-8 by 214-3 mm, pitted; embryo c. 7 mm; cotyledons ovate, broadly emarginate at one side towards the top, c. 6'4 by 514 mm. Distr. Malesia: Philippines (Sulu Is., Basilan I., W. Mindanao: Zamboanga Prov.). Fig. 10: 6. Ecol. Forest and forest-edges; 0-500 m. Fi. Sept.—Dec., fr. Aug. & Jan. Vern. Sabugok, Basilan I. Uses. According to GORDON (PNH 82005) the fleshy arils are edible. Notes. Related to A. cordifolia, but distinguish- ed by the stronger 3—S—plinerved leaves and the subglobose, thick-valved fruits. Leaves of juvenile specimens are deeply 3—lobed with a much reduced middle lobe, and the base + peltate. Field notes. Fresh flowers are pale (greenish) yellow, when dry finely purple-red spotted; fresh fruits are shining red. 3. HOLLRUNGIA K.Scu. Bot. Jahrb. 9 (1888) 212; BoeRL. Handl. | (1890) 571; Harms, Bot. Jahrb. 15 (1893) 578 (anat.); in E. & P. Nat. Pfl. Fam. 3, 6a (1893) 79, 86; ibid. ed. 2, 21 (1925) 484, 494; STEEN. Reinwardtia 1 (1952) 480; Acta Bot. Neerl. 15 (1966) 40— 44, f. 1-7; Hutcu. Gen. FI. Pl. 2 (1967) 370. Liana. Leaves simple, entire, pinninerved; petiole with or without glands; stipu- 432 FLORA MALESIANA [ser. I, vol. 7? Z 7 Ey Fig. 11. Hollrungia aurantioides K. Scu. a. Habit of 2 specimen, x %, b. detail of node, showing axil- lary tendril and serial, supra-axillary ramification, x %, c. ¢ flower in longitudinal section, «4%, d. 2 flower in longitudinal section, x 4%, e. fruit, x %, f. seed, x2%(a, d BSIP 4071, b Brass 3283, c Brass 27551 e KasEwskI 1915, f KAsEwSKI 1913). 1972] PASSIFLORACEAE (de Wilde) 433 les minute. Inflorescences axillary or slightly supra-axillary, shortly peduncled, 1-15-flowered, mostly without a tendril. Bracts minute. Flowers polygamous (bisexual or functionally unisexual). Hypanthium saucer-shaped. Sepals and petals free; sepals 5, greenish; petals 5, resembling the sepals, white, membranous. Corona double; outer corona consisting of 2(-3) rows of filamental appendages, inner corona an erect tube with complicatedly folded, curved and laciniate edge; nectary ring or disk obscure, at the bottom of the hypanthium. Androgynophore 14-2 mm. Stamens 5, filaments free or connate to more than halfway into a tube enveloping (not connate with) the gynophore. Anthers ellipsoid-sagittate, blunt, + dorsi- fixed, + versatile. Gynophore 114-6 mm. Ovary subglobose to ellipsoid; styles 3, c. Y, mm, free; stigmas lobed-papillate. Fruit globose, pericarp coriaceous to woody. Distr. Malesia: monotypic, from the Moluccas eastwards to the Solomons. Ecol. Rain-forest, from the lowland up to c. 1700 m. Taxon. Distinctly related to the monotypic genus Tetrapathaea, from New Zealand, which differs by 4-merous flowers and the absence of a separate inner corona. 1. Hollrungia aurantioides K. Scu. Bot. Jahrb. 9 (1888) 212; Harms in E. & P. Nat. Pfi. Fam. 3, 6a (1893) 86, f. 25 E-F; ibid. ed. 2, 21 (1925) 495, f. 218 E-F; K. Sco. & Hoitr. FI. Kais. Wilh. Land (1889) 82; K. Scu. & Laut. FI. Schutzgeb. (1901) 456; MeRR. & PERRY, J. Arn. Arb. 24 (1943) 210; ibid. 29 (1948) 160; ibid. 30 (1949) 44; STEEN. Reinwardtia 1 (1952) 480; Acta Bot. Neerl. 