& Krukopy FLORA mu yo MALESIANA SERIES I- SPERMATOPHYTA Flowering Plants Vol. 9, part 1 Revisions ~ INDEX TO REVISED FAMILIES Aceracea@ . «9% ) 4: 3, 592 Actinidiaceae s. str. 4: 37 Aizoaceae.. 6s. ae ees 4: 267 Alismataceae S317 262905 Amaranthaceae 4: 69, 593; 6: 915 Anacardiaceae. . . . . 8: 395 Ancistrocladaceae . . 4: 8 Aponogetonaceae 4: 11; 7: 213 Araliacede-02! 2 t eoeae Lael | Balanophoraceae. . . .7: 783 Basellacede, |) 2) nice ies oe 5: 300 | Batidaceae: = (oo saesk 5: 414 Betulaceae 5: 207; 6: 917 Bignoniaceae ... . .8: 114 Bixacese's/sth.* 2 +s ce ag Burmanniaceae 7443 135592 Burseraceae 5: 209, 567; 6: 917; 7: 820 Butomaecede =s | ee 5: 118 Byblidaceaes.) Atoms we at 7 ils}s) Callitiehaceae = <1. aie 4: 251 Campanulaceae 6: 107, 928 Cannabinaceae ... .4: 223 Capparidaceacs.) ia ae 6: 61 Caprifoliaceae 4: 175, 598; 6: 928 Cardiopteridaceae . . .7: 93 Celastraceae 6: 227, 389, 930 - Centrolepidaceae . 5: 421 Ceratophyllaceae. . . .4: 41 Chenopodiaceae 4: 99, 594; 6: 932 Clethraceae ees S 12139 Cochlospermaceae . . .4: 61 Combretaceae 4: 533: 5: 564; 6: 932 Connaraceae 5: 495; 6: 933 Convolvulaceae 4: 388, 599; 5: 558; 6: 936; 7: 823 COMACEIEY “Pecks oe 8: 85 Corynocarpaceae A: 26272152 55) Crassnlaceae™ 7 isha 4: 197 Crypteroniaceae . .. .8:187 Cyperaceae . . . 7: 435; 9: 107 Datiscaceae (82h. Sar: 4: 382 Dichapetalaceae 45: 305; 6: 941 Dilleniaceae. . 4: 141; 7: 824 Dioscoreaceae..... 4: 293 Dipsacaceae as ee 4: 290 Droseraceae: = .. -453772)52 557 Blatinaceae foe cee 4: 203 Epachidaccae, fb ene 6: 422 Bricdceae) 222+, 2 162 4695943 Erythroxylaceae . . . 5: 543 Papackaee a har ee 7: 265 Ficoidaceaes «. Sehr 4: 267 Flacourtiaceae 5: 1, 565; 6: 943; 7: 827 Flagellariaceae seheagae, Feorte 4: 245 Geraniacede sient noe 6: 445 Gnetaceae. . . 4: 336; 6: 944 Gonystylaceaei=. 44. te . 4: 349 Goodeniaceae 5: 335, 567; 6: 949; 7: 827 Haemodoraceae.. 9 .5.. sada) Haloragaceae serie 73 239 Hamamelidaceae. . . . 5: 363 Hippocrateaceae . . . . 6: 389 Hydrocaryaceae . . .4: 43 Hydrocharitaceae 53381; 67952 Hydrophyllaceae. . . . 4: 207 IB hyoyotenrereuys A EP Ae Se IeACINACEAeH aan ates ea WidaCeAe a ape eae-t eres ona, Jugiandacede™<..) 20. 6: 143 JURGACEAE 7.00.5" hy sk 8 4: 210 Juncaginaceaey, w-cae lone. 4: 57 Eabiatae. geeican. neo ies SO IGeGACEAG. VA) ecg aes 7 (ES) Wemnaccden ae. ea eee 72219 Lentibulariaceae . . . .8:275 Lillaceae=l in: wee ees 9: 189 Loganiaceae . 6: 293, 953 Lophopyxidaceae .. .7: 89 Malpighiaceae. .... 5S: 125 Martyniaceae’ 205. . 4: 216 Molluginaceae 3. 2) ..). 4: 267 Moringacedeo. 95 ee 4: 45 Myoporacedae, 2 2. 4: 265 Myricaceaegren.. + Gieme 4: 277 INajadacede vey 9 0y.1 a 6: 157 Nyctaginaceae..... 6: 450 INyssacede Sie. err et ets 4: 29 Mehnaceae se ees! 7297 Onagracedesy 5.f wate. 8: 98 Oxalidaceaets si. sl)’ AAT! Papavetacede: a. siase 5: 114 ~ at —~ LIBRARY RHE NEW YORK S74) CAL GARDEN ~ BRONX, NEW YORK 10458 by 8. keute| WITH THE COMPLIMENT FROM J. VAN STEENIS. Di. C.G.G~ Passifloraceae: 1). 7: 405 Pedaliaceae) ee 4: 216 Pentaphragmataceae . . 4: 517 Pentaphylacaceae nope ha PA Philydraceaeuint eee a0 5 Phytolaccaceae . 1 aD) Pittosporaceae.s)2° rae 5: 345 Plumbaginaceae . . . . 4: 107 Podostemaceae 4: 65; 6: 963 Polemoniaceae. .... 4: 195 Pontedeniaccae ara 4: 255 Portulacaceae™ wee TA Primulaceae*. eee 6: 173 Proteaceaes <>. eee 5: 147 Punicaceae: : came 4: 226 Restionacede) y.0.eee 5: 416 Rhizophoraceae 5: 429; 6: 965; 8: 550 - SalicdaGeaceric ie ieee 5: 107 Salvadoracedeu: a caaaee 4: 225 Sarcospermaceae. . . .4: 32 Saururaceaer: =.aee eee 4: 47 Scyphostegiaceae : 5: 297; 6: 967 Simaroubaceae 6: 193, 968 Sonneratiaceae 4: 280, 513; 6: 973 Sparganiacede) = onan 4: 233 Sphenocleaceae A] Stackhousiaceae . . . .4: 35 Staphyleaceseaw ase 6: 49 Stylidiacede" i= eee 4: 529 Styracaceaeui.) te aamcereune 4: 49 Symplocaccaci sa. samen 8: 205 Taccacedes.. sa. ee 7: 806 Thymelaeaceae 4: 349; 6: 1, 976; 7: 830 Trapaceaein..:) taste 4: 43 Trigoniacéde eae 4: 59 Turneraceae. cane ane 4: 235 Typhacedés 4) aes ee 4: 243 Ulmaceae; 3 eee Si Umbelliferae 4: 135 595: 5: 555; 6: 983; 7: 830 Valerianaceae' . 3.7). 4: 253 Violacéae’. ..s) 2 dele eo Xyridaceae 4: 366, 598 Zygophyllaceae 4: 64 24(r] x B.A. Kruko@ P.0. Box 352 ARALIACEAE—I (W. R. Philipson, ChristcM{0#tgwn. N.Y. 11787 Trees, shrubs, lianas, woody epiphytes or (extra-Mal.) more rarely herbs. Branches usually stout with leaves clustered at their ends; armed or unarmed; glabrous or with a tomentum of stellate or simple hairs; buds either covered by the stipular sheaths of leaves or by cataphylls. Leaves spiral or rarely opposite or in whorls; petiole usually clasping the stem; stipules either distinct or united into a ligule or absent (in Osmoxylon the petiole bears + elaborate crests around its base); lamina digitately compound or pinnate, sometimes to the second or third degree, or simple, when either entire or pinnately or palmately lobed, margin entire or dentate. Inflorescence terminal or more rarely lateral; either simple or compound racemes or spikes, or more commonly of umbels or capitula, either solitary or arranged in compound umbels or panicles; bracts usually small and caducous; pedicel either articulated with the flower or continuous with it. Flowers hermaphrodite or hetero- sexual, sometimes dioecious; actinomorphic. Calyx lobes small, or reduced to a rim, or rarely absent. Petals 3 to numerous, often 5, sometimes fused into a calyptra, or forming a tube with spreading lobes (Osmoxylon), valvate or imbricate in bud, usually with a broad base but rarely narrowed below. Stamens usually as many as the petals and alternating with them, or twice as many, or indefinite; filaments inserted at the edge of the disk; anthers dorsifixed, introrse, pollen sacs 4 or rarely 8. Ovary inferior, half inferior, or very rarely (extra-Mal.) superior, 1- to many-celled, the top of the ovary usually a fleshy disk; styles and stigmas as many as the cells, either connate or wholly or partially free. Ovules solitary, pendulous, anatropous, with the raphe ventral. Fruit baccate or drupaceous, exocarp usually fleshy, endocarp forming cartilaginous or membranaceous pyrenes around the seeds. Seeds one per pyrene, with a small embryo within smooth or ruminate endosperm. Distribution. About 50 genera with a roughly estimated 1150 species, ranging mainly in the warmer parts of both hemispheres (especially in montane zones), a small number in or extending to cool-temperate regions. With the exception of SE. Asia, the family and its centres of distribu- tion are largely found within the land masses derived from ancient Gondwanaland. In Malesia 17 genera with a total (excluding Schefflera) of 117 species in 16 genera. (The largest genus, Schefflera, with an estimated 250 species for the region, is omitted from this account.) Three genera are endemic to Malesia (or nearly so). One of these, Anakasia (related to Polyscias) is of very local distribution in West New Guinea; the two others, Harmsiopanax (Java, Lesser Sunda Is., Celebes, to New Guinea) and Aralidium (Malay Peninsula, Sumatra and Borneo, with an extension to Thailand) are more widely distributed. Five other genera have their main centres of distribution within Malesia. Two of these extend further eastwards: Mackinlaya ranges from the Philippines and Celebes through New Guinea to the Solomon Is. and NE. Australia; Osmoxylon occurs from Botel Tobago (Taiwan) and the Marianas to Borneo, the Philippines, Celebes, the Moluccas, New Guinea to the Solomon Is. and the New Hebrides. A third, Trevesia, is confined to western Malesia (Lesser Sunda Is., Java, Borneo, Sumatra, and Malaya), with an extension into the Asian mainland. Wider ranges are recorded for Gastonia (widespread in Malesia to the Solomon Is.) with an additional range in the Seychelles, Mascarenes, and Madagascar (but not East Africa as previously reported) and Arthrophyllum (all over Malesia with extensions to Thailand, Laos, the Nicobar Islands, and New Caledonia. A significant element in the Malesian representation of the family consists of six genera which occur mainly in SE.-E. Asia, two of which further extending to the Americas, viz Aralia (in (1) The genus Schefflera is omitted and will be treated separately by Dr D. G. Frodin (University of Papua New Guinea). I enjoyed his assistance in drawing the general chapters. (1) 2 FLORA MALESIANA [ser. I, vol. 9? America south to Mexico) throughout Malesia, and Dendropanax (tropical America) in West Malesia (Sumatra, Malay Peninsula, Borneo). The remaining four genera range in Malesia as follows: Macropanax and Brassaiopsis in Malay Peninsula, Sumatra and Java, Acanthopanax in Malay Peninsula, Sumatra and the Philippines, and Pentapanax only in East Java. Only one genus, Delarbrea, has its main centre of distribution to the east of Malesia (Melanesia, mostly New Caledonia) and Queensland, and extends through New Guinea to the Moluccas and Lesser Sunda Is. as far west as Timor. Two other genera have wider distributions. Schefflera, including several segregate genera, is pantropical although with but few representatives (13) on the African mainland. However, the several sections of Schefflera as redefined by FRopIN (1970, 1975) often show distinctive regional distributions paralleling those of several of the other genera. The type section ranges from the New Hebrides to Samoa and New Zealand, including Fiji and New Caledonia. Polyscias is widely dispersed in the Old World tropics from the African mainland eastwards to the Society Is. and Australia, but with only a few species in SE. Asia and western Malesia. As in Schefflera, the diverse series of species comprising the genus show distinctive regional distribution patterns, with the type series being mainly Melanesian and East Malesian (in West Malesia it is only cultivated or adventive). As said above several Malesian genera extend into, or have their main centres in, mainland Asia, but only Polyscias (in Sri Lanka) and Schefflera are shared with Africa and the same genera (with Gastonia) with Madagascar. Only Polyscias and Gastonia occur in the Mascarene Islands and Gastonia and Schefflera in the Seychelles. Two other genera are restricted to mainland Africa. Hedera of temperate Eurasia extends to the Canary Islands. The Americas have two (or three) distinctive endemic genera with recognizable affinities, as well as disjunct groups of the Austral- asian Pseudopanax, the Asiatic Pentapanax and Dendropanax and (in North America) Oplopanax, Aralia and Panax, all very closely related to those in East Asia; there are also distinctive sections of Schefflera in the neotropics which are nearer those in Africa than in Asia. Many distinctive endemic genera (or parts of the larger genera) occur in Oceania, the New Zealand region and Australia, some of them taxonomically very isolated. The ratio of species : genus is nere estimated as 23 : 1, but if the very large genus Schefflera is not accounted for this reduces to 13 : 1. Some 30 genera have five or fewer species; in view of the considerable insular endemism at generic or infrageneric level, weak intercontinental links, and great distinctiveness of many genera (even though small), the family is surely of great antiquity, although much of the available palaeobotanical evidence requires re-evaluation (cf. DILCHER & Do.pu, 1970). Many fossils previously referred to Oreopanax must now be placed in Platanaceae (DoyLe, pers. comm.). Dendropanax has reliably been reported from Tertiary deposits in both Europe and North America where it is now absent (DILCHER & DOLPH, /.c.). References: DiLcHER & Dotpu, Amer. J. Bot. 57 (1970) 153-160; Fropin, The complex of Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K. (1970); J. Arn. Arb. 56 (1975) 427-448. Ecology. Malesian Araliaceae are usually small trees, shrubs, or lianas, with a number being sometimes or always epiphytic (especially in Schefflera), and where terrestrial usually in the under- growth or lower stories of rain-forest, seldom reaching over 20 m. A remarkable exception is provided by the two species of Gastonia; of these G. spectabilis (HARMS) PHILIPSON of Papuasia can attain the great height of 40 m with a stem of 1.75 m @; it is the largest araliad known and is of very striking appearance. Fig. 27. Araliads are almost always found scattered in forest and other vegetation, at least in Malesia. However, a notable exception is provided by the tree Schefflera rugosa (BL.) HARMS in Java; Lam (1924) recorded its gregarious occurrence on the volcanic cone of Mt Slamet in Central Java where it is co-dominant with the pyrophilous A/bizia lophantha BTH. in elfin forest between 2500- 3050 m, above which it gives way to the open, rocky, treeless slopes below the summit (3428 m). It is also gregarious on Mt Tjeremai in West Java, where Lam (1925) noted that above some 2650 ma low forest dominated by this species replaces the high forest of Dacrycarpus imbricatus ; this low forest extends to c. 3000 m. On the Gedeh-Pangrango complex above Puntjak Pass in West Java Schefflera rugosa is frequent in forest borders. On Mt Ulu Kali in Malaya, east of Kuala Lumpur, S. nervosa (KiNG) Vic. is common in young regrowth along the main road 1979] ARALIACEAE—I (Philipson) 3 below the Genting Highlands hotel/casino complex from 1300-1600 m. This pioneering tendency is shared by a number of other species in the genus, especially in montane parts of Papuasia. Many of these species are terrestrial as pioneers or in secondary formations, but epiphytic in closed forest (and then often much more scattered). Gregarious occurrence, usually in pioneering situations or in forest borders, has also been observed in the various species of Harmsiopanax ; STEUP (1938) observed H. aculeatus (BL.) WARB. ex BOERL. as a characteristic pioneer in grass thickets on hills in SW. Celebes. H. harmsii K. Scu. behaves similarly around Wau, Bulolo, and Sogeri (Rouna) in Papua New Guinea, especially in narrow intermontane valleys, while H. ingens PHILIPSON can be locally abundant in open situa- tions in the highlands of New Guinea. Gastonia spectabilis (HARMS) PHILIPSON can be locally frequent as a pioneer in hill areas, e.g. around Bulolo and on the Madang-Ramu Divide; while Polyscias elegans (C. Moore & F.v.M.) HARs is frequently seen in stable monsoon scrub and forest borders in the Port Moresby region and P. /edermannii HARMS can be an exceedingly com- mon regrowth tree in cut-over montane forest subject to frequent cloudiness and rain. In the subalpine zone of New Guinea Brass (1941) found two species of Schefflera conspicuous in the stunted forest of Xanthomyrtus-Vaccinium-Papuacedrus-Phyllocladus between 3200-3800 m on Mt Wilhelmina. From Lake Habbema upwards, the characteristic overtopping tree is S. alti- gena FRODIN (sect. Brassaia), with its large foliage contrasting sharply with the surrounding microphyllous vegetation; at higher altitudes it gives way to S. pagiophylla HARMs (S. falcata PHILIPSON), a species of uncertain affinities. Schefflera chimbuensis FRODIN and S. straminea FRopIn are likewise often seen in forest borders on Mt Wilhelm and Mt Giluwe respectively in Papua New Guinea. In contrast to the large and conspicuous species of Schefflera, many others are more or less shade-loving epiphytes or vines of lower storeys within the forest, although they may occur in sunnier situations where clouding is frequent (S. singularis B. C. STONE on Mt Ulu Kali). One group of species in New Guinea is almost limited to perhumid moist or wet rain-forests and usually are small epiphytic shrubs or even herbs (S. gemma FRODIN). A few species are con- spicuous rosette-trees of the forest understorey (S. stahliana (WARB.) FRODIN). Climate. In Malesia Araliaceae for the most part shun regions subject to a seasonal climate; all species are evergreen. A few exceptions include Schefflera thaumasiantha HaARMs from open savanna in the hill zone in SE. New Guinea and S. actinophylla (ENDL.) HARMS as a gallery tree in seasonal parts of southern New Guinea, but especially Harmsiopanax aculeatus whose range is for the greater part subject to an annual drought period; in addition to SW. Celebes, it is also frequent on old lava-streams on Mt Idjen in East Java together with Wightia, Casuarina junghuhn- iana, Dodonaea, and Wendlandia, and is one of the few araliads in the Lesser Sunda Islands. With respect to altitude, most Araliaceae occur in the lowland, hill and montane zone below 2300 m. The only genera of which all Malesian representatives occur in the hills and mountains above 1000 m are Pentapanax and Dendropanax, but even these do not exceed 3000 m. Only certain species of Schefflera and Harmsiopanax continue upwards much higher, especially in New Guinea where the highest known record belongs to S. pagiophylla HARMs on the Carstensz complex, where F. J. WissEL found it in 1936 at 3900 m. Flower biology. Little has been recorded about the floral biology of the family in Malesia, but BECCARI’s account (1878) of the ‘false fruits’ of Osmoxylon (including Boerlagiodendron) serving to attract doves which are assumed to effect pollination has become a classic description and example of ornithophily. _ Heterosexual fiowers, usually involving combinations of perfect and male flowers, occur frequently but understanding will require intensive study in the living state. Though flowers are not generally showy and often veritably inconspicuous, the disk produces abundant nectar. They also may spread a rather disagreeable scent, somewhat spermatic, that in Schefflera rugosa resembling that of Ligustrum. For this species DocTERS VAN LEEUWEN (1933) observed on Mt Gedeh, West Java, only rare visits (notably by Diptera), but assumed that cross- pollination will be the rule. Flowers (at least those of Fatsia) will also be visited by Hymenoptera. In Schefflera stahliana (WaRB.) FRoDIN, the thick fleshy flowers, numerous stamens, and position of the inflorescence below the rosette of leaves all point to bat pollination. Dispersal in the family takes place generally by fruit-fall; but as fruits are baccate or (more 4 FLORA MALESIANA [ser. I, vol. 91 usually) drupaceous, they will also be eaten by birds (for the most part) and bats, as recorded by Ripey (1930) for Aralia, Hedera, and Schefflera. The black fruits of Schefflera sect. Brassaia in New Guinea and Australia are especially popular with birds. The hooked mericarps of Harmsio- panax are exceptional. Seed germination is most likely after the seed having passed the gut of a bird or after mastica- tion of the fruit by a bat. References: BECCARI, Malesia 1 (1878) 193-198; Brass, J. Arn. Arb. 22 (1941) 271-342, esp. 318, 320, 323, 327; DocTERS VAN LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. 2, 31 (1933) 195; H. J. Lam, Trop. Natuur 13 (1924) 20; ibid. 14 (1925) 6; RipLey, Disp. (1930); StEup, Trop. Natuur 27 (1938) 142. Morphology. Most erect Araliaceae are sparingly branched or even unbranched; their limbs are upright and the often massive, easily broken twigs generally bear rosettes of big, long- stalked leaves which leave large scars when they fall. Few form a true crown (Arthrophyllum, Gastonia, some species of Schefflera and Polyscias), while the others often look like elongated and grotesque shrubs with a candelabrum-like framework. Branches in most Araliaceae are entirely orthotropic; generally speaking, the plants would fit into the so-called Holttum, Corner, Tomlin- son, Chamberlain and Leeuwenberg models of HALLE c.s. (1970, 1978), or their intermediates. Aralia scandens (MERR.) HA is a true climber. Epiphytic species only occur in the genus Schefflera; it is not yet recorded that any of them may appear to turn into a hemi-epiphytic habit. Hedera (not native in Malesia) is almost the only genus with differentiated shoots and marked vegetative dimorphism. A preliminary account of shoot-morphology in the family has been given by PHILIPSON (1978), but much further field work is required in this area. Vegetative buds may be either proleptic or sylleptic; in the resting phase such buds may be covered by the clasping bases of foliage leaves (Osmoxylon, Schefflera) or (more rarely) specialized cataphylls (Acanthopanax). In both types of bud the primordia and young leaves may be covered by exudations of resinous slime. There is so far little recorded evidence of the changes in leaf shape and configuration during the somatic phase of the life cycle known in many araliads in other parts of the world; however, recent observations in New Guinea made by FRODIN suggest that heteroblastism does occur, although its manifestation is not constant for a given species. Distinctive juvenile and intermediate foliage has been found in Schefflera eriocephala Harms and (to a lesser extent) in S. stolleana Harms. Other examples are seen in Brassaiopsis, Trevesia and Schefflera subg. Agalma (S. aroma- tica (BL.) Harms; S. nervosa (KING) ViG.); juvenile leaves of some of these are preserved in Herbarium Bogoriense. In Harmsiopanax, the configuration of the leaves changes abruptly just below the inflorescence, a phenomenon paralleled in some other genera although less dramati- cally. In Mackinlaya celebica (HARMS) PHILIPSON and M. schlechteri (HARMS) PHILIPSON, leaf polymorphism is very marked with the result that in the past several ‘paper species’ have been described on too limited a range of material; in this revision many reductions have been made. References: HALLE & OLDEMAN, Essai sur l’architecture et la dynamique de croissance des arbres tropicaux, Paris (1970); HALLE, OLDEMAN & TOMLINSON, Tropical trees and forests: an architectural analysis, Berlin (1978); PHiLipson in Tomlinson & Zimmermann (eds.), Tropical trees as living systems (1978) 269-284. Anatomy. General accounts of the vegetative anatomy of the ivy family are given by GUssow (1900), ViGurER (1906, 1909) and METCALFE & CHALK (1950). Secretory canals are characteristic of the stems and leaves, but are absent from Aralidium. A comparison of the wood anatomy of Araliaceae and Cornaceae is made by PHILIPSON (1967), and an extensive account of vegetative anatomy in the context of woody Umbellales was provided by RODRIGUEZ (1957, 1971). The xylem of the former family is characterized by fibres with small, simple pits, and the presence of scalariform and reticulate perforation plates in the vessel elements (as opposed to simple perforations) is thought to be a less advanced feature. Recent special reports bearing on taxonomy include: on stomatal development (INAMDAR c.s., 1969); on sievetube plastids (BEHNKE, 1972), and on epidermal papillae (Bul, 1974). Floral anatomy is discussed by BAUMANN-BODENHEIM (1955), PHILIPSON (1967, 1970) and especially by EypE & TsENG (1971). Embryological characteristics of the family have been re- 1979] ARALIACEAE—I (Philipson) 5 viewed by Davis (1966) as well as by RAo (1972). The single pendulous anatropous ovule has the funiculus and ovular vascular bundle axial (PHILIPSON, 1970). The embryo is small in a mass of endosperm (MARTIN, 1946; GRUSHVITZKY, 1967). References: BAUMANN-BODENHEIM, Bulli. Soc. Bot. Suisse 65 (1955) 481-510; BEHNKE, Bot. Rev. 38 (1972) 155-197; Bu1 NGoc-SANH, Bull. Mus. Hist. Nat. Paris III, Bot. 18 (1974) 85-91 (whole no 271); Davis, Systematic embryology of the angiosperms, New York (1966); EyDE & TsENG, J. Arn. Arb. 52 (1971) 205-239; GrusHvitzky, Proc. Int. Symp. Physiol. Ecol. & Bio- chem. of Germination (ed. H. Borriss) (1967); GUssow, Beitrage zur vergleichende Anatomie der Araliaceae, Thesis, Breslau (Wroclaw) (1900) 67 pp., illus.; INAMDAR, GOPAL & CHOHAN, Ann. Bot. n.s. 33 (1969) 67-73; MARTIN, Amer. Midl. Nat. 36 (1946) 513-660; METCALFE & CHALK, Anatomy of the dicotyledons II, Oxford (1950); PHiLipson, New Zeal. J. Bot. 5 (1967) 134-165; in Robson, Cutler & Gregory (eds.), New research in plant anatomy, London (1970) 87-100; Rao, Phytomorphology 22 (1972) 75-87; RoDRIGUEZ, Univ. Calif. Publ. Bot. 29 (1957) 145-318; in Heywood (ed.), The biology and chemistry of the Umbelliferae, London (1971) 63-91; VicuieER, Ann. Sci. Nat. Bot. IX, 4 (1906) 1-209; ibid. IX, 9 (1909) 305-405. Palynology. Palynological studies of Malesian Araliaceae include: on Gastonia (TsENG, 1971); on Tupidanthus and Plerandra (= Schefflera) (TSENG, 1973); on Osmoxylon (as Boer- lagiodendron) (TSENG, 1974) and on Schefflera (TSENG & SHOUP, 1978). A detailed consideration of the relations of Araliaceous pollen to those in other orders is given by HiDEAUX & FERGUSON (1976), and of the affinities of Klotzschia (Umbelliferae/Hydrocotyloideae) to Araliaceae by SHOUP & TSENG (1977). To date, much useful new evidence has been made available, but better correla- tion with other classes of attributes is required. References: HIDEAUX & FERGUSON in Ferguson & Muller (eds.), The evolutionary significance of the exine, London (1976); SHourp & TsENG, Amer. J. Bot. 64 (1977) 461-463; TsENG, Amer. J. Bot. 58 (1971) 505-516; Grana 13 (1973) 51-56; Amer. J. Bot. 61 (1974) 717-721; TsENG & SHoup, Amer. J. Bot. 65 (1978) 384-394. Chromosome numbers. Lists of chromosome numbers for members of Araliaceae are given by DARLINGTON & WYLIE (1955), SHARMA & CHATTERJI (1964) and BOLKOVSKIKH c.s. (1969). The family shows considerable constancy of the basic number of x = 12, although x = 11 has been recorded for one non-Malesian group of Schefflera. The implications of chromosome data on the wider relationships of the Araliaceae are discussed by Moore (1971). References: BOLKOVSKIKH c.s., Chromosome numbers of flowering plants, Leningrad (1969); DARLINGTON & WYLIE, Chromosome atlas of flowering plants, ed. 2, London (1955); Moore in Heywood (ed.), The biology and chemistry of the Umbelliferae, London (1971) 233-255; SHARMA & CHATTERJI, Cytologia 29 (1964) 1-12. Phytochemistry. Information on the chemistry of the Araliaceae should be sought in HEGNAUER (1964, 1978) where references to original sources are given. The family is characterized by the occurrence of essential oils and resins in canals and by the presence of polyacetylenic compounds (especially falcarinone-type), triterpenic sapogenins of the oleanene-, ursene- and dammarene-types, seed oils with petroselinic acid, and by the absence of true tannins. The chemistry of the family fully confirms its close relationship with the Umbelliferae and also more distantly with the Pittosporaceae and the Compositae (HEGNAUER, 1969, 1971; BOHLMANN, 1971). The rareness of flavones and the predominance of flavonols in Araliaceae suggests closer relationship with two of the three subfamilies of the Umbelliferae, flayones having not yet been found in Hydrocotyloideae and Saniculoideae (HARBORNE, 1971). The absence of iridoid sub- stances and true tannins and the presence of polyenes, petroselinic acid and isoprenylated coumarins in the Umbellales contrasts with the Cornales (HEGNAUER, 1969; JENSEN c.s., 1975), and this led to a suggestion that the Umbellales and Cornales had to be separated (BATE-SMITH c.s., 1975), an argument with increasing support from other lines of inquiry. However, insufficient evidence appears to be yet available for the detection of possible lines of relationship within the Araliaceae. References: BATE-SMITH c.s. Biochem. Syst. Ecol. 3 (1975) 79-89; BOHLMANN in Heywood (ed.), The biology and chemistry of the Umbelliferae (1971) 279-291; HARBORNE, /.c. 293-314; HEGNAUER, Chemotaxonomie der Pflanzen 3 (1964); in Harborne & Swain (eds.), Perspectives in phytochemistry (1969) 121-138; in Heywood (ed.), The biology and chemistry of the Umbelli- 6 FLORA MALESIANA [ser. I, vol. 9# ferae (1971) 267-277; in Cauwet-Marce & Carbonnier (eds.),, Les Ombelliféres. Contributions pluridisciplinaires a la systématique. Perpignan (1978) 335-363; JENSEN, NIELSEN & DAHLGREN, Bot. Notis. 128 (1975) 148-180. Taxonomy. The Araliaceae are on all grounds closely connected with the Umbelliferae, a very large but mainly temperate and tropical-montane group comprising mostly aromatic herbs with a restricted floral scheme. It seems likely that the ancestors of the ivy family were the woody tropical stock from which the herbaceous Umbelliferae evolved under the rigours of cooler climates (CORNER, 1940) with the arborescent Heteromorpha of upland Africa and Myrrhidendron in Central and South America, both in the subfamily Apioideae, perhaps representing relics of the transition on account of the presence of a number of attributes primitive for the family (RODRIGUEZ, 1957); on the other hand, the umbellifers may have originated as megaherbs on tropical mountains (PHILIPSON, 1978). A few Araliaceous genera exhibit some features charac- teristic of Umbelliferae, such as Harmsiopanax, Mackinlaya, and especially Myodocarpus (from New Caledonia) and Stilbocarpa (from southern New Zealand and associated ‘subantarctic’ islands). This led HARMs (1898) to show three different lines leading from Araliaceae to Umbelliferae, suggesting that considerable overlap between the families existed; more recently RODRIGUEZ (1971) again called attention to this phenomenon and THORNE (1968, 1973) has gone further by merging Umbelliferae into Araliaceae (a step also advocated by HALLIER f. in 1905) and assuming the three subfamilies of the former to have arisen separately, perhaps in different parts of the world at different times, from proto-Araliaceous ancestors. However both he and RODRIGUEZ have concluded in agreement with CORNER (/.c.) and BAUMANN-BODENHEIM (1946) that the Araliaceae sensu stricto, ‘“‘because of their greater evolutionary breadth and their retention of many more primitive features, would seem closest to the proto-araliad stock’? (THORNE, 1973). Nevertheless, the concept of Araliaceae as a separate natural family appears to serve a useful purpose and has been retained for this Flora. An isolated, doubtfully included genus is the West Malesian Aralidium; it is the only genus lacking resin ducts but it would be equally anomalous in the Corndceae to which it has also been referred. It shows some resemblance with the New Zealand genus Griselinia, usually relegated to the Cornaceae, be it as a marginal member (PHILIPSON, 1967). Subdivision. Since the first significant family monograph by SEEMANN (1868), several systems have been proposed which, taken together, are notable for their lack of consistency. This results from a lack of agreement on the relative importance of the comparatively minor structural and gross anatomical features of reproductive parts traditionally used and by conflicting claims on the relative ‘antiquity’ of polymery versus pentamery. Until recently, there has been for a priori reasons (cf. EYDE, 1975) little recognition of the potential value of vegetative features and their acceptance as valid evidence for a system; and information from wood anatomy, floral histology, palynology, karyology, phytochemistry, and other areas is only beginning to be utilized. While woodiness is generally accepted as a primitive feature in Araliaceae, the impact of the work of CORNER and others on tree structure dnd growth rhythms (summarized in HALLE, OLDEMAN & TOMLINSON, 1978; see also BORCHERT, 1969; HLADIK, 1970; PHILIPSON, 1978) has still to be fully assimilated. Much more work is also required on inflorescences, although FRODIN (1970), PHILIPSON (1970b) and others have made a beginning. The monothetic interpretation of most attribute states usual in systems of the family was first challenged by BAUMANN-BODENHEIM (/.c.) who considered that phyletic changes could have taken place in parallel; and EyDE & TSENG (1969) showed that at least some supposedly unidirectional sequences were reversible. This has tended to reduce the supposed importance of many of the traditional attributes, with a conse- quent reduction in the number of genera. Sufficient evidence is not yet available, however, for the construction of a more balanced, polythetically based system of the family, and the long-standing subdivision proposed by HARMS (1898) into three tribes, Schefflereae, Aralieae, and Mackinlayeae, based monothetically on the structure and aestivation of the perianth, is retained. In spite of its now recognized imperfections, it is more valid than the systems of VIGUIER (1906) and HUTCHINSON (1967) and remains the most widely accepted. Some steps towards the formulation of a new system on polythetic principles have been taken by EyDE & TSENG (1971: 221) who make a fundamental distinction based on 1979] ARALIACEAE—I (Philipson) 7 basic leaf-organization (either pinnate or palmate) and recognize the heterogeneity of HARMs’ Aralieae; however, this system was deliberately not completely developed. Generic delimitation. Generic limits within the Araliaceae have long been unstable. As in the Umbelliferae, the flower conforms to a simple and relatively uniform pattern throughout most of the family and systematists have resorted to small technical differences to delimit genera. Harms recognized 51 genera, while ViGuier, little more than a decade later, recognized 80; HUTCHINSON distinguished 84 genera but with criteria very differently weighted as compared with VicuieR. Nevertheless, in Malesia as elsewhere there are several distinctive and very natural genera, including Harmsiopanax, Aralidium, Osmoxylon, Anakasia, Mackinlaya, Arthrophyllum, Delarbrea, and Trevesia. Many other genera in the family, including those Malesian ones not noted above, are mutually less distinctive and delimitation is based on various combinations of a number of attributes which have been subject to many different standards of weighting and inter- pretation. These include: petals valvate or imbricate; pedicel articulated or not; leaves digitately compound, pinnate, or simple; stamen and locule number and the relationship of these numbers; style free or connate; endosperm ruminate or not; and thorns present or not. The present treatment is characterized by a number of generic reductions or exclusions; only one genus described since 1900 has been retained (Anakasia). Hederopsis is united with Macro- panax, a genus with a very similar facies and (in part) overlapping range and which was separated merely on the number of cells in the ovary and variations in inflorescence structure. Wardenia has been united with Brassaiopsis because apart from its simple palmately veined leaves there are no differences; better material collected in recent years has provided evidence that the ovary is in fact 2-locular, but that one ovule aborts and the fruit is as a result 1-seeded. Acanthophora differs from Aralia only in habit (VAN STEENIS, 1948). PHILIPSON (1951) already regarded Anomo- panax as insufficiently distinct from the older Mackinlaya, a union retained here. The same author (1973) combined Boerlagiodendron with the earlier described Osmoxylon as a number of species intermediate between the two genera had come to light. The Malesian species formerly included in Tetraplasandra as well as Peekeliopanax were likewise by PHILIPSON (1970a) reduced to two species of Gastonia; but it should be noted that in Malesia as on the SW. Indian Ocean islands this genus is very close to Polyscias, particularly the very similarly ranging sect. Eupteron (P. nodosa, P. ledermannii, etc.). The two largest Malesian genera are herein both treated in a wider sense. Polyscias includes as its type section a distinctive Melanesian/Micronesian group of species which in the wild state extends into eastern Malesia and the Philippines, and in cultivation further west. The remainder of the genus as represented in Malesia comprises some rather distinctive species, several of which have been given generic rank (Eupteron, Kissodendron, and Palmervandenbroekia). If the name Polyscias were to be confined to the type-section of the genus it would be necessary to recognize a considerable number of small genera. Retention of a wide concept for the genus therefore appears to be preferable. Schefflera, by far the largest genus of the family in Malesia, is likewise more broadly conceived than in the past, although in our region only a small number of species from segregate genera are involved (i.e. those formerly in Brassaia, Plerandra, Scheffleropsis, and Tupidanthus). On a world-wide basis, FRoDIN (1975) recommended the reduction of 12 segregate genera; improved knowledge of the genus required that, as in Polyscias, this step be taken or have the genus split into a number of differently organized smaller genera with more serious nomenclatural conse- ‘quences. A broad concept of Schefflera has therefore been adopted. It may be noted here that the flowers in both the former genera Plerandra and Tupidanthus, characterized by numerous stamens, are thought by FRoDIN to represent a secondary development related to bat pollination, and the large fruits for dispersal by bats or larger birds; this is supported by the position of the inflorescences, which are beneath the leafy rosettes as a result of retarded development although the axes remain sympodial and the shoot units mostly orthotropic. As divisions of the larger genus Schefflera, they are not at all closely related, conforming to the views of TsENG (1974) on the pollen morphology and contrasting with the views of HARMs (1898) and EypDE & TsENG (1971), who regarded them (as genera) as of close affinity and relatively primitive within the family. In fact, rather different levels of specialization are represented overall by the two taxa, not just in the pollen morphology (TsENG, /.c.). 8 FLORA MALESIANA [ser. I, vol. 91 Specific delimitation. This has offered many problems, but fortunately the very abundant material now available from many areas has revealed that many of the species described up to 1951 are in fact conspecific. Especially was the great plasticity in vegetative and inflorescence development not sufficiently appreciated in the past, partly due to imperfect field knowledge of the plants. This has led to considerable reductions in Arthrophyllum, Aralia, Osmoxylon, Gastonia, and parts of Schefflera and Polyscias. Other species have been reduced through a regional approach to the genera. However, in Osmoxylon and Schefflera, many species from the Philippines, Celebes, western New Guinea, and to a lesser extent Borneo and Sumatra are still known only from very few or even only a single collection; this has meant a rather tentative treatment in many cases. The same applies in a more limited way in some of the other genera. Certain species are rather polymorphic, and at least in Schefflera several ‘species-complexes’ have been discerned. References: BAUMANN-BODENHEIM, Bull. Soc. Bot. Suisse 56 (1946) 13-112; BORCHERT, Amer. J. Bot. 56 (1969) 1033-1041; CorNER, Ways. Trees Malaya (1940) 153; Eype, Amer. Sci. 63 (1975) 430-437; EYDE & TSENG, Science 166 (1969) 506-508; J. Arn. Arb. 52 (1971) 205-239; FRoDIN, The complex of Cephaloschefflera in Schefflera (Araliaceae), Thesis, Cambridge, U.K. (1970); J. Arn. Arb. 56 (1975) 427-448; HALLE, OLDEMAN & TOMLINSON, Tropical trees and forests: an architectural analysis, Berlin (1978); HALLIER f. New Phytol. 4 (1905) 151-162; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1898) 1-62; HLApDIK, Adansonia 10 (1970) 383-407; HuTCHINSON, Gen. FI. Pl. 2 (1967) 52-81; PHiLipson, Bull. Brit. Mus. Nat. Hist. Bot. 1 (1951) 3-20; New Zeal. J. Bot. 5 (1967) 134-165; Blumea 18 (1970a) 490-495; ibid. 18 (1970b) 497-505; ibid. 21 (1973) 81-89; in Tomlinson & Zimmermann (eds.), Tropical trees as living systems (1978) 269-284; RODRIGUEZ, Univ. Calif. Publ. Bot. 29 (1957) 145-318; in Heywood (ed.), The biology and chemistry of the Umbelliferae (1971) 63-91; SEEMANN, Revision of the natural order of Hederaceae, repr. from J. Bot. London (1868); VAN STEENIS, Bull. Bot. Gard. Btzg III, 17 (1948) 390-391; THORNE, Aliso 6 (1968) 57-66; Notes R. Bot. Gard. Edinb. 32 (1973) 161-165; TsenG, Amer. J. Bot. 61 (1974) 717-721; ViGuter, Ann. Sci. Nat. Bot. [X, 4 (1906) 1-209. Uses. A variety of minor local uses are reported by BURKILL (1966), HEYNE (1927), and OcHsE & BAKHUIZEN VAN DEN BRINK (1931) as well as in the notes under individual species in this Flora. The only species that form articles of trade are the taxa of Polyscias sect. Polyscias popularly grown as foliage and hedge plants; however, many other species in a variety of genera are of actual or potential ornamental worth, with Schefflera actinophylla (ENDL.) HARMS and S. longifolia (BL.) ViG. being particularly widely used. In Papua New Guinea, Gastonia spectabilis is cut for timber and the wood used for light carpentry, boxes, etc. Monkeys are fond of the flush of some aromatic species of Schefflera, as observed in West Java and North Sumatra. References: BURKILL, Dict. rev. ed. (1966); HEYNE, Nutt. Pl. (1927); OcHsE & BAKHUIZEN VAN DEN BRINK, Veget. D.E.I. (1931). Notes. Nomina nuda and invalidly published names have only been quoted if they have been cited in Index Kewensis. Notes for collectors. Many araliads present problems to collectors because of the size of their leaves and inflorescences. It is often advisable to select leaves of medium size, but the maximum size of leaves should be recorded on the label. It is important to preserve the junction of leaf and stem and also sufficient of the leaf to allow reconstruction of the whole. Likewise, with inflores- cences the base, main axis and some primary branches should be preserved so that the whole can be visualized — ultimate branches alone are insufficient. Fruiting material is as useful as a flower- ing specimen. Collectors should be alert to note the existence of vegetative heteroblasty and floral dimorphism and document these with specimens and notes; the form in immature plants is important. Rapid drying is essential or all parts will disarticulate and very fragmentary specimens result. KEY TO THE GENERA 1. Petals imbricate. Tribe ARALIEAE. 2. Leaves simple, palmately or pinnately lobed, or entire. 3) Leaves palmately lobed; tomentose’... = = « ss sonst eel oo nS 1. Harmsiopanax 3. Leaves pinnately lobed or entire, glabrous’ =< 2. = =). 0) meee 2. Aralidium 1979] ARALIACEAE—I (Philipson) 9 2. Leaves pinnately compound (or bi- or tripinnate). A eAVeSUWICe) (OL INOTe) PINNATC erie ie eS ss, os) wont fo obs Seeds 4 v0) Fig. 1. Harmsiopanax ingens PHILIPSON ssp. ingens. a. Lea lule, x 4, d. developing fruit 1979] ARALIACEAE—I (Philipson) 11 minute rim. Petals 5, free, valvate with a broad base. Stamens 5, dorsifixed, versa- tile, introrse. Ovary inferior, narrowly obconic, densely bristly; cells 2; disk conical, deeply cleft between the two subulate styles. Fruit consisting of 2 dry mericarps, each 3-ribbed and bearing a persistent slightly hooked style. Distr. Malesia: 3 spp. from Java, the Lesser Sunda Is., Celebes, and New Guinea. Ecol. Montane and mossy forest and in regrowth on grassy hillsides. Notes. Harmsiopanax is a small structurally isolated genus confined to Malesia. The three species are uniform both in their vegetative and their reproductive features. It has long been recognized that some of the characters of this genus are anomalous within Araliaceae and a return to its earlier position within Umbelliferae would have something in its favour. The monocarpic habit is unknown elsewhere in Aralia- ceae, but is not uncommon in Umbelliferae. The character of the fruit, which splits into two dry mericarps, closely approaches the fruit structure of Umbelliferae, and the vascularization of the gynoecium is also characteristic of that family. However, the structure of the leaf-base, the woody habit, and the shape of the petals all incline towards Araliaceae. KEY TO THE SPECIES 1. Upper surface of leaves uniformly setulose. PemUMBCIWIESISESSIIC) cha bc Islas eis oe se Zumbellules peduncled. . . 2. . ws ws 1. Upper surface of leaves with many (or rarely few) larger spines among the setulose hairs 1. Harmsiopanax aculeatus (BL.) WARB. ex BOERL. Hand. 3 (1900) 88; Koorp. Exk. Fl. Java 2 (1912) 719; Atlas 4 (1916) f. 668 & 669; Baxn. f. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1946) fam. 159, p. 19; Back. & BAKH. f. Fl. Java 2 (1965) 171; STEEN. Mt. FI. Java (1972) pl. 3-2; PHILIPSON, Blumea 21 (1973) 82. — Schubertia aculeata Bi. Bijdr. (1826) 885. — Horsfieldia aculeata (BL.) DC. Prod. 4 (1830) 87; BENN. PI. Jav. Rar. (1840) 123, t. 26; K. & V. Bijdr. 7 (1900) 57; BoERL. Handl. 1 (1890) 647. — Horsfieldia peltata BTH. in B. & H. Gen. Pl. 1 (1862) 937. — Fig. 2. Tree up to 4 m, with a slender spiny trunk. Young stems covered more or less densely with woolly hairs, bristles, and spines with bulbous bases, the spines enlarging on older stems. Leaves rounded, variable in size, often 60 cm or more in @, deeply palmately lobed, usually peltate in mature leaves, sinuses between the lobes broad or narrow, lobes 7-10, usually sharply and irregularly incised and toothed, apex acute, upper surface rather sparsely covered with evenly-spaced, appressed, sometimes branched hairs (denser on the main veins), underside densely clothed with a soft, woolly tomentum, often with some bristles on the main veins; petiole c. 60 cm, 1 cm @ at base, terete with clasping base, densely covered with woolley hairs, bristles, and some spines. Jnflores- cence up to c. 70 cm long, main branches rather sparsely covered with a short tomentum and, when young, bearing numerous bracts similar to the leaves but smaller, not peltate, and often 3-lobed or entire; ultimate branchlets slender and often woolly-tomentose, bearing minute linear bracts which subtend the sessile umbellules. Umbellules ee ee I ee CO ee ee ee ee Tk a 3. H. ingens about 4 mm @ in flower, the broadly ovate outer bracts forming a more or less distinct involucre. Flowers hermaphrodite or male, either mixed in an inflorescence, or separate, c. 10-15 per umbellule, each subtended by a lanceolate receptacular bract c. 2 mm long. Pedicel c. !/, mm long. Calyx rim fringed. Petals strap-shaped, c. 11/, mm long at anthesis. Filaments c. 2 mm; anthers c. 0.3 mm long, orbicular. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps long-ciliate, crowned with the divergent styles. Distr. Malesia: Java, Lesser Sunda Is. (Bali, Lombok, Sumbawa, Flores, Timor), southern half of Celebes. There is a single KORTHALS sheet in L, ticketed from Central Sumatra, but this is presum- ably wrongly localized. Ecol. Usually in rather dry, open localities, but also in forest, in secondary forest, also pioneering on rocks, in grasslands and on lava-streams, 300-1800 m. Fi. fr. April-Nov. ScHMutTz found it in Flores flowering in October, but leaves had fallen. Vern. Java: djankurang, d. tjutjuk, djogloran- grang, S, gabus, garang, g. lanang, gungrang, udulan laki, J. Note. In this species lateral shoots usually appear below the infructescences so that the trees are not normally monocarpic. 2. Harmsiopanax harmsii K. Sco. in K. Sch. & Laut. Nachtr. (1905) 329; Harms, Bot. Jahrb. 56 (1921) 413. Tree up to 7 m, with a slender trunk. Young stem covered with bristles, hairs, and spines, older stems with smooth bark with small rounded lenti- cels and numerous spines. Monocarpic. Leaves 12 FLORA MALESIANA [ser. I, velo" Fig. 2. Habit of Harmsiopanax aculeatus (BL.) WARB. ex BOERL. Coarse shrub on old lava-streams in E. Java (Mt Idjen) at c. 900 m altitude (Photogr. VAN STEENIS). rounded, up to 30 by 40 cm, deeply palmately lobed, cordate at base, lobes 5—9 with broad sinuses between them, margin unevenly and sharply den- tate, apex acute, upper surface densely covered with evenly spaced bristles of varying size (larger on the main veins), appressed and directed towards the leaf margin, often with woolly hairs inserted on their enlarged bases, the underside very densely woolly and with many bristles, usually bearing crisped hairs on their enlarged bases; petiole 50 cm, '/, cm @ at base, terete with clasping base, densely covered with bristles, woolly hairs, and spines. Panicle at first with numerous leaf-like bracts, the principal branches with some spines, rather sparsely covered with bristles and hairs, ultimate branches slender and tomentose, bearing linear bracts c. 4 mm long subtending peduncled umbellules; peduncles up to 5 mm, slender, tomen- tose, bearing 2 minute bracts. Umbellules spheri- cal, c. 4-5 mm @ in flower, outer bracts not form- 1979] ARALIACEAE—I (Philipson) 13 ing a distinct involucre. Flowers hermaphrodite, maturing in basipetal succession, the lower bracts of a branch either with sterile umbellules or lacking flowers; up to 60 in an umbellule, each subtended by a lanceolate ciliolate bract c. 1 mm long, and borne on a glabrous pedicel c. 1'/, mm long. Calyx rim fringed with many lacerate filaments. Petals ovate, c. 1 mm long. Filaments c. 1 mm; anthers c. '/, mm long. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps with rounded ribs, long-ciliate, crowned by the diver- gent styles. Distr. Malesia: Papua New Guinea (Madang Distr., Western Highlands, Morobe Distr. & Central Distr.). Ecol. Forested hills, grassy slopes, and road- sides, 100-1800 m. Vern. Opme, Ganja, Mt Hagen, mafiong, Sattelberg, Morobe Distr. Note. Information about the habit is inade- quate. The stalked spherical umbellules are very distinctive. 3. Harmsiopanax ingens PHILIPSON, Blumea 21 (1973) 84. ssp. ingens. — Fig. 1, 3. Unbranched tree up to 18 m witha thick or some- times slender trunk densely covered, except towards the base of mature specimens, with long, sharp, upwardly directed spines and marked with leaf-scars. Monocarpic. Leaves usually peltate, rounded, up to 1 m @, deeply palmately lobed, lobes usually with minor lobes and coarsely dentate, apex acute, upper surface bearing few to many long spines, especially on the midrib and principal veins between which the surface is often rugose and glabrous except for the remains of a tomentum of branched hairs, or with many bristles often with woolly hairs on their bases, the under- surface also with few to many long spines and usually clothed with a fawn or greyish woolly tomentum of branched hairs, or densely furnished with bristles usually with woolly hairs on their bases, or occasionally glabrous between the spines Fig. 3. Harmsiopanax ingens PHILIPSON. Left: apex of leafy stem; right: the large inflorescence (Photogr. FRODIN, Murmur Pass, 1971). 14 FLORA MALESIANA [ser. I, vol. 91 except for a few bristles; petiole up to 1 m and 3 cm @, terete with clasping base, covered with woolly hairs and bearing many spines. Panicle up to 5 m long and 5 m wide, leafless or with lobed bracts c. 10-20 cm long, principal branches spiny especially below, ultimate branches slender, tomentose, bearing linear bracts c. 1 cm long sub- tending peduncled or sessile umbellules; peduncles elongating as the fruit ripens, up to 4 mm, rather stout, tomentose, bearing 1 or 2 minute bracts. Umbellules bowl-shaped, c. 6-10 mm @ in flower, enlarging slightly in fruit, with an involucre of about 8 ovate bracts, 2-4 mm long and ciliolate distally. Flowers hermaphrodite, maturing in basipetal succession, terminal branches bearing maturing fruit while lower branches bear flowers or unopened buds; usually c. 12-16 (8-20) in an umbellule each subtended by an involucral bract or a narrower receptacular bract and borne on a glabrous pedicel 1-2 mm long. Calyx rim fringed with many lacerate filaments. Petals ovate, 1-2 mm long. Filaments 2-3'!/, mm; anthers +/,/, mm long. Ovary covered with cilia which lengthen as the fruit ripens. Mericarps with rounded ribs, long- ciliate, crowned by the divergent styles. Distr. Malesia: New Guinea (NW. Irian and extending along the central mountains from the Orion Mts to the Owen Stanley Range, Murray Pass). ; Ecol. Montane and mossy forest and second- growth forest, 2000-3600 m, occasionally rather lower. Vern. Papua: mauku, Huli; Mandated Terr.: Sepik Distr., kamul, Hindenburg Ra.; Western Highlands: murri, Hagen, tolsan, Minj, mauri, Melpa, mai, Mendi, kinogore, makua, makw, Enga; Eastern Highlands: kimu, Ka, ollu, Chimbu. Notes. A striking, single-trunked, monocarpic tree bearing immense inflorescences. The bark is described as grey brown and the wood white with a wide pith. The inflorescence has the appearance of bearing female flowers above and male flowers below, but this is evidently due to a basipetal sequence of anthesis. The terminal flowers have stamens when freshly opened and all those on lower branches bear styles. Apparently, the female organs of the lower branches are functional because branches from mature inflorescences bear fruit uniformly. Nevertheless, herbarium specimens cannot adequately represent such a large inflores- cence so that the possibility remains that some female-sterile flowers occur in this species. Variation occurs in both tomentum and in- florescence characters. For example, most speci- mens from West Irian have small umbellules and fewer leaf-spines. In the eastern part of the Eastern Highlands District a number of gatherings display a series of variations: the under-leaves give the appearance of being glabrous between bristles, the inflorescence branches bear small leafy bracts, the umbellules are sessile, with rather numerous {c. 18-21) small flowers subtended by rather broad bracts. Specimens from Mt Otto show all these features combined, but other specimens from this region diverge from the typical state in only some of these characters. No specimens of this subspecies are known from the Finisterre Range and only one from the Owen Stanley Range. ssp. moniliformis PHILIPSON, Blumea 21 (1973) 86. Umbellules disposed irregularly along the branches, singly or in small groups, with bare spaces intervening, sessile; flowers usually c. 20-30 per umbellule, floral parts smaller than in ssp. ingens; fruiting heads rather small (c. 5 mm 2). Distr. Malesia: Papua New Guinea (districts bordering on the Huon Gulf). Vern. Morobe Distr.: mobian, Finschhafen. Note. This subspecies occurs at lower altitudes than is usual for ssp. ingens {1500-2000 m). No specimens of either subspecies have been collected from higher altitudes in the mountains north of the Markham River and the Huon Gulf. At higher altitudes in the Owen Stanley Range ssp. ingens is known from one gathering. The most south-easterly gathering at present known (CARR 13603) has a distinctive appearance due to the straight rigid in- florescence branches with small sessile umbellules. 2. ARALIDIUM Mia. Pl. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762, t. 13; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 25; Bru. in B. & H. Gen. PI. 1 (1865) 936; HEMSL. in Hook. Ic. Pl. 16 (1886) t. 1549; Borer’. Handl. 1 (1890) 631; HArRMs in E. & P. Nat. Pf. Fam. 3, 8 (1894) 60; Hutcu. Gen. FI. Pl. 2 (1967) 59. — Fig. 4. Unarmed tree or shrub with simple, exstipulate pinnately lobed, irregularly incised, or entire leaves. Inflorescence a large panicle, with cymules of small flowers arranged racemosely on the branches. Pedicels articulated below the ovary. Dioecious. Male flowers: calyx 5; petals 5, imbricate; stamens 5, anthers dorsifixed. Female flowers: calyx and corolla similar to male; staminodes 5; ovary with 3 1979] ARALIACEAE—I (Philipson) 15 Fig. 4. Aralidium pinnatifidum (SUNGH. & DE VrigsE) Mia. a. Habit, x 2/s, b. ¢ flower and bud, c. 2 flower and bud, both x 8, d. fruit, seed, and CS, slightly enlarged (a VAN BALGooy 2185, b COCKBURN FRI 8376, c SINCLAIR 9884, d fresh material). Drawn by W. R. PHILIPSON. 16 FLORA MALESIANA [ser. I, vol. 91 locules (2 abortive) and 1 ovule, styles 3-4, tapering from broad bases, stigmas terminal. Fruit drupe-like. Seed solitary, pendulous from a thickened funicle, 4-5-2rooved; endosperm deeply ruminate. Distr. Monotypic. Peninsular Thailand and Malesia: Malay Peninsula, Sumatra, and Borneo. The record from Java by MIQuEL (Ann. Mus. Bot. Lugd.-Bat. 1, 1863, 25) is erroneous (see K. & V. Bijdr. 7, 1900, 2). Ecol. Primary and secondary forest, from sea-level to c. 1250 m. Notes. The single species forms a genus with several unique features, namely the large, simple, charac- teristically pinnately lobed leaves, the absence of resin ducts, the diffuse panicles of male or female flowers, the 3-carpellate ovary with a single surviving loculus resulting in a single-seeded fruit, the dorsal raphe, the deeply ruminate endosperm, and the enlarged funicle. The genus is treated here as a member of the Araliaceae mainly as a matter of convenience. Sometimes it has been placed in the Cornaceae (e.g. RiDL. Fl. Mal. Pen. 1, 1922, 894, and VicuterR, Ann. Sc. Nat. Bot. 4, 1906, 171), and on full investigation it may well prove to be better placed in that family. The absence of resin ducts and the dorsal raphe strongly support a relationship with the Cornaceae and its immediate allies, though the absence of borders to the pits of the xylary fibres is characteristic of Araliaceae. Several genera formerly placed in the Cornaceae have now been elevated to the rank of family. If this course is followed then Aralidium should also be segregated. Many of the features of Aralidium approach those of Griselinia (segregated as Griseliniaceae) and possibly these two genera should be united as a single family. 1. Aralidium pinnatifidum (JUNGH. & DE VRIESE Mia. FI. Ind. Bat. 1, 1 (1856) 763; HEMsL. in Hook. Ic. Pl. 16 (1886) t. 1549; BoerL. Handl. 1 (1890) 631; RipL. Fl. Mal. Pen. 1 (1922) 895; PHILIPSON, J. Bot. 78 (1940) 118. — Aralia pinnatifida JUNGH. & DE VRIESE, Ned. Kruidk. Arch. 1 (1846) 15; Ann. Sc. Nat. III, 6 (1846) 115. — A. dentatum Mia. Sum. (1861) 340. — A. integrifolium HEINE in Fedde, Rep. 54 (1951) 245. — Fig. 4. Shrub or small tree up to c. 10 m, rarely reaching 20 m and 25 cm @, glabrous in its vegetative parts; buds enclosed in long sheathing leaf-bases. Leaves spaced with distinct internodes, usually c. 30 by 22 cm or more, + regularly pinnately incised, frequently as deep as the midrib, lobes oblong- acuminate and decurrent on the midrib, c. 21/, cm wide or more, the lobing sometimes irregular, and occasionally the blade entire and broadly ovate (up to 25 by 20 cm) or rarely lanceolate, leaf mar- gin either entire or coarsely dentate, especially on the terminal lobe; petioles 5-12 cm, broadly channelled above, clasping the stem with a slightly dilated base, exstipulate. Panicles terminal, or occasionally in the upper axils, to 50 cm long, pendulous, puberulous; main bracts caducous, but the minute bracteoles often persisting until anthesis. Flowers numerous, small (buds c. 2!/, mm long), fragrant, creamy or red-tinged, ovary, calyx lobes and petals densely covered in a minute but coarse puberulence. Male flowers with the corolla persistent during anthesis, petals c. 11/, mm long, strap-shaped, spreading, stamens c. 1 mm with flattened filaments and round anthers; stylopodium a succulent disk with a concave centre; styles absent, the ovary 1'/, mm long, narrowly turbinate, without a loculus. Female flowers with the corolla caducous at anthesis together with the staminodes, styles divergent from their gibbous bases, ovary ovate with a single loculus (two abort early); ovule pendulous. Fruit usually obliquely ellipsoid, tapering to the apex and c. 3-4'/, cm long, but rarely subspherical, white when immature, ripening to purplish or black, juicy ; exocarp fleshy, endocarp chartaceous. Seed broadly ellipsoid, 2—21/, cm long, with the surface patterned with deep ruminations. Distr. Peninsular Thailand; in Malesia: Malay Peninsula (from Kedah southwards common; Singapore), throughout Sumatra (also in Simalur I.), Anambas Is. (Siantan) and throughout Borneo. Ecol. Frequent in evergreen primary rain- forest, also in open bamboo forest and secondary growths, from sea-level to c. 1250 m, in Borneo up to 1500-1800 m. FI. fr. Jan.—Dec. Uses. The only use, once mentioned, is from Brunei, as “‘leaves make good ghost medicine”’. Vern. Malay Peninsula: /émpédu buaya, (poko) balai, pungar, sahalat, sébalai tingal, sibilai, tébalai, M; Sumatra: (kayu) attarodan, Asahan, Batak lang., ségéntut, Gajo, médung, M, maneél silai, mannel dotan, sukun dotan, M, Simalur; Anambas Is.: ballok, M, Siantan; Borneo: daun tutchol antu, Brunei, Iban lang. Note. Entire leaves are not infrequent through- out the range of the species, so that the recognition of a second species using this character is not justified. Coarsely dentate leaf-margins were also employed as a specific character but are merely a minor variation. Some specimens from Mt Kina- balu have rather small globose fruits, but the typical form of fruit also occurs on that mountain. 1979] ARALIACEAE—I (Philipson) 17 3. ARALIA Linn, Gen. Pl. ed. 5 (1754) 134; Sp. Pl. (1753) 273; DC. Prod. 4 (1830) 257; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6; Bru. in B. & H. Gen. Pl. 1 (1865) 936; BoeRL. Handl. 1 (1890) 629; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 56; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 391; Hutcu. Gen. FI. Pl. 2 (1967) 63; Stone, Gard. Bull. Sing. 30 (1977) 134; PHILIPson, /.c. 97. — Acanthophora MERR. Philip. J. Sc. 13 (1918) Bot. 316, non LAMOUREUX, 1813 (Algae); STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 390. — Fig. 5, 6. Sparingly branched shrubs or small trees, or climbing, rarely (extra-Mal.) herbaceous, glabrous or hairy, often prickly. Leaves pinnate to tripinnate, usually with leaflets at the insertion of the lateral pinnae; leaflets serrate; petiole with a sheathing base. Inflorescence a terminal panicle; flowers sessile or pedicelled, with an articulation below the flower; calyx with 5—6 teeth; petals 5-6, imbricate; ovary 2-6-celled; styles 2-6 free or shortly connate below. Fruit a fleshy drupe; pyrenes cartilaginous, compressed; endosperm uniform. Distr. More than 30 spp. in North America (S. to Mexico) and East Asia, 6 spp. in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Is. (Sumba), Borneo, Celebes, Philippines, and West New Guinea. Ecol. Usually on scrubby hillsides and in secondary growth, often in ravines or near streams, or in thickets near or above the limit of tree-growth, at low altitude (100 m), but usually in the montane zone, up to 3000 m. Note. For a discussion of specific distinctions see VAN STEENIS, /.c. 391. Hut-Lin Li in Sargentia 2 (1942) 101, treated some species that extend into Malesia. MERRILL considered that the climbing habit and recurved spines of Acanthophora justified its separation as a distinct genus, but more recent authors have not agreed. KEY TO THE SPECIES 1. Flowers sessile (capitate) or very shortly pedicelled. 2. Flowers sessile, underside of leaf + densely tomentose, hairs of the branches and inflorescence + appressed and felted, bracts around the capitula enveloped in hairs 1. A. dasyphylla 2. Flowers short-pedicelled, underside of leaf sparsely tomentose, hairs of the branches and inflorescence + patent, bracts around the capitula less densely tomentose 2. A. javanica 1. Flowers !/,-1!/, cm pedicelled (umbellate). 3. Climbing or scrambling liana, spines curved 3. Erect shrubs or small trees, spines straight. 4. Leaflets glaucous beneath, margins with few crenations. Fruit small (c. 3 mm long) 4. A. bipinnata 4. Leaflets green (or with fawn pubescence) beneath, margins serrate. Fruit rather larger (4-6 mm long). 5. Young parts and undersurface of leaves glabrous (but with small spines) 5. A. ferox 5. Young parts and undersurface of leaves pubescent 6. A. montana 3. A. scandens 1. Aralia dasyphylla Mra. FI. Ind. Bat. 1, 1 (1856) 751; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9, incl. var. strigosa Miq. et var. latifolia M1Q.; BOERL. Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 53; Koorp. Exk. Fl. Java 2 (1912) 718; Atlas 4 (1916) f. 673 A-K; Hut-Lin Lt, Sargentia 2 (1942) 20; BAKH. f. Blumea 6 (1947) 367, incl. var. urticifolia (BL. ex Mig.) BAKH. f.; BAKH. f. & OostTsTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 18; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 391; Ncoc-SANH Bul, Adansonia 4 (1964) 464; Back. & BAKH. f. Fl. Java 2 (1965) 170; STEEN. Mt. FI. Java (1972) pl. 3-1; PHiLrpson, Gard. Bull. Sing. 30 (1977) 98; Y.-R. Lina, Acta Phytotax. Sin. 15 (1977) 86. — A. chinensis (non L.) BL. Bijdr. (1826) 870. — A. urticifolia BL. ex Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9; BoeRL. Hand. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 55; Koorp. Atlas 4 18 FLORA MALESIANA [ser. I, vol. 91 DD» S CRY) DOS, BG ~\ av. AY a Wy BD S ve S ps g.@ S O5 m ae |g QO Sy 39, 2A BZA gata eyeiKs 3 \V2 2 7a \ at S 8 oe a 7 Fig. 5. Aralia bipinnata BLANco. a. Upper branches of inflorescence, b. pinna, c. base of petiole, all x 2/,, d. flower bud, e. fruit, both x 3 (JAcoss 7017). Drawn by W. R. PHILIPSON. 1979] ARALIACEAE—I (Philipson) 19 (1916) f. 673 L-N. — A. beccarii Riv. J. Mal. Br. R. As. Soc. 1 (1923) 64. — Fig. 6. Prickly shrub or small tree, often unbranched, to c. 5 m; young parts densely brown pubescent. Leaves forming large rosettes at the summit of the stems, c. 1 m long (or more), bi- or tripinnate, with a pair of leaflets (occasionally pinnate) at each division of the rachis, the petiole, rachis and lateral rachides prickly or unarmed, densely pubescent; leaflets subsessile or petiolule c. 5 mm long (or longer), usually densely pubescent on the lower surface, less dense above, ovate to oblong-ovate, c. 5—14(-18) by 3—S(—10) cm, base rounded to sub- cordate, apex acuminate, margin finely or some- times coarsely serrulate; petiole c. 40 cm, with an elongated sheathing base and a small ligule. Inflorescence a large terminal panicle, 70 cm or more long, densely brown pubescent, rachis bearing several secondary branches c. 30-40 cm long, with ultimate branches arranged racemosely, bracts ligulate, ending in heads of several sessile flowers, surrounded by an involucre of small usually densely pubescent bracts. Calyx with 5 usually obtuse teeth; petals 5, c. 11/, mm long, glabrous; stamens 5; ovary c. 2 mm long, glabrous, 5-celled; styles 5, slightly connate below. Fruit globose, c. 31/, mm @, ribbed when dry. Distr. Northwards to southern China; in Malesia: Sumatra, Malay Peninsula, West and Central Java. Ecol. Primary forest and secondary growths in deep ravines or open hillsides, from low altitude (c. 100 m) to 2500 m. Vern. Sumatra: kaju burle lasét, k. sépak- sipang, k. si marsuga-suga, k. sipang-sipang, sami- mpadan, M; Java: gorang, osangsing, J, pangang tjutjuk, S. Note. The capitulate flowers are characteristic (see also under A. javanica). The presence of this species in the Malay Peninsula has often been overlooked, though it extends into southern China. Its variability was discussed by VAN STEENIS (1948, hes): 2. Aralia javanica Mia. Pl. Jungh. 3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 749; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9; BoeRL. Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 55; Koorp. Exk. FI. Java 2 (1912) 718; Atlas 4 (1916) f. 670; Baku. f. & Ooststr. in Back. Bekn. FI. Java (em. ed.) 7 (1948) fam. 159, p. 17; Back. & Baku. f. Fl. Java 2 (1965) 170; PHtLipson, Gard. Bull. Sing. 30 (1977) 98. A shrub or small tree, often unbranched, young parts covered with brown pubescence which per- sists on the stems and rachides of the inflorescence as + patent hairs. Leaves tufted at the ends of the branches, bipinnate with a pair of leaflets at the divisions of the rachis; leaflets variable in size, subsessile or the petiolules up to c. 2 cm, blade ovate to elliptic, up to 18 by 8 cm (usually smaller), both surfaces with sparse, short, appressed, bristly hairs, base cuneate to truncate, apex acuminate, margin finely and unevenly serrulate; petiole 20-30 cm. Inflorescence a large terminal panicle, rachis bearing several secondary branches c. 30-40 cm long, with the ultimate branches bearing heads (or subumbellules) of c. 10 flowers, sur- rounded by an involucre of small linear bracts. Calyx with 5 small teeth; petals 5; stamens 5; ovary c. 2 mm long, glabrous, 5-celled; styles erect at anthesis. Fruit ovoid, c. 5 mm long, ribbed when dry, with the persistent styles recurved. Distr. Malesia: West and Central Java (Mts Papandayan, Malabar, Diéng, Surakarta). Ecol. Mountain forests, 2000-3000 m. Note. This imperfectly known species may prove to be a form of the widespread A. dasyphylla, from which it appears to differ in the shortly pedicelled flowers, the sparser leaf-tomentum, the more patent hairs on the inflorescence branches and the bracts of the umbellules less thickly enveloped in hairs. 3. Aralia scandens (MERR.) Ha, Nov. Sist. Vyssh. Rast. 11 (1974) 229; Stone, Gard. Bull. Sing. 30 (1977) 276, f. 1; PHiipson, /.c. 99. — Acantho- phora scandens Mer. Philip. J. Sc. 13 (1918) Bot. 316; En. Philip. 3 (1923) 236; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 390. — A. ferox (MIaQ.) Kina, J. As. Soc. Beng. 67, ii (1898) 45; Koorp. Minah. (1898) 498; Merr. En. Born. (1921) 458; RIwDi. Fl. Mal. Pen. 1 (1922) 872; Masam. En. Phan. Born. (1942) 564. Prickly scandent shrub, glabrous, reaching a height of 10 m or more, stems c. 2!/, cm @. Leaves dispersed (c. 30 cm apart), up to 11/, m long, tri- or quadripinnate with a pair of leaflets at each division of the rachis, prickly on the petiole, rachides, and sometimes on the leaf veins; leaflets ovate to elliptic-ovate or ovate-lanceolate, petio- lules 3-10 mm, blade 5-14 by 2!/,-5 cm, base rounded or subcordate, apex acuminate, margins finely spinulose-denticulate; petiole to 35 cm, with an elongated sheathing base and a small ligule. Inflorescence a large terminal spiny panicle, the main rachis to c. 60 cm, bearing secondary branches singly or in whorls, up to 50 cm; ultimate branches 1-4 cm, subtended by lanceolate bracts, racemosely arranged, ending in umbellules; umbellules c. 10—20-flowered, pedicels slender, 10-12 mm, with lanceolate bracts 2-3 mm long, articulated below the flower. Calyx with 5-6 short acute teeth; petals 5-6, with a broad base, imbri- cate; stamens 5-6, filaments c. 4 mm long, anthers c. 1 mm long; ovary turbinate c. 2'/, mm long, 5—6-celled, styles 5—6, free or only slightly connate below, at first erect. Fruit ellipsoidal, c. 5 mm long, purple to blue-black, deeply furrowed when dry, crowned by the persistent radiating styles. Distr. Malesia: Malay Peninsula (Perak, Selan- gor, Pahang), Sabah (Mt Kinabalu), Philippines 20 FLORA MALESIANA [ser. I, vol. 9! Fig. 6. Aralia dasyphylla Mia. Habit, Tjibodas, on slope of Mt Gedeh, W. Java, c. 1400 m altitude (Photogr. PHILIPSON, 1973). 1979] ARALIACEAE—I (Philipson) 2t (Luzon, Panay, Catanduanes, Mindanao), Celebes (Menado, Buton I., S. Celebes). Ecol. Thickets on slopes and mountainsides, often near streams, or among secondary growths, 180-1550 m. Uses. In Mindanao is reported that scrapings of the bark are applied to wounds and a decoction of the boiled bark is drunk to releave internal pain. Vern. Philippines: simbar, Bag. Note. The only species with the habit of a liana, with spaced leaves, and recurved spines. The flowers are whitish or yellowish, and slightly fra- grant and are visited by numerous small bees. The fruit is purple and fleshy. cwangayan, Mindanao, 4. Aralia bipinnata BLANCO, FI. Filip. (1837) 222; MerR. Sp. Blanc. (1918) 294; En. Philip. 3 (1923) 235; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 392, incl. f. inermis STEEN.; PHILIPSON, Gard. Bull. Sing. 30 (1977) 99. — A. hypoleuca PrEsL, Epim. (1851) 250; Mia. Fl. Ind. Bat. 1, 1 (1856) 751; F.-ViILL. Nov. App. (1880) 101; VmpaL, Phan. Cuming. Philip. (1885) 117; Rev. Pl. Vasc. Filip. (1886) 144; Harms, Bot. Jahrb. 23 (1896) 18; MERR. Philip. J. Sc. 5 (1910) Bot. 369; En. Philip. 3 (1923) 235. — A. javanica (non Mia.) F.-VILL. Nov. App. (1880) 101. — A. glauca MeErR. Philip. J. Sc. 2 (1907) Bot. 291; En. Philip. 3 (1923) 236. — A. apoensis ELMER, Leafl. Philip. Bot. 7 (1914) 2325; MERR. En Philip. 3 (1923) 235. — Fig. 5. A shrub or small, sparsely branched tree to 7 m, with prickly stems. Leaves to 11/, m or more long, forming large crowns at the ends of the branches, bipinnate, with a pair of pinnae at each division of the rachis, with some prickles, especially on the petiole or unarmed, the rachis swollen and articu- lated at the nodes; leaflets sessile or with a short petiolule, ovate to lanceolate, usually 4-5 by 2-2'/, cm, but variable in size, apex acute or acumi- nate, base rounded to cordate, usually markedly oblique in lateral leaflets, margin conspicuously crenate, upper surface green, glabrous, lower sur- face glaucous, pubescent along the veins and some- times sparingly on the mesophyll, sometimes only in the angles of the lower veins, or almost glabrous throughout, primary and secondary veins con- spicuous; petiole to 30 cm, base long sheathing and slightly ligulate. Inflorescence a large terminal panicle 30-70 cm long (or more), peduncle and also usually the main branches prickly, the whole either almost glabrous or pubescent; peduncle 5-18 cm long, stout; main rays c. 5-10, mostly clustered at the apex of the rachis, 25-65 cm long, bearing many short tertiary branches along their length; tertiary branches usually 5-10 cm long, ending in umbellules, and bearing a small number of lateral umbellules, or branches, minute lanceo- late bracts subtending the branches of the third or higher orders; umbellules with c. 20-30 radiating pedicels ; pedicels 5-10 mm. Calyx lobes 5, rounded, 1/, mm long; petals 5, 1'/, mm long; stamens 5; ovary 5-celled, styles subulate, free. Fruit spheroi- dal, c. 3 by 4 mm, strongly 5-ribbed when dry, persistent styles spreading. Distr. Taiwan; in Malesia: Philippines (Luzon, Leyte, Negros, Mindoro, Mindanao) and West New Guinea (Vogelkop Peninsula, possibly also in Swart Valley). Ecol. In rather open forests, ravines, and in thickets and secondary growths, (700—)1000- 2450 m. Vern. Philippines: badbaranai, C.Bis., dasanat, Neg., Karugi, Buk., magkasau, Bis., mara-bauya, Bag., papang, Bon., sugsuga, lg. Note. VAN STEENIS /.c. discussed the vari- ability in pubescence and the development of spines. 5. Aralia ferox Mia. Fl. Ind. Bat. 1, 1 (1856) 750; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 9; BoeRL. Handl. 1 (1890) 629; K. & V. Bijdr. 7 (1900) 49; Koorp. Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 671; Fl. Tjib. 2 (1923) 229; Baku. f. & OoststrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 18; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 394; Back. & BAKH. f. Fl. Java 2 (1965) 170; PHILIPSON, Gard. Bull. Sing. 30 (1977) 99. — A. filicifolia RID. J. Fed. Mal. St. Mus. 8 (1917) 42, non C. Moore, 1876. Spiny shrub or small tree, usually unbranched, to c. 10 m. Leaves forming a large rosette at the summit of the stem, up to c. 1 m long, or shorter below the inflorescence, bi- or tripinnate, with a pair of leaflets (often pinnate) at each division of the rachis, prickly on the petiole, main rachis, and often on the lateral rachides; leaflets sessile or petiolules to c. 5 mm, ovate or ovate-oblong, usually c. 3 by 13/, cm, but variable in size, base truncate to rounded or cuneate, apex acute acumi- nate, margin sharply serrate, both surfaces with small bristle-like spines, especially on the veins, sometimes with small spines on the underside of the midrib; petiole to c. 25 cm, with an elongated sheathing base and a small ligule. Inflorescence a large terminal panicle, 25-SO cm long, glabrous, the main rachis rather short, bearing a few lateral or a terminal cluster of branches c. 15-25 cm long; tertiary branches disposed singly or in subverticils, ending in umbellules and bearing a variable number of lateral umbellules. Flowers c. 10-12 per umbellule; pedicels c. 5-6 mm, articulated below the ovary; calyx a rim bearing 5 narrow or trian- gular teeth; petals and stamens 5; ovary turbinate c.2 mm long; styles 5, erect at first, soon spreading, scarcely united at the base. Fruit spheroidal, c. 6 by 5 mm, deeply furrowed when dry, the persistent styles radiating. Distr. Malesia: Central W. Sumatra (Mt Kerintji), W. Java (Mts Gedeh, Patuha and Tangkuban Prahu). 22 FLORA MALESIANA [ser. I, vol. 91 Ecol. In montane scrub and among scattered trees, 1900-2900 m. Vern. Pabong, pangang njirvan, panggang- tjérmé, S. forma nana STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 394, f. 1. Smaller, probably '/.-1 m, leaves tripinnate, 30 cm long, spiny all over; leaflets 4-13 by 2-7 mm, rachides of the ultimate pinnae winged. Distr. Malesia: Central W. Sumatra (Mt Talang). Ecol. Growing about 2500 m. Note. VAN STEENIS considered this interesting dwarf form to represent the extreme of a series in size variability. 6. Aralia montana BL. Bijdr. (1826) 870; Mia. FI. Ind. Bat. 1, 1 (1856) 750; Ann. Mus. Bot. Lugd.- Bat. 1 (1863) 9, incl. var. acutata MIQ.; BOERL. Handl. 1 (1890) 646; K. & V. Bijdr. 7 (1900) 51; Koorp. Exk. Fl. Java 2 (1912) 718; Atlas 4 (1916) f. 672; STEEN. Bull. Bot. Gard. Btzg III, 17 (1948) 391; BAKH. f. & OoststrR. in Back. Bekn. FI. Java (em. ed.) 7 (1948) fam. 159, p. 17; BAKH. f. Blumea 6 (1950) 367, incl. var. crassifolia BAKH. f.; BACK. & BAKH. f. Fl. Java 2 (1965) 170; PHILipson, Gard. Bull. Sing. 30 (1977) 100. — A. bipinnata REINW. ex BL. Cat. (1823) 43, nomen; ex DE VRIESE, PI. Ind. Bat. Or. (1857) 84, nomen in synon. — Panax armatus WALL. [Cat. (1832) n. 4933, nomen] ex G. Don, Gen. Syst. 3 (1834) 386. — A. decomposita REINW. ex DE VRIESE, Pl. Ind. Bat. Or. (1857) 84, nom. illeg. altern. — A. armata (WALL.) SEEM. J. Bot. 6 (1868) 134; CLARKE, FI. Br. Ind. 2 (1879) 723; KING, J. As. Soc. Beng. 67, ii (1898) 44; RIDL. Fl. Mal. Pen. 1 (1922) 873; Hut-Lin L1, Sargentia 2 (1942) 106. — A. thomsonii SEEM. J. Bot. 6 (1868) 134; CLARKE, FI. Br. Ind. 2 (1879) 723; KiNG, J. As. Soc. Beng. 67, ii (1898) 44; RipL. Fl. Mal. Pen. 1 (1922) 873; Hut-Lin Li, Sargentia 2 (1942) 12: Shrub or small tree, frequently unbranched, with prickly stems, occasionally attaining a height of 12 m. Leaves to 1 m or more long, forming large crowns at the ends of the branches, bipinnate, with a pair of simple or occasionally pinnate leaflets at each division of the rachis, usually with some prickles, especially on the petiole, or unarmed, the rachis constricted at the joints; leaflets sessile or with a short petiolule, ovate, up to 14 by 7 cm, apex acute to acuminate, base truncate or rounded, oblique in lateral leaflets, margin sharply serrate, upper surface with the remains of a strigose tomen- tum, often + rugose, lower surface often witha + velvety tomentum, or with more harsh hairs + confined to the veins; petiole to 30 cm, its base sheathing and ligulate. /nflorescence a large ter- minal panicle, peduncle and branches tomentose, prickles, if any, confined to the peduncle and main rachis, small usually persistent linear or ovate bracts c. 11/, cm long subtending the branches and also spaced along the peduncle; bracts of tertiary branches similar but smaller; secondary branches at intervals along the main rachis, c. 35 cm long, bearing numerous tertiary branches along their length; tertiary branches usually c. 6 cm long, ending in umbellules, and often bearing a number of lateral umbellules; umbellules with c. 20-30 radiating pedicels; pedicels usually 12-15 mm, occasionally shorter, pubescent. Flowers herm- aphrodite; calyx lobes 5, triangular or rounded; petals 5, c. 2 mm long; stamens 5; ovary 5-celled, glabrous; styles subulate, connate below, free and spreading above. Fruit spheroidal, up to c. 4 by 4 mm, strongly 5-ribbed when dry, surmounted by the reflexed styles. Distr. Malesia: Malay Peninsula, Sumatra, Java, Borneo (Sarawak, Sabah), Celebes, Lesser Sunda Is. (Sumba). Ecol. Primary and secondary forest, bamboo forest and low-lying moist ground, from near sea- level to 2600 m. Vern. Gorang, panggang tjutjuk, S; Malay Peninsula: poko dulang-dulang; Sarawak: tepa paluk. Notes. This species is considered to include all West Malesian examples with pedicelled flowers and pubescent leaves. This broad concept is contrary to former treatments which have recog- nized several species (A. thomsonii, A. armata). The alliance with A. chinensis L. and A. decaisneana HANczE is also very close. The application of names to this and other Javanese species has been very confused. The position is ably discussed by VALETON (in K. & V. Bijdr.) and by VAN STEENIS (1.c.). In most specimens the lower leaf surface and the pedicels are densely tomentose, but there are specimens in which the leaf is only sparsely hairy and the pedicels may be glabrous. A. armata appears to be within the range of variation of the complex although this plant is very spiny, its leaves and inflorescences always being provided with numerous short spines. The leaflets also are thinner and smoother and, like the pedicels, are less densely pubescent (see NGoc-SANH Bul, Adan- sonia 9, 1969, 461). However, A. armata (if distinct) has been collected only very rarely in the Malay Peninsula and only in the extreme north. The most aberrant specimens are those with glabrous umbel- lules which are mostly from Sumatra but also from Java: they may indicate that A. foliolosa SEEM. should also be included in this complex. Excluded Aralia capitulata JUNGH. & DE VRIESE, Ned. Kruidk. Arch. 1 (1846) 17; Ann. Sc. Nat. III, 7 (1846) 116 is, cf. SLEUMER, FI. Males. I, 7 (1971) 24 = Gomphandra capitulata (JUNGH. & DE VRIESE) BECC. (Icacinaceae). 1979] ARALIACEAE—I (Philipson) 23 Fig. 7. Delarbrea collina ViEILL. a. Habit, x ?/;, b. flower bud, c. flower and ditto in LS, d. petal, e. stamen, f. flower after anthesis, x 13, g. fruit and ditto in CS, x 3 (a-e, g RIDSDALE NGF 36736, f SOEKMA S.i.). Drawn by HELENE MULDER. 24 FLORA MALESIANA [ser. I, vol. 91 4. DELARBREA VIEILL. Bull. Soc. Linn. Norm. 9 (1865) 342, 393; Bru. in B. & H. Gen. Pl. 1 (1865) 935; BRITTEN in Forbes, Nat. Wand. (1885) 506 (see also p. 354); HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 61; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 485; Hutcu. Gen. FI. Pl. 2 (1967) 63. — Fig. 7. Glabrous unarmed shrubs or small trees. Leaves large, imparipinnate, with a stipular sheath; leaflets alternate or opposite, entire or indistinctly dentate. Flowers in umbellules grouped in large terminal panicles; pedicels articulated below the flower. Calyx lobes 5. Petals 5, imbricate, obovate, narrowed towards the base. Stamens 5, filaments stout, anthers dorsifixed. Ovary inferior, 2-celled, disk fleshy, obconic, crowned by two erect styles with clavate stigmas. Fruit ovoid, crowned by the small calyx lobes and the recurved style arms (which eventually fall); exocarp thin, fleshy, with peripheral oil vesicles; endocarp papery; endosperm with shallow longitudinal grooves not ruminate. Distr. Queensland, Melanesia and East Malesia, 3 or 4 spp., from the Lesser Sunda Is. eastwards to New Guinea (also New Britain), Queensland, Solomons, New Caledonia, and New Hebrides. In Malesia 1 sp. Ecol. Lowland to montane forest. Note. The corolla is distinctive, the petals being strongly imbricate and narrowed towards their insertion. The fruit also has a characteristic appearance, since the calyx and stylopodium, although persistent, do not enlarge as in most other araliads. The fruit, therefore, is a smooth ellipsoid berry without a prominent rim around the apex. 1. Delarbrea collina VIEILL. Bull. Soc. Bot. Norm. 9 (1865) 342; Puiipson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 18. — D. sp. HEMsL. Rep. Challenger, Bot. 1, pt 3 (1885) 155. — D. paradoxa (non VIEILL.) BRITTEN in Forbes, Nat. Wand. (1885) 506. — D. lauterbachii Harms in K. Sch. & Laut. Fl. Schutzgeb. (1900) 485. Sparsely branched shrub to 5 m high, with the multijugate leaves clustered at the ends of the branches. Leaves c. 70-100 by 30-40 cm; rachis not articulated; leaflets alternate or in pairs, c. 7 on each side; petiolules c. 1 cm long; lamina c. 17-20 by 4-61/, cm, lanceolate, ovate, oblong or elliptic, gradually tapered to an acute apex, base truncate, rounded or cuneate, usually oblique, margin entire; petiole c. 17-20 cm, terete, lenticellate, with a heavily lenticellate clasping base with mem- branous margins. Inflorescence a terminal panicle of umbellules, rachis up to 60 cm long, bearing well-spaced secondary branches 6-25 cm long, bracts caducous; tertiary branches c. 2-8 cm long, terminating in a circlet of broadly ovate bracts (mostly caducous) surrounding the umbellules, sometimes with smaller (male) lateral umbellules; umbellules c. 2 cm @ at anthesis, with c. 30-40 flowers. Pedicels c. 5 mm (elongating to 10-15 mm in fruit), pustulate. Calyx lobes 5, obtuse, united below into a tube. Petals 5, c. 11/, by 3/, mm, keeled within. Stamens 5, 1 mm long. Ovary sometimes prominently ribbed when dry, c. 2 mm long; disk and styles c. 1 mm high at anthesis. Fruit 16 by 10 mm, purplish black when mature. Distr. Solomon Is. to New Caledonia and Queensland; in Malesia: Lesser Sunda Is. (Timor, Wetar, Babar), Moluccas (Tenimber Is., Banda), New Guinea (Aru Is., Kar Kar Is., Madang, New Britain). Fig. 8. Ecol. Rain-forest, from sea-level to 1000 m. Vern. Don, Madang. Notes. The most wide-ranging of any Malesian araliad. It was collected in Malesia by ForBEs in Fig. 8. Range of Delarbrea collina ViEMLL., localities in Malesia dotted. 1979] ARALIACEAE—I (Philipson) 25 Timor in 1882, when it was incorrectly identified as the New Caledonian species D. paradoxa VIEILL. Eight years earlier it had been collected in the Aru Is. during the Challenger Expedition (HEMSLEY, /.c.) and also been referred as close to D. paradoxa. Nearly twenty years later LAUTER- BACH collected it in the Moluccas, when HARMS described it as a new species. The statement by HarMs that the genus was known previously only from New Caledonia cannot be reconciled with his note in the Pflanzenfamilien in which he recorded BRITTEN’s report of it in Timor. Solomon Islands collections were identified as D. collina VIEILL. by PHILIPSON in 1951. The arrangement of the um- bellules differs in the panicles of D. collina and D. paradoxa. All the material from Malesia con- forms to the characters of D. collina. The plant is evidently rare, few collections having been made in spite of its wide distribution. It was formerly cultivated in the Botanic Garden at Bogor until about 1958, having been introduced from Banda. 5. PENTAPANAX SEEM. J. Bot. 2 (1868) 294; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 55; Koorp. Bull. Jard. Bot. Btzg III, 1 (1919) 181; Hutcu. Gen. FI. Pl. 2 (1967) 63; NGoc-SANH Bul, Adansonia 9 (1969) 389; PHILIPSON, Austrobaileya | (1977) 23. — Fig. 9. Trees or shrubs, often scandent, unarmed. Leaves imparipinnate, exstipulate, glabrous. Flowers in racemes or umbels, which are arranged in panicles or com- pound umbels, pedicels articulated below the ovary. Calyx 5. Petals 5, imbricate in the bud. Stamens 5. Ovary inferior, 5-celled, disk -_ conical, surmounted by the styles which are united their whole length or become free down to half their length. Fruit globose; exocarp leathery, enclosing crustaceous pyrenes. Seeds compressed, endosperm smooth. Distr. About 14 spp. in India, Thailand, Vietnam, Ceylon, Burma, southern China, Taiwan, in Malesia: 1 sp. locally in E. Java. South American species formerly included are best excluded, and the 2 Queensland spp. are now referred to Polyscias. Ecol. Forest and scrub. 1. Pentapanax elegans Koorp. Bull. Jard. Bot. Btzg III, 1 (1919) 182, pl. 16 & 17; Baku. f. & OoststTrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 17; BAckK. & BAKH. f. Fl. Java 2 (1965) 169. — Fig. 9. var. elegans. Epiphytic scrambler or terrestrial shrub up to 10 m, with unarmed branches, leaf and flower buds separate, enclosed in + persistent imbricated cataphylls. Leaves disposed along the branches; petiole c. 6-10 cm, flattened above, base scarcely dilated, and sometimes minutely fimbriated, articu- lated with the rachis, and the rachis articulated with the petiolules, articulations minutely fimbriate, rachis to 5 cm, petiolules of lateral leaflets up to 5 mm, of terminal leaflet to 20 mm, leaflets 5 or fewer, ovate to oblong-elliptic, the lateral some- times oblique, up to 10 by 7 cm, usually c. 5 by 2'/, cm, apex acute, base rounded or cuneate, margin entire or with subulate teeth, glaucous beneath. Inflorescence terminal with persistent cataphylls at the base of the main axis, umbels solitary or 1-4 smaller (apparently male) lateral umbels arising from the axils of minute bracts on the rachis; rachis 8-16 cm, slender, glabrous; terminal umbel 3-6 cm @, many-flowered; pedicels 1'/,-3 cm, filiform, glabrous, with minute brac- teoles surrounding the articulation below the flower. Calyx lobes ligulate, obtuse, c. 1 mm long; petals triangular to ligulate c. 2 mm long; filaments yellow, 3 mm, anthers purple, '/, mm long. Ovary broadly obconic, surmounted by a stylar column, 2 mm long. Fruit globose, 3-4 mm @, disk broadly conical, crowned by the persistent calyx lobes and an awl-shaped stylar column which may divide at apex. Distr. Malesia: E. Java (Mts Ardjuno and Tengger), the variety in Thailand. Ecol. Uncommon, in light forest or scrub, including Casuarina junghuhniana forest, 1700- 2600 m. Note. A remarkably isolated species of a genus otherwise unrepresented in Malesia. var. pubescens Koorp. Bull. Jard. Bot. Btzg III, 1 (1919) 183 (as var. puberula in f. 17); NGoc-SANH Bui, Adansonia 9 (1969) 389. 26 FLORA MALESIANA [ser. I, vol. 91 EAN Px 7, IN, Fig. 9. Pentapanax elegans Koorb. a. Habit, 1/3, b. flower bud, c. flower in anthesis, d. ovary in CS, enlarged (VAN STEENIS 10879). Drawn by P. PRENDERGAST. 1979] ARALIACEAE—I (Philipson) 27 Peduncle and pedicels tomentose. Note. Since all specimens from Mts Tengger Distr. Thailand; in Malesia: E. Java (Mt Jang). and Ardjuno lack pubescence, the retention of the Ecol. In Casuarina junghuhniana forest, scat- variety appears justified. However, more collections tered, 1900-2300 m. are required from all localities. 6. MACKINLAYA F.v.M. Fragm. 4 (1864) 119; HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 62; Bot. Jahrb. 56 (1921) 413; PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 3; Hutcu. Gen. FI. Pl. 2 (1967) 65. — Anomopanax HaArMs [in Dalla Torre & Harms, Gen. Siph. (1903) 364, nomen;] Ann. Jard. Bot. Btzg 19 (1904) 13; in K. Sch. & Laut. Nachtr. (1905) 332; in E. & P. Nat. Pfl. Fam. Nachtr. 3 (1908) 255; Bot. Jahrb. 56 (1921) 414; Hutcu. Gen. FI. Pl. 2 (1967) 59. — Fig. 10. Glabrous unarmed shrubs, often unbranched (sympodial). Leaves with a petiole having a dilated sheath encircling the stem and (in dried material) a constriction at the apex, and with a leaf-blade either unifoliolate or digitately compound, the central leaflet, or the three central leaflets, sometimes digitately lobed or compound. Inflorescence terminal (but sympodium often continued by axillary branching), the peduncle bearing umbellately arranged branches which terminate either in umbel- lules or in cymes. Flowers male or hermaphrodite, the male flowers either in distinct inflorescences or towards the periphery of mixed inflorescences. Pedicel articulated below the flower. Calyx lobes 5—6, triangular or lanceolate. Petals 5—6, narrowed below into a distinct claw, and above into a long incurved process. Stamens 5-6; anthers subglobose. Ovary inferior, with two uni-ovulate cells. Disk prominent, with a crenulate margin. Styles 2, subulate, free, recurved in fruit. Fruit strongly compressed, 2-seeded (or one aborted), with a longitudinal furrow between the seeds; exocarp leathery, endocarp cartilaginous. Endosperm smooth. Distr. 5 spp., Solomon Is., Queensland, and in Malesia (3 spp.): Philippines, Celebes and throughout New Guinea. Ecol. Understorey of rain-forest and montane forest, or epiphytic. Also in secondary growth. Notes. The leaves are palmately divided or they may be reduced to a single leaflet, especially on the upper branches of M. schlechteri. The central leaflet, or the three central leaflets, are either lobed or com- pound, a character rarely found in other genera of this family. Another foliar character rare in the family is the insertion of the leaf-sheath round the whole circumference of the stem. This character occurs also in a section of Polyscias, but is more typical of Umbelliferae. The narrow base of the petal is also very rare in the family, but is characteristic of Umbelliferae. The constantly 2-celled ovary is also typical of Umbel- liferae, but other characters of the fruit appear to justify the retention of these plants in the Araliaceae. Reasons for regarding Anomopanax as congeneric with Mackinlaya are given by PHILIPSON (i.c.). KEY TO THE SPECIES 1. Ultimate branches of the inflorescence in irregularcymes............ 1. M. celebica 1. Ultimate branches of the inflorescence in umbellules. 2. Elimary rays of the inflorescence many (30-50). « ..-..9h 0. 6 2 sam we 2. M. radiata 2. primary taysioL. the inflorescence ¢ 15 or fewer . .. 52 2... + + wes = 3. M. schlechteri 1. Mackinlaya celebica (HARMS) PHILIPSON, Bull. (1904) 15. — Anomopanax warburgii HARMS, /.c. Br. Mus. Nat. Hist. Bot. 1 (1951) 8.— Anomopanax 15. — M. amplifolia HEMSL. Kew Bull. (1909) celebicus Harms, Ann. Jard. Bot. Btzg 19 (1904) 260; Harms, Bot. Jahrb. 56 (1920) 413. — Anomo- 14; Ic. Bog. 2 (1906) t. 176 & 177. — Anomopanax panax arfakensis Gipps, Arfak (1917) 163. — philippinensis Harms, Ann. Jard. Bot. Btzg 19 Anomopanax digitata MeRR. Philip. J. Sc. 17 28 FLORA MALESIANA [ser. I, vol. 91 Fig. 10. Mackinlaya celebica (HARMS) PHILIPSON. a. Part of inflorescence, x 2/s, b. leaf, c. leaflet, x 2 IRs d. flower cluster, e. flower, x 7, f. petal, x 13, g. fruit, x 5/, (BRASs 28056). Drawn by HELENE MULDER. 1979] ARALIACEAE—I (Philipson) 29 (1920) 301. — Polyscias cibaria WHITE & FRANCIS ex LANE-POOLE, For. Res. (1925) 129, descr. angl. minim. — Anomopanax variifolius C. T. WHITE, J. Arn. Arb. 10 (1929) 256. — M. digitata (MERR.) PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 7. — M. warburgii (HARMS) PHILIPSON, /.c. 8. — Fig. 10. Shrub or small sparsely branched tree to 6 m. Leaves very variable in size and complexity; petiole up to 52 cm, 1*/,; cm wide, terete, striate, with a membranous base ensheathing the stem; leaflets 5 (rarely 3) or the central petiolule (or the central 3 petiolules) frequently dividing to bear three, or more rarely 5 leaflets; lateral petioles short (c. 1-2 cm), the three central longer (up to 12 cm); lamina elliptic or ovate, up to 48 by 22 cm, base abruptly attenuated into the petiolule or subcordate, apex acuminate or gradually narrowed, acute, margin entire, denticulate or coarsely serrate, especially towards the apex, membranous, lamina of the terminal leaflet (and less frequently of the central three leaflets) sometimes deeply 3-lobed or with 3-5 separate leaflets, of which the lateral are markedly oblique at the base. Inflorescence a terminal compound umbel, very variable in size, either entirely of male flowers or with male and hermaphrodite flowers, often overtopped by sym- podial growth; peduncle terete, striate, stout, up to 30(-45) cm, 3/, cm @, bearing lanceolate bracts below the rays; primary rays c. 9-18, 10-20 cm, striate, with distal small linear bracts; secondary trays about 5-10, 3-6 cm, dividing again (often repeatedly) either umbellately or in an irregular cymose manner, the central ray frequently more strongly developed. Calyx lobes 5, triangular, c. 1 mm long. Petals 5, obovate c. 1!/, mm long. Filaments c. 11/, mm, anthers small. Ovary obconic, 1-2 mm long, narrowly turbinate in male flowers, ovoid and quickly swelling in female flowers. Fruit up to 2!/, by 3 cm, compressed, rotund, constricted in the mid-axis, the two halves frequently unevenly developed. Distr. Solomon Is.; in Malesia: New Guinea (incl. New Britain and Aru Is.), Celebes, and Central & S. Philippines. Ecol. Rain-forest, open hill forest, and montane forest, also in secondary growths, from sea-level to 1450 m. Uses. LANE-POOLE (/.c.) recorded that at Mt Obree leaves and flowers are cooked with coconut oil and put in armlets in dances. Vern. Philippines: binlaon, C.Bis., pararau, Bag., tagima, Sub., lumot-lumot, Mindanao; New Guinea: bugini, wale, yam bonga, Sepik Distr., lak-lak, W. Highlands, po’undo, S. Highlands, Papua, nere, Central Distr., Papua, narona, New Britain. Notes. The leaves have a strong parsley-like odour. The flowers are creamy white, and the fruits blue to purple with a glaucous bloom. Salt is said to be obtained from the ashes of the leaves. Although collected frequently in the Philippines and New Guinea, this species is unrecorded for the Moluccas. There is considerable variation in the size of the leaves and of the inflorescence. A few New Guinea specimens are intermediate in charac- ter between this and the equally common M. schlechteri, and are interpreted as hybrids. 2. Mackinlaya radiata PHILIPsON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 6. Slender shrub to 5 m. Petiole c. 20 cm, terete and finely striate, base ensheathing the stem, mem- branous. Leaflets 5, or the central petiolule bearing three leaflets, the two lateral petiolules short (1- 1'/, cm), the three central longer (6-8 cm, or the midpetiolule to 11 cm); lamina of the lateral leaflets elliptic or ovate, up to 20 by 12 cm, base abruptly attenuated into the petiolule, apex gradually narrowed, acute, margin entire or minutely denticulate towards the apex, mem- branous; lamina of the central leaflet similar or deeply 3-lobed or with 3 separate leaflets of which the lateral are strongly oblique at the base. /n- florescence a terminal compound umbel; peduncle terete, striate, stout, from 20 cm to considerably longer, 4-6 mm @, bearing lanceolate bracts below the rays; primary rays numerous (c. 30-50), 9-18 cm, slender, striate, with distal minute, linear, caducous bracts; secondary rays (pedicels) numerous (35-130), filiform, 1-2 cm; outer flowers male, central hermaphrodite. Calyx lobes 5, narrowly triangular, c. !/, mm long. Petals 5, obovate, c. 1 mm long. Ovary narrowly obconic in male flowers, ovoid in hermaphrodite flowers, c. 0.7 mm long. Fruit (immature) ovate, com- pressed. Distr. Malesia: New Guinea (NW. Irian; Sepik Distr.). Ecol. Montane rain-forest and mossy forest, 900-1200 m. Vern. Apiyetimber, Sepik Distr. Note. Flowers creamy white. 3. Mackinlaya schlechteri (HARMS) PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 7. — Anomo- panax schlechteri HARMS in K. Sch. & Laut. Nachtr. (1905) 332, t. 13. — Anomopanax versteegii Harms, Nova Guinea 8 (1910) 276. — M. versteegii (HARMS) PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 7. — M. brassii Puitipson, l/.c. 6. — M. klossii Putieson, /.c. 6. — M. subulata PHILIPSON, /.c. 7. Slender shrub to 6 m. Leaves 1-3-(rarely 4-)folio- late. Petiole usually less than 10 cm, but occasion- ally longer (to 20 cm), especially in compound leaves, terete and finely striate, base ensheathing the stem, membranous. Lamina elliptic obovate, or oblong, occasionally irregularly lobed, up to 28 by 12 cm but usually considerably smaller, base cuneate or truncate (of lateral leaflets often 30 FLORA MALESIANA [ser. I, vol. 91 K Fig. 11. Osmoxylon novoguineense (SCHEFF.) BECC. a. Leaf half, b. base of petiole, both x 2/., c. inflores- cence, x ‘/s, d. flower and CS of ovary, x 4, e. false fruit and ditto in CS (CRAVEN & SCHODDE 789). Drawn by W. R. PHILIPSON. 1979] ARALIACEAE—I (Philipson) 31 oblique), apex shortly acuminate, acute, margin entire or dentate towards the apex, membranous or chartaceous. Inflorescence a terminal compound umbel, often overtopped by a lateral branch at its base; peduncle terete, striate, c. 10-20 cm, bearing small lanceolate bracts below the rays; primary rays 6-15, c. 4-7 cm, striate, with minute distal caducous bracts; secondary rays (pedicels) c. 10-20, filiform or rather rigid, usually 5-12 mm; male flowers towards the outside of the umbellules. Calyx lobes 5, triangular to subulate, +/,-1 mm long. Petals 5, obovate. Ovary narrowly turbinate, in male flowers obconic or ovoid, c. 3/, mm long in hermaphrodite. Fruit large, 15 by 22 mm, com- pressed, rotund, constricted above and below on the central axis; styles persistent, recurved. Distr. Malesia: New Guinea (along the Central Ranges, from the Star Mts east to Meyamya), also in New Britain. Ecol. Rain-forest and montane forest, 600— 2300 m. Uses. The cut stem exudes a viscous sap which is an irritant. The leaves are aromatic. The plant is reported to be poisonous and to have a number of medicinal uses. The boiled leaves are eaten to reduce fever and to relieve ‘korima’. Pieces of leaf placed in a cavity relieve toothache. The leaves are wrapped around taro at planting to encourage growth. Vern. Dako, Wissel Lakes, kolobang, kulbang, Sepik Distr., auke, kenata, muklofo, E. Highlands, narona, New Britain. Notes. The flowers are white and the ripe fruit mauve to purple with a glaucous bloom. A large number of collections made in recent years throughout New Guinea all have regularly compound umbels with the flowers borne on branches of the third degree in the form of strict umbels. In two of the earliest gatherings (SCHLECH- TER 14365 and VERSTEEG 1419) the third degree branches frequently divide again either umbellately or cymosely. These two specimens were described as species by HARMS. PHILIPSON later (1951) kept the forms with regular umbellules separate (de- scribing three species). All these five entities are now considered conspecific, the SCHLECHTER and VERSTEEG specimens being regarded as rare anomalies in a widespread and abundant species. It is possible that the VERSTEEG plant is a hybrid with M. celebica. 7. OSMOXYLON Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; BTH. in B. & H. Gen. Pl. 1 (1865) 944; Becc. Malesia | (1878) 193; BogrL. Ann. Jard. Bot. Btzg 6 (1887) 123; O. K. Rev. Gen. Pl. 1 (1891) 645; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; Bot. Jahrb. 56 (1920) 384; Hutcu. Gen. FI. Pl. 2 (1967) 73; PHILIPSON, Blumea 23 (1976) 99. — Eschweileria Zipp. ex BoERL. Ann. Jard. Bot. Btzg 6 (1887) 112, non Eschweilera Mart. 1828; Handl. 1 (1890) 640. — Pseudosandalum O. K. Rev. Gen. Pl. 1 (1891) 271 (‘Pseudosantalum’), nom. illeg. — Boerlagiodendron HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484; Bot. Jahrb. 56 (1920) 377; Hutcu. Gen. FI. Pl. 2 (1967) 72. — Fig. 11, 13-16. Unarmed, glabrous or tomentose shrubs or trees. Leaves palmately lobed or simple, rarely digitately compound; stipules forming a ligule, and the base of the petiole furnished with one to several spiral or transversal crests or collars (very rarely absent). nflorescence a terminal compound umbel; peduncle short; primary rays each terminating into three branches; the central branch bearing a head or umbellule of almost always sterile bacciform flowers (‘pseudo-fruits’); the two lateral branches each bearing a head or umbellule of hermaphrodite flowers. Calyx an obsolete rim or 0. Corolla with few to many lobes above, tubular below. Stamens 4-30, filaments thick, anthers oblong, exserted. Ovary inferior, not articulated with the pedicel, cells 1-many; disk flat with a central raised boss bearing the pustulate stigmas. Fruit subglobose (ribbed when dry); exocarp fleshy, endocarp crustaceous. Seeds compressed, endosperm smooth or wrinkled. Distr. About 50 spp., of which 40 occur in Malesia, extending from Borneo and the Philippines east- wards through Celebes and Moluccas to New Guinea, the remainder lying further north and east in Taiwan, Micronesia, Melanesia and the New Hebrides. Fig. 12. Sy FLORA MALESIANA [ser. I, vol. 91 Ecol. Mainly understorey trees in primary rain-forest, also in second growth forest, usually at low altitudes, especially in shaded situations and near rivers, 15. O. borneense a characteristic rheophyte. Notes. The foliage, inflorescence and flowers of this genus are all unique within the family. The base of the petiole often bears a spiral crest, or this may form a simple collar; the blade varies from simple to elaborately compound; the inflorescence is composed of trifid rays, the central branch bearing sterile bacciform flowers; the corolla is tubular. The central bacciform flowers (pseudo-fruits) are sterile, except in 12. O. yatesii, in which apparently mature seed was once found. By exception a specimen of 40. O. luzoniense had apparently fertile flowers on the central branches of the inflorescence. KEY TO THE SPECIES 1. Leaves simple, without lobes. 2: Petiole base without crests\(New Guinea)i =): < 552-55) cin 1. O. miquelii 2. Petiole base with 1 or more crests. 3: Petiole) base withiseveralcrests (Philippines), =) 5.c)) ee 8. O. oblongifolium 3. Petiole base with a single collar-like crest. 4. Fertile flowers pedicelled (Philippines). 5. Leaf: obovate to’ oblanceolate; petiole:3)cm:or less) - =n). Gee . 2. O. dinagatense 5.) Leatselliptic, petiole: Sicmyormorey-sea a | 6 6 oe oe 3. O. simplicifolium 4. Fertile flowers sessile (or subsessile). 6, Leaf broadlyvobovate (Solomons)ijsesaes | i een eee 4. O. spathipedunculatum 6. Leaf lanceolate or narrowly obovate. 7. Pseudo-iruits pedicelled'(Moluccas)i. a2) =| =) .eelese ka 5. O. articulatum 7. Pseudo-fruits sessile. 82 Inflorescenceyc, 60em:@u(Moluccas) i.) =) sa) ence 6. O. umbelliferum 8: Inflorescence ¢) 30cm!) Gi (New Ireland))\) 2). oe ee 7. O. lanceolatum 1. Leaves lobed or digitately compound. 9. Leaves digitately compound. 10° Ovary 10=16-celled;@New Guinea)” “22. Gee. ee 13. O. geelvinkianum 10. Ovary 4-5-celled. 11. Leaflets lobed (Philippines). 125) Primary rays’ of, inflorescencee: 10\or fewer... = ane een 9. O. catanduanense 12. Primary rays of inflorescence more than 20. 13s Inflorescencec. 16—20ien. Sc ne 10. O. caudatum 13. Inflorescence ¢..5=6.6M Oi. « coh ech ec es cs Eee 11. O. heterophyllum 11. Leaflets entire. 14° Leaflets elliptic-oblong to ovate (Philippines). < 7% 2). cactus 12. O. yatesii 14. Leaflets linear-lanceolate. 15. Leaflets 4—7, petiole 4—6)cmi(Philippines)-) 5 «2 = re) ee 14. O. lineare 15. Leaflets 9 or more, petiole longer than 10 cm (Borneo). ........ 15. O. borneense 9. Leaves lobed (sometimes smaller simple leaves below inflorescence). 16. Pseudo-fruits sessile. 17. Petiolar crests long-pectinate. Bracts of the peduncle heavily setose (Philippines) 16. O. pulcherrimum 17. Petiolar crests fimbriate or undulate. Bracts of the peduncle not setose (Moluccas). 18. Lateral inflorescence branches with opposite, persistent bracts close to the base 17. O. soelaense 18. Lateral inflorescence branches with scars of opposite caducous bracts near the middle 18. O. globulare 16. Pseudo-fruits pedicelled. 19. Lateral inflorescence branches without an articulation. 20. Petiolar base with a single collar-like crest (Borneo) ......... 19. O. kostermansii 20. Petiolar base with several crests. 21. Petiolar crests long-pectinate. 22. Fertile flowers pedicelled. Stamens at least 7 (KeyIs.)......... 20. O. barbatum 22. Fertile flowers sessile (or subsessile). Stamens 5 (Bismarck Arch.)...... 21. O. pfeilii 21. Petiolar crests fimbriate or undulate. 23. Stamens and ovary cells more than 5. Flowers usually pedicelled (Moluccas) 22. O. palmatum 23. Stamens and ovary cells 4. Flowers sessile or subsessile (Philippines). . . . 23. O. ramosii 19. Lateral inflorescence branches with an articulation (2 bracts or bract-scars). 1979] ARALIACEAE—I (Philipson) 33 24. Fertile flowers pedicelled. 25. Pedicels of fertile flowers longer than 7 mm (New Guinea) ..... 24. O. novoguineense 25. Pedicels of fertile flowers 5 mm or shorter. 26. Ovary cells 7 or more. Dipmteecamobes elliptic (Celebes) Mise tee o ehactes OM. LON a se 25. O. teysmannii 2 pabeamlobes! pinnatilobed (Batian)iere. . . Sie Bin) oe ee: 41. O. insigne 26. Ovary cells 4-5. 28 Petiolaricrests'=-entire (Philippines)). . 5... ss 6's 2% as ee 26. O. humile 28. Petiolar crests with long setae. 29. Leaves coriaceous, margin thickened, teeth obtuse (Philippines). . . .27. O. pectinatum 29. Leaves membranaceous, margin not thickened, teeth setulose (Celebes) 28. O. masarangense 24. Fertile flowers sessile (or if subsessile bracteoles longer than the very short pedicels). 30. Ovary cells 10 or more (New Guinea). 31. Flower buds large (c. 9-12 mm long). Primary rays of inflorescence very strong 29. O. insidiator 31. Flower buds smaller (c. 4-6 mm long). Primary rays of inflorescence more tenuous. 32> Headsioftertileifiowers'+- discoid’atianthesis . ; 3) .-..5.... 30. O. boerlagei BeACAGS OmleruleniOwers GlIOHDOSE ere =) ce le) es lier ae 31. O. sessiliflorum 30. Ovary cells 6 or fewer. 33. Leaf lobes lanceolate. 34. Leaf lobes 4-7; petiole 4-6 cm (Philippines) .........2.... 14. O. lineare 34. Leaf lobes 9 or more; petiole longer than10cm (Borneo) ...... 15. O. borneense 33. Leaf lobes broader. 35. Leaf surfaces retaining some trace of setulose tomentum. Umbellules of pseudo-fruits small (Simmrcrorless)i(NewsGuinea) saa1. «sae eee sa ee 38. O. micranthum 35. Leaf surfaces glabrous at maturity. Umbellules of pseudo-fruits larger (1 cm @ or more). 36. Ovary cells 3 (Philippines). Sieseeolamcrests)-|entire seems ert se Sk ey eee 32. O. camiguinense See beriolanicrestsvonp-pectinate: (5.5 6. 5s ee a a 33. O. fenicis 36. Ovary cells 4 or more. 38. Umbellules of ps€udo-fruits large (3-5 cm @) (Philippines). 39. Primary rays of the inflorescence 9-12 cm long. Leaves with strong radiating veins, usually MOTehans wee! Seer SB aes eee 34. O. eminens 39. Primary rays of the inflorescence 4-5 cm long. Leaf-veins less strongly developed, usually SeWernal ae! ae Se aie. Ate es eee 35. O. serratifolium 38. Umbellules of pseudo-fruits smaller (2!/, cm @ or less). 40. Leaf with a small triangular lobe below the middle of each side of the blade (not strictly paimatelyslobed)i(ehilippines)"— — 2 seemeeeee oe ene ee 8. O. oblongifolium 40. Leaf palmately lobed. 41. Petiolar crests long-pectinate (Celebes)». 6. 2. 2 OS. 36. O. celebicum 41. Petiolar crests fimbriate, entire, or undulate. 42. Articulation of lateral branches of umbels close to the base (Talaud Is.) 37. O. talaudense 42. Articulation of lateral branches of umbels near the middle. 43. Inflorescence over 20cm @ (New Guinea) ......... 31. O. sessiliflorum 43. Inflorescence under 20 cm @ (Philippines). 44. Leaf usually 3-lobed. Inflorescence rays delicate, indistinctly setose to glabrous 39. O. trilobatum 44. Leaf usually 5—7-lobed. Inflorescence rays sturdy, markedly setose 40. O. luzoniense 1. Osmoxylon miquelii BoeRL. Ann. Jard. Bot. Sparsely branched tree, 15 m. Leaves glabrous, Btzg 6 (1887) 125, t. 16; Harms, Bot. Jahrb. 56 simple, subcoriaceous; stipules small forming a (1920) 384; Puitipson, Blumea 23 (1976) 103. — __ bicuspid ligule; petioles long (to 19 cm), swollen O. amboinense Mig. Ann. Mus. Bot. Lugd.-Bat. 1 distally; blade oblong-elliptic, 22-36 by 9-12 cm, (1863) 6, p.p.; BEcc. Malesia 1 (1878) 194, p.p.— subrounded at base and apex or mucronulate, Gastonia simplicifolia Zipp. ex SEEM. J. Bot. 3 midrib prominent below, secondary veins arched- (1865) 75, nomen in synon.; ex BoERL. Ann. Jard. ascending and uniting, c. 1-2 cm apart, margin Bot. Btzg 6 (1887) 125, nom. inval. in synon. —_ entire to undulate. Umbel terminal, sessile, with Pseudosandalum miquelii (BOERL.) O. K. Rev. Gen. many (28-32) radiating rigid, angular, trifid Pl. 1 (1891) 271. branches c. 7 cm long to first joint. Central 34 FLORA MALESIANA [ser. I, vol. 91 ajo * get 0 -4 Fig. 12. Species density of Osmoxylon Mia. in Malesia; above the hyphen the number of endemic species, below it the non-endemics. branches unknown. Lateral branches c. 5 cm long, articulate near the base. Flowers 20-30, sessile on the expanded ends of the inflorescence branches. Corolla and stamens unknown. Drupes crowded, subrotund, c. 4mm @ (dry), c. 8-10 ribbed when dry, crowned by a semiglobose entire stigma, 8-10-celled. Seeds with slightly ruminate endo- sperm. Distr. Malesia: West New Guinea. Only known from the type (coll. ZIPPELIUS). 2. Osmoxylon dinagatense (MERR.) PHILIPSON, Blumea 23 (1976) 103. — Boerlagiodendron dinagatense MERR. Philip. J. Sc. 17 (1920) 301; Me_rr. En. Philip. 3 (1923) 222. Glabrous shrub, c. 2 m. Leaves crowded at the ends of the branches; petiole 2-3 cm, channelled above, with a small triangular base, bearing a short stipular ligule (2-3 mm long) and extending around the base of the petiole as a single narrow collar; blade obovate to oblanceolate, to 23 by 8'/, cm; base narrowed into the petiole, apex rounded with a very short apiculum, margin slightly revolute, entire to obscurely undulate with minute teeth, coriaceous. Inflorescence a terminal compound umbel; peduncle c. 1!/, cm, with 1-few bracts (reduced leaf-bases) with small triangular bracts (3 mm long) among the primary rays; primary rays about 15, 8-10 mm long, flattened, bearing opposite bracts (2 mm long) at the apex, each bearing three branches; central branch c. 4 mm long, bearing a head of sessile, bacciform flowers c. 21/, mm @; lateral branches 11/,-2 cm long with two opposite small bracts about the middle and ending in an involucre of minute rounded bracts around a terminal umbellule of c. 7-10 flowers; pedicels 1*/,-2'/, mm long. Calyx a minute rim. Corolla and stamens unknown. Ovary 3-4-celled. Ripe fruit unknown. Distr. Malesia: Philippines (Dinagat I.). Note. A species clearly demonstrating the con- generity of Osmoxylon and Boerlagiodendron. 3. Osmoxylon simplicifolium (ELMER) PHILIPSON, Blumea 23 (1976) 103. — Boerlagiodendron simplicifolia ELMER, Leafl. Philip. Bot. 7 (1914) 2329; MErRR. En. Philip. 3 (1923) 224. Glabrous shrub, to 5 m, with numerous Jeaves clustered near the ends of the branches; petiole 5-7 cm, 2 mm wide, terete, with a small clasping base, an inconspicuous stipular ligule, and a single broad disk-like crest around the lower part of the petiole; blade simple, elliptic, base broadly cuneate, apex acute to apiculate, to 20 by 61/, cm, coriaceous, margin thickened, coarsely dentate, midrib prominent, principal nerves c. 8-10 mm apart. Inflorescence a terminal compound umbel, spherical, c. 7 cm @; peduncle 11/,-2 cm; primary rays c. 25-30, c. 1 cm long with two small obtuse bracts at the apex, ending in three branches; the central branch c. 6 mm long, bearing a subglobose umbel of c. 10-12 sterile bacciform flowers (2 mm @), 3-4 mm pedicelled; lateral branches c. 2—21/, cm long, articulated about the middle, terminating in an umbel of c. 10-20 flowers; pedicels c. 2'/, mm. Calyx rim obsolete. Corolla 3—4-lobed, tubular below, 2 mm long. Stamens 3-4, exserted, 3 mm long. Ovary subcylindric, 2-4-celled, 1 mm long. Drupe spherical, c. 5 mm @ (dry), 2-4-ribbed when dry; surface of endosperm shallowly wrinkled. Distr. Malesia; Philippines (Mindanao: Agusan Prov., Cabadbaran). Ecol. On wind-swept ridge at 1750 m, on moss- covered soil with stones. Vern. Bolauanon, Mbo. Note. A wide-spreading shrub. Bark thick, yellowish, becoming grey. Wood soft, yellowish. Twigs repeatedly branched, the leafy portion sub- erect, leaves mostly ascending, rigidly coriaceous. Inflorescence branches green. Flowers orange, odourless. Berries becoming purple-black. 4. Osmoxylon spathipedunculatum (PHILIPSON) PHILIPSON, Blumea 23 (1976) 103. — Meryta spathipedunculata PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 12. Glabrous tree, to 20 m, with spreading branches. Leaves crowded at the ends of the branches; petiole to 14 cm with a small clasping base bearing a stipular ligule and a rim-like collar around the base of the petiole; blade obovate 17-30 by 10-15 cm, attenuate at the base, apex obtuse, margin entire, midrib prominent, secondary veins arched and uniting, 1'/,-2 cm apart. Inflorescence a terminal compound umbel; primary rays c. 12, stout, com- pressed 9-17 cm long, bearing three branches at the apex; central branches and sterile flowers unt- known; the two lateral branches 9-14 cm long with an articulation c. 1-2 cm from base, bearing helmet-shaped bracts which fall to reveal the terminal head of c. 12 flowers sessile on an ex- panded receptacle with an involucral rimc.1cm @. Calyx rim 0. Corolla split into 5 lobes above, tubular below. Stamens 5. Ovary subcylindr:c, + 1979] ARALIACEAE—I (Philipson) 35 9-celled; disk raised in the centre to the pustulate stigmas. Drupes in a spherical head, globose, c. 12 mm @, c. 9-ribbed when dry, crowned by the prominent, persistent, confluent stigmas. Distr. Solomon Islands (Bougainville and Guadalcanal). Ecol. Rain-forest, 800-1200 m. 5. Osmoxylon articulatum PHILIPSON, Blumea 23 (1976) 103. Tree with stout branches, glabrous. Leaves well spaced towards the ends of the branches; petiole 10-13 cm, 2 mm broad, narrowly channelled above, with a small triangular base, bearing a short stipu- lar ligule (c. 2 mm long) and extending around the base of the petiole as a collar; blade obovate, to 27 by 9 cm, base narrowly cuneate, apex rounded or acute and shortly apiculate, margin thickened, remotely dentate towards the apex, midrib promi- nent, principal lateral veins c. 1'/,—11/, cm apart. Inflorescence a terminal compound umbel, almost sessile, saucer-shaped bracts caducous; primary rays c. 24, 8-11 cm long, flattened, c. 4 mm broad, bearing three branches at the apex; central branch 4-6'/, cm long, the apex expanded and bearing an umbel of c. 10 sterile bacciform flowers c. 6 by 6 mm (when dry) apparently 1-celled, c. 9-14 mm pedicelled; the two lateral branches c. 7 cm long at anthesis with an articulation c. 8-10 mm above the base, bearing helmet-shaped bracts which fall to reveal the terminal head of c. 15—18 flowers, sessile on an expanded receptacle with an involucral rim c.6mm 2. Calyx rim 0. Corolla split into c. 4 lobes above, tubular below, c. 2'!/, mm long. Stamens 5, exserted. Ovary subcylindric, c. 11/; mm long, 7-8-celled, disk with a pustulate central stigmatic boss. Drupes in a spherical head c. 2 cm @ (when dry), strongly 7-8-ribbed (when dry), c. 8 mm @; stigmas persistent, prominent. Distr. Malesia: Moluccas (Halmaheira: Ake Mumar to upper reaches of the Kakatua-matawe). Fr. Sept. Note. This plant is evidently similar to 6. O. umbelliferum described by RumpuHIus. However, a number of differences make it unlikely that it is the same species. The diameter of the inflorescence of the Halmaheira plant is only about half that given by Rumpuius; the lateral rays of the inflores- cence are distinctly articulated near the base, a feature now shown in Rumpuius’ figure; and the sterile bacciform flowers are long-pedicelled, where- ‘as Rumpuius described and figured his as borne in capitula. 6. Osmoxylon umbelliferum (LAMK) Merr. Int. Rumph. (1917) 406; PHitipson, Blumea 23 (1976) 104. — Pseudo-Sandalum amboinense RuMPH. Herb. Amb. 2: 54, t. 12. — Aralia umbellifera LAMK, Encycl. 1 (1783) 225. — Hedera umbelli- ferum (LAMK) DC. Prod. 4 (1830) 262. — Gilibertia saururoides DC. l.c. 256. — Gastonia saururoides Roxs. [Hort. Beng. (1814) 90, nomen;] FI. Ind. ed. Carey 2 (1832) 408 (‘sasuroides’). — O. amboi- nense Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 6, p.p. — Pseudosandalum umbelliferum (LAMK) O. K. Rev. Gen. Pl. 1 (1891) 271. According to RUMPHIUs: Tree with stout trunk, the branches marked with prominent round leaf- scars. Leaves clustered at the ends of the branches, glabrous; petioles long; blades simple, lanceolate (30-36 by 10-12 cm), base rounded, apex acute, margin dentate. Flowers in large spreading umbels, the radiating branches tripartite, c. 30 cm long, each ending in a capitulum. Distr. Malesia: Moluccas. Infrequent on Am- bon, but said to be more numerous in Ceram and the Sula Islands. Only known from RUMPHIUS’ excellent plate and description; not yet re- collected in Ambon. Ecol. Evidently in forest in the hills of Ambon, and also planted at the time of RUMPHIUs. Vern. Sasuru, Leytimor, tonokuku, Hitu. Notes. Valued for the perfume of its wood and foliage. Since this plant is known only from a description and a figure, some uncertainties remain as to its specific characters. In the description it is stated that the young leaves possess a few small teeth of which some signs remain on the older leaves. It is not clear whether the teeth are best developed on distinctive juvenile foliage, or whether the newly expanded normal foliage is intended. The leaves in the figure have prominent teeth, but as they are not shown associated with the inflorescence, they may be from a juvenile shoot. The description of the size of the flower buds is confusing, and it seems likely that sterile pseudo-fruits were mistaken for flower buds. Nevertheless, most characters of the plant are adequately portrayed and there can be no doubt that this species is distinct from the other simple-leaved species with a single, collar-like, petiolar crest. 7. Osmoxylon lanceolatum PHILIPSON, Blumea 23 (1976) 104. — Fig. 13. Small tree with few branches, up to 16 m, glabrous. Many /eaves clustered towards the ends of the branches; petiole 8-15 cm, terete, with a small triangular base, bearing a short stipular ligule (c. 2mm long) and extending around the base of the petiole as a collar; blade oblanceolate, to 33 by 7'/, cm, base narrowly cuneate, apex acute or slightly apiculate, margin entire, midrib promi- nent, lateral veins arched ascending, c. 2-3 cm apart. Inflorescence a terminal compound umbel, almost sessile, saucer-shaped; bracts caducous; primary rays c. 15, c. 10 cm long, flattened, c. 4-5 mm broad, bearing three branches at the apex; central branch 5-6 cm long, the apex expanded and bearing a spherical head of c. 8-12 sessile, sterile, bacciform flowers c. 5 by 5 mm (when dry), 1—2-celled; the two lateral branches 36 FLORA MALESIANA [ser. I, vol. 94 Fig. 13. Osmoxylon lanceolatum PuiLirson. Above: habit of inflorescence and leaves; below, left: twig showing collar-like crests at the base of the petioles; below, right: the trifid branches of the inflorescence (New Ireland, SANDs 795). 1979] c. 5 cm long at anthesis, with an articulation immediately above the base, bearing two helmet- shaped bracts which fall to reveal the terminal head of c. 8-10 flowers, sessile on an expanded receptacle with an involucral rim c. 7 mm @. Calyx rim 0. Corolla known only in bud, c. 2?/, mm long. Stamens c. 5. Ovary gibbous, c. 1'/, mm high, 4-celled. Fruit unknown. Distr. Malesia: New Ireland (Namatanai Subdistr., Danfu R. area, inland from Manga). Ecol. Understorey tree in ridge top forest on limestone, 750-850 m. Note. The bark is pale grey, -- smooth with fine cracks. The twigs and cut branches are strongly aromatic. The wood is soft and dark straw- coloured. The central branches of the inflorescence rays are held + horizontally or depressed and come to maturity before the lateral branches which are held erect. 8. Osmoxylon oblongifolium PHILIPSON, Blumea 23 (1976) 105. Shrub c. 2 m, glabrous when mature, setulose on young parts. Leaves clustered at the ends of the branches; petiole to 16 cm, channelled above, 4mm broad, with a clasping base prolonged upwards as a stipular ligule 11/,-2 cm long, and with entire or fimbriate crests encircling the lower part of the petiole; blade simple, oblong-ovate, occasionally with a small triangular lobe on each side below the middle, to 46 by 17(-—24) cm, base rounded to truncate, apex shortly acuminate, margin serrate, midrib prominent, principal lateral veins c. 3-4 cm apart (at broadest part of leaf). Inflorescence a terminal compound umbel, hemispherical, c. 13 cm @; peduncle 34 cm, c. 6 mm wide, bearing fimbriate, lanceolate bracts (to 2 cm long) along its length and around and among the primary rays; primary rays c. 20, rather short and stout (16-20 by 3-4 mm) with small opposite caducous bracts at apex, each ray ending in three branches; the central branch c. 4 mm long; sterile flowers un- known; the two lateral branches 3!/,-4 cm long, articulated below the middle, terminating in a head of c. 20-30 sessile or subsessile flowers. Flowers unknown. Fruits crowded in a spherical head, drupes c. 5 mm @ (when dry) on pedicels c. 1 mm, 4-ribbed, pyrenes 4, cartilaginous; endosperm with faint reticulate ridging. Distr. Malesia: Philippines (Samar). _ Ecol. In dipterocarp forest, along creek bank, at 200 m. 9. Osmoxylon catanduanense (MERR.) PHILIPSON, Blumea 23 (1976) 105. Boerlagiodendron catanduanense MERR. Philip. J. Sc. 13 (1918) Bot. 318; En. Philip. 3 (1923) 222. Shrub c. 1 m, glabrous except for parts of the inflorescence. Leaves clustered at the ends of the branches; petiole c. 30 cm, terete, 3 mm wide, base with a short ligule, and inconspicuous ARALIACEAE—I (Philipson) a7 recurved crests; blade digitately compound, leaflets 7, chartaceous to subcoriaceous; petiolules 4-5 cm, the lateral shorter; leaflets lanceolate, mid-leaflet to 26 cm long, base cuneate, apex + caudate; irregularly lyrately lobed, the sinuses reaching to within c. 8 mm of the midrib, lobes 2-4 cm long, patent, margin slightly thickened, entire or obscurely dentate. Inflorescence a terminal compound umbel, c. 10 cm @, peduncle stout, bearing ovate bracts c. 1 mm long; primary rays c. 7-10, 2-2'/, cm long, to 2 mm wide, minutely pubescent, with 2 broadly ovate bracts 4-5 mm long at apex; central branch 3 mm or less, bearing a head of sterile flowers; lateral branches 2!/, cm long, bearing 2 broad bracts near the middle, and ending in a spherical head of c. 15 sessile flowers. Calyx an obsolete rim. Petals and stamens un- known. Ovary 4-celled. Fruit ovoid, 4-ridged when dry, 7 by 5 mm. Distr. Malesia: Philippines (Catanduanes). Ecol. On forested slopes, at c. 350 m. Note. Together with 10. O. caudatum and 11. O. heterophyllum this species forms a small group with leaves composed of lyrate leaflets. Although the foliage is similar (though not identical), the inflorescences are distinctive. In O. heterophyllum the rays are short, resulting in a compact compound umbel; in O. caudatum there are few rays (10 or fewer); in O. catanduanense there are many, relatively long rays, resulting in a large, diffuse compound umbel. Since O. catanduanense and O. caudatum are known only from the type collec- tions, the range of variation of these species is not known, but the inflorescence differences justify the retention of all three species. 10. Osmoxylon caudatum (MERR.) PHILIPSON, Blumea 23 (1976) 105. Boerlagiodendron caudatum Merk. Philip. J. Sc. 14 (1919) 440; En. Philip. 3 (1923) 222. Erect shrub, c. 2 m, becoming glabrous. Leaves clustered towards the ends of the branches; petiole c. 45 cm, terete, striate, 4-5 mm wide, base with a stipular ligule c. 11/, cm long, and 2-3 fimbriate or pectinate crests; blade digitately compound, leaflets 5-9, subcoriaceous; petiolules of the central leaflets to 5 cm long, lateral leafiets + sessile; leaflets elliptic-lanceolate, mid-leaflet to c. 25 cm long, base decurrent on the slightly winged petio- lule, apex caudate-acuminate, the larger leaflets with 1-3 pairs of lyrate lobes reaching almost to the midrib, lobes ascending, margin slightly revolute with often prominent usually incurved teeth. Inflorescence a terminal compound umbel, c. 18 cm @; peduncle stout, c. 2 cm, bearing few ligulate bracts c. 11/,-2 cm long, rough, with short setae on the back; primary rays c. 25, c. 4-5 cm long, glabrous and striate, subtended by ligulate bracts, similar opposite bracts at the apex, c. 1 cm long; central branch 1-1'/, cm long, glabrous, ending in a whorl of obtuse rough coriaceous bracts 38 FLORA MALESIANA [ser. I, vol. 91 c. 3 mm long and a head (c. 1 cm @) of sterile bacciform flowers c. 3 mm @, on pedicels 3-8 mm; lateral branches 4-5 cm long, with opposite fimbriate bracts near the middle, bearing an ellipsoid head of c. 20 sessile flowers, each flower subtended and + enclosed by an ovate fimbriate bract 3-4 mm long. Calyx an obsolete rim. Petals and stamens unknown. Ovary 2-21/, mm long, turbinate, obscurely angled, 4-celled. Fruit unknown. Distr. Malesia: Philippines (Luzon: Ilocos Norte Prov., Mt Palimlin). Ecol. On forested slopes near the summit, at c. 1000 m. Note. Apparently never re-collected. For a dis- cussion of distinctive features see under 9. O. catanduanense. 11. Osmoxylon heterophyllum (MERR.) PHILIPSON, Blumea 23 (1976) 106. — Boerlagiodendron heterophyllum Merr. Philip. J. Sc. 9 (1914) Bot. 329; En. Philip. 3 (1923) 223. Erect tree, c. 5 m, glabrous. Leaves clustered at the ends of the branches; petiole to 25 cm, base clasping, prolonged as a stipular ligule to 2'/, cm long, and bearing several pectinate crests (becom- ing recurved when the bristles may be obscured or shed); blade to 35 cm long, digitately com- pound (or some leaflets imperfectly separated); leaflets 3-7, unequal in size, oblong-ovate to broadly oblong-oblanceolate, acuminate, lyrately lobed and irregularly dentate, the base gradually narrowed to the petiolule; petiolule up to 7 cm. Inflorescence a dense terminal compound umbel; peduncle stout, c. 2 cm, bearing many lanceolate bracts 2-3 cm long; primary rays c. 30, 10-15 mm long, subtended by lanceolate c. 21/, cm long bracts, sometimes with bristles on the back, and bearing similar opposite terminal bracts 8-10 mm long, each ending in three branches; central branch short (not seen fully developed) terminating in an umbellule of c. 15 sterile bacciform flowers (c. 3 mm long), pedicels 3-4 mm; lateral branches 8-10 mm long (? fully developed), articulation present ending in heads (c. 1 cm @) of numerous flowers. Calyx rim obsolete. Corolla lobes 4, 2 mm long. Stamens 4. Ovary 4-celled. Fruit unknown. Distr. Malesia: Philippines (Samar, Biliran and Mindanao). Ecol. Primary forest, under shade near creek, 100-550 m. Vern. Aranas, Bis., kayuang, Mbo, magusayag, C.Bis. Note. The description is partially based on the original publication as I have seen only immature inflorescences. For a discussion of distinctive features, see under 9. O. catanduanense. 12. Osmoxylon yatesii (MERR.) PHILIPSON, Blumea 23 (1976) 106. — Boerlagiodendron yatesii MERR. Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923) 225. Shrub, 1 m, glabrous, except for the inflores- cence. Leaves clustered towards the ends of the stout branches; petiole to 38 cm, channelled above, 5 mm @, base with a stipular ligule 1 cm long, and 1—2 inconspicuous non-fimbriate crests around the back of the petiole; blade digitately compound, leaflets S—7; petiolule 2-7 cm (the lateral shorter); blade elliptic-oblong to ovate, mid-leaflet to 23 by 8 cm, base gradually tapered, apex acuminate- caudate, margin dentate or somewhat undulate. Inflorescence a terminal compound umbel, sub- sessile or peduncle stout 1—3 cm, bearing few ovate bracts 1 cm long; primary rays 5-10, tomentose, c. 3 cm long and 3 mm wide, subtended by ovate bracts 6 mm long, similar opposite bracts at apex; central branch 2-3 mm long, pubescent, ending in a whorl of blunt bracts (3 mm long) and an umbellule of c. 10 sterile flowers 4 mm @, 2-4- celled, pedicels 3-8 mm; lateral branches 2—2!/, cm long, pubescent, with small opposite bracts about the middle, bearing a head of 10-15 sessile flowers, subtended by ovate ciliate bracts. Calyx rim obsolete, sometimes fimbriate. Corolla 4—5- lobed, tubular below, 11/, mm long in bud. Stamens 4-5. Ovary 4-celled. Young fruit (MERRILL, /.c.) shortly pedicelled (2-3 mm). Distr. Malesia: Philippines (Luzon and Catan- duanes). Ecol. In rain-forest and mossy forest, from low altitude to 1250 m. Vern. Magalayag, Dinagat. Note. The leaves are unlike any other Osmoxy- lon, resembling those of Macropanax or Schefflera. The flowers are described as yellow, and the fruit black. The only instance of a pseudo-fruit con- taining apparently normal seeds occurred in this species. 13. Osmoxylon geelvinkianum BeEcc. Malesia 1 (1878) 196; PHiLipson, Blumea 23 (1976) 106. — Eschweileria geelvinkiana (BECC.) BOERL. Ann. Jard. Bot. Btzg 6 (1886) 120. — Trevesia geelvin- kiana (BEcc.) O. K. Rev. Gen. Pl. 1 (1891) 272. — Boerlagiodendron geelvinkianum (BECC.) HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. — Eschweiler(i)a elegans Riv. Trans. Linn. Soc. II, 9 (1916) 63. — Boerlagiodendron elegans (RIDL.) Harms, Bot. Jahrb. 56 (1920) 380. — Boerlagioden- dron stenolobum HARMS, I.c. 382, f. 1 k-t. Glabrous shrub with few slender branches, up to 3 m high. Leaves clustered near the ends of the branches; petiole up to 22 cm, usually shorter, 2-3'/, mm wide, narrowly channelled above, with a sheathing base prolonged as a membranous stipular ligule up to 4 cm or longer and with fimbriate or + entire crests encircling the lower part of the petiole; blade up to 30 cm @ (usually 20 cm or less) very deeply 5-11-lobed, or with distinct digitately arranged leaflets, the lobes or 1979] leaflets linear-lanceolate to lanceolate-obovate, entire or irregularly pinnatifid with narrow finely- tapering lobes, base gradually narrowed, apex narrowly caudate, margin serrate; leaves below the inflorescence sometimes reduced to a single leaflet. Inflorescence terminal hemispherical, c. 12-20 cm @; peduncle 1 cm or less, with caducous lanceolate bracts mostly clustered below the primary rays, 1-2 cm long; primary rays rather few, spaced, 2'/.-6 cm long, slender, bearing two caducous lanceolate bracts at the apex, up to 1'/, cm long, each ray ending in three branches; the central branch c. 4-6 mm long bearing a whorl of lanceo- late caducous bracts and an umbel of c. 7-12 sterile, globose or ovoid bacciform flowers (c. 7 mm @ when dry) with pedicels c. 5 mm long and 6-9- celled; the two lateral branches c. 3-4 cm long at anthesis, articulated about the middle, terminating in a small head of 10-20 sessile or subsessile flowers. Calyx rim obsolete; corolla splitting into c. 4 irregular lobes above, tubular below, c. 2!/, mm long. Stamens 10-14, exserted, 3 mm long, anthers small. Ovary cylindric, c. 2 mm long, 10—16-celled; disk with a central raised boss formed by the pustulate stigmas. Fruit globose, fleshy (ribbed when dry), c. 10 mm @. Distr. Malesia: New Guinea (Irian Jaya, to Sepik and Fly R. areas). Ecol. Primary forest, along creeks and river banks, flood-resistant, from near sea-level to 850 m. Vern. Amamutapu, Kamora, korinki, Orne, ida’ pforpforsami, Kutubu. Note. The narrow leaf segments, almost or quite separated at their bases, are characteristic, even though variable in outline. The flowers are described as orange or reddish, and the soft fleshy fruits as dark purple, dark blue, or black. 14. Osmoxylon lineare (MERR.) PHILIPSON, Blumea 23 (1976) 106. — Boerlagiodendron lineare MERR. Philip. J. Sc. 3 (1908) Bot. 253; En. Philip. 3 (1923) 223. Glabrous, erect shrub, c. 3 m. Leaves crowded near the ends of the branches; petiole 4-6 cm, the base with a small stipular ligule (5 mm long) and a few fimbriate crests; blade to 20 cm @, digitately compound with 4~7 leaflets (or very deeply divided into as many lobes); leaflets linear- lanceolate, c. 1-11/, cm wide, the base decurrent on the winged petiolule, apex attenuated, margin thickened, denticulate especially above. Jnflores- cence a terminal compound umbel; peduncle short (c. 1 cm), bracteate; primary rays c. 10, 2—23/, cm long, each ending in three branches; the central branch 4-5 mm long bearing a spherical head of numerous fimbriate bracts (sterile flowers fallen); the lateral branches 3—3!/, cm long with a pair of minute fimbriate bracts about the middle, ending in a capitulum of c. 20 sessile flowers subtended by small fimbriate bracts, c. 7 mm @. Calyx rim minute. Corolla and stamens unknown. Ovary ARALIACEAE—I (Philipson) 39 5-celled. Fruit ovoid c. 3 mm long, 5-ridged (when dry). Distr. Malesia: Philippines (Luzon). Note. Apparently never re-collected. The original specimen is without field information. The foliage is similar to that of 15. O. borneense but with shorter petioles and fewer leaflets. 15. Osmoxylon borneense SEEM. J. Bot. 6 (1868) 141; PHILIPsON, Blumea 23 (1976) 107. — O. helleborinum Becc. Malesia 1 (1878) 198. — Eschweileria helleborina (BECC.) BOERL. Ann. Jard. Bot. Btzg 6 (1887) t. 13. — Trevesia helleborina (BECC)) (O02 KU Rev. Gen. PI 1 (1891), 272. Boerlagiodendron helleborinum (BEcc.) HARMs in E. & P. Nat. Pfil. Fam. 3, 8 (1894) 31. — Boerlagio- dendron borneense (SEEM.) MERR. En. Born. (1921) 456. Glabrous, spreading shrub, up to 3 m. Leaves clustered near the ends of the branches; petiole up to 24cm, narrowly channelled above, 2-3 (or 5) mm wide, with a sheathing base prolonged as a mem- branous stipular ligule up to 2'/, cm long, and usually with fimbriate, + entire, or more rarely long-setose crests encircling the lower part of the petiole; blade up to 20 cm @ digitately compound (or the bases of the leaflets joined by a very short web of tissue); leaflets 9-13, linear-lanceolate to lanceolate, gradually narrowed to the base and apex, up to 20 by 3 cm, usually much narrower, margin serrate, principle veins numerous, c. 5— 10 mm apart. Jnflorescence terminal, hemispherical, c. 6-13 cm @; peduncle 1-2 cm or shorter, with caducous lanceolate-ovate entire or fimbriate bracts mostly clustered below the primary rays, 1-11/, cm long; primary rays rather few (5-12), 2'/.-4 cm long, with 2 ovate bracts at the apex, c. 8 mm long, each ray ending in three branches; central branch c. 5-12 mm, bearing an umbel of c. 6-16 sterile, globose, bacciform flowers, c. 3- 5 mm @ (when dry) with pedicels 4-5 mm long and 5-celled; the two lateral branches 11/,—3 cm long at anthesis, articulated about the middle, terminating in a small head of c. 20-25 sessile flowers with minute rounded bracts. Calyx rim obsolete; corolla splitting into few irregular lobes above, tubular below, c. 11/.-2 mm long; stamens 5-6, slightly exserted; ovary turbinate, angled, c. 1 mm long, 5-celled, disk with a central raised boss formed by the pustulate stigmas. Fruit globose, fleshy (ribbed when dry) c. 5mm @. Distr. Malesia: Borneo (Sarawak, Sabah and Kalimantan). Ecol. Characteristic of rocky river banks, not beyond flood-level, often in deep shade, from near sea-level to 950 m. Vern. Medong, Kayan, empasia abor, Iban, kayan, Tamang, koung, Kinabalu, bungor, Murut Bokan, salimpangaya, Murut Kalabakai. Notes. The leaves of some specimens of 13. O. geelvinkianum (New Guinea) resemble this 40 FLORA MALESIANA [ser. I, vol. 91 species closely. This species can be distinguished by the more compact and smaller fertile flowers, and by its leaflets being uniformly unlobed. Both species are characteristic of river banks, and O. borneense has a low spreading habit, with the branches often rooting, resulting in extensive patches of this low shrub. It is a characteristic rheophyte confined to below flood-level. The flowers are described as greenish white or cream and the inflorescence branches are frequently dark purple. 16. Osmoxylon pulcherrimum VIDAL ex F.-VILL. Nov. App. (1880) 102; Sinopsis Atlas (1883) 28, t. 55, f. B; PHimipson, Blumea 23 (1976) 107. — Eschweileria pulcherrima (VIDAL) BOERL. Ann. Jard. Bot. Btzg 6 (1887) 123. — Trevesia pulcher- rima (VIDAL) O. K. Rev. Gen. Pl. 1 (1891) 272. — Boerlagiodendron pulcherrimum (VIDAL) HARMs in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32; Merr. Philip. J. Sc. 3 (1908) Bot. 254; En. Philip. 3 (1923) 224. — Boerlagiodendron sibuyanense ELMER, Leafl. Philip. Bot. 7 (1914) 2328; MERR. En. Philip. 3 (1923) 224. Erect, sparsely branched tree, up to 10 m, glabrous when mature, except for the inflorescence. Leaves crowded at the ends of the branches; petiole to 1 m, channelled above, clasping base heavily lenticellate, prolonged as-a broad stipular ligule c. 2 cm long, usually with strong bristles on the back, and with strong long-pectinate crests encircling the base of the petiole; blade coriaceous, fan-shaped, c. 40 cm long, base broadly cuneate to truncate, palmately 7-11-lobed, lobes extending to within c. 12 cm from the base, lanceolate, coarsely serrate, sometimes irregularly lobulate, slightly narrowed towards the base, apex acute, sinuses rounded. Inflorescence a terminal compound umbel, c. 18 cm @; peduncle very short, bearing heavily setose bracts; primary rays 15-20, c. 3-4 cm long, 3-4 mm broad, setulose, at the apex bearing opposite, ovate-lanceolate bracts 10-15 mm long, each ending in three branches; central branch c. 15-20 mm long, terminating in a globular head (c. 12 mm @) of c. 20-30, sessile, sterile, bacciform flowers (3-4 mm @) 3-celled, subtended by small ovate-lanceolate bracts; lateral branches c. 5!