15 (1966) 40-44, f. 1-7. — Passiflora moluccana (non BL.) MERR. & PERRY, J. Arn. Arb. 30 (1949) 44. — Fig. 11. Liana to 45 m. Serial buds distinct, with acute bud-scales (cataphylls) 14-1 mm. Leaves subcoria- ceous, elliptical to lanceolate, top subobtuse to acute, up to 1 cm acuminate, sometimes 1-2 mm mucronate, base rounded to acute-acuminate, 7-23 by 3-12 cm, pinninerved; nerves 4~6(-8) pairs, arching to above, the basal nerves weaker; petiole 114-4 cm. Glands on lamina 0 or 1-—2(-6), mostly + paired, either at the very blade-base, or halfway the lower nerves or scattered, large but often inconspicuous, flat, roundish or irregular in shape, 2-6 mm 9; glands on petiole 0 or 1(-2) pairs, inserted halfway or mostly atc. 44 (and 2% from the base, 14-2 mm 9. Inflorescences axillary or up to 5 mm supra-axillary, 1-10(—15)—flowered; peduncles 14-6 cm, pedicels 1—-5(—10) mm; bracts and bracteoles narrowly triangular, acute, 14-1 mm. Tendrils 0 or rarely the central flower re- placed by a 1-5 cm long tendril; sterile tendrils 10-15 cm, sparse, in the axils of leaves. — ¢ Flow- ers 15-20 mm 9; stipe 14-20 mm. Hypanthium saucer-shaped, 3-5 mm wide. Sepals lanceolate, 8-10 by 3-4 mm, obtuse.Petals 7-9(-10) by 3-314 mm, (sub)entire, obtuse. Corona double; outer corona + spreading, consisting of densely set appendages in 2(-—3) rows, 2-9 mm, the outer longest; inner corona an erect membranous tube with complicatedly folded and inward curved much divided (laciniate) upper %, 214-4 mm high, 3—S(-6) mm wide. Disk entire, flat or some- times irregularly wrinkled, covering the bottom of the hypanthium, 2-3!4 mm 9, c. 2 mm thick. Androgynophore 14-2 mm. Filaments flat, 5—6 mm, free or up to 4 mm connate into a tube; anthers elliptic, obtuse, (1!4—)2-3 by (1-)11%4-2%4 mm, the filaments dorsally attached to the fila- ments, but anthers sagittate and toppled over in anthesis. Gynophore 2-6 mm; vestigial ovary oblong, + 3—angular, smaller than the anthers, 1-114 by 4-25 mm, vestigial stigmas sessile, + brush-shaped, each c. 4 mm 9, or as a single irregular disk, c. 1 mm 9. — § Flowers as 3 flowers but ovary ellipsoid, subcircular in cross-section, 114-214 by 1-2 mm. Styles c. % mm; stigmas papillate, each 14-1 mm @. — 2 Flowers as 3 and 3 flowers or smaller, 10-20 mm 9; stipe 8-18 mm. Sepals and petals 514—-8(—10) mm. Outer corona filaments 114-5 mm, inner corona 2-3 mm high; disk as in 3 flowers. Androgynophore 14-114 mm. Filaments 2-4 mm, free or up to 1 mm connate; vestigial anthers 1-114 by 14(-1) mm. Gynophore 1-2 mm; ovary ellipsoid to subglobose, 11-2 by 1-2 mm; styles c. 14 mm; stigmas papillate, each 1-2 mmo. Fruits 1-2, globose, excl. the (5—)10 mm long gynophore 2-444 cm @; pericarp thickly coriaceous to woody, 1-4 mm thick, greenish yellowish when fresh, sometimes + fleshy inside. Seeds 15-50 ellipsoid-obovate, 514-7 by 4-6 by 2-3 mm, 5-7 pits or grooves 9; embryo 4-6 by 314-514 mm; cotyledons obovate, subtruncate, 3(—5)-plinerved; radicle c. ?/; mm. Distr. E. Malesia: Moluccas (Ternate), W.—E. New Guinea, New Britain (Wiriai Subdistr.), New Ireland (inland from Lavongai), Misima I., Solomon Is. (Bougainville, Shortland, Ronongo, New Georgia, Santa Ysabel, Santa Ana, Rennell). Fig. 12. Ecol. Primary and secondary rain-forest, lo- cally common; 0-1700 m. Fi. fr. Jan—Dec. Vern. Aa, Vogelkop (Maibrat lang.). Notes. Polygamous, apparently largely (func- FLORA MALESIANA [ser. I;-walkk7* Hollrungia aurantioides Fig. 12. Distribution of Hollrungia aurantioides K. SCH. tionally) dioecious. The 3 and & flowers are usually somewhat larger than the 2 flowers. In ¢ flowers always a distinct vestigial ovary is present, in © flowers stamens with distinct, reduced, sterile anthers. In one specimen (CLEMENS 5435) some of the 2 flowers contained beside 4 reduced anthers one well-developed fertile anther. In the flowers a short androgynophore is always present; the filaments are either free or to a various degree connate into a tube enveloping the gyno- phore. The variability in the flowers as well as in the size of the fruits and the place and presence or absence of glands suggested that several taxa might be involved (VAN STEENIS, 1966); a thorough investigation of the flowers of the rather abundant recently collected specimens, however, proved that all the material belongs to a single species. In the original description the stigma was erro- neously described as single, undivided, cap-shaped, but it appeared (VAN STEENIS, 1966) that this was due to the young stage of the flowers in the type material. Australasian Passifloras differ by the absence of a distinct gynophore and by the globular er club- shaped stigmas. In Adenia the corona is absent or composed of but a single row of short hairs, where- as the disk is composed of 5 separate lingulate or strap-shaped appendages; the anthers are nar- row; petiolar glands are absent or restricted to auricles at the very top of the petiole. Field notes. The bark is green with brown corky lenticels, in old specimens dark brown, fissured. Slash: wood soft whitish cream, bark soft, reddish brown. Big sterile tendrils reported as present only on the main stems, not on the fruiting lateral branches. Leaves glossy. The flowers are greenish white or yellow-green; sepals pale green or greenish yellow, petals white; outer corona filaments whit- ish yellow; inner corona yellow-green, tipped yellow; filaments white or greenish; anthers yellow; ovary green; stigmas yellow-green. Once recorded with faint smell, once with nasty smell (¢ fl.). The fruits are light green or yellowish green. 1 v 7) ae WO | 3 5185 00202 6928 FLORA MALESIANA under the Auspices of Lembaga Biologi Nasional (Botanic Gardens of Indonesia), Bogor, Indonesia, and the Rijksherbarium, Leyden, Holland, executed by Foundation Flora Malesiana Scientific Communications concerning the Flora Malesiana should be addressed to the General Editor, Dr. C. G. G. J. VAN STEENIS SUBSCRIPTION REQUEST AND RELATED CORRESPONDENCE SHOULD BE DIRECTED EXCLUSIVELY TO WOLTERS-NOORDHOFF PUBLISHING P.O. Box 58, Groningen, The Netherlands For sale only - Not for exchange Series I - Spermatophyta (Flowering Plants) Volume 1. Cyclopaedia of collectors & collections. 1950. pp. clii + 639 Volume 2. Malesian vegetation. (In preparation) Volume 3. Malesian plant geography (In preparation) Volume 4. General chapters and revisions. 1948-1954. pp. ccix + 631 Volume 5. Bibliography, specific delimitation, & revisions. 1955-1958. pp. cccxlii + 596 Volume 6. Systematic revisions. 6 parts, 1960-1972. pp. 20 + 1023. Volume 7. Systematic revisions. Parts 1-2. 1971-1972. pp. 434 ANI W OLTERS-NOORDHOFF PUBLISHING, GRONINGEN, THE NETHERLANDS PRINTED IN THE NETHERLANDS ISBN 90 01 31814 2