/, cm long (7'/, cm in fruit), with opposite bracts (c. 3-4 mm long) near the middle, terminating in a globose head of c. 40-50 sessile flowers, bracts between the flowers very small, setulose. Calyx rim obsolete. Corolla 4-lobed, tubular below, 2 mm long in bud. Stamens 4. Ovary 4-celled. Fruit globose c. 6-8 mm long, 4-ribbed (dry). Distr. Malesia: Philippines (Luzon, Mindoro and Sibuyan), recorded also from Formosa and Micronesia (Palau), cf. KANEHIRA, En. Micron. Pl. (1935) 384. Ecol. Damp primary forests, 225-800 m. Vern. Cf. MERRILL: paladukai, Bik., salapak, Neg.; cf. ELMER: palad-amok, Vis. Notes. The fan-shaped leaves with several nar- row lobes and prominent main veins resemble those of 34. O. eminens but are less strikingly developed. The inflorescence is considerably smaller with the pseudo-fruits forming a compact head borne on a comparatively long peduncle. The heads of true flowers, and of the fruits, are considerably smaller than those of O. eminens. Although VIDAL’s material is no longer avail- able, the figure and description relate well to later collections. The specimens on which ELmerR based his Boerlagiodendron sibuyanense have the lobes of the leaf rather simpler in outline than is usual, but the fragments of young inflorescence are quite typical of the taxon and the name is reduced to synonymy. 17. Osmoxylon soelaense PHILIPSON, Blumea 23 (1976) 108. A glabrous shrub or small tree. Petiole c. 32 cm, broadly channelled above, clasping base prolonged as a Stipular ligule 11/, cm long, and bearing c. 3 fimbriate crests; blade c. 40 cm long, broadly cuneate at the base, palmately 7-lobed to within c. 12 cm from the base, lobes narrowly ovate to oblong-elliptic, slightly narrowed towards the rounded sinuses, apiculate, margin denticulate. Inflorescence a terminal compound umbel; peduncle c. 2 cm, bearing lanceolate bracts c. 2 cm long (similar smaller bracts subtend the primary rays); primary rays c. 11/, cm long, 5 mm wide, flattened, bearing opposite, terminal, persistent bracts 10-12 mm long, with lenticels and branched bristles on the back, ending in three branches; central branch c. 10 mm long terminating in a head 1 cm @ of 10-15 sessile sterile bacciform flowers (4 by 3 mm when dry; 3-celled) surrounded by an involucre of ovate bracts (3 mm long) and with minute bracts interspersed; lateral branches 4'/.-5 cm long, bearing opposite ovate bracts (4 mm long) c. 8 mm above the base, terminating in a dense head c. 1 cm @ of 25-30 sessile flowers interspersed with inconspicuous obtuse fimbriate bracts. Calyx rim obsolete. Corolla 5-lobed above, tubular below. Stamens 5. Ovary 4—5-celled. Fruit unknown. Distr. Malesia: Moluccas (Sula Is.: Taliabu and Sulabesi). Note. Fora discussion of the distinctive features, see under 31. O. sessiliflorum. 18. Osmoxylon globulare PHiLipson, Blumea 23 (1976) 108. Shrub to 4 m, furfuraceous on the young parts. Petiole to 55 cm long, broadly channelled above, c. 1 cm wide, clasping base heavily lenticellate, prolonged as a stipular ligule 4 cm long sometimes scaly on the back, and bearing numerous irregular undulate crests on the base of the petiole often continued up the petiole, as rough fascicles of bristles as far as the blade; blade 45 cm long, base 1979] ARALIACEAE—I (Philipson) 41 cordate or emarginate, with some bristles under- neath, palmately 7-lobed to within c. 10-15 cm from the base, lobes narrowly ovate to oblong- elliptic, slightly narrowed towards the broadly rounded sinuses, apex acute, margin serrate. Inflorescence a terminal compound umbel, spheri- cal, c. 15 cm @; peduncle stout, 2-3 cm, bracts together with those among the primary rays caducous; primary rays 30-40, rigid only slightly flattened (subterete), 2-4 cm long, 2—2!/, mm wide, bearing opposite bract-scars at the apex, ending in three branches; central branch 8-10 mm long, terminating in a head c. 13 mm @ of c. 20 sub- sessile sterile bacciform flowers (5 by 4 mm when dry, 2—3-celled); pedicels to 2 mm interspersed with minute bracts; lateral branches 3-4!/, cm long, with the scars of opposite bracts near the middle, terminating in a dense head 1-1!/, cm @ (in bud) of 20-30 sessile flowers interspersed with inconspicuous bracts. Calyx rim obsolete. Corolla 6-8-lobed above, tubular below, 2!/, mm long (in bud). Stamens 6-8. Ovary turbinate, obscurely ribbed, 5-8-celled (varying on the same plant). Fruit unknown. Distr. Malesia: Moluccas (Halmaheira, Moro- tai). Ecol. In forest from sea-level to 800 m. Said to be rare in Halmaheira but common in Morotai. Vern. Bungan-gutu, saha-sasate, Djailolo. Note. For a discussion of the distinctive features see under 31. O. sessiliflorum. 19. Osmoxylon kostermansii PHILIPSON, Blumea 23 (1976) 108. Glabrous, small tree, 8 m. Leaves clustered near the ends of the branches; petiole up to 35 cm, narrowly channelled above, c. 3 mm broad, witha sheathing base prolonged as a stipular ligule c. 2 cm long, continued around the back of the leaf-base as a single wide crest with an entire recurved margin; biade up to 30 cm @, base cordate, deeply 5-7-lobed, lobes elliptic, slightly narrowed towards the sinuses and with a short acute apiculum, margin minutely serrate and sometimes with small sub-lobes, sinuses rounded. Inflorescence a terminal compound umbel, hemispherical, c. 14 cm @ at anthesis; peduncle c. 1 cm long, bearing small lanceolate bracts (c. 3 mm long) below and among the numerous (c. 20-24) primary rays; primary rays 4-5 cm long and 1 mm broad, with opposite bracts (2 mm long) at the apex, each ending in three branches; the central branch 5-6 mm long, bearing a spherical umbel of c. 20 small, sterile, bacciform flowers (2 mm @) on pedicels c. 5-7 mm long, 2-celled; the two lateral branches c. 31/,-4 cm long, with no articulation or bracts except for a minute involucre around the terminal umbellule of c. 10-14 flowers; pedicels c. 2-3 mm. Calyx rim obsolete; corolla 4-lobed, 2 mm long in bud; stamens 4; ovary subcylindric, angled, c. 1 mm long, 5-7-celled, disk with a central stylar boss. Fruit spherical, c. 6 mm long, strongly ribbed when dry. Distr. Malesia: Borneo (Kalimantan: Sang- kulirang Distr., Mt Medadam). Ecol. On limestone at 450 m. Note. The foliage is similar to that of 22. O. pal- matum, except for the distinctive petiolar crest. The inflorescence is also similar to O. palmatum, especially in the lack of an articulation on the rays below the umbellules. 20. Osmoxylon barbatum Becc. Malesia 1 (1878) 197; PHILIPSON, Blumea 23 (1976) 109. — Esch- weileria barbata (BECC.) BoERL. Ann. Jard. Bot. Btzg 6 (1886) 117. —Trevesia barbata (BECC.) O. K. Rev. Gen. Pl. 1 (1891) 272. — Boerlagio- dendron barbatum (BECC.) HARMS in E. & P. Nat. Pfi. Fam. 3, 8 (1894) 31. Small, glabrous tree. Leaves clustered near the ends of the branches; petiole to 32 cm, channelled above, 4-5 mm broad, with a sheathing base prolonged as a stipular ligule 2-3 cm long, and with several long setose crests encircling the lower part of the petiole; blade up to 43 cm @, base cordate to truncate; deeply 5-7-lobed; lobes elliptic-lanceolate, narrowed towards the sinuses and tapered to an acuminate apex, margin minutely serrate, sinuses broadly rounded. /nflorescence a terminal compound umbel, hemispherical, to 12 cm @; peduncle 1-2 cm; primary rays c. 20, 3-5 cm long, with three branches at the apex; the central branch c. 1 cm long, bearing a subglobose umbel of c. 15-20, sterile, bacciform flowers (3-4 mm @) on pedicels 5—7 mm long, 2-4-celled; the two lateral branches about 4 cm long with no clear articulation but 1 or 2 obsolescent bracts, terminating in an umbellule of c. 15-20 flowers; pedicels c. 3-4 mm. Calyx rim obsolete; corolla 4-lobed above, tubular below, c. 4 mm long; stamens 7 or more, rarely fewer, exserted; ovary subcylindric c. 3 mm long, angled, 7-celled; disk with a central stigmatic boss. Fruit unknown. Distr. Malesia: SE. Moluccas (Key Is.). Note. Distinguished from the closely related 22. O. palmatum by the long-setose petiolar crests. For discrimination from 21. O. pfeilii see that species. 21. Osmoxylon pfeilii (WARB.) PHILIPSON, Blumea 23 (1976) 109. — Eschweileria pfeilii WARB. Bot. Jahrb. 13 (1891) 396. — Boerlagiodendron pfeilii (WarB.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. A tree developing a crown when mature, up to 16 m, glabrous when mature, young parts slightly setulose. Leaves in terminal clusters; petiole up to 60 cm, channelled above, c. 6 mm broad, with a sheathing base prolonged as a membranous stipular ligule 2-4 cm long, and with many pro- nounced, long-setose crests encircling the lower part of the petiole; blade up to 75 cm @, deeply 42 FLORA MALESIANA [ser. I, vol. 9! 7-11-lobed, base cordate or emarginate; lobes lanceolate to narrowly elliptic-oblong, narrowed towards the sinuses and tapered to an attenuate apex, margin sharply and remotely serrate, sinuses broadly rounded. Inflorescence a terminal com- pound umbel, hemispherical, to 20cm @; peduncle short (2-3 cm); primary rays numerous (c. 30-40), 4—5 cm long, glabrous, with obsolete bracts at the apex, each ending in three branches; central branch c. 10 mm long, bearing a subglobose umbel of c. 20, small, sterile, bacciform flowers (1—11/, mm @) on pedicels c. 3-4 mm long, 2-5-celled; the two lateral branches c. 3-4 cm long, with no clear articulation but 1 or 2 obsolescent bracts, termi- nating in a head of c. 12-16 subsessile flowers (pedicel c. 1 mm, becoming longer in fruit) sur- rounded by an inconspicuous involucral rim. Calyx rim obsolete; corolla 5-lobed, 11/, mm long in bud; stamens 5; ovary subcylindric, angled, 11/, mm long at anthesis, 5—16-celled, disk with a central boss formed by the united pustulate stigmas. Fruit spherical, fleshy, c. 8 mm @, ribbed when dry, the stigmatic boss persistent and prominent; pyrenes cartilaginous. Distr. Malesia: Bismarck Archipelago (New Britain, Duke of York Group and New Ireland). Ecol. Primary rain-forest, from near sea-level to 600 m. Vern. Sare, sasare, sare a lauvolau, New Britain, Pomio; a ibalur, New Ireland. Notes. The bark is grey-brown and pustular, the wood straw-coloured and soft. The flowers are orange, and the ripe fruit dark red-violet. In the original description the ovary is recorded to possess 10-14 cells. However, some other specimens have as few as 5 cells in the ovary, but in other respects agree with specimens with the large number of seeds. Since the inflorescence, leaf-shape, and especially the nature of the petiolar crests, as well as the distribution, are all highly distinctive within the genus, all the specimens can be accepted as examples of one species with a highly variable number of carpels. This species is very close to 20. O. barbatum of the Key Islands. The original diagnostic character of the number of cells in the ovary has been found to be unreliable. However, since the primary rays in the inflorescence are more numerous and the pedicels of the fertile and sterile flowers are shorter this geographically distinct species is maintained. 22. Osmoxylon palmatum (LAMK) PHILIPSON, comb. nov. — Folium polypi mas (et femina?) RUMPH. Herb. Amb. 4: 101, t. 43. — Aralia palmata Lax, Encycl. 1 (1783) 224, type, non Lour. 1790, nec R. & S. 1820. — Trevesia moluccana Mia. FI. Ind. Bat. 1, 1 (1856) 748; Bonplandia 4 (1856) 137. — Trevesia zippeliana Mia. Ann. Mus. Bot. Lugd.- Bat. 1 (1863) 11. — Unjala bifida REINW. ex DE VrieEsE, Pl. Ind. Or. (1867) 83, nomen in synon.; ex Boeri. Ann. Jard. Bot. Btzg 6 (1887) 166, in synon. — O. moluccanum (MiqQ.) Becc. Malesia 1 (1878) 195; PHiLipson, Blumea 23 (1976) 109. — O. zippelianum (Miq.) BEcc. Malesia 1 (1878) 195. — Eschweileria palmata Zipp. ex BOERL. Ann. Jard. Bot. Btzg 6 (1887) 116, t. 14. — Boerlagio- dendron palmatum (Zipp. ex BOERL.) HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31; Merr. Int. Rumph. (1917) 407. — Boerlagiodendron moluc- canum (Miq.) BAKH. f. & OoststTR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 3; Blumea 6 (1950) 367; BACK. & BAKH. f. Fl. Java 2 (1965) 163. Small, glabrous tree to 15 m. Leaves clustered near the ends of the branches; petiole up to 40 cm, channelled above, 4-5 mm broad, with a sheathing base prolonged as a stipular ligule up to 5 cm long, and with several fimbriate crests encircling the lower part of the petiole; blade up to 45 cm @, base cordate, deeply 5—9-lobed, lobes elliptic, slightly narrower towards the sinuses, acuminate, margin serrate and sometimes with small sub-lobes, sinuses rounded. Inflorescence a terminal, compound umbel, subspherical, to c. 20 cm @ at anthesis; peduncle c. 1-2 cm, with small caducous bracts below and among the numerous (20-60) primary rays; primary rays 3-5 cm long, 2-3 mm broad, with two opposite caducous bracts at the apex, each ending in three branches; central branch 6-10 mm long, bearing a spherical umbel of c. 20-30 small, sterile, bacciform, flowers (2-4 mm @) on pedicels 4-6 mm long, 4—7-celled; two lateral branches c. 2!/,-5 cm long, rigid and straight, with no articulation (indistinct scars of bracts may occur below the involucre), bearing a minute involucre around the terminal umbellule of c. 12-20 flowers; pedicels to c. 5 mm (occasionally flowers subsessile). Calyx rim obsolete. Corolla irregularly 4~-5-lobed above, tubular below, c. 5 mm long. Stamens 6-9, exserted, filaments stout. Ovary subcylindric, angled, c. 2 mm long, 6-9-celled, disk with a raised central stigmatic boss. Fruit globose, fleshy, c. 10 mm @, strongly ribbed when dry. Distr. Malesia: Celebes (once, not localized) and Moluccas (Buru, Ceram, Ambon, Banda, Tenimber Is.). Also cultivated in the Bogor Botanic Garden. Ecol. An understorey tree in primary rain- forest. Uses. The leaves are used for culinary and medi- cinal purposes (against gonorrhoea). Vern. Daun gurita, pelenda darat, saha-saha, Moluccas, fumala-alas, Tenimber Is. Note. The spherical inflorescence is characteris- tic, having straight rigid rays with no articulation on the secondary branches, and the pseudo-fruits are well separated from the true flowers. MIQUEL distinguished Trevesia zippeliana because the collector noted that its ovary was 4-celled. Possibly this number related to the sterile flowers. In two gatherings from Ambon (Waai, TEYSMANN; G. Salhutu, BOERLAGE 179) the fertile flowers are 1979] ARALIACEAE—I (Philipson) 43 subsessile in heads, but otherwise conform to the characters of this species. The only record of this genus from the Tenimber Is. consists of leaves only, but their characters conform to this species. 23. Osmoxylon ramosii (MeERR.) PHILIPSON, Blumea 23 (1976) 110. — Boerlagiodendron ramosii MERR. Philip. J. Sc. 11 (1916) Bot. 27; Me_rr. En. Philip. 3 (1923) 224. Erect, unbranched, or sparingly branched, 4 m high shrub, becoming glabrous. Petiole to 40 cm, the clasping base prolonged as a stipular ligule c. 2 cm long, and with few to several prominent recurved, obscurely fimbriate, crests surrounding the lower part of the petiole; blade to 30 cm long, base emarginate, palmately 3-7-lobed, lobes extending to within 3-8 cm from the base, oblong- ovate, margin serrate, sometimes lyrately lobulate, apex acuminate, sinuses broadly rounded. Jnflores- cence a terminal compound umbel, 10-15 cm @; peduncle stout, with lanceolate bristle-bearing bracts; primary rays c. 15, 2-3 cm long, 2-3 mm wide, flattened, subtended by lanceolate bracts c. 2 cm long, with bristles on the back and bearing similar opposite terminal bracts c. 1 cm long, each terminating in three branches; central branch c.4 mm long, slightly pubescent, terminating in an umbellule (c. 11/, cm @) of c. 10-15 sterile, bacci- form flowers 4-5 mm @, 2-celled, pedicels 5-8 mm long, subtended by caducous bracts; laterai branches 2!/, cm long (slightly longer as fruits form), without any articulation or bracts except for a caducous small involucre around the terminal head of c. 25-35 sessile or very short-pedicelled flowers, c. 1 cm @, bracts among the flowers obscure. Calyx rim obsolete. Corolla 4-lobed above, tubular below. Stamens 4, exserted, filaments broad. Ovary subcylindric, obscurely angled, 4-celled. Fruit spherical 9 mm @, 4-ribbed when dry. Distr. Malesia: Philippines (Luzon). Ecol. On low-lying, wet ground, in forest, or on forested slopes, 700-800 m. Notes. The flat-topped inflorescence is c. 15 cm @ with much-reduced leaves below it. The in- florescence rays are dark purplish and the flowers orange-yellow. The bark is grey and the wood soft. This is the only species in the Philippines without opposite bracts on the lateral branches of the inflorescence rays. In this respect it resembles 22. O. palmatum and a few other species. 24. Osmoxylon novoguineense (SCHEFF.) BECC. Malesia 1 (1878) 197; PHILIPSON, Blumea 23 (1976) 110. — Trevesia novo-guineensis SCHEFF. Ann. Jard. Bot. Btzg 1 (1876) 26. — Eschweileria novoguineensis (SCHEFF.) BOERL. ibid. 6 (1886) 118. — Boerlagiodendron novoguineense (SCHEFF.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31; Back. & BAKH. f. Fl. Java 2 (1965) 163. — Boerlagiodendron lauterbachii HARMS in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484. — Fig. 11, 14. Fig. 14. Osmoxylon novoguineense (SCHEFF.) BECC. Petiolar base with ligule and collar-like crests (Photogr. PHILIPSON, Wantoat, 1968). 44 FLORA MALESIANA [ser. I, vol. 91 Tree or shrub, unbranched or sparingly Trevesia teysmannii (BOERL.) O. K. Rev. Gen. Pl. 1 branched, up to 16 m, the young parts rufous- (1891) 272. — Boerlagiodendron teysmannii furfuraceous, glabrescent. Large leaves forming (BOERL.) HARMs in E. & P. Nat. Pfl. Fam. 3, 8 terminal crowns; petiole up to 1 m, stout (1-2 cm (1894) 31. broad), flattened above, with a sheathing base prolonged as a strong stipular ligule up to 7 cm long, and with fimbriate crests encircling the lower part of the petiole; blade up to 1.20 m @, with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, the lobes in turn deeply lobed and incised, the central lobes especially being strongly pinnatisect or digitately tripartite, apices acute, margins serrate; upper leaves associated with inflorescences may be smaller, more simply lobed, or entire. Inflorescence terminal, a large compound umbel, bowl-shaped, up to 35 cm @; peduncle up to 10 cm, stout, with lanceolate caducous bracts (c. 4 cm long) below and among the numerous (c. 50-70) primary rays; primary rays c. 12-15 cm long at anthesis, c. 3 mm @, bearing two caducous bracts (1 cm long) at the apex, each ray ending in three branches; central branch c. 2 cm long, bearing an umbel of c. 20-40, sterile, bacciform flowers (c. 6 mm @ when dry) on pedicels c. 10 mm, and 2-6-celled; the two lateral branches c. 4-6 cm, with two opposite or sub- opposite bracts about the middle, terminating in a subspherical umbel 2!/,-3 cm @ of 30-50 flowers on pedicels c. 8-10 mm long. Calyx rim obsolete, undulate. Petals with irregular erect lobes, tubular below. Stamens 6-10 exserted. Ovary turbinate somewhat angled; glabrous, 6—14-celled; disk flat with a central double row of pustulate stigmas. Fruits on stiff radiating pedicels, ovoid or spherical, fleshy, ribbed when dry. Distr. Solomon Is.; in Malesia: throughout New Guinea and in the Bismarck Archipelago. Ecol. Primary and second-growth forest, from sea-level to 1600 m. Vern. Lebe, Mooi, teresakui, Manikiong, akriek, Biak, hoppung, Hottam, uger, Wagu, faliifalii, Tifal, ap gan dandam, aimaini, Mamig, ida’ pfopforsami, Kutubu, pulaka, Gazelle Peninsula. Notes. The foliage is similar to that of 30. O. boerlagei, but the pedicelled flowers of O. novo- guineense distinguish it readily from that species. The ripe fruits are usually ovoid, but in the Solomon Is. they are characteristically spheroidal, and this feature recurs in some specimens from the Bis- marck Archipelago and the adjacent coast of New Guinea. The fawn bark is pustulate with many lenticels. An orange exudate flows from the cut stems. The wood is soft and straw-coloured. The inflorescence branches are dark purple, the corolla and stamens usually deep red, and the ripe fruit shining purple or blue-black. 25. Osmoxylon teysmannii (BOERL.) PHILIPSON, Blumea 23 (1976) 111. — Eschweileria teysmannii BoerL. Ann. Jard. Bot. Btzg 6 (1887) 119. — A small, glabrous tree, 6 m. Leaves clustered at the ends of the branches; petiole to 40 cm, chan- nelled above, 4 mm broad, with a sheathing base prolonged as a stipular ligule 2—2'/, cm long, and with several fimbriate or entire crests on the lower part of the petiole; blade c. 30 cm @, membranous, cordate at the base, deeply 7-lobed, lobes elliptic, slightly narrowed to the broadly rounded sinuses, narrowed to a fine apiculum at the apex, margin finely serrulate. Inflorescence a terminal compound umbel, c. 10cm @; peduncle c. 1 cm, bearing ovate bracts (ligules of reduced leaves) and terminating in a cluster of bracts (c. 10 mm long) below and among the primary rays; primary rays c. 12-15, c. 3-4 cm long, with a pair of lanceolate bracts at the apex (c. 1 cm long); central branch c. 1 cm long, terminating in an umbellule of c. 5-8 sterile bacci- form flowers (c. 4 mm @ when dry) on pedicels 6-9 mm long interspersed with linear bracts 5 mm long; two lateral branches c. 31/, cm long, articu- lated about the middle, terminating in an umbellule surrounded by caducous linear bracts (leaving a rim-like involucre); flowers c. 8-12, pedicels c. 2-3 mm. Calyx rim obsolete. Corolla 3 mm long, with 7-8 lobes above, tubular below. Stamens 7-8, filaments stout, anthers exserted. Ovary sub- cylindric, angled, c. 11/, mm long, 7—8-celled; disk flat, with a central stigmatic boss. Fruit unknown. Distr. Malesia: SW. Celebes (Tjamba, Kosali- Porema) and NW. Central Celebes (Palu-Parigi and Mt Nokilalaki). Ecol. In rain-forest, 800-1000 m.. 26. Osmoxylon humile (ELMER) PHILIPSON, Blumea 23 (1976) 111. — Boerlagiodendron humilis ELMER, Leafl. Philip. Bot. 7 (1914) 2327; Merr. En. Philip. 3 (1923) 223. Erect, small, sparsely branched shrub, up to 11/, m. Leaves clustered at the ends of the branches; petiole to 25 cm, terete, base prolonged as a stipular ligule c. 1 cm long, and with c. 3 entire crests at the base; blade palmately 5-lobed, 24 cm long, base truncate or cordate, lobes reaching to within 3—6 cm from the base, elliptic, 4-6 cm wide, narrowed towards the broadly rounded sinuses, tapered to an acute apiculum, margin serrate in the upper part, the outer lobes with a lobule on the lower edge. Inflorescence a terminal compound umbel, 9 cm @, subtended by a few foliaceous bracts; peduncle stout, 2-3 cm, with furfuraceous, oblong bracts; primary branches crowded, numer- ous, 21/, cm, furfuraceous, flattened, striate with opposite minute bracts at the apex; central branch c. 3 mm, bearing an umbellule of sterile, bacciform flowers 3!/, mm @, pedicels 4!/, mm; lateral branches 3-31/, cm, articulated about the middle, terminating in an umbellule of c. 15 flowers, bracts 1979] ARALIACEAE—I (Philipson) 45 inconspicuous, fimbriate, pedicels 11/,-2 mm. Calyx an obsolete rim. Corolla and stamens not known. Ovary 1'/,-2 mm, 4-celled, with a flat disk and a raised central stigmatic boss, 4-celled. Fruit 6 by 4 mm (dry) 4-ribbed; pyrenes crusta- ceous; endosperm rugose. Distr. Malesia: Philippines (Mindanao). Ecol. Damp fertile ground in dense forest, on south side of Baruring R., at 1000 m, Vern. Sardng-ka-mdno, Bag. 27. Osmoxylon pectinatum (MERR.) PHILIPSON, Blumea 23 (1976) 111. — Boerlagiodendron pectinatum Merk. Philip. J. Sc. 3 (1908) Bot. 253, 424; En. Philip. 3 (1923) 224; KANEHIRA, Form. Trees rev. ed. (1936) 520, f. 480; Hur-Lin Lt, Woody FI. Taiwan (1963) 666, f. 273. Shrub or small glabrous tree up to 8 m. Leaves clustered at the ends of the branches; petiole to 18 cm, with a clasping base prolonged as a short acute stipular ligule, and with several basal crests fringed with 1-2 cm long bristles; blade to 25 cm @, base truncate to broadly cuneate, palmately 5-7- lobed, lobes reaching to about the middle of the lamina, sinuses narrow-rounded, lobes oblong- elliptic, usually slightly narrowed below, obtuse to acute, margin thickened, coarsely dentate, coria- ceous. Inflorescence a terminal compound umbel; peduncle c. 1 cm, with small bracts (3 mm) below and among the primary rays; primary rays c. 25- 35, c. 2-3 cm long, with opposite ovate caducous bracts at the apex, each terminating in three branches; central branch c. 8-11 mm long, ending in an involucre of minute bracts (1 mm) surround- ing an umbellule of c. 15-20 ovoid sterile flowers (c. 3 mm long, 3-celled), pedicels 5-6 mm long; lateral branches c. 2!/, cm long at anthesis, with an articulation about the middle, ending in an umbel- lule c. 1 cm @ with minute fimbriate bracts; flowers c. 30, pedicels c. 11/, mm (elongating slightly in fruit). Calyx rim obsolete. Corolla lobes 4-5, tubular below, 2 mm long. Stamens 4-5. Ovary turbinate, 1!/, mm long, 4—6-celled. Fruits globose, 5 by 5 mm (dry), 4~6-ribbed when dry. Distr. Taiwan (Botel Tobago and Lutao I., east off Taiwan proper); in Malesia: N. Philippines (Batan I.). Ecol. Forested slopes at 650 m. Vern. Narapan, Iv. 28. Osmoxylon masarangense PHILIPSON, Blumea 23 (1976) 111. Small tree, 5 m, the young parts setulose, becoming + glabrous. Leaves in terminal clusters; petiole c. 17 cm, rather narrow (1'/,-2 mm @), channelled above, with a small clasping base, prolonged as a stipular ligule, 1-11/, cm long, setulose on the back, and with a number of long- setulose crests encircling the lower part of the petiole; blade c. 18 by 22 cm, deeply 3—5-lobed (or below the inflorescence sometimes simple), the base truncate or emarginate, lobes oblong or elliptic, slightly narrowed to the broadly rounded sinuses, apex with a short apiculum, membranous, margin finely setulose-serrate, sinuses c. 6 cm frob- base of the blade. Inflorescence a terminal sum sessile compound umbel; primary rays c. 10, setulose, 13/,-2 cm long, each ray ending in three branches; central branch 4-5 mm long, ending in an umbellule, pedicels 6 mm, sterile flowers not known; two lateral branches 2-2!/, cm long, 1 mm broad, with two bract scars about the middle but usually not opposite, terminating in an umbellule with c. 10 pedicels 3—5 mm long (in fruit). Flowers unknown. Fruit (when dry) ovoid, 6 by 4 mm, 5-seeded. Distr. Malesia: N. Celebes (Minahasa, Tomo- hon, Mt Masarang). Ecol. Secondary forest at edge of crater lake, at 1200 m. Note. This species is similar in aspect to the Philippine 39. O. trilobatum, but the petiolar crests are distinctive. 29. Osmoxylon insidiator BEcc. Malesia 1 (1878) 195; PHILIPSON, Blumea 23 (1976) 112. — O. carpo- phagarum Becc. Malesia 1 (1878) 196. — Esch- weileria insidiatrix (BECC.) BOERL. Ann. Jard. Bot. Btzg 6 (1886) 120. — Eschweileria carpophagarum (BEcc.) BoERL. /.c. 121, t. 15. — Trevesia insidiator (BEcc.) O. K. Rev. Gen. Pl. 1 (1891) 272. — Trevesia carpophagarum (BEcc.) O. K. lc. — Boerlagiodendron insidiator (BECC.) HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32. — Boerlagiodendron carpophagarum (BECC.) Harms, /.c. — Boerlagio- dendron pachycephalum Harms, Nova Guinea 8 (1910) 271. Small tree to 12 m, young parts with uniform scurfy tomentum. Large Jeaves forming terminal crowns; petiole up to 80 cm, stout (1-2 cm @), broadly channelled above, with a sheathing base prolonged as a strong stipular ligule c. 9 cm long, and with moderately developed irregular (not fimbriate) crests encircling the lower part of the petiole; blade up to 85 cm @, with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, lobes in turn deeply lobed and incised, the median often digitately tripartite, apices long acuminate, margin irregu- larly and remotely serrate, subglabrous when mature or showing remnants of the tomentum. Inflorescence a terminal hemispherical compound umbel, c. 15 cm high by 30 cm wide; peduncle short, stout (11/,-2 cm @), with lanceolate bracts 4-6 cm long below and among the numerous (15-20) primary rays; primary rays 6-10 cm long, c. 5-12 mm wide, rigid, bearing 2 lanceolate bracts (2-3 cm long) at the apex, each ray ending in three branches; central branch c. 1*/, cm long, bearing an umbel of c. 30 sterile bacciform flowers (c. 5— 12 mm @ when dry), the pseudo-fruits and their pedicels + rufous tomentose, pedicels 10-12 mm, 46 FLORA MALESIANA [ser. I, vol. 9 and 6-celled, surrounded by an involucre of short ovate bracts (3-8 mm long); two lateral branches c. 6 cm at anthesis, rigid, slightly flattened, to 8 mm broad, bearing a pair of bracts (c. 11/2 cm long) about the middle, terminating in a subglobose head 3!/,-4 cm @ of c. 30-40 sessile flowers, and surrounded by an involucre of ovate bracts c. 10-14 mm long. Calyx rim fimbriate. Petals irregularly 4~-S-lobed, 7-8 mm long, connate below to form a fleshy tube, pubescent on the outer surface. Stamens c. 15-26, filaments strap- like, projecting beyond the corolla, anthers c. 4 mm long. Ovary shortly turbinate, 2-4 mm long, angled, furfuraceous, c. 13—25-celled; disk flat, with a central boss formed of the pustulate stigmas. Fruits in a compact spherical head, the individual drupes angled by mutual pressure, and bearing the persistent stigmas on the exposed face, c. 10-14 mm long, the numerous pyrenes compressed and flat; cartilaginous. Distr. Malesia: throughout New Guinea, but local; also Waigeo I. Ecol. Primary rain-forest and regrowths, frequently beside streams, from sea-level to 350 m. Vern. Angit, kangit, Waigeo, pennifogo, Orakawa, Papua. Notes. The bark is greyish brown, slightly fissured with many lenticels. The wood is soft and white. The flowers are reddish-brown or purple, with orange-red filaments and the fruit purple. BECCARI provided a detailed description of the living plant, and noted that the fruits are eaten by various species of pigeon. Boerlagiodendron pachycephalum Harms has very strongly developed umbels and leaves, but apart from size, it does not differ from this species. Since a range in stature is shown by the several gatherings now available, the whole is best re- garded as a single species. Similarly, the very short pedicels of O. carpo- Phagarum, which BEccCARI used to distinguish it from O. insidiator, can in fact be matched on several specimens of that species. 30. Osmoxylon boerlagei (WARB.) PHILIPSON, Blumea 23 (1976) 112. — Eschweileria boerlagei Wars. Bot. Jahrb. 13 (1891) 395. — Boerlagioden- dron warburgii HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 32, nom. illeg. superfi.; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 484. — Boerlagiodendron boerlagei (WARB.) Harms, Bot. Jahrb. 56 (1920) 382. — Fig. 15. Small to fairly large tree, unbranched or sparingly branched, up to 24 m, glabrous, at least when mature. Large leaves forming terminal crowns; petiole up to 1 m, stout (1-2 cm @), Fig. 15. Osmoxylon boerlagei (WaARB.) PHILIPSON. Showing that each ray ends into a central umbel of bacciform flowers and two lateral umbels with normal flowers (Photogr. PHILIPSON, Kassam Pass, E. New Guinea, 1968). 1979] ARALIACEAE—I (Philipson) 47 broadly channelled above, with a_ sheathing, heavily lenticellate base prolonged as a strong stipular ligule up to 7 cm long, and with lacerate crests encircling the lower part of the petiole; blade up to 1.15 m @, with 5-7 strong ribs radiating from the top of the petiole, deeply lobed almost to the base of these ribs, lobes in turn deeply lobed and incised, the central lobes especially being strongly pinnatisect or digitately tripartite, apices acute, margin undulate or indistinctly serrate. Jnflores- cence terminal (or overtopped by a lateral leafy branch), a large compound umbel, bowl-shaped with a slightly convex top, up to 60 cm @; peduncle c. 10 cm, stout, with lanceolate bracts below and among the very numerous radiating primary rays; outer primary rays c. 20 cm long at anthesis (elongating in fruit), inner rather shorter, woody, bearing two caducous bracts at the apex, each ray ending in three branches; central branch c. 4 cm, bearing an umbel of c. 20 sterile bacciform flowers (c. 8 mm @ when dry) with rigid pedicels c. 1'/, cm long, and 5—6-celled; two lateral branches c. 9 cm long at anthesis, articulated about the middle, terminating in a button-like head of c. 20-30 sessile flowers and surrounded by ovate bracts which soon fall leaving a bowl-shaped involucre, c. 1/, cm @. Calyx rim obsolete. Petals c. 13, bud flat-topped, angled, minutely pubescent, apparently falling as a calyptra. Stamens 8-13. Ovary shortly turbinate, angled, glabrous, 10—14-celled; disk flat, with a central double row (or ellipse) of pustulate stigmas. Fruits spreading to form a + spherical head obscuring the involucre, each c. 9 by 7 mm (when dry) with prominent persistent stigmas. Distr. Malesia: throughout New Guinea. Ecol. Primary forest, and secondary growths on old cultivations, from near sea-level to 1800 m. Vern. Eunya, Gimi, apiatambay, Washkuk, ma-korr-korr, Jal, teresahui, Manikiong. Note. Bark yellow grey with shallow fissures and many pale corky pustules; wood straw- coloured, fibrous. The large terminal inflorescence is shallowly convex on top and is surrounded by several large leaves. Flowers reddish brown. Fruit purplish black and succulent. 31. Osmoxylon sessiliflorum (LAUT.) PHILIPSON, Blumea 23 (1976) 113. — Boerlagiodendron sessiliflorum LAUT. Nova Guinea 8 (1910) 272. _ §mall tree, up to 18 m, glabrous when mature, or tomentum persistent on the inflorescence. Leaves crowded at the ends of the branches; petiole to 60 cm, with the clasping base prolonged as a stipu- lar ligule up to 6 cm long, and with few to several strong or weak crests around the base of the petiole, margin undulate or fimbriate; blade to 50 cm long, base cordate, palmately 5—9-lobed, lobes extending to near the base, elliptic, coarsely serrate, often irregularly lobulate, apex acute, sinuses rounded. Inflorescence a terminal compound umbel; peduncle to c. 4 cm, bearing lanceolate bracts to 21/, cm long, caducous or persistent, occasionally with some bristles on the back, primary rays c. 20-30, c. 9 cm long, pubescent, bearing opposite caducous or rarely persistent bracts at the apex; central branch variable in length (2-18 mm), bearing an umbellule or head 1-2!/, cm @ of sterile bacciform flowers (4-5 mm @), 4-8-celled, subtended by minute bracts, pedicels variable in length (5-18 mm); lateral branches 3—7 cm, articu- lated near the middle, terminating in a globose head of c. 20-30 sessile flowers (or pedicels 11/, mm long), bracts between the flowers very small. Calyx rim obsolete. Corolla few- to many-lobed, tubular below, 1'/.4 mm long in bud. Stamens 6-17. Ovary 5-18-celled. Fruit a globose head of drupes; drupes c. 10 by 6 mm, obovoid, ribbed when dry. Distr. Malesia: throughout New Guinea. Ecol. Rain-forest, especially along the muddy banks of rivers, from sea-level to 100 m. Vern. Akriek, Biak, korinki, Orne, kwita-kwita, Milne Bay, sapi-ai, Jense, terrasahui, Manikiong. Notes. Unbranched or sparsely branched with crowns of large leaves. The bark is light brown and the wood cream. The inflorescence branches are purple, the flowers red, and the succulent ripe fruits black. The variation in the numbers of floral parts is considerable. Most specimens have more than 10 stamens and the same number of cells in the ovary, or more. Three gatherings have from 5—7 stamens and cells. These may possibly require to be segregated as a distinct species, but other evidence to support this course is lacking. An even more dis- tinctive gathering has central branches to 3 cm long with the pseudo-fruits on short pedicels (less than 5 mm) and flowers with 17 stamens and 25 cells in the ovary. These characters have not been included in the specific description as this specimen is only very tentatively referred to this species. This species forms an eastward extension of a complex of species, represented in the Moluccas by 37. O. talaudense, 17. O. soelaense and 18. O. glo- bulare.‘Several similar species occur in the Philip- pines. They are characterized by dense spherical heads of flowers. Osmoxylon talaudense resembles some specimens of O. sessiliflorum rather closely, and the difficulty of preserving the characters of these large-leaved plants in an herbarium probably obscures several good diagnostic features. The most reliable charac- ter to distinguish these two species is the position of the articulation on the lateral branches of the inflorescence rays. In specimens from the Talaud Is. this is close to the base (below the apex of the central umbellule of pseudo-fruits) and the bracts are persistent, whereas in the New Guinea material it is near (or above) the middle, and is usually marked by two inconspicuous scars which fre- quently are not opposite. In both species the pseudo-fruits are pedicelled, whereas in the other 48 FLORA MALESIANA [ser. I, vols9" two Moluccan species the pseudo-fruits are sessile or subsessile forming spherical heads. In O. globu- lare (from Morotai and Halmaheira) the lateral branches are rigid and only slightly flattened with the articulation near the middle and the bracts caducous. In O. soelaense the articulation is much nearer the base, the bracts are persistent, and the branches are broader and much flattened. 32. Osmoxylon camiguinense (MERR.) PHILIPSON, Blumea 23 (1976) 113. — Boerlagiodendron camiguinense Merk. Philip. J. Sc. 3 (1908) Bot. 252; En. Philip. 3 (1923) 222. Shrub to 2 m, glabrous except for the inflores- cence. Leaves clustered at the ends of the branches; petiole to 18 cm, with a sheathing base prolonged as a stipular ligule 1 cm long, and with several entire crests encircling the lower part of the petiole; blade 20 by 22 cm, base truncate, 3—5-lobed to about the middle, lobes oblong, scarcely nar- rowed towards the base, apex acuminate, sinuses broad, rounded, margin dentate, coriaceous. Inflorescence a terminal compound umbel; primary rays c. 2-2'/, cm, pubescent with opposite lanceo- late bracts (c. 12 mm long) at the apex; central branch c. 4 mm long, bearing a globose umbellule (c. 12 mm @) of sterile bacciform flowers (c. 2 mm @), pedicels c. 3 mm, subtended by numerous ligulate bracts; lateral branches c. 3 cm long, with opposite lanceolate bracts 4 mm long, ending in a head of c. 20-30 sessile flowers. Corolla and stamens not seen, described by MERRILL as 3-merous. Fruit globose 6 mm @ (dry), 3-seeded. Distr. Malesia: Philippines (Babuyan Is.: Camiguin I.). Ecol. On slopes in forest, at 500 m. 33. Osmoxylon fenicis (MERR.) PHILIPSON, Blumea 23 (1976) 114. — Boerlagiodendron fenicis MERR. Philip. J. Sc. 13 (1918) Bot. 44; En. Philip. 3 (1923) 223. — Boerlagiodendron tayabense MERR. Philip. J. Sc. 13 (1918) Bot. 45; En. Philip. 3 (1923) 224. Erect shrub or treelet a few m high, glabrous except for the inflorescence. Petiole to 45 cm, clasping base prolonged as a broad stipular ligule, 11/,-3 cm long, several prominent long pectinate petiolar crests extending up the petiole as oblique groups of bristles; blade to 30 cm long, base cordate or truncate, deeply palmately 3-7-lobed, lobes extending to within c. 7 cm of the base, broadly elliptic to oblong, entire or with subsidiary lobes, somewhat narrowed towards the sinuses, apex abruptly apiculate, margin remotely denticu- late or serrate, sinuses very broadly rounded. Inflorescence a terminal, compound, furfuraceous umbel, hemispherical, c. 10-12 cm @; peduncle 2-3 cm, densely enclosed in broadly ovate bracts c. 2 cm long, bearing dense fascicles of strong bristles on their blades; primary rays 15-30, c. 2'/,-3 cm long, 2!/, mm wide, pubescent, sub- tended by large bristly bracts c. 11/, cm long, bearing opposite terminal usually bristly bracts 6-15 mm long, each ending in 3 branches; central branch c. 4-10 mm, pubescent, bearing a terminal umbellule of c. 10-20 sterile bacciform flowers 2-3 mm @, 2-celled, pedicels 3-4 mm tomentose sometimes with a ruff of hairs around the pseudo- fruits, subtended by early-caducous small fimbriate bracts; lateral branches 21/,-3 cm long, with two opposite bracts (2-3 mm) about the middle, bearing a terminal head (8 mm @ without corollas) of c. 15-30 sessile flowers, subtended by incon- spicuous ovate tomentose bracts. Calyx an obsolete rim. Corolla in bud c. 1 mm long. Stamens 3. Ovary 3-celled. Fruiting head 10-12 mm @; drupes c. 6 mm long, crowded, sessile, 3-angled; seeds 3. Distr. Malesia: Philippines (Luzon). Ecol. In primary dipterocarp forest, on rocky slopes near streams, 300-400 m. Note. Closely allied to 40. O. luzoniense and 27. O. pectinatum, but the combination of long- fringed petiolar crests, pubescent inflorescence branches, excessively bristly bracts and 3-merous flowers is distinctive. 34. Osmoxylon eminens (BULL) PHILIPSON, Blumea 23 (1976) 114. — Trevesia eminens BULL, Cat. New Plants (1884) 17; Retail List (1885) 64, fig. — Boerlagiodendron mindanaense MerRR. Philip. J. Sc. 3 (1908) Bot. 154. — Boerlagiodendron eminens (BULL) Me_erR. En. Philip. 3 (1923) 223. Small tree, up to 10 m, with few stout branches. Leaves large, forming terminal crowns, glabrous when mature, young inflorescence densely fur- furaceous; petiole to 1 m, stout (to 2 cm Q), flattened above, base clasping the stem, heavily lenticellate, prolonged as a stipular ligule 2 cm long, bicuspid, often with scales or bristles on the back, and bearing few to several entire, fimbriate or occasionally long setose crests; blade to 60 cm long, palmately 10-19-lobed, base cordate, lobes reaching to near the base, lanceolate to oblong, up to 15 cm wide, in outline either strap-shaped or irregularly pinnatisect, or the central lobe occa- sionally distinctly tripartite, margin coarsely and irregularly dentate, apex acuminate. Inflorescence a terminal compound umbel c. 40 cm @; peduncle stout c. 6 cm, 11/, cm wide, bearing many lanceo- late scaly bracts 3-6 cm long; primary rays numerous, rigid, flattened, 9-12 cm long, 6-10 mm broad, bearing opposite oblong scaly bracts (2-3 cm long) at the apex, each ending in three branches; central branch '/,-1 cm long, terminating in an umbellule (3-5 cm @) of c. 20-40 sterile bacciform flowers, 7 mm @, 2-3-celled, pedicels 1-2 cm long, surrounded by an involucre of small bracts (to 8 mm long); lateral branches c. 12 cm long, with opposite bracts (6-10 mm long) near the middle, terminating in a head of c. 50-60 sessile flowers, 1!/,-2 cm @ (with corollas), heads spherical at anthesis, ovoid after corollas absciss; 1979) ARALIACEAE—I (Philipson) 49 bracts between the flowers very small. Calyx rim obsolete. Corolla 5-6-lobed, tubular below, 4-5 mm long. Stamens 4-6, filaments 7 mm long, anthers 1!/, mm long. Ovary 2-3 mm long (at anthesis), 5—6-celled. Fruits crowded in dense ovoid heads 3-4 by 2!/.-3 cm, drupes c. 9 by 5 mm, 5—6-angled by mutual pressure, narrowed to the base, crowned by the persistent stigmatic boss; pyrenes crustaceous; endosperm wrinkled. Distr. Micronesia: Carolines; in Malesia: throughout the Philippines. Ecol. In primary forest from low altitude (100 m) to ridge forest and mossy forest at 950 m, often in shady ravines. Vern. Cf. MERRILL: apalong or apulong, Bis., bunglui-babde, pina-pina, Sul., mangunpulun, Bag., palad-ulot, S.L.Bis., ulo-ulo, C.Bis.; in addition: lolobongan, Lan. Notes. This is the most widespread and most frequently collected species in the Philippines. It is also the most striking. Its large, many-lobed, fan- shaped leaves and the strong inflorescences, with globular flower-heads and large clusters of pseudo- fruits are distinctive. Only 16. O. pulcherrimum resembles it somewhat in its leaf characters, but the central branches of the inflorescence rays of that species are much longer and its pseudo-fruits are sessile. The inflorescence branches are described as dull reddish brown, the flowers as light orange, and the fruits as indigo-black. 35. Osmoxylon serratifolium (ELMER) PHILIPSON, Blumea 23 (1976) 114. — Boerlagiodendron serratifolium ELMER, Leafl. Philip. Bot. 2 (1908) 505; MErRR. En. Philip. 3 (1923) 224. Sparingly branched shrub to 5 m. Petiole to 50 cm long, channelled above, clasping base pro- longed as an obtuse stipular ligule, and with few narrow + fimbriate crests around the base of the petiole; blade to 50 cm long, base cordate, pal- mately lobed (up to 11 lobes), lobes extending to within about '/,; from the base, narrowly elliptic, margin serrate (or slightly lobulate), apex acumi- nate, sinuses narrowly rounded. Inflorescence a terminal compound umbel c. 30 cm @; peduncle stout, bracteate; primary rays 20-30, 4-5 cm long, 5-6 mm wide, flattened, subtended by lanceolate bracts 3-5 cm long, sometimes with bristles on the back, and bearing similar opposite terminal bracts 2'/, cm long, each terminating in three branches; central branch 12-15 mm long, terminating in an umbellule (3-4 cm @) of c. 20-25 sterile bacciform flowers 5-6 mm @, 3-4-celled, pedicels to 10 mm, interspersed with persistent small bracts; lateral branches c. 9 cm long, with opposite bracts (c. 6 mm long) 2-3 cm from the base, terminating in a spherical head of c. 30 sessile flowers c. 11/, cm @ (in bud), bracts between the flowers very small, obtuse, fimbriate. Calyx rim obsolete. Corolla 5-7-lobed above, tubular below, 5 mm long. Stamens 5-6, exserted, filaments 7 mm long, anthers 2 mm long. Ovary 3 mm long, 5-celled. Fruit unknown. Distr. Malesia: Philippines (Leyte, Camiguin, Panay). Ecol. ELMER noted that this species was rare in the low hills of Leyte. Notes. The flowers are orange-yellow (salmon), the fruits dark purple. The inflorescence is very similar to that of 34. O. eminens, but the leaves lack the many strong fan-like ribs of that species. 36. Osmoxylon celebicum PuHiLiPson, Blumea 23 (1976) 115. — Boerlagiodendron celebicum HARMS ex Koorpb. Minah. (1898) 489, nomen. A small, sparsely branched tree, 6 m high. Large leaves forming terminal crowns; petiole 50 cm by 8 mm, flattened above, with a sheathing base prolonged as a strong stipular ligule 3 cm long with branched fibrous setae on the outer surface, and with several crests bearing similar setae on the lower part of the petiole; blade 50 cm @, base emarginate, deeply 9-lobed, lobes narrowly elliptic to lanceolate, narrowed towards the sinuses, apex broadly cuneate, margin minutely and remotely serrate, sinuses broadly rounded. Inflorescence a terminal compound subspherical umbel c. 20cm @; peduncle short, stout (15 mm wide) with large setose bracts (c. 4 cm long) below and among the primary rays; primary rays c. 15, c. 6 cm long, 5 mm broad, with a pair of large setose bracts (22 by 10 mm) at the apex, each ending in three branches; central branch c. 6 by 2 mm, terminating in an involucre of setose ovate bracts (c. 4 mm long) and an umbel of c. 20-30 sterile bacciform flowers (c. 3 mm @ when dry, 2-3-celled, on pedicels 6-10 mm long) interspersed with bracts covered with crisp reddish-brown setulae; the two lateral branches c. 2 cm long, with opposite setulose bracts (c. 1 cm long) below the middle, terminating in a spherical head of c. 30-40 sessile flowers each sub- tended by a reddish brown setulose cymbiform bract. Calyx rim obsolete. Corolla c. 2'/, mm long in bud (not seen in open condition). Stamens 5. Ovary subcylindric, c. 1 mm long in bud, 5-celled; disk with a central stigmatic boss. Fruit unknown. Distr. Malesia: Celebes (Minahasa, Manado). Ecol. On rich volcanic sand, at 10 m. Vern. Sinomaha. Note. The flower buds are orange and the fruits deep purple. 37. Osmoxylon talaudense PHILIPSON, Blumea 23 (1976) 115. Shrub or small tree, to 6 m, glabrous. Leaves at the ends of the stout branches; petiole to 60 cm, broadly channelled above, clasping base prolonged as a Stipular ligule c. 2 cm long, and bearing 2-3 fimbriate crests; blade c. 50 cm long truncate to cordate at the base, palmately 7-11-lobed to within 50 FLORA MALESIANA [ser. I, vol. 91 Fig. 16. Osmoxylon micranthum (HARMS) PHILIPSON. a. Habit, x 4/2, b. flower, c. false fruit and ditto in CS, x 15, d. CS of fruit, x 6 (a—-c Kanis 1384, d PULLEN 428). Drawn by W. R. PHILIPSON. 1979] ARALIACEAE—I (Philipson) 51 + 4/, of the base, lobes elliptic oblong slightly narrowed towards the rounded sinuses, apiculate, margin denticulate to undulate. Inflorescence a terminal compound umbel, peduncle 1-2 cm, bearing broad ovate bracts 2-3 cm long, with bristles on the back, (similar persistent bracts subtend the primary rays); primary rays c. 15, c. 34 cm long, 4 mm wide, flattened, bearing opposite terminal persistent bracts 11/,-2 cm long, sometimes with a few bristles on the back, ending in three branches; central branch 8-10 mm long, terminating in an umbellule 2 cm @ of 15-20 sterile bacciform flowers (4 mm @, 2-celled) surrounded by an involucre of obtuse bracts 1-2 mm long, pedicels 5—7 mm; lateral branches 4'/,-5 cm long, bearing opposite ovate persistent bracts (3-7 mm long) c. 5-10 mm above the base, terminating in a dense head c. 1 cm @ of 30-40 sessile flowers inter- spersed with inconspicuous obtuse bracts. Calyx rim obsolete. Corolla 5-lobed. Stamens 5. Ovary turbinate, 11/, mm long, 5-celled. Fruit in spherical heads 2 cm @ (when dry); drupes c. 9 by 6 mm, obovoid, 5-ribbed. Distr. Malesia: N. Moluccas (Talaud Is.: Karekelong and Salebabu). Ecol. Common in forest, besides streams, from near sea-level to 100 m. Vern. Laripatu, Talaud. Note. The flower is yellow-orange and the fruit dark purple. For a discussion of the distinctive features, see under 31. O. sessiliflorum. 38. Osmoxylon micranthum (HARMS) PHILIPSON, Blumea 23 (1976) 115. — Boerlagiodendron micranthum Harms, Bot. Jahrb. 56 (1920) 379. — Boerlagiodendron sayeri HarRMs, I.c. 379, f. 1 a-j. — Eschweileria gawadensis BAKER f. J. Bot. 61 (1923) 22. — Boerlagiodendron tricolor PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 11. — Fig. 16. A sparsely branched shrub to 8 m, sometimes trailing or semi-scandent, young parts uniformly setulose, buds without cataphylls. Leaves in terminal clusters; petiole up to 30 cm, rather narrow (2-4 mm @), becoming sparsely setulose, channelled above, with a sheathing base prolonged as a membranous stipular ligule up to 3 cm long, and with a number of lacerate crests encircling the lower part of the petiole; blade deeply 3—5- or more rarely 7-lobed, or below the inflorescence sometimes simple, base cordate or emarginate, the central lobe up to 30 cm long, the lobes oblong, lanceolate or broadly elliptic, entire or irregularly lobed or incised, or with small sub-lobes, apices long cuspidate, acute, margin serrate, sinuses between the lobes broad and rounded, surfaces become sparsely setulose to subglabrous. /nflores- cence a terminal compound umbel, often appearing subterminal by growth of a leafy brafich at the base of the peduncle; peduncle short (1-2 cm), heavily setulose, occasionally with a flowering ray arising from the axils of bracts on or below the peduncle, bearing distally many lanceolate bracts 5-10 mm long; primary rays 12-18, 10-20 mm long, setulose, with two lanceolate bracts at the apex, each ray ending in three branches; the central branch very short (2-3 mm) bearing a subglobose umbel of many (c. 40) small sterile bacciform flowers (c. 1/, by 1 mm) with filamentous pedicels c. 2 mm long, and 1-2 cells each with 1 abortive ovule; the two lateral branches c. 2 cm long, with two minute bracts about their middle, setulose, terminating in a head of c. 20 sessile flowers sur- rounded by an involucre of small rounded bracts. Calyx rim obsolete. Corolla 4(—5)-lobed, united below, c. 2 mm long. Stamens 4(—5), filaments ribbon-like elongating beyond the corolla tube at anthesis, 3-4 mm, anthers small. Ovary shortly subcylindric, c. 1 mm high, faintly angled, glabrous, 1-5-, usually 4-celled, disk fleshy, rising slightly to the central stigmas. Fruit an ellipsoid drupe with 1-5 cartilaginous pyrenes; seeds with smooth endosperm. Distr. Malesia: New Guinea (Irian Jaya: Idenburg R. to Milne Bay Distr.). Ecol. In primary forest from the foothills to the montane mossy forest, often in swampy or deeply shaded situations, 700-2400 m. Vern. Diande, Chimbu, kenata, Okapa. Note. The inflorescence branches are often red or purple, and the flowers either orange or reddish with yellow anthers. The ripe fruits are deep purple or black. The shape and size of the leaf can vary greatly, even on the same plant. The ovary usually has 4 cells, but plants with 3, 2 and 1 occur. Since these are alike in other respects they have been treated as a single species. Although the type of Boerlagiodendron tricolor has an ovary with 5 cells and is from much further west than other gather- ings, it is not considered to be specifically distinct. 39. Osmoxylon trilobatum (MERR.) PHILIPSON, Blumea 23 (1976) 116. — O. cumingii SEEM. J. Bot. 6 (1868) 141, nomen. — Boerlagiodendron trilo- batum Merk. Philip. J. Sc. 2 (1907) Bot. 289; En. Philip. 3 (1923) 224. Slender shrub or small tree up to 5 m, becoming glabrous except for slight pubescence on the inflorescence. Leaves clustered near the ends of the branches; petioles to 25 cm, c. 3 mm wide, clasping base prolonged as a broad stipular ligule c. 1 cm long, and with 2-3 entire, or obscurely fimbriate, often recurved crests surrounding the base of the petiole; blade 3- or occasionally 5-lobed (leaves below the inflorescence sometimes simple), to 30 by 28 cm, base broadly cuneate, rounded or trun- cate (emarginate in 5-lobed leaves), lobes about 1/,7/, of the blade, narrowly or broadly oblong, often slightly narrowed below and sharply acumi- nate to caudate, margin serrate. Inflorescence a terminal compound spherical umbel, 7-15 cm 2, either rather compact or branches lax; peduncle 2-3 cm with broad ovate bracts; primary rays 8-20 52 FLORA MALESIANA [ser. I, vol. 9 or more, slightly pubescent, 11/,-4 cm long, sub- tended by ovate bracts 5-10 mm long, opposite ovate bracts at the apex, 1-3 mm long; central branch 1!/,-6 mm long, pubescent, bearing an umbel (1-1!/, cm @) of sterile bacciform flowers up to 5 mm @, 1-4-celled, pedicels 2-6 mm long, subtended by ovate bracts 1-3 mm long; lateral branches 1!/,-3 cm with opposite small bracts about the middle, bearing a terminal head, c. 1 cm @ of c. 8-20 flowers, surrounded by an involucre of small rounded pubescent bracts, pedicels c. 1 mm or less (up to 3 mm in fruit). Calyx rim obsolete. Corolla 4-5-lobed above, tubular below, 2-3 mm long. Stamens 4-5, exserted. Ovary subcylindric, 4-5-celled. Fruit a spherical drupe (when dry 4-5- ribbed, 7 by 5 mm). Distr. Malesia: widespread in the Philippines (Luzon to Mindanao). Ecol. In primary forest, frequently beside streams in damp ravines, from the lowland at 75 m to 750 m. Vern. Kamay-kamay, Tag., ayum, C.Bis. Note. Sparingly branched but wide-spreading slender shrub, with yellowish bark, at first heavily dotted with brown lenticels. The flowers are white and the ripe fruit smooth and purple. 40. Osmoxylon luzoniense (MERR.) PHILIPSON, Blumea 23 (1976) 116. — Boerlagiodendron luzoniense MERR. Philip. J. Sc. 3 (1908) Bot. 252; En. Philip. 3 (1923) 223. — Boerlagiodendron clementis MERR. Philip. J. Sc. 3 (1908) Bot. 155; En. Philip. 3 (1923) 222. — Boerlagiodendron agusanense ELMER, Leafl. Philip. Bot. 7 (1914) 2330; MerrR. En. Philip. 3 (1923) 222. — Boer- lagiodendron diversifolium MERR. Philip. J. Sc. 10 (1915) Bot. 333; En. Philip. 3 (1923) 223. Erect, unbranched or sparsely branched shrub or tree to 8 m, becoming glabrous except for the inflorescence. Leaves clustered at the ends of the branches; petiole to 40 cm, with a clasping base prolonged as a short stipular ligule, and with several basal entire or shortly fimbriate crests; blade to 33 cm @, base truncate or cordate, deeply palmately 3—7-lobed, sinuses broad, rounded, lobes elliptic, usually narrowed below, entire or with subsidiary lobes (the central lobe especially often narrow below and strongly pinnately lobed), apex acute, margin coarsely serrate, coriaceous; upper- most leaves often reduced and simple. Inflorescence a terminal compound, subsessile umbel 10-15 cm @; primary rays c. 20-30, 2-3 cm long, subtended by lanceolate furfuraceous and + fimbriate bracts, furfuraceous villose or + hirsute, with opposite rounded or lanceolate hirsute bracts at the apex, each ending in three branches; central branch c. 2-8 mm long, hirsute, ending in an involucre of minute bracts (1 mm) surrounding a globose (2 cm @) umbellule of c. 15-20 sterile flowers (c. 6 by 6 mm, 2-3-celled), pedicels 2-3 mm, hirsute; lateral branches c. 2'/,-31/, cm long at anthesis, with an articulation about the middle, + hirsute, ending in a globose head, 2 cm @ (with open corollas), with ovate tomentose obtuse bracts, c. 2 mm long; flowers 30-40, + sessile (rarely pedicels to 2 mm). Calyx rim obsolete. Corolla 4-5-lobed above, tubular below, 3'/.4 mm long. Stamens 4-5, exserted, filament stout, 5 mm, anther 1 mm long. Ovary 4-5-celled. Fruit 6 by 5 mm (dry), strongly 4~5-ribbed. Distr. Malesia: widespread in the Philippines (Luzon to Mindanao), also in N. Celebes. Ecol. In forests, often by streams and on ridge in mossy forest, 280-1650 m. Vern. Philippines: bolwang hi inalahan, If., iyangnok, Mbo, malakapdayas, 8.L.Bis., molonpolon, Buk., tafigan-tafigan-batu, Buk., tachung, vafigang, Ig. Notes. Inflorescence yellow to red (salmon), fruits blue-black or purple. MERRILL did not liken his Boerlagiodendron diversifolium (from Mindanao) to this species, no doubt because he gave importance to the occur- rence of variable leaves and the 5-merous flowers. However, specimens from Luzon may possess simple leaves below the umbel, and both 4- and 5-merous flowers occur in both Luzon and Mindanao. The greater range of material now available establishes the identity of the two species. Similarly, no features seem to distinguish Boer- lagiodendron clementis, and B. agusanense though a greater range of collections would be desirable. The species is treated here in a broad sense. The inflorescence characters of most specimens are uniform, being hirsute and with the bracts at the apex of the primary rays obtuse and short. Some specimens (CURRAN 5088, ELMER 16762) have longer lanceolate bracts with some bristles on the back. The northernmost specimen, from Ilocos Norte, has finer and less hairy inflorescence rays (recalling 39. O. trilobatum), but the foliage agrees with this species. Leaf-shape is more variable, even on the same specimen. Other species which resemble O. /uzoniense in some respects are: 32. O. camiguinense with broader, shallower lobing, a more delicate. inflorescence, and _ tri-merous flowers; 27. O. pectinatum with glabrous inflores- cence branches and long-pectinate petiolar crests; and 26. O. humile with pedicelled flowers forming less dense heads. A specimen from Surigao Province (BS 83562) has a most interesting abnormal structure. The central branches of the inflorescence rays bear heads of apparently fertile flowers, with lobed corollas and exserted stamens. 41. Osmoxylon insigne (Miq.) Becc. Malesia 1 (1878) 195; PHiLreson, Blumea 23 (1976) 117. — Trevesia insignis M1q. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 222. — Trevesia palmata var. insignis CLARKE, FI. Br. Ind. 2 (1879) 732, pro nomen. — Eschweileria insignis (M1Q.) BOERL. Ann. Jard. Bot. 1979] ARALIACEAE—I (Philipson) a3 Btzg 6 (1887) 122. — Boerlagiodendron insigne (Miq.) Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 31. A glabrous tree. Leaves large, palmately lobed; petiole to 45 cm, 8-10 mm wide, flattened above, with a sheathing base prolonged as a stipular ligule, numerous fimbriate crests around the base of the petiole, and irregular tufts of bristles along the whole length of the petiole; blade to 55 cm @, deeply 5—7-lobed, the sinuses broadly rounded and c. 4-5 cm from the base of the blade, lobes pinnatilobed, with a narrow base and an attenuated apex, margins serrate. Inflorescence a terminal compound umbel c. 16 cm @; peduncle stout, with lanceolate bracts (1!/,-2 cm long) subtending the primary rays; primary rays 25-30, c. 4 cm long, bearing 2 caducous lanceolate bracts (c. 1 cm long) at the apex, each ray ending in 3 branches; central branch c. 12-16 mm long, bearing an umbel (c. 2 cm @) of c. 20 sterile ovoid bacciform flowers (c. 4 mm long when dry) with pedicels c. 4-6 mm long, and 3-celled; two lateral branches c. 31/, cm long, with an articulation about the middle, termi- nating in an umbellule (c. 2 cm @) of c. 10-15 flowers on short stout pedicels c. 2 mm long, umbellules surrounded by a receptacular rim after caducous bracts have abscissed. Ca/yx rim minute. Petals c. 3!/, mm long in bud (when dry), with 8-9 lobes above, tubular below. Stamens 8-9, with stout filaments. Ovary cylindric, 8—9-celled; disk with a central double row of pustulate stigmas. Fruit unknown. Distr. Malesia: Moluccas (Batjan). Note. Tufts of bristles along the entire length of the petiole together with the pinnatifid lobes of the leaf are distinctive. SEEMANN (J. Bot. 4, 1866, 353) referred to 5-flowered umbels with 5-angled drupes, but this probably relates to the New Guinea specimen which he included under this name. Insufficiently known Boerlagiodendron ledermannii HArMs, Bot. Jahrb. 56 (1920) 383; PHILIPSON, Blumea 23 (1976) 117. — Type: LEDERMANN 12293. HarMs compared this species with Boerlagio- dendron geelvinkianum. The size of the foliage and flowers prevents it from being included within that species. If it represents a local species, it has not been re-collected since the original gathering of LEDERMANN in 1912. The type specimen, which was incomplete, was destroyed during the war. Boerlagiodendron monticola HARMs in K. Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 330; PxHiLip- SON, Blumea 23 (1976) 117. — Type: SCHLECHTER 14471. The incomplete type specimen, gathered by SCHLECHTER, was destroyed during the war. This species was evidently similar to 38. Osmoxylon micranthum, but the ovary was possibly 10-celled. I have tentatively identified Ropspins 1644 as this species: it is close to O. micranthum but its ovary, with 8 cells, is outside the range of variation of that species and the pedicels are longer (in fruit). Its distribution (foothills of Adalbert Range) is not dissimilar to that of Boerlagiodendron monticola (Torricelli Mts). 8. ARTHROPHYLLUM BL. Bijdr. (1826) 878; DC. Prod. 4 (1830) 266; Mig. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27; BTH. in B. & H. Gen. Pl. 1 (1865) 944; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 54; Koorp. Atlas 4 (1916) f. 675 & 676; Hutcu. Gen. FI. Pl. 2 (1967) 80; STONE, Gard. Bull. Sing. 30 (1977) 276; PHiLipson, /.c. 299, f. 1-16; Adansonia 17 (1978) 329. — Mormoraphis JACK ex WALL. Cat. (1831) n. 4931, nomen. — Eremopanax BAILL. Adansonia 12 (1878) 158. — Fig. 17, 19-23. Unarmed, sparingly branched trees or shrubs. Leaves on vegetative shoots and lower leaves spirally arranged, imparipinnate, often crowded at the end of the branches, those on flowering branches often opposite, smaller, or reduced to a single leaflet; petiole terete; rachis articulated at the insertion of the pinnae and leaflets; leaflets entire; stipular sheath clasping, small, ligule a mere rim. /nflores- cence consisting of compound umbels, either solitary and terminal or more commonly borne on a cluster of specialized leafy branches arising from the axils of the uppermost leaves; pedicels not articulated. Flowers bisexual. Calyx an undulate rim, sometimes with indistinct teeth, persistent. Petals 4-6, valvate in bud. Stamens 4-6, anthers curved, basifixed. Ovary turbinate, 1-celled; disk FLORA MALESIANA [ser. I, vol. 91 Fig. 17. Habit of Arthrophyllum diversifolium BL. (Photogr. HOOGLAND, Bogor). 1979] ARALIACEAE—I (Philipson) 55 fleshy, rising in the centre to the + sessile capitate stigma. Fruit ovoid or spheroidal, often oblique; exocarp leathery; endocarp cartilaginous. Seed solitary, pendulous; endosperm deeply transversely ruminate. Distr. About 31 spp. (17 in Malesia) extending from the Nicobar Is. and Indo-China to the Philippines, New Guinea and New Caledonia. Fig. 18. Notes. The principal distinctive characters of this isolated genus are the single-celled ovary and the arrangement of the umbels on specialized lateral branches in the majority of the species. The present treatment, following that which I gave in 1977, /.c., remains tentative until widespread field studies can be undertaken. KEY TO THE SPECIES 1. Inflorescence becoming paniculate by the successive development of branches below the umbellules. Fig. 19a 1. A. proliferum is heen ee hel em Cmte ese ae e olhijal Nels «© (© ©, (© \¢ Je .e8sd © © 2 « © .» .e 1 1. Inflorescence a compound umbel. 2. Inflorescence with four orders of branching. 3. Leaflets lanceolate. 4. Leaflets c. 4-7 cm long. Fig. 20f 4. Leaflets much longer 3. Leaflets broader. 5. Pedicels (at early anthesis) c. 10 mm long. 6. Peduncles of umbellules with bracts or their scars. Fig. 21a 6. Peduncles of umbellules without bracts or their scars. Fig.21b. . ... . 2. A. ashtonii 3. A. angustifolium 4. A. ahernianum 5. A. engganoense 5. Pedicels (at early anthesis) c. 5 mm long, or shorter. 7. Leaves associated with the umbels rotund. Fig.2le. .........2... 6. A. collinum 7. Leaves (or leaflets) associated with the umbels ovate or elliptic. 8. Leaves associated with the umbels ovate, + fleshy, with the lower surface smooth (lateral veins obscure). Fig. 21f cr Sra ye ee CY a CeCe te ee SP Cae) oC ee et Seem en er 7. A. crassum 8. Leaves (or leaflets) associated with the umbels + elliptic, coriaceous or chartaceous, veins visible. 9. Young parts glabrous 9. Young parts with rufous tomentum. 10. Umbels at anthesis with numerous filamentous pedicels. Fig. 21c. 10. Umbels at anthesis with fewer stout pedicels. Fig. 21d So ees ae ee Pe eee eee cee 8. A. pacificum 9. A. diversifolium 10. A. macranthum 2. Inflorescence with three (or fewer) degrees of branching. 11. Leaflets membranaceous or chartaceous. 12. Mid-leaflets c.16cmlong. ....... 12. Mid-leafiets c. 8 cm long or shorter. 11. A. papyraceum 13. Leaflets usually 5-7 (Fig. 20a). Primary inflorescence branches usually short (c. 3-6 cm) and without articulations 12. A. maingayi 13. Leaflets more numerous (Fig. 20d). Primary inflorescence branches longer (10-20 cm), with one or more leafy nodes 11. Leaflets coriaceous. 14. Leaflets c. 5 14. Leaflets more numerous. 15. Petals and stamens 6 15. Petals and stamens 4 or 5. 16. Leaflets 6 cm long, or longer. Fig. 20b . 16. Leaflets shorter. Fig. 20c 1. Arthrophyllum proliferum PuHrLipson, Gard. Bull. Sing. 30 (1977) 302, f. 3-4. — Fig. 19a-b. Medium-sized, glabrous tree. Leaves multijugate, up to 90 by 24 cm, of the flowering branches smaller with fewer pinnae or usually simple; petioles up to 28 cm, 5 mm @; petiolules 1-1!/, cm; leaflets obovate-oblong, c. 16 by 7 cm, chartaceous, margin slightly revolute, base broadly cuneate to truncate, Lee OP Cer ae ee eee ge Cie Ow ee Ly ee et POPE pm eet =) Cua a te Che ys ite Ae eee OM re oC) or Cemecee bel "wt eee: Lay Wel e ter. ie sep re! =e S26!) ale ee ee ae 13. A. kjellbergii Be get ck a RAE ges My teen BIR. pel 14. A. cenabrei Suemesy (sh echt ees va. Xo Von ee Ga eis 15. A. pulgarense 16. A. montanum 17. A. alternifolium sometimes oblique, apex acute. Inflorescences terminating specialized plagiotropic shoots, bear- ing axillary flowering branches and ending in umbellules of a few flowers below which pairs or whorls of branches continue the growth of the inflorescence to produce an elongated panicle of umbellules; pedicels 7-8 mm (slightly longer in fruit). Calyx often with 5 indistinct teeth. Petals 5, 56 FLORA MALESIANA [ser. I, vol. 91 Le — + Fig. 18. Species density of Arthrophyllum BL. in Malesia; above the hyphen the number of endemic species, below it the non-endemics. Complete range of genus encircled; 17 of the total of 31 spp. occur in Malesia. 3'/, mm long, narrowly triangular. Stamens 5, filaments c. 2 mm, anthers reniform, c. !/, mm long. Ovary obconical, c. 2 mm long; disk fleshy, cushion-like; stigma capitate + sessile. Fruit ovoid, fleshy, capped by the calyx and the enlarged beak-like stylopodium, c. 10 by 5 mm when dry. Distr. Malesia: E. New Guinea (Morobe Distr.). Ecol. Mid-mountain rain-forest, reaching the canopy, on steep slopes, 300-1200 m. Notes. The thick outer bark is grey-brown, fissured, and peeling in small flakes. Wood straw- coloured. Cut stems exude brown latex. The thick petals are yellow-green. The flower and fruit are typical of this well- defined genus, but the branching of the inflores- cence is unlike that found in all other species. 2. Arthrophyllum ashtonii PHILIPSON, Gard. Bull. Sing. 30 (1977) 303, f. 12. — Fig. 20f. Slender small tree, to 5 m, with the leaves dis- persed for some distance from the apex of the branches, young parts with brown scurfy tomen- tum which persists on the umbellules. Leaves multijugate, to c. 30 cm long; of the flower-bearing branches smaller with fewer pinnae, or simple; petioles slender, c. 7-9 cm; petiolules c. 4-7 mm; leaflets lanceolate to broadly lanceolate, 3—71/, by 1-2 cm, thinly coriaceous, margin revolute, base broadly cuneate, apex tapered to subcaudate, veins channelled above, visible beneath. Jnflorescence a terminal cluster of specialized leafy branches; main rays variable in length in the same inflorescence, the longest from c. 16-25 cm long with a pair of opposite simple leaves about the middle (with flowering branches in their axils) and ending in a whorl of secondary rays subtended by simple leaves; secondary rays 4-12 cm long, bearing simple leaves, usually in an opposite pair and terminating in compound umbels; umbellules with c. 8-10 flowers; pedicels 4-10 mm, furfuraceous. Flower buds c. 2 mm long, calyx a furfuraceous rim. Petals 5, broadly triangular. Stamens 5, anthers curved, basifixed. Ovary glabrous, obconical, c. 1 mm long; stigma on a raised stylopodium at the centre of a flat disk. Fruit spheroidal, with a persistent stylopodium, c. 6 mm long when dry. Distr. Malesia: Borneo (Sarawak and Brunei). Ecol. Mossy forest on sandstone ridge, and in kerangas forest, 1000-1550 m. Note. The small narrow leaflets are very dis- tinctive. No other species with small leaflets has inflorescences which branch to the fourth degree. 3. Arthrophyllum angustifolium RIDL. J. Fed. Mal. St. Mus. 10 (1920) 136; Fl. Mal. Pen. 1 (1922) 885; PHILIPSON, Gard. Bull. Sing. 30 (1977) 304. Shrub or small tree, up to 5 m, young parts rufous-tomentose, glabrescent. Lower Jeaves multi- jugate, rachis dilated, c. 70-90 by 30-40 cm; petioles c. 15-22 cm, 3-5 mm wide, ligule a rim c. 2 mm long; petiolules c. 7-10 mm long; leaflets coriaceous, lanceolate, c. 15-22 by 11/,-2!/, cm, tapering to an acute or obtuse apex, base cuneate, margin slightly revolute; upper leaves reduced, mostly unifoliolate, opposite, broader, with petioles 2-4 cm long. Jnflorescence a terminal cluster of specialized leafy branches; main rays 30 cm (or more) long, bearing simple leaves in opposite pairs with small flowering branches in their axils, ending in a whorl of c. 10-12 secondary rays subtended by simple leaves; secondary rays 8-12 cm long, bearing pairs of simple leaves, and terminating in an umbel of 5-12 tertiary rays c. 2-3 cm long, with bract scars about the middle, and ending in an umbellule of c. 8-12 flowers, pedicels c. 5 mm. Petals 5, 2-3 mm long in bud. Stamens 4. Ovary turbinate, inconspicuous at anthesis; disk fleshy; stigma + sessile. Fruit spheroidal, c. 5 by 5 mm when dry, calyx and stylopodium small. Distr. Malesia; Malay Peninsula (Perak) and Borneo (Brunei). Ecol. Forest and old regenerated forest on peat swamp at low altitude or on ridges. Note. The lanceolate leaflets are unlike those of any other species. The grey bark is minutely fissured and bears many small orange lenticels. The wood is soft and white. The Malayan and Bornean specimens are similar, except that the flower buds are larger in the Brunei plant. 4. Arthrophyllum ahernianum Me_rr. Philip. J. Sc. 1 (1906) Suppl. 109; En. Philip. 3 (1923) 235; PuHILipson, Gard. Bull. Sing. 30 (1977) 304, f. 13. — A. pinnatum (non CLARKE) F.-VILL. Nov. App. (1880) 103; VIDAL, Sinopsis Atlas (1883) 28, t. 55 f. c. — Macropanax sp. VIDAL, Rev. Pl. Vasc. Filip. (1886) 145. — A. sablanense ELMER, Leafl. Philip. Bot. 1 (1908) 331. — A. borneense Merk. Pl. Elm. Born. (1929) 231, non BAKER, 1896. — A. elmeri Merr. Webbia 7 (1950) 319. — A. merrilliana Furtapo, Gard. Bull. Sing. 19 (1962) 185. — Fig. 21a. 1979] ARALIACEAE—I (Philipson) 57 Fig. 19. Arthrophyllum proliferum PHILipson. a. Part of inflorescence in fruiting state, x 1/2, b. ditto, showing detail of ultimate branches in flowering stage, nat. size. — A. maingayi PHILIPSON. c. Terminal inflorescence, x !/,.— A. montanum RwwL. d. Single flowering shoot, '/; (Courtesy Gard. Bull. Sing. 30, 1977). Tree up to 15 m, young parts with rufous tomen- tum. Leaves clustered at the ends of the branches, multijugate, up to 200 by 60 cm; of the flower- bearing branches smaller with fewer pinnae, or simple; petiole stout, up to 35 cm; petiolules 5-20 mm; leaflets ovate-oblong, occasionally oblong-lanceolate, up to 35 by 12 cm, membrana- ceous or chartaceous, margin revolute, base cuneate to rounded, usually oblique, apex short acuminate. Inflorescence a whorl of specialized leafy branches forming a terminal crown; main rays up to 150 cm (or more), bearing pinnate leaves usually in 1-2 opposite pairs, and with flowering branches in the upper axils, ending in a whorl of secondary rays subtended by pinnate or more rarely simple leaves; secondary rays up to 30 cm bearing simple or pinnate leaves in opposite pairs, and terminating in compound umbellules; um- bellules with c. 10-20 flowers c. 3 cm @; peduncles with 1-2 pairs of small simple, often caducous leaves; pedicels c. 1 cm (at anthesis) subtended by minute caducous bracts. Petals 5, 4 mm long. Stamens 5, anthers curved. Ovary turbinate; disk fleshy; stigma + sessile. Fruit c. 10 by 7 mm, ellipsoidal, calyx and stylopodium forming a prominent beak. 58 FLORA MALESIANA [ser. I, vol. 91 Fig. 20. Foliage leaves of Arthrophyllum spp. a. A. maingayi PHILIPSON, b. A. montanum RIDL., c. A. pulga- rense ELMER, d. A. kjellbergii PHILIPSON, e. A. alternifolium MAINGAY ex RIDL., f. A. ashtonii PHILIPSON. All = 1/3; (Courtesy Gard. Bull. Sing. 30, 1977). Distr. Malesia: N. Borneo and throughout the Philippines to the northern Moluccas (Talaud, Ternate). Ecol. Primary and second-growth forest, from the lowlands to 1000 m. Vern. Philippines, cf. MERRILL: alabihig, dokloi, P.Bis., binalayon, C.Bis., danipo, \g., higin, Mang., malapapaya, puyga-puygahan, Tag., pama- latafgen-a-purau, Wk. Additional names: Philip- pines: /ulpo, Luzon, mayari, Mindoro, bungyo, Palawan; Moluccas: /angator’a, Talaud. Notes. This species replaces the more westerly 9. A. diversifolium which it closely resembles. It is characteristically larger in all its parts, particularly in the size of the individual flowers and the length of their pedicels. There are fewer flowers in an umbellule. The distinction between these two species is not always easy to make, especially when the material is fragmentary; PHILIPSON, /.c. A few specimens from the Philippines appear very similar to A. diversifolium, and it is possible that this species extends beyond Borneo. I have regarded them as part of the range of variation of A. ahernianum. Similarly, at least one specimen from S. Borneo approaches A. ahernianum in appearance. 5. Arthrophyllum engganoense PHILIPSON, Gard. Bull. Sing. 30 (1977) 305, f. 14. — Fig. 21b. Tree to 21 m high, becoming glabrous. Lower leaves imparipinnate, multijugate, 60 cm long or more; petiole 24 cm, 6 mm wide, rachis articulated at the insertion of the leaflets; petiolules 10-18 mm; leaflets broadly elliptic to elliptic-oblong, c. 12-15 by 6-7 cm, base rounded with a short asymmetrical cuneate centre, apex shortly apiculate, margin entire, often undulate chartaceous. Flowering branches c. 40 cm; leaves opposite, simple, or unifoliolate, petiolules c. 5-7 cm, leaflets ovate, c. 13 by 6 cm, with inflorescence branches in their axils; ending in a whorl of simple leaves surround- ing a compound umbel to 30 cm @; secondary rays c. 8, c. 10-15 cm long at flowering, slender (2mm @) and striate, each bearing a pair of small leaves about the middle (sometimes with inflorescences in their axils) and ending in an umbel; tertiary rays c. 8, slender, c. 20-40 mm long, without bracts; tertiary rays pedicels c. 5-10 per umbellule, c. 10-15 mm at anthesis. Ca/yx an undulate rim. Petals 5, c. 2 mm long in bud. Stamens 5, anthers curved. Ovary turbinate, obscurely ribbed, c. 2'/, mm long at anthesis. Fruit ellipsoid, c. 10 by 7 mm, with a rather small persistent calyx and stylopodium. Distr. Malesia: S. Sumatra (Enggano I.), two collections. Ecol. Forest at low altitude, up to c. 100 m. Vern. Langkapu utan kaauh, Enggano. Note. The two known collections of this species are very similar and contrast with the widespread 9. A. diversifolium because of the few-flowered umbellules with long, spreading pedicels. 6. Arthrophyllum collinum PHILIPsON, Gard. Bull. Sing. 30 (1977) 30S, f. 17. — Fig. 21e. Sparingly branched shrub or small tree up to 12 m, all young parts with dense, rufous tomentum which persists on flowers and_ inflorescences. Leaves multijugate, up to 60 (or more) by 32 cm; of the flowering branches smaller with fewer pinnae or more frequently unifoliolate, leaflets more rotund and with longer petioles; petiole up to 20 cm; petiolules c. 1 cm; leaflets oblong, broadly elliptic or rotund, up to 16 by 7 cm, coriaceous, margin slightly revolute, base truncate to rounded, unequal, apex rounded, obtuse, or shortly and bluntly apiculate, midrib prominent, lateral veins usually clearly visible below, upper surface fre- quently rugose. /nflorescence a cluster of specialized leafy branches forming a terminal crown; main rays up to 60 cm long, bearing one or more, rarely two, opposite pairs of usually unifoliolate rotund leaves often with flowering branches in their axils, and ending in a whorl of secondary rays, subtended by usually unifoliolate leaves; secondary rays up to 17 cm, bearing a pair of unifoliolate leaves with flowering branches in their axils, and terminating in compound umbellules; tertiary rays c. 5-10, ARALIACEAE— I (Philipson) 59 Ib Fig. 21. Umbellules of some Arthrophyllum spp. a. A. ahernianum MErRR., 6. A. engganoense PHILIPSON, c. A. diversifolium BL., d. A. macranthum PHILIPSON. All nat. size. — Bracts of two Arthro- phyllum spp. e. A. collinum PHILIPSON, f. A. crassum PHILIPSON. Both x !/; (Courtesy Gard. Bull. Sing. 30, 1977). c. 3-6 cm long, articulated near the middle; pedicels c. 10, c. 2-3 mm long, rufous tomentose. Petals 5, broadly triangular, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, rufous-tomen- tose, c. 1'/, mm long at anthesis; disk fleshy, stigma sessile. Fruit ellipsoidal, c. 9 by 5 mm (when dry), the calyx and stylopodium prominent. Distr. Malesia: Borneo (Sarawak and Sabah). Ecol. Forest and scrub, sometimes growing as an epiphyte in the crowns of trees, 600-2700 m. Notes. The bark is grey and smooth, the wood pale and soft, and the cut stems exude a yellowish or orange latex. This species is characterized by the rotund, eoriaceous leaves on the flowering branches. 7. Arthrophyllum crassum PHILIPsON, Gard. Bull. Sing. 30 (1977) 305, f. 18. — Fig. 21f. Sparingly branched shrub or small tree up to 8 m, occasionally epiphytic, all young parts with dense, rufous tomentum which persists on the 60 FLORA MALESIANA [ser. I, vol. 9} flowers and inflorescence. Leaves multijugate, up to 100 (or more) by 50 cm, occasionally with 4 leaflets inserted at some of the lower articulations; of the flowering branches smaller with fewer pinnae or more frequently unifoliolate; petiole up to 25 cm; petiolules 10-20 mm; leaflets elliptic, oblong or lanceolate, up to 24 by 7!/, cm, coria- ceous, margin strongly revolute, base cuneate to rounded, often oblique, apex narrowed to an acute often caudate apiculum, midrib prominent, lateral veins faint to obscure. Inflorescence a cluster of specialized leafy branches forming a terminal crown; main rays up to 70 cm, bearing an opposite pair (or rarely more pairs) of unifoliolate or (less frequently) pinnate leaves, often with flowering branches in their axils, and ending in a whorl of secondary rays, subtended by usually unifoliolate leaves; secondary rays up to 25 cm, bearing a pair of usually unifoliolate leaves with flowering branches in their axils, and terminating in com- pound umbellules; tertiary rays c. 10, c. 2-3 cm long, articulated near the middle; pedicels c. 12-14, c. 3-4 mm long, rufous-tomentose. Petals 5, broadly triangular, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, rufous-tomentose, c. 1'/, mm long at anthesis; disk fleshy, stigma sessile. Fruit spheroidal, c. 6 by 5 mm (when dry), calyx and stylopodium prominent. Distr. Malesia: Borneo (Sarawak and Kaliman- tan). Ecol. Swampy peat forest and heath woodland, both primary and disturbed, from sea-level to c. 150m. Some fragmentary collections from higher altitude (1000 m) further inland may belong to this species. Note. The leaves associated with the in- florescence are distinctively fleshy, have a strongly revolute margin and a smooth lower surface with indistinct lateral veins, and are ovate with rather acute apex. The leaflets of the pinnate leaves on the vegetative shoots are also rather leathery with indistinct lateral venation. 8. Arthrophyllum pacificum PuHILIPSON, Gard. Bull. Sing. 30 (1977) 306. A slender, glabrous tree to 14 m. Leaves multi- jugate, c. 60 by 24 cm; of the flower-bearing branches smaller with fewer pinnae or simple; petioles c. 22 cm, 3-4 mm @; petiolules c. 1 cm; leaflets elliptic, oblong or ovate, c. 8-10 by 4-5 cm, rather membranaceous, margin entire, very slightly revolute, base abruptly cuneate, often oblique, apex obtuse, acute, or slightly apiculate (in a specimen from Morotai attenuated). Inflorescence a whorl of specialized leafy branches forming a terminal crown to the vegetative shoots; main rays 35 cm, bearing small pinnate leaves in opposite pairs and with flowering branches in the axils, ending in a whorl of secondary rays, subtended by a whorl of simple leaves (bracts); secondary rays 10-24 cm long, bearing simple or trifoliolate leaves usually in opposite pairs, and terminating in umbellules; umbellules c. 7—-12-flowered; pedicels c. 5 mm at anthesis, slightly elongating in fruit, minute bracts caducous. Petals 5, broadly triangu- lar, 2'!/,-3 mm long. Stamens 5, anthers reniform. Ovary obconical, 1'/, mm long; disk fleshy, furrowed when dry; stigma + sessile. Fruit ellipsoidal, fleshy, somewhat oblique, conical beak with calyx and stigma, c. 9 by 5 mm when dry. Distr. E. Malesia: Moluccas (Morotai) to the Bismarck Archipelago. Ecol. Primary forest on mountain slopes, attaining Nothofagus mossy forest, 500-2000 m. Reported as common in most localities. Vern. Kainsoka, Ambai, Japen I. Notes. All specimens from islands to the north of New Guinea are similar, in having more delicate foliage than 10. A. macranthum from the mainland of New Guinea. The specimens from Morotai are sterile, and have narrower more tapering leaflets. The bark is light brown and the cut branches exude a clear latex. The flowers are light green with yellow anthers, and the fruit is black. 9. Arthrophyllum diversifolium BL. Bijdr. (1826) 879; DC. Prod. 3 (1830) 266; Mia. Fl. Ind. Bat. 1, 1 (1856) 767; Sum. (1861) 340, incl. var. lanceolata Miq.; CLARKE, FI. Br. Ind. 2 (1879) 733; K. & V. Bijdr. 7 (1900) 46; Koorp. Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 675 & 676; BACK. & BAKH. f. Fl. Java 2 (1965) 169; Stone, Gard. Bull. Sing. 30 (1977) 135; Puttieson, /.c. 306, f. 15. — A. javani- cum BL. Bijdr. (1826) 879; DC. Prod. 4 (1830) 266; Back. & BAKH. f. Fl. Java 2 (1965) 169. — A. ellipticum BL. Bijdr. (1826) 879; DC. Prod. 3 (1830) 266. — Mormoraphis sumatrana JACK ex WALL. Cat. (1831) n. 4931, nomen. — A. blumeanum Z. & M. Syst. Verz. (1846) 41, nom. illeg.; Miq. Fl. Ind. Bat. 1, 1 (1856) 768; Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27, incl. var. oblongatum Miq., var. ellipticum (BL.) Mia. et var. ovalifolium (JUNGH. & DE VRIESE) Mig. — A. ovatifolium JUNGH. & DE VRIESE, Ned. Kruidk. Arch. 1 (1846) 19; Ann. Sc. Nat. Paris III, 6 (1846) 117; Mia. Fl. Ind. Bat. 1, 1 (1856) 768, t. 14 (‘ovalifolium’); Rip. Fl. Mal. Pen. 1 (1922) 885; Stone, Gard. Bull. Sing. 30 (1977) 136, f. 2. — A. dilatatum Mio. FI. Ind. Bat. 1, 1 (1856) 768. — A. borneense BAKER, Kew Bull. (1896) 23; Merr. En. Born. (1921) 457. — A. congestum Riwv. J. Fed. Mal. St. Mus. 10 (1920) 137; Fl. Mal. Pen. 1-(1922) 887; STONE, Gard. Bull. Sing. 30 (1977) 136. — A. havi- landii RL. Kew Bull. (1933) 494. — A. rufosepa- lum Rip. Kew Bull. (1946) 40. — A. rubiginosum RIDL. /.c. 41. — Fig. 17, 21c, 22, 23. Small tree, up to 14 m, young parts with rufous tomentum. Leaves clustered at the ends of the branches, imparipinnate or bipinnate (rarely tripinnate) with leaflets at the insertion of the lateral rachides, multijugate, 150 by 45 cm (wider in bipinnate leaves); usually in opposite pairs on 1979 ] yy Ae ve GR Se Ei) a ARALIACEAE—I (Philipson) 61 MT gs a I, L—~) Cw as Zs SY 2 AS Roy eae CASS » BS AS Sli eS OS LS WS LOPES” _ as 3 q QS" _ SY Fig. 22. Arthrophyllum diversifolium BL. Diagrammatic sketch of habit, showing vegetative and flowering branches, < 1/39 (Courtesy Gard. Bull. Sing. 30, 1977). the inflorescence-bearing branches and smaller with fewer pinnae or unifoliolate; petiole up to 40 cm; petiolules '/,—1'/, cm; leaflets ovate-oblong or elliptic, up to 24 by 11 cm (leaflets of bipinnate leaves usually c. 10 by 5 cm), + coriaceous or (especially in bipinnate leaves) somewhat mem- branaceous, margin slightly revolute, base truncate, rounded, or cuneate, often oblique, apex shortly acuminate, veins usually 5-7 pairs. Inflorescence a cluster of specialized leafy branches forming a terminal crown which abscisses after fruiting; main rays up to 150 cm, bearing pinnate (or more rarely unifoliolate) leaves mostly in opposite pairs and usually with flowering branches in the axils of the upper leaves, ending in an umbel of secondary rays subtended by a whorl of pinnate or unifoliolate leaves; secondary rays up to c. 30 cm, bearing mainly simple leaves in opposite pairs with flowering branches in their axils and terminating in compound umbellules each subtended by a whorl of often caducous bracts; tertiary rays (peduncles) c. 5 cm, articulated about the middle; umbellules with c. 30-40 flowers, c. 17-20 mm @; pedicels c. 20, c. 5 mm at anthesis (longer in fruit), 62 FLORA MALESIANA (ser. I, vol. 91 Fig. 23. Arthrophyllum diversifolium BL. Schematic drawing of a single flowering shoot, with four degrees of branching, * 1/; (Courtesy Gard. Bull. Sing. 30, 1977). 1979 ] with an involucre of minute caducous bracts. Petals 5, c. 2 mm long. Stamens 5, anthers curved. Ovary turbinate, often inconspicuous at anthesis; disk fleshy, rising in the centre to a sessile stigma. Fruit ellipsoidal, c. 9 by 7 mm; calyx and stylo- podium forming a conspicuous beak. Distr. Malesia: Sumatra, Malay Peninsula, Java, Borneo, Celebes. Ecol. In a wide variety of habitats, on dry sandy soil to swampy humus, in primary lowland and montane rain-forest and also in secondary forest, heath-forest and waste land, from sea-level to 1600 m. Uses. Concoctions of the root and bark are reported to have medicinal properties, including a remedy for syphilis, and the plant has stupifying and poisonous properties. Vern. Sumatra: antjaneudeung uding, bidju, bolu bolu, bulu, (kayu) abang-abang, k. attu turut, légung, Riouw Arch., mapang, miu, obang, potah, silanta, ténjam dakan, Banka, tocrah. Malay Peninsula: chindangan utan, jolok hantu, lupa dahan, (pokok) restong, ségan budahan, susun kélapa, tum bong ninyor, tusum perpah. Java: dajo, délahan, déleg, délek, djangkorrang, gombong, (kayu) gompang, kédrja, kidjangkurang, kingom- pang, ki ompong, kléntjang, krépang, langitj, malas bérdahan, pongporang, putjangan. Borneo: merje- meh, Sarawak, karadjungjung, Kalimantan. Celebes: kambabah, susangkangan. Notes. The very widespread A. diversifolium is variable in many characters, and may comprise a number of geographic subspecies, but no basis for this is apparent at present. Most individuals have the lower leaves simply imparipinnate, whereas others have bipinnate, or rarely tripinnate, leaves. The flowers and inflorescences of these forms appear to be identical, though rapid changes in the umbellules after flowering produce a deceptively distinctive appearance in specimens at different stages of development. Field experience over the whole range of the species will be required to understand this interesting leaf-polymorphism. In treating all forms as one species I am partly influenced by the fact that most authors who have been familiar with the plants in Java (where both forms occur) have regarded the complex as a single species (the fact that some authors have recognized the variant from Mt Salak as a distinct species does not affect the problem of leaf-polymorphism). Apart from the strikingly different leaf forms just discussed, certain iocal variants may eventually be shown to justify specific rank. A form growing on Mt Salak (near Bogor) has often been regarded as distinct (see, for example, HOCHREUTINER, Candollea 2, 1925, 481, and BACKER & BAKHUIZEN VAN DEN BRinK f. FI. Java 2, 1965, 169). Indeed this form is the basis of the name A. diversifolium. I retain this name in preference to the other two names published simultaneously by BLUME because it has been most consistently adopted since ARALIACEAE—I (Philipson) 63 it was first used in this comprehensive sense by CLARKE, /.c. On the evidence available I do not consider the Salak plants any more distinctive than many other local variants. It might be considered that 5. A. engganoense is also no more than another such variant, but its facies is so marked that specific rank appears justified. It is possible that RIDLEY was correct in distin- guishing A. congestum, but the material is not good and appears inadequate to confirm specific status. Five collections from Brunei and a neighbouring district of Sarawak are all very alike and sufficiently distinct from both A. diversifolium and 7. A. crassum to suggest that they represent a separate taxon, but for the present they are tenta- tively retained as a form of A. diversifolium. Similarly, the two collections described by RIDLEY as A. rubiginosum and A. rufosepalum are based on collections which are not altogether typical of A. diversifolium, but which come closest to that species. In the absence of more supporting material, it is advisable not to retain them as species. The first of these names (A. rubiginosum) has been widely used in identifications of Bornean specimens, but the specimens concerned are either typical A. diversifolium or belong to the distinctive 7. A. crassum. Specimens from Mt Kinabalu described by RIDLEY as A. havilandii have bipinnate leaves, and appear to conform well with A. diversifolium. This form was again collected on. Mt Kinabalu by CLEMENS and is also known from Sarawak. The smooth bark is whitish to greyish brown with pustulate lenticels; the wood is cream, with a colourless aromatic exudate. The flowers are yellowish with a sickly sweet scent. Seedlings have simple and trifoliolate leaves. 10. Arthrophyllum macranthum PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 18; Gard. Bull. Sing. 30 (1977) 308, f. 16. — A. diversifolium (non BL.) Harms, Bot. Jahrb. 56 (1920) 413. — Fig. 21d. Tree up to 25 m, sparsely branched with leaves crowded at the ends of the branches, all young parts with dense rufous tomentum which may persist on the flowers and inflorescence. Leaves multijugate, up to 100 by 30 cm; of the flower- bearing branches smaller with fewer pinnae, or simple; petioles stout, up to 40 cm, clasping base heavily lenticellate; petiolules '/,-2 cm; leaflets ovate to oblong, up to 16 by 8 cm, coriaceous, mar- gin revolute, base rounded, truncate, or cordate, very rarely cuneate, often oblique, apex obtuse or bluntly apiculate, principal veins arched-ascending, reticulations visible especially beneath (pinnae of leaves on the flowering branches usually elliptic with a cuneate base). Inflorescence a whorl of specialized leafy branches forming a terminal crown; main rays up to 60 cm, bearing pinnate leaves often in opposite pairs and with flowering 64 FLORA MALESIANA [ser. I, vol..91 branches in the upper axils, and ending in a whorl of secondary rays subtended by a whorl of usually simple leaves (bracts); the secondary rays c. 10- 20 cm, bearing simple leaves, usually in opposite pairs, and terminating in compound umbels; umbellules (c. 2 cm) with c. 10-15 flowers; pedicels stout, 3-5 mm to 10 mm or more in fruit, sub- tended by minute bracts. Flower buds 5 mm or more long; calyx a rim or with 5 indistinct teeth. Petals 5, triangular, c. 4 mm long, fleshy. Stamens 5. Ovary obconical, c. 2'/, mm long; disk fleshy (hemispherical in living material, conical and furrowed when dry); stigma capitate, ++ sessile. Fruit ellipsoid, sometimes slightly oblique, c. 12 by 8 mm, the stylopodium forming a conical beak with the persistent stigma; exocarp fleshy, endo- carp cartilaginous. Distr. Malesia: New Guinea (from Irian Jaya to Milne Bay Distr. and the Bismarck Archipelago). Ecol. Usually a sub-canopy tree of rain-forest ranging from the lower montane zone to mossy subalpine woodland and scrubland, occasionally in second growth, usually above 1000 m (up to 2700 m), but occurring also on the coastal scarps of the Astrolabe Range. Vern. Maguva, W. Sepik, agagwa, agare, agugwa, alolo, angga, engga, pooli, tipilan, W. High- lands, arua, hagegoa, wonkurumeh, E. Highlands, kolom, S. Highlands. Note. Trees become very different in appearance when in flower or fruit: the spiral pinnate foliage leaves are surmounted by tufts of branches which end in inflorescences and bear much smaller leaves. The foliage leaves are fleshy, leathery and glossy. The ripe fruit is purple and shining. The bark is grey, at first smooth with many leaf-scars and lenticels, but small longitudinal fissures develop. The cut branches exude a brown latex and a scent of celery. The soft wood is white or straw-coloured. 11. Arthrophyllum papyraceum PHILIPSON, Gard. Bull. Sing. 30 (1977) 308. Shrub, rufous-tomentose on the young parts. Leaves alternate, imparipinnate; petiole 15-20 cm; leaflets c. 7, membranaceous, elliptic, c. 12-24 by 5-10 cm, base broadly cuneate, apex finely acuminate, margin slightly revolute. Inflorescence a terminal compound umbel; primary rays few (2), 2-3 cm long, 2 mm wide, without bracts (caducous), secondary rays few (3), c. 13-18 mm long, articu- lated about the middle, ending in an umbellule of c. 10-12 flowers; pedicels 2-3 mm, slightly fur- furaceous. Petals 5, c. 2 mm long in bud. Stamens 5, anthers curved. Ovary turbinate, glabrous, obscurely ribbed. Fruit unknown. Distr. Malesia: E. Sumatra (East Coast Res. near Aek Sordang), one collection. Ecol. Primary rain-forest. Note. Known from a single collection (with no duplicates) this species resembles 12. A. mainyayi in its simple inflorescence, and the few pinnae of its foliage leaves. However, the large size of the leaflets precludes its inclusion in that species. 12. Arthrophyllum maingayi PHILIPSON, Gard. Bull. Sing. 30 (1977) 309, f. 7. — A. pinnatum CLARKE, Fl. Br. Ind. 2 (1879) 734, excluding synonyms [see also SEEM. J. Bot. 4 (1866) 294]; RIDL. Fl. Mal. Pen. 1 (1922) 886. — Fig. 19c, 20a. Low shrub or slender tree, rarely as high as 10 m, rufous-tomentose on the very young parts, soon becoming glabrous. Leaves tufted at the ends of the branches, usually with 5—7 leaflets (but up to 15), up to 30 by 18 cm; petiole to 12 cm (usually shorter), 2 mm @; petiolules c. 0-10 mm; leaflets elliptic or elliptic-lanceolate (occasionally the lowermost pair of leaflets is replaced by pinnate leaf segments), c. 8 by 31/, cm, rather thin, base cuneate or rounded, apex apiculate or caudate, margin slightly revolute, lateral veins faint and obscure; leaves below the flowering branches sometimes reduced to 3 or 1 leaflet(s). Inflorescence consisting of a number of primary branches radiating from the end of a leafy shoot (which forms a longer or shorter peduncle); primary branches, often rather few, usually 3-6 cm long and devoid of leaves except for a few terminal simple or trifoliolate leaves around the compound umbels, occasionally the branches bear pairs of opposite leaves when they may be up to 30 cm long; secondary rays c. 2-3 cm long; pedicels 4-10 cm. Petals 5, 1/,-2 mm long in bud. Stamens 5. Ovary turbinate, disk fleshy. Fruit spheroidal, c. 8 by 6 mm, calyx and stylopodium rather inconspicuous. Distr. Malesia: Central W. Sumatra (Mt Kerintji), throughout the Malay Peninsula (incl. Penang) and Borneo. Vern. Poko minta anak, Kedah, karon baru, Sumatra. Notes. The name ‘A. pinnatum’, misapplied by CLARKE, /.c., has been in general use for this species. However, this name was based on Panax pinnatum LAMK, which in turn was based on the Rumphian name ‘Scutellaria secunda’ ; this is quite a distinct plant (see under 6. Polyscias cumingiana). The three species A. maingayi, 16. A. montanum and 17. A. alternifolium are similar in having simpler inflorescences than 9. A. diversifolium. A. montanum can be distinguished from A. main- gayi by its more leathery leaves with more promi- nent nervation, and by the more woody and leafy flower-bearing primary branches of the inflores- cence. A. alternifolium is distinguished from both these species by its small, coriaceous leaflets with incon- spicuous nervation. Most specimens can be readily distinguished, but a few can be assigned to a species only doubtfully, usually because the material is inadequate. This is usually due to the junction between the vegetative (spiral phyllotactic) shoots and the flower-bearing branches being omitted. 1979 ] 13. Arthrophyllum kjellbergii PHiLipson, Gard. Bull. Sing. 30 (1977) 309, f. 10. — Fig. 20d. Small tree, 10 m, branches c. 1 cm @, young parts covered by rufous tomentum. Leaves clustered towards the ends of the branches, multijugate, 25-30 by 16 cm; petioles c. 8 cm, 2 mm wide; petiolules c. 3-8 mm; leaflets elliptic or ovate- oblong, up to 8 by 3!/, cm, chartaceous, margin minutely revolute, base rounded to cuneate often oblique, apex tapered to a blunt apiculum, or rounded and mucronate. Inflorescence a terminal cluster of c. 5-10 specialized branches (primary rays); primary rays c. 15-20 cm, 1'/,-2 mm wide, bearing near the middle an opposite pair of simple leaves or sometimes trifoliolate leaves with short flowering branches in their axils, and sometimes with a second pair higher up, and 2-3 similar leaves below the terminal cluster of secondary rays; secondary rays c. 12, c. 2'!/,-3'/, cm long, each subtended by a small bract and bearing opposite caducous bracts near the middle, terminating in an umbellule of c. 12 flowers surrounded by an involucre of caducous bracts (1 mm long). Flowers known only in young bud. Fruit spheroidal, c. 5 by 4 mm, calyx and stylopodium prominent; pedicel 5-6 mm. Distr. Malesia: SE. Celebes (Kendari). Ecol. Primary rain-forest, 50-150 m. Note. The small leaves and relatively simple inflorescences are distinctive. 14. Arthrophyllum cenabrei Merr. Philip. J. Sc. 20 (1922) 417; En. Philip. 3 (1923) 235; PHILIPSON, Gard. Bull. Sing. 30 (1977) 309. Glabrous tree, c. 10 m, ultimate branches c. 5mm @. Upper /eaves pinnate, up to 10 cm long, leaflets mostly 5, sometimes 3, or the uppermost reduced to simple leaflets, the rachis and petiole c. 4 cm; leaflets mostly elliptic, 4'/,-6 by 21/,-31/, cm, chartaceous to subcoriaceous, very shortly and obtusely acuminate, base acute, brownish oliva- ceous and slightly shining when dry, nerves 3-4 pairs, slender; petiolules 5-10 mm. Peduncles c. 4 cm, umbellately arranged at the tops of the branchlets, usually however with solitary inflorescences in the axils of the uppermost leaves, thus forming a somewhat leafy inflorescence. Fruits 5-8 in each umbel, ovoid, c. 7 mm @; pedicels 8-10 mm. Distr. Malesia: Philippines (Cebu; FB 28343, type, not seen). Ecol. On slopes at 600 m. Vern. Bingliu, C.Bis. Note. No specimen of this species has been located. The above description is taken from MERRILL’s original account. In placing this species in the key, it has been assumed that the inflorescence branching is relatively simple. 15. Arthrophyllum pulgarense ELmer, Leafl. Philip. Bot. 7 (1915) 2551; Merr. En. Philip. 3 (1923) 235; ARALIACEAE—I (Philipson) 65 PHILIPSON, Gard. Bull. Sing. 30 (1977) 311, f. 9. — Fig. 20c. Small tree, branches c. 1 cm @, young parts with red tomentum, becoming glabrous except on the ovaries. Leaves clustered towards the ends of the branches, leaflets c. 6 pairs, c. 22 by 8 cm; petioles c. 6 cm, 3 mm @; petiolules 5-6 mm; leaflets elliptic to rotund, c. 4 by 23/, cm, coriaceous, margin revolute, rounded to broadly cuneate, apex rounded or abruptly tapered to a short obtuse apiculum. Inflorescence a terminal cluster of specialized branches (primary rays); primary rays c. 6-10 cm, 3-4 mm @, bearing near the middle an opposite pair of simple rotund leaves, sometimes with flowering branches in their axils, and with a whorl of similar leaves below the terminal cluster of secondary rays; secondary rays c. 6-8, 2-4 cm long, articulated about the middle with scars of bracts or bearing a pair of small simple leaves terminating in an umbellule of c. 8-12 flowers; bracts caducous. Calyx with indistinct teeth. Petals 6, 2'/, mm long (in bud). Stamens 6, anthers curved. Ovary obconical, 2 mm long, furfuraceous. Fruit ellipsoid, c. 8 by 5 mm, the stylopodium forming a beak with stigma and calyx; exocarp fleshy. Distr. Pulgar). Ecol. Common in montane forest on Mt Pulgar. Note. The coriaceous, small, often rotund leaf- lets are characteristic. Malesia: Philippines (Palawan: Mt 16. Arthrophyllum montanum RuIDL. J. Fed. Mal. St. Mus. 4 (1909) 24; Fl. Mal. Pen. 1 (1922) 886; Puiieson, Gard. Bull. Sing. 30 (1977) 311, f. 8. — A. nitidum Rip. J. Fed. Mal. St. Mus. 7 (1916) 42; Fl. Mal. Pen. 1 (1922) 886. — A. ovatum RIDL. J. Fed. Mal. St. Mus. 7 (1916) 42; Fl. Mal. Pen. 1 (1922) 886. — Fig. 19d, 20b. Shrub or small tree to 6 m, unbranched or sparingly branched, rufous-tomentose on the young parts, becoming glabrous. Leaves tufted at the ends of the branches, multijugate, c. 30-55 by 12-22 cm; petiole 9-21 cm, 3 mm @; petiolules c. 10-15 mm; leaflets elliptic or oblong, 6-10 by 21/,4 cm, coriaceous or chartaceous, base cuneate, apex with a short blunt apiculum, margin entire, revolute, the few principal lateral veins usually rather prominent; the leaves associated with the umbels usually unifoliolate, broadly ellip- tic to rotund, with a petiole to 4'/, cm. /nflores- cences on specialized leafy branches either in terminal clusters or axillary in the upper leaves; branches 10-30 cm, leaves mostly simple in opposite pairs, usually without flowering branches in their axils, branches ending in a whorl of simple leaves surrounding a compound umbel; primary rays c. 5—-15(-20), 2-6 cm, with scars of caducous bracts; pedicels 4-8 mm. Petals (4-)5, 2 mm long in bud. Stamens (4~-)5. Ovary turbinate; disk fleshy, stigma + sessile. Fruit spheroidal, c. 8 by 66 FLORA MALESIANA [ser. I, vol. 9! Fig. 24. Gastonia serratifolia (M1Q.) PuiLipson. a. Habit, x '/3, b. lower bud, c. flower, d. fruit, both x 5 (a—c SCHMUTZ 3612, d VAN ROYEN 4090). Drawn by HELENE MULDER. 1979] 5 mm, calyx and stylopodium rather prominent. Distr. Malesia: Malay Peninsula (Kedah to Selangor). Ecol. Primary forest ascending to the montane zone, and in second-growth, 250-1500 m. Note. Similar to 12. A. maingayi but dis- tinguished by the more leathery leaves and by the leafy inflorescence branches which are usually absent in A. maingayi. 17. Arthrophyllum alternifolium MAINGAyY ex RIDL. FI. Mal. Pen. 1 (1922) 886; PHILIPSON, Gard. Bull. Sing. 30 (1977) 311, f. 11. — A. pinnatum CLARKE, FI. Br. Ind. 2 (1879) 734, p.p., excl. basionym; KiNG, J. As. Soc. Beng. 67, ii (1898) 59, p.p. — A. alternifolium MAINGAY ex CLARKE, FI. Br. Ind. 2 (1879) 734, nomen in synon. — Fig. 20e. Slender, sparingly branched shrub to 2 m, rufous-tomentose on the young parts, becoming glabrous. Leaves tufted at the ends of the branches, multijugate, c. 20-25(-30) by 9-12(-15) cm; petiole terete, c. 3-6(-9) cm, 2-3 mm @; petiolules c. 2 mm; leaflets ovate, elliptic or lanceolate, 31/,-4(-6) by 1-2(C2!/4) cm, coriaceous, base cuneate, apex acuminate to caudate, obtuse, margin revolute, veins obscure; leaves associated with the umbels (if any) reduced, with fewer leaflets or unifoliolate, sometimes broadly ovate. Inflorescence usually a terminal compound umbel, occasionally a whorl of leafy branches (5-14 cm long) (leaves usually simple in opposite pairs), each ARALIACEAE—I (Philipson) 67 ending in a compound umbel; peduncle 1'/,— 4'/, cm, with one or more usually caducous simple (or trifoliolate) leaves at the apex; primary rays c. 5, 3-4'/, cm, with scars of caducous leaves about the middle, each ending in an umbellule of c. 12-25 flowers, pedicels 5-8 mm. Calyx a rim or minutely 4—5-dentate. Petals 4-5, 2 mm long in bud. Stamens 4-5. Ovary turbinate; disk fleshy, stigma + sessile. Fruit spheroidal, c. 5 by 5 mm when dry, calyx and stylopodium small. Distr. Malesia: Malay Peninsula (Johore: Mt Ophir; Pahang, Selangor, and Malacca). Ecol. In shady montane forest, with Rhodo- dendron and Dacrydium, 900 m and above. Note. The small, coriaceous, often apiculate leaflets are characteristic. Although collected most frequently on Mt Ophir it occurs on other high ridges in southern Malaya. Excluded Arthrophyllum ceylanicum Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27 (type in L), is according to VAN STEENIS, Rec. Trav. Bot. Néerl. 24 (1927) 819 = Oroxylum indicum (L.) KURZ (Bignoniaceae). Arthrophyllum reticulatum BL. ex MiqQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27 (type in L), is according to MIQUEL (/.c. 318) and VAN STEENIS (vide supra) = Oroxylum indicum (L.) Kurz (Bignoniaceae). 9. GASTONIA Comm. ex LAMK, Encycl. 2 (1786) 610; Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43; Hutrcu. Gen. FI. Pl. 2 (1967) 68; PHILIPSON, Blumea 18 (1970) 491, 497, f. 1-10. — Tetraplasandra (non A. GRAY) Mia. Ann. Mus. Bot. Lugd.-Bat. | (1863) 4; HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 29, p.p.; Nachtr. 2 (1900) 253. — Indokingia HEMSL. in Hook. Ic. Pl. (1906) t. 2805. — Peekeliopanax Harms, Notizbl. Berl.-Dahl. 9 (1926) 478, fig. — Fig. 24, 26, 27. Trees unarmed with thick branches, glabrous or tomentose. Leaves large, im- paripinnate, exstipulate; rachis articulated; leaflets in pairs entire or crenate; petiole terete, with clasping base. Flowers in umbellules which are arranged race- mosely, or in verticils, on strong inflorescence branches; pedicels not articulated below the ovary. Calyx forming a continuous rim with an entire or indistinctly denticulate margin. Corolla of 5-13 free petals or calyptrate, fleshy, valvate. Stamens either equal in number to the petals or up to several times as many; filaments usually rather short and thick; anthers large, often irregularly lobed, dorsifixed. Ovary inferior, broadly obconic, cells 7-22; disk fleshy with stylar processes equal in number to the cells arising from its centre. Fruit a spherical berry (strongly ribbed when dry), with an indistinct calyx rim, a flattened disk, and a 68 FLORA MALESIANA [ser. I, vol. 91 prominent stylopodium bearing a ring of radiating stigmatic arms; exocarp fleshy, endocarp crustaceous. Endosperm with smooth surface. Distr. About 10 spp. in East Africa, Madagascar, the Seychelles and Mascarenes, Malesia, and the Solomon Is. Ecol. Primary and second-growth forest, or in open country, from sea-level to lower montane zone. Taxon. I have amply analyzed and discussed the affinities of Gastonia (Blumea 18, 1970, 497). I have come to the conclusion that within its alliances Gastonia is the only genus west of Samoa; in Polynesia there are three other closely allied genera. KEY TO THE SPECIES 1. Corolla with free petals. Ovary cells and style arms 6—-9(-12). Whole plant glabrous. Leaflets usually entire or sparsely crenate. Main inflorescence branches borne along an elongated axis with caducous bracts; peduncles of the umbellules mostly aggregated into pseudo-whorls 1. G. serratifolia 1. Corolla calyptrate. Ovary cells and style arms 12-18(—22). Young parts with scurfy tomentum, per- sisting on the ovary and bracts. Leaflets strongly crenate. Main inflorescence branches borne sub- umbellately on a short axis with persistent bracts; peduncles of the umbellules scattered 1. Gastonia serratifolia (MIQ.) PHILIPSON, comb. nov. — Arthrophyllum serratifolium Mi1Q. Sum. (1861) 341, type from Sibolga, leg. TEYSMANN (in U). — G. papuana Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 5; Harms, Bot. Jahrb. 56 (1921) 408; Puitipson, Blumea 18 (1970) 492, 500 f. 3. — Tetraplasandra paucidens Mia. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 4; Koorp. Minah. (1898) 488. — G. eupteronoides T. & B. Nat. Tijd. N. I. 25 (1863) 416. — Polyscias papuana (MiIQ.) SEEM. J. Bot. 3 (1865) 181. — Tetraplasandra koordersii Harms, Ann. Jard. Bot. Btzg 19 (1904) 12; Ic. Bog. 2 (1906) t. 178. — Tetraplasandra philippinen- sis MerRR. Philip. J. Sc. 1 (1906) Suppl. 219; En. Philip. 3 (1923) 222. — G. winkleri Harms in Fedde, Rep. 15 (1917) 20. — Tetraplasandra solomonensis PHILIPSON, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 11. — Fig. 24. Shrub or small tree, sometimes epiphytic, up to 27 m, with clear bole to 15 m, dbh 34 cm, crown sparsely branched with leaves crowded at the ends of the branches, glabrous. Leaves up to 80 by 20 cm; petiole c. 13 cm; leaflets c. 10 pairs, petiolules up to 1 cm; blade oblong, ovate or lanceolate, middle leaflets 8-14(-18) by 2'/,- 31/,(-8) cm, chartaceous, entire and subrevolute, or with a few obscure, more rarely several promi- nent crenations, apex rounded and bluntly apicu- late or tapering and acute, base broadly cuneate, midrib prominent. Inflorescence terminal, glab- rous, with a stout primary axis 15-25 cm long, bearing scattered or clustered branches along its length and ending in an umbel of c. 10 branches; bracts caducous; secondary branches 30-40 cm, bearing numerous subverticillate peduncles along their length and ending in an umbel; peduncles c. 3 cm, somewhat longer in fruit; pedicels 1-2'/, cm, forming umbellules of c. 10 flowers. Flower buds (when dry) c. 7 by 3 mm. Calyx rim 2. G. spectabilis undulate. Petals 5-9, slightly fleshy, fully separated. Stamens variable in number, (7—)14—55; filaments short; anthers broad and irregularly lobed, variable in size. Ovary glabrous, smoothly rounded below, slightly constricted below the calyx; cells 6-12, usually c. 9, disk with a prominent rim and, at anthesis, a central boss formed by closely appressed subulate style arms equal in number to the ovary cells. Fruit c. 9 by 7 mm (without stylopodium), the flattened stylopodium ending in a ring or double row of radiating subulate stigmatic arms, black when ripe, the fleshy exocarp enclosing com- pressed crustaceous pyrenes. Distr. Solomon Is.; in Malesia: Central W. Sumatra (Sibolga, Enggano I.), Malay Peninsula (Johore), Sunda Straits (islet Dwars in den Weg), West Java, Lesser Sunda Is. (Sumba, Timor, Wetar, Flores), W. Borneo (east of Pontianak, Karimata), N. Borneo (Sabah), Philippines Fig. 25. Distribution of Gastonia in Malesia. Known localities of G. serratifolia (MIQ.) PHILIP- son: black dots; range of G. spectabilis (HARMS) PHILIPSON: broken line. 1979 ] ARALIACEAE—I (Philipson) 69 2 = " Sr Cre 77 ote oN em = % Fig. 26. Gastonia spectabilis (HARMS) PHILIPSON, two inflorescences visible below the leaves (Photogr. PHILIPSON, Kassam Pass, E. New Guinea, 1968). 70 FLORA MALESIANA [ser. I, vol. 9? (Palawan, Balabac), Celebes (Lepo-Lepo, Luwuk, Minahassa), N. Moluccas (Talaud Is.), New Guinea (Vogelkop, NW. & SW. Irian, Schouten I., Waigeu I.). Fig. 25. Ecol. Primary and secondary forest, or in open country, usually at low altitude and often on the shore or sea-cliffs, but ascending to 1000 m. EYMA noted it to be a characteristic tree on Mt Tam- bunan, Luwuk, E. Celebes. Vern. Bajur talang ékoaho, M, Enggano, jarum, Sabah, raka, Sumba, kre, wangka, Flores, lampo pad, bungku, Celebes, bufgio, Palawan, /antora, Talaud Is., mansnongoree, mantsenongor, Schouten I., raauwrack, ara-orach, Vogelkop, Maibrat. Note. The entire or serrate nature of the leaf margins, the number of ovary cells, and especially the number of stamens are variable characters but show no discernable geographical segregation. The outer bark is described as light brown, with small oblong brittle scales. Inner bark, leaves and inflorescences with copious sticky juice. Wood soft white. The petals are variously described as purple, light green, and white. 2. Gastonia spectabilis (HARMS) PHILIPSON, Blumea 18 (1970) 494, pl. 1. — Peekeliopanax spectabilis Harms, Notizbl. Berl.-Dahl. 9 (1926) 478, fig.; PHILIPSON, Blumea 18 (1970) 500 f. 2. — G. bori- diana Harms, Bot. Jahrb. 69 (1938) 282. — Fig. 26, 27. Tree up to 40 m high, clear bole to 28 m, dbh 1.75 m, crown sparsely branched with the branches whorled or regularly forked and the leaves crowded at the ends of the branches, all young parts with scurfy indumentum more evident in dried material. Leaves up to 80 by 30 cm; petiole c. 15 cm, with some tomentum remaining at the joints; leaflets c. 11 pairs on petiolules c. 2-8 mm, oblong or elliptic, middle leaflets 10-15 by 4-6 cm, charta- ceous when dry, prominently crenate, apex nar- rowed to a short blunt apiculus, base rounded or truncate. Inflorescence in forks well below the leaves, when in bud covered with large scurfy cataphylls, and when mature with radiating branches forming clusters c. 130 cm @. Primary axis short (5-10 cm) with persistent bracts; secondary branches numerous, radiating, up to 65 cm, bearing small, persistent, scurfy bracts and peduncles scattered along their length and clustered in a terminal umbel; peduncles c. 11/,-5 cm; pedicels +/,-1!/, cm, forming umbellules of c. 5-12 flowers. Flower buds c. 8 by 5 mm when dry (fresh c. 12 by 8 mm). Calyx rim straight. Petals 6-12, very fleshy, incompletely separated (often splitting into c. 5 lobes). Stamens 25-66, often c. 35, filaments short; anthers broad and irregularly lobed, variable in size. Ovary with a short dense indumentum, smoothly rounded below, cells usually c. 16, very rarely fewer than 10, as many as 22; disk at anthesis with a prominent rim and a central boss formed by closely appressed subulate Fig. 27. Gastonia spectabilis (HARMS) PHILIPSON. Young tree, free bole 19 m, 47 cm @, crown 10 m, longest leaves 2 m, not yet flowering (Photogr. G. PEEKEL, New Ireland, Ugana, 1940). styles equal in number to the cells. Fruit c. 8 by 10 mm (dry and without stylopodium), the flattened disk with a prominent stylopodium ending in an elliptical ring of radiating subulate stigmatic arms; the fleshy exocarp enclosing compressed crusta- ceous pyrenes. Distr. Solomon Is.; in Malesia: New Guinea (Vogelkop, NW. Irian, Papua New Guinea), Bismarcks, and New Ireland. Fig. 25. Ecol. Primary and secondary rain-forest, and in cultivated areas, 200-2000 m. Vern. Amoriga, bekuak, djak, ntjier, tuju, Vogelkop, bohko, boinga, gabiel, jamwa, mestic, sikoto, tubat, tumbala, Madang Distr., waki, W. Highlands, aita, E. Highlands, kuhuh, Papua. Note. Possibly the largest araliad known. The regular habit of branching results from the vegeta- 1979] ARALIACEAE— I (Philipson) 71 Fig. 28. Polyscias nodosa (BL.) SEEM. a. Leaf and part of inflorescence, x */s, b. leaflet, x 7/35 c. branch of inflorescence, x 2/3, d. umbellule, e. flower bud and ditto in LS, x 8, f. ovary and ditto in CS, x 8, g. Stamens (VERSTEEGH BW 3868). Drawn by HELENE MULDER. fp FLORA MALESIANA [ser. I, vol. 9} tive shoots springing in pairs or whorls from below terminal inflorescence buds. Several flushes of growth occur as an inflorescence bud matures, so that at anthesis the inflorescences are situated in forks well below the leafy crown, with a succession of younger inflorescence buds in higher forks. Bole without buttresses or with buttresses 1 m high and 2 m wide. Outer bark brown with prominent pustular lenticels and small shallow fissures. Exudate from cuts abundant, clear and aromatic. Wood soft. Flowers cream, stamens yellow. Ripe fruit dark red-brown. 10. POLYSCIAS J. R. & G. Forster, Char. Gen. (1776) 63, t. 32; DC. Prod. 4 (1830) 257; SEEM. J. Bot. 3 (1865) 179; Bru. in B. & H. Gen. Pl. 1 (1876) 941; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 43; Koorpb. Atlas 4 (1916) f. 677-680; Merr. Int. Rumph. (1917) 408; Harms, Bot. Jahrb. 56 (1921) 409; Merr. En. Philip. 3 (1923) 233; Hutcu. Gen. FI. Pl. 2 (1967) 75; BERNARDI, Candollea 26 (1971) 13; PHILIPsON, Blumea 24 (1978) 169. — Eupteron Mia. Pl. Jungh. 3 (1855) 423; Fl. Ind. Bat. 1, 1 (1856) 762; Hutcu. Gen. FI. Pl. 2 (1967) 68. — Nothopanax Mia. Pl. Jungh. 3 (1855) 425; Bonplandia 4 (1856) 139; FI. Ind. Bat. 1, 1 (1856) 765; SEEM. FI. Vit. (1866) 114; Merr. Int. Rumph. (1917) 409; En. Philip. 3 (1923) 233. — Irvingia F.v.M. Fragm. 5 (1865) 17, non Hook. f. 1860. — Kissodendron SEEM. J. Bot. 3 (1865) 201; ibid. 6 (1868) 129; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; Bot. Jahrb. 56 (1921) 412; Hurcu. Gen. FI. Pl. 2 (1967) 75. — Palmervandenbroekia Gipss, Arfak (1917) 162; HutcH. Gen. FI. Pl. 2 (1967) 75. — Gelibia Hutcu. Gen. Fl. Pl. 2 (1967) 57. — Fig. 28, 30. Unarmed shrubs or trees, glabrous or furfuraceous. Leaves imparipinnate or 2-3-pinnate (rarely unifoliolate) with an elongated or short sheathing base; rachis articulated; petiole terete; leaflets in pairs, entire, crenate or dentate. Inflorescence terminal, often large, a panicle, corymb, or compound rarely simple umbel. Flowers in umbellules, capitula, or racemose; pedicels articulated below the ovary. Calyx a rim with an undulate or dentate margin. Petals 4-5(-8 or more), valvate. Stamens equal in number to the petals; anthers dorsifixed. Ovary inferior, 4-5(-8 or more)-celled; disk fleshy; styles either free and recurved (at least in fruit) or joined to form a beak-like stylopodium. Fruit a spherical or ovoid drupe, crowned by the persistent calyx rim and the styles or stylopodium; exocarp fleshy, endocarp chartaceous. Endosperm with an uneven surface, fissured, or rarely smooth. Distr. About 100 spp. throughout the tropics of the Old World (incl. Australia), and the Pacific Islands; in Malesia 23 spp. of which 3 adventive: rare in West (none native in Sumatra and Malaya), more common in East, the majority in New Guinea. Fig. 29. Ecol. Primary or second-growth forest, from sea-level to 2650 m. Note. Araliads with pinnate leaves and an articulated pedicel are here regarded as forming one genus, divided into several sections. These sections have formerly been segregated as genera, but I follow BERNARDI, /.c., in uniting them. The most distinctive is sect. Polyscias, characterized by an elongated leaf- sheath. Several of the species of this section are cultivated and have a number of cultivars. KEY TO THE SECTIONS 1. Leaf-sheath elongated, extending along the petiole for about '/, of its length (Spp. 1-7) 1. Sect. Polyscias 1. Leaf-sheath short or obsolete, restricted to the base of the petiole. 2. Style arms spreading, at least in fruit. 1979] 3. Flowers arranged racemosely (Sp.8)... . 3. Flowers arranged in umbellules or capitula (Spp. 9-14)... .. 2... 2. Style arms erect, fused, forming a beak in fruit. 4. Inflorescence large, + as long as the leaves (Spp. 15-19) 4. Inflorescence much shorter than the leaves (Spp. 20-23). ARALIACEAE—I (Philipson) 73 2. Sect. Gelibia . . 3. Sect. Eupteron Ae as ee 4. Sect. Kissodendron 5. Sect. Palmervandenbroekia 1. Section Polyscias PHILIPSON, Blumea 24 (1978) 169. Aromatic, glabrous shrubs or small trees, often cultivated. Leaf-sheath elon- gated along the petiole for '/,;—'/, of its length. Styles spreading, at least in the fruit. Distr. Polynesia, Queensland, Malesia, and SE. Asia. KEY TO THE SPECIES HPRCAVES 2—S-PINNAtC.. uc es ee se 1. Leaves imparipinnate, or unifoliolate. 2. Ovary 2-celled (or predominantly so). 3. Main inflorescence branches diffusely branched 3. Main inflorescence branches with verticils of short branches oh) fap) ole oles gs Mleings. Bie te oe ae 6 le ee 1. P. fruticosa 2. P. macgillivrayi 3. P. verticillata 2. Ovary variable, but many flowers with more than 2 cells. 4. Leaflets orbicular or reniform. Leaves unifoliolate or trifoliolate. ...... 4. P. scutellaria 4. Leaflets ovate, oblong, or elliptic (cultivated forms often laciniate or lanceolate). Leaves with 3 or more pairs of leaflets. 5. Leaf margin sharply serrate (blade often rhomboidal and variegated with light yellow) 5. Leaf margin entire, or obscurely dentate. 5. P. guilfoylei 6. Peduncles of the ultimate umbellules bearing 1 or more pairs of small bracts (reduced flowering branches sometimes present in their axils) Stein Stel au) lie). ial. fel ey. GREK) 7 6) yey wD oe 6. P. cumingiana 6. Peduncles of the ultimate umbellules either without bracts, or with 1 or more small bracts inserted singly 1. Polyscias fruticosa (L.) HARMs in E. & P. Nat. Pfil. Fam. 3, 8 (1894) 45; K. & V. Bijdr. 7 (1900) 2; Harms in K. Sch. & Laut. Fl. Schutzgeb. (1900) 486; Koorp. Exk. Fl. Java 2 (1912) 716; Atlas 4 (1916) f. 680; BamLEy, Rhodora 18 (1916) 153, incl. var. plumata (Hort.) BAILEY; BACK. & BAKH. f. Fl. Java 2 (1965) 168; PuHittpson, Blumea 24 (1978) 169. — Scutellaria tertia RUMPH. Herb. Amb. 4: 78, t. 33. — Panax fruticosum LINNE, Sp. PI. ed. 2 (1763) 1513; BL. Bijdr. (1826) 830; DC. Prod. 4 (1830) 254; BLANco, FI. Filip. ed. 2 (1845) 156; ed. 3, 1 (1877) 281, t. 78; CLARKE, FI. Br. Ind. 2 (1879) 725; F.-ViLL. Nov. App. (1880) 101; Boer. Handl. 1 (1890) 647; F.v.M. Descr. Pap. Pl. 9 (1890) 60; Wars. Bot. Jahrb. 13 (1891) 396; Koorpb. Minah. (1898) 488; Merr. Philip. J. Sc. 3 (1908) Bot. 84. — Panax obtusum BL. Bijdr. (1826) 880; DC. Prod. 4 (1830) 254. — Aralia tripinnata BLANCO, FI. Filip. (1837) 223, cf. MERR. Sp. Blanc. (1918) 295. — Nothopanax fruticosum (L.) Mia. Pl. Jungh. 3 (1855) 425; Fl. Ind. Bat. 1, 1 (1856) 765; SEEM. FI. Vit. (1866) 114, 115; J. Bot. 4 (1866) 363; MeErRR. Fl. Manila (1912) 358, incl. var. plumatum (Hort.) MERR. et var. victoriae (Hort.) 7. P. javanica Me_rr.; Int. Rumph. (1917) 410; Sp. Blanc. (1918) 295; HARMS, Bot. Jahrb. 56 (1921) 412; Merr. En. Philip. 3 (1923) 233; HEYNE, Nutt. Pl. (1927) 1209; OcuseE & BAKH. Veg. D.E.I. (1931) 64, f. 37; CorRNER, Ways. Trees (1940). 150. — Nothopanax obtusum (BL.) Mia. FI. Ind. Bat. 1, 1 (1856) 766; SEEM. FI. Vit. (1866) 114. — P. obtusa (BL.) HARMS in E. & P. Nat. Pfi. Fam. 3, 8 (1894) 45, nom. illeg., non BLANCO, 1837, quae est Schefflera odorata (BLANCO) MerRR. & ROLFE; Koorp. Exk. Fl. Java 2 (1912) 716; Atlas 4 (1916) t. 679 S; BACK. & BAKH. f. Fl. Java 2 (1965) 168. Glabrous shrub or small tree, up to 5 m, with branches bearing spirally arranged leaves towards their ends. Leaves tripinnate, variable in size to c. 75 cm long; petiole up to c. 25 cm, with a sheath- ing base to 5 cm long; pinnae to c. 25 cm long; leaflets shortly petiolate, very variable in shape and size, oblong or linear-lanceolate, or 1-12 by '/,- 4 cm, more or less deeply serrate or lobed, or irregularly pinnatisect, apex attenuate, acuminate or rounded, base cuneate, truncate or subcordate, midrib and lateral veins evident. Inflorescence a diffuse panicle; primary axis up to c. 60 cm with 74 FLORA MALESIANA [ser. I, vol. 9} secondary branches mostly in verticils at intervals along its length; secondary branches up to 30 cm bearing umbellules in an irregularly branched system towards their extremities; umbellules with 12-20 flowers on pedicels c. 3 mm long. Calyx a minute rim. Petals 5, 2 mm long. Stamens 5. Ovary turbinate, c. 1 mm high, 2—3(—4)-celled; styles at first erect, later spreading. Fruit subglobose, fleshy, c. 5S mm @ when dry. Distr. Native country not properly known, cultivated throughout the region, and in other parts of the Indo-Pacific tropics. Ecol. Grown at low and moderate altitudes (c. 1000 m) as an ornamental or hedge shrub and for culinary use. Vern. (from various sources). Sumatra: orang aring, Medan, kédongdong mekka, Palembang; Malaya: daun girang, siku kluang, M; Java: kédongdong alus, k. batur, k. laut, M, imba, kédongdong laki, k. tjina, randa nunut, S, tjakar kutjung, t. tjikri, J, kadongdung, k. laut, k. petédhan, Md; Celebes: boka ula risé, kéndém rinték, Minahasa, Alfur lang., bombu, Makassar; Philip- pines: papua, Tag., Bik., bani, makan, Bik.; Moluccas: daun papeda papua, pagar pagar, Ambon, guarbati, tampusong, Ternate. The name kedongdong belongs properly to species of the fruit tree genus Spondias, but is sometimes also applied to trees of other families with pinnate leaves; papua means ‘curly’. Notes. The foliage of this plant is extremely variable in size and form. Typical plants are figured by Koorpers (1916, /.c.) and by OCHSE & BAK- HUIZEN VAN DEN BRINK (i.c.). These may be readily identified by the intricately compound leaf. The individual leaflets are characteristically ovate-lanceolate and serrate to deeply pinnatifid. However, broader leaflets with simpler outlines are not uncommon. The more rotund, blunter leaflets of P. obtusa are considered here to be an extreme form of this species (the type is bipinnate). Fig. 29. Species density of Polyscias J. R. & G. FORSTER in Malesia; above the hyphen the number of endemic species, below it the non-endemics. Only the native species. Occasional specimens have leaves so much reduced that they are simply pinnate or unifoliolate when they approach forms of 6. P. cumingiana. A number of names have been applied to horti- cultural forms belonging to this section of the genus (cf. BAILEY, Rhodora 18, 1916, 153), but the interrelations of these will be understood only after intensive biosystematic study of the many cultivars. 2. Polyscias macgillivrayi (SEEM.) HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; PHiLieson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 14; Blumea 24 (1978) 169. — Nothopanax macgillivrayi SEEM. FI. Vit. (1866) 114, nomen. — Panax macgillivraei (SEEM.) BrH. FI. Austr. 3 (1866) 382. — P. grandi- folia VOLKENS, Bot. Jahrb. 31 (1901) 471; KANEHIRA, En. Micron. Pl. (1935) 385. — Tieghe- mopanax macgillivrayi VIGUIER, Bull. Soc. Bot. Fr. 52 (1905) 313. — Polyscias sp. C. T. WHITE, J. Arn. Arb. 10 (1929) 255. Glabrous shrub or small tree, up to 15 m, with few branches bearing terminal clusters of large leaves. Leaves imparipinnate, up to 1 m (or more) long, uppermost leaves smaller; petiole c. 15 cm, with a sheathing base extending for 6-10 cm along the petiole; petiolules c. 10-15 mm; lamina oblong often broader near the base (lower and the terminal pinnules more ovate) c. 20-25 by 8-10 cm, slightly succulent when fresh, margin entire, revolute or occasionally minutely dentate, apex rounded and shortly apiculate, base rounded, truncate or sub- cordate, midrib and widely spaced laterals promi- nent. Inflorescence a large panicle; primary axis stout, rather short (c. 3-10 cm), often bearing reduced leaves or cataphylls basally, and a few lateral inflorescence branches, terminating in an umbel of long diffusely branched rays; lateral branches and rays c. 40-50 cm, with secondary branches c. 6-12 cm borne singly or in subverticils along their length; secondary branches with numer- ous tertiary branches bearing lateral and terminal umbellules (or the branching may be of a high order); umbellules with c. 6-12 flowers on delicate pedicels c. 3 mm long. Calyx a minute rim. Petals 5, 2'/, mm long. Stamens 5, anthers oblong 11/, mm long, filaments delicate, 11/, mm long. Ovary turbinate in bud, 1!/, mm long, rapidly becoming rotund, compressed with prominent veins, 2-celled; styles 2, divergent. Fruit fleshy, black, compressed, c. 5 by 6 mm; styles persistent, recurved. Distr. Micronesia, Solomon Is., Queensland; in Malesia: New Guinea (Papua, Eastern and Milne Bay Distr.; Territory of New Guinea, Morobe Distr., Musi I.), also in the D’Entrecasteaux and Trobriand Is., New Britain, and Louisiades. Ecol. Strand vegetation and littoral rain-forest, often behind mangrove. Vern. Aikove, Mimific lang., ane, D’Entre- casteaux Is., gambou, Musi I., Morobe Distr., nakaigwoo, Trobriand Is., raumonas, Onjob lang. 1979] 3. Polyscias verticillata STONE, J. Arn. Arb. 47 (1966) 272, f. 1; PHILIPSON, Blumea 24 (1978) 169. Glabrous, small tree to 7 m with few branches bearing spirally arranged leaves towards their ends. Leaves imparipinnate, up to 1 m long; petiole c. 24-30 cm, terete with a sheathing base c. 6-7 cm long; petiole 5-20 mm, leaflets oblong often broader near the base, c. 16-27 by 5-13 cm, margin entire, slightly revolute or sparsely denticulate, apex acuminate, base subcordate or truncate, often oblique, midrib and lateral veins evident. Jnflores- cence a large panicle; primary axis stout, rather short, with broad cataphylls, bearing several long radiating secondary branches in a sub-umbel; secondary branches c. 50 cm, with numerous tertiary branches borne in well defined verticils and in a terminal umbel, bracts triangular c. 8 mm long, caducous; tertiary branches c. 4-7 cm with small bracts near the middle; umbellules with c. 10-15 flowers, on pedicels 1-4 mm long. Calyx a minute rim. Petals 5, 21/; mm long. Stamens 5, anthers oblong, c. 1-1'/, mm long. Ovary turbinate, c. 1 mm long, 2-celled; styles 2, at first erect, later recurved. Fruit globose, fleshy, black, compressed, c. 4 by 7 mm when dry; styles persistent. Distr. Solomon Is.; in Malesia: New Guinea (New Britain, New Ireland, New Hanover, Admiralty Is., and Bagabag I.). Ecol. Usually near the beach or in lowland forest, to 140 m. Uses. The young foliage is eaten fresh or boiled. Vern. Babagula, valagur, Kuana dial., /a bara kiku, Nakanai, tauwol, Gasmata, palu, Vairamana, vela vela, Pomio. Note. Cut surfaces produce a sweet scented sticky exudate. The petals are violet within, the fruits purple-black. 4. Polyscias scutellaria (BURM. f.) Foss. Un. Hawaii Occ. Pap. 46 (1948) 9; Stone, Taxon 14 (1965) 284; PHimipson, Blumea 24 (1978) 169. — Scutellaria prima RumMPH. Herb. Amb. 4: 75, t. 31. — Scutellaria secunda latifolia RumpPH. lI.c. 76. — Crassula scutellaria BurM. f. Fl. Ind. (1768) 78. — Aralia cochleata LAMK, Encycl. 1 (1783) 224. — Panax scutellarioides REINW. ex BL. Bijdr. (1826) 880; SPAN. Linnaea 15 (1841) 208. — Panax cochleatum (LAMK) DC. Prod. 4 (1830) 253; BorrRL. Hand. 1 (1890) 647. — Panax conchifolium Roxs. Fl. Ind. ed. Carey 2 (1832) 77. — Notho- panax cochleatum (LAMK) Mia. PI. Jungh. 3 (1855) 425; Fl. Ind. Bat. 1, 1 (1856) 766; SEEM. J. Bot. 4 (1866) 296; FI. Vit. (1866) 116; Koorp. Minah. (1898) 490; Exk. Fl. Java 2 (1912) 717; Atlas 4 (1916) f. 697. — Nothopanax tricochleatum Mia. Fl. Ind. Bat. Suppl. (1860) 135, 340; Merr. Int. Rumph. (1917) 409; En. Philip. 3 (1923) 234; OcHsE & BAKH. Veg. D.E.I. (1931) 69, f. 39A. — Panax rumphii Hassk. Abh. Naturf. Ges. Halle 9 (1866) 220. — Nothopanax scutellarium (BURM. f.) Me_rr. Int. Rumph. (1917) 409; En. Philip. 3 ARALIACEAE—I (Philipson) obs 1923) 234; HEYNE, Nutt. Pl. (1927) 1209; OcHsE & BAKH. Veg. D.E.I. (1931) 67, f. 39; CoRNER, Ways. Trees (1940) 156; Back. & BAKH. f. Fl. Java 2 (1965) 169. — P. tricochleata (MiQ.) Foss. Phyto- logia 5 (1955) 290. Glabrous shrub or small tree, up to 6 m, with branches bearing spirally arranged leaves towards their ends. Leaves simple or trifoliolate, variable in size; petiole often c. 6 cm, but as long as 28 cm, with a sheathing base 1-6 cm long; lamina rotund or reniform, often c. 8 cm @ but as wide as 28 cm, margin usually serrate or becoming sub-lobed towards apex, in larger leaves the serrations often inconspicuous, apex rounded, base + cuneate, midrib and lateral veins evident. Inflorescence a diffuse panicle; primary axis usually long (up to 1 m) with secondary branches (mostly in verticils, at intervals along its length; secondary branches 15-30 cm, bearing umbellules in an irregularly branched system towards their extremities; umbellules with c. 8-16 flowers, on pedicels c. 3 mm long. Calyx a minute rim. Petals 4-5, 2 mm long. Stamens 4-5, anthers oblong, 1 mm long. Ovary turbinate, c. 1 mm high, (2—)3-4-celled, styles at first erect, later recurved. Fruit subglobose, fleshy, c. 5mm @ when dry. Distr. Native country not properly known, possibly East Malesia; cultivated throughout the region, extending through the tropical Pacific. Ecol. Grown at low and moderate altitudes (800 m) as an ornamental or hedge shrub. Uses. The foliage is aromatic and is used to furnish perfume. Forms with variegated foliage are in cultivation. Used medicinally as a diuretic, against breast cancer, and to prevent baldness (HEYNE, /.c.). Also for culinary purpose (OCHSE & BaAKH. /.c. 67). Vern. Cf. HEYNE: memangkokan, (pohon) mangkok, Mal. (after the dish(= mangkok)-shaped leaves which are in the Moluccas sometimes used as dishes); Java: godong mangkokan, M, mama- mékan, S, puring mangkok, Md; Lesser Sunda Is.: lanido, ndalido, ndari, ramido, Roti; Celebes: (daun) mangko, tuwo mangku, M, bobohang, boku ula, kéndém wéwéné, woworan, Manado, angko mangko, Bug. & Mak.; Philippines: salapiin, Mindanao; Moluccas: daun koin, d. papéda, M, Ambon, ai /aun niwél, ai lohoi, Alf., Ambon, goma ma tari, Gal., Halmaheira, sawoko, Loda, Halmah.., rau paroro, Ternate. Note. This species is usually readily distin- guished by its simple, orbicular, saucer-shaped leaves, but plants with some or all of their leaves trifoliolate occur. STONE (Micronesica 2, 1965, 51) advanced evidence for uniting these trifoliolate plants with P. pinnata (= 6. P. cumingiana) while retaining the unifoliolate plants as a distinct species. Possibly all are forms of one polymorphous species complex, but here it is considered convenient to adhere to the view which unites all forms having 76 FLORA MALESIANA [ser. I, vol. 91 orbicular usually bowl-shaped leaves under the concept P. scutellaria. 5. Polyscias guilfoylei (COGN. & MARCHE) L. H. BAILEY, Rhodora 18 (1916) 153; STONE, Micro- nesica 2 (1965) 57; PHILIPSON, Blumea 24 (1978) 169. — Aralia guilfoylei [BULL, Cat. (1873)] CoGn. & MARCHE, PI. Ornam. 2 (1874) t. 58. — Notho- panax guilfoylei (COGN. & MARCHE) MErRR. Philip. J. Sc. 7 (1912) Bot. 242; Fl. Manila (1912) 357; En. Philip. 3 (1923) 234. Glabrous shrub to 3 m high, with few branches, bearing spirally arranged leaves towards their ends. Leaves imparipinnate, with 3-4 pairs of leaflets, c. 60 cm long; petiole c. 18 cm, terete, with a sheathing base 3-4 cm long; petiolules c. 11/,— 2'/, cm; leaflets rotund, oblong or rhomboidal, c. 10-14 by 6-7 cm, rather thin and flaccid, some- times rugose, margin sharply serrate to broadly cuneate (often decurrent on the petiole), often oblique, midrib and lateral veins evident. Inflores- cence a diffuse panicle; primary axis short (c. 3- 4 cm) with one or few lateral flowering branches, terminating in an umbel of long much-branched rays; lateral branches and rays c. 40-50 cm, with secondary branches (c. 5-8 cm) singly or mainly in subverticils along their length and in a terminal umbel; secondary branches with one or more pairs of small bracts and ending in umbellules with a few lateral tertiary branches also ending in umbel- lules; umbellules with c. 8-12 flowers on pedicels c. 8-10 mm long. Calyx a minute rim. Petals 5, 2'/, mm long. Stamens 5, anthers oblong, 1'/, mm long; filaments 2 mm. Ovary turbinate, c. 1 mm high, usually 3-celled, styles at first erect, but soon elongating and recurved. Fruit fleshy, globose, c. 4 by 5 mm. Distr. Native country unknown, possibly from East Malesia. Cultivated throughout the region, but less commonly than other cultivated species of this section. General throughout the tropical Pacific. Ecol. Usually grown as a hedge plant. Flowers only when left untrimmed or when growing as an escape from cultivation. Note. Recognizable by the shape, texture and serrations of the leaflets, which are usually varie- gated with whitish or yellowish blotches near the margins. The flowers are brown in bud but yellow- green when open. 6. Polyscias cumingiana (PRESL) F.-VILL. Nov. App. (1880) 102; PHiLirson, Blumea 24 (1978) 169. — Scutellaria secunda angustifolia RUMPH. Herb. Amb. 4: 76, t. 32. — Panax pinnatum LAMK, Encycl. 2 (1788) 715, non P. pinnata J. R. & G. Forster, 1776; DC. Prod. 4 (1830) 254; SPAN. Linnaea 15 (1841) 208; Mio. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 15; BoerL. Handl. 1 (1890) 647; Koorpb. Minah. (1898) 15. — Panax secundum SCHULT. Syst. 6 (1820) 215, nom. superft. illeg. — Panax bandanense Zipp. ex SPAN. Linnaea 15 (1841) 208, nom. inval. in synon. — Paratropia cumingiana PRESL, Epim. (1851) 250. — Notho- panax pinnatum (LAMK) Mia. Bonplandia 4 (1856) 139; Mia. FI. Ind. Bat. 1, 1 (1856) 766; Merr. Int. Rumph. (1917) 409; En. Born. (1921) 458; HEYNE, Nutt. Pl. (1927) 1209; OCHSE & BAKH. Veg. D.E.I. (1931) 67, f. 38; CORNER, Ways. Trees (1940) 156. — Nothopanax cumingii (PRESL) SEEM. Fi. Vit. (1865) 114. — Aralia filicifolia C. Moore, Ill. Hortic. 23 (1876) 72, t. 240. — Arthrophyllum pinnatum (LAMK) CLARKE, FI. Br. Ind. 2 (1879) 734, pro basionym. — Panax cumingiana (PRESL) ROLFE, J. Linn. Soc. Bot. 21 (1884) 310; VipAL, Phan. Cuming. (1885) 117. — ? Aralia naumannii E. MarCHAL, Bot. Jahrb. 7 (1886) 469. — Panax crispatum BULL, Cat. (1888) 9. — Panax ornatum BULL, /.c. — P. cumingii (PRESL) HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45. — P. rumphiana Harns, /.c. 45; in K. Sch. & Laut. Fl. Schutzgeb. (1900) 485; Baku. f. & OoststrR. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 13; BAcK. & BAKH. f. Fl. Java 2 (1965) 167. — Nothopanax crispatum (BULL) MerRR. Philip. J. Sc. 7 (1912) Bot. 241. — Nothopanax ornatum (BULL) MERR. l.c. — P. filicifolia (C. Moore) BAILEY, Rhodora 18 (1916) 153. — P. sorongensis Gipss, Arfak (1917) 216. — Anomopanax cumingianus (PRESL) MERR. Philip. J. Sc. 17 (1920) 300; En. Philip. 3 (1923) 236. Glabrous shrub or small tree, to c. 4 m, un- branched or with few branches bearing spirally arranged leaves towards their ends. Leaves impari- pinnate, up to 100 cm; petiole to 20 cm, with a sheathing base c. 5-6 cm long; petiolules to 3 cm; leaflets ovate-oblong or elliptic, 10-30 by 2-13 cm, apex attenuated or acuminate, base rotund, trun- cate or broadly cuneate, often oblique, margin entire or minutely and distantly dentate (sometimes pinnatilobed or almost pinnatipartite); midrib and lateral veins evident. Inflorescence a large terminal panicle, or with flowering branches also in the axils of the upper leaves; primary axis up to 140 cm with secondary branches mostly in verticils at intervals along its length; secondary branches up to 120 cm, bearing umbellules in an irregularly branched system towards their extremities, peduncles of the umbellules with one or more pairs of small bracts; umbellules with c. 10-20 flowers; pedicels 4-8 mm. Calyx a minute rim with 4-5(-6) teeth. Petals 4~5(-6), 2!/,-31/, mm long. Stamens 4—-5(-6), 2 mm long. Ovary turbinate, c. 2 mm long, 2-5-celled; styles at first erect, later spreading. Fruit subglobose, fleshy, 3-S mm @ when dry. Distr. Cultivated throughout the region, and also apparently growing as part of the indigenous vegetation. Ecol. Rain-forest and secondary growth, usually at low altitude but ascending to 1700 m, native range not very clear. Uses. A common ornamental shrub, especially 1979] the cultivars with dissected foliage. Also for culinary use. Vern. (from various sources). Celebes: daun grisik, d. mangko, d. papéda pandang, M, Manado; Philippines: bani, Luzon, Albay; Moluccas: kéndém, Ternate, gurabati, M; papua, Jappen I.; New Britain: awalagu, Gazelle Pen. OcHSE & BAKHUIZEN VAN DEN BRINK record that the vernacular names applied to P. fruticosum also include P. cumingianum. Notes. A complex of forms requiring intensive biosystematic study. I adhere to the view that the Indo-Malayan material is specifically distinct from the Polynesian Polyscias pinnata J. R. & G. Forster; cf. HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; Puiieson, Bull. Br. Mus. Nat. Hist. Bot. 1 (1951) 9, but some cultivated forms included here may have been derived from the Pacific rather than the Indo-Malayan species. Existing herbarium material, though voluminous, is inadequate to determine whether the complex includes species with more limited ranges. As in 1. P. fruticosa, cultivated forms with dis- sected foliage occur. See also under 4. P. scutellaria for a discussion of the relationship with that species. I have not seen the type of Aralia naumannii MARCHAL which, from the description, I assume to be a synonym. 7. Polyscias javanica K. & V. Bijdr. 7 (1900) 13; Koorp. Atlas 4 (1916) f. 679 A-R; Baku. f. & Ooststr. in Back. Bekn. Fl. Java (em. ed.) 7 (1948) fam. 159, p. 12; BAcK. & BAKH. f. Fl. Java 2 (1965) 167; PHILIpsoN, Blumea 24 (1978) 169. ARALIACEAE—I (Philipson) 77 Glabrous shrub or sparsely branched small tree, bearing spirally arranged leaves towards the ends of the branches. Leaves imparipinnate, c. 70 cm long; petiole c. 17 cm, with a wide membranous sheath- ing base (c. 7 cm long); petiolules c. 1 cm; leaflets ovate to elliptic-oblong, up to 22 by 8 cm, mem- branous, apex attenuated, base rounded to broadly cuneate, often oblique, margin entire or minutely distantly dentate, slightly revolute, midrib and lateral veins prominent. Inflorescence a panicle; rachis 40 cm, with caducous bracts (reduced leaves or leaf-sheaths), bearing few secondary branches and terminating in a compound umbel; secondary branches with one or more verticils of tertiary branches; peduncles of the umbellules with one obscure bract or none; umbellule of c. 10-20 flowers, pedicels c. 5 mm. Calyx an undulate rim, indistinctly S-dentate. Petals 5, 2!/,-3 mm long. Stamens 5, with very short filaments. Ovary turbi- nate, 1'/, mm high, 5-celled, styles at first erect, later recurved. Fruit a globose or obovoid fleshy drupe (prominently 5-ribbed when dry) c. 7 by 5 mm; persistent styles united below, strongly reflexed above. Distr. Malesia: East Java, Lesser Sunda Is. (Lombok, Sumbawa, Sumba). Ecol. In forest up to 1650 m. Uses. The root is said to be made into a face powder in Sumbawa. Vern. Lesser Sunda Is.: pulosari, Sumbawa, amdari, Sumba. Note. The vegetative parts are rather similar to those of 6. P. cumingiana, but the smaller and simpler inflorescence is distinctive. 2. Section Gelibia (HUTCH.) PHILIPSON, Blumea 24 (1978) 169. — Gelibia Hutcu. Gen. FI. Pl. 2 (1967) 57. Tree with bipinnate leaves. Flowers arranged racemosely; style arms free and divergent after anthesis. Distr. In New South Wales, Queensland, and Malesia (New Guinea) | sp. 8. Polyscias elegans (C. Moore & F.v.M.) HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45; J. Arn. Arb. 20 (1939) 323; PHitipson, Blumea 24 (1978) 169. — Panax elegans C. Moore & F.v.M. Trans. Phil. Inst. Vict. 2 (1858) 68; Bru. FI. Austr. 3 (1866) 383; C. Moore, Handb. FI. N.S.W. (1893) 215. — Nothopanax elegans (C. Moore & F.v.M.) Seem. FI. Vit. (1866) 114; J. Bot. 4 (1866) 294. — Tieghemopanax elegans (C. Moore & F.v.M.) VIGUIER, Bull. Soc. Bot. Fr. 52 (1905) 308. — P. branderhorstii HARMS, Nova Guinea 8 (1910) 274; Bot. Jahrb. 56 (1920) 42; J. Arn. Arb. 20 (1939) 323. — Gelibia branderhorstii (HARMS) Hutcu. Gen. FI. Pl. 2 (1967) 57. — Gelibia elegans (C. Moore & F.v.M.) Hutcn. /.c. 58 Tree to 20 m high, with a spreading crown of thick branches bearing terminal clusters of large leaves, young parts with fawn to grey scurfy tomentum. Leaves bipinnate, becoming glabrous, up to 110 by 50 cm, or larger; petiole c. 13 cm, with a slightly clasping base; petiolules up to c. 1/, cm; leaflets ovate or elliptic, c. 6 by 3 cm, chartaceous to somewhat leathery, margin entire and slightly revolute, apex with an obtuse apiculum, base cuneate, midrib prominent, lateral veins rather obscure. Inflorescence a large panicle, 78 FLORA MALESIANA [ser. I, vol. 91 tomentum persistent especially on the pedicels; primary axis stout, c. 30 cm, bearing secondary axes along its length and in a terminal subumbel- late cluster, bracts caducous; secondary axes c. 20-30 cm, bearing tertiary axes (c. 10 cm) along their length; flowers borne racemosely along the tertiary axes on pedicels 1-2 mm long. Calyx a minute rim. Petals 5, oblong, rather fleshy, spreading at anthesis and soon falling. Stamens 5, 2 mm long on short filaments, soon falling. Ovary scurfy, at first turbinate, globose at anthesis, c. 4 mm high, 2-celled, disk rising to the 2 styles; style arms at first short and pressed together, elongating and diverging after the anthers have fallen. Fruit leathery, globose, compressed, c. 5 by 5 mm and irregularly ribbed when dry. Distr. New South Wales and Queensland; in Malesia: New Guinea (West Irian, Southern Distr. ; Papua, Western & Central Distr.). Ecol. Gallery and second growth forest in (seasonal) monsoon regions, at low altitudes, up to 300 m. Notes. The racemose flower arrangement has sometimes been considered sufficiently distinctive to justify the accommodation of this species in a separate genus. The bark is described as grey or patched grey- brown; the wood as soft and pale; and the cut surfaces having a clear sticky exudate which has a pleasant fragrance. The inflorescence branches are purplish, the petals maroon, the anthers cream to yellow on reddish filaments, and the fruit purplish and smooth. 3. Section Eupteron (MiQ.) PHILIPSON, Blumea 24 (1978) 170. — Eupteron Mia. Pl. Jungh. 3 (1855) 420; Fl. Ind. Bat. 1, 1 (1856) 762. Trees or shrubs with imparipinnate or bipinnate leaves. Flowers in umbellules or capitula; style arms divergent at least in fruit. Distr. In Malesia 6 spp., but the section probably includes the majority of the genus in other parts of the Old World tropics. Note. With the exception of 14. P. philipsonii, the Malesian spp. of sect. Eupteron have umbellules or capitula arranged racemosely along the main inflorescence branches. The type species of the section (P. nodosa) has its flowers in capitula, but this is not considered a sufficient difference to require a separate section. KEY TO THE SPECIES 1. Flowers in capitula 1. Flowers in umbellules. oer ee ie Neya Ge Gtolmweghcie? Veliuo chemise (ai 2. Umbellules arranged racemosely along the principal rays. 3. Leaf margins crenate. 4. Leaflets oblong (Luzon) 4. Leaflets lanceolate (New Guinea) . ' i i Oey CO Ck © “ey Yo, ves “elauloumninn ye mea 3. Leaf margins entire (rarely with few minute dentations). 5. Leaflets ovate (Sabah, Palawan) ..... 5. Leaflets elliptic (New Guinea) 2. Umbellules in a short corymbose compound umbel i \ Y Fp g g Sigs Vz p Pee § | ‘N Zp & q CJ , jo y Fig. 39. Acanthopanax malayanus HENDERSON. a. Habit, x 1/,, b. flower, < 7, c. fruit, x 4 (a, b DE WILDE c.s. 16725, c DE WILDE c.s. 15925). Drawn by W. R. PHILIPSON. 1979) lets rather stout (c. 5 mm @); bark greyish brown, glabrous, smooth; buds with small brown cata- phylls. Leaves crowded at the ends of the branch- lets; petiole terete, striate, glabrous, with a slightly dilated base, up to 12'/, cm; leaflets 3—5; petiolules up to 7 mm, articulated with the petiole, channelled above; blade up to 17 by 5'/, cm, ovate-lanceolate to narrowly elliptic, apex acute, base cuneate or one side truncate, chartaceous, the principal veins pinnately arranged and conspicuous, the minor veins forming a dense reticulation, margin with numerous spinulose teeth. Jnflorescence a terminal, sessile, compound umbel, often on short lateral shoots, apparently dioecious; primary rays c. 3-7, slender, terete, glabrous, striate, c. 4-8 cm; secondary rays (pedicels) of male flowers numer- ous, slender, 1'/,-2 cm, of female flowers stouter. Calyx a rim with 4-5 minute teeth. Petals 4-5, ligulate, c. 2'/, by 11/. mm long. Stamens 4-5, filaments c. 23/, mm long. Ovary turbinate, 3—4- celled; styles 2, at first connate, but becoming free to about the middle. Fruit an oblate spheroidal ARALIACEAE—I (Philipson) 105 drupe, c. 10 by 8 mm when dry, with a small stylo- podium bearing a persistent bifid style; endocarp cartilagineous. Endosperm surface strongly undu- late. Distr. Malesia: Malay Peninsula (Pahang; Cameron Highlands), N. Sumatra (Gajo Lands). Ecol. Montane rain-forest and mossy forest, 1400-2600 m. Vern. Berlaki, Sakai lang. Notes. The interpretation of this species as having sexually dimorphic inflorescences cannot be proved with the available material. This appears to consist of twigs bearing either inflorescences of male flowers, or infructescences. Appearances suggest that the putative male flowers have rudi- mentary ovaries. There is no evidence whether the fruiting flowers had produced pollen. The tree is said to be conspicuous when flowering by reason of its delicate feathery foliage, which is pale green with a reddish tinge, the petioles and inflorescence also being reddish. Excluded Meryta colorata F. M. BAILEY, Queensl. Agric. J. 3 (1898) 283; HARms, Bot. Jahrb. 56 (1920) 384. The type specimen of this species cannot be located, but the original description does not seem to relate to a member of the Araliaceae, and certainly not to a Meryta. Panax ? anisum DC. Prod. 4 (1830) 254. — Anisum moluccanum RumpPH. Herb. Amb. 2: 131, t. 42. — Nothopanax ? anisum Mia. FI. Ind. Bat. 1, 1 (1856) 766; SEEM. FI. Vit. (1866) 114. — Polyscias anisum HARMS in E. & P. Nat. Pfl. Fam. 3, 8 (1894) 45. According to MERRILL (Int. Rumph. 1917, 289) these names are all based on the Rumphian description and plate and must refer to some species of Fagara, possibly F. avicennae LAMK = Zanthoxylum avicennae (LAMk) DC. (Rutaceae). aed) ; une Gee ike pwoR! Carat | ies ith ak” etd ath shee - 2 ( Galpi,s ota : Sf — J = 7 ca | ey on } —" ; ee oer 4 oe , . sicread ~ e 5 a A ' al 7 ee fart e >= 2 W ee Cn ¥ ive 29 = + 7 oat J . af i : © nm a) — i t : é we a (a2 @p af <« =) Se H , ».D or ‘ wy: ae ity z / = = ip Ss i e tin? inj \agh CYPERACEAE—IDI (J. H. Kernt & H. P. Nooteboom, Leyden) 28. CAREX LINNE, Gen. Pl. ed. 5 (1754) 420; Sp. Pl. (1753) 972; ENDL. Gen. (1836) 110; KuNnrTH, En. 2 (1837) 368; Steup. Syn. 2 (1855) 182; Mig. FI. Ind. Bat. 3 (1856) 346; Boort, Illustr. Genus Carex I-IV (1858-67); Boeck. Linnaea 39 (1875) 14; ibid. 40 (1876) 327; ibid. 41 (1877) 145; B. & H. Gen. Pl. 3 (1883) 1073; Pax in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 122; CLarkg, FI. Br. Ind. 6 (1894) 699; J. Linn. Soc. Bot. 37 (1904) 1; Philip. J. Sc. 2 (1907) Bot. 107; Kix. Pfl. R. Heft 38 (1909) 67; Philip. J. Sc. 6 (1911) Bot. 57; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 232; NELMES, Kew Bull. (1946) 5; Reinwardtia | (1951) 221; ibid. 2 (1954) 373; RAYM. Mém. Jard. Bot. Montréal n. 53 (1959) 17; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 149; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 487. — Fig. 118-135. Perennial herbs with tufted or creeping rhizome, monoecious, rarely dioecious. Stems arising centrally or laterally, erect or obliquely erect, mostly triquetrous or trigonous, rarely subterete, solid or sometimes hollow, often clothed at the base by persistent leaf-sheaths or their fibrous remains. Leaves tristichous, usually narrowly linear, sheathing at the base, with a ligule at the junction of blade and sheath, rarely lanceolate or elliptic with a more or less distinct petiole and eligulate, mostly basal and subbasal, 0-several higher on the stem, the lower ones often reduced to bladeless sheaths; sheaths of the stem-leaves and bracts closed. Inflorescence paniculiform, racemiform or spiciform, more rarely reduced to a single spikelet. Spikelets 1-very numerous, terete, sessile or peduncled, few- to many-flowered, wholly male, wholly female, or bisexual (androgynous when male flowers above, gynaecandrous when female flowers above). Bracts foliaceous or glume-like, often sheathing, sometimes wanting. Base of the branches of the inflorescence usually with a utriculiform or ocreiform bracteole (cladoprophyllum) surrounding it. Flowers unisexual, naked, solitary in the axils of the spirally arranged glumes; male flowers consisting of 3 free or rarely more or less connate stamens; anthers linear; female flowers consisting of a single pistil enclosed in a bottle-shaped prophyll (utricle, perigynium). Style either continuous with the ovary and persistent, or articulated with it and deciduous, straight or flexuous, often incrassate at the base; stigmas 2 or 3, protruding through the small terminal orifice of the utricle. Vestigial rachilla (see Uncinia) rarely present. Utricles membranous, chartaceous, or coriaceous, bicarinate, sometimes winged, sessile or stipitate, beakless to strongly beaked, nerveless, nerved, or ribbed, glabrous, or pubescent or hispid, papillose or puncticulate or smooth, sometimes spongy at the base; beak truncate, obliquely cleft, bidentate, or bifurcate at the top. Nut trigonous (when stigmas 3), or lenticu- lar (plano-convex or biconvex; when stigmas 2), enclosed within the utricle. Distr. A large genus with 600 to 1000 spp., the majority of them outside the tropics. However, the most primitive section, Vigneastra with a compound, paniculate inflorescence and andro- gynous spikelets, occurs mainly in the tropics of the Old World, from sea-level up to 3000 m. This section is represented in Malesia with 11 spp., and is there by far the largest section. Subg. Carex is, besides with the 11 spp. of sect. Vigneastra, represented in Malesia by 25 sections and 49 spp., subg. Vignea with 6 sections and 6 spp. (1) Continued from volume 7, part 3 (1974) p. 753. (107) 108 FLORA MALESIANA [ser. I, vol. 9} Of the 66 spp. in all, 12 spp. are endemic and mostly confined to one island. Of the other 54 spp. the majority occur also in the northern hemisphere, either widely or only in SE. and/or E. Asia. Of these, 25 spp. extend without noticeable disjunctions into Malesia, either to its western part or further eastward. Another 4 of the E. Asian spp. occur in Formosa and in Malesia only in the Philippines (mostly only in Luzon), viz 12. C. satzumensis, 30. C. rhynchachaenium, 32. C. dolichostachya, and 33. C. formosensis. Among the Asian spp. 4 show a distinct disjunction between SE. Asia and Java, viz 11. C. vesi- culosa, 14. C. helferi, 63. C. nubigena, and 58. C. longipes (which is also once found in Celebes and once in New Guinea); all are mountain species. A similar disjunction is found in the lowland species 10. C. stramentitia and 38. C. tricephala; this disjunction is understandable as both are bound to a distinct seasonal climate which shows a similar pattern (see VAN STEENIS, Reinwardtia 5, 1961, 420-429). Still larger disjunctions are shown by 5 E. Asian spp. which are in Malesia almost only found in New Guinea, Formosa being mostly the nearest station, viz 46. C. brachyathera, 47. C. finitima (also in N. Sumatra), 48. C. michauxiana, 56. C. bilateralis, and 61. C. duriuscula; all are mountain plants. Not a few Carex spp. occur in both the northern and southern hemisphere (Australia, often Tasmania and New Zealand) and are also found in the intervening tropical zone in Malesia. There are 11 of such species in all. Among these the following 7 are found in more than one island, viz 6. C. indica, 31. C. breviculmis, 39. C. maculata, 40. C. capillacea, 57. C. brunnea, 5. C. hors- fieldii, and 64. C. echinata, although the latter two are very rare. Four others are also transtropical but are in Malesia only found in New Guinea, and thus show almost the same disjunction between Formosa and New Guinea as mentioned above; these are 43. C. brownii, 49. C. pseudo- cyperus, 55. C. gaudichaudiana, and 66. C. curta. They occur in Malesia only in bogs and marshes on (high) mountains. The southern hemisphere has contributed very little to the Malesian Carices, as there is only one southern species from Australia, New Zealand, and New Caledonia, viz 62. C. appressa, which is found in Malesia, and then again only in New Guinea at very high altitude. Summarizing, one cannot withdraw oneself from concluding that only the presence of moun- tain bogs, marshes and grasslands gives the opportunity for the maintenance of Carex in the Malesian tropics. This would explain why the big mountainous island of New Guinea — where this habitat is most abundantly represented, much more than in any other island — is so rich in species. It is then also understandable that this is the place where relict areas could maintain their last stand in the tropical mountains of Malesia. Ecol. Most species prefer moist or wet localities, some are found in rain-forest, others in open habitats like in mountain marshes, grassland, heaths, etc. Only 22 spp. grow below 1000 m, 8 of them belonging to sect. Vigneastra, 13 are found below 500 m, 4 of them of sect. Vigneastra; 14 spp. occur exclusively above 2000 m. 57. C. brunnea, which is found from 80 m up to 3400 m, has the largest altitudinal range. The majority of the Carex spp. is growing on more or less acid soils. Three species of the lowland tropics occupy a different habitat, and are bound to a seasonal climate, viz 10. C. stra- mentitia, 38. C. tricephala, and 37. C. speciosa. The first two occur only in Java or Madura I., amongst others with the teak-forest; the third of these drought species occurs also in some local dry spots in E. Sumatra and SE. Borneo, and is obviously adapted to a slightly less severe dry season. Of the three mentioned species, C. tricephala and C. speciosa are confined to limestone, like one other species, 35. C. malaccensis, an endemic of N. Malaya (the Langkawi Is.). One species, 59. C. teinogyna, seems in Malesia to be confined to banks of swift streams and occupies a rheophytic habitat. Notes. The distinction of the sections is not always unanimously agreed on, and several species were put into different sections by different authors. Some sections are closely allied, the only difference being sometimes 2 versus 3 carpels. As this has great influence on the shape of the nut, it often is considered a very important character. In my opinion the difference is due to a minor genetic difference, and is taxonomically not very important. Besides, in some collections 2-carpelled and 3-carpelled fruits are found in the same inflorescence, e.g. in species of sect. 1979] Cy PERACEAE—II (Kern & Nooteboom) 109 Graciles, which has normally 2 carpels, and of sect. Oligostachyae (Decorae) which has normally 3 carpels (Noort.). Acknowledgement. The manuscript of Carex was for the greater part elaborated by Dr. J. H. KERN before he died in 1974. The obstacle for finishing it in time was his worry about the status cq. evaluation of a number of names in sect. Vigneastra, which also prohibited the finishing of the practical key to the species. Dr. H. P. NooTEBoom has done this evaluation and he made the final draft of the key, for which he is responsible. He is also responsible for the treatment offered here for several species, viz 3. C. cruciata, 18. C. perakensis, 19. C. turrita, 20. C. verticil- lata, and 31. C. breviculmis, and for some critical notes which have been marked ‘“‘Noor.” between brackets. The division in subgenera and sections is largely from Dr. KERN. The revision of Uncinia is entirely by Dr. NooTEBOom. (Ed.) Explanation of some terms. The term ‘stems’ refers to the flower-bearing stems, except when stated otherwise, whereas ‘inflorescence’ means the entire flower-bearing stem including all the spikelets. The terms used for the inflorescence (‘panicle’, ‘raceme’) indicate the disposition of the spikelets, not of the individual flowers. “Stems central’ means that the leaves are arranged around the flower-bearing stem (usually towards the base). ‘Stems lateral’ means that the leaves are placed on a vegetative shoot, and that the flower-bearing stems arise separately from the rhizome and bear few short-bladed leaves. The flowering stems can also be situated in the axils of the leaves. This situation is sometimes difficult to distinguish from ‘stems central’, because the leaves are often crowded in a basal rosette. The bracts are inserted on the stem and they are sheathing or not; they bear in their axils a spikelet or a partial inflorescence, the peduncles of which are often basally surrounded by a cladoprophyllum; this is a bracteole which is utriculiform or ocreiform. If the cladoprophyllum is ocreiform, it usually is hidden entirely in the sheath of the bract. Each flower is solitary in the axil of a glume; several flowers together form a spikelet which can be 4, 2, or bisexual; in the latter case the 3 flowers are usually placed only at the base or at the apex of a spikelet. Gynaecan- drous spikelets have 2 flowers above, 3 below, androgynous spikelets have ¢ flowers above and 2 below. The figures 118-123, depicting utricles and nuts of nearly all the species, and some glumes (fig. 132), are given to facilitate use of the keys. KEY TO SUBGENERA AND SECTIONS as represented in Malesia 1. Stigmas 3 and nut trigonous, or, when stigmas 2 and nut lenticular (plano-convex or biconvex) some or all of the spikelets peduncled or terminal spikelet ¢. Spikelets either dissimilar in appearance (the terminal one ¢ and some or all the lateral ones 9), or similar in appearance (all bisexual). Base of the peduncles of the lateral spikelets or of the branches of the inflorescence surrounded by a utriculiform or ocreiform bracteole (cladoprophyllum) ; inflorescence sometimes reduced to a single terminal spikelet I. SuBG. CAREX 2. Stigmas 3; nut trigonous. 3. Inflorescence consisting of more than one spikelet. 4. Lateral spikelets arising from a gaping, utriculiform, fertile (i.e. nut-bearing) cladoprophyll. Spikelets androgynous, up to 15 mm long. Bracts not sheathing. Stems 3-20 cm tall. Sp. 12 2. Sect. Japonicae 4. Lateral spikelets arising from a sterile cladoprophyll, or sessile without cladoprophyll. 5. Stems scapiform, surrounded at the base by spadiceous sheaths in place of leaves. Leaves wide, conduplicate-petioled below. Secondary panicles contracted, very dense. Utricles with very long linear beak and very oblique mouth. Sp.14............ . . 4. Sect. Mapaniifoliae 5. Otherwise. 6. Spikelets peduncled, in fascicles in the axils of at least some of the sheathing bracts, all androgy- nous or some wholly ¢ or 9. Utricles long-beaked. Spp.17-20 . . . . . 6. Sect. Oligostachyae 6. The peduncled or sessile spikelets or secondary panicles single or binate in the axils of the bracts. 7. Spikelets similar in appearance, androgynous (upper flowers 3, lower ones 9). 8. Lower bracts not or only shortly (some millimeters) sheathing. 9. Apex of nut truncate, !/,-3/, mm @, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. Spp. 27-30. . . 10. Sect. Lageniformes 110 FLORA MALESIANA [ser 1, vol, 9? 9. Apex of nut not truncate. ees base pyramidally thickened, often broader than apex of nut. S7p135—S Omen . 5 « ees seo sh joao) IDES ecommitadicales 8. Lower bracts long sheathing. 10. Flowering stems arising from the axils of the leaves which are crowded on a short stem and much longer than the flowering stems. Style flexuous. Nut with irregularly excavated sides, seated ona spongy, disk-like stipe. Sp.26 .. . . . . 9. Sect. Cryptostachyae 10. Flowering stems arising either from the centre of the leaves, or from lateral shoots bearing a few short-bladed leaves. 11. Spikelets small, up to 15(-20) mm long. 12. Stems central. Spikelets numerous, ovoid or oblong-ovoid, arranged in often diffuse, compound panicles. Bracts long. Spp.1-l1l1 .. . . . . 1. Sect. Vigneastra 12. Stems lateral. Spikelets few, globose-ovoid, in a simple panicle. Bracts shorter than 8 cm, the sheaths not included Sp: 21, 2 = = = 3 = 22 = <) -oeme Eee ecerSurculosae 11. Spikelets longer. 13. Utricles densely many-nerved. Spikelets 1—8(—14). 14. Utricles ellipsoid, short-beaked, more or less ciliate on the margins, spongy-thickened at the base. Spikelets 1-3(-4). Spp. 35-38... . . . . . 12. Sect. Radicales 14. Utricles narrowly ellipsoid, long-beaked, more or less hispidulous on the faces, not spongy-thickened at the base. Spikelets 2-8(-14). Spp. 17-20 . 6. Sect. Oligostachyae 13. Utricles distinctly but not densely nerved. Leaves with reddish sheaths splitting into fibres) Spikelets’ 6-numeroussSppsl5—l6) sess neue ee 5. Sect. Polystachyae 7. Spikelets dissimilar in appearance, unisexual (or rarely gynaecandrous), the terminal one or some upper ones 4, the remainder 9 (occasionally the 9 spikelets may have a few ¢ flowers, and the ¢ spikelets a few 9 flowers). Or most of the spikelets 2, shortly cylindrical, the ¢ spikelets lateral, 1-5 just below some of the terminal 2 spikelets. 15. Leaves not crowded in a basal rosette but inserted all over the stem, the upper ones merging into leafy bracts, the lower ones gradually decreasing in length and merging into bladeless sheaths. 16. Inflorescence a compound panicle consisting of 5-8 secondary panicles each containing numerous @ spikelets and 1-5 gd spikelets. Spikelets small, 4-8 mm long. Utricles glabrous, with short, !/, mm long beak. Leaves sparsely hairy beneath. Sp. 13 3. Sect. Hypolytroides 16. Inflorescence simple, racemose, consisting of 4-9 spikelets; terminal spikelet ¢ (sometimes a second, smaller one added), remaining spikelets 2, 11/,-4 cm long. Utricles densely hispid, with distinct, c. 1 mm long beak. Leaves glabrous. Sp. 50... . . . 21. Sect. Occlusae 15. Leaves crowded at the base of the stem, often also some higher on the stem. 17. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, hastiform, i.e. cylindric, 1/,—'/; mm long beak expanding into the annulate apex. Spp. 22-25 8. Sect. Rhomboidales 17. Nut without hastiform beak. 18. Apex of nut truncate, 3/,-1 mm wide, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. Spp. 27-30 10. Sect. Lageniformes 18. Apex of nut not truncate, at most 1/, mm wide. 19. Nut mitrate, i.e. contracted at the apex and then expanded into a discoid annulus. Utricles ovoid or ellipsoid. Spp. 31-34 ............. 1. Sect. Mitratae 19. Nut not mitrate. 20. Utricles densely papillose, erostrate or but shortly beaked. Sp. 39 13. Sect. Trachychlaenae 20. Utricles not papillose, distinctly beaked. 21. 2 Spikelets broadly ovoid or subglobose, with few narrowly lanceolate, c. 1 cm or more long utricles. Bracts sheathing. Rhizome short. Sp.48 . . . . 19. Sect. Folliculatae 21. 2 Spikelets not broadly ovoid-subglobose. Utricles smaller. 22. Spikelets linear-cylindric, loosely flowered. Utricles nerveless (except for 2 submar- ginal nerves) or obscurely nerved, with long beak and oblique mouth. Bracts sheathing. 23. Utricles hispidulous, brownish. Sp.46 ..... .. . . 17. Sect. Ferrugineae 23. Utricles glabrous, light green. Sp.47 ... . .... . . 18. Sect. Sylvaticae 22. Spikelets oblong or cylindric. Utricles many- -nerved. 24. Spikelets short-cylindric, few-flowered. Utricles fusiform-ellipsoid or rhomboid, at least towards the base many-nerved. Flowering stems lateral or central. Spp. 22-25 8. Sect. Rhomboidales 24. Spikelets cylindric, densely many-flowered. 1979] Cy PERACEAE—II (Kern & Nooteboom) 111 25. Utricles firm, of thick texture, short-beaked. Rhizome emitting strong stolons. Leaves conspicuously septate-nodulose. AREER erect. Bracts not or scarcely sheathing. Sp.60 .. . . . . . 26. Sect. Paludosae 25. Utricles membranous, with long beak (except in C. brownii ssp. brownii). 26. Spikelets erect. Utricles patulous, subinflated, many-nerved, with shortly bidentate beak. Style straight. Rhizome often creeping. Spp. 42-45 . 16. Sect. Anomalae 26. Lower spikelets cernuous. Utricles widely patent to reflexed, closely many-ribbed, with bifurcate beak. Style flexuous. Rhizome cespitose. Sp. 49 20. Sect. Pseudocypereae 3. Inflorescence consisting of a single terminal spikelet. 27. Utricles with whitish pubescent margins or hispidulous, densely and strongly many-nerved, ASTOMMMMAONSMSPDN GO=—S08 cee lh, Glo sGak sh Reils. nels wl. «ll 2oSect Radieales 27. Utricles glabrous, 1!/,-5 mm long. 28. Utricles rather large (41/,-5 mm long), with slender, linear, bidentate beak. Sp. 41 15. Sect. Rhizopodae 28. Utricles small (1'/.-4 mm long), with short, subentire or slightly emarginate beak. Sp. 40 14. Sect. Capitellatae 2. Stigmas 2; nut lenticular (plano-convex or biconvex). 29. Bracts sheathing. Spikelets often in fascicles. Utricles often hispidulous, membranous, nerved, usually long-beaked. Stigmas often very long. Spp. 56-59 ...... ... 25. Sect. Graciles 29. Bracts not sheathing (or very shortly). 30. Lateral spikelets sessile or the lowest shortly Seieeg erect. Terminal spikelet 3. Glumes MULICOUSS SPyoD) = ae: aE IO TS PA Sect Carex 30. Spikelets peduncled, cernuous. Glumes often awned. 31. Spikelets 3-8, single at the nodes, cylindric, short peduncled. Terminal spikelet 3, or (often with exception df basal spikelets) spikelets gynaecandrous. Spp. 52-54. . . . 23. Sect. Praelongae 31. Spikelets 6—50, single or binate at the nodes, the lower ones long-peduncled, all androgynous or some 3, linear-cylindric. Sp. 51... . . . . 22. Sect. Longispicae 1. Stigmas 2 and nut lenticular. Spikelets similar in 1 appearance, bisexual, sessile. Cladoprophylla usually AlDSC Ube athe) sn & See ee SUBGSVIGNEA 32. Spikelets androgynous (upper flowers “a lower. ones 2). 33. Rhizome long-creeping. Spikelets 4-7. Utricles coriaceous, J spores sharp-edged but not wingedsSp. 61 47. =: pie: Ds Lae27s Sect. Divisae 33. Rhizome short, forming dense tufts. Spikelets numerous or ‘very numerous. 34. Utricles coriaceous, with rounded margins. Bracts inconspicuous. Sp. 62 28. Sect. Paniculatae 34. Utricles membranous, with acute or winged margins. Lower bracts foliaceous. Sp. 63 29. Sect. Multiflorae 32. Spikelets gynaecandrous (upper flowers, 9, lower ones 3). Utricles membranous. 35. Utricles with rounded margins, densely whitish-puncticulate, beakless or very shortly beaked. Spikelets ebracteate (bracts glumiform). Sp. 66 ........ ... . .32. Sect. Heleonastes 35. Utricles with acute or winged margins, not puncticulate. 36. Utricles obliquely erect, also when mature. Spikelets more or less distinctly bracteate. Sp. 65. 31. Sect. Elongatae 36. Utricles widely spreading or reflexed when mature. Spikelets not or inconspicuously bracteate GEARS a EMILOLIN) SDs 104). oc 2. shine or oes SYNTH Ee Pay ee SO Sect. Stellulatae KEY TO THE SPECIES Only complete specimens with nearly or fully mature utricles are identifiable. As the distribution of sexes is variable, especially in the species with a terminal male spikelet, it is advisable to examine several specimens if possible. In the measurements of the length of the utricles the beak is included. Unless stated otherwise ‘glumes’ refers to the glumes subtending the female flowers. 1. Inflorescence consisting of a single (androgynous) spikelet terminating the stem. Stigmas 3; nut trigonous. 2. Utricles with whitish pubescent margins, or hispidulous, or winged and scabrid margined, 4-8 mm long. Glumes (many-)nerved. Style pyramidally thickened towards the base, persistent on the nut. Leaves (2—)3-10 mm wide. 3. Spikelets 2-8 cm long. Utricles with whitish pubescent margins . ... . . . . 37. C. speciosa 3. Spikelets 6-12 mm. Utricles hispidulous or glabrous with scabrid margins. 112 FLORA MALESIANA [ser: I, vol.*9" 4. Utricles hispidulous in the upper 2/3, 4-6 mm, with conic-cylindric, 1—2!/, mm long beak. 38. C. tricephala 4. Utricles glabrous with scabrid, distinctly winged HS 6-8 mm, gradually narrowed into the 3mm long beak .. . . . 36. C. ramosii . Utricles glabrous, membranous and otherwise different, Be 5 mm n long. Glumes 3-nerved i in the centre, otherwise nerveless. Style not or but slightly thickened towards the base. Leaves 1/,-3 mm wide. 5. Utricles (11/,-)2!/,-31/,(-4) mm long, rather gradually tapering into a very short, subentire or slightly emarginate beak. Leaves !/,-2 mm wide. Glumes 1!/,-3-mm long . . . 40. C. capillacea 5. Utricles 4!/,-5 mm long, abruptly narrowed into a slender, linear, bidentate, c. 11/, mm long beak. Leaves 2-3 mm wide. Glumes 3-4 mm long ............. . 41. C. eremostachya 1. Inflorescence consisting of 2 or more spikelets. 6. Terminal and most of the other spikelets 2, shortly cylindrical, the ¢ spikelets lateral, 1-5, just below some of the terminal 2 spikelets. Inflorescence a compound panicle. Leaves all over the stem 13. C. hypolytroides 6. Terminal spikelet bisexual or wholly 2. 7. Terminal spikelet wholly 3 (exceptionally in some specimens with a few 2 flowers), or when spikelets in fascicles, 1-2 spikelets of the upper fascicle wholly ¢. 8. Stigmas 3. 9. Leaves inserted all over the stem, the upper ones merging into leafy bracts, the lower ones gradu- ally decreasing in length and merging into bladeless sheaths. Inflorescence simple, racemose, consisting of 4—9 spikelets. Spikelets eres cm long. Utricles densely hispid, with distinct, c. 1 mm long beak. Leaves glabrous. . . . ss 8 wap ay 3509 @jmaubertiana 9. Leaves basal and often also some higher o1 on the stem. 10. Nut excavated in the middle, the apex abruptly narrowed into a conspicuous, cylindric, 1/,-1'/, mm long beak expanding into the annular apex. . . . . . . . 22. C. anomocarya 10. Nut different. 11. Apex of nut truncate, 3/,-1 mm wide, hollowed out. Style centred in the apical hollow of the nut. Utricles lageniform or rhomboid-lageniform. 12. Utricles 5-7 mm long. Apex of nut contracted into a cylindric, truncate, ?/,-1 mm long and wide neck. 13. Leaves 5-10 mm wide. Beak of utricles 1!/,-2 mm long. Stems arising from basal leaf axils 28. C. gracilispica 13. Leaves 1-3 mm wide. Beak of utricles */,-1 mm long. Stems central 30. C. rhynchachaenium 12. Utricles 3'/,-4'/, mm long. Apex of nut hardly or not contracted into a cylindric neck 27. C. breviscapa 11. Apex of nut not truncate, at most '/, mm wide. 14. Utricles densely papillose, subabruptly contracted into a very short, up to '/, mm long beak 39. C. maculata 14. Utricles not papillose, nut either mitrate or distinctly beaked. 15. Nut mitrate, i.e. contracted at the apex and then expanded into a discoid annulus. 16. Glumes of ¢ spikelets cup-shaped, the margins more than halfway connate in front. Stamens monadelphous (not always so in the lowermost flowers). Utricles many-nerved 34. C. tristachya 16. Glumes of 3 spikelets with free margins. Stamens free. 17. Nut rhomboid with deeply concave faces. . . ..... =... . 33. C. formosensis 17. Nut (long-)ellipsoid to ovoid or obovoid. 18. Leaves 3-10 mm wide. Stems arising from basal leaf-axils. Utricles strongly many-nerved 32. C. dolichostachya 18. Leaves 1-4(-6) mm wide. Stems central. Utricles nerveless to multinerved. 31. C. breviculmis 15. Nut not mitrate. = 19. Spikelets in fascicles of 3-20 from the axils of at least one of the bracts. 20. Utricles narrowly ellipsoid, 4-6!/, mm long, gradually tapering below into a stipe-like, 3/,-1 mm long base, above into a 1-3 mm long beak . . . . . . . 20. C. verticillata 20. Utricles ovoid or ellipsoid, 2'/,—31/, mm long, scarcely stipitate; beak 3/,-1 mm long. 17. C. celebica 19. Spikelets solitary (exceptionally binate) from the axils of the bracts, or crowded at the apex of the stem. 21. Utricles (8-)12-13 mm, many-nerved. 2 Spikelets broadly ovoid to subglobose, 15-25 mm long and wide, with few, divergent, narrowly lanceolate utricles. Bracts long-sheathing 48. C. michauxiana 1979] CYPERACEAE—II (Kern & Nooteboom) 113 21. Utricles at most 7!/, mm long. Other characters not so combined. 22. Broader leaves 1-2 cm wide. Spikelets 5'/,-16 cm long. Glumes vinaceous to dark red with a green, central stripe. Stoloniferous . . . see cl oot se aoe. OUVACER 22. Broader leaves at most 10 mm wide. Spikelets usually much shorter. 23. Stems lateral. Lateral spikelets 6-12 by 4-7 mm, 2 or androgynous. Utricles at least below many-nerved, 5-7 mm long. Glumes 2!/;-4 mm. Nut 3-4 mm. 24. Leaves 3-7 mm wide. Glumes mess mm, with a 1'/,-6 mm long awn. Lateral Spikeletsnca ane , hse pee 6244 GC lateralis 24. Leaves 11/,— 3 mm wide. Glumes oe 4 mm long, muticous or mucronulate. Lateral spikelets androgynous ... . ts. |, utes 4G, loners 23. Stems central. Lateral spikelets more en 10 mm oe ? or rarely androgynous, when shorter than 12 mm nut 2!/,-3 mm. 25. Utricles 5—7'/, mm long; nut at least 2'/, mm long. 26. Glumes 2-2!/, mm, with an antrorsely scabrous, 2-4 mm long awn. Utricles with deeply furcate 1'/,-2 mm long beak, strongly and densely many-nerved, when ripe widely spreading or reflexed. Lower bracts much overtopping the inflorescence, scarcely sheathing. . . . .. . . . . 49, C. pseudocyperus 26. Glumes 3-6 mm long, whether c or not awned. 27. Utricles nerveless or obscurely few-nerved, except for the 2 marginal veins, 5—7!/, mm long. 2 Spikelets linear-cylindric, lax-flowered, 2-9 cm by 3-7 mm. Glumes 3!/,— 6 mm, oblong to ovate, acute to very obtuse, translucent, with broad white margins and greenish midrib, the latter not reaching the apex, but sometimes excurrent into ashorvawn up) tol l(2)immilong 95". Sa se oesee 2) a 4 475) GS finitima 27. Utricles (strongly) many-nerved. 28. Utricles 5'/,-7!/, mm, fusiform. Glumes oblong-ovate, acute, very thin, dirty white with 3-nerved greenish central stripe, 3—5'/, mm long, muticous or mucronulate, rarely witha shortawn .. . Sere 25-7 ackiana 28. Utricles S—5S!/, mm. Glumes ovate, deeply emarginate at the top, pale with purplish margins, c. 3 mm long, the strong midrib excurrent into a firm, smooth or scabrid awn as long asthe glume... . BS the LY C. sp. (§ Paludosae) 25. Utricles 3-5(—7) mm long; nut at most 2), mm iGhie: but mostly shorter; if the utricles are longer than 5 mm, the nut is at most 17/; mm long. 29. Utricles sparsely to rather densely hispidulous, nerveless except for 2 submarginal nerves, 3—4'/, mm long, with stout, 1-1!/, mm long beak. 2 Spikelets linear-cylindric, 1'/,-6 cm by 2!/,-31/, mm. Glumes 3—4'/, mm, with an hispidulous awn up to 1 mm long . Si cba ke on cto 2 46. Cobrachyathera 29. Utricles glabrous, many-nerved ¢ or ribbed. 30. Utricles 4-S(—7) mm long, strongly many-nerved. Glumes 2-2!/, mm, with a 2-4 mm long awn. 2 Spikelets 2!/,-5 cm by 7-15 mm. Leaves 6-10 mm wide. 49. C. pseudocyperus 30. Utricles 3-4 mm (up to 6 mm in C. brownii ssp. transversa), strongly many-ribbed or plurinerved. Glumes 1!/,-2!/, mm, excurrent into a !/,-3!/, mm long awn. 2 Spikelets 1!/,-12 cm by 4-6 mm. Leaves 2-10 mm wide. 31. Utricles strongly many-ribbed. 2 Spikelets short-cylindric to cylindric, 1'/,-3 cm by 5-6 mm. Glumes 1-2 mm with a !/,—-3!/, mm long awn. Nut 2!/,—2!/, mm. Leaves 3-4 mmwide .. es 43...) DFO Wilk 31. Utricles plurinerved. 9 Spikelets (narrowly) cylindric, ( 1 ye yoy 12 cm by 4-6 mm. Nut 11/,-2!/, mm. Leaves 2-10 mm wide. 32. Plant densely cespitose, without stolons. Lower bracts long sheathing. Utricles olive-brown to fuscous. 2 Spikelets 2!/,-12 cm long .. 44. C. oedorrhampha 32. Plant stoloniferous. Bracts not sheathing. Utricles pale greenish to stramineous. Orspikelets; (le /-—)2—-sicmn | ee ee eas, en oe Ca LOpeCHrolges 8. Stigmas 2. 33. Utricles densely covered with raised glandular papillae, slenderly nerved or nerveless. At least the lower spikelets subcernuous to pendulous. Lowest bract much overtopping the inflorescence. 34. Glumes acute to obtusish, muticous or only the lower ones excurrent in a short, up to 1 mm long awn. Utricles slenderly nerved, 3-4 mm long . . . . . 53. C. pruinosa 34. Glumes truncate or bilobed-emarginate, distinctly awned (awn ayet cr mm long). Utricles mostly nerveless, 21/,-31/, mm. . . A, eee enzo. Ce DRACOES 33. Utricles not papillate, whether minutely punctulate o or not. 114 FLORA MALESIANA [ser. I, vol. 91 35. Stems 3—40(-75) cm a 1-1'/, mm. Utricles distinctly 8-14-nerved, 2—3!/, mm. Spikelets erect or suberect, terminal g, remainder 2 or with a few 3 flowers at the top, 1-6 cm. Leaves 1/5—Aummipwides ) -) 4: ... . . . 55, C. gaudichaudiana 35. Stems 30-110 cm by up to 4 mm i(kclow): Spikelets pedis all lateel ones androgynous or 1(-2) at the base of the uppermost much smaller and wholly 3, (3-)6-13 cm long. Leaves 3-12: mmiwide.. 4 «6.8. «es a ele 2 & © D ag Rae roraeiieana 7. Terminal spikelet always bisexual. 36. Terminal spikelet (often all the spikelets) gynaecandrous (i.e. 2 flowers above, 3 ones below); 3 flowers often so few that the spikelets have an entirely 2 appearance. 37. Stigmas 2. Nut lenticular (biconvex or planoconvex). 38. Spikelets all sessile, 1/,-11/, cm long. 39. Utricles distinctly winged on the margins. Lower bracts foliaceous, long, far exceeding the up to 15 cm long inflorescence . . . . . 65. C. remota 39. Utricles wingless. Bracts glumiform, the lowest exceptionally subulate to herbaceous, some- times exceeding the 11/,-31/,(-6) cm long inflorescence. 40. Utricles (3—)4-5(-5!/,) mm long, when mature widely patent to reflexed, distinctly beaked, not whitish-punctulate =~ > . =). curta 38. At least the lowest apikelen peduricledt cernuous, 3- me 8) cm iones sryokyk al S4iuG; teres 37. Stigmas 3. Nut trigonous. 41. Utricles hispidulous or sparsely pubescent. Nut mitrate (i.e. contracted at the apex and then expanded into a discoid annulus). 42. Utricles many-nerved. Glumes of ¢ flowers infundibuliform ...... 34. C. tristachya 42. Utricles obscurely nerved or nerveless. Glumes of ¢ flowers with free margins 31. C. breviculmis 41. Utricles glabrous. Nut not mitrate. 43. Lower bracts sheathing. Plants cespitose. 44. Utricles broadly ellipsoid to subglobose, 3-4 mm long or ovoid-ellipsoid, 6 mm long 43. C. brownii 44. Utricles fusiform or fusiform-ellipsoid, 5—7!/, mm long. 45. Utricles strongly many-nerved ..... . .. .. 23. C. jackiana 45. Utricles nerveless except 2 marginal nerves or ‘obscurely few-nerved . . . 47. C. finitima 43. Bracts not sheathing. Plants stoloniferous . . . . . . 42, C. alopecuroides 36. Terminal spikelet and all or most of the other spikelets androgynous (i.e. 3 flowers above, 2 ones below). 46. Stigmas 3. 47. Leaves pseudopetiolate, 1!/,-3!/, cm wide, the base conduplicate. Stems central, erect, scapiform, the base surrounded by some brown, infundibuliform, up to 10 cm long leaf sheaths. Utricles very long beaked, the beak curved, bulbous at the base (to hold the thickened conical style-base), nearly linear towards the very oblique mouth, the whole beak 3-3'/, mm long, the utricle including the beak 6-7 mm long. Inflorescence paniculate with head-like very dense partial inflorescences. -- 20°. 0) SO ee Se 14. C. helferi 47. Leaves not pseudopetiolate (except i in C. commixta). Other characters not so combined. 48. Spikelets peduncled, in fascicles from at least one of the sheathing bracts, sometimes com- pound. 49. Utricles densely Tee Ds 5-91/, by 1-2!/; mm. cigars 3-8 mm thick. Nut 3-4 mm longi. . . . . 18. C. perakensis 49. Utricles nerveless or r slenderly few- nerved, ca 6 by yp mm. Spikelets 2-3'/, mm thick. Nut 11/,-3'1/, mm long . . . . » woes « » 19); turrita 48. Inflorescence racemose or BpICatE end epiteters pedanciedt or cee single or binate at the nodes or inflorescence a panicle and the secondary panicles single or binate at the nodes. Bracts sheathing or not. 50. Flowering stems arising from the axils of the basal leaves, which are crowded on a short stem and much longer than the flowering stems. Inflorescence racemiform or paniculiform; spikelets 6-30, 8-30 mm long. Bracts usually shorter than the spikelets, with funnel-shaped sheaths and short blade. Nut irregularly rhomboid-ellipsoid or rhomboid-obovoid with faces excavated at base and apex; style flexuous . . .. . . 26. C. eryptostachys 50. Flowering stems not arising from the axils of the basal leaves. 1979 ] Cy PERACEAE—II (Kern & Nooteboom) 115 51. Flowering stems developing from lateral shoots. Inflorescence a narrow, simple panicle, consisting of 6-12 head-like or racemose partial inflorescences each containing 1-8 spikelets 4-10 mm long. Utricles strongly many-nerved, 2'/,-4 mm long. . . . 21. C. oligostachya 51. Flowering stems central. 52. Nut with a stout, cylindric, '/,-*/, mm wide neck. Style centred in the apical hollow of the nut. 53. Inflorescence spiciform, with 2-10 sessile ‘aa Utricles rhomboid-lageniform, 4'/,-Smmlong ... . . . . 29. C, palawanensis 53. Inflorescence a slender compound panicle with numerous spikelets. Utricles ellipsoid- rhomboid,c.4mmlong . . . . .1. C. cirrhulosa 52. Nut without stout cylindric neck. Style not centred i in an ‘apical hollow. 54. Lower bracts not or only very shortly sheathing. 55. Inflorescence with 12 to numerous spikelets, spiciform, the lateral spikelets arising from a utriculiform, gibbous prophyll ene a 2 flower or a nut. eae 3—20(—30) cm oil Le . . . . 12. C. satzumensis 55. Inflorescence with 1-4 spikelets, or 1-4 heads, each consisting of a crowded sessile spikelets. The lateral spikelets not arising from a fertile prophyll. 56. Utricles densely hispidulous in the upper 2/3. Glumes densely setulose. Lateral spikelets of the inflorescence sessile, ovoid or subglobose (sometimes absent) 38. C. tricephala 56. Utricles at most scabrous at the margins towards the apex, the rest glabrous. Sie shies 6-8 mm long, with a c. 3 mm baa beak. Glumes 3-4'/, mm long, with a 1-137/, mmlongawn.. . . . 36. C. ramosii 57. Utricles 4—6'/, mm long, with 2 a 1 9 mm 1 long beak. Glumes ‘2/,— Le long, with a Een OnDtAWH a) ets ot SW, TD PIO aS, ee ae . C. malaccensis 54. Lower bracts long-sheathing. 58. The longer spikelets at least 20 mm. 59. Spikelets many. Utricles 3-41/, mm. Nut 2-3 mm. 60. Beak of utricles straight. Utricles not inflated, ellipsoid or ellipsoid-obovoid, trigo- nous, pale to castaneous, slenderly nerved, sparsely to subdensely hispidulous, at least towards the apex. Leaves 2-10 mm broad .. . en = S165. Cc myosnras 60. Beak of utricles curved. Utricles inflated, obscurely trigonous, ovoid to subglobose, strongly nerved, glabrous except the hispidulous margins towards the apex, at first yellowish green, ultimately red and more or less succulent. Leaves 5-18 mm broad 15. C. baccans 59. Spikelets 1-many. Utricles 4-91/, mm (if utricles less than 5 mm, spikelets few). Nut 3-4 mm. 61. Spikelets 2-many, branched (or when not branched the beak of the utricle 2-3 mm). Utricles 5—9'/, mm long, the beak 1!/,-3 mm. Style-base slightly thickened 18. C. perakensis 61. Spikelets not branched, 1—3(—4), 5-20 cm distant. Utricles 4-7 mm, the beak shorter. Style-base pyramidically thickened. . . . “ os 2 372 Gospeciosa 58. Spikelets never becoming longer than 20 mm. 62. Leaves inserted all over the stem... . ...... . 13. C. hypolytroides 62. Leaves (sub)basal, often 1-3 higher on the stem. 63. Utricles densely pale to oon BERNE 5- oe oy mm BRON: Spikelets at least 8 mm long. . Se: . . . 18. C. perakensis 63. Utricles different, 33/,— 6 mm ‘long. 64. Nut narrowly discoid-annulate at the apex. Utricles whitish setulose, not inflated. Secondary panicles mostly spiciform .. . . . . 8. C. nodiflora 64. Nut not narrowly discoid-annulate at the apex, or utricles glabrous, inflated. 65. Leaves often pseudopetiolate, the broader leaves more than 15 mm wide. Spikelets + patent, rather few-flowered, 5-10 mm long. Glumes ovate or oblong-ovate, obtuse or slightly emarginate, 2-3 mm long, the awn !/,-2 mm. Utricles distinctly trigonous, ellipsoid, with prominent angles and flattish faces, membranous, c. 5- nerved on each face, 4—5(—6, extra-Mal.) by 11/,-13/, mm, subabruptly beaked, the beak 1!/,-2(-3, extra-Mal.) mm mi Soaad bidentulate on the ventral side with very oblique mouth ... . 7s 221Gecommixta 65. Leaves not pseudopetiolate, rarely more than 151 mm wide. 66. Utricles glabrous, whether inflated or not, shiny and conspicuously 2—4-nerved on each face and then glumes muticous or mucronulate, or dull, 3-5-spongy-ribbed or strongly 5-10-nerved on each face, with oblique, often scabrid beak. 116 FLORA MALESIANA [ser. I, vol. 91 67. Utricles shiny, triquetrous, not inflated, conspicuously few-(c. 2-4-)nerved on each face, olivaceous to reddish-castaneous, 2!/,—5 by '/.-1 mm, with slender, subulate or scarcely tapering beak which is about as long as the body or slightly longer (1-2!/, mm long). Glumes 11/,-2'/,(-4!/,) mm, about as long as the body of the utricle, muticous or rarely mucronulate ... . . able ie Lae aerec. filicina 67. Utricles rather dull, inflated or subinflated, or at feast not triquetrous, strongly spongy 3—5-ribbed on each face or strongly 5-10-nerved. (If different proceed under 66 second lead.) 68. Utricles spongy 3—5-ribbed on each face, stramineous or brown, 2-4 by 1(-1!/,) mm, abruptly beaked, the beak Sina MAES with oblique mouth often becoming bifid, 37/,-11/, mmlong. . . . L .. . 3. C. cruciata 68. Utricles strongly many-nerved (5-7 or c. 10 nerves on each face), 31/,-5 by 1-21/, mm. 69. Utricles distinctly inflated, obscurely trigonous, c. 10 nerves on each face. The rather compact ¢ part of the spikelets as long as to usually much longer than the @ part, rarely shorter. Glumes a mm, with an awn to 3!/, mm. Nut Often!rostrate =.) se ee 1 eOn'Ce indica 69. Utricles rhomboid, Pivenone se 7 nerves on eacel fe Nut erostrate. 70. The (often plumose) ¢ part of the spikelets shorter than to about as long as the 2 part. Glumes 1'/,-2 mm, with an awn of 1-3 mm. . 5. C. horsfieldii 70. The 3 part longer than the @ part of the spikelets. Glumes 2—23/, mm, with an aw Ofs/,—2 mm =. ... . . 10. C. stramentitia 66. Utricles glabrous, or hairy, at ated obectrel to conspicuously nerved or spongy-ribbed, the beak bidentate or bidentulate. 71. Glumes distinctly awned, broadly ovate. At least in the lowest glumes of a large part of the spikelets the awn longer than the glume. Utricles with (3—)4-6 strong nerves on each face, glabrous and smooth (glossy), or hispidulous on the margins in the upper part, light green to stramineous, (2!/,-)3-4 by 1-1!/, mm, the beak compresséd, shorter to about as long as the body of the utricle, 1—-1'/, mm long. (If the utricles are hispidulous, see 3. C. cruciata.) . . . 7. C. lamprochlamys 71. Glumes whether awned or not, awn never longer than glume. 72. Glumes muticous or mucronulate, rarely awned. Utricles obscurely nerved, only 2 nerves more prominent, glabrous at the base, otherwise densely scabrid- pubescent, blackish fuscous or greenish, 2/,—4'/, by c. 1 mm; the beak straight, compressed, deeply bidentate, but often originally oblique, blackish fuscous, the mouth with whitish hyaline margins . . . . . . . 9. C. sarawaketensis 72. Glumes often awned. Beak of utricles not blackish fuscous, or utricles not scabrid-pubescent except at the base, or characters otherwise combined. 73. Glumes ovate or broadly ovate, acutish or obtuse. Utricles obtusely trigonous, strongly spongy 3—5-ribbed on each face, glabrous, except sometimes on the margins’ sees eel 7s Cr cruciata 73. Glumes ovate or aorowiy, orate gui. Witieles distinctly trigonous or triquetrous, more or less distinctly several-nerved to obscurely nerved, glabrous or hairy. 74. The 3 and 9 part of the spikelets of same length. Glumes 11/,-2'/. mm 3. C. cruciata 74. In the longer spikelets the ¢ part much one than the few-flowered @ part. Glumes 2!/,-44/, mm ... . kee C. vesiculosa 46. Stigmas 2. 75. Rhizome long-creeping, slender. Leaves canaliculate or convolute, c. 1 mm wide. Spikelets approximate, forming an oblong, 1—1'/, cm long head. Utricles nerveless, glabrous 61. C. duriuscula 75. Rhizome short, plant forming dense tufts. Leaves usually wider than 1 mm. 76. Spikelets very numerous, forming a slender, 5—25 cm long, oblong-cylindric, contracted, spike- like 1-2 cm broad panicle. Leaves with very scabrous margins, 3-10 mm wide. Utricles coria- ceous . . . . . . . 62, C, appressa 76. Spikelets 4-10(- 25), forming a ‘head- like or ‘spike- like 1 ae 5) cm long inflorescence, or spikelets up to 50, in a lax raceme or in (2-)4-8 fascicles of 2—7 spikelets each and inflorescence up to 50 cm. Utricles membranous. 1979] Cy PERACEAE—II (Kern & Nooteboom) 117 77. Spikelets all sessile. Cladoprophylls (see below) absent. Utricles ovate or ovate-lanceolate, not cordate at the base, winged on the margin, 3'/,-4'/, mm long. Glumes mucronate 63. C. nubigena 77. At least the lower spikelets distinctly peduncled. Peduncles of the lateral spikelets at the base surrounded by an ocreiform or utriculiform bracteole (cladoprophyllum) which is often hidden in the sheath of the bract subtending the spikelet. 78. Bracts not sheathing. Spikelets linear-cylindric, very ae flowered, (2—)6-13 cm long. 79. Utricles densely beset with ferrugineous papillae . . F hoe: 2 . 52. C. phacota 79. Utricles epapillate . . . . 51. C. graeffeana 78. Bracts sheathing. Spikelets loosely or somewhat densely flowered, up to 4'/, cm long. 80. Inflorescence very lax, only 1 spikelet at each node. Glumes with a stoutish, up to 5 mm long awn. Utricles glabrous except for the sparsely hispid beak (rarely the margin setulose) 58. C. longipes 80. Spikelets often in fascicles at the nodes. Glumes muticous or minutely apiculate (rarely an awn up to 2 mm present). Utricles setulose at least on the margins. 81. Lower bract setaceous, suddenly widening into a spathaceous, reddish brown base clasping the base of the spikelets. Glumes 4-6 mm 56. C. bilateralis 81. Lower bracts foliaceous. 82. Glumes distinctly shorter than to about as long as the utricle, 2-4 mm, rarely some of them to 5 mm. Stigmas shorter than to about as long as the utricles, up to c. 5 mm 57. C. brunnea 82. Glumes about as long as the utricles, 3!/,-5 mm. Stigmas very long (7-12 mm), always muchwoncemthannthe:utticless). «sft | fis ee eee 59. C. teinogyna I. Subgenus Carex Carex subg. Indocarex BAILL. Hist. Pl. 12 (1893) 345. — Primocarex KUK. Verh. Bot. Ver. Brandenb. 47 (1905) 204. — Carex subg. Primocarex KUK. Pfl. R. Heft 38 (1909) 68. Type species: Carex acuta L. 1. Section Vigneastra Tuckerm. En. Meth. (1843) 10. — Sect. Vigneastra [grex} Indicae [TUCKERM. I.c., nomen]; ex BAILEY, Proc. Am. Ac. 22 (1886) 98. — Sect. Indicae CLARKE, FI. Br. Ind. 6 (1894) 713; Kix. Pfl. R. Heft 38 (1909) 260; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 38; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 491, in nota. — Sect. Polystacheae CLARKE [ser.] Stramentitiae, Cruciatae & Filicinae CLARKE, J. Linn. Soc. Bot. 37 (1904) 4. — Sect. Stramentitiae (CLARKE) NELMES, Reinwardtia 1 (1951) 250, p.p. — Sect. Cruciatae (CLARKE) NELMES, /.c. 275. — Sect. Filicinae (CLARKE) NELMES, /.c. 286; RAYM. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 48. — Sect. Indicae subsect. Indicae KoYAMA, J. Fac. Sc. Un. Tokyo Ill, 8 (1962) 151. Type species: Carex indica L. (lectotype). 1. Carex cirrhulosa Nees in Hook. J. Bot. Kew Misc. 6 (1854) 29; NeLMgs, Reinwardtia 1 (1951) 262. — ? C. densiflora PRESL, Rel. Haenk. 1 (1828) 204; F.-ViLL. Nov. App. (1882) 310. — C. fuire- noides (non GAUDICH.?) F.-VILL. Nov. App. (1882) 310; CrarKeE, J. Linn. Soc. Bot. 37 (1904) 11, pep:> Philip: J. Se: 2. (1907) Bot: 107, p.p. — — C. fibrata Boott ex VIDAL, Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 286, nomen. — C. fuirenoides Gaupicu. var. cirrhulosa KUK. Pfl. R. Heft 38 (1909) 287; Philip. J. Sc. 6 (1911) Bot. 61; MerRR. En. Philip. 1 (1923) 138, p.p. — Fig. 118. Inflorescence a slender, compound panicle, continuous above, interrupted below, c. 18 cm long; secondary panicles 6, single at the nodes, erect, oblong, rather dense, 1!/,-3'/, cm long, their lower branches again branched into several sessile, patent, 118 FLORA MALESIANA [ser. I, vol. 9! crowded spikelets on trigonous, smooth or sparsely scabrid peduncles; rachis sparsely scabrid above. Lower bracts foliaceous, much exceeding the in- florescence, stiff, flat or with revolute margins, long-attenuate, long-sheathing, 5-8 mm wide, upper ones much reduced, shortly sheathing. Spikelets numerous, androgynous, oblong, ovoid, or subglobose, dense, 4-5 mm long, their 3 part about as long as the 9, their bracteoles glumiform, with hispid, often curved awns, c. 10 mm long. Glumes ovate or suborbicular, translucent, erose- ciliolate at the apex, otherwise glabrous, 3/,-1 mm long, the midnerve excurrent in a stout, flat, sparsely hispid, straight or slightly curved, 11/,-2'/, mm long awn. Utricles trigonous, ellipsoid: rhomboid, subcoriaceous, patent, strongly many- nerved, glabrous, smooth or very sparsely scabrid at the apex, usually slightly curved, somewhat spongy-thickened at the base, subabruptly narrowed into the beak, pale stramineous to brown, 4 by 1-1!/, mm; beak scarcely tapering, compressed, sparsely scabrid, straight or slightly curved, bidenticulate, with straight mouth, 11/, mm long. Nut triquetrous, with prominent angles and concave faces, ellipsoid-rhomboid, stipitate, the apex narrowed into a short, thick neck expanding into a discoid ring c. '/; mm diam., 2!/, by 1- 1'/, mm. Style-base broadly pyramidal, persistent on the nut, centred in the apical hollow of the nut. Stigmas 3. Distr. Malesia: Philippines (Cebu). Note. Only known from the type collection (CUMING 1764, collected in 1841), which is re- presented in BM and K by very defect specimens (rhizome, leaves and parts of the inflorescence missing), so that it is difficult to ascertain its status. CLARKE considered it synonymous with C. fuire- noides GAuDICH. from the Marianas, which is C. indica L. NELMES supposed affinity with C. cryptostachys BRONGN. and the species of sect. Lageniformes (OHW1) NELMES, wrongly as I think. 2. Carex commixta Steup. [in Zoll. Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 207; Mia. FI. Ind. Bat. 3 (1856) 349; Kern, Blumea 15 (1967) 427, f. 1; in Back. & Bakh. f. Fl. Java 3 (1968) 491. — C. horsfieldii (non Boott) Miq. Fl. Ind. Bat. 3 (1856) 349, p.p: (quoad specim. Jungh.); Kix. Pfl. R. Heft 38 (1909) 273; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 67. — C. spatiosa Bootr, Ill. 2 (1860) 86, t. 246; Boeck. Linnaea 40 (1876) 349; CLarKeE, J. Linn. Soc. Bot. 37 (1904) 12, incl. var. bogorensis CLARKE; KUK. Pfl. R. Heft 38 (1909) 265; Camus, Fl. Gén. I.-C. 7 (1912) 188; NELMES, Kew Bull. (1946) 21, 23; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 111; RAymM. Mém. Jard. Bot. Montréal n. 53 (1959) 40. — C. blepharolepis NELMES, Kew Bull. (1946) 18, 23; Reinwardtia 1 (1951) 265. — C. smitinandii RayM. Dansk Bot. Ark. 23 (1965) 255, f. 1. — Fig. 118. Rhizome shortly creeping, stout, woody. Stems loosely tufted, triquetrous, smooth, 40-100 cm by 2-3 mm, the base clothed with membranous, pale or fuscous bladeless sheaths or their fibrous remains. Leaves herbaceous, subbasal and a few higher on the stem, equalling to much longer than the stems, linear-lanceolate, long-attenuate, taper- ing below (often into a pseudo-petiole), flat, light green, scabrid on the margins, 1'/,-3 cm wide. Inflorescence a pale, compound, erect, much inter- rupted, 15-40 cm long panicle; secondary panicles 2-6, single at the nodes, erect, pyramidal, loose, 3-7'/, by 1'/,-5 cm, upper continuous, lower (when more than 2) distant, on exserted, hispidu- lous peduncles; rachis densely whitish hispid. Lower bracts foliaceous, exceeding the inflores- cence, long-sheathing, upper reduced. Spikelets androgynous, subsessile, patulous, rather few- flowered, 5-10 mm long, ¢ and 9 parts about equal in length. Glumes ovate or oblong-ovate, thinly membranous, obtuse or slightly emarginate, slenderly nerved, ciliolate at least at the top, other- wise glabrous or minutely adpressed-hispidulous, pale stramineous to light brown, 2—3 mm long, the midnerve excurrent in an antrorsely scabrid, 1/,-2 mm long awn. Utricles distinctly trigonous, ellipsoid with prominent angles and flattish faces, membranous, not inflated, patulous, many-nerved (nerves c. 5 on each face), glabrous or very sparsely hispidulous, rounded at the base, straight or slightly recurved, subabruptly beaked, 4-5(-6) by 11/,-13/, mm; beak sparsely scabrid on the margins, or smooth, 11/,-2(—3) mm long, bidenticulate on the ventral side, with very oblique mouth. Nut trique- trous, ellipsoid-rhomboid, not cuneate at the base, erostrate, dark brown with yellowish angles, 21/,- 23/, by 1/,-14/; mm. Style-base pyramidally thickened, subpersistent on the nut. Stigmas 3. Distr. Burma, N. Thailand, Tonkin, Annam, Hainan; in Malesia: Sumatra (Bencoolen, Lam- pongs), West and Central Java. Ecol. Moist places in primary and, less fre- quently, secondary forests, 800-1500 m, along water-courses sometimes descending to 300 m. Vern. Java: ilat harashas, S. Note. The type of C. spatiosa is from Annam (GAUDICHAUD 67). According to CLARKE, l.c., the Malesian plants should differ by the ovoid utricles with a scabrid beak 1/,-7/, as long as the body, and according to NELMEs (1946, /.c.) by the shorter, glabrous glumes and the shorter utricles. On the whole the Indochinese specimens have longer utricles (up to G6 mm) because of the slenderer beak, but some of them are indistinguish- able from the Malesian plants, which were described as C. spatiosa var. bogorensis CLARKE and C. blepharolepis NELMES. 3. Carex cruciata WAHLENB. Vet. Akad. Handl. Stockh. 24 (1803) 149; CLarkeg, FI. Br. Ind. 6 (1894) 715; J. Linn. Soc. Bot. 37 (1904) 9; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 118; KUxK. 1979] Cy PERACEAE—II (Kern & Nooteboom) 119 E VIJSMA ’78 Fig. 118. Utricles and nuts of Carex. Species numbered as in the text (J CumING 1764, 2 Jacoss 8046, 3a CHEW, CORNER & STAINTON 1706, 36 (left) VERHEIJEN 2585, 3b (right) VAN Ooststroom 13154, 4 Hartcey 11122, 5 NGF 21261, 6 Riwiey 15721, 7 Brass 24684, 8 B. F. HERNAyY 652, 9 PULLEN 6106, 10 BAKHUIZEN VAN DEN BRINK 6565, J] VAN STEENIS 4140). All x 10. 120 FLORA MALESIANA [ser: 1; vol: 91 Pfi. R. Heft 38 (1909) 265; Camus, Fl. Gén. I.-C. 7 (1912) 189, f. 27, 5-9; Rip. Fl. Mal. Pen. 5 (1925) 185; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 11 (1936) 460; Kuk. Bull. Jard. Bot. Btzg III, 16 (1940) 315; Netmes, Reinwardtia 1 (1951) 277; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 129; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 46; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 153; RayM. Dansk Bot. Ark. 23 (1965) 254; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — Fig. 118, 124a-d. Further synonyms under the varieties. Rhizome short or shortly creeping, stout, woody. Stems tufted, stout, trigonous, smooth, up to 200 cm by 2-5 mm, surrounded below the leaves by bladeless, dark reddish to blackish sheaths and their fibrous remains. Leaves (sub)coriaceous, mostly basal but 1-3 higher on the stem, much exceeding to shorter than the stems, linear, long- attenuate, flat or with revolute margins, scabrid on the margins and the nerves, (3-)5-14 mm wide. Inflorescence a slender, usually much elongated, interrupted, decompound, ferrugineous or strami- neous panicle up to 70 cm long; secondary panicles 3-12, at 3-8 nodes, all or the middle ones binate at the nodes or all single, erect or suberect, oblong- linear or (broadly) oblong-lanceolate or ovoid- subpyramidal, loose to dense, 5-10 cm long, the upper approximate, the lower distant on unequal scabrid or smooth peduncles; rachis scabrid-hispid or hispidulous on the angles. Lower bracts folia- ceous, equalling or exceeding the inflorescence, long-sheathing, upper much reduced; bracteoles excurrent in a slender, ciliolate-scabrid, often recurved awn. Spikelets (very) numerous, andro- gynous, obliquely patent or divaricate, 4-8 (-10) mm long, ovoid to oblong, the ¢ part as long as or somewhat longer than the @ part. Glumes broadly ovate, ovate or ovate-lanceolate, acute to very obtuse, membranous, (sub)translu- cent, glabrous or sparsely hispidulous, with reddish brown to dark brown streaks, 1'/,-3 mm long, several-nerved, the midnerve of the lower ones acute or excurrent in a smooth or hispidulous, up to 1 mm long awn. Utricles distinctly or obtusely trigonous, not inflated or subinflated, (broadly) ellipsoid to ovoid, membranous or subcoriaceous, patulous or patent, straight or slightly recurved, often finally recurved, more or less distinctly several-(spongy-)nerved, glabrous or sparsely hispidulous, (sub)abruptly beaked, stramineous or brown with reddish brown streaks and spots, 2-4 by */,—1(—-1'/,) mm; beak slender, smooth or scabrid on the margins, (*/,—)1—1'/, mm long, with oblique but soon bidenticulate to bifid mouth. Nut distinctly trigonous, with prominent angles and flat or concave sides, (ovoid-)ellipsoid, not or scarcely beaked, scarcely stipitate, stramineous to brown or blackish, 11/,-2!/, by 3/,-1 mm. Style- base not or slightly thickened. Stigmas 3. Distr. Widely spread, from India through SE. Asia to S. China, Formosa and the Ryu Kyu Is.; throughout Malesia. Note. Several collections from Sumatra: vAN BorRSSUM WAALKES 2753, BUNNEMEIJER 2530 (type of C. buennemeijeri NELMES), 3644, 3880, 4126, 4651, are intermediate between var. cruciata and var. rafflesiana. KEY TO THE VARIETIES 1. Leaves usually much exceeding the stem. Inflorescence stramineous. Utricles strongly spongy 3—5-ribbed on each face a. var. cruciata 1. Leaves usually as long as the stems or shorter. Inflorescence ferrugineous. Utricles more or less distinctly 3-7-nerved on each face b. var. rafflesiana a. var. cruciata. — C. cruciata WAHLENB. Vet. Akad. Handl. Stockh. 24 (1803) 149; Ripv. FI. Mal. Pen. 5 (1925) 185, incl. var. condensata RIDL. — C. bengalensis Roxs. FI. Ind. ed. Carey 3 (1832) 572; Boortt, Il. 2 (1860) 85, t. 240-243; Bogck. Linnaea 40 (1876) 346, excl. varieties. — C. con- densata NEES in Wight, Contr. (1834) 123; CLARKE, Fl. Br. Ind. 6 (1894) 716; Boorrt, Ill. 2 (1860) 86, t. 247-248. — C. vacua Boott ex Boeck. Linnaea 40 (1876) 343, p.p. (excl. pl. jav.). — C. valida NEES in Wight, Contr. (1834) 123, p.p.; KUNTH, En. 2 (1837) 513, p.p. — C. continua CLARKE, FI. Br. Ind. 6 (1894) 717. — ? C. repanda CLARKE var. implumis CLARKE, J. Linn. Soc. Bot. 37 (1904) 9. — C. galactolepis NELMES, Kew Bull. (1946) 20; Reinwardtia 1 (1951) 279. — C. spongocrepis NELMES, Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 128. — Fig. 118, 124a-b. Rhizome shortly creeping. Stems 40-150 cm. Leaves coriaceous, usually much exceeding the stems. Inflorescence stramineous, continuous above, usually interrupted below, 15-60 cm long; secondary panicles 3-11, at 3-8 nodes, usually all single at the nodes, sometimes binate at the middle nodes, erect, broadly lanceolate or ovoid-sub- pyramidal, rather dense to dense, on stiff, smooth or scabrid peduncles long-exserted from the sheaths, up to 10 cm long; rachis hispidulous on the angles. Lower bracts slightly to much exceeding the inflorescence. Spikelets very numerous, divari- cate, rather dense, 5—8(—10) mm long, the ¢ part as long as or somewhat longer than the @ part. Glumes ovate or broadly ovate, acutish to very obtuse, thinly membranous, translucent, several- nerved, glabrous, with ferrugineous to dark brown streaks, 13/,-3 mm long. Utricles obtusely trigo- nous, subinflated, ovoid or broadly ellipsoid, membranous, patent, finally recurved, strongly spongy-ribbed (nerves 3-5 on each face), glabrous, rarely sparsely setulose at the apex, often with spongy-thickened base, abruptly beaked, strami- neous or brown, purplish spotted, 2'/.-3(-4) by Uijisy Bddiia ii Eire ot ~ hua COESE ES Riccar Sek AEA LVS LNTT I>) 9 seeatnadiantiaed a Soe V IPR IRIN they Strata, AF tlh re Fig. 119. Utricles and nuts of Carex. Species numbered as in the text (J2 Santos 5776, 13 BUNNEMEUER 9820, 14 BACKER 22996, 15 ELBERT 62, 16 VAN OosTSTROOM 13293, 17 NGF 39555, 18a VAN STEENIS 8277, 18b BROOKE 8561, 18c VAN STEENIS 8461, 19a ANU 15532, 19b SANTOS 5785, 20 MEIJER 6127). All x 10. {22 FLORA MALESIANA [ser. I, vol: 91 1(-1'/,) mm; beak usually straight, smooth or sparsely scabrid on the margins, with oblique mouth often becoming bifid, (*/,-)1—-11/, mm long. Nut with flat or slightly concave sides, ovoid- ellipsoid, olive-brown to blackish, 2—2!/, by °/19- 1 mm. Style-base slightly thickened. Distr. Widely spread, from India through SE. Asia to S. China, Formosa, the Ryu Kyu Is., and Queensland; in Malesia: Malay Peninsula, Sumatra, N. Borneo. Ecol. In savannahs, open places in mountain forests, forest-borders, along river-banks, 1100- 1500 m. Vern. Rija-rija. Note. Very variable; sometimes hard or im- possible to distinguish from var. rafflesiana, especially in Sumatra. The type specimens of C. galactolepis (KiNG’s coll. 106) and C. repanda var. implumis (WRAY 1982) are too poor to be certain of their affinity. b. var. rafflesiana (BootT) Noort., stat. nov. — C. rafflesiana Boott, Trans. Linn. Soc. 20 (1846) 132; Ill. 1 (1858) 12, t. 33; CLARKE, J. Linn. Soc. Bot. 37 (1904) 10, incl. var. tenuior CLARKE; KUK. Pfl. R. Heft 38 (1909) 282, incl. var. macro- thyrsa KUK. et var. scaberrima KUx.; Philip. J. Sc. 6 (1911) Bot. 59, incl. var. scaberrima KUK. et var. continua KUK.; MERR. En. Philip. 1 (1923) 140, incl. var. scaberrima KUK. et var. continua KUK.; KUK. in Hochr. Candollea 6 (1936) 430, incl. var. minor KUK.; NELMES, Reinwardtia 1 (1951) 290, incl. var. macrothyrsa KUK. et var. virgata NELMES; ibid. 2 (1954) 376; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 153, incl. var.; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — C. macrothyrsa Mia. Fl. Ind. Bat. 3 (1856) 351. — C. virgata Mia. l.c., non SOL. ex Boott, 1853. — C. pentacarpa BOECK. Flora 58 (1875) 265. — C. vacua Bootr ex BOECK. Linnaea 40 (1876) 343, p.p. (pl. jav.). — C. benga- lensis var. virgata BOECK. et var. scaberrima Boeck. /.c. 347. — C. continua [non CLARKE, FI. Br. Ind. 6 (1894) 717] CLarKeE, J. Linn. Soc. Bot. 37 (1904) 11; Philip. J. Sc. 2 (1907) Bot. 107; Kx. Pfi. R. Heft 38 (1909) 282, p.p.; NELMES, Rein- wardtia 1 (1951) 299; ibid. 2 (1954) 376. — C. gem- bolensis CLARKE var. timorensis CLARKE, J. Linn. Soc. Bot. 37 (1904) 10. — C. scaberrima CLARKE, l.c. 10; Philip. J. Sc. 2 (1907) Bot. 107; Kew Bull. add. ser. 8 (1908) 72. — C. clarkeana KUx. Bull. Herb. Boiss. II, 4 (1904) 52, ex descr.; Pfl. R. Heft 38 (1909) 282; RipL. Fl. Mal. Pen. 5 (1925) 183; NELMES, Reinwardtia 1 (1951) 288. — C. pycno- thyrsos KUK. Philip. J. Sc. 6 (1911) Bot. 60; MerRR. En. Philip. 1 (1923) 140; Ne_mes, Reinwardtia 1 (1951) 282; ibid. 2 (1954) 374, descr. — C. semi- glomerata KUx«. Bull. Jard. Bot. Btzg III, 16 (1940) 315; in Fedde, Rep. 53 (1944) 106; NeELMes, Reinwardtia 1 (1951) 285; ibid. 2 (1954) 376. — C. sarawaketensis KUx. var. brevirostris KUK. Bot. Jahrb. 70 (1940) 464. — C. timorensis (CLARKE) NELMES, Kew Bull. (1946) 24; Reinwardtia 1 (1951) 242, 287. — C. spongoneura NELMES, Kew Bull. (1946) 18; Reinwardtia 1 (1951) 281. — C. xestogyne NELMES, Kew Bull. (1946) 16; Reinwardtia 1 (1951) 311. — C. buennemeijeri NELMES, Kew Bull. (1950) 191; Reinwardtia 1 (1951) 283; ibid. 2 (1954) 376. — C. oblonga NELMES, Kew Bull. (1950) 192; Reinwardtia 1 (1951) 297. — C. ceramica NELMES, Kew Bull. (1950) 193; Reinwardtia 1 (1951) 319; ibid. 2 (1954) 376. — Fig. 118, 124c-d. Rhizome short. Stems up to 200 cm. Leaves subcoriaceous, mostly as long as or shorter than the stems. Inflorescence ferrugineous, interrupted, up to 70 cm long; secondary panicles 8-12, at 4-6 nodes, all or the middle ones binate at the nodes, or single in depauperate specimens, erect or suberect, oblong-linear or oblong-lanceolate, loose or rather dense, 5-10 cm long, the lower on unequally exserted scabrid peduncles; rachis scabrid-hispid on the angles. Lower bracts equal- ling or exceeding the inflorescence. Spikelets numerous, obliquely patent, 4-7 mm long, the ¢ and @ part about equal in length. Glumes ovate or ovate-lanceolate, acute, membranous, subtrans- lucent, glabrous or sparsely hispidulous, with ferrugineous streaks, 11/,-2!/, mm long. Utricles distinctly trigonous, not inflated, ellipsoid or ovoid-ellipsoid, membranous or subcoriaceous, patulous, straight or slightly recurved, more or less distinctly 3—7-nerved, glabrous or sparsely his- pidulous, subabruptly beaked, stramineous with reddish brown streaks and spots, 2-4 by */,-1 mm; beak slender, scabrid on the margins, 1-1!/, mm long, with oblique but soon bidenticulate mouth. Nut with concave sides, ellipsoid, stramineous to brown, 1'/,-2 by 3/,-1 mm. Style-base not or scarcely thickened. Distr. Thailand, Formosa (Kotosho Queensland; throughout Malesia. Ecol. Primary forests, forest edges, grassy slopes, sometimes in dry sunny places, 500-2400 m. Vern. Java: ilateun, S; Philippines: chidak, Ig., ikidsan, Klg., talayid, tamalang, Bag. IS5))5 4. Carex filicina NEEs in Wight, Contr. (1834) 123; KuntTH, En. 2 (1837) 510; Boorr, Ill. 3 (1862) 105, t. 311-318; Boeck. Linnaea 40 (1876) 352; O. K. Rev. Gen. Pl. 2 (1891) 748, incl. var. laevis O. K. et var. ciliata O. K.; CLARKE, Fl. Br. Ind. 6 (1894) 717; J. Linn. Soc. Bot. 37 (1904) 11; Kix. Pfi. R. Heft 38 (1909) 274, incl. var. saturata (CLARKE) KUk.; Camus, FI. Gén. I.-C. 7 (1912) 191; Merr. En. Philip. 1 (1923) 137; Netmes, Reinwardtia 1 (1951) 304, incl. var. angustifolia NELMES et var. zipelii NELMES; ibid. 2 (1954) 376; AKryAMA, Car. Far East Reg. Asia (1955) 138, t. 124; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 48; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 152. — C. nilagirica Steup. Syn. 2 (1855) 207. — C. neoguineensis CLARKE, J. Linn. Soc. Bot. 37 123 FE VIJSMA ’78 E ” ame 7 sate Fig. 120. Utricles and nuts of Carex. Species numbered as in the text (27 VANOVERBERGH 323, 22 DE WILDE c.s. 13535, 23 VAN STEENIS 4579, 24 KAUDERN 333, 25 BS 45002, 26 MEUER 688, 27 LORZING 6678, 29 ELMER 13146, 30 MERRILL 512). All x 10. 124 FLORA MALESIANA [ser. I, vol. 9} (1904) 12; Kuk. Pfl. R. Heft 38 (1909) 280; S. T. BLAKE, J. Arn. Arb. 28 (1947) 105; NELMEs, Reinwardtia 1 (1951) 308; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — C. saturata CLARKE, J. Linn. Soc. Bot. 37 (1904) 12; Nemes, Rein- wardtia 1 (1951) 302. — C. filicina NEEs f. saturata KUK. Philip. J. Sc. 6 (1911) Bot. 59. — C. sclerioides RipL. Trans. Linn. Soc. Bot. II, 9 (1916) 247; NELMES, Reinwardtia 1 (1951) 307. — C. ceylanica BoECK. var. saturata KUK. in Hochr. Candollea 6 (1936) 431. — C. sarawaketensis KUK. var. glabrinux KUK. Bot. Jahrb. 70 (1940) 464. — Fig. 118. Rhizome short, stout, woody. Stems loosely tufted, slender to stout, trigonous, smooth, up to 150 cm by 6 mm near the base, surrounded below the leaves by bladeless, reddish or blackish sheaths. Leaves herbaceous to subcoriaceous, mostly basal but some spaced on the stem, as long as or shorter than the stems, linear, long-attenuate, flat or with revolute margins, scabrid on the margins, (2—)7- 20 mm wide. Jnflorescence a slender, interrupted or continuous, decompound, fuscous panicle up to 60 cm long; partial panicles 5-13, at 4-8 nodes, single and (middle ones) binate, oblong-lanceolate or pyramidal, erect, rather loose to very dense, up to 12 by 2-5 cm, the upper approximate, the lower usually distant on smooth or scabrid, unequally exserted peduncles; rachis smooth towards the base, hispidulous above. Lower bracts foliaceous, shorter than to slightly exceeding the inflorescence, long-sheathing, the upper much _ reduced; bracteoles excurrent into a hispidulous awn. Spikelets very numerous, androgynous, oblong, rather loose, 4-15(-20) mm long, the ¢ part shorter than to about as long as the @ part. Glumes ovate-lanceolate, acuminate, muticous, rarely minutely mucronulate, membranous, sub- translucent, glabrous or the lower ones hispidulous, finely nerved, ferrugineous, often darker streaked, 11/,-2'/,(-4'/,) mm long (about as long as the body of the utricle). Utricles triquetrous, not inflated, ellipsoid, membranous, patent, eventually arcuately recurved, conspicuously 2—4-nerved on each face, glabrous, smooth, often shiny, sub- abruptly beaked, olivaceous to reddish-castaneous, (2'/,-)3-5 by */,-1 mm; beak slender, subulate (scarcely tapering), smooth, or very sparsely scab- rid on the margins, (1-)11/,-2(-2!/,) mm long (about as long as the body or slightly longer); mouth very oblique (the base '/,—*/, mm from the apex). Nut trigonous, ellipsoid, beaked, closely filling the utricle, 11/,-17/, by '/.-1 mm. Style- base slightly thickened. Stigmas 3. Distr. Ceylon and India to S. China, Formosa, Tonkin, Laos and Annam; probably throughout Malesia, in the Lesser Sunda Is. only known from Lombok and in the Malay Peninsula from Pahang. Ecol. Wet openings in primary forests, in sub- alpine shrub vegetation, on grassy slopes, along river-banks, 1350-3750 m. Vern. Philippines: silak, tdan, Ig., taldyig, Bag. Notes. In the circumscription here accepted very variable in all its parts. Numerous segregates have been described, but typical C. filicina from India and the aberrant forms from New Guinea are connected by all sorts of intermediates. Typical C. neoguineensis is characterized by its dense, often darker inflorescence, and the utricles more scabrid on the beak (usually smooth or almost so in typical C. filicina). The high variability of C. filicina also in India is amply discussed and excellently figured by Bootrt, [Le The few specimens on which C. sarawaketensis var. glabrinux KUxk. was based are more or less transitional to 9. C. sarawaketensis. 5. Carex horsfieldii Boott, Proc. Linn. Soc. Lond. 1 (1845) 257; Ill. 1 (1858) 11, t. 32; Mig. FI. Ind. Bat. 3 (1856) 349, p.p. (quoad specim. Horsf.); CLARKE, J. Linn. Soc. Bot. 37 (1904) 11; NELMegs, Reinwardtia 1 (1951) 269 (incl. var. major NELMES?); ibid. 2 (1954) 374; RayM. Mém. Jard. Bot. Mon- tréal n. 53 (1959) 44; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — C. fleckeri NELMES, Kew Bull. (1939) 313, fide Netmes 1951. — Fig. 118, 124e-¢g. Rhizome very short, stout, woody. Stems loosely tufted, obtusely trigonous, smooth, 50-100 cm by 2-4 mm, the base clothed with fuscous, bladeless sheaths and comose by their fibrous, fuscous to blackish remains. Leaves herbaceous, subbasal and a few higher on the stem, exceeding the stems, linear, long-attenuate, flat or with revolute mar- gins, greyish-glaucescent, scabrid on the margins in the upper part, 5-15 mm wide. Inflorescence a pale, decompound, erect, much interrupted, 20-40 cm long panicle; secondary panicles 5-8, single at the nodes, erect, oblong-ovoid, loose, distant, up to 10 cm long; peduncles smooth below, scabrid above, the lower much exserted from the sheaths, upper shortly; rachis hispid on the angles. Spikelets androgynous, widely patent, finally almost squar- rose, sessile, 5-15 mm long, the ¢ part shorter than to about as long as the 2 part. Glumes ovate, obtuse (the lower ones truncate to emarginate), membra- nous, glabrous or sparsely hispidulous, not ciliate, slenderly nerved, whitish, 11/,-2 by 1-1'/, mm, the midnerve excurrent in an antrorsely scabrid, curved, 1-3 mm long awn. Utricles distinctly trigo- nous, rhomboid, with prominent angles and flat faces, membranous, not inflated, strongly many- nerved (nerves c. 5-7 on each face), glabrous, patent, arcuately recurved, gradually tapering to the base, suddenly narrowed above into the beak, olive-brown, 3!/,-4'/, by 1-11/, mm; beak strongly recurved, smooth or very sparsely scabrid, 11/,- 2 mm long; mouth dorsally very oblique, not bidentate. Nut trigonous, rhomboid-ellipsoid, erostrate, dark brown with prominent pale angles, 1979] Cy PERACEAE—II (Kern & Nooteboom) 125 E VIISMA ’?78 Fig. 121. Utricles and nuts of Carex. Species numbered as in the text (3/a BRAss 4697, 3/b EyMa 864, 31c BRASS 9032, 32 SANTos 5394, 33 EDANOo 17857, 34 ROBBINS 5.n., 35 ROBINSON 6186, 36 RAMos 20990, 37 LORZING 16274, 38 BACKER 20462, 39 DE WILDE c.s. 13325, 40 KALKMAN 5213, 4/ Brass 10255, 42 JERMY 4274). All x 10. 126 FLORA MALESIANA [ser. I, vol. 91 2-2'/, by 1-11/; mm. Style-base pyramidally thick- ened at the base. Stigmas 3. Distr. Burma, Thailand, Annam (var. annamica RayM.), Queensland (C. fleckeri NELMES); in Mal- esia: West to East Java, Moluccas, New Guinea. Ecol. Primary and secondary forests, 100- 1100 m. Vern. Java: irissan, J; New Guinea: homuma, Garaina. Notes. C. horsfieldii var. major NELMES, from Tanimbar Is., P. Jamdena (“‘utricles 5.2-5.8 mm long, straight or slightly recurved, beak 2.5-— 2.8 mm long’’), was based on a young, very poor collection. KUKENTHAL and others misapplied the name C. horsfieldii to the Javan plants of C. commixta STEUD., while C. horsfieldii itself has often been misidentified as C. indica L., from which species it chiefly differs in the shorter ¢ part of the spikelets, and the strongly recurved, distinctly trigonous, not inflated, less densely nerved utricles. 6. Carex indica LINNE, Mant. 2 (1771) 574; Mia. Fl. Ind. Bat. 3 (1856) 350; Boortr, Ill. 2 (1860) 87, t. 250-254; Boeck. Linnaea 40 (1876) 347; CLARKE, Fl. Br. Ind. 6 (1894) 714, incl. var. ? laetebrunnea CLARKE ef var. milnei BooTT ex CLARKE; J. Linn. Soc. Bot. 37 (1904) 8; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 117; Kx. Pfl. R. Heft 38 (1909) 262, f. 40, incl. var. fissilis (BOOTT) KUK., saltem quoad specim. males.; Camus, Fl. Gén. I.-C. 7 (1912) 187; Merr. En. Philip. 1 (1923) 138; RiDt. Fl. Mal. Pen. 5 (1925) 185; Ne_mes, Reinwardtia 1 (1951) 271; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 42; Dansk Bot. Ark. 23 (1965) 253; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — C. fuirenoides GAUDICH. Freyc. Voy. Bot. (1826) 412. — C. moritzii Steup. [in Zoll. Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 207; Mia. Fl .Ind. Bat. 3 (1856) 350; Boeck. Linnaea 40 (1876) 350. — C. dietrichiae Boeck. Flora 58 (1875) 122; CLARKE, J.. Linn. Soc. Bot. 37 (1904) 8; S. T. BLAKE, J. Arn. Arb. 28 (1947) 102; NeELMEs, Reinwardtia 1 (1951) 273; Kew Bull. 2 (1955) 303. — C. fuirenoides (non GAUDICH.?) CLARKE, J. Linn. Soc. Bot. 37 (1904) 11; Philip. J. Sc. 2 (1907) Bot. 107, p.p.; KUx. Pfl. R. Heft 38 (1909) 287, p.p.; MEerRR. En. Philip. 1 (1923) 138, p.p. — Fig. 118, 124j-k, 125. Rhizome shortly creeping, woody. Stems loosely tufted, triquetrous, smooth, (15—)60-100 cm by 1-3 mm, surrounded below the leaves by bladeless, brownish to blackish sheaths and their fibrous remains. Leaves subcoriaceous, mostly basal but some higher on the stem, as long as or longer than the stems, linear, long-attenuate, flat or with slightly revolute margins, scabrid on the margins and the nerves above, 5-15 mm wide; sheaths blackish-nerved. Inflorescence a decompound, erect, interrupted panicle occupying the upper half of the stem; secondary panicles 3-8, single at the nodes, erect, oblong-pyramidal, rather loose to dense, the upper approximate, the lower distant on long-exserted peduncles, up to 10 cm long; rachis sparsely hispidulous on the angles. Lower bracts foliaceous, exceeding the inflorescence, long- sheathing, their sheaths glabrous, or hispidulous at the mouth; upper bracts much reduced; brac- teoles with a filiform, usually long and recurved awn. Spikelets divaricate, androgynous, rather dense, (S—)10-20 mm long, the ¢ part as long as to (usually) much longer than the 9 part. Glumes thinly membranous, ovate or ovate-lanceolate, acute or obtuse, sometimes slightly emarginate, nerved, glabrous, stramineous to brownish, 2-3'/, mm long, the midnerve excurrent into an antrorsely scabrid, recurved, up to 31/, mm long awn. Utricles inflated, obsoletely trigonous, broadly ellipsoid to subglobose, patent, straight or almost so, subcoriaceous, strongly many-nerved (nerves c. 10 on each face), glabrous, scarcely stipitate, abruptly beaked, olive-brown, 31/,-5 by 17/,-2'/; mm; beak straight or slightly bent, 1'/,-2 mm long; mouth oblique, not bidentate. Nut triquetrous, with very prominent angles and concave faces, broadly ellipsoid-obovoid to sub- globose, pyriform, or rhomboid, scarcely stipitate, sometimes slightly discoid-annulate at the apex, often with a curved or straight, dark brown to blackish rostrum, 2-3 by 1!/,-2 mm. Style-base pyramidally thickened, subpersistent on the nut. Stigmas 3. Distr. Widely distributed, from Ceylon and India through Burma, Thailand, Indo-China and S. China to Queensland, New Caledonia, Caro- lines, Solomon and Fiji Is.; in Malesia: Malay Peninsula, W. Java (only once collected near Tjikoya by ZOLLINGER), Borneo, SE. & N. Celebes (2 collections), Philippines (Luzon, Samar, Palawan), New Guinea. Ecol. Moist places in forests, along streams, at low and medium altitudes, up to 1000 m. Vern. Bundung, sesayak, M. Notes. Very variable, especially in the length of the awns on bracteoles and glumes. When KoyAMA, Micronesica 1 (1964) 108-109, says that in C. indica the glumes gradually taper to the cuspidate apex without any conspicuous awn, in contradistinction to the Micronesian C. fuirenoides GAUDICH. with truncate or shallowly emarginate glumes with a long, scabrous awn, he cannot have had true C. indica before him. Segregation of C. dietrichiae (= C. indica var. laetebrunnea) on account of the darker glumes appears to be impossible. See RAYMOND 1959, /.c., who is in all probability right in supposing that in general NeLMEs referred young plants to C. indica, and those with mature fruits, in which the glumes have often become darker, to C. dietrichiae. 7. Carex lamprochlamys S. T. BLAKE, J. Arn. Arb. 28 (1947) 104, f. 2A; NeLMeEs, Kew Bull. (1949) 1979] Cy PERACEAE—II (Kern & Nooteboom) Fig. 122. Utricles and nuts of Carex. Species numbered as in the text (¢3a WomERSLEY 5122, 43b ANU 614, 44 BRAss 4867, 45 MEIJER 6656, 46 BRASS 9803, 47 LAE 61655, 48 SCHODDE 1992, 49 EymMa 4709, 50 KOSTERMANS 14005, 5/7 SINCLAIR 9797, 52 JERMY 4634). All x 10. 128 FLORA MALESIANA [ser. I, vol. 91 379, incl. var. diplocolea NELMES. — C. rafflesiana var. continua [non (CLARKE) KUK.] KUK. Bull. Jard. Bot. Btzg III, 16 (1940) 315. — C. continua (non CLARKE) S. T. BLAKE, J. Arn. Arb. 28 (1947) 104, p.p. — C. papuana NELMES, Kew Bull. (1949) 379; Reinwardtia 1 (1951) 314. — C. tyttholepis NELMES, Kew Bull. (1949) 388; Reinwardtia 1 (1951) 320. — Fig. 118, 124h-i. Rhizome short. Stems tufted, rather stout, triquetrous, smooth, 50-100 cm by 3 mm at the base. Leaves subcoriaceous, mostly basal but some- times one halfway up the stem, shorter than the stems, linear, long-attenuate, with strongly revo- lute margins, vesiculose-asperous by whitish vesicles especially on the upper surface, (3-)7- 10 mm wide; lower sheaths reddish brown, densely but minutely hispidulous, eventually fraying into fibres. Inflorescence a slender, interrupted or sub- continuous, decompound panicle up to 50 cm long; partial panicles 4-12, at up to 8 nodes, mostly single, sometimes binate, oblong-lanceolate, erect, rather dense, 3-9 cm long, upper ones approximate, lower ones distant on slender, rigid, scabrid peduncles more or less exserted from the sheaths; rachis hispid. Lower bracts foliaceous, as long as or longer than the inflorescence, long- sheathing, upper much reduced; bracteoles glumi- form, with hispidulous awns. Spikelets numerous, androgynous, oblong or ovoid, 5-12 mm long, the 3 part shorter than to about as long as the ° part. Glumes broadly ovate, often broader than long, obtuse, translucent, glabrous or the lower ones hispidulous, pale fulvous to dark reddish, finely several-nerved, (3/,-)1-17/; mm, the midnerve excurrent into a strong, antrorsely scabrid, up to 2 mm long awn. Urtricles trigonous, not inflated, ellipsoid or slightly obovoid, membranous, patent, straight or more or less recurved, with (3—)4-6 strong nerves on each face, glabrous and smooth, or hispidulous on the margins in the upper part, subabruptly beaked, light green to strami- neous, (2'/,-)3-4 mm by 1-1!/, mm; beak slender, compressed, somewhat tapering, straight or slightly curved, scabrid on the margins, shorter than to about as long as the body of the utricle, (1-)1!/, mm long; mouth bidentate, not or scarcely oblique. Nut ellipsoid or slightly obovoid, triquetrous with shallowly concave sides, brown, 1!/,-2 by 9/,,.— 1 mm. Style-base slightly thickened. Stigmas 3. Distr. Malesia: throughout New Guinea, also known from some adjacent islands. Ecol. Primary forests (often Fagaceae forests), secondary growths, rarely in wet grassland, 850- 2700 m. Vern. Anifi, Wapi lang. Note. NeLMes distinguished C. papuana from C. lamprochlamys by the narrower leaves and the darker glumes and utricles of the former. To me those differences are insufficient for specific separation. The type collection of C. tyttholepis I cannot distinguish from C. lamprochlamys. 8. Carex nodiflora Boeck. Bot. Jahrb. 5 (1884) 516; KUx. Pfl. R. Heft 38 (1909) 288; Philip. J. Sc. 6 (1911) Bot. 61; Merr. En. Philip. 1 (1923) 140; NeLMes, Reinwardtia 1 (1951) 260; ibid. 2 (1954) 374. — C. cumingii [Boott’ in VIDAL, Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 286, nomen]; ex CLARKE, J. Linn. Soc. Bot. 37 (1904) 11; Philip. J. Sc. 2 (1907) Bot. 107. — C. vulcanica ELMER, Leafl. Philip. Bot. 10 (1938) 3526. — Fig. 118. Rhizome shortly creeping, woody, covered with the fibrous remains of old scales. Stems tufted, triquetrous, smooth, surrounded below the leaves by a few bladeless, brown sheaths or their fibrous remains, 50-70 cm by 1!/,-2 mm. Leaves basal, sometimes 1-2 higher on the stem, much exceeding the stems, rigid, flat, long-attenuate, scabrid on the margins in the upper part, 7-16 mm wide; cauline leaves long-sheathing. Inflorescence a_ slender, compound panicle, continuous above, interrupted below, 15-35 cm long; secondary panicles 4-6, single at the nodes, erect, spiciform or the middle ones again branched, oblong, dense, 2-6 by 1-2 cm; lower peduncles more or less exserted from the sheaths, smooth, or scabrid above; rachis hispidu- lous especially above. Lower bracts foliaceous, overtopping the inflorescence, long-sheathing, upper much reduced. Spikelets numerous, androgynous, ovoid, oblong, or subglobose, 5-10 mm long, the 2 part rather longer than the ¢ part. Glumes ovate-lanceolate, acutish or obtuse, membranous, nerved, shortly setulose above or glabrescent, sparsely ciliolate, greenish white or light brown with whitish hyaline margins, 2 mm long, the midnerve excurrent in a smooth or his- pidulous, '/, mm long awnlet. Utricles much overtopping the glumes, trigonous, ellipsoid- rhomboid, with prominent angles and flattish faces, membranous, suberect, many-nerved, straight, whitish setulose (except at the base), scarcely stipitate, rather abruptly beaked, finally dark brown, 31/,-4'/, by 11/,-2 mm; beak oblong- conical, bidenticulate, with slightly oblique mouth, 1 mm long. Nut triquetrous, ellipsoid-rhomboid, sessile, narrowly discoid-annulate at the apex, ferrugineous to blackish, 2!/,—2'/, by 11/,-13/, mm. Style-base pyramidally thickened, persistent on the nut. Stigmas 3. Distr. Malesia: Philippines (Luzon; according to MERRILL, /.c., also in Alabat and Mindanao: Agusan). Ecol. Primary forests at low and medium alti- tudes. Note. According to NELMEs (1951: 262) in this species it is the style-base which is discoid-annulate, not the apex of the nut. I do not see any difference with the nut and style-base of the allied species. 9. Carex sarawaketensis KUx. Bot. Jahrb. 69 (1938) 262; ibid. 70 (1940) 464, incl. var. minor; NELMES, Reinwardtia 1 (1951) 316. — C. melano- 1979] E VIJSMA ’78§ Fig. 123. Utricles and nuts of Carex. Species numbered as in the text (53 VAN STEENIS 4624, 54 DE WILDE c.s. 13329, 55 Brass 9284, 56 Brass 9515, 57 EyMA 3862, 58 herb. VAN Soest 88, 59 VAN STEENIS 9804, 62 HOOGLAND & SCHODDE 7470, 63 JEswIET 39, 64 DE WILDE c.s. 13323, 65a KOORDERS 43403, 66 BRAsS 9539). All x 10. FLORA MALESIANA [ser. I, vol. 91 phora S. T. BLake, J. Arn. Arb. 28 (1947) 106, f. 2B. — Fig. 118. Rhizome emitting slender stolons covered with brown or blackish scales. Stems erect or somewhat curved, solitary or tufted, slender, obtusely trigo- nous, smooth or scaberulous at the top, 3-40 (—60) cm by '/,-1 mm, the base surrounded by the fibrous, reddish-black remains of old leaf-sheaths. Leaves basal, rarely 1 higher on the stem, rigid, shorter than the stems, linear, long-attenuate, flat or with revolute margins, conspicuously keeled, scabrid on the margins and keel in the upper part, 2-4 mm wide; lower sheaths fuscous to blackish- purple. Jnflorescence a decompound, narrow panicle interrupted at the base, 10—20(-30) cm long; partial panrcles 3-14, at 2-8 nodes, single and binate, oblong, dense, 1-5 cm long, upper ones approximate, lower on slender, erect, scabrid, up to 8 cm long peduncles exserted from the sheaths. Lower bracts foliaceous, not or scarcely over- topping the inflorescence, long-sheathing, upper ones much reduced; bracteoles glume-like, hispidulous, awned; rachis hispid. Spikelets androgynous, suberect or appressed, oblong, 4-10 by 2-2!/, mm, the ¢ part inconspicuous, mostly shorter than the 2 part. Glumes ovate, membranous, acute, muticous or minutely mucro- nulate, rarely awned, slenderly nerved, glabrous or the lower ones minutely hispidulous, blackish- fuscous with white-hyaline margins, 2-3 mm long. Utricles trigonous, not inflated, ellipsoid, narrowed at both ends, suberect, membranous, straight, obscurely nerved (only 2 nerves more prominent), glabrous at the base, otherwise densely scabrid- pubescent, gradually narrowed into the beak, blackish-fuscous, 23/,-4'/, by 4/,.-11/; mm; beak straight, compressed, deeply bidentate (teeth 1/,-?/, mm) but often originally oblique, scabrid on the margins, 1—13/, mm long, the mouth with whit- ish-hyaline margins. Nut triquetrous with shallowly concave sides, ellipsoid or broadly ellipsoid, stipitate, conspicuously apiculate, stramineous- brown, 11/,-13/, by 4/;,-1 mm. Style scabrid, not or scarcely incrassate at the base. Stigmas 3. Distr. Malesia: New Guinea (W. New Guinea: Carstensz Mts, Mt Wilhelmina, Hellwig Mts; NE. New Guinea: Mt Sarawaket, Lake Naho). Ecol. Wet places in alpine grassland, steep mountain slopes, open places in mountain forests, moist grassy cliffs, 2700-3950 m. 10. Carex stramentitia Bootr ex Boeck. Linnaea 40 (1876) 351; CLarKE, FI. Br. Ind. 6 (1894) 717; J. Linn. Soc. Bot. 37 (1904) 9; Kix. Pfl. R. Heft 38 (1909) 264; Camus, FI. Gén. I.-C. 7 (1912) 188; NELMES, Kew Bull. (1950) 191; Reinwardtia 1 (1951) 267; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 120; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 45, 99, f. 28 (map); Dansk Bot. Ark. 23 (1965) 254; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — Fig. 118, 124m-n. Rhizome shortly creeping, stout, woody. Stems loosely tufted, trigonous, scaberulous below the nodes to almost smooth, 30-130 cm by 11/,-3 mm, the base clothed with fuscous, bladeless sheaths and comose by their fibrous remains. Leaves subcoriaceous, subbasal and 1-2 higher on the stem, exceeding the stems, linear, long-attenuate, flat or with revolute margins, scabrid on the mar- gins in the upper part, pale green, 5—-10(-15) mm wide. Inflorescence a pale, compound, erect, much interrupted, narrow, 10-40 cm long panicle; secondary panicles 2-4, single at the nodes, erect, lanceolate or oblong, very dense, distant, up to 10 cm long; peduncles smooth or scaberulous, the lower ones much exserted from the sheaths, the upper ones scarcely so; rachis hispid. Lower bracts foliaceous, exceeding the inflorescence, long- sheathing, hispidulous in front or at the mouth, upper reduced. Spikelets androgynous, sessile, suberect, 5-15 mm long, the 2 part few-flowered, rather shorter than the ¢ part. Glumes oblong or oblong-ovate, thinly membranous, obtuse or slightly emarginate, slenderly nerved, glabrous or sparsely hispidulous, not ciliate, pale stramineous to whitish, 2—23/, mm long, the midnerve excurrent into an antrorsely scabrid, 3/,-2 mm long awn. Utricles distinctly trigonous, rhomboid-ellipsoid, with prominent angles and flattish faces, mem- branous, not inflated, patulous, many-nerved (nerves 5-7 on each face), glabrous, straight to slightly recurved, curved-tapering below into a cuneate basal part, suddenly narrowed above into the beak, greenish to light brown, 4-5 by c. 11/, mm; beak straight or slightly curved, often somewhat inflated at the base, glabrous or very sparsely scabrid, 11/,-2 mm long; mouth dorsally very oblique, not bidentate. Nut trigonous, rhomboid- eilipsoid, erostrate, curved-tapering below into a stout cuneate part, with prominent angles and concave faces, brown, 2!/,-3 by 1'/,-14/; mm. Style-base pyramidally thickened, persistent on the nut. Stigmas 3. Distr. India, Lower Burma, S. China (Kwei- chow), Thailand, Laos, Tonkin, Annam; in Malesia only known from a single collection from W. Java (Krawang: Pléréd near Purwakarta); see map in RAYMOND, 1959, /.c. Ecol. In Krawang abundant in jungle at low altitude under seasonal climatic conditions. Vern. Lilisungan, S. Note. In spite of its different facies because of the stiff, linear leaves, C. stramentitia is very near C. commixta, which it closely resembles in the size and shape of the utricles. According to NELMES the leaves of C. stramentitia can reach a width of 20 mm. 11. Carex vesiculosa Boort, III. 3 (1862) 107, t. 323; Boeck. Linnaea 40 (1876) 345; CLARKE, FI. Br. Ind. 6 (1894) 717, incl. var. paniculata CLARKE; KUx. Pfl. R. Heft 38 (1909) 283, f. 43, inc’ var. 1979] - CYPERACEAE—II (Kern & Nooteboom) 131 Fig. 124. Inflorescence (nat. size), spikelets and (occasionally) venation (x 2). — 3a. Carex cruciata WAHLENB. var. cruciata, a-b. — 3b. var. rafflesiana (BootT) Noor., c—d. — 5. C. horsfieldii Boott, e-g. — 7. C. lamprochlamys S. T. BLAKE, h-i. — 6. C. indica L., j-k. — 13. C. hypolytroides Rip. |. — 10. C. stramentitia Boott, m-n. — 11. C. vesiculosa Boott, o—p. (a—b CHEW, CORNER & STAINTON 1706, ¢ VAN OosTSTROOM 13154, d VERHEIJEN 2585, e-g NGF 21261, A-i Brass 24684, j-k RipLey 15721, / BUNNEMEJJER 9820, m—n BAKHUIZEN VAN DEN BRINK 6565, 0-p VAN STEENIS 4140). 132 FLORA MALESIANA [ser. E voli 9" congesta KUK.; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 68; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 492. — C. impunctata Bootr, Ill. 3 (1862) 107, t. 326 (pl. depauperata); BoECcK. Linnaea 40 (1876) 342. — C. gembolensis CLARKE, J. Linn. Soc. Bot. 37 (1904) 10; Nexmes, Rein- wardtia 1 (1951) 294, incl. var. crebra NELMES; ibid. 2 (1954) 376. — C. rhizomatosa var. impunctata Kuk. Pfi. R. Heft 38 (1909) 291. — C. vesiculosa var. latifolia KUK. in Hochr. Candollea 6 (1931) 341. — Fig. 118, 1240-p. Rhizome shortly creeping, woody, covered with the fibrous remains of decayed sheaths. Stems loosely tufted, stiff, trigonous, smooth, 30-150 cm by 2-3 mm, surrounded below the leaves by blade- less, reddish or fuscous sheaths and their fibrous, often reticulate remains. Leaves coriaceous, basal and 1-2 higher on the stem, usually much shorter than the stems, narrow, long-attenuate, keeled, with strongly revolute margins when dry, scabrous on the margins, often vesiculose-asperous above especially towards the apex, 2-8 mm wide. /nflores- cence a decompound, interrupted, ferrugineous, 15-40 cm long panicle; secondary panicles 5-10, at 4-6 nodes, lowest 1-2 often single, remainder usually unequally binate, erect, often with nodding top, oblong-pyramidal, dense or very dense, rarely loose, up to 10 by 5 cm, upper ones approxi- mate, lower distant on exserted, smooth or scabrid, up to 15 cm long peduncles; rachis hispidulous on the angles. Lower bracts foliaceous, narrow, shorter than the inflorescence, long-sheathing, the upper ones much reduced; bracteoles pilose, excurrent in a curved, hispidulous awn. Spikelets numerous, androgynous, obliquely patent, 5- 15 mm long, the ¢ part in the longer spikelets much longer than the few-flowered 2 part. Glumes lanceolate-ovate, acutish, sometimes lightly emar- ginate, membranous, translucent, glabrous or somewhat hispidulous, ferrugineous or castaneous, 2'/,-4'/, mm long, the midnerve in the upper glumes excurrent in a hispid, up to 13/, mm long awn. Utricles triquetrous, narrowly ellipsoid, sub- coriaceous, not inflated, obliquely erect, straight or but slightly recurved, slenderly or obscurely nerved, rather densely setulose in the upper 3/4, scarcely stipitate, rather abruptly beaked, reddish with castaneous flecks, 3-5 by 1—13/, mm; beak slender, scabrid on the margins, mouth not oblique, 11/,— 2 mm long. Nut triquetrous, ellipsoid or slightly obovoid, with concave faces, shortly stipitate, abruptly beaked, 1!/,-21/, by 1-11/, mm. Style- base scarcely thickened. Stigmas 3. Distr. Nepal to Thailand; in Malesia: Java (in W. Java on Mt Papandajan, not rare in the moun- tains of Central and E. Java), Lesser Sunda Is. (Bali, Lombok, Flores, Timor). Ecol. In dry grass-lands, open stony places, mountain scrub, Casuarina forests, 1200-3500 m. Notes. In old specimens the leaves are often less asperous because the vesiculose (bulbous- based) hairs have broken off. The former presence of vesicles is indicated by small, circular scars. If C. gembolensis is specifically distinct from C. vesiculosa, its correct name is C. impunctata Boott, based on ZOLLINGER 2563 from Java. This collection NeLMes rightly included in C. gembolen- sis. Var. congesta KUK. (= C. gembolensis var. crebra NELMES) has no taxonomic value. 2. Section Japonicae KUK. Pfl. R. Heft 38 (1909) 252. — Sect. Paciricae OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 458; NELMEs, Reinwardtia 1 (1951) 329; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 52. — Sect. Indicae subsect. Japonicae (KUK.) Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 152. Type species: Carex nikoensis FRANCH. & SAVAT. 12. Carex satzumensis FRANCH. & SAVAT. En. Pl. Jap. 2 [(1877) 132, nomen] (1878) 558; FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 8 (1896) 259 (t. 4, f. 1 ut C. nikoensis); AKIYAMA, J. Fac. Sc. Hokk. Imp. Un. V, 2 (1932) 84, f. 36; Car. Far Fast. Reg. Asia (1955) 135, t. 121; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 458; NELMEs, Reinwardtia 1 (1951) 330; KoyAMA, Nat. Canad. 82 (1955) 197; Contr. Inst. Bot. Un. Montréal n. 70 (1957) 10; YosHikawa, Ic. Jap. Carex 2 (1958) 236, t. 118; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 52. — C. nikoensis FRANCH. & SAvAT. En. Pl. Jap. 2 [(1877) 132, nomen] (1878) 558; KUx. Pfl. R. Heft 38 (1909) 252, f. 38A-E (‘nikkoensis’); Philip. J. Sc. 6 (1911) Bot. 59; Me_rr. En. Philip. 1 (1923) 140. — C. contracta Boeck. Cyp. Nov. 2 (1890) 34. — Fig. 119. Rhizome long-creeping, clothed with large, brown scales. Stems erect, triquetrous, ribbed, smooth, or scabrid just below the inflorescence, 3-20(-30) cm by 1-1'/, mm, surrounded at the base by the fibrous remains of old leaf-sheaths. Leaves subbasal, rigid, flattish to conduplicate, longer than the stems, often recurved, scabrid on the margins and on the upper surface towards the long-attenuate apex, 2-5 mm wide. Inflorescence 1979] Cy PERACEAE—II (Kern & Nooteboom) 133 Fig. 125. Carex indica L. a. Habit, x */3, b-c. glumes, d. fruit in utricle, e. fruit, all x 7 (SCHODDE 2972). simple, spiciform, sometimes slightly branched at the base, cylindrical-conical, acute, 2-8 cm by 1—2'/, cm at the base; rachis smooth. Lower bracts subulate to subfoliaceous, shorter than the in- florescence, not sheathing, upper filiform. Spikelets 12-numerous, androgynous, - sessile, patent, ovate to oblong, densely flowered, lower 5-14 mm, upper 3-7 mm long, all the lateral ones arising from a + utriculiform, gibbous prophyll containing a @ flower. Glumes lanceolate-ovate, acute to obtuse, glabrous, pale, slenderly nerved, 2-3 mm long, the midrib sometimes slightly excur- rent. Urricles oblong-ellipsoid, obtusely trigonous, hardly inflated, glabrous, slenderly plurinerved, greenish, 2!/,-3'/, by 3/,-1 mm, rather gradually narrowed into a long, conical, bilobed beak with 134 FLORA MALESIANA [ser. 1, velo" oblique mouth. Nut oblong-obovoid, triquetrous, shortly beaked, dark brown, 1'/,-1'/, mm long. Style pyramidally thickened at the base. Stigmas 3. Distr. Japan, Formosa, Tonkin; in Malesia: Philippines (Luzon: Benguet). Ecol. Along trails in the mossy forest, at c. 2300 m, also along roadsides under pine-trees at 400 m. Notes. Readily recognizable by its spike-like inflorescence with non-sheathing bracts. Carex nikoensis FRANCH. & SAVAT., with slightly larger spikelets in denser spikes and somewhat longer beak of the utricle than in typical C. satzu- mensis, was already reduced to C. satzumensis by FRANCHET (1896). The specific epithet is frequently spelt “satsumen- sis’, but the spelling ‘satzumensis’, employed when the species was validly published, must be retained. In an abnormal inflorescence of SANTOS 5776 from Luzon I found in all branches between the fertile cladoprophyll and the normal utricles some deeply split utricles containing one or some ¢ flowers besides the 2 one, a situation normal in Schoenoxiphium and Kobresia. 3. Section Hypolytroides NELMES, Kew Bull. (1951) 121; Reinwardtia | (1951) 246; RAymM. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 21. Type species: Carex hypolytroides RIDL. 13. Carex hypolytroides Rip_. J. Fed. Mal. St. Mus. 8 (1917) 124; NeL_mes, Reinwardtia 1 (1951) 246; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 96; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 21, f. 3, 102, f. 31 (map). — C. hypolytroides Gross & Mattr. Notizbl. Berl.-Dahl. 14 (1938) 190. — C. hypolytropsis Gross in Fedde, Rep. 50 (1941) 213. — Fig. 119, 1241, 126. Rhizome woody, emitting strong stolons covered with lanceolate, fuscous sheaths. Stems rigid, erect, rather acutely trigonous, smooth, 100-250 cm by 3-5 mm, surrounded at the base by a few reddish brown, bladeless sheaths. Leaves at regular intervals throughout the stem, shorter than the stem, stiff, flattish, or with revolute margins when dry, smooth, or scabrid on the margins, sparsely to rather densely covered with long, pale hairs beneath, 4-8 mm wide; sheaths smooth, hairy above, membranous in front, with concave mouth; ligule elongate, hairy, ferrugineous. Inflorescence a compound, interrupted panicle, 25-40 cm long; secondary panicles 5-8, erect, broadly pyramidal, single at the nodes, or lowest sometimes binate, rather loose, upper approximate, lower distant, on exserted, hispidulous peduncles with patent branches. Lower bracts foliaceous, slightly exceeding the inflorescence, long-sheathing, upper reduced. Spikelets usually unisexual; 2 ones numerous, shortly cylindrical, very dense, 5-8 by 31/,-4'/, mm, sometimes with a few ¢ flowers at the top; gd ones inconspicuous, few, lateral, 1-5 just below some of the terminal 2 spikelets, sessile, ellipsoid, 4 by 1-2 mm. Cladoprophylls of branches and peduncles utriculiform, hairy, those of the 3 spikelets sometimes enclosing a $ flower, those of the partial panicles ocreiform. Glumes thinly membranous, ovate-lanceolate, acute, hairy, slen- derly nerved, with narrow whitish-hyaline margins and shortly excurrent midnerve, 2—23/, mm long, in ripe spikelets almost completely hidden among the utricles. Utricles obtusely trigonous, obovoid, not or but slightly inflated, horizontally patent, straight, glabrous or very sparsely hispidulous, slenderly nerved (2 nerves more prominent), fuscous, densely reddish glandular-spotted, 11/,— 1?/, by c. 1 mm, subabruptly narrowed into a very short, bidenticulate beak. Nut triquetrous, obovoid, filling the utricle, sessile, brown, whitish papillose, 11/,-1'/, by °/;>-1 mm. Style-base thickened. Stigmas 3. Distr. Tonkin, Annam; in Malesia: Central Sumatra (Mt Kerintji), Borneo (Mt Kinabalu: Lumu-lumu, Mesilau Cave). See the distribution map by RAYMOND, /.c. Ecol. In wet spots in forests, on Mt Kerintji between 2200 and 2750 m, on Mt Kinabalu at 1800 m. Notes. The infructescence so strongly recalls a Hypolytrum that RipLey and Gross independently of each other chose the epithet hypolytroides for it. The ¢ spikelets are inserted laterally, as they are borne from a utriculiform prophyll, and there- fore cannot have degraded from an originally terminal position as was supposed by NELMES, Reinwardtia 1 (1951) 248. Carex hypolytroides is closely related to C. moupinensis FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris II, 10 (1888) 102; ibid. III, 8 (1896) 257, t. 7 f. 2; CLARKE, J. Linn. Soc. Bot. 36 (1904) 209; Kuk. Pfl. R. Heft 38 (1909) 289; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 21, f. 2. — Homalostachys sinensis Boeck. Cyp. Nov. 1 (1888) 38, non C. chinensis Retz. — Scleria sinensis H. Preirr. in Fedde, Rep. 26 (1929) 263, only known from China (Hupeh, Szechuan, Yunnan). 1979] CY PERACEAE—II (Kern & Nooteboom) 135 Res Fig. 126. Carex hypolytroides Rv. a. Habit, x */3, b. terminal ? and lateral 3 spikelet, x 3, c-d. fruit in utricle, e. fruit, both x 13 (JAcoss 4332). the slightly larger (2 by 1'/; mm), much inflated utricles truncate or depressed at the top, and the nut not filling the utricle. The latter species is much lower, with narrower leaves, and usually all the spikelets of the terminal partial panicle 3; it is especially characterised by 136 FLORA MALESIANA [ser. I, vol. 91 4. Section Mapaniifoliae NeELMEs & AiRY SHAW in Hook. Ic. Pl. 35 (1943) t. 3434; NeELMgs, Reinwardtia 1 (1951) 248; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 27; 93, f. 23 (map). — Sect. Indicae subsect. Scaposae Kux. Pfl. R. Heft 38 (1909) 285, p.p. Type species: Carex helferi BOECK. 14. Carex helferi Boeck. Linnaea 40 (1876) 365; CLARKE, FI. Br. Ind. 6 (1894) 714; Ktx. Pfl. R. Heft 38 (1909) 286; NeELMEs in Hook. Ic. Pl. (1947) t. 3468; Kew Bull. (1950) 189; Reinwardtia 1 (1951) 248; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 99; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 30; Dansk Bot. Ark. 23 (1965) 252; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 491. — C. mapaniifolia RipL. J. Fed. Mal. St. Mus. 10 (1920) 124; Ne_tmes, Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 98; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 30. — Fig. 119. Rhizome short, stout, woody, clothed with some brown bladeless sheaths or their fibrous remains. Stems central, erect, scapiform, trigonous or sub- terete, smooth, or scabrid on the angles, (5—)20— 40 cm by 1-2 mm, the base enveloped by some brown, infundibuliform, up to 10 cm long sheaths. Leaves overtopping the stems, broadly linear, conduplicate below (narrowed into a pseudo- petiole), otherwise flat, long-acuminate, minutely scabrid on the margins in the upper part, with the midnerve and 2 mid-lateral nerves prominent, 11/,-31/, cm wide; ligule elongate-triangular, brown-bordered, 1!/,-2 cm long. Inflorescence paniculate, consisting of (2—)3-7 partial inflores- cences, up to 25 cm long; partial inflorescences erect, oblong-ovoid, very dense, head-like, upper approximate, lower distant, single at the nodes, with hispid rachis, 2—31/, by 1—2!/, cm, on exserted, smooth or finely scabrid peduncles. Bracts sheath- like, ampliate, infundibuliform, short-bladed, light brown or spadiceous. Spikelets densely crowded, androgynous, ovoid, patent, 6-10 by 5-8 mm, the ¢ part usually much longer than the few-flowered 92 part. Glumes thinly membranous, oblong-ovate, obtuse to truncate-bilobed, many- nerved, glabrous or hispidulous, pale brown with broad, whitish hyaline margins, 2-4 mm long, the midnerve excurrent in a 1-4 mm long, scabrous, often recurved awn. Utricles trigonous, ellipsoid, with shallowly concave faces, patulous, glabrous at the base, hispidulous above, shortly stipitate, strongly 6—-8-nerved on each face, stramineous to brown, 6—7 by 1'/,-1*/,; mm (the long beak included); beak hispidulous, curved, bulbous at the base (to hold thickened style-base), nearly linear above, with very oblique mouth (its base 11/,- 13/, mm from the apex), 3—-3!/, mm long. Nut tri- gonous, ellipsoid or slightly obovoid, scarcely stipitate, shortly beaked and suddenly expanded into the conical style-base, dark brown with pale angles, 21/,-23/, by 11/,-13/, mm. Style-base pyramidally thickened, subpersistent on the nut. Stigmas 3. Distr. Lower Burma, SE. & Peninsular Thailand; in Malesia: W. Java (Priangan). Wrongly recorded for Sumatra and Borneo by NELMES (1950, p. 100). Ecol. In forests, 1100 m. Notes. This is the only Malesian member of the wide-leaved sect. Mapaniifoliae, which section appears to be almost confined to the Indo- Chinese Peninsula, where it is represented by several species. The few Javan specimens hitherto collected are vegetatively much less developed than those from the continent, but agree with them in floral and fruit characters. By its broad leaves, scapiform flowering stems, and shape of the inflorescence Carex helferi deceivingly resembles Hypolytrum humile (STEUD.) Boeck. It can be distinguished by the long-beaked utricles, the 3 stigmas, and the trigonous nuts. 5. Section Polystachyae [TUCKERM. En. Meth. (1843) 10, nomen (‘Polystachae’)|; ex KUx. Bot. Jahrb. 27 (1899) 517, quoad basion.; Pfl. R. Heft 38 (1909) 257; NeLMgs, Reinwardtia 1 (1951) 322; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 49. — Sect. Acrarrhenae Fries Sippe Longebracteatae PAX in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 124. — Sect. Polystacheae CLARKE [ser.] Longispicae CLARKE, J. Linn. Soc. Bot. 37 (1904) 4, p.p. — Sect. Extensae Frits subsect. Baccantes KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 151. Type species: Carex myosurus NEES. 1979] 15. Carex baccans NEEs in Wight, Contr. Bot. Ind. (1834) 122; Kuntu, En. 2 (1837) 513; Boortt, Ill. 2 (1860) 83, t. 234-236, 238, 239; Boeck. Linnaea 40 (1876) 339; O. K. Rev. Gen. Pl. 2 (1891) 747, incl. var. nigra O. K.; CLARKE, Bot. Mag. III, 49 (1893) t. 7288; Fl. Br. Ind. 6 (1894) 722; J. Linn. Soc. Bot. 37 (1904) 14, incl. var. siccifructus CLARKE; Philip. J. Sc. 2 (1907) Bot. 108; Kix. Pfl. R. Heft 38 (1909) 258, f. 39E-H; Philip. J. Sc. 6 (1911) Bot. 59; Koorp. Exk. Fl. Java 1 (1911) 211; Camus, FI. Gén. I.-C. 7 (1912) 185; Merr. En. Philip. 1 (1923) 136; Riv. Fl. Mal. Pen. 5 (1925) 184; Onwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 462; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 68; NELMES, Kew Bull. (1950) 194; Reinwardtia 1 (1951) 322; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 50; Koyama, J. Fac. Sc. Tokyo Un. III, 8 (1962) 216; Dansk Bot. Ark. 23 (1965) 257; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 493; STEEN. Mt. FI. Java (1972) pl. 15-1. — C. curvirostris KUNZE, Suppl. Riedgr. (1840-50) 79, t. 20; Mia. Fl. Ind. Bat. 3 (1856) 350; CLARKE, J. Linn. Soc. Bot. 37 (1904) 14; Steup. [in Zoll. Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 207 (‘recurvirostra’). — C. javanica BOECK. Cyp. Not. 1 Cy PERACEAE—II (Kern & Nooteboom) a7 (1888) 43. — C. walkeri (non ARN. ex Bootr) Kix. Pfl. R. Heft 38 (1909) 546; cf. BAcK. Brittonia 3 (1938) 76. — Fig. 119, 127, 131i. Rhizome short, stout, woody. Stems usually robust, loosely tufted, erect, triquetrous or trigonous, smooth, 60-150 cm by up to 5 mm, the base surrounded by reddish, bladeless sheaths splitting in front into reticulate fibres. Leaves all over the stem, often overtopping the inflorescence, long-attenuate, coriaceous, flat or with revolute margins, asperous on the upper surface in the apical part, with scabrous margins, 5-18 mm wide; sheaths long, often splitting up in front. Jnflores- cence paniculate, compound or decompound, oblong, occupying '/,;—'/, of the stem, erect or somewhat nodding at the top; partial panicles 5-8, single at the nodes, much branched, upper approxi- mate, lower distant on long-exserted, smooth or scabrid peduncles. Lower bracts usually much overtopping the inflorescence, foliaceous, long- sheathing, upper much reduced. Spikelets very numerous, androgynous, suberect to patent, 21/,-8 cm long; 9 part cylindrical, densely flowered, usually longer than the slenderer 3 part. Glumes ovate or oblong-ovate, acute to subobtuse, glab- Fig. 127. Carex baccans NEES in pioneer tufts with sand-binding capacity in the sand plain and dunes of volcanic ash of the Tengger Sandsea, East Java, c. 1900 m altitude (CLAsoN). 138 FLORA MALESIANA [ser. I, vol. 91 rous or hispidulous, strongly nerved, purplish or castaneous with whitish-hyaline margins, muticous or the midrib excurrent in a scabrid mucro, 2'/,-31/. mm long. Urtricles inflated, obscurely trigonous, obovoid to subglobose, patent, sub- coriaceous, strongly nerved, glabrous except for the hispidulous margins at the apex, shining, at first yellowish green, ultimately red and more or less succulent, 31/,—4!/, mm long, abruptly beaked; beak recurved, bidentate. Nut triquetrous, ellip- soid, with flattish or shallowly concave faces, dark brown, 2°/,-3 mm long. Style-base not thickened. Stigmas 3. Distr. Ceylon, India (Sikkim and Khasia), S. China and Formosa, and through Thailand and Indo-China to Malesia; in Malesia: Sumatra, Malay Peninsula (Pahang), Java, Lesser Sunda Is. (Bali, Lombok), Philippines (Luzon), New Guinea. Ecol. In damp thickets, open places in forests, in the mossy forest on the higher mountains, some- times in mountain savannahs, or as pioneer in the Sandsea and on fresh landslides, on volcanoes, talus; (600—)1000-—3300 m. Vern. Ladingan, ria-ria-batu, sukét kérisan, J, ilat beureum, ilateun téki, S; Philip.: gihidsan, Bon., mankat, silak, Ig.; New Guinea: djigudjigufa, Dunantina, djugudjuguha, Asoro: Kefamo, pul, Chimbu: Masue, koimin, Wahgi: Minj. Note. This beautiful species seems to be some- times cultivated as an ornamental in temperate regions; see NICHOLSON, Ill. Dict. Gard. 1 (1885- 89) 267, f. 367). 16. Carex myosurus NEEs in Wight, Contr. Bot. Ind. (1834) 122; KuNTH, En. 2 (1837) 507; STEuD. Syn. 2 (1855) 207; Boorr, Ill. 2 (1860) 82, t. 229, 230, 232; Boeck. Linnaea 40 (1876) 334; CLARKE, Fl. Br. Ind. 6 (1894) 723, incl. var. eminens (NEES) CLARKE; KUK. Pfl. R. Heft 38 (1909) 258; MeErR. En. Philip. 1 (1923) 139; NELMEs, Kew Bull. (1950) 195, incl. var. celebica NELMES; Reinwardtia 1 (1951) 325; ibid. 2 (1954) 377; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 51; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 493; STEEN. Mt. FI. Java (1972) pl. 14-2. — C. eminens NeEs in Wight, Contr. Bot. Ind. (1834) 122. — C. longibracteata StTeuD. [in Zoll. Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 205 (‘longebracteata’), non SCHLEICH. Cat. Pl. Helv. ed. 4 (1821) 11, nomen; Mia. FI. Ind. Bat. 3 (1856) 348, incl. var. major MiQ.; BOECK. Linnaea 39 (1875) 108; CLarkE, J. Linn. Soc. Bot. 37 (1904) 15; KUK. Pfl. R. Heft 38 (1909) 260, incl. f. distans Kix. et f. angustifolia KixK.; Bull. Jard. Bot. Btzg IIT, 16 (1940) 314, incl. var. gigantea KUK.; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 70; NELMEs, Reinwardtia 1 (1951) 327; ibid. 2 (1954) 377. — C. floribunda Boeck. Linnaea 40 (1876) 335. — C. kuntzeana Borck. Cyp. Nov. 1 (1888) 51. — C. composita (non Boott) CLARKE, J. Linn. Soc. Bot. 37 (1904) 14. — Fig. 119. Rhizome short, stout, woody. Stems slender, loosely tufted, erect, trigonous, smooth, (20—)50— 150 cm by up to 4 mm at the base, clothed at the base with fuscous to purplish sheaths or their fibrous, reticulate remains. Leaves mainly subbasal, a few widely spaced on the stem, equalling or exceeding the inflorescence, long-attenuate, coria- ceous, flat or with revolute margins, asperous on the upper surface in the upper part, with scabrous margins, (2—)5-10 mm wide; sheaths long, often reddish, frayed in front into reticulate fibres. Inflorescence from almost simple to decompound, narrow, (10—)25—100 cm long, erect or somewhat nodding at the top; secondary panicles 5-9, single at the nodes, upper approximate, lower distant on exserted, more or less scabrid peduncles, sometimes all reduced to simple spikelets. Lower bracts foliaceous, long-sheathing, exceeding the stem, upper much reduced. Spikelets androgynous, sub- erect to patulous, the longer ones on a plant 3 to 10 cm long, ¢ and 2 parts often about equal in length, but sometimes the terminal spikelet almost wholly $ and some of the lateral almost wholly 2. Glumes oblong or oblong-ovate, acute to obtuse, glabrous, or sparsely hispidulous, slenderly nerved, pale to castaneous, with whitish-hyaline margins, 2'/,-4 mm long, the midrib usually excurrent in a hispidulous awn up to 2 (rarely 3) mm long. Utricles trigonous, ellipsoid or ellipsoid-obovoid, patulous, membranous, slenderly nerved, sparsely to subdensely hispidulous at least towards the apex, straight or slightly bent, pale to castaneous, 3-41/, mm long, subgradually to subabruptly narrowed into a hispidulous-margined, bidentate, 3/,-1 mm long beak. Nut triquetrous, ellipsoid, oblong-ellipsoid, or slightly obovoid, with flattish to slightly concave sides, dark brown, abruptly beaked, 2-3 mm long; stipe and beak straight to bent at the base. Style-base not thickened. Stigmas a Distr. India, Burma, Indo-China; in Malesia: Sumatra (Atjeh, W. Coast Res.), Java, Lesser Sunda Is. (Lombok: Mt Rindjani), Philippines (Luzon: Bontoc), SW. Celebes (Mt Bonthain). Ecol. In open or lightly shaded places, on forested slopes, in mountain scrub, heath forma- tions, a pioneer on sterile crater soil, sometimes dominant; 1700-3300 m. Notes. Carex myosurus in the wide circumscrip- tion here accepted is extremely polymorphous. CLARKE, FI. Br. Ind. 6 (1894) 723, distinguished between typical C. myosurus (from Nilghiri Mts and Coromandelia) and C. myosurus var. eminens (NEES) CLARKE (occurring throughout the Hima- layas), the latter mainly characterized by the shorter and usually broader utricles with often rather deeply bifid beak, although in some Sikkim and Bhotan plants (distinguished by BOECKELER as C. floribunda) the beak is not more notched than in the Nilghiri plants. Carex spiculata BoottT, with narrower leaves, denser spikelets with obliquely ascending utricles and more rigid panicles he 1979] considered specifically distinct. Obviously he had not seen Malesian specimens of C. /ongibracteata Steup. (cf. J. Linn. Soc. Bot. 37, 1904, 15). KUKENTHAL, Pfl. R. Heft 38 (1909) 259, distin- guished var. eminens by the shorter and broader utricles and the usually fuscous glumes, and var. floribunda (BoECK.) Kix. by its profusely branched inflorescences; C. spiculata was reduced to sub- specific rank. On the other hand, C. longibracteata was upheld as a species. As in numerous Indian myosurus specimens the inflorescence is paniculate or even spicate (see KUKENTHAL’s description!) it is clear that C. longibracteata cannot be separated from C. myosurus on account of the characters used in KUKENTHAL’s key (“‘inflorescentia subdepanicu- lata, spiculae numerosae”’ in C. myosurus, versus “inflorescentia paniculata rarius spicata, spiculae haud numerosae” in C. /ongibracteata). No more can I understand in what way NELMES Cy PERACEAE—II (Kern & Nooteboom) 139 distinguished between C. myosurus and C. longi- bracteata, for the former is said to have leaves 5—10 mm wide and secondary panicles composed of 3 to rather numerous spikelets, whereas the latter should differ in having leaves 2-6 mm wide and secondary panicles composed of 1-7 spikelets. Besides, in the Lombok specimens referred to C. myosurus, the leaves are only 2—3 mm wide. Carex longibracteata is very similar to C. myo- surus var. eminens; usually the inflorescence is less compound (but profusely branched inflorescences occur, see KERN 8376!) and the utricles are still somewhat broader. In the Celebes and Lombok specimens referred by NELMEs to C. myosurus the utricles are considerably narrower. The only Javan collection Nemes referred to C. myosurus (VAN STEENIS 12267) was gathered together with VAN STEENIS 12269, which was named C. longibracteata for reasons I do not understand. 6. Section Oligostachyae CLARKE, J. Linn. Soc. Bot. 37 (1904) 4. — Sect. Polystachyae CLARKE [ser. | Longis- picae CLARKE, /.c., p.p. — Sect. Frigidae Fries subsect. Decorae KUx. Pfl. R. Heft 38 (1909) 541. — Sect. Decorae (KUK.) OHwi, Mém. Coll. Sc. Kyoto Imp. Un. B11 (1936) 338; NeLMES, Reinwardtia 1 (1951) 332; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 71; KoyAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 154. — Sect. Borneenses NELMES, Kew Bull. (1951) 121; Reinwardtia 1 (1951) 347; Ray. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 82. Type species: Carex borneensis CLARKE (lectotype). 17. Carex celebica KUx. Bot. Jahrb. 70 (Jan. 1940) 465; Bull. Jard. Bot. Btzg III, 16 (Feb. 1940) 318; NELMES, Kew Bull. (1949) 389; Reinwardtia 1 (1951) 345. — C. constricta S. T. BLAKE, J. Arn. Arb. 28 (1947) 112, f. 4A. — Fig. 119. Rhizome creeping, emitting slender stolons. Stems solitary or somewhat tufted, erect or curved, triquetrous, smooth, often hidden in the leaf- sheaths, 2-25 cm by c. 1'/, mm, surrounded below the leaves by purplish cataphylls or their fibrous remains. Leaves subbasal, often recurved, longer than the stem, flattish or canaliculate, rigid, scabrid in the upper part, gradually attenuate into a firm point, 1'/,-5 mm wide; sheath purplish. Jnflores- cence consisting of 5-15 spikelets, 2-10 cm long. Lower bracts foliaceous, much exceeding the inflorescence, shortly sheathing, upper much reduced. Spikelets suberect, '/,-3 cm long, upper approximate on hardly or not exserted; smooth peduncles; lower sometimes distant on longer peduncles, terminal dg, linear, 1-2 mm thick, lateral 2 or witha few ¢ flowers at the top, cylindric, 3-4 mm thick, lowest solitary, upper in fascicles of 2-3 (at least one of the fascicles of 3). Glumes ovate or oblong-ovate, obtuse to emarginate, more rarely acute, reddish brown with whitish-hyaline margins, slenderly few-nerved, 2-3 mm long, the midrib excurrent in a mucro or in an up to 3 mm long awn. Utricles triquetrous, ovoid or ellipsoid, with flat faces, nerveless (except for two marginal nerves), glabrous, smooth, patulous, slightly recurved, finally dark brown, 2!/;—3'/, by ?/;).>—-1 mm, subabruptly narrowed into a smooth or sparsely hispidulous-margined */,-1 mm long beak; mouth subentire or bidenticulate. Nut ellipsoid or ovoid, triquetrous with flat faces, densely puncticulate, dilate-annulate at the apex, 1!/.-11/, by */,-1 mm. Style-base thickened. Stigmas 3. Distr. Malesia: Sumatra (G. Leuser), SW. Celebes (Mt Pokapindjang), New Guinea (W.: Carstensz Mts; NE.: Mt Hagen, Mt Michael, Mt Piora, Mt Wilhelm, Mt Sarawaket, Sattelberg; Papua: S. Highlands, Mt Ambua, Mt Giluwe, Mt Victoria, Owen Stanley Range. Ecol. Open places, mountain heaths, alpine grasslands and swamps; 2400-4040 m. Vern. New Guinea: tudik, Mendi language. Note. The original descriptions of both Carex celebica and C. constricta were based on dwarfy specimens a few cm tall. Collections recently made 140 FLORA MALESIANA [ser. I, vol. 9} in New Guinea have shown that the species can reach a height of 25 cm and the leaves a width of 5 mm. Stout specimens are very similar to 20. C. verticillata, a close ally of C. celebica, but easily distinguished by the long stipitate 4'/,-6'/, mm long utricles in the latter. 18. Carex perakensis C. B. CLARKE, FI. Br. Ind. 6 (1894) 720; J. Linn. Soc. Bot. 37 (1904) 9; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 116; FI. Mal. Pen. 5 (1925) 184; Urrr. Rec. Trav. Bot. Néerl. 32 (1935) 201; NeELMEs, Kew Bull. (1950) 189; Reinwardtia 1 (1951) 253; Mém. Mus. Hist. Nat. Paris n.s. 4 (1955) 114; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 74, f. 18; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 156; Raym. Dansk Bot. Ark. 23 (1965) 259. — ? C. arridens CLARKE, FI. Br. Ind. 6 (1894) 726; Kux. Pfl. R. Heft 38 (1909) 548; RipLt. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 117; Fl. Mal. Pen. 5 (1925) 184; NELMES, Reinwardtia 1 (1951) 333; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 139; RAaym. Mém. Jard. Bot. Montréal n. 53 (1959) 74. — C. tonkinen- sis FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 8 (1896) 251; KUxK. Pfl. R. Heft 38 (1909) 292; NELMES, Kew Bull. (1950) 190; Reinwardtia 1 (1951) 254; Raym. Nat. Canad. 82 (1955) 165, f. 5. — C. wightiana NEES var. perakensis KUK. Pfl. R. Heft 38 (1909) 288. — C. leucostachys RipL. Kew Bull. (1928) 77; NELMEsS, Reinwardtia 1 (1951) 251; Mém. Mus. Hist. Nat. Paris n.s. 4 (1955) 113. — C. nodiflora (non Boeck.) KUK. Bull. Jard. Bot. Btzg III, 16 (1940) 316. — C. pseudorivulorum Kix. l.c. 319, quoad specim. cit., non C. rivulorum RIDL. (basionym). — C. setulifolia NELMES, Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 116. — Fig. 119, 131b-d. Further synonyms under the varieties. Rhizome short or shortly creeping (thick) woody. Stems tufted, erect, trigonous or triquet- rous, smooth or sparsely scabrid on the angles, especially above, 15-175 cm by '/,-4 mm below, surrounded below the leaves by brown to purplish brown bladeless sheaths or their fibrous remains. Leaves basal and subbasal, rarely spaced through- out the stem, shorter than to exceeding the stem, flattish, rather stiff, scabrid or smooth on the Margins, asperous towards the long acuminate apex, (2-)3-16 mm wide; sheaths often dark nerved, hispidulous or puberulous at the mouth. Inflorescence usually consisting of 2—7 fascicles, or spikelets single, binate or ternate at each node; the fascicles formed by single spikelets (2-14 spikelets in the whole inflorescence), or by up to 20 panicles (in the whole inflorescence); the panicles consist of a very lax raceme of up to 10 secondary spikelets, each secondary spikelet subtended by a whether or not sheathing glumiform, often long awned bractlet, and surrounded at the base by an ocrei- form cladoprophyll. The lower fascicles or spike- lets distant, on long exserted smooth peduncles, the higher approximate. Lower bracts foliaceous, long sheathing, the sheaths often ampliate; higher bracts much reduced, when glumiform the inflores- cence is terminated by a seemingly more compound panicle. Spikelets androgynous, cylindric, but the 3 part tapering, up to 10 in each panicle, the 2 part mostly lax flowered, from much shorter to longer than the ¢ part, 3-8 mm thick. Glumes oblong to ovate or lanceolate acute to obtuse or acuminate, glabrous or hispidulous, nerved, light or red brown, or whitish to fulvous, usually with whitish hyaline margins, 2!/,-7 mm long, the midrib just below the apex apiculate or excurrent into an up to 5 mm long awn. Utricles trigonous, oblong to narrowly ellipsoid, or obovoid, 5—9!/, by 1-21/; mm, sub- erect to patulous, straight or obliquely bent at the apex, many-nerved, hispidulous, shortly stipitate or sessile, subabruptly beaked or gradually narrowed, beak bidentate, often with oblique mouth, 1!/,-3 mm long. Nut trigonous with flat to slightly concave faces, ellipsoid to oblong ellipsoid or obovoid, shortly stipitate and beaked, 3-4 mm long. Style-base slightly thickened. Stigmas 3. Distr. S. China (Yunnan and Kwangsi), For- mosa, Tonkin, Annam, Laos, Lower Burma, Thailand; in Malesia: Sumatra (Atjeh, East Coast, Palembang), Malay Peninsula (Perak, Larut, Pahang, Selangor), Borneo, SW. & SE. Celebes. Ecol. See under the varieties. Notes. The species is very variable, three varie- ties being distinguished. Carex borneensis seems at first sight to be a different species, although closely allied to C. vansteenisii s.s., because in most of the material of C. borneensis the spikelets are not com- pound. In the Malay Peninsula, however, this difference fades away, the spikelets of C. borneensis becoming also compound. Carex vansteenisii, a very rare plant, possesses exactly the same inflores- cence as C. perakensis, the utricles are narrower, and their length overlaps with the range of those of C. perakensis (7-9 mm in C. vansteenisii, 5— 6(—8) mm in C. perakensis). NELMES distinguished also between C. borneensis and C. kinabaluensis, although there are no differences at all between the two. The Celebes collection (C. eymae NELMES) is young and differs hardly from the Bornean specimens of C. borneen- sis. Carex kinabaluensis was considered an ally of 57. C. brunnea, as STAPF made a mistake in describing it as having 2 stigmas (in fact the number of stigmas is the only difference between the two sections!). In the specimens of C. borneensis from the Malay Peninsula, as mentioned in the first note, some of the spikelets are branched (into 2-4 secondary spikelets). On account of this NELMES maintained C. breviglumis RIDL. as a species distinct from the Bornean plants of C. borneensis, in which the spikelets are usually unbranched. However, this difference does not always hold. NELMES considered C. tonkinensis and C. leuco- 1979 Cy PERACEAE—II (Kern & Nooteboom) 141 stachys specifically distinct from C. perakensis s.s., but I could not find differences, and his key characters are unfit for discrimination. Carex leucostachys has been recorded from Pahang (P. Tioman), S. Sumatra (G. Pesagi), and Tonkin. In the type collection the inflorescence is a dense, head-like panicle, and the utricles are about 8 mm. The inflorescence is still too young for good description, but it shows resemblance with C. van- steenisii s.s. in the utricles, and several other collections show transitions to C. perakensis s.s. CLARKE based C. arridens on two collections (Lower Burma, KuRzZ, n.v., and Perak, K1NG’s coll. 2801). The Perak specimen, the only one ever collected in Malesia (in 1882) is very young with quite undeveloped flowers. It has the dark-nerved basal sheaths and the pale indumentum of the young utricles of C. perakensis s.s. The glumes are darker than is usual. In 1951 Nemes placed it in subg. Carex, but C. perakensis, C. tonkinensis and C. leucostachys in subg. Indocarex. The two sub- genera he distinguished by their cladoprophylls (utriculiform in subg. Indocarex, ocreiform in subg. Carex). He described the cladoprophylls of C. perakensis as “‘utriculi-glumiform’’, those of C. tonkinensis as “‘more or less ocreiform” of C. leucostachys as ‘‘subocreiform below, glumiform above” and of C. arridens as “‘subutriculiform below, glumiform above’. In 1955 he removed C. tonkinensis to subg. Carex, C. leucostachys and C. perakensis he left in subg. Indocarex. KEY TO THE VARIETIES 1. Spikelets single or binate at the nodes, not com™ pound. Utricles 5—9!/, mm, gradually tapering into the 2-3 mm long beak. Leaves 2-8 mm wide b. var. borneensis 1. Spikelets compound, or when simple in fascicles at the nodes. 2. Utricies densely pale to golden hispidulous, subabruptly beaked, sessile, ellipsoid or ellipsoid-obovoid, 5—6(-8) mm long. Glumes translucent, whitish, or fulvous with broad whitish margins, 3!/,-5!/, mm long. Inflores- cence consisting of up to 20 panicles which are single, binate or ternate at the nodes. Each panicle consists of a raceme of up to 10 shortly peduncled 8-40 mm long secondary spikelets which is sometimes branched. The ¢ part of the spikelets from much shorter to longer than the 2 part. Number of 92 flowers usually less Phan lO) TR. «aero . a. var. perakensis 2. Utricles brown or olivaceous, gradually beaked, shortly stipitate, narrowly ellipsoid or fusiform, 5—91/, mm. Glumes light brown or reddish brown, the margin often whitish hyaline, 2!/,-7 mm long. Inflorescence con- sisting of 2-14 spikelets branched into 2-4 secondary spikelets, single or binate at the nodes, or 4~7 fascicles of spikelets or panicles. 3. Inflorescence consisting of 2-14 spikelets, which are often branched into 2-4 secondary spikelets, the g part shorter to much longer than the 2 part. Glumes 2'/,-4 mm long, light brown with whitish hyaline margins b. var. borneensis 3. Inflorescence consisting of 4~7 fascicles of spikelets or panicles. The panicles consisting of a very lax raceme of up to 7 secondary spikelets. Spikelets (or secondary spikelets) up to 10 in each fascicle, lax flowered, the 2 part much longer than the 3 part. Glumes brown or reddish brown, 5—7 mm long c. var. vansteenisii a. var. perakensis. — All synonyms under the species. — Fig. 119, 131¢c-d. Stems tufted, erect, trigonous or triquetrous, smooth or sparsely scabrid on the angles above, 60-175 cm by 2-4 mm. Leaves 4-12(—16) mm wide. Spikelets 1-2(-4) cm long, the 3 part much shorter than the 5-7 mm thick 2 part. Awn of glumes up to 17/, mm long. Beak of the 1*/,-2'/, mm wide utricles 11/,-2(—-3) mm long, often obliquely bent. Distr. China (Yunnan, Kwangsi), Formosa, Tonkin, Annam, Thailand, Lower Burma; in Malesia: Sumatra (Atjeh, E. Coast, Palembang), Malay Peninsula (Perak: Larut; Pahang, Selangor), Borneo (Sarawak: Mt Dulit; Sabah: Mt Kinabalu), SW. Celebes (Mt Poka Pindjang). Ecol. In swampy localities and primary forest, 750-1700 m, on Mt Kinabalu up to 2700 m. b. var. borneensis (CLARKE) Noot., comb. nov. — C. borneensis CLARKE, J. Linn. Soc. Bot. 37 (1904) 14; Kew Bull. add. ser. 8 (1908) 76; KUxK. Bull. Jard. Bot. Btzg III, 16 (1940) 320, incl. var. clemensii (KUK.) KUxK. f. angustifrons KUK.; NELMES, Reinwardtia 1 (1951) 351. — C. fusiformis (non Nees) STAPF, Trans. Linn. Soc. II, 4 (1894) 246. — C. fusiformis NEES var. borneensis (CLARKE) KK. Pfi. R. Heft. 38 (1909) 598. — C. kinabaluen- sis STAPF, J. Linn. Soc. Bot. 42 (1914) 183; NELMES, Kew Bull. (1950) 200; Reinwardtia 1 (1951) 349; ibid. 2 (1954) 377; RaymM. Mém. Jard. Bot. Montréal n. 53 (1959) 82. — C. rivulorum Rip. J. Fed. Mal. St. Mus. 6 (1915) 195, non Dunn, 1908. — C. breviglumis RD. Fl. Mal. Pen. 5 (1925) 183; NeELMEs, Reinwardtia 1 (1951) 348. — C. clemensii Kix. in Fedde, Rep. 29 (1931) 202. — C. pseudorivulorum Kwux. Bull. Jard. Bot. Btzg III, 16 (1940) 319, quoad basionym. — C. eymae NeLmes, Kew Bull. (1950) 199; Reinwardtia 1 (1951) 352. — Fig. 119, 131b. Stems erect, triquetrous, smooth or slightly scabrid, (15—)40-140 cm. Leaves shorter than to as long as the stem, (2-)3—5(-8) mm wide. Inflores- cence narrow, up to 30 cm long, consisting of 2 to 14 spikelets. Spikelets up to 5 cm long, single or binate at the middle nodes, erect or suberect, simple or branched into 2-4 secondary spikelets, the 3 142 FLORA MALESIANA [ser. I, vol. 91 part shorter to much longer than the lax-flowered 3-5 mm thick 2 part. Glumes light brown with whitish hyaline margins, 2!/,-4 mm long, apiculate or midrib excurrent into an up to 5 mm long awn. Utricles olivaceous, 5—9!/, by 1-2 mm, gradually tapering into a hispidulous margined, 2-3 mm long beak. Distr. Annam (Prov. Kontum: Ngoe Pang); in Malesia: Malay Peninsula (Pahang: G. Tahan, G. Jasar, G. Berumban), Borneo (Sabah: Mt Kinabalu, Mt Tambuyokan, G. Alab; Sarawak: Mt Poé, Mt Dulit; SE. Borneo: W. Kutei), SE. Celebes (Enrekang). See the map by RAYMOND, Eexpai03: Ecol. Wet places in forests and in shady places along streams; 800-3300 m. Note. Specimens with very regularly puckered- bullate leaves were twice collected on Mt Kinabalu. The strange abnormality of the leaves may be due to some disturbance during the growth period. The same phenomenon has been observed in some Aponogeton, Cryptocoryne, and Halophila species. c. var. yansteenisii (KUK.) Noot., comb. nov. — C. vansteenisii KUK. Bull. Jard. Bot. Btzg III, 16 (1940) 320, incl. var. brevispiculosa KUx. I.c. 321; NELMES, Reinwardtia 1 (1951) 444; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 141; Ravym. Mém. Jard. Bot. Montréal n. 53 (1959) 73. — Fig. 119. Stems rather stout, erect, triquetrous, smooth, c. 150 cm by 3-4 mm. Leaves shorter than the stem, stiff, 3-15 mm wide, sheaths shining membranous in front. Inflorescence consisting of 4~7 fascicles of panicles consisting of a very lax raceme of up to 7 secondary spikelets. Spikelets lax-flowered, 2-6 cm long, 4-8 mm thick, the 2 part much longer than the ¢ part, from c. 4- to c. 10-flowered. Glumes (reddish) brown, 5-7 mm long, awn 0-1 mm long. Utricles exceeding the glumes, suberect to patulous, gradually narrowed into the 2-3 mm long beak, 7-9 by 1-1!/, mm. Distr. Tonkin, Laos; in Malesia: N. Sumatra (Atjeh: Losir massif). Ecol. Mountain heaths, 2100-2500 m. In the Losir area this variety is found at higher altitude than var. perakensis. 19. Carex turrita C. B. CLARKE, J. Linn. Soc. Bot. 37 (1904) 13; Philip. J. Sc. 2 (1907) Bot. 108; Kew Bull. add. ser. 8 (1908) 74; Kix. Philip. J. Sc. 5 (1911) Bot. 63; Merr. En. Philip. 1 (1923) 142; NELMES, Reinwardtia 1 (1951) 335. — Fig. 119, 131e-f. See for synonyms under the varieties. Rhizome short or shortly creeping, woody. Stems erect, trigonous, smooth, up to more than 1 m by ?/,-3 mm below. Leaves basal and up to 3 higher on the stem, shorter than the stem, stiff, flat or with slightly revolute margins, with scabrid margins and asperous upper surface towards the long attenuate apex, 2-7 mm wide; sheaths usually reddish or purplish, glabrous, with concave to convex mouth, the basal ones fraying into fibres. Inflorescence an often interrupted slender panicle to 50 cm long, consisting of 3-8 fascicles, lower fas- cicles distant, upper approximate. Lower bracts foliaceous, shorter to longer than their fascicles but much exceeded by the stem, long sheathing, upper much reduced; sheaths similar to those of the stem. Lower spikelets often single or binate, upper 2-6 together, often branched into 2-6 secondary spikelets, all androgynous, the ¢ part from much shorter to longer than the 9 part, 2-8 cm by 2-3!/, mm, much exserted from their sheaths. Glumes oblong-ovate, obtuse to emargi- nate, glabrous or sparsely hispidulous towards the apex, dark red with wide white hyaline margins, nerved, 2-3'/, mm long, the midrib excurrent into an up to 2 mm long hispidulous awn. Utricles trigonous, oblong to ellipsoid, slenderly nerved to nerveless except 2 submarginal nerves, dark red, reddish or brown, straight or slightly curved, glabrous or hispidulous, at least above, the margins sparsely hispidulous from the middle or above the middle upwards, 3-6 by 3/,—1 mm, gradually taper- ing into a hispidulous margined, 1-2 mm long bidentate beak. Nut triquetrous, ellipsoid, with flattish or concave faces, brown or fulvous, stipi- tate, 1'/,-3!/, mm long. Style-base slightly thickened. Stigmas 3. Distr. Malesia: Philippines and New Guinea. a. var. turrita. — C. turrita C. B. CLARKE. — C. walkeri ARN. ex BooTT var. turrita (CLARKE) KO. Pfi. R. Heft 38 (1909) 546. — C. atrosanguinea NELMES, Kew Bull. (1950) 197; Reinwardtia 1 (1951) 339. — Fig. 119, 131e-f. Glumes 3-3!/, mm long, with an awn of 2/,- 2 mm. Utricles slenderly nerved, 5-6 mm long, with a beak of 2 mm. Nut 3-3!/, mm long. Distr. Malesia: Philippines (Luzon: Abra, Bontoc, Ifugao, Benguet), New Guinea (W. New Guinea: Mt Treub; NE. New Guinea: Morobe Distr.; Papua: W. Highlands, Kubor Range, E. Highlands, Central Div.). Ecol. Open places in mossy forest, montane rain-forest, 1500-3040 m. Vern. Philippines: tangtafgo, Bon. Notes. NeLMEs distinguished the New Guinean C. atrosanguinea from the Philippine C. turrita by the simple spikelets of the former. Additional collections from New Guinea have shown that also here specimens with branched spikelets occur; it must, however, be admitted that in general the spikelets in the Philippines are more com- pound. I have not seen C. gibbsiae RENDLE, J. Linn. Soc. Bot. 39 (1909) 180, from Fiji, which is, according to NELMES, closely related to his C. atrosanguinea, but looking ‘“‘very distinct because of its different colouring’’. I suspect it to be also conspecific with C. turrita. 1979] b. var. merrillii (KUK.) Noort., comb. nov. — C. merrillii KUK. in Fedde, Rep. 8 (1910) 7; Philip. J. Sc. 6 (1911) Bot. 63; Merr. En. Philip. 1 (1923) 139; Netmes, Reinwardtia 1 (1951) 336; ibid. 2 (1954) 377. — C. pullei NeELMEs, Kew Bull. (1950) 198; Reinwardtia 1 (1951) 338. — Fig. 119. Glumes 2-2'/, mm long, awn up to '/, mm. Utricles merveless to slenderly few-nerved, 3-33/, mm long, with a beak of 1-1'/, mm. Nut 11/,-13/, mm long. Distr. Malesia: Philippines (Luzon, Ifugao, Benguet), New Guinea (W. New Guinea: Mt Treub; NE. New Guinea: Morobe Distr.). Ecol. Mossy forest, open places along trails, also on steep slopes; 1800-2490 m. Note. Kern (in manuscript) reduced C. pullei to C. turrita with a question-mark. He noted: “It seems to differ mainly by the smaller utricles only 3-4 mm long, and may represent a depauperated state’. NeLMES described the utricles as being “almost nerveless to rather slenderly 1—3-nerved”’. As these were the only differences between C. tur- rita and C. merrillii, 1 have reduced the latter to a variety of the former. 20. Carex verticillata ZoLL. & Mor. in Mor. Syst. Verz. (1846) 98; StEuD. Syn. 2 (1855) 222; Mia. Fl. Ind. Bat. 3 (1857) 353; Boeck. Linnaea 41 (1877) 256; NeLMEs, Kew Bull. (1950) 195, incl. var. havilandii (CLARKE) NELMES et var. lutescens NELMES; Reinwardtia 1 (1951) 340; Raym. Mérn. Jard. Bot. Montréal n. 53 (1959) 74; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 488. — C. hypsophila Mia. Fl. Ind. Bat. 3 (1857) 354; Boeck. Linnaea 41 (1877) 257; CLarKE, J. Linn. Soc. Bot. 37 (1904) 13; KUK. Pfl. R. Heft 38 (1909) 546, f. 89, incl. var. havilandii (CLARKE) KUK. et var. verticillata (ZOLL. & Mor.) KuK.; in Hochr. Candollea 6 (1936) 432. — C. walkeri (non Bootr) Boeck. Linnaea 40 (1876) 332, p.p.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 65. — C. tartarea RIDL. J. Bot. 23 (1885) 35. — C. havilandii CLARKE, J. Linn. Soc. Bot. 37 (1904) 13; Kew Bull. add. ser. & (1908) 75. — C. sumatrensis CLARKE, J. Linn. Soc. Bot. 37 (1904) 13; Kew Bull. add. ser. 8 (1908) 75. — C. turrita (non CLARKE) KUxK. Bull. Jard. Bot. Btzg III, 16 (1940) 319. — C. decora Bootr var. losirensis KK. l.c. — C. phacelostachys NELMES, Kew Bull. (1950) 195; Reinwardtia 1 (1951) 344, incl. var. losirensis (KUK.) NELMES; RAYM. Mém. Jard. Bot. Montréal n. 53 (1959) 72. — Fig. 119. Rhizome creeping, woody. Stems solitary or somewhat tufted, erect, triquetrous, smooth, 25-100 cm by 1-2 mm, surrounded below the leaves by dark reddish, entire or fibrous remains of old leaf-sheaths. Leaves basal and 1-2 on the stem proper, shorter than to equalling the stem, flattish with often strongly revolute margins, stiff, coria- ceous, with scabrid margins, asperous above especially towards the long-attenuated apex, Cy PERACEAE—II (Kern & Nooteboom) 143 3-8 mm wide; sheaths of the lower leaves reddish to blackish red, upper ones pale. Jnflorescence a lax to dense panicle, 15-50 cm long, consisting of 4-7 fascicles of up to 20 spikelets, lower fascicles rather distant, upper approximate. Bracts of the lower fascicles foliaceous, equalling or exceeding their fascicles but usually much exceeded by the whole inflorescence, long-sheathing, upper much reduced; sheaths ampliate, glabrous, the mouth often prolonged into a short tongue. Spikelets erect or somewhat nodding, slenderly cylindric, lax- to rather dense-flowered, up to 6 cm long, simple or longest ones sometimes branched into 1-3 short secondary spikelets, on slender, smooth peduncles more or less exserted from the sheaths; 1-2 spikelets of uppermost fascicle (sometimes of all the fascicles) 3, remaining ones wholly 2 or with a few ¢ flowers at the top; ¢ spikelets 1-2 mm thick, 2 3-5 mm. Glumes oblong-lanceolate, ob- tuse, glabrous, fulvous to dark reddish with wide whitish-hyaline margins, 3—4°/, mm long, the midrib often excurrent in a smooth or hispidulous awn up to 1 mm long. Utricles trigonous, narrowly ellipsoid, tapering at both ends, glabrous but the margins often hispidulous, nerveless or faintly nerved (except for the marginal or submarginal nerves), suberect to patulous, often somewhat recurved, long-stipitate, stipe (/,-)°/,-1 mm, subabruptly beaked, blackish red to golden, 4-6'/, by */,-1 mm; beak almost linear, sparsely hispidulous-margined, bidenticulate, (1-)2-3 mm long. Nut triquetrous with flattish faces, ellipsoid, brown, 13/,-2 mm long. Style not or scarcely thickened at the base. Stigmas 3. Distr. Laos, Tonkin; in Malesia: Sumatra, Malay Peninsula (Perak: G. Kerbau), Java, N. Borneo (Mt Kinabalu). Ecol. Damp open places in forests, open places in the subalpine region, near craters; 2000-3750 m. Notes. Variable as to the colour of glumes and utricles. In most of the Mt Kinabalu speci- mens the glumes and utricles are very dark; they were distinguished as var. havilandii (CLARKE) NELMES. A collection from E. Java (G. Semeru), in which the glumes and utricles are pale golden to fulvous, was distinguished as var. /utescens NELMES. Carex phacelostachys NeELMES only could be distinguished by the utricles which are faintly nerved against nerveless in C. verticillata. The collection VAN STEENIS 9624 from Atjeh has been identified as C. verticillata by both NELMES and KERN; after careful examination of the utricles I could not find any difference with C. phacelostachys. I examined many utricles of C. verticillata with translucent light. Most of them are nerveless indeed, but others are faintly nerved, often only at the base, at least in plants from Sumatra and Borneo. — (Noort.) The species might also be conspecific with C. walkeri ARNOTT ex Boott. — (Noot.) 144 FLORA MALESIANA [ser. I, vol. 91 7. Section Surculosae Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 20, 21. — Sect. Scabrellae Kuk. Pfi. R. Heft 38 (1909) 286, p.p.; NELMES, Reinwardtia 1 (1951) 243. Type species: Carex oligostachya NEES (lectotype). 21. Carex oligostachya NEEs in Hook. J. Bot. Kew Misc. 6 (1854) 29; NELMEs, Reinwardtia 1 (1951) 244; ibid. 2 (1954) 373; Kew Bull. (1955) 301; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 95; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 22. — C. cumingiana STEUD. Syn. 2 (1855) 206; Mia. FI. Ind. Bat. 3 (1856) 349; Boorrt, Ill. 3 (1862) 107, t. 324, 325; Boeck. Linnaea 40 (1876) 367; F.-VILL. Nov. App. (1882) 310. — C. rhizomatosa Steup. [in Zoll. Syst. Verz. 1 (1854) 60, nomen] Syn. 2 (1855) 206; Mia. Fl. Ind. Bat. 3 (1856) 348; CLARKE, FI. Br. Ind. 6 (1894) 721; J. Linn. Soc. Bot. 37 (1904) 12; Philip. J. Sc. 2 (1907) Bot. 108; Kux. Pfi. R. Heft 38 (1909) 289, f. 44, excl. var. impunctata (BOOTT) KUK.; Philip. J. Sc. 6 (1911) Bot. 61; Camus, Fl. Gén. I.-C. 7 (1922) 193; Merr. En. Philip. 1 (1923) 141; KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 316, incl. var. aristulata Kwx.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 107; NELMES, Kew Bull. (1949) 378, 387. — C. bukaénsis PALLA in Rechinger, Bot.-Zool. Ergebn. Samoa & Solomon Ins. (1913) 58. — C. breviceps KUxK. Bot. Jahrb. 69 (1938) 263. — Fig. 120. Rhizome shortly creeping, woody, like the stem- base clothed with brown or blackish, fibrous remains of old leaf-sheaths. Flowering stems lateral, approximate, subscapiform (bearing a few short- bladed leaves), triquetrous, smooth, 20-90 cm by 3/,-2 mm. Leaves of the sterile shoots shorter than the flowering stems, stiffish, flat, rather abruptly acuminate, scabrid in the upper part, glaucous, 2-5 mm wide. Inflorescence a narrow panicle consisting of 6-12 head-like or racemose partial inflorescences, 10-50 cm long; partial inflores- cences erect, subglobose to oblong-ovoid, with 1-8 spikelets, the lower ones single at the nodes, distant, on long-exserted, setaceous peduncles, the upper ones more approximate, unequally binate (rarely ternate); axis and top of peduncles scabrid. Bracts foliaceous with blades shorter than 8 cm, their sheaths ampliate, membranous near the mouth, the upper ones much reduced, infundibuli- form. Spikelets androgynous, ovoid to subcylindric, sessile, 4-10 mm long, the ¢ and 2 parts usually about equal in length. G/umes ovate or lanceolate, acutish to slightly notched, usually sparsely hispi- dulous towards the apex, 5—10-nerved, brown, often with reddish spots, whitish hyaline-margined, 2-3 mm long, the midnerve usually excurrent in a short, scabrid awn up to 1 mm long. Utricles obtusely trigonous, ellipsoid, subinflated, patent, straight, glabrous or sparsely hispidulous in the upper !/,, strongly many-nerved, rather abruptly beaked, fuscous, 2!/,-4 by 1-2 mm; beak 3/,-1 mm long, bidenticulate. Nut trigonous, ellipsoid or obo- void, minutely stipitate, 2-2'/, by 1-11/, mm. Style-base not thickened. Stigmas 3. Distr. Assam, Upper Burma, S. China, Tonkin, through Malesia to the Solomon Is.; in Malesia: Sumatra (Atjeh, Tapanuli), Lesser Sunda Is. (Sum- ba, Sumbawa, Flores), Philippines (Luzon, Negros, Mindanao), SE. Celebes (Buton I.), New Guinea. According to MERRILL, /.c., also in the Moluccas. Ecol. In Imperata-fields, on open grassy slopes, often in places where the grass is burned annually; from low altitude up to 1400 m. Vern. New Guinea: simboro, Orokaiva lang. Notes. The utricles vary from wholly glabrous (C. breviceps Kix.) to sparsely hispidulous above. I have not seen C. breviceps var. recurvirostris KUK. Bot. Jahrb. 70 (1940) 464, based on CLEMENS 8032 from NE. New Guinea, Morobe, Sattelberg. Stems 90 cm tall. Leaves 5-10 mm wide. Panicle 35 cm long. Spikelets longer, broadly oblong. Utricles green, subabruptly excurrent in a long, recurved beak. 8. Section Rhomboidales KUK. Pfi. R. Heft 38 (1909) 622; NeLMEs, Reinwardtia | (1951) 383; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 66. Type species: Carex chinensis RETZ. (lectotype). 22. Carex anomocarya NELMES, Kew Bull. (1950) 202; Reinwardtia 1 (1951) 383; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 163; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 66, f. 16, map p. 102, f. 32; Dansk Bot. Ark. 23 (1965) 259; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489. — C. har- landii (non Bootr) Merr. & CHUN, Sunyatsenia 2 (1935) 208. — C. harlandii BootT var. angustior KUx. ex Back.; Gross, Notizbl. Berl.-Dahl. 14 (1938) 193; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 65. — C. harlandii Bootr f. longibracteata Gross, Notizbl. Berl.-Dahl. 14 (1938) 193. — C. manca Bootrt var. contigua GROSS, lic. — Fig. 120, 128. 1979] Fig. 128. Carex anomocarya NELMES. a. Habit, x 2/3, 6. glume, c. fruit in utricle, d. fruit, all x 6 (RAHMAT st BoEEA 10601). 146 FLORA MALESIANA [ser. I, vol. 91 Rhizome short, not creeping. Stems central, erect, triquetrous, smooth, 30-70 cm by 1?/,- 2 mm. Leaves basal (rarely one in the middle of the stem), flat, weak, scabrid on the margins, 6-15 (-20) mm wide. Jnflorescence with 3-5 spikelets, erect; terminal spikelet g, cylindric, 21/,-5 cm by c. 2 mm, peduncled, lateral ones 9, oblong- cylindric, approximate with one another and with the 3 spikelet, but lowest often distant, subdense- flowered, on shortly exserted peduncles, 4-8 cm by 6-9 mm; peduncles stoutish, smooth. Bracts foliaceous, lower usually much exceeding the inflorescence, shortly sheathing; sheaths pale, membranous in front or only at the mouth, sub- ampliate. Glumes of the 9 spikelets oblong, sub- truncate to slightly emarginate, dirty white with hyaline margins and 3-nerved centre, ciliolate at the apex, awned, 3-4'/, mm long, those of the g spikelets narrower, also long-awned; awns scabrid, up to 1 cm long. Utricles ellipsoid to rhomboid- lageniform, obtusely trigonous, surface uneven with concavities and convexities corresponding to the surface of the nut, densely many-nerved, coriaceous, glabrous, straight, suberect, shortly stipitate, stramineous, 7—8(-10) by 2-23/, mm, rather abruptly narrowed into a 2!/,-4 mm long, conical, bidentate beak; teeth 1-1'/, mm long, diverging, sparsely hispidulous at the mouth. Nut unevenly ellipsoid or ovoid, distorted-trigonous, excavate on the angles at the centre, with concave faces below, stipitate, abruptly beaked, 4—5!/, mm long; beak cylindric, 1/,-11/, mm long, expanding into the annulate apex. Style-base pyramidally thickened, persistent. Stigmas 3. Distr. N. Burma, NE. Thailand, Tonkin, Annam, Hainan; in Malesia: N. Sumatra (E. Coast: Dolok Singgalang and G. Batu Lopang; Toba), W. Java (Priangan: Tjadas Malang S. of Tjiandjur; Mt Bésér near Tjidadap). Ecol. In damp shady forest, in W. Java 1000- 1200 m, in N. Sumatra at 1400-1700 m. Note. Very near to C. harlandii Bootrt, Ill. 2 (1860) 87, t. 255; Kix. Pfl. R. Heft 38 (1909) 630, f. 107, and possibly better treated as a race of that species. In C. harlandii the leaves are broader (up to 3 cm wide), the bracts much shorter, often not or hardly overtopping spikelet, the glumes of the 3 spikelet rounded at the apex and muticous, those of the 2 spikelets but shortly awned, the teeth of the utricles shorter (c. 1/, mm long), and the beak of the nut slenderer, c. 11/, mm long. It is known from South and Central China. 23. Carex jackiana Boott, Proc. Linn. Soc. 1 (1845) 260; Trans. Linn. Soc. 20 (1846) 132; Ill. 1 (1858) 9, t. 25; Sreup. Syn. 2 (1855) 226; Mia. FI. Ind. Bat. 3 (1856) 353; Boeck. Linnaea 41 (1877) 277; CLARKE, FI. Br. Ind. 6 (1894) 735, excl. var. 8; J. Linn. Soc. 37 (1904) 15; Kix. Pfl. R. Heft 38 (1909) 638; in Hochr. Candollea 6 (1936) 432, excl. var. tumens KUK.; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 66; NELMEs, Reinwardtia 1 (1951) 384; Koyama, Bot. Mag. Tokyo 70 (1957) 352, f. 10 & 12 A-A’; ibid. 72 (1959) 303; J. Fac. Sc. Un. Tokyo III, 8 (1962) 230; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490; STEEN. Mt. FI. Java (1972) pl. 14-3. ssp. jackiana. — Fig. 120. Rhizome short. Stems central, tufted, erect, triquetrous (angles prominent to narrowly winged), smooth, 15-100 cm by 1—2 mm, surrounded at the base by a few brownish sheaths or their fibrous remains. Leaves subbasal, sometimes one higher on the stem, shorter than to as long as the stem, flat, rather weak, scabrid at the top, 3-10 mm wide; sheaths pale to whitish. Inflorescence with 3-7 spikelets, erect; terminal spikelet ¢ (sometimes gynaecandrous), slenderly cylindric (or clavate), 1-3 cm by 2-4 mm, lateral ones 2 (rarely andro- gynous), oblong-cylindric, sublax-flowered, 1-3 cm by 5-8 mm, upper erect, crowded with the ¢ spikelet, sessile or very shortly peduncled, lower distant on included to long-exserted, slender, smooth peduncles, often with 1-3 shorter spikelets branching from their peduncles. Bracts foliaceous, exceeding the inflorescence, lower long-sheathing, upper much shorter, scarcely to shortly sheathing. Glumes oblong-ovate, acute, very thin, dirty white with 3-nerved, greenish, central stripe, muticous or mucronulate, rarely with a short awn, 3—5!/, mm long. Utricles fusiform-ellipsoid, trigonous, sub- coriaceous, densely and strongly many-nerved, glabrous, straight, suberect, shortly stipitate, olivaceous, 51/,-7'!/, by 1'/,-2 mm, gradually narrowed into the conical, straight, bidentate beak; teeth 1/,-9/, mm long, hardly diverging, smooth. Nut oblong-obovoid to suborbicular, triquetrous with prominent angles, with faces shallowly concave below, very shortly stipitate, abruptly shortly beaked, 2!/,-3 mm long. Style- base not thickened. Stigmas 3. Distr. Ceylon, India (Assam, Khasia), Yunnan; according to NELMES (1951) 386 also in Australia; in Malesia: Sumatra (W. Coast: Mt Kerintji), West and Central Java. Distribution maps in Bot. Mag. Tokyo 72 (1959) 303; Phytologia 17 (1968) 408. Ssp. parciflora (BOOTT) KUK., often considered specifically distinct, differs from ssp. jackiana by its smaller utricles and shorter glumes; in S. Sachalin, Japes, S. Kuriles, Kyushu, and S. Korea. Ecol. In marshes, swamps, and grassy plains; 1350-2550 m. = 24. Carex lateralis Kix. Pfl. R. Heft 38 (1909) 639; Bull. Jard. Bot. Btzg III, 16 (1940) 322; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 66; NeELMES, Kew Bull. (1950) 204; Reinwardtia 1 (1951) 388; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 488. — C. jackiana BooTtrT var. minor CLARKE, Fl. Br. Ind. 6 (1894) 735, p.p. typ. — C. elmeri Kux. in Fedde, Rep. 8 (1910) 326; 1979 | Emer, Leafl. Philip. Bot. 3 (1910) 853; Kuk. Philip. J. Sc. 6 (1911) Bot. 64; Merr. En. Philip. 1 (1923) 137; KUK. Bull. Jard. Bot. Btzg III, 16 (1940) 322. — Fig. 120. Rhizome short, tufted. Flowering stems lateral, obliquely erect or somewhat cernuous, very slender to subfiliform, weak, triquetrous, smooth, (5—)20— 50cm by !/,-1 mm, surrounded at the base by a few brown sheaths or their fibrous remains, and bearing a few small, bract-like or subfoliaceous leaves. Leaves of the sterile shoots much over- topping the stems, flat, revolute on the margins, stiff, with very rough margins, asperous towards the apex, 3-7 mm wide. Spikelets 3-4, crowded at the apex of the stem, subsessile on shortly exserted or wholly included peduncles, sometimes one 1-3 cm lower down; terminal spikelet 3, cylindric, 5-10 by !/,-1 mm, usually very inconspicuous when the lateral spikelets have developed utricles, few- flowered, lateral ones 9, oblong, few- (up to 5-)flowered, erect, 7-12 by 5-7 mm. Bracts folia- ceous, much exceeding the inflorescence, shortly sheathing. G/umes oblong or oblong-ovate, subacute to rounded, thin, whitish with 3-nerved greenish central stripe, 2'/,-3'/, mm long, excurrent in a flat, scabrous awn 1'/,-6 mm long. Utricles ovoid-ellipsoid, trigonous, membranous, suberect, many-nerved below but few nerves extending more than halfway towards the apex, sparsely pubescent to glabrous, stipitate, shiny, pale green, 5-7 by 2-2'/, mm, subabruptly contracted into a long, conical, somewhat recurved, bidentate beak. Nut ovoid-ellipsoid, triquetrous, with faces concave in the lowest '/3, brown to fuscous, stoutly stipitate, scarcely or not beaked at the rounded apex, 3-4 mm long. Style thickened at the base. Stigmas 3. Distr. Ceylon, India; in Malesia: S. Sumatra (Palembang: Air Njuruk), W. Java (Priangan: Mt Papandajan), Central Celebes, Philippines (Luzon, Mindoro, Negros), Lesser Sunda Is.: Flores (Mt Ranaka). Ecol. In forests, on road-sides; 1400-2200 m. Note. The type of this species is THWAITES CP 3198 p.p., not CLARKE 11061 as cited by NELMES. CLARKE’s description of C. jackiana var. minor, based on this THWAITES collection and on CLARKE 11061 (!) refers to the species described above. Carex jackiana var. minor is therefore synonymous with C. /ateralis and cannot be main- tained as a variety of C. jackiana alongside of C. lateralis, as was done by KUKENTHAL. BOECKELER’S description of C. jackiana (Linnaea 41, 1871, 277), cited by CLARKE in the synonymy of his var. minor, obviously refers to typical 23. C. jackiana. VELDKAMP 7130 has the entirely hairy utricles of C. lateralis, but lacks the awn on the glume; in that respect it fits in 25. C. loheri. The leaves are also Cy PERACEAE—II (Kern & Nooteboom) 147 intermediate between C. /oheri and C. lateralis. When more material becomes available, it is not impossible that the two species will have to be united. — (Noort.) See also note under 25. C. loheri. 25. Carex loheri CLARKE, J. Linn. Soc. Bot. 37 (1904) 14; Philip. J. Sc. 2 (1907) Bot. 108; Kix. Pfl. R. Heft 38 (1909) 487; Philip. J. Sc. 6 (1911) Bot. 64, incl. f. grandimascula KUK.; MERR. En. Philip. 1 (1923) 139; Ne_mes, Reinwardtia 1 (1951) 386. — Fig. 120. Rhizome short, tufted. Flowering stems lateral, Suberect or cernuous, filiform, weak, triquetrous, smooth, 10-40 cm by '/, mm, surrounded at the base by a few brown scales or their fibrous remains. Leaves crowded at the base, shorter than to exceed- ing the stems, flat with revolute margins, stiff, greyish or glaucous-green, with scabrid margins, asperous towards the apex, 11/,-3 mm wide. Spike- lets 3-6, terminal 3, cylindric, 7-15 by 1-2 mm, lateral androgynous, 6-10 by 4-6 mm, with g and 2 parts about equal in length and few-flowered, upper one often approximate with the ¢ spikelet, lower on long, subbasal ones pendulous on very long, setaceous, smooth peduncles. Bracts of the lower spikelets subbasal leaves, of the other lateral spikelets subfoliaceous, rather long-sheathing. Glumes oblong-ovate, acute, thin, many-nerved, muticous or mucronulate, whitish, c. 4 mm long. Utricles ovoid-ellipsoid, trigonous, membranous, suberect, many-nerved, shining, glabrous below, sparsely to densely pubescent above, shortly stipitate, shining, pale green, 6-7 by 13/,-2 mm, subabruptly contracted into a long, conical, some- what recurved, bidentate beak. Nut ovoid or ellip- soid, triquetrous with concave faces, brown to fuscous, shortly stipitate, hardly beaked, 31/,-4 mm long. Style thickened at the base. Stigmas 3. Distr. Malesia: Philippines (Luzon: Lepanto, Bontoc, Benguet, Rizal, Zambales, Laguna, Tayabas). Ecol. Mossy forest, 1300-2400 m. Vern. Silak, Ig. Notes. Very similar to 24. C. /ateralis, to which it is certainly closely related, though KUKENTHAL placed it in a different section. Carex tatsutakensis HAYATA, Ic. Pl. Form. 6 (1916) 133, f. 45; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 396; Koyama, Nat. Canad. 82 (1955) 204; Contr. Inst. Bot. Un. Montreal n. 70 (1957) 19, not rare in Formosa and, according to KoyaMa, also in Tonkin (not mentioned for this country by RAYMOND!) is very near to C. loheri. Also C. sublateralis KOYAMA, Jap. J. Bot. 15 (1956) 180, f. 9, from E. China (Kiangsu) belongs to this group of very closely related species. 148 FLORA MALESIANA [ser. I, vol. 91 9. Section Cryptostachyae (OHw1) NELMES, Reinwardtia | (1951) 363; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 52, 61. — Sect. Praecoces CHRIST subsect. Cryptostachydeae FRANCH. ex OuwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 340. Type species: Carex cryptostachys BRONGN. 26. Carex cryptostachys BRONGN. in Duperrey, Voy. Coq. Bot. (1828) 152, t. 25; Mia. FI. Ind. Bat. 3 (1856) 352; Boorr, Ill. 3 (1860) 103, t. 310; BENTH. FI. Hongk. (1861) 403; Boeck. Linnaea 40 (1876) 327; CLARKE, FI. Br. Ind. 6 (1894) 714 (‘eyrtostachys’); J. Linn. Soc. Bot. 36 (1903) 281; ibid. 37 (1904) 8; Ripi. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 116; Kix. Pfl. R. Heft 38 (1909) 471; Camus, FI. Gén. I.-C. 7 (1922) 195, f. 27, 10-13; Merr. En. Philip. 1 (1923) 137; Riu. Fl. Mal. Pen. 5 (1925) 181, f. 221; KUK. in Hochr. Candol- lea 6 (1936) 432; Onwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 342; Exmer, Leafi. Philip. Bot. 10 (1938) 3526; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 66; NELMEs, Reinwardtia 1 (1951) 363; AktyAMa, Car. Far East. Reg. Asia (1955) 176, t. 177; Raym. Nat. Canad. 82 (1955) 151, f. 2; Mém. Jard. Bot. Montréal n. 53 (1959) 61, map p. 102, f. 34; Dansk Bot. Ark. 23 (1965) 258; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 491. —Fig. 120. Rhizome elongate, horizontal or ascending, woody, clothed with fibrous remains of sheaths. Flowering stems arising from the axils of the leaves, and often almost hidden among them, single or binate, scapiform, suberect, flexuous, obtusely trigonous, smooth, 10-50 cm tall, with sheaths and peduncles almost from the base. Leaves crowded on a short stem, much longer than the flowering stems, firm, flat, long-acuminate, scabrid on the margins, grey- or glaucous-green, 3-18 mm wide, surrounded at the base with fibrous remains of older leaves. Inflorescence racemiform or paniculiform, with 6-30 spikelets; spikelets androgynous, oblong- lanceolate, lax-flowered, 8-30 by 3-5 mm; ¢ part much shorter than the 2 one. Bracts usually shorter than their spikelets, with funnel-shaped sheaths and short blades; peduncles sparsely scaberulous, lower exserted, upper included. Glumes ovate, amplexicaul, many-nerved, obtuse, ciliolate, often minutely appressed-hairy, whitish or stramineous, apiculate or mucronulate, 2!/,-23/, mm_ long. Utricles oblong-rhomboid to obovoid-fusiform, obscurely trigonous, suberect, densely many- nerved, subcoriaceous, sparsely puberulous, ciliolate-scabrid on the margins, stramineous- green, stoutly stipitate, abruptly shortly beaked, with oblique, bidentate mouth, 31/,-51/, by 11/,- 2 mm. Nut irregularly rhomboid-ellipsoid or oblong-obovoid, trigonous, with faces excavated at base and apex, broadly stipitate, seated on a spongy, disk-like body, c. 2 mm long; beak short, strongly deflexed; style bent upwards, thickened above, papillose. Stigmas 3. Vestigial rachilla sometimes present (according to KUKENTHAL). Distr. Formosa, Hainan, S. China (Kwang- tung), Tonkin, Annam, Thailand, Queensland; in Malesia: Sumatra (also Banka), Malay Peninsula, W. Java, E. Borneo, Philippines (Luzon, Minda- nao), Aru Is., New Guinea; the specimen in L from the Lesser Sunda Is. (leg. PLoEM) is probably mis- labelled. Ecol. In primary forests, on banks in woods, in open jungle, at low and medium altitude; in Malaya between 350 and 1200 m, in Banka at 40 m, in the Aru Is. at a few m above sea-level, in W. Java between 500 and 1250 m, in the Philippines ascending to 1000 m. Vern. Rumput ringgin, Mal. Pen.; ilat kampadan, Ss 10. Section Lageniformes (OHw1) NELMES, Reinwardtia | (1951) 366; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 63. — Sect. Praecoces Curist subsect. Lageniformes OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 340. Type species: Carex formosensis Liv. & VAN. (lectotype). 27. Carex breviscapa C. B. CLARKE, FI. Br. Ind. 6 (1894) 736; Kix. Pfl. R. Heft 38 (1909) 474; Evmer, Leafi. Philip. Bot. 10 (1938) 3525; MERR. En. Philip. 1 (1923) 137; Kix. Bot. Jahrb. 69 (1938) 265; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 64; NeLMes, Kew Bull. (1949) 390; Reinwardtia 1 (1951) 369; ibid. 2 (1954) 380; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 63, f. 14, map p. 102, f. 33; Dansk Bot. Ark. 23 (1965) 258; KoyAMA, Phytologia 17 (1968) 405, t. 14; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489. — C. jackiana Bootr var. breviculmis THw. En. P1. 1979] Cy PERACEAE—II (Kern & Nooteboom) 149 Zeyl. (1864) 356. — C. curtisii RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 117; Fl. Mal. Pen. 5 (1925) 183. — C. lutchuensis Onw1, Mem. Coll. Sc. Kyoto Imp. Un. BS5 (1930) 270; ibid. B11 (1936) 343, f. 5, t. 9 f. 8; AkryAMA, Car. Far East. Reg. Asia (1955) 177, t. 178. — Fig. 120. Rhizome short, cespitose. Stems central, some- times more stems together, more or less hidden amongst the leaves, suberect, triquetrous, smooth, 5—20(—30) cm by c. 1 mm. Leaves basal, very much longer than the stems, flat, long-acuminate, scabrid on the margins and upper surface, 3-6(—8) mm wide; lower leaves reduced to bladeless brown sheaths or their fibrous remains. Spikelets usually 5-7, single at the nodes, erect or suberect, cylindric, lax-flowered, lower ones somewhat distant; termi- nal spikelet g¢, 1-2 cm long, 1 mm thick, often Overtopped by some of the @ spikelets; lateral spikelets 9 or with some ¢ flowers at the top, 1-3 cm by 3-4 mm; peduncles scaberulous, lower ones exserted. Lower bracts foliaceous, much exceeding the inflorescence, shortly sheathing, upper much reduced. Glumes broadly ovate to oblong-ovate, usually rounded at the erose apex, much shorter than the utricles, slenderly nerved, with not or slightly (up to 1 mm) excurrent mid- nerve and hyaline margins, ciliolate, 2-3 mm long. Utricles rhomboid-lageniform, trigonous, broadest about the middle, subcoriaceous, straight, suberect, strongly multinerved, glabrous or sparsely puberu- lous above, stramineous or greenish, stipitate, 31/,—4'/, by 11/,-1!/,mm; beak hispid, bidenticulate, 1/,-1 mm long. Nut narrowly rhomboid, triquetrous with concave faces, stipitate, truncate and hollowed out at the apex, dark brown with pale stipe, apex, and angles, 2!/,-3 by 1'/,-11!/. mm. Style-base slightly thickened. Stigmas 3. Distr. Ceylon, NE. Thailand, Annam, Ryu Kyu Is., Formosa, N. Queensland; in Malesia: Sumatra (E. Coast Res., Lampongs), Malay Peninsula (Penang, Pahang, Negri Sembilan), W. Java, Borneo (Sarawak), Philippines (Luzon: Sorsogon; Palawan), W. & NE. New Guinea. Ecol. In secondary forest, on forested ridges, from low altitude up to 1250 m. Vern. /lat daun eurih, S. Note. In some of the ¢ flowers I observed stamens with connate filaments. — (Noort.) 28. Carex gracilispica HAYATA, Ic. Pl. Form. 10 (1921) 62, f. 39; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 345; AktyAma, Car. Far East. Reg. Asia (1955) 178, t. 179; RAym. Mém. Jard. Bot. Montréal n. 53 (1959) 65. — C. ligata (non Boott) Ripv. Fl. Mal. Pen. 5 (1925) 181. — ? C. tristachya THuNB. var. pseudopocilliformis Gross, Notizbl. Berl.-Dahl. 14 (1938) 191. — C. malayana NeELMES, Kew Bull. (1950) 209; Reinwardtia 1 (1951) 366. Rhizome short, cespitose. Flowering stems arising from basal leaf-axils, erect, compressed- trigonous, smooth or scaberulous just below the inflorescence, 10-30 cm by 3/,-1 mm. Leaves basal, much longer than the stems, plicate to flat, long- attenuated, with scabrid margins, 5-10 mm wide, surrounded at the base by brown, fibrous remains of old leaf-sheaths. Spikelets 4-6, approximate or lowest somewhat distinct, erect or suberect, cylindric, 1-31/, cm long; terminal spikelet 3, 1 mm thick, lateral ones single at the nodes, wholly 2 or with some ¢ flowers at the top, lax-flowered, 2-3 mm thick, their peduncles shortly exserted, smooth or scaberulous. Bracts foliaceous, lower as long as or exceeding the inflorescence, shortly sheathing, upper much reduced. G/umes oblong- ovate, acute to obtuse, slenderly nerved, with not or slightly excurrent midnerve, whitish hyaline margins, and erose-ciliate apex, 3-4 mm long. Utricles \ageniform, obscurely trigonous, sub- coriaceous, broadest 2-2!/, mm from the base, straight, suberect, strongly multinerved, sparsely puberulous, stramineous or greenish, shortly stipitate, 5-7 by 11/,-1*/, mm; beak 1!/,-2 mm long, bidenticulate. Nut oblong, triquetrous, with faces concave below, stoutly stipitate, slightly constricted at the apex and re-expanded into a cylindric, truncate neck ?/,-1 mm long and wide, the whole nut 3-4 mm long. Style-base thickened, centred in the apical hollow of the nut. Stigmas 3. Distr. Formosa, Indo-China (Annam, ?Tonkin); in Malesia: Malay Peninsula (Pahang: Mt Tahan). Ecol. On Mt Tahan in damp woods by streams, at c. 1700 m. 29. Carex palawanensis Kix. in Elmer, Leafi. Philip. Bot. 4 (1911) 1169; Merr. En. Philip. 1 (1923) 140; Netmes, Reinwardtia 1 (1951) 263; ibid. 2 (1954) 374. — Fig. 120. Rhizome probably shortly creeping. Stems triquetrous, sparsely scabrid just below the inflorescence, otherwise smooth, 25-50 cm by 1/,-1 mm, surrounded below the leaves by the fibrous remains of old leaf-sheaths. Leaves basal and one half-way up the stem, exceeding the stem, long-attenuate, rigid, with revolute margins, scabrid in the upper part, grey-green with a metallic hue, 3-7 mm wide; cauline leaf long-sheathing. Inflorescence spiciform (or almost so when its lowest node bears 2 spikelets), 2-4 cm long. Lowest bract foliaceous, patent, much overtopping the inflorescence, shortly sheathing, upper one(s) much reduced. Spikelets 2-10, androgynous, subglobose, dense, sessile or on very short included peduncles, 5-8 mm long, the ¢ part about as long as the 9, but inconspicuous when the utricles are fully developed. Glumes ovate or ovate-lanceolate, acutish or ob- tuse, nerved, minutely ciliolate, otherwise glabrous, brownish with whitish hyaline margins, 1'/.-2 mm long, the midnerve excurrent in a hispidulous, 3/,-1 mm long awn. Urricles much overtopping the glumes, trigonous, rhomboid-lageniform, with FLORA MALESIANA [ser. I, vol. 91 150 prominent angles and somewhat concave faces, subcoriaceous, patulous, strongly many-nerved, straight, glabrous, smooth except for a few setae in the upper part, not stipitate, curved-tapering to the base, rather gradually narrowed into the beak, greenish brown, 4!/,—5 by 11/,-13/, mm; beak com- pressed, bidentate, 1'/,-2 mm long, with slightly oblique mouth. Nut trigonous, broadly rhomboid, with thickened angles and concave faces, curved- tapering downwards to a short cylindric stipe and upwards to a stout, cylindric, !/,-"/, mm long and 1/,-3/, mm wide neck, truncate at the apex, c. 21/, by 11/, mm. Style-base slightly thickened, centred in the hollowed apex of the nut. Stigmas 3. Distr. Malesia: N. Borneo (Sabah: Lahad Datu, Mt Silam; Sandakan) and SW. Philippines (Palawan: Mt Pulgar). Ecol. Common in wet, sandy, gravelly soilamong shrubs bordering streams in the hills at 150-600 m. Note. Only known from the type collection, ELMER 13146 from Palawan and SAN 37905 and 43845 from Sabah. 30. Carex rhynchachaenium CLARKE in Merr. Publ. Gov. Lab. Philip. n. 35 (1906) 5; Philip. J. Sc. 2 (1907) Bot. 108; Kew Bull. add. ser. 8 (1908) 79; KUK. Pfl. R. Heft 38 (1909) 480; Philip. J. Sc. 6 (1911) Bot. 62; Merr. En. Philip. 1 (1923) 142; NELMES, Reinwardtia 1 (1951) 368; RAym. Mém. Jard. Bot. Montréal n. 53 (1959) 65. — C. hatusi- mana OHWI, Jap. J. Bot. 7 (1934) 196; Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 344, f. 6, t. 9 f. 7; KoyaAMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 21, t.3; AkryAMA, Car. Far East. Reg. Asia (1955) 178; RaymM. Mém. Jard. Bot. Montréal n. 53 (1959) 65, f. 15, map p. 102, f. 29. — Fig. 120. Rhizome short, cespitose. Stems central, more or less hidden amongst the leaves, erect, triquetrous, scaberulous above, 5—10(-30) cm by #4/,—/, mm, surrounded below the leaves by brown, fibrous remains of old leaf-sheaths. Leaves basal, much longer than the stems, plicate to flat, long-attenuate, scabrid on the margins and upper surface, 1-3 mm wide; sheaths pubescent. Spikelets 3-6, upper approximate, lower distant, erect or suberect, cylindric, lax-flowered; terminal spikelet 4, 1(-2) cm long, 1 mm thick, lateral ones 2 or with some ¢ flowers at the top, 1/,-2 cm long, 3-4 mm thick; peduncles smooth or scaberulous, lower exserted. Lower bracts foliaceous, much exceeding the inflorescence, upper much reduced. Glumes elliptic-ovate to oblong-lanceolate, acute or with rounded apex, slenderly nerved, with not or scarcely excurrent midnerve and hyaline margins, erose-ciliate towards the apex, 2-3 mm long. Utricles lageniform, trigonous, broadest c. 2 mm from the base, subcoriaceous, straight, suberect, strongly multinerved, more or less puberulous above, stramineous or greenish, shortly stipitate, 5'/,-6'/. by 1'/,-1'/. mm; beak 3/,-1 mm long, bidenticulate. Nut rhomboid, tri- quetrous, with faces concave below, stoutly stipitate, the apex subabruptly contracted into a cylindric, truncate neck c. 1 mm long and 3/, mm broad, the whole nut c. 4 mm long. Style-base scarcely thickened, centred in the apical hollow of the nut. Stigmas 3. Distr. Formosa, Tonkin, Annam; in Malesia: Philippines (Luzon: Kalinga, Pampanga, Bataan, Benguet, Laguna; Mindanao: Bukidnon, Zam- boanga). Ecol. On forested ridges in and near the mossy forest, 800-2100 m. Note. I have not seen CLEMENS 34431 from Borneo (Mt Kinabalu?) which may belong here (cf. NELMES, /.c. 369). 11. Section Mitratae KUK. Pfi. R. Heft 38 (1909) 458; NELMEs, Reinwardtia | (1951) 371; RAymM. Mém. Jard. Bot. Montréal n. 53 (1959) 52, 60. — Sect. Praecoces CHRIST, Bull. Soc. Bot. Belg. 24 (1885) 14, nomen; MACKENZ. N. Am. FI. 18 (1935) 183; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 339. — Type species: Carex mitrata FRANCH. Note. In this section many species have been described which differ only in slight characters and in my opinion several must be combined or deserve at most varietal rank. With the existing keys of KUKENTHAL, OHwI, and Koyama identification appears often impossible. — (Noor.). 31. Carex breviculmis R. Br. Prod. (1810) 242; Boorr, Ill. 4 (1867) 181; Boeck. Linnaea 41 (1877) 209; BENTH. FI. Austr. 7 (1878) 445; CLARKE, F]. Br. Ind. 6 (1894) 746; Kix. Pfl. R. Heft 38 (1909) 469; RipL. Trans. Linn. Soc. Bot. II, 9 (1916) 247; S. T. BLaxe, J. Arn. Arb. 28 (1947) 112; NELMEs, Kew Bull. (1949) 383; Reinwardtia 1 (1951) 373; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489. — Fig. 121. For synonyms see under the varieties. Rhizome short (obliquely descending, woody). Stems tufted, slender, erect or oblique, (obtusely) trigonous, smooth, or scaberulous above, 1-40 cm by '/,-1 mm, clothed at the base by old leaf sheaths or their fibrous remains. Leaves basal and subbasal, from much shorter to much longer than the stems, often thickish, rigid, flat or with recurved or revolute margins, gradually attenuate to the triquetrous tip, smooth or mostly scabrid on margins and keel, 1-4(-6) mm wide. Inflorescence 1979] simple, erect, with 3-7 approximate spikelets (sometimes lowest spikelet remote on a peduncle from the sheath of a basal leaf; in small plants sometimes only one ¢ and one @ spikelet, the other ® spikelets single on a long peduncle arising from the centre of the basal leaves together with the main inflorescence); terminal spikelet 3, sometimes gynaecandrous (in var. perciliata), linear, 5—12(—20) by 1-2 mn, lateral ones 9, sessile or (very) shortly peduncled, subglobose to shortly cylindrical, 5-15 (—30) by 3-4 mm, peduncles smooth or scab- rous; lower bracts usually overtopping the in- florescence, foliaceous or setaceous, shortly sheath- ing, upper much reduced. G/umes ovate or oblong- ovate, acute to obtuse, whitish or brownish, excurrent into a scabrid awn in var. breviculmis, muticous or with a short mucro in the other varieties, whitish or brownish and then with white hyaline margins, 3-nerved in the centre, 2—3!/, mm long. Utricles obtusely or obsoletely trigonous, lanceolate to ellipsoid or obovoid, membranous, except the 2 marginal nerves nerveless or obscurely nerved to more distinctly nerved, subabruptly to gradually beaked, from densely hispidulous to only hispidulous on the margins or glabrous, not or scarcely stipitate, light brown to stramineous or fulvous, 2'/,4 by c. 1 mm. Nut triquetrous, with faces concave below, ellipsoid or ovoid to obovoid, stramineous to fuscous, 1!/,-2!/, by c. 1 mm, suddenly contracted into a very short neck and then suddenly expanded into an annulate apex. Style-base thickened, persistent on the annulus (but when the nuts are falling the style-base probably has disappeared). Stigmas 3. Distr. Widely distributed from the Himalayas to China, Japan and Formosa through Malesia to Australia and New Zealand. Note. The length of the stem varies consider- ably, even on the same plant. The same holds for the nerves on the utricles and for their hairiness, but these characters are rather constant in a single specimen. For these reasons I consider C. brevicul- mis R. BR., C. perciliata NELMES, and C. montivaga S. T. BLAKE to be conspecific, although it is possible to discriminate between them on varietal rank. KEY TO THE VARIETIES 1. Glumes whitish, 2-3 mm long, excurrent into a scabrid awn. Utricles greenish becoming light brown, subabruptly beaked, several-nerved, usually more or less pubescent or hispidulous, 25-3 | mmilong 2: a. var. breviculmis 1. Glumes brownish with white hyaline margin, acute or obtuse, muticous or with a small mucro. Utricles stramineous to fulvous, gradually beaked, nerveless or faintly nerved, hispidu- lous to glabrous, 2!/,-4 mm. 2. Utricles usually hispidulous, at least above on the margins, 2'/,-3 mm . b. var. perciliata Cy PERACEAE—II (Kern & Nooteboom) 2. Utricles glabrous, rarely obscurely hispidulous on the margins above, 3'/,-4 mm long c. var. montivaga a. var. breviculmis. — C. /euchochlora BUNGE, En. Pl. Chin. Bor. (1833) 68; Koyama, Act. Phytotax. Geobot. 16 (1955) 9; YosHikAwa, Ic. Jap. Car. 1 (1957) 104, t. 52; RAym. Mém. Jard. Bot. Montréal n. 53 (1959) 60. — C. royleana Nees in Wight, Contr. (1834) 127; Boorr, Ill. 1 (1858) 6, t. 19; Camus, FI. Gén. I.-C. 7 (1922) 195. — C. eggytera STEUD. Syn. 2 (1855) 220. — C. breviculmis ssp. royleana Kix. Pfl. R. Heft 38 (1909) 469; Philip. J. Sc. 6 (1911) Bot. 62, incl. var. kingiana KUxK.; Me_rr. En. Philip. 1 (1923) 137. — C. jackiana Bootr var. tumens KUxK. in Hochr. Candollea 6 (1936) 432. — C. rugata (non OHw1) NELMES, Reinwardtia 1 (1951) 378. — C. conorrhyncha NELMES, Kew Bull. (1956) 182. — Fig. 121. Distr. As for the species; in Malesia: E. Java (Mt Tengger and Mt Jang), SW. Celebes (Mt Bon- thain), N. Borneo (Mt Kinabalu), Philippines (Luzon), and New Guinea. Ecol. Open grassy slopes, among shrubs, 1750- 3900 m. b. var. perciliata KUxK. Pfl. R. Heft 38 (1909) 469. — C. breviculmis: CLARKE, J. Linn. Soc. Bot. 37 (1904) 16. — C. bulbostylis var. ciliato-marginata KUK. et var. hispidula KUxK. Bot. Jahrb. 70 (1940) 464; Bull. Jard. Bot. Btzg III, 16 (1940) 318. — C. tricholoma S. T. BLake, J. Arn. Arb. 28 (1947) 110, f. 3A. — C. brevis S. T. BLAKE, /.c. 111, f. 3B; NELMES, Kew Bull. (1950) 202; Reinwardtia 1 (1951) 375. — C. perciliata (KUK.) NELMES, Kew Bull. (1946) 26; ibid. (1949) 383, 391; Reinwardtia 1 (1951) 374. — Fig. 121. Distr. Malesia: N. Borneo (Kinabalu), SW. Celebes (Latimodjong Range: Mt Rante Mario), New Guinea (W. New Guinea: Star Mts; Papua and Territory of New Guinea). Ecol. Rock crevices, open bare ground, wet grassland, forest glades, 2400-4200 m. c. var. montivaga (S. T. BLAKE) Noor., comb. nov. — C. montivaga S. T. BLAKE, J. Arn. Arb. 28 (1947) 109; NeLMes, Kew Bull. (1949) 383; Reinwardtia 1 (1951) 377. — C. bulbostylis Kx. Bot. Jahrb. 69 (1938) 264; Bull. Jard. Bot. Btzg III, 16 (1940) 317, excl. var. et specim. Born., non MACKENZ. 1915. — Fig. 121. Distr. Malesia: New Guinea (Lake Habbema, Mt Wilhelmina, Star Mts, Mt Sarawaket). Ecol. Bogs, alpine grassland, wet grassy slopes, along water, also in mossy forest, 3200-3450 m. 32. Carex dolichostachya HAYATA, Ic. Pl. Form. 10 (1921) 61, f. 38; O#w1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 375; KoyaMaA, Bull. Arts & Sc. Div. Ryukyu Un. (Math. & Nat. Sc.) n. 3 (1959) 72; AkryaMA, Car. Far East. Reg. Asia (1955) 204, 152 FLORA MALESIANA [ser. I, vol. 91 t. 208, f. 2. — C. ligata Bootr var. nexa KUK. Philip. J. Sc. 6 (1911) Bot. 63; MerR. En. Philip. 1 (1923) 39, quoad specim. cit., non C. nexa BOOTT. — C. multifolia OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B5 (1930) 264; ibid. B11 (1936) 373; NELMEs, Reinwardtia 1 (1951) 371; AktyAMa, Car. Far East. Reg. Asia (1955) 202, t. 206; YOSHIKAWA, Ic. Jap. Carex 1 (1957) 122, t. 61. — C. foliosissima (non F. SCHMIDT) FRANCH. Carex As. Or. (1898) n. 232; Kuk. Pfi. R. Heft 38 (1909) 478. — Fig. 121. Rhizome short, cespitose, or shortly stoloni- ferous. Stems arising from basal leaf-axils, trigo- nous, smooth or sparsely scaberulous above, (15—)30-60 cm by 1/,-1 mm, bearing a few short- bladed, bract-like leaves. Basal /eaves about as long as the stems, flat, with scabrid margins and under surface above, 3-10 mm wide; sheaths reddish brown to spadiceous, older fibrous. Spikelets 3-7, distant, erect or suberect; terminal spikelet g, linear, (1'/,-)3-6 cm by 1'!/,-2 mm, lateral 2 or with a few ¢ flowers at the top, lax- flowered, (11/,-)3-5 cm by 2'/,-3'/, mm, on exserted, smooth or slightly scaberulous peduncles. Bracts vaginiform with a short blade or reduced to long, subampliate, bladeless sheaths. Glumes oblong or oblong-obovate, truncate or rounded at the erose-ciliate apex, translucent, whitish to brownish, 21/,-3 mm long, the 3-nerved centre excurrent in a short, hispidulous awn c. 1/, mm long. Utricles fusiform, trigonous with flat faces, straight, membranous, suberect, strongly many- nerved, hispidulous especially above, greenish to light brown, 3-4 by c. 1 mm; beak conical, 1 mm long, bidenticulate. Nut ellipsoid to oblong- ellipsoid, triquetrous with shallowly concave faces, stoutly stipitate, contracted at the apex and re- expanded into a discoid, !/, mm broad annulus, c. 2 by 1 mm. Style-base pyramidally thickened. Stigmas 3. Distr. Japan, Ryu Kyu Is., Formosa; in Malesia: Philippines (Luzon: Kalinga, Bontoc, Rizal). Ecol. Clearings and along edges of primary forest, 1200-1600 m. Notes. The distinction between C. multifolia OHWI, common in the montane regions of Japan, and C. dolichostachya HAYATA, occurring from the Ryu Kyu Is. and Formosa to Luzon, is mainly made on account of the different colour of the basal leaf-sheaths. I have followed KoyAMA, who united the two as neither this character nor the other slight differences are constant. HATUuSIMA, Mem. Fac. Agric. Kagoshima Un. 5, 3 (1966) 59, referred specimens from the Batan Is. (N. Luzon), mentioned above as C. ligata var. nexa KUK. (non C. nexa Boott), to C. sociata Boott in A. Gray, Bot. Jap. (1859) 420; Ill. 4 (1867) 200; OHwI, Cyp. Japon. 2 (1943) 376. If this identity is true, C. sociata Bootr would be the correct name for this species. A specimen from Sarawak (Mt Murud, 2400 m, NooTEeBooM 2030) undoubtedly belongs to sect. Mitratae, possibly in the vicinity of C. dolichos- tachya. The utricles, however, are longer together with the nuts, and the leaves are smaller. — (Noot.) 33. Carex formosensis LEv. & VAN. Mém. Soc. Nat. Sc. Nat. Math. Cherb. 35 (1906) 216, et in Fedde, Rep. 5 (1908) 31; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 345; NeELMEs, Rein- wardtia 2 (1954) 379; AKIYAMA, Car. Far East. Reg. Asia (1955) 178, t. 180. — C. ligata Bootr var. formosensis (LEV. & VAN.) KUx. Pfl. R. Heft 38 (1909) 474; Merr. En. Philip. 1 (1923) 139. — Fig. 121. Rhizome short, cespitose. Stems central, erect, trigonous, smooth, 10—-25(—50) cm by 1/,-1 mm. Leaves basal, shorter to longer than the stems, flat, long-attenuate, with scabrid margins, 2-4(-6) mm wide, surrounded at the base by brown, fibrous remains of old leaf-sheaths. Spikelets 3-7, sub- approximate or lower more distant, erect or sub- erect, cylindric; terminal spikelet usually 4, 1-2 cm long, 2-3 mm thick, lateral ones single at the nodes, usually gynaecandrous, with very few 3 flowers at the base, subdensely flowered, 3!/,-5 mm thick, their peduncles smooth, upper shortly, lower rather long-exserted. Lower bracts folia- ceous, shorter to longer than the inflorescence, lower long-sheathing, upper much reduced. Glumes oblong, truncate to bilobed-emarginate, 11/,-2 mm long, nerveless except for a greenish, strongly 3-nerved central stripe excurrent in a hispid, up to 1'/, mm long awn. Utricles rhomboid, slightly lageniform or fusiform, trigonous, subcoriaceous, broadest below the middle, straight, suberect, multinerved, very sparsely puberulous, strami- neous or greenish, shortly stipitate, c. 31/. mm by 1 mm; beak c. 1 mm long, bidentate. Nut rhom- boid, trigonous, with deeply concave faces, shortly stipitate, slightly constricted at the apex and slightly re-expanding into a rather discoid-annulate 1/,—1/, mm wide apex, the whole nut c. 2(—2!/.) mm long. Style-base slightly thickened. Stigmas 3. Distr. Korea, Japan (Kyushu, Hondo), For- mosa; in Malesia: Philippines (Luzon: Benguet Subprov., Ilocos Norte). Ecol. Forested ridges, along streams and trails, on cliffs and steep banks in and near the mossy forest, 1000-2400 m. Vern. Egegedan, Bon., silak, Ig. _ 34. Carex tristachya THUNB. FI. Jap. (1784) 38; ScHKuHR, Riedgr. 2 (1806) 48, t. Ww f. 109; Boott, Ill. 4 (1867) 131, t. 424; Kix. Pfl. R. Heft 38 (1909) 471; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 363; AkryAMA, Car. Far East. Reg. Asia (1955) 195, t. 198; YosHIKAwa, Ic. Jap. Carex 1 (1957) 118, t. 59. The few Malesian collections belong to: 1979] Cy PERACEAE—II (Kern & Nooteboom) 153 var. pocilliformis (Bootr) KUx. Pfl. R. Heft 38 (1909) 473, t. 75, f. A-F; Philip. J. Sc. 6 (1911) Bot. 62; Merr. En. Philip. 1 (1923) 142; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 364; Yosui- KAWA, Ic. Jap. Carex 1 (1957) 120, t. 60. — C. pocilliformis Boor, Ill. 4 (1867) 175, t. 593; NeELMES, Kew Bull. (1949) 391; Reinwardtia 1 (1951) 381; AKryAMA, Car. Far East. Reg. Asia (1955) 196, t. 199. — Fig. 121. Rhizome short. Stems densely tufted, very slen- der, erect, trigonous, smooth, 10-40 cm by 1/,-1 mm, clothed at the base with dark brown, more or less fibrous remains of old leaf-sheaths. Leaves basal and subbasal, as long as or shorter than the stems, rather rigid, flat, gradually attenu- ate, 11/4 mm wide. J/nflorescence with 3-7 spikelets, erect, fastigiate or lower spikelets more distant on exserted peduncles; terminal spikelet gd or gynaecandrous, cylindrical, '/,-3 cm by 1/,-1 mm, lateral ones 2, slenderly cylindrical, rather loosely flowered, 1-3 cm by 2-2'/, mm; peduncles smooth. Lower bracts foliaceous, shorter than to slightly exceeding the inflorescence, upper reduced. G/umes of the 3 spikelet cup-shaped (the margins more than halfway connate in front) but not so in gynaecandrous spikelets, muticous, those of the 2 spikelets ovate or oblong-ovate, rounded at the ciliolate apex, glabrous, muticous or mucronulate, light brown with whitish hyaline margins and 3-nerved central stripe, 2 mm long. Stamens monadelphous (filaments connate almost throughout their length), not always so in lower flowers. Utricles trigonous, ellipsoid, membranous, many-nerved, sparsely pubescent, suberect, re- curved at the top, shortly stipitate, gradually tapering above, green to brownish, 2!/,-3 by 3/,-1 mm; beak subconical, glabrous or his- pidulous-margined, bidenticulate, !/, mm _ long. Nut oblong-ovoid or oblong-ellipsoid, triquetrous with faces concave below, stipitate, c. 2 mm long, stramineous to dark brown, rounded above and abruptly contracted into a short neck which is suddenly expanded into the annulate apex. Style- base pyramidally thickened, persistent. Stigmas 3. Distr. Var. tristachya occurs in Japan, Korea, Central and Eastern China, var. pocilliformis is also in Japan and Korea, but extends more southwards, through the Ryu Kyu Is. and Formosa to Malesia: Philippines (Luzon: Benguet: Mt Pulog), Borneo (Mt Kinabalu), and New Guinea (NE.: Mt Sarawaket). Ecol. Open meadows and grassy slopes, 2200- 2700 m. Notes. Carex tristachya is remarkable for its monadelphous anthers, a feature very rare in the genus (according to BARRos also found in C. acaulis D’URV. of the Falkland Is. [cf BRONGN. in Duperrey, t. 28, f. A; RoEPER, Zur Flora Mecklenb. 2 (1844) 16!)). There is no agreement on the taxonomical status of C. pocilliformis Bootr. Whereas CLARKE (J. Linn. Soc. Bot. 36, 1904, 315) did not distinguish it from C. tristachya, it was reduced to varietal rank by KUKENTHAL, OHWwI, and others, but main- tained as a distinct species by NELMES and AKIYAMA. The main difference is to be found in the glumes of the 3 flowers, which are said to have free margins in typical C. tristachya. I think the difference is only gradual, as in the latter the margins of the glumes are also connate, though at the very base only. Glumes with united margins are very rare in the genus; they are also known in some North American species belonging to sect. Phyllostachyae TUCKERM. (see MACKENZ. N. Am. FI. 18, 1935, 174). Carex tristachya is very near to C. mitrata FRANCH. & SAV., the main difference being in the 3 spikelets, of which the glumes are infundibuli- form and the filaments connate in C. tristachya. But in C. tristachya var. tristachya the margins of the glumes are connate at the very base only, and sometimes they are free. Besides, even in var. pocilliformis, 1 found perfectly free filaments and anthers, especially in the lower flowers (e.g. MERRILL 6629). It is very well possible that the mentioned characters in the ¢ spikelets are not very important, and that C. tristachya and C. mitrata have to be considered as subspecies of a single species. — (Noort.) 12. Section Radicales (KUK.) NELMES, Reinwardtia | (1951) 389; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 75. — Sect. Digitatae Fries subsect. Radicales Kix. Pfl. R. Heft 38 (1909) 480. Type species: Carex radicalis BooTt. 35. Carex malaccensis C. B. CLARKE, FI. Br. Ind. 6 (1894) 722; J. Linn. Soc. Bot. 37 (1904) 9; RipL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 116; Kix. Pfil. R. Heft 38 (1909) 289; Rip. Fl. Mal. Pen. 5 (1925) 183; Netmes, Reinwardtia 1 (1951) 257. — Fig. 121. Rhizome shortly creeping, woody, covered with brown, readily fraying scales. Stems tufted, slender, triquetrous, very narrowly winged above, smooth below, slightly scabrid on the angles above, up to 50 cm by 1-1'/, mm. Leaves basal and subbasal, shorter to longer than the stems, stiff, strongly 154 FLORA MALESIANA [ser. I, vol. 91 Fig. 129. Carex speciosa KUNTH. a. Habit, x '/,, b. spikelet, lower part 2, upper part g, x 11/2, c. glume, d. fruit in utricle, e. fruit, f. young stamens in glume, g. glume, 4. seemingly connate filaments, i. filaments, all x 13 (WAITz s.n.). 1979] Cy PERACEAE—II (Kern & Nooteboom) 155 revolute when dry, long-attenuate, greyish green or glaucous, scabrous-asperous in the apical part, 2-6 mm wide; sheaths brown, membranous in front, outer ones bladeless. /nflorescence racemiform, consisting of 1-4 remote, subglobose or pyramidal heads on short (the lowest rather long), smooth or scabrid peduncles, each head composed of 1-4 crowded, sessile spikelets. Lower bracts foliaceous, much exceeding the inflorescence, not auricled, not or lowest very shortly sheathing, ultimately horizontally spreading, upper much reduced. Spikelets androgynous, the 3 part about as long as or longer than the 9, ovoid, patent, 5-10 mm long and wide. G/umes broadly ovate to oblong-ovate, obtuse, membranous, strongly nerved, ciliolate above, otherwise glabrous, white or light brown, 2'/,-3 mm long, the midnerve excurrent in a stoutish, scabrid, '/,-1 mm long awn. Utricles trigonous, rhomboid or obovoid, mem- branous, suberect or patulous, strongly many- nerved, rather abruptly contracted into the beak, glabrous, narrowly marginate or winged, scabrid on the margins at the apex, stramineous, 4—6%/, by 1*/,—21/, mm; beak linear-conical, flattened, curved inwards, bidenticulate, with oblique mouth 1-2 mm long. Nut triquetrous, pyriform, scarcely stipitate, brown but densely overlain white-scurfy, 2—21/, by 13/, mm. Style-base pyramidally thickened, per- sistent on the nut. Stigmas 3. Distr. Malesia: Malay Langkawi Is.). Ecol. On limestone along the sea-shore, on limestone rocks, at low altitude. Notes. Like Fimbristylis malayana Onwi1 this interesting species is apparently endemic in the Langkawi Is., here exclusively growing on lime- stone. RIDLEY (/.c.) recorded it for Thailand, but it is not mentioned for that country either by NELMES or RAYMOND. Its nearest ally is C. leucantha ARN. ex BooTrt, Proc. Linn. Soc. 1 (1845) 257; Trans. Linn. Soc. 20 (1846) 135; Ill. 1 (1858) 10, t. 28, from Ceylon and India, which differs by the long-sheathing lower bracts and the hairy utricles. Peninsula (Kedah, 36. Carex ramosii KUxk. in Fedde, Rep. 8 (1910) 8; Philip. J. Sc. 6 (1911) Bot. 63; Merr. En. Philip. 1 (1923) 141; Netmes, Reinwardtia 1 (1951) 259; ibid. 2 (1954) 374. — Fig. 121. Rhizome shortly creeping, woody, covered with brown, readily fraying scales. Stems very slender, erect but often curved, triquetrous, slightly incrassate just below the inflorescence, smooth, or slightly scabrid at the top, surrounded below the leaves by a few, bladeless, brown sheaths, 10-25 cm by '/,-1 mm. Leaves basal, up to twice as long as the stems, rigid, flat, with revolute margins when dry, long-atienuate, greyish green, scabrid in the apical part, 2-5 mm wide. Inflorescence either a single head of 1-3 sessile spikelets, ovoid or oblong-ovoid, 1-2 cm long, or spiciform with 1-2 additional subsessile heads near the base of the stem in the axil of a leafy bract very similar to and as long as the leaves. Bracts not sheathing, amplexi- caul by ciliolate auricles, the lowest of the terminal head foliaceous, 5—-10(-25) cm long. Spikelets androgynous, the ¢ part longer than the 2 but finally hidden by the fully developed utricles, 6-10 by 5-7 mm. Glumes ovate-lanceolate, obtuse or acutish, membranous, strongly nerved, glabrous, stramineous with white margins, 3—4'/, mm long, the midnerve excurrent in a minutely scabrid, 1-17/, mm long awn. Utricles trigonous, ellipsoid- rhomboid, subcoriaceous, suberect, strongly many- nerved, distinctly winged almost all round, scabrid- margined, gradually narrowed into the beak, glabrous, stramineous or brownish, 6-8 by 2- 2?/, mm (wings included); beak compressed, sub- conical, bidenticulate, with oblique mouth, 3 mm long. Nut triquetrous, rhomboid-ellipsoid, sessile, livid, 3 by 13/,-2 mm. Style-base pyramidally thickened, persistent on the nut. Stigmas 3. Distr. Malesia: Philippines (Luzon, Mindanao). Ecol. In forest at medium altitude. 37. Carex speciosa KUNTH, En. 2 (1837) 504; Mia. FI. Ind. Bat. 3 (1856) 348; Boorrt, Ill. 1 (1858) 53; Boeck. Linnaea 40 (1876) 388, incl. var. minor BoeEck.; CLARKE, FI. Br. Ind. 6 (1894) 729; KUkK. Pfl. R. Heft 38 (1909) 481, incl. var. courtallensis KUkK. et var. angustifolia KUx.; Camus, Fl. Gén. I.-C. 7 (1922) 197; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 63; NELMES, Reinwardtia 1 (1951) 390; ibid. 2 (1954) 381; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 160, incl. var. angustifolia; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 80; Dansk Bot. Ark. 23 (1965) 260; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 488, 492. — C. concolor Nees in Wight, Contr. (1834) 125, non R. Br. 1823. — C. courtallensis NEES ex Boott, Ill. 1 (1858) 52, t. 138, incl. var. angustifolia Bootr; CAMus, FI. Gén. I.-C. 7 (1922) 197, f. 28, 1-8; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 78, f. 19. — C. longispica Boeck. Allg. Bot. Zeitschr. 2 (1896) 174. — C. speciosa var. abscondita Kix. Bull. Jard. Bot. Btzg III, 16 (1940) 318. — C. stenura Newtmes, Kew Bull. (1950) 202; Reinwardtia 1 (1951) 392. — Fig, 121, 129. Rhizome short, descendent, woody. Stems tufted, erect or suberect, slender, triquetrous, often narrowly winged at the top, smooth or finely scaberulous, 5-50 cm by !/,-1!/, mm, surrounded below the leaves by fuscous sheaths and fibres. Leaves subbasal, longer to much longer than the stems, stiff, flat (but margins revolute when dry), scabrous on the margins, asperous on the upper surface towards the long-attenuate apex, greyish or glaucous-green, 3-10 mm wide. Spikelets 1-3(-4), 5-20 cm distant from one another, lowest some- times arising from a basal leaf-sheath, androgy- nous, cylindric, 2-8 cm long, rather laxly 6—-18- flowered, on shortly exserted smooth peduncles, 156 FLORA MALESIANA [ser. I, vol. 91 the 2 part 3-7 mm thick, half as long to about as long as the slender ¢ part. Bracts foliaceous, usually overtopping the inflorescence, sheathing. Glumes of the ¢ flowers with free margins or rarely the margins connate in front in the lower 2/;—'/,, those of the 2 flowers triangular-broadly ovate, rounded to subacute, translucent, many-nerved, with prominent midrib, muticous, minutely cilio- late above, long-persistent, 2!/,-3 mm _ long. Utricles ovoid-ellipsoid or ovoid-pyramidal, tri- quetrous, coriaceous, suberect, strongly many- nerved, straight or slightly curved, with whitish- pubescent margins, greenish-stramineous, 4-7 by 1—2'/, mm, spongy at the base, gradually narrowed into the short, bidenticulate beak. Nut obovoid or ellipsoid, triquetrous, shortly stipitate, abruptly shortly beaked, pale to dark brown with promi- nent pale angles, 3-4 by 1'/,-2 mm. Style-base pyramidally thickened, ciliolate, persistent. Stig- mas 3. Distr. Widely distributed in India (from Nepal to Sikkim, also in S. India), S. China (Yunnan), Thailand, Indo-China; in Malesia: Sumatra (Atjeh, E. Coast Res.), Malay Peninsula (Ulu Kelantan: Bertam, Gua Musang), SE. Borneo (N. of Bandjermasin, once), and Java (rare in W. and E., more common in Central Java), also in Kangean I. Ecol. Primary forest, often in teak-forest, distinctly preferring seasonal climatic conditions, in Borneo on dry serpentine rock (once), at low and medium altitude, from sea-level up to 1500 m. Vern. Java: ilat djepun, S. Notes. According to RAYMOND, /.c., the leaves in C. speciosa are 11/,-2 mm wide, the number of spikelets 1-4, and the utricles 4-5 mm long. He kept C. courtallensis apart from C. speciosa on account of its broader leaves, the greater number of spikelets, and the larger utricles. The type- collection of C. speciosa (WALLICH 3391) does not answer RAYMOND’s description of this species. I agree with Nevmes that C. courtallensis is con- specific with C. speciosa; the specimens from Sumatra and Java agree very well with the type of C. courtallensis (W1GHT 991 in K). The Bornean specimens are remarkable by the narrow leaves, the long, tail-like ¢ part of the spikelets, and especially by the infundibuliform glumes of the ¢ flowers (the last character not mentioned by Netmgs). I have not seen the Tonkin specimens NELMEs (1955) referred to C. speciosa var. angustifolia (BootT) Kix. They have leaves only 1'/,-2 mm wide, and utricles 3!/,-4 mm long. It is not clear how Newmes distinguished between this variety and his C. stenura. Another member of sect. Radicales from Lower Burma was described as C. pterocaulos NELMES in Hook. Ic. Pl. 35 (1947) t. 3467; Mém. Mus. Nat. Hist. Paris B4 (1955) 159, according to NELMES a very distinct species because of its widely winged stems, its sometimes empty bract, and the very stout 2 part of its spikelets. In some specimens collected in Thailand, Chiengmai, Doi Chiengdao, at 1200 m (SLEUMER & SMITINAND 1063, L), the stems are strikingly winged (4 mm broad) and the bracts very long and sometimes empty, but the glumes are only 3-31/, mm long, the (young) utricles 4 mm (not 4-5 mm and 6-7 mm respec- tively); the 2 part of the spikelets is therefore not stouter than in C. speciosa. I doubt whether C. pterocaulos is specifically distinct from C. speciosa. The polymorphism of the group is badly in need of special study. The record of C. speciosa var. angustifolia for NE. New Guinea, based on CLEMENS 7909a (KUK. Bot. Jahrb. 70, 1940, 465) is very doubtful for plant-geographical reasons. The filaments are often coherent to such a degree as to give the impression of their being connate. 38. Carex tricephala Boeck. Flora 58 (1875) 263; KU. Pfl. R. Heft 38 (1909) 289; Koorp. Exk. FI. Java 1 (1911) 205; KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 316; Net~mes, Kew Bull. (1950) 190; Reinwardtia 1 (1951) 256; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 117; KoyAMaA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 15, t. 1; RAymM. Mém. Jard. Bot. Montréal n. 53 (1959) 76; Dansk Bot. Ark. 23 (1965) 260; KERN in Back. & Bakh. f. FI. Java 3 (1968) 491. — C. madoerensis CLARKE, J. Linn. Soc. Bot. 37 (1904) 15; Kew Bull. add. ser. 8 (1908) 77. — C. thorelii CAMus, Not. Syst. 1 (1910) 295; Fl. Gén. I.-C. 7 (1922) 196; NELMEs, Kew Bull. (1946) 15, 23. — C. plesiocephala Turr. Kew Bull. (1910) 385; ibid. (1912) 427. — C. his- pidangula KoyAMA, Nat. Canad. 82 (1955) 200, t. 1. — Fig. 121. Rhizome shortly creeping, woody, covered with the fibrous, fuscous remains of old scales. Stems slender, triquetrous, narrowly winged just below the inflorescence, smooth below, antrorsely scabrous-setulose above, 10-50 cm by 1 mm (up to 2 mm in the winged part), the base clothed with the fibrous remains of old leaf-sheaths. Leaves basal and 1-2 cauline, shorter than to much exceeding the stems, rigid to rather weak, flat, with revolute margins when dry, long-attenuate, scabrid in the upper part, light green, 3-10 mm wide; cauline leaves long-sheathing; ligule short, triangular. Inflorescence spiciform, consisting of (1-)3(-4) dense, sessile, ovoid or subglobose spikelets 0-5 cm distant, and 6-10(—12) mm long and wide. Lower 1-2 bracts foliaceous, usually much exceed- ing the inflorescence, not sheathing, amplexicaul by ciliolate auricles, upper much reduced. Spikelets androgynous, the ¢ part about as long as the 2, but inconspicuous when the utricles are fully developed. Glumes oblong-ovate to ovate, membranous, nerved, densely setulose, white, or brownish with white margins, 2'/,4'/, mm long; midnerve excurrent in a scabrid, '/,-1 mm long awn. 1979] Cy PERACEAE—II (Kern & Nooteboom) 157 Utricles trigonous, ovoid or rhomboid-ovoid, membranous, patent, slenderly nerved, rather abruptly narrowed into the beak, densely hispidu- lous in the upper 2/3, whitish or brownish, 4-6 by 14/,-2'/, mm; beak conic-cylindric, bidentate, with slightly oblique mouth, 1-2'/, mm long. Nut triquetrous, ellipsoid, rhomboid, or slightly obo- void, scarcely stipitate, cinereous, dark brown on the angles, 2'/,-3 by 13/,-2 mm. Style-base pyramidally thickened, persistent on the nut. Stig- mas 3. Distr. Yunnan, Upper Burma, Thailand, Indo- China; in Malesia only known from Madura I. off NE. Java (type locality !). Ecol. In grassy fields on heavy calcareous marl, in muddy places in teak-forests, at low altitude (up to 200 m), obviously bound to a distinct seasonal climate and its range apparently coin- ciding with that of teak. See Netmes 1951, /.c. Note. NELMEs placed this species, together with C. malaccensis and C. ramosii in sect. Stramentitiae, but as RAYMOND 1959, /.c., pointed out, it has very little in common with the other members of that section. 13. Section Trachychlaenae DREJER, Symb. Caric. (1844) 9; KUK. Pfl. R. Heft 38 (1909) 415. — Sect. Anomalae (non CAREY) NELMES, Reinwardtia | (1951) 413; RAymM. Mém. Jard. Bot. Mon- tréal n. 53 (1959) 54, 71. Type species: Carex hispida WILLD. (lectotype). 39. Carex maculata Boott, Trans. Linn. Soc. 20 (1846) 128; Ill. 1 (1858) 9, t. 26; Boeck. Linnaea 41 (1877) 191; BentH. FI. Austr. 7 (1878) 447; CLARKE, Fl. Br. Ind. 6 (1894) 735; Kix. Pfl. R. Heft 38 (1909) 427, incl. var. neurochlamys KUK.; OxnwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 434; Kx. Bull. Jard. Bot. Btzg III, 16 (1940) 317, incl. var. sanguineo-squamata KOK. et f. humilior KUuKk.; Onwi & KoyAMA, Misc. Rep. Nat. Sc. Mus. Tokyo n. 5 (1952) 1, t. 1; AKryAMA, Car. Far East. Reg. Asia (1955) 120, t. 105, f. 1; YosHIKAWA, Ic. Jap. Carex 2 (1958) 206, t. 103; NELMES, Kew Bull. (1950) 205; Reinwardtia 1 (1951) 414; Ray. Mém. Jard. Bot. Montréal n. 53 (1959) 71; Dansk Bot. Ark. 23 (1965) 259; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489; STEEN. Mt. FI. Java (1972) pl. 14-5. — C. neurochlamys F.v.M. Fragm. 8 (1874) 258; NeELMEs, Kew Bull. (1949) 391; Rein- wardtia 1 (1951) 413. — C. samoensis BOECK. Bot. Jahrb. 25 (1898) 588. — C. elibates NELMES, Kew Bull. (1937) 353; ibid. (1950) 205; Reinwardtia 1 (1951) 416. — C. pruinosa (non Boott) Kx. Bull. Jard. Bot. Btzg III, 16 (1940) 317. — C. pruinosa Boott f. tristigmatosa Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 62. — Fig. 121. Rhizome short, woody. Stems densely tufted, slender, erect, trigonous, smooth, (5—)15-—60 cm by 1-1'/, mm, surrounded below the leaves by a few ferrugineous or brownish, bladeless sheaths or their fibrous remains. Leaves subbasal, shorter to longer than the stems, flat with often revolute margins, greyish green, scabrous towards the _ long- attenuated apices, (1-)3-8 mm wide; sheaths pale to ferrugineous, membranous in front. Lower bracts sheathing, leaf-like, usually overtopping the inflorescence. Spikelets (2-)3—5(—6), upper approxi- mate, lower distant, cylindric, densely flowered; terminal spikelet ¢, sessile, pale, 1-2 mm thick, lateral ones 9, suberect, dark, 1-4 cm by 3-5 mm, upper sessile or shortly peduncled, lower on short to long, firm, smooth peduncles. Glumes ovate or oblong-ovate, acute or subobtuse, thin, translucent, often minutely ciliolate, ferrugineous to castaneous with narrow whitish-hyaline margins and 3-nerved, pale green central stripe, muticous or scarcely mucronulate, 2-3 mm long. Urtricles ellipsoid, compressed-trigonous, membranous, distinctly several-nerved, suberect to patulous, straight, glabrous but densely ferrugineous-papillose, scarcely stipitate, 2-3(-4) by 1-1%/, mm, sub- abruptly contracted into a very short, up to '/, mm long, entire or slightly emarginate beak. Nut ellipsoid or obovoid, triquetrous, shortly stipitate and beaked, 11/,—2 by 3/,-11/, mm, both stipe and beak usually bent. Style-base oblique, somewhat thickened. Stigmas 3 (rarely 2 in some fruits). Distr. Widely distributed from Ceylon and India through Thailand, China, and Korea to Formosa, Japan and the Ryu Kyu Is., and through Malesia to Queensland, New South Wales, New Caledonia and Samoa; in Malesia: Malay Penin- sula (Pahang), N. Sumatra (Atjeh), Java (West: Mt Papandajan; Central: Diéng Plateau; East: Jang plateau), Lesser Sunda Is. (Flores), Celebes (Minahassa, Poso), and New Guinea. Distr. map: MEUSEL, Vergl. Arealkunde 2 (1943) Karte 32c. Ecol. In swamps, marshes, boggy meadows, wet mountain heaths, open places in mossy woods, 1600-3500 m. Vern. New Guinea: sisik, Tomba, toni, ititu, Mendi, era, teleleme, Onim. Notes. Very variable. I cannot follow NELMES in assigning specific rank to C. neurochlamys and C. elibates. His descriptions are fairly well covered by that of C. maculata. C. elibates has darker coloured glumes and often stiffer and narrower 158 FLORA MALESIANA [ser. 1, wok"9! leaves than typical C. maculata, and for this reason KUKENTHAL distinguished it as var. sanguineo- squamata. The other characters given by NELMES are valueless: in all Carices with a 3 terminal spikelet this spikelet may bear some flowers; the very short beak of the utricle in C. maculata is variable, in length, and so it is in C. elibates. F.v.MUELLER distinguished C. neurochlamys from C. maculata mainly by the non-papillose, strongly nerved utricles, a mistake apparently due to inaccurate observation. According to NELMES the glumes in C. neurochlamys are usually mucronulate, not usually so in C. maculata; J fail to see any difference. KUKENTHAL reduced C. neurochlamys to varietal rank and distinguished it by the stouter stems, the leaves much overtopping the inflores- cence, and the many-nerved, less densely papillose utricles. These characters are not found in the New Guinea specimens referred to C. neurochlamys by NELMES. Carex maculata is closely related to species of sect. Praelongae (C. phacota and C. pruinosa). One collection of N. Sumatra (DE WILDE c.s. 13251) has several fruits with only 2 styles, and becoming as long as 4 mm, thus exactly resembling the fruits of C. pruinosa. These fruits occur in the same spikelet as the ‘normal’ 3-styled fruits. Another deviating feature in this collection is that some of the utricles are not glabrous, but minutely scabrid on the margins. — (Noot.) 14. Section Capitellatae MEINSH. Act. Hort. Petrop. 18 (1901) 280, 309; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 437; NELMES, Reinwardtia 1 (1951) 404; Raym. Mem. Jard. Bot. Montréal n. 53 (1959) 17, 18. — Sect. Rarae CLARKE, Kew Bull. add. ser. 8 (1908) 143. — Sect. Unciniaeformes KUK. subsect. Capitellatae (MEINSH.) KUK. Pfl. R. Heft 38 (1909) 100. — Sect. Extensae Frits subsect. Capitellatae (MEINSH.) KoyAma, J. Fac. Sc. Un. Tokyo III, 8 (1962) 202. Type species: Carex capitellata Botss. & BAL. 40. Carex capillacea Boortr, Ill. 1 (1858) 44, t. 110; Boeck. Linnaea 39 (1875) 37; BENTH. FI. Austr. 7 (1878) 438; CLARKE, FI. Br. Ind. 6 (1894) 713; J. Linn. Soc. Bot. 37 (1904) 7; Philip. J. Sc. 2 (1907) Bot. 107; OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 441; S. T. BLake, J. Arn. Arb. 28 (1947) 101; NELMES, Kew Bull. (1949) 381; Reinwardtia 1] (1951) 405; ibid. 2 (1954) 381; AKryAMA, Car. Far East. Reg. Asia (1955) 42, t. 6, f. 1 A-D; Yosut- KAWA, Ic. Jap. Carex 2 (1958) 218, t. 109; Ray. Mém. Jard. Bot. Montréal n. 53 (1959) 18; KoyAMa, J. Fac. Sc. Un. Tokyo III, 8 (1962) 206; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 488; STEEN. Mt. Fl. Java (1972) pl. 14-8. — C. simplicis- sima F.v.M. Fragm. 9 (1875) 191. — C. rara (non Boott) STAPF, Trans. Linn. Soc. II, Bot. 4 (1894) 246; CLARKE, J. Linn. Soc. Bot. 37 (1904) 7; KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 313; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 64. — C. rara Bootr ssp. capillacea KiK. in Mats. Index Pl. Jap. 2 (1905) 130, quoad synon.; Pf. R. Heft 38 (1909) 102; Philip. J. Sc. 6 (1911) Bot. 58; Merr. En. Philip. 1 (1923) 141; STEEN. Trop. Natuur 19 (1930) 87, 89 f. 14. — Fig. 121, 130. Rhizome short, woody. Stems densely tufted, slender, erect, triquetrous, smooth or slightly scaberulous just below the inflorescence, 5—50 cm by 1/,-3/, mm, clothed below the leaves by brown- ish, bladeless sheaths and their fibrous remains. Leaves subbasal, much shorter than to slightly exceeding the stems, flat or canaliculate, smooth or nearly so, erect, '/,-2 mm wide; sheaths pale, membranous in front. Spikelet solitary, terminal, androgynous, ovoid or shortly cylindric, densely but rather few-flowered, 5-15 mm long, the g part c. 1 mm thick, shorter than to about as long as the 2 part, the latter becoming 4-6 mm thick. Glumes ovate or oblong-ovate, obtuse to acutish, muticous, translucent, ferrugineous to brown with narrow paler margins and 3-nerved, pale central stripe, caducous, 1!/,-3 mm long. Utricles oblong-ovoid or oblong-ellipsoid, rounded at the base, obscurely trigonous, subinflated, membranous, finally widely patent to somewhat reflexed, glabrous, several- nerved, scarcely stipitate, often minutely reddish resinous-dotted, (11/,-)21/,-31/,(-4) by c. 1 mm, rather gradually tapering to a very short, subentire or slightly emarginate beak. Nut ellipsoid or oblong-ellipsoid, triquetrous, shortly stipitate, abruptly short-beaked, stramineous to brownish, 11/,-2'/, by c. 1 mm. Style-base slightly thickened. Stigmas 3. . Distr. From the Himalayan region to Annam, Manchuria, Korea, Japan, and extending as far north as Sachalin (var. sachalinensis Onwt); through Malesia to Australia (New South Wales) and New Zealand; in Malesia: Sumatra (North: Atjeh; Central West: G. Singalang), W. Java (Mt Papandajan), N. Borneo (Mt Kinabalu), SW. Celebes (Mt Rante Mario), Philippines (Luzon), and New Guinea. 1979] Cy PERACEAE—II (Kern & Nooteboom) 159 Ecol. Swamps, wet alpine grasslands, open seepages, wet borders of small streams, marshy places in forests, 2000-4000 m. Vern. New Guinea: ere, tep, Mendi lang. Notes. Closely related to the Indian C. rara Boott, Proc. Linn. Soc. 1 (1845) 284; Ill. 1 (1858) 44, t. 109. Bootr distinguished C. capillacea from C. rara by the capillary stems and leaves, the shorter, ovoid spikelet, the smaller, reflexed, puncticulate utricles, and the caducous glumes. Except for the much longer, linear spikelets (¢ part much longer than the 2 one), and the suberect utricles of C. rara, these characters are unreliable. The two might possibly better be regarded as subspecies of a single species. In typical C. capillacea (var. capillacea) the stems and leaves are filiform and the utricles 2—2!/, mm long. The above description comprises also stouter plants with leaves up to 2 mm wide and utricles 3-4 mm long, which have been distin- guished as: var. sachalinensis (F. ScHmMrpT) OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 442; AxryaMa, Car. Far East. Reg. Asia (1955) 42, t. 6, f. 1 E-H; YOSHIKAWA, Ic. Jap. Carex 3 (1960) 386, t. 193. — C. nana Bootrt in A. Gray, Mem. Am. Ac. n.s. 6 (1859) 418; Ill. 4 (1867) 139, t. 44, f. 2, non CHAM. ex STEUD. 1855. — C. uda MAxio. var. sachalinensis F. SCHMIDT, Reisen Amur]. (1868) 191. — C. capil- lacea var. nana FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 8 (1896) 198. — C. rara ssp. capil- lacea var. nana KUx. Pfl. R. Heft 38 (1909) 103. — C. capillacea var. major NELMES, Kew Bull. (1949) 381; Reinwardtia 1 (1951) 406. Known from Sachalin, Japan, Korea, and Malesia. The Malesian specimens are often difficult to refer to one of the varieties. 15. Section Rhizopodae Ouw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 443; NELMEs, Reinwardtia 1 (1951) 411; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 17. — Sect. Extensae FRIES subsect. Rhizopodae (OHW1) KoyaMa, J. Fac. Sc. Un. Tokyo III, 8 (1962) 202. Type species: Carex rhizopoda MAXIM. 41. Carex eremostachya S. T. BLAKE, J. Arn. Arb. 28 (1947) 99, f. 1A; Ne~mMes, Kew Bull. (1949) 381; Reinwardtia 1 (1951) 412. — Fig. 121. Rhizome creeping, slender, 1-2 mm_ thick, covered with brown, striate scales. Stems approxi- mate or loosely tufted, erect or suberect, slender and rather flaccid, triquetrous, scabrid at the top, 20-45 cm by !/,-1 mm. Leaves subbasal, as long as to longer than the stems, flat, shortly attenuate, glabrous, scabrid on margins and nerves towards the apex, 2-3 mm wide, the lower ones reduced to brown, entire, bladeless or short-bladed sheaths. Spikelet 1, terminal, erect, androgynous, oblong- ellipsoid to shortly cylindric, densely many- flowered, 10-15 by 5-6 mm, the ¢ part shorter than the 2 one. Glumes ovate or ovate-lanceolate, acute or the upper obtusish, thin, glabrous, reddish brown with 3-nerved, pale, central stripe, 3-4 mm long, lower mucronate (mucro up to 1 mm), upper muticous. Urricles ellipsoid, triquetrous, angled dorsally, obliquely erect, membranous, glabrous and smooth, distinctly 4-5-nerved on each face, pale green, 41/,-5 by 11/,-1'/, mm, shortly stipitate, rather abruptly narrowed into a slender, linear, bidentate, c. 11/, mm long beak. Nut obo- void, triquetrous, apiculate, c. 2 by 11/; mm. Style- base scarcely thickened. Stigmas 3. Distr. Malesia: W. New Guinea, 9 km NE of Lake Habbema; only known from the type collection. Ecol. Massed on open beaches of small stream in forest, 2800 m. Note. Closely allied to C. rhizopoda Maxim. from Japan, which is however clearly distinct by its more flaccid, usually broader leaves, and its longer, oblong-cylindric spikelet with pale green glumes much shorter than the utricles. 16. Section Anomalae [CaREY in Gray, Man. Bot. N. Un. St. (1848) 557, pro subsect.]; MACKENZ. N. Am. FI. 18 (1935) 339. — Japonicae FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 10 (1898) 107, nomen. — Sect. Tumidae Kix. Pfl. R. Heft 38 (1909) 611; AKIYAMA, J. Fac. Sc. Hokkaido Imp. Un. V, 2 (1932) 188; NELMES, Reinwardtia | (1951) 394; non MeEtnsu. 1901. — Sect. Dispalatae OHW1, Mem. Coll. Sc. Kyoto 160 FLORA MALESIANA [ser. I, vol. 91 Imp. Un. B11 (1936) 480; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 54, 82. — Sect. Confertiflorae FRANCH. ex OHWI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 474; AKIyAMA, Car. Far East. Reg. Asia (1955) 144. — Sect. Molliculae OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 450; Koyama, Bot. Mag. Tokyo 72 (1959) 307; Quart. J. Taiwan Mus. 13 (1960) 226. — Sect. Extensae FRIES ser. Tumidae (KUK.) Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 202. Type species: Carex scabrata SCHWEIN. (lectotype). 42. Carex alopecuroides D. Don, Trans. Linn. Soc. 14 (1825) 332; Prod. Fl. Nepal. (1825) 43; Boorrt, Ill. 2 (1860) 88, t. 258; Boeck. Linnaea 41 (1877) 2s In Malesia only: var. chlorostachys (D. Don) CLARKE, J. Linn. Soc. 36 (1903) 271 (‘chlorostachya’); KoyAMA, Nat. Canad. 82 (1955) 199; Contr. Inst. Bot. Un. Montréal n. 57 (1957) 14; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 85; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490. — C. chlorostachys D. Don, Trans. Linn. Soc. 14 (1825) 330; Prod. Fl. Nepal. (1825) 43; non STEVEN, 1813. — C. doniana SPRENG. Syst. 3 (1826) 825; NEEs in Wight, Contr. (1834) 128; Dreyer, Symb. Caric. (1844) 26, t. 13; S. T. BLake, J. Arn. Arb. 28 (1947) 115; Kew Bull. (1950) 204, incl. var. cacuminis NELMES; NELMES, Reinwardtia 1 (1951) 399; ibid. 2 (1954) 381; AxryAMA, Car. Far East. Reg. Asia (1955) t. 161. — C. japonica (non THUNB.) Boott, Ill. 2 (1860) 88, t. 257, excl. var. minor BootT; Boeck. Linnaea 41 (1877) 283, excl. syn. SCHKUHR t. Ww f. 110; CLarKe, FI. Br. Ind. 6 (1894) 736; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 70. — C. japonica var. chlorostachys KtK. ex Matsuo. Ind. Pl. Jap. 2 (1905) 116; Pfl. R. Heft 38 (1909) 620; Merr. En. Philip. 1 (1923) 139; KoyaAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 212. — C. subtransversa CLARKE, Philip. J. Sc. 2 (1907) Bot. 108; Kew Bull. add. ser. 8 (1908) 92; Kix. Pfl. R. Heft 38 (1909) 614, in nota; Philip. J. Sc. 6 (1911) Bot. 63; Merr. En. Philip. 1 (1923) 142; S. T. BLAKE, J. Arn. Arb. 28 (1947) 115; NELMEs, Reinwardtia 1 (1951) 401. — C. japonica var. mesogyna KUK. Bot. Jahrb. 69 (1938) 265. — C. japonica ssp. subtransversa (CLARKE) KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 213. — Fig. 121. Rhizome emitting long, slender stolons covered with pale brown sheaths. Stems tufted, slender, erect, very acutely triquetrous to narrowly winged, smooth or scabrid on the angles above, (2-)30- 90 cm by 1-2 mm, surrounded below the leaves by a few pale, bladeless sheaths. Leaves subbasal and 1 higher on the stem, shorter to much longer than the stem, flat, stiffish, long-acuminate, scabrid on the margins and often asperous on the upper surface, 2-10 mm wide. Spikelets (2-)3-6, erect, cylindric, (1'/,-)2-5 cm long, approximate and fastigiate, or lower 1-2 somewhat distant, sessile, or lower shortly peduncled; terminal spikelet ¢ (not rarely gynaecandrous), pale, 1-3 mm thick, lateral ones 2, greenish or stramineous, 4-6 mm thick. Lower bracts foliaceous, overtopping the inflorescence, not sheathing, upper much reduced. Glumes oblong-ovate, acute to obtuse, 2—2!/, mm long, whitish with 3-nerved, green, central stripe excurrent in a scabrid, (?/,-)1-17/, mm long awn. Utricles ellipsoid, obscurely trigonous, subinflated, membranous, plurinerved, glabrous, ultimately patent, straight, greenish to stramineous, 3-4 by 1-11/, mm, rather gradually narrowed into a 1-13/, mm long, linear-conic, bidenticulate, smooth or almost smooth beak. Nut ellipsoid, obovoid, or oblong-obovoid, triquetrous, 11/,-2 by 3/,-1 mm. Style straight or slightly bent at the scarcely thickened base. Stigmas 3, about half as long as the utricle. Distr. Nepal, Sikkim, and Khasia to Central and S. China, Korea, Japan, Ryu Kyu Is., Formosa, Tonkin and Annam, and Malesia: Sumatra (Atjeh; W. Coast: Mt Kerintji), Java (Mt Wajang in W., Mt Tengger: Ranu Regulo in E.), Central Celebes (Mt Kambuno), Philippines (Luzon), and New Guinea. Ecol. Margins of lakes, riversides, openings in the mossy forest, also on dry, open slopes, stated to be a common trackside sedge in New Guinea; 1500-2900 m. Vern. New Guinea: akele, Lalibu. Notes. The only collection known from Celebes (EYMA 1395) was distinguished as C. doniana var. cacuminis NELMES, which has no taxonomic value; the dwarfish habit has apparently been caused by continuous grazing of anoas, the small wild buffalo of Celebes, Bos (Bubalus) depressicornis. Both C. alopecuroides and its var. chlorostachys are treated as varieties of C. japonica THUNB. by KUKENTHAL, and also recently by KoyAmMA, but C. japonica seems to be distinct by its short- cylindric, not crowded, peduncled, usually pendu- lous spikelets, not distinctly awned glumes, and its very long stigmas. In typical C. alopecuroides the leaves are usually narrower than in var. chloro- stachys, the spikelets very slender, and the utricles somewhat narrower, less inflated, but the two are very near to each other and connected by inter- mediates. C. subtransversa CLARKE was described from N. Luzon, where also C. alopecuroides var. chlorostachys occurs. I fail to see differences 1979] Cy PERACEAE—II (Kern & Nooteboom) 161 Fig. 130. Carex capillacea Boott. Habit, x '/; (from Mt Papandajan, VAN STEENIS). between the two but for the narrower leaves of the former, which is said to differ also by the suberect— not patent — utricles; the specimens are not fully ripe and the degree of reflexion of the utricles depends upon maturity. KUKENTHAL placed C. subtransversa under C. brownii and its var. trans- versa (BooTtr) KUK., with which it is certainly not closely related. 43. Carex brownii TUCKERM. En. Meth. Car. (1843) 21 (or 15? ‘brownii’?); Boott, Ill. 4 (1867) 161, t. 532 (pl. dextrae); Boeck. Linnaea 41 (1877) 151, incl. var. viridis BOECK.; BENTH. Fl. Austr. 7 (1878) 447; KK. Pfil. R. Heft 38 (1909) 612, f. 104 A-D; Ouwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 477; S. T. Biake, J. Arn. Arb. 28 (1947) 115; NeELMES, Kew Bull. (1949) 384; Reinwardtia 1 (1951) 398; Koyama, Act. Phytotax. Geobot. 18 (1959) 23; J. Fac. Sc. Un. Tokyo III, 8 (1962) 215; Phytologia 17 (1968) 409, t. 16. — C. striata R. BR. Prod. (1810) 243; KunTH, En. 2 (1837) 458; STEUD. Syn. 2 (1855) 226; DreJeR, Symb. Caric. (1844) 28, t. 15; non Micwx, 1803. — C. nippo- sinica OnwI, Act. Phytotax. Geobot. 11 (1942) 255; AKIYAMA, Car. Far East. Reg. Asia (1955) 144, t. 133; YosHIKAWa, Ic. Jap. Carex 2 (1958) 256, t. 128. a. ssp. brownii. — Fig. 122. Rhizome short, without stolons. Stems slender, erect, triquetrous, smooth or almost so, 25-75 cm by 1-1!/, mm. Leaves basal and often 1 higher up the stem, flat, weak, shorter than the stems, long- acuminate, scabrous on nerves and margins, 3-4 mm wide, lowest ones reduced to bladeless, reddish-brown sheaths. Spikelets 3-4(—5), erect, short-cylindric to cylindric, upper approximate, fastigiate, lowest often more or less distant; terminal spikelet g¢, or rarely gynaecandrous, subsessile, pale, (?/,-)1-2!/, cm by 2(-3) mm; lateral spikelets 2, subsessile or lowest sometimes on a long, exserted peduncle, short-cylindric, obtuse, densely flowered, 1'/,-2'/,(-3) cm by 5-6 mm. Lowest bract foliaceous, overtopping the inflorescence, more or less sheathing, upper much shorter, not or hardly exceeding the inflorescence. Glumes ovate or oblong-ovate, obtuse, scarious, 1-2 mm long, whitish with 3-nerved green central stripe excurrent in a scabrous, '/,-3'/, mm long awn. Urtricles broadly ellipsoid to subglobose, obscurely trigonous, inflated, membranous, strongly many-ribbed, glabrous, straight, patent, dark olive-green to dark brown, contrasted against the white glumes, 3-4 by 1'/;-1°/, mm, suddenly contracted into a very short, bidenticulate beak. Nut obovoid, triquetrous, yellowish, 2/,-2'/, by 1'/;-1?/, mm, with a very short, (sometimes indistinctly) bent stipe dnd beak. Style-base slightly thickened. Stigmas 3. Distr. Japan, Korea, Formosa, E. China (Yangtze R. valley), Australia (New South Wales, Victoria), New Zealand; in Malesia: New Guinea (W. New Guinea: Balim R. valley; Terr. of New Guinea: W. Highlands, lower Tale valley and 162 FLORA MALESIANA [ser. I, vol. 9! 3 WISSMA 778 f Fig. 131. Inflorescence (nat. size), spikelets and (occasionally) venation (x 2). — 48. Carex michauxiana BOoECK., a. — 18b. C. perakensis CLARKE var. borneensis (CLARKE) Noot., b. — 18a. var. perakensis, c—d. — 19a. C. turrita CLARKE var. turrita, e-f. — 50. C. maubertiana Bootr, g. — 64. C. echinata Murr., h. — 15. C. baccans Nees, i. — 51. C. graeffeana BoECK., j (a SCHODDE 1992, b BROOKE 8561, c-d VAN STEENIS 8277, e-f ANU 15532, g KosTERMANS 14005, A DE WILDE c.s. 13323, i ELBERT 62, j SINCLAIR 979’). 1979] es * aes EE a b c d Cy PERACEAE—II (Kern & Nooteboom) " pete € Fig. 132. Glumes, x 10. — 46. Carex brachyathera OHWI, a. — 49. C. pseudocyperus L. var. fascicularis (SOLAND. ex Boott) Boott, b. — 52. C. phacota SPRENG., c. — 53. C. pruinosa Boott, d. — 54. C. teres Bootrt, e. — 56. C. bilateralis HAyATA, f. — 57. C. brunnea THUNB., g (a BRAss 9803, b EyMa 4709, c JERMY 4634, d VAN STEENIS 4624, e DE WILDE c.s. 13329, f BRAss 9515, g EYMA 3862). Sirunki; E. Highlands, Aiyura, and Kainantu, Morobe Distr., Langiman R.). Distr. map in Acta Phytotax. Geobot. 18 (1959) 24. Ecol. In wet places, pools, drains, etc., also open grassy area; 1500-2500 m. Vern. Kariandend, Enga lang. b. ssp. transversa (BooTT) KERN, stat. nov. — C. transversa Boott, Perry Exp. 2 (1857) 324; Ill. 4 (1867) 202; Franco. & Sav. En. Pl. Jap. 2 (1879) 149, incl. var. dissociata FRANCH. & SAV.; FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 10 (1898) 48, t. 3, f. 1; CLARKE, J. Linn. Soc. Bot. 36 (1904) 314; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 478; Akiyama, Car. Far East. Reg. Asia (1955) 145, t. 134; YosuikAwa, Ic. Jap. Carex 2 (1958) 258, t. 129. — C: brownii var. transversa KUK. ex Matsum. Index Pl. Jap. 2 (1905) 103; Kux. Pfl. R. Heft 38 (1909) 614. — C. furusei KOYAMA, J. Jap. Bot. 30 (1955) 135, pro C. brownii x transversa. — C. brownii var. dissociata KoyaAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 215. —.Fig. 122. Utricles ovoid-ellipsoid, up to 6 mm long, gradually narrowed into a c. 2 mm long beak. Distr. Japan, Ryu Kyu Is., Korea, China (Yangtze valley region); in Malesia: New Guinea (W. Highlands, Sirunki). Ecol. In New Guinea near water-course on clay soil, at c. 2500 m. Note. The long-beaked utricles are so different from those of C. brownii s.s., that I prefer to treat this taxon as a subspecies, not as a variety of C. brownii. 44. Carex oedorrhampha NeELMes, Kew Bull. (1939) 659; ibid. (1949) 384; Reinwardtia 1 (1951) 396; ibid. 2 (1954) 381; RaymM. Mém. Jard. Bot. Montréal n. 53 (1959) 83; Dansk Bot. Ark. 23 (1965) 261; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490. — C. tumida Bootr, Ill. 1 (1858) 66, t. 181; Boeck. Linnaea 41 (1877) 243; CLARKE, FI. Br. Ind. 6 (1894) 741; J. Linn. Soc. Bot. 37 (1904) 16; KUx. Pfl. R. Heft 38 (1909) 615; Bull. Jard. Bot. Btzg III, 16 (1940) 321, non BEILSCHM. 1850. — C: olivacea (non Boott) KUK. Bot. Jahrb. 70 (Jan. 1940) 467; Bull. Jard. Bot. Btzg III, 16 (Feb. 1940) 321, incl. var. altissima KUK.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 70, p.p. — C. oedorrhampha var. arfakiana OHWI, Bot. Mag. Tokyo 56 (1942) 214. — C. oedorrhampha var. microcarya NELMES, Kew Bull. (1950) 204; Reinwardtia 1 (1951) 398. — Fig 122. Rhizome short, in large tussocks, without stolons. Stems tufted, slender, erect, triquetrous, smooth, 50-125 cm by 2-3 mm below. Leaves subbasal and 1-2 higher up the stem, about as long as the stem, flat, stiffish, long-acuminate, scabrous on margins and main nerves, 4-8 mm wide, few lowest reduced to reddish purple sheaths readily fraying into thin strips; ligula elongate, up to 1'/, cm long. Spikelets 4-6, erect or subcernuous, upper approximate and fastigiate, lower distant; terminal spikelet 3, long-linear, subsessile, not or slightly exceeding the uppermost lateral spikelet, pale, 2-6 cm long, 1-2 mm thick, lateral spikelets 2, narrowly cylindric, densely flowered, 2'/,-12 cm long, 4-6 mm thick, upper on shortly, lower on long-exserted, scabrid peduncles. Lower bracts 164 FLORA MALESIANA [ser. I, vol. 9! foliaceous, much exceeding the inflorescence, long- sheathing, upper more or less reduced. Glumes oblong to ovate-lanceolate, obtuse, translucent, glabrous or sparsely hispidulous, 13/,-2 mm long, whitish with 3-nerved, green, central stripe excur- rent in a scabrid awn 1-2 mm long. Utricles ellipsoid, membranous, somewhat inflated, pluri- nerved, glabrous, obliquely erect to subpatent, olive-brown to fuscous, 3—33/, by 1-1'/, mm, rather gradually narrowed into a 1-1!/, mm long, conic-linear, smooth, minutely notched beak which is often somewhat swollen at or below the middle. Nut ellipsoid, triquetrous, yellow to brownish, shortly stipitate, distinctly beaked, (11/,—-)2-2!/, by c. 1 mm. Style straight, thickened at the base. Stigmas 3. Distr. E. Himalaya, S. China (Yunnan), N. Thailand, Tonkin, Assam; in Malesia: Sumatra (Atjeh; W. Coast Res.: Mt Kerintji), W. Java (Mt Papandajan), Moluccas (Buru), New Guinea. Ecol. Wet places in forests, lake margins, swampy grasslands, 1200-2400 m. Notes. Boott’s figure, showing a distinct swelling in the middle of the beak (hence his specific epithet) is, according to CLARKE, taken from a not fully ripe collection. In mature utricles the lower half of the beak is cylindric, somewhat swollen, the upper conic. In the specimens from Mt Kerintji the beak is scarcely swollen; they were distinguished as var. microcarya NELMES. Besides to C. olivacea the species is closely related to C. ischnostachya StEuD. from Japan, which is easily distinguishable by the muticous glumes, but otherwise very similar. 45. Carex olivacea Boott, Proc. Linn. Soc. 1 (1846) 286; Ill. 1 (1858) 56, t. 149; CLARKE, FI. Br. Ind. 6 (1894) 741; J. Linn. Soc. Bot. 37 (1904) 15; KUK. Pfl. R. Heft 38 (1909) 617; BAck. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 70, p.p.; NELMES, Reinwardtia 1 (1951) 395; Koyama, Bot. Mag. Tokyo 72 (1959) 304; J. Fac. Sc. Un. Tokyo III, 8 (1962) 215; KERN in Back. & Bakh. f. FI. Java 3 (1968) 490. ssp. Olivacea. — Fig. 122, 133. Rhizome stout, woody, emitting long, stout stolons. Stems stout, erect, triquetrous, smooth or scabrid on the angles above, 50-110 cm by up to 5 mm below. Leaves basal and subbasal, much exceeding the stems, flat, stiffish, long-acuminate, scabrous on the margins towards the top, the broader ones 1-2 cm wide; sheaths stramineous, somewhat spongy, deeply concave in front, ligule elongate, up to 5 cm long. Spikelets 4-9 (according to Boott up to 14), erect, long-cylindric, upper approximate, fastigiate, lower somewhat distant; terminal spikelet ¢ (often with some @ flowers at the base), peduncled, 51/,-16 cm long and 5 mm thick, often a second, shorter 3 spikelet added at Fig. 133. Carex olivacea Bootr in the Kerintji area, Westcoast of Sumatra, Rawal Bento, 1400 m altitude (MEWER 6656, Aug. 1956). 1979 the base, remaining spikelets 2 (but usually with short ¢ apices), very densely flowered, sessile but lowest sometimes distinctly peduncled, up to 16 cm by 5-8 mm. Lower bracts foliaceous, much over- topping the inflorescence, semi-amplexicaul by dark auricles, upper shorter, usually none sheathing (lowest sometimes more or less sheathing). Glumes oblong, obtuse, vinaceous to dark red with 3-nerved, green, 1'/,-3 mm long, central stripe excurrent in a scabrid awn ?/,—-3 mm long. Utricles obovoid or ellipsoid, membranous, much inflated when ripe, rugose when dry, slenderly plurinerved, patent or reflexed, glabrous, olive-brown, 3'/,— 4?/, by 1'/,-2 mm, suddenly narrowed into a short, conic, often recurved, minutely bidenticulate beak ciliate at the mouth. Nut obovoid or oblong- obovoid, triquetrous, beaked, stramineous to yellowish, c. 2 by 1 mm. Style straight, not or hardly thickened at the base. Stigmas 3. Distr. NE. India; in Malesia: W. Sumatra (Mt Kerintji), W. Java (Telaga Bodas, once collected by H. O. Forses in 1880). The record for Cy PERACEAE—II (Kern & Nooteboom) Indo-China (Reinwardtia 1, 1951, 396) refers to a collection of C. nemostachya Steup. Distr. map in Bot. Mag. Tokyo 72 (1959) 304, f. 26 (the occur- rence in Java erroneously indicated as comprising the whole western and central part of that island, the Sumatran locality unknown at the time). Ecol. On Mt Kerintji in swamp at 1400 m altitude, at Telaga (= lake) Bodas ‘“‘in warm water, 1500 m.” Notes. Closely related to 44. C. oedorrhampha NELMES, but much stouter, and distinguished from all other members of its section by its very wide leaves and very long spikelets. Ssp. confertiflora (BooTT) KoyAMA, Bot. Mag. Tokyo 72 (1959) 307; Phytologia 17 (1968) 413, t. 17 (C. confertiflora Boott in A. Gray, Bot. Jap. 1859, 418; Ill. 4, 1867, 184; C. olivacea var. minor KUkK. Pfi. R. Heft 38, 1909, 618), from Japan, and ssp. recurvisaccus (KOYAMA) KoyAMaA, /.c. (C. recur- visaccus KOYAMA, J. Jap. Bot. 15, 1956, 166, f. 2), from China, Kwantung, differ but slightly from the typical subspecies. 17. Section Ferrugineae [TUCKERM. En. Meth. (1843) 12, nomen]; ex BAILEY, Proc. Am. Ac. 22 (1886) 92, as group; NELMES, Reinwardtia | (1951) 409. — Sect. Frigidae Frits subsect. Ferru- gineae (BAILEY) KUK. Pfl. R. Heft 38 (1909) 559. Type species: Carex ferruginea Scop. 46. Carex brachyathera Ouwi, Jap. J. Bot. 7 (1934) 190; Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 332, f. 4, t. 8 f. 5; AKIYAMA, Car. Far East. Reg. Asia (1955) 133, t. 118; KoyAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 158, incl. var. brevispiculosa KoyAMa. — C. breviculmis var. perciliata (non KUK.) RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 247. — C. tricuspidata KK. Bot. Jahrb. 70 (1940) 466, incl. var. brevispiculosa KUK. et var. minor KUK.; S. T. BLake, J. Arn. Arb. 28 (1947) 113; NELMES, Kew Bull. (1949) 384; Reinwardtia 1 (1951) 410. — Fig. 122, 132a. Rhizome thick, woody, obliquely descending, sometimes elongated. Stems erect, somewhat cernuous at the top, triquetrous, smooth or slightly scaberulous just below the inflorescence, (5—)20- 70 cm by 1-1'/, mm, at the base covered with a thick mass of fibrous, brownish remains of leaf- sheaths. Leaves basal or 1-2 higher on the stem, shorter than the stem, flat, or involute when dry, stiffish, asperous, long-attenuate, 1—3!/, mm wide. Spikelets 3—5, single at the nodes, erect to cernuous, often subfastigiate, rather densely flowered above, lax-flowered below, 1'/,-6 cm long, terminal one 36, subclavate, 1-2 mm thick, long-peduncled, lateral ones 9, linear-cylindric, 2!/,—-3'/, mm thick, on exserted, slender, smooth or antrorsely scabrid peduncles. Lower bracts foliaceous, shorter to slightly longer than their spikelet, long-sheathing, upper reduced. Glumes oblong, often asymmetri- cal, truncate or emarginate, thin, brown, whitish hyaline at the top, 3-4'/, mm long, with strong midrib excurrent in a hispidulous, up to 1 mm long awn. Utricles ellipsoid or ellipsoid-fusiform, obtusely compressed-trigonous, angled ventrally, membranous, nerveless except for 2 submarginal nerves, sparsely to rather densely subappressed- hispid, suberect, stipitate, 3-4'/, by c. 1 mm, rather gradually narrowed into a stout, straight or slightly bent, bidentate, 1-1!/, mm long beak with oblique, white-hyaline mouth. Nut ellipsoid, triquetrous, c. 2 by 1 mm. Style-base pyramidally thickened. Stigmas 3 (according to NELMES 2 or 3). Distr. Ryu Kyu Is., Formosa; in Malesia: New Guinea (Mt Carstensz, Mt Wilhelmina in W., Mt Sarawaket in E.). Ecol. In mountain grasslands, on steep rocks, seepages, sandy margins of streams, 3000-4000 m. Notes. The New Guinea collections exhibit a high degree of variability in size, length of bracts, spikelets, and utricles, probably due to their growing at high altitude, often in uncongenial habitats. I do not see any reason to treat them as varietally distinct from the Formosan plants, as was 166 FLORA MALESIANA [ser. I, vale 9* done by KoyAMa, on account of the longer bracts and — according to NELMES — sometimes digynous flowers. To judge from its description and excellent figure, C. drepanorhyncha FRANCH. Pl. David. 2 (1888) 141; Nouv. Arch. Mus. Hist. Nat. Paris III, 9 (1897) 178, t. 4, f. 1; KUxK. Pfl. R. Heft 38 (1909) 563, from Szechuan, is very near to C. brachyathera. 18. Section Sylvaticae [TUCKERM. En. Meth. (1843) 12, nomen]; BooTT ex MACKENZ. N. Am. FI. 18 (1935) 283; NELMES, Reinwardtia 1 (1951) 402. — Sect. Hymenochlaenae DREJER subsect. Debiles (CAREY) KUK. Pfl. R. Heft 38 (1909) 594, p.p. Type species: Carex sylvatica HUDs. 47. Carex finitima Bootr, Ill. 1 (1858) 44, t. 112; Boeck. Linnaea 41 (1877) 247; CLARKE, FI. Br. Ind. 6 (1894) 736; KUx. Pfl. R. Heft 38 (1909) 598, f. 101 E-H; Netmes, Kew Bull. (1949) 385, 391; ibid. (1950) 204; Reinwardtia 1 (1951) 403; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 218. — C. remotiflora HAYATA, Ic. Pl. Form. 10 (1921) 68, f. 45; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 446; AktyAMaA, Car. Far East. Reg. Asia (1955) 155, t. 148, f. 2. — C. fusiformis NEES var. enervosa KUK. Bot. Jahrb. 70 (1940) 467. — C. atjehensis Kx. Bull. Jard. Bot. Btzg III, 16 (1940) 314; in Fedde, Rep. 53 (1944) 105 (‘atjehien- sis’). — Fig. 122. Rhizome short. Stems tufted, erect, triquetrous, smooth, (10-)30-90 cm by 1-2 mm below, sur- rounded below the leaves by a few reddish bladeless sheaths. Leaves basal and 1-2 higher up the stem, shorter than to slightly exceeding the stem, flat, smooth except for the minutely scaberulous mar- gins, long-attenuate, 2-8 mm wide. Spikelets 4-8(-12 according to Boott), linear-cylindric, terminal one ¢ or rarely gynaecandrous, peduncled, 1'/,-41/, cm by c. 1 mm (sometimes a smaller second ¢ spikelet added), remaining ones 9, lax- flowered, erect or lower cernuous, upper approxi- mate, lower remote, 2-9 cm by 3-7 mm; peduncles very slender, smooth or sparsely scaberulous above, lower long-exserted. Lower bracts foliaceous, slightly shorter than to much exceeding the in- florescence, long-sheathing, upper much reduced; sheaths pale to reddish brown. Glumes oblong- ovate, acute to very obtuse, 31/,-4(-6) mm long, translucent, with broad, white margins and greenish midrib, the latter not reaching the apex but sometimes excurrent below the apex in a short, up to 1(-2) mm long awn. Utricles fusiform, tri- gonous, membranous, with 2 marginal nerves, otherwise nerveless or obscurely few-nerved, glabrous, smooth, obliquely erect, shining, light green, 5—7!/, by 1-2 mm, rather gradually narrowed into a linear-cylindric, long, smooth beak with oblique, scarious mouth. Nut ellipsoid or oblong- ellipsoid, triquetrous, finally dark brown, shortly stipitate, abruptly beaked, 2'/,-3 by 1-1'/, mm. Style thickened at the base. Stigmas 3, long, slender. Distr. E. Himalaya, S. China (Szechuan, Yun- nan), Formosa; in Malesia: N. Sumatra (Atjeh: Gajolands), E. New Guinea. Ecol. Grassy and bushy slopes, alpine grass- lands, 2400-3900 m. 19. Section Folliculatae MACKENz. in Britt. & Brown, Ill. Fl. ed. 2, 1 (1913) 353; N. Am. FI. 18 (1935) 426. — Sect. Orthocerates Kocu subsect. Folliculatae (MACKENZ.) KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 234. Type species: Carex folliculata L. 48. Carex michauxiana Boeck. Linnaea 41 (1877) 336; CLARKE, J. Linn. Soc. Bot. 36 (1904) 298; KUK. Pfil. R. Heft 38 (1909) 705; MACKENZz. N. Am. Fl. 18 (1935) 427; N. Am. Car. 2 (1940) t. 489; FERN. in Gray’s Man. Bot. ed. 8 (1950) 375, f. 742; GLEASON, New Britt. & Brown Ill. Fl. 1 (1952) 358 f. — C. rostrata Micux, Fl. Bor.-Am. 2 (1803) 173; Boortrt, Ill. 2 (1860) 91, t. 267; non STOKES, 1787. — C. abacta L. H. BAILEY, Bull. Torr. Bot. Club 20 (1893) 427. The Asiatic plants are distinguished as: var. asiatica (HULTEN) OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 491; AkryAMa, Car. Far East. Reg. Asia (1955) 163, t. 158 f. 2; YOsHI- KAWA, Ic. Jap. Carex 2 (1958) 268, t. 134; KOYAMA, 1979] CyYPERACEAE—II (Kern & Nooteboom) 167 J. Fac. Sc. Un. Tokyo III, 8 (1962) 235; Kern, Blumea 13 (1965) 125. — C. michauxiana ssp. asiatica HULTEN, Kung]. Svenska Vet. Ak. Hand. 5 (1927) 207, f. 14, t. 223; Phytologia 17 (1968) 417, t. 19. — C. michauxiana f. asiatica (HULTEN) AKIYAMA, J. Fac. Sc. Hokkaido Imp. Un. 5 (1932) 220, f. 162. — C. dolichocarpa C. A. MEY. ex Krecz. in Komar. Fl. U.R.S.S. 3 (1935) 458, 623. — Fig. 122, 131a. Rhizome short, thick. Stems densely tufted, stiffly erect, slender, trigonous, smooth, at the base clothed with the remains of old leaf-sheaths, 20-60 cm by 1-2 mm. Leaves subbasal, shorter to longer than the stems, firm, flat, long-attenuate, scabrid towards the apex, light green, 2-5 mm wide; sheaths tight, white-hyaline ventrally, ligule as long as wide. Inflorescence consisting of 1 terminal ¢ spikelet and (1—)2-4 9 spikelets. Bracts long-sheathing, leaf-like, the lower overtopping the inflorescence; sheaths concave at the mouth. 3 Spikelet (see note) shortly peduncled or subsessile, few-flowered, 8-20 by 1'/,-2!/, mm; glumes oblong-ovate, acute, yellowish brown with 3-nerved green centre and hyaline margins; upper @ spikelets subsessile, crowded and sometimes hiding the g one (see note), the lower 1—2 remote, on slender, more or less exserted, erect, smooth peduncles, sub- globose or broadly ovoid, 5-—20-flowered, 15- 25 mm long and wide. G/umes ovate, acutish, hyaline with conspicuously 4—5-nerved centre, 1/,1/, as long as the utricle. Ufricles at first appressed-ascending, soon divergent, lanceolate- subulate, obtusely trigonous, subcoriaceous, glab- rous, many-nerved, stipitate, spongy at the base, shining yellowish green, (8—)12—13 by 1'/,-2 mm, gradually tapering into the scabrid bidentate beak; teeth erect, ?/; mm. Nut trigonous with convex sides, oblong-ovoid, shortly stipitate, shining yellow, c. 3 by 1'/, mm. Sty/e continuous with the nut, tortuous, subincrassate at the base. Stigmas 3. Distr. The typical variety in northeastern N. America, the var. asiatica in eastern Asia: S. Kam- chatka, Kuriles, Yezo, Hondo, China (Shanghai); in Malesia: E. New Guinea. Ecol. In swamps, bog grasslands, 2250-2650 m. Vern. Tudik, Mendi lang., koale, Enga lang. Notes. The differences between the American plants and the Asiatic ones are but slight, and it is questionable whether recognition of two geographi- cal races is justified. In the few American specimens I could examine the 3 spikelet does not overtop the upper 2 ones (but this is not rarely also the case in Asiatic specimens!), the leaves are slightly narrower (2-3 mm wide), the 9 glumes 3-nerved, and the utricles in general somewhat smaller. I fail to see the difference in the relative length of the glumes mentioned by Koyama. If the detailed description by MACKENZIE, /.c., refers to American plants only, the distinction of two varieties seems unjustified, as the Asiatic plants almost completely fall within the limits of variation there given. It is worth noticing that most of the terminal spikelets in the Mt Giluwe specimens are not strictly g, but bear some @ flowers at the top. Sometimes the ¢ spikelet is either inconspicuous or absent. A northern element in the New Guinea moun- tain flora. 20. Section Pseudocypereae [TucKeRM. En. Meth. (1843) 13, nomen]; ex BAILEY, Proc. Am. Ac. 22 (1886) 69, as group; Kix. Pfl. R. Heft 38 (1909) 693. — Sect. Orthocerates KOCH subsect. Pseudocypereae (BAILEY) Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 234. Type species: Carex pseudocyperus L. 49. Carex pseudocyperus LINNE, Sp. Pl. 2 (1753) 978; Boott, Ill. 4 (1867) t. 451, 452; Boeck. Linnaea 41 (1877) 321; KUK. Pfl. R. Heft 38 (1909) 695. In Malesia only: var. fascicularis (SOLAND. ex Boott) Bootrt, Ill. 4 (1867) 41; Kix. Pfl. R. Heft 38 (1909) 696; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 234; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 490. — C. fascicularis SOLAND. ex Boott in Hook. f. FI. Nov. Zel. 1 (1853) 283; Boortrt, III. 1 (1858) 53, t. 139, 140; S. T. BLake, J. Arn. Arb. 28 (1947) 116; NeELMEs, Reinwardtia 1 (1951) 393. — C. pseudo- cyperus (non L.) R. Br. Prod. (1810) 243; BENTH. FI. Austr. 7 (1878) 448; CLARKE, J. Linn. Soc. Bot. 37 (1904) 16. — Fig. 122, 132b. Rhizome very short, woody. Stems stout, tufted, erect, triquetrous with flat or slightly concave sides, smooth except just below the inflorescence, surrounded below the leaves by a few bladeless, somewhat fibrous sheaths, 50-150 cm by 3-4 mm. Leaves subbasal, longer than the stems, flat, stiffish, rough-margined, conspicuously septate-nodulose, long-acuminate, light or yellowish green, 6-10 mm wide. Spikelets 3-7, usually close together (lowest 1-2 sometimes distant), the terminal 3, peduncled, 3-7 mm thick, the lateral 2, cylindrical, very densely flowered, 2!/,-5 cm long and 7-15 mm thick, the upper shortly peduncled, the lower on slender, FLORA MALESIANA [ser: I, vols 9! 168 smooth or scabrid peduncles at length pendulous. Lower bracts leaf-like, far exceeding the inflores- cence, scarcely sheathing (occasionally long- sheathing in distant spikelets), upper subfoliaceous to setaceous, not sheathing. Glumes oblong or oblong-spathulate, obtuse, translucent, eventually ferrugineous, with 3-nerved green centre, ciliate above, 2—2!/, mm, excurrent into a flat, antrorsely scabrous awn 2-4 mm long. Utricles ovoid or ovoid-lanceolate, obscurely trigonous, strongly and densely many-nerved, coriaceous, glabrous, when ripe widely spreading or even reflexed, somewhat inflated, distinctly stipitate (stipe 7/,-1 mm, in some Australian specimens up to 1!/, mm), greenish to stramineous, 4-5(—7) by 1—1!/, mm, tapering above into a 1!/,-2 mm long, smooth, deeply 2-cleft beak (teeth c. 1 mm, divergent). Nut obovoid or ellip- soid, triquetrous, scarcely stipitate, golden or brown, 1'/,-17/; by 1(-11/3) mm. Style continuous with the nut, contorted. Stigmas 3. Distr. Carex pseudocyperus in the circum- scription here accepted shows an almost world- wide distribution. The typical variety is widely distributed in the northern hemisphere (Eurasia from W. Europe to Kashmir and Japan, N. Africa, eastern N. America). Other varieties occur in North and South America; var. fascicularis in Australia (Tasmania to Queensland) and in New Zealand, in Malesia: New Guinea (W. New Guinea: Lake Habbema, Wissel Lake region; Terr. of New Guinea: W. Highlands, Kandep valley; Yobobos grassland area; Lake Inin). Specimens of, this variety in BM and K are labelled “Java, Horsfield”’ ; recent collections from this island are not extant. Ecol. In pools and swamps, along lakes, river- sides; 1750-3225 m. Vern. Kwai’a’reh, koali, Enga lang., Kepilam. Notes. Var. fascicularis differs from var. pseudocyperus in its darker, ferrugineous glumes, its subinflated utricles with more divergent teeth to their beaks and longer stipe, and its contorted style. The varieties certainly represent geographical races and might therefore better be considered subspecies. They are often treated at specific level, but the differences are so slight that it is here preferred to follow Bootr (1867) and KUKENTHAL (1909). 21. Section Occlusae CLARKE, Kew Bull. add. ser. 8 (1908) 147; Raym. Mém. Jard. Bot. Montréal n. 53 (1959) 55, 87; KoyaMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 169. — Sect. Scleriiculmes NELMES, Kew Bull. (1951) 121; Reinwardtia 1 (1951) 407;, Koyama, J. Jap. Bot. 29 (1954) 44; Act. Phytotax. Geobot. 16 (1955) 108. Type species: Carex maubertiana BooTt. 50. Carex maubertiana Boott, Ill. 1 (1858) 45, t. 114; CLarkeE, J. Linn. Soc. Bot. 36 (1904) 297; Camus, FI. Gén. I.-C. 7 (1922) 200, f. 29, 5-8; NELMES, Reinwardtia 1 (1951) 408; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 87; KoyAma, Act. Phytotax. Geobot. 16 (1955) 39; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 489. — C. hebecarpa C. A. MEY. var. lachnosperma CLARKE, FI. Br. Ind. 6 (1894) 747, quoad specim., non C. lachnosperma Nees. — C. hebecarpa var. maubertiana FRANCH. Nouv. Arch. Mus. Hist. Nat. Paris III, 10 (1898) 70; KOK. Pfl. R. Heft 38 (1909) 745. — C. hebe- carpa var. ligulata (non Kix.) Back. Bekn. FI. ae (em. ed.) 10 (1949) fam. 246, p. 69. — Fig. 122, 1g. Rhizome very short. Stems tufted, erect, tri- quetrous, smooth, for the greater part hidden by the leaf-sheaths, 40-60 cm by 2-3 mm. Leaves of normal length only in the upper half of the stem, exceeding the inflorescence, flattish to strongly revolute, stiff, greyish green, asperous above towards the long-attenuated apices, 3-7 mm wide, merging above into the foliaceous lower bracts and below into short-bladed to bladeless, purplish, rather tight sheaths; ligule obtuse, membranous, ferrugineous. Spikelets 4-9, erect or suberect, cylindric, dense-flowered, 1'/,-4 cm long, upper approximate, fastigiate, on included or shortly exserted peduncles, lower somewhat distant, on scarcely to long-exserted, scabrid peduncles; ter- minal spikelet g, 1-2 mm thick, sometimes a second, smaller ¢ spikelet added; remaining spikelets 9, 4-6 mm thick. Lower bracts foliaceous, exceeding the inflorescence, their sheaths hispidu- lous especially near the mouth, upper much reduced. Glumes ovate, obtuse to acutish, trans- parent, glabrous or hispidulous at the top, pale with hyaline margins and 3-nerved,- greenish central stripe, covered with resinous flecks and streaks, muticous or mucronulate, 1'/,-2!/, mm long. Urtricles ellipsoid, trigonous, submem- branous, obliquely patent, with 2 marginal nerves displaced on to the dorsal face, densely white- hispid, broadly stipitate, 3-33/, by 1'/,-13/, mm, abruptly narrowed into a c. 1 mm long, bidentate, finally somewhat upcurved beak. Nut ellipsoid, triquetrous, c. 2 mm long. Style-base thickened. Stigmas 3. 1979] Cy PERACEAE—II (Kern & Nooteboom) 169 Distr. Himalaya, Annam, China (Hupeh); in Malesia: Sumatra (Atjeh; W. Coast Res.: Mt Kerintji), W. and E. Java (Priangan; Besuki), E. Borneo (Berouw: Mt Ilas Mapulu), New Guinea (Morobe Distr., near Bulolo). Distr. map in Act. Phytotax. Geobot. 16 (1955) 38. Ecol. Primary forests, forest-borders, bushy slopes, 400-2100 m. 22. Section Longispicae CLARKE, J. Linn. Soc. Bot. 37 (1904) 3; NELMEs, Reinwardtia | (1951) 417. Type species: Carex graeffeana BOECK. 51. Carex graeffeana Boeck. Flora 58 (1875) 123; CLARKE, J. Linn. Soc. Bot. 37 (1904) 5; Philip. J. Sc. 2 (1907) Bot. 107; Kuk. Pfl. R. Heft 38 (1909) 403; Philip. J. Sc. 6 (1911) Bot. 62; in Hochr. Candollea 6 (1936) 433; Bot. Jahrb. 69 (1938) 264; Merk. Philip. J. Sc. 5 (1910) Bot. 335; En. Philip. 1 (1923) 138; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 62; NELMES, Kew Bull. (1938) 109, incl. var. samoensis NELMES; ibid. (1955) 317; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN. Mt. Fl. Java (1972) pl. 14-9. — C. rechingeri PALLA, Oest. Bot. Z. 57 (1904) 424. — C. philip- pinensis NELMES, Kew Bull. (1938) 109; ibid. (1949) 385, 392; Reinwardtia 1 (1951) 419; ibid. 2 (1954) 381. — C. exploratorum NELMES, Kew Bull. (1938) 108; Reinwardtia 1 (1951) 418. — C. pandanus Oxuwit, Bot. Mag. Tokyo 56 (1942) 214. — C. euph- lebia S. T. BLAKE, J. Arn. Arb. 28 (1947) 108, f. 3C. — Fig. 122, 131}. Rhizome short, woody, forming large clumps. Stems densely tufted, stoutish, erect, triquetrous, scabrid on the angles at the top, 30-110 cm by up to 4 mm below, surrounded below the leaves by shining brown to blackish red sheaths splitting into herring-bone-shaped fibres. Leaves basal and subbasal, often 1-2 higher up the stem, stiff, subcoriaceous, flat but margins often revolute, scabrid on the nerves, long-attenuate, greyish or glaucous-green, much overtopping the stems, 3-12 mm wide. Bracts not sheathing, 2-3 lower ones foliaceous, semi-amplexicaul and blackish red auricled at the base, middle ones subfoliaceous, upper ones reduced to long-awned glumes. Spike- lets (6—)10-15(—S0), single or binate at the nodes, upper approximate, fastigiate, subsessile, lower more distant, on capillary, scabrid, up to 5 cm long peduncles, more or less nodding, all androgynous or 1(—2) at the base of the uppermost much smaller and wholly g, exceptionally terminal spikelet wholly 3, linear-cylindric, (3—)6-13 cm long, very densely flowered, the ¢ part occupying the upper 1/,—'/,, 1-3 mm thick, the 2 part 4-6 mm thick. Glumes ovate or oblong-ovate, obtusish, purplish black, with wide, 3-nerved, greenish central stripe and very narrow hyaline margins, 11/,-2(-2!/,) mm long, excurrent in a hispid, short mucro sometimes 1 mm long. Utricles elliptic or obovate-elliptic, biconvex or plano-convex, multinerved, longer than (rarely about as long as) the glumes, finally divaricate, not stipitate, somewhat acuminate at the top but scarcely beaked, often puncticulate, some- times minutely and sparsely setulose at the truncate or emarginate mouth, 11/,-21/, by °/,.-1!/, mm. Nut elliptic or obovate, biconvex, brown, 11/,—1%/, by 4/,-11/, mm. Sty/e short, not thickened towards the base. Stigmas 2. Distr. W. Polynesia (Samoa, Fiji Is.) and Malesia: New Guinea (incl. New Britain), Philip- pines (Luzon, Negros, Leyte, Mindanao), N. Borneo (Mt Kinabalu), Lesser Sunda Is. (Flores), W. Java (Mt Gedeh). Ecol. On slopes in open places, on open banks of streams, in peaty alpine grasslands, and in mossy forest; on Mt Gedeh at 1600-1800 m, in the Philip- pines at 1500-2200 m (once collected at 400 m), on Mt Kinabalu at 1200 m, in New Guinea between 800 and 3800 m. Vern. Jlateum, S; Philippines: a/asas, Buk., bagibi, giron, Bag., kigid, Bon., sidak, silak, Ig. Notes. NELMES (1938) distinguished between C. graeffeana (Fiji) with its var. samoensis (Samoa), C. philippinensis (Java, Philippines, New Guinea), and C. exploratorum (Borneo), mainly using width of leaves, number and length of spikelets, and size of utricles as specific characters. For the New Guinea specimens two more specific names are available, C. pandanus Onwi and C. euphlebia S. T. BLAKE. From NeEwMEes’s later publications it is clear that the characters used for discrimination are by far not so constant as was originally supposed. There is indeed considerable variation in the specimens collected in New Guinea or in the Philippines, and even in those from the only Javanese locality. The utricles are remarkably small in the Fiji specimens, but not in those from Samoa (the latter NeLMES in 1955 no longer treated as varietally distinct.) Carex exploratorum is only known from the type collection (Mt Kinabalu, CLEMENS 34297), in which the much compressed, sterile, and elongate utricles are apparently diseased. The few well-developed, nut- bearing utricles hardly differ from those of the other Malesian materials. The glumes are larger than usual, but not until more Kinabalu specimens are available will it be possible to judge of their taxonomical value. On Mt Pulog (Luzon) the lower spikelets are often branched into a raceme of secondary spikelets, the total number of spikelets reaching up to c. 50. 170 FLORA MALESIANA [ser. I, vol. 9! 23. Section Praelongae (KUK.) NELMES, Reinwardtia | (1951) 421; RAymM. Mém. Jard. Bot. Montréal n. 53 (1959) 52, 55. — Sect. Acutae Fries subsect. Praelongae Kuk. Pfl. R. Heft 38 (1909) 345. Type species: Carex praelonga CLARKE. 52. Carex phacota SPRENG. Syst. 3 (1826) 826; NEES in Wight, Contr. (1834) 126; KuNTH, En. 2 (1837) 420; DREJER, Symb. Caric. (1844) 15, t. 4, excl. syn. C. notha et C. punctata; Bootr, Ill. 1 (1858) 63, t. 168; Boeck. Linnaea 40 (1876) 434; CLARKE, FI. Br. Ind. 6 (1894) 708; J. Linn. Soc. Bot. 37 (1904) 6 (‘phacodes’); KUx. Pfl. R. Heft 38 (1909) 350, f. S6A—C; Philip. J. Sc. 6 (1911) Bot. 62; MerRR. En. Philip. 1 (1923) 140; KUk. in Hochr. Candollea 6 (1936) 431; Bull. Jard. Bot. Btzg III, 16 (1940) 317; OHw1, Bot. Mag. Tokyo 56 (1942) 214; S. T. Brake, J. Arn. Arb. 28 (1947) 107; BACK. Bekn . Fl. Java (em. ed.) 10 (1949) fam. 246, p. 62; NELMES, Reinwardtia 1 (1951) 426; ibid. 2 (1954) 382; RaymM. Mém. Jard. Bot. Montréal n. 53 (1959) 55; Dansk Bot. Ark. 23 (1965) 257; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN. Mt. Fl. Java (1972) 48a, pl. 14-7.— C. lenticularis D. Don, Trans. Linn. Soc. 14 (1824) 331; Prod. Fl. Nepal. (1825) 43; non MicnHx, 1803. — C. platycarpa Hocust. ex STEUD. Syn. 2 (1855) 214. — C. hexasticha REINW. ex Mia. FI. Ind. Bat. 3 (1857) 353, p.p. — C. pruinosa var. aristata O. K. Rev. Gen. PI. 2 (1891) 748. — Fig. 122, 132c. Rhizome short. Stems tufted, slender, erect, triquetrous, smooth or sparsely scaberulous just below the inflorescence, 20-120 cm by 1-3 mm, surrounded below the leaves by a few ferrugineous to dark brown, bladeless sheaths split into fine, sometimes reticulate fibres. Leaves subbasal and occasionally 1-2 higher up the stem, shorter to much longer than the stems, stiff, with strongly revolute margins, gradually attenuated, 3-8 mm wide. Spikelets 4-6, rarely more, subapproximate or the lowest more distant, cylindric, 2-8(—-10) cm, terminal wholly ¢ or with a few @ flowers, erect, 1'/,4'/,. mm_ thick, remainder androgynous (g part short) or wholly 9, 4-6 mm thick, lower usually nodding on slender, smooth or scabrid, long peduncles. Lower bracts foliaceous, much overtopping the inflorescence, upper ones much smaller, none sheathing, but dark brown auricled at the base. Glumes oblong, obtuse, truncate, or bilobed-emarginate, thinly membranous, ferru- gineous to castaneous, or pale with reddish flecks, 1'/,-3 mm long, with 3-nerved central stripe excurrent in a wide, scaberulous-margined awn */2-2 mm long. Utricles elliptic, obovate, or sub- orbicular, compressed-biconvex, membranous, nerveless, narrowly marginate, glabrous, densely beset with ferrugineous or reddish, raised glandu- lar papillae, obliquely erect, shortly stipitate, 2"/4-3(—3'/2) by 11/,-21/, mm; beak extremely short, entire. Nut pyriform to suborbicular, compressed- biconvex, broadly stipitate and beaked, 13/,-2 by 11/,-13/, mm. Style thickened at the base. Stigmas 2. Distr. Ceylon, Himalayan region, N. Thailand, S. & E. China, Korea, Formosa, Ryu Kyu Is., and Japan; in Malesia: Sumatra, Java, Lesser Sunda Is. (Flores), Philippines (Luzon), N. Celebes, and New Guinea. Ecol. In open damp places, swampy grassland, 1500-2700 m. On Mt Diéng (Java) VAN STEENIS (1972, /.c.) observed in the shallow crater marsh Telaga Pangonan that it formed in a stand of Scirpus mucronatus L. large, solid, hummocky tussocks, 75 cm high and 50 cm thick, similar as do C. pani- culata L. and the grass Molinia caerulea (L.) MOoeENCH. in Europe and C. secta Bootr (‘nigger- head’) in New Zealand. This peculiar habit is obviously for a large part developing with a fluctuating water-level. — (Ed.) Vern. New Guinea: int, koole, kwajare, Enga, tsineme, Mendi, gogowe, Kapauku, yaguogufa, Okapa, tsiri ku, Upper Kangel valley. 53. Carex pruinosa Boott, Proc. Linn. Soc. 1 (1845) 255; Trans. Linn. Soc. 20 (1846) 131; Ill. 1 (1858) 65, t. 174; SrEuD. Syn. 2 (1855) 213; Mia. Fl. Ind. Bat. 3 (1856) 352; CLARKE, Fl. Br. Ind. 6 (1894) 709; J. Linn. Soc. Bot. 34 (1898) 111; ibid. 37 (1904) 7, excl. syn.; KUxK. Pfl. R. Heft 38 (1909) 352; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 62, excl. f. tristigmatosa BACK.; NELMES, Reinwardtia 1 (1951) 428; Koyama, Bot. Mag. Tokyo 72 (1959) 300, 306; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 56; Dansk Bot. Ark. 23 (1965) 258; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494. — C. hexasticha REINW. ex Mia. FI. Ind. Bat. 3 (1857) 353, p.p. — C. pruinosa f. sub- mutica O. K. Rev. Gen. Pl. 2 (1891) 748. — Fig. 123, 132d. Closely related to 52. C. phacota SPRENG., from which it is distinguished by the following characters: Glumes \anceolate or elliptic, acute to obtusish (but not truncate-emarginate), muticous or (the lower ones) excurrent in a short awn up to 1 mm long. Utricles slightly larger, 3-4 mm long, dorsally slenderly 2—S-nerved, ventrally 1-3-nerved, the glandular papillae usually whitish. Spikelets more approximate, fastigiate, and suberect to subcernu- ous, 3/4,-5'/, cm long. Leaves greyish green to glau- cous, 2-6 mm wide; sheaths not splitting into fibres. Distr. Assam, Annam, NE. Thailand; in Malesia: Java (West: several localities in Priangan; 1979] Central: Diéng plateau; East: Jang plateau). In the Leyden Herbarium there is a sheet labelled “Sumatra, WaITz’’, but WAITz probably did not visit this island. See Fl. Males. I, 1 (1950) 554. KUKENTHAL’s record for the Moluccas (Buru) refers to a collection of C. phacota; see there. The rather different ssp. maximowiczii (MIQ.) KUK. (C. maximowiczii MiqQ.) in Japan and Korea; see Koyama, /.c., with map. Ecol. Swamps, damp grassy places, along streams, 1500-2500 m. Notes. According to BACKER /.c., 52. Carex Phacota and C. pruinosa would be connected by intermediates, but I have not seen any transitional forms and I find the two clearly distinct. Carex pruinosa f. tristigmatosa BACK. belongs to 39. C. maculata BootTrt. 54. Carex teres Bootr, Ill. 1 (1858) 62, t. 167, excl. utriculo imo; BoECK. Linnaea 40 (1876) 393; CLARKE, FI. Br. Ind. 6 (1894) 707; J. Linn. Soc. Bot. 34 (1898) 110; KUx. Pfl. R. Heft 38 (1909) 348; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494. — C. teres var. spathulata Kix. Bull. Jard. Bot. Btzg III, 16 (1940) 316. — C. phacota (non SPRENG.) KUK. /.c. — C. spathulata NeLMes, Kew Bull. (1950) 207; Reinwardtia 1 (1951) 423. — C. kemi- riensis NELMES, Kew Bull. (1950) 206; Reinwardtia 1 (1951) 422. — C. petecticalis NELMES, Kew Bul!. (1950) 205; Reinwardtia 1 (1951) 424. — Fig. 123, 132e. Probably loosely tufted on a woody, ascending rhizome. Stems erect, stiff, triquetrous, smooth, 50-120 cm by 2-3 mm, at the base surrounded by bladeless to short-bladed, brownish to dark sheaths fraying into fine reticulate fibres. Leaves subbasal, crowded in the lower !/; of the stem, flattish with revolute margins, stiff, 4-7(-11) mm wide. Spikelets 4-8, cernuous, upper subapproxi- mate, fastigiate, lower more separated, gynaecan- drous usually with short ¢ base, or lower wholly 2, cylindric, dense-flowered but often laxer at the base, 3—6(—8) cm long, the 2 part 5-9 mm thick, the 3 part 2—-3(-S) mm, upper on short, lower on long, smooth peduncles. Lower bracts foliaceous, much to little exceeding the inflorescence, upper setaceous to glumiform, none sheathing but dark-auricled at the base. G/umes elliptic-oblong or oblong-obovate, subobtuse to very obtuse, truncate or bilobed, 2'/,-3'/, mm long, dark red with wide, 3-nerved, pale central stripe excurrent in a flat, more or less Cy PERACEAE—II (Kern & Nooteboom) 171 hispidulous awn up to 13/, mm long. Utricles ovate to elliptic, plano-convex, membranous, obscurely to distinctly 3—5-nerved on each face, straight, patulous, stramineous, usually densely covered with purplish flecks and minutely granular- puncticulate, scarcely stipitate, gradually beaked, 3-4(—-4'/,) by 11/,-2 mm;; beak short, '/,-*/,mm long, entire or slightly emarginate. Nut broadly elliptic or suborbicular, compressed-biconvex, 1'/,-2'/, mm long, brown, shortly stipitate and beaked. Style slightly thickened at the base. Stigmas 2, short. Distr. E. Himalaya; in Malesia: Sumatra (Atjeh: Mt Kemiri, Mt Losir, Senubong Mts; W. Coast: Mt Ophir, Mt Singgalang, Mt Kerintji), W. Java (Mt Papandajan). I have not seen KERR 21031 from Laos, Pu Bia, the type collection of C. kerrii NELMES, Kew Bull. (1939) 304; ibid. (1946) 7, 28; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 175; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 56. From the description I infer that it is not specifically distinct from C. teres. Ecol. In marshes, damp mountain meadows and heaths, in ericoid forest, in Sumatra between 2100 and 3500 m, in Java at 2300 m. Notes. Variable, particularly as to the size of spikelets and utricles, and the nervation of the lat- ter, but I do not see how to distinguish between the three “‘endemic species” into which NELMES split up the rather scanty Malesian materials. According to him C. spathulata differs from C. teres princi- pally in its stouter and shorter spikelets, C. petecti- calis from C. spathulata by its longer inflorescence (11-13 versus 7-10 cm), longer spikelets (3-6 versus 2-41/, cm), and shorter utricles (23/,—3'/, versus 31/,-4'/, mm) with shorter beak. Carex kemiriensis, only known from a single collection, is said to differ by the densely papillose utricles, a difference I am unable to find. KUKENTHAL thought the Sumatran and Javan plants to be varietally distinct from the continental Asian ones by the reticulate-fibrous basal sheaths, the shorter spikelets, the spatulate glumes, and slightly larger utricles. However, in the type collection of C. teres the basal sheaths are distinctly fibrous-reticulate and the glumes variable in shape, just like in the Malesian plants (see also Boortt’s figure!). Provisionally I prefer to include both the Indian and Malesian materials in the variable C. teres, a mountain species with a broken-up area. 24. Section Carex Sect. Vulgares (ASCHERS.) NELMES, Reinwardtia | (1951) 429. Type species: Carex acuta («) L. 55. Carex gaudichaudiana KUNTH, En. 2 (1837) 417; CLarKE, J. Linn. Soc. 37 (1904) 6; Kuk. Pfl. R. Heft 38 (1909) 312; Bot. Jahrb. 69 (1938) 264, incl. var. humilior Kix.; OHwI1, Bot. Mag. Tokyo 56 (1942) 215; S. T. Biake, J. Arn. Arb. 28 (1947) 107; Koyama, Act. Phytotax. Geobot. 18 (1959) FLORA MALESIANA [ser. I, vol. 91 22, f. 28. — C. caespitosa (non L.) R. Br. Prod. (1810) 243. — C. vulgaris FR. var. gaudichaudiana Boorr, Ill. 4.(1867) 169, t. 567; BENTH. Fl. Austr. 7 (1878) 442. — C. vulgaris (non FR.) F.v.M. Fragm. 8 (1874) 257. — C. lacerans Kux. Pfl. R. Heft 38 (1909) 326; NeLMEs, Kew Bull. (1949) 385, 392; Reinwardtia 1 (1951) 430. — Fig. 123. Rhizome emitting short to rather long, horizon- tal stolons clothed with reddish, ribbed scales. Stems erect, triquetrous, scabrid above, 3-40 (-75) cm by 1-1'/, mm. Leaves subbasal, often longer than the stem, somewhat rigid, flat or canali- culate, scabrid on the margins, 1!/,-4 mm wide, with involute margins when dry, the lower reduced to reddish to blackish red sheaths. Spikelets 3-6, approximate to rather distant, cylindric, terminal 3, 1-4 cm by 2-3 mm, peduncled, sometimes a second smaller one immediately under it also g, remainder 2 (or sometimes with a few 3 flowers at the top), sessile or lowest very shortly peduncled, suberect, (1-)2-4(-6) by 3-4(-6) mm, densely flowered. Lower bracts foliaceous, usually over- topping the inflorescence, not sheathing, dark- auricled at the base, upper much reduced. Glumes ovate or oblong-ovate, obtuse to rather acute, reddish to blackish red, with a central 3-nerved pale stripe, 2-3!/, mm long, the midrib sometimes very shortly excurrent. Utricles elliptic, ovate, or ovate-lanceolate, compressed, plano-convex, obliquely erect, submembranous, strongly 5—7(-9)- nerved on the dorsal, 3—S-nerved on the ventral face, glabrous, green, becoming brownish, often minutely reddish-puncticulate especially towards the apex, scarcely stipitate, subabruptly beaked, 2-3(—31/,) by 1-1'/. mm; beak short, entire or bidenticulate. Nut elliptic, obovate, or suborbicular, compressed, biconvex or plano-convex, shortly stipitate and apiculate, brown, 1!/,-2 mm long. Style not or scarcely thickened at the base. Stigmas Dr Distr. Australia (Tasmania to Queensland) and New Zealand to E. China and Japan; in Malesia: throughout New Guinea. Distr. maps: DURIETZ, Act. Phytogeogr. Suec. 13 (1940) 219; Koyama, Act. Phytotax. Geobot. 18 (1959) 24. Ecol. Open marshes, edges of lakes, marshy banks of streams, boggy alpine grasslands, 1450-3800 m. Vern. New Guinea: kwajare, lebandili, Enga, taua-tane, Tari. Notes. Very similar to C. nigra (L.) REICH., a common European and N. American species, and sharing with it an extreme polymorphism. Carex gaudichaudiana is characterized by the strongly nerved, more distinctly beaked utricles, and the lower bracts usually exceeding the terminal spikelet. The type of C. gaudichaudiana is from Australia. The Asiatic plants have been distinguished as var. thunbergii (STEUD.) KUK. Pfl. R. Heft 38 (1909) 313. — C. thunbergii StEuD. Flora 29 (1846) 23; Syn. 2 (1855) 221 (type from Japan). The differences between the Australian plants and the Asiatic ones are but slight; see KOYAMA, /.c. Carex appendiculata (TRAUTV.) KUK. Pfl. R. Heft 38 (1909) 338 from E. Siberia, according to Koyama also only a variety of C. gaudichaudiana, is C. nigra (L.) REICH. 25. Section Graciles [TUCKERM. En. Meth. (1843) 10]; ex KUK. Bot. Jahrb. 27 (1899) 516, quoad basion. ; OuwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 464; NeLMes, Reinwardtia 1 (1951) 353; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 52, 57; KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1962) 161. — Sect. Hymenochlaenae DREJER subsect. Graciles (KUK.) KUK. Pfl. R. Heft 38 (1909) 599. Type species: Carex gracilis R. Br. 56. Carex bilateralis HAyATA, Mat. FI. Form. (1911) 380; Ic. Pl. Form. 6 (1916) 127, f. 40 e-i; OHwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 471, p.p.; AKIYAMA, Car. Far East. Reg. Asia (1955) 105, t. 84, f. 2; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) 163. — C. subteinogyna Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 469, f. 17 & t. 15, f. 22; Aktyama, Car. Far East. Reg. Asia (1955) 102. — C. spathaceo-bracteata Kix. Bot. Jahrb. 70 (1940) 466; Necmes, Reinwardtia 1 (1951) 356. — C. acrophila S. T. Buake, J. Arn. Arb. 28 (1947) 114; Nemes, Kew Bull. (1949) 382; Reinwardtia 1 (1951) 354. — C. asperinervis Koyama, Act. Phytotax. Geobot. 16 (1955) 6, t. 3, f. U-V & f. W (ut C. subteinogyna). — Fig. 123, 132f. Rhizome very short, woody. Stems tufted, slender, erect, trigonous, smooth except on the angles above, 20-90 cm by 3/,—11/, mm, surrounded below the leaves by shining, fuscous to blackish, bladeless sheaths tending to split in front into reticulate fibres. Leaves shorter to longer than the stems, rigid, conduplicate, keeled, often flexuous at the long-attenuate top, scabrid, 1—21/. mm wide when flattened out. Inflorescence with 4-6 fascicles of (1-)2-3(-5) spikelets, narrow, 5-15 cm long; fascicles approximate or lower somewhat distant. Spikelets erect (or some possibly slightly cernuous), 1979 | linear-cylindric, simple, rather densely flowered, androgynous (9 part considerably longer than to subequalling the 3 part), 1-4 cm long, upper sessile or subsessile on included peduncles, lower on included to long-exserted, smooth or scaberu- lous peduncles. Lower bracts setaceous, shorter to longer than their fascicles but usually much shorter than the inflorescence, suddenly widening into a spathaceous, strongly nerved, reddish brown base with membranous margins clasping the base of the spikelets, with ampliate, short or long sheaths; upper bracts much reduced. Glumes slightly shorter than utricles, oblong-lanceolate, acute to very obtuse, translucent, glabrous or very sparsely hispidulous on the midrib towards the apex, 3-nerved in the centre, ferrugineous to fuscous with rather wide, whitish-hyaline margins, muti- cous or apiculate, 4-6 mm long. Utricles narrowly elliptic, plano-convex (occasionally trigonous), membranous, slenderly multinerved, suberect, reddish brown, whitish-hispidulous on the margins in the upper half and often on the nerves, 31/,—5'/, (-6!/,) by 1-1'/, mm, contracted below into a stout, stipe-like base '/,-1 mm long, rather gradually narrowed into a bidenticulate, 11/,-13/, mm long beak. Nut elliptic, oblong-elliptic, or oblong- obovate, compressed-biconvex (occasionally trigonous), not stipitate, shortly beaked, brown, 13/,-2'/, by 1-1'/; mm. Style-base _ slightly thickened. Stigmas 2 (occasionally 3), 3-7 mm long. Distr. Malesia: New Guinea (Carstensz Mts, Lake Habbema, Mt Wilhelmina, Star Mts in W., Morobe Distr., Mt Sarawaket in E.). Ecol. Alpine grasslands, marshy hollows, high mountain ridges, (2100?—)3000-4050 m. 57. Carex brunnea THUNB. FI. Jap. (1784) 38; SCHKUMR, Riedgr. 2)(1806) 16, t. Xx f. 111; Boeck. Linnaea 39 (1875) 145; CLARKE, FI. Br. Ind. 6 (1894) 705; J. Linn. Soc. Bot. 37 (1904) 5; Philip. J. Sc. 2 (1907) Bot. 107; Kuk. Pfl. R. Heft 38 (1909) 599; in Fedde, Rep. 8 (1910) 8, incl. var. subteinogyna KUx.; Philip. J. Sc. 6 (1911) Bot. 63; Camus, FI. Gén. I.-C. 7 (1922) 194; Merr. En. Philip. 1 (1923) 137; Onwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 466; Netmes, Kew Bull. (1950) 201, incl. var. dolichocarpa NELMES; Rein- wardtia 1 (1951) 357; Kew Bull. 2 (1955) 309; RAYMOND, Mém. Jard. Bot. Montréal n. 53 (1959) 58; KoyAMA, Micronesica 1 (1964) 109, incl. var. meyenii (NEES) KOYAMA; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 493. — C. gracilis R. BR. Prod. (1810) 242, non Curt. 1777; KunTH, En. 2 (1837) 513; Boortrt, Ill. 1 (1858) 59, t. 154-156; BENTH. Fl. Austr. 7 (1878) 442. — C. meyenii NEES, Nova Acta Nat. Cur. 19, Suppl. 1 (1843) 123; Krauss, Pac. Sc. 4 (1950) 264-267, f. 8-9. — C. hattoriana NAKAI ex TuyAMA, Bot. Mag. Tokyo 49 (1935) 508, t. 15; On#w1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 466; AxryamMA, Car. Far East. Reg. Asia (1955) 103, t. 82, f. 1; KoyAMA, J. Fac. Sc. Un. Cy PERACEAE—II (Kern & Nooteboom) 173 Tokyo III, 8 (1962) 164. — C. kanehirae Onwt, Act. Phytotax. Geobot. 8 (1939) 67; J. Jap. Bot. 18 (1942) 138. — C. spadiceo-vaginata OHWI, Bot. Mag. Tokyo 56 (1942) 215. — C. teinogyna (non Boott) BAcK. Bekn. Fl. Java (em. ed.) 1 (1949) fam. 246, p. 61. — C. buruensis NELMES, Kew Bull. (1950) 201; Reinwardtia 1 (1951) 360; ibid. 2 (1954) 379. — C. megacarpa KOYAMA, Bot. Mag. Tokyo 69 (1956) 210, f. 2. — Fig. 123, 132g. Rhizome very short, woody. Stems tufted, slender, erect, triquetrous, smooth, or scaberulous on the angles above, (10-)30-100 cm by !/,- 1'/, mm, surrounded below the leaves by dull brown bladeless sheaths tending to split in front into reticulate fibres. Leaves subbasal, shorter to longer than the stems, rigid, flattish or condupli- cate, rarely subfiliform, usually asperous above, gradually attenuated towards the apex, (1—)2-6 mm wide. Inflorescence with (2—)4-8 fascicles of 2-7 spikelets (rarely all the spikelets solitary), narrow, erect or more or less nodding, c. 5-50 cm long, upper fascicles or spikelets approximate and some fastigiate, lower rather distant, one of the spikelets at each node (especially at the lower ones) usually longer than the others and with some smaller spikelets branching from it. Spikelets erect or suberect, cylindric or narrowly cylindric, rather densely to rather loosely flowered, androgynous (2 part usually much longer and thicker than the 3 part, 2-4 mm thick), 1-4'/, cm long, upper sessile or subsessile on included or shortly exserted peduncles, lower on long-exserted, smooth or scaberulous peduncles. Lower bracts foliaceous or subfoliaceous, longer than their fascicles but usually shorter than the inflorescence, long-sheathing, upper bracts much reduced. Glumes from much shorter than to almost as long as the utricles, ovate to ovate-lanceolate, acute to obtuse, glabrous, obsoletely nerved but distinctly 3-nerved in the centre, muticous or apiculate, rarely an awn to 2 mm present, light castaneous, sometimes with whitish-hyaline margins, 2-4(—5) mm long. Utricles ovoid to oblong-elliptic, plano-convex, mem- branous, prominently multinerved, shortly whitish setulose at least on the margins, suberect, cinna- momeous to castaneous, cuneately tapering to a 1/,-1 mm long stipe, subabruptly beaked, 21/,—5(—6) by 1-1!/, mm; beak !/,-2 mm, bidenticulate. Nut ovate or oblong-ovate, compressed-biconvex, scarcely stipitate and beaked, stramineous, 11/,-21/, by 1-11/, mm. Style-base _ slightly thickened. Stigmas 2 (or rarely 3), up to about as long as the utricle. Distr. Widely distributed from Madagascar and the Mascarene Is. to Ceylon, India, Farther India, China, Japan, the Hawaiian Is., New Caledonia, Australia (New South Wales and Queensland); in Malesia: New Guinea, Moluccas (Buru), Philip- pines (Luzon, Bohol, Negros, Mindanao), Celebes, Lesser Sunda Is. (Lombok, Sumbawa, Flores), NW. Borneo (Sarawak), Java (a few localities in 174 FLORA MALESIANA [ser. I, vol. 9} W., Central & E.), N. Sumatra (Pajakumbuh; Gajolands: Ketambe). Ecol. Primary forest, mossy forest, exposed ridges, grassy slopes, (80—)800—2800(—3100) m, in Lombok in monsoon forest scrub on dry calcareous soil, 200-400 m, in W. Sumatra on limestone hills. Vern. Philippines: selak, Ig., tavalid, Bag. Notes. As will be seen from the above des- cription C. brunnea in the sense accepted here is extremely polymorphic, which is to be expected in so wide-ranging a species. Especially Japanese taxonomists have split it up into several micro- species and infraspecific taxa among which I do not see sharp distinctions. To me it is even very doubt- ful whether 59. C. teinogyna and 56. C. bilateralis, both closely related to C. brunnea, but which I have decided to treat provisionally as separate species, can be upheld as such in future. Typical C. brunnea, described from Japan, has small, broadly ovate, c. 2'!/, mm long utricles sub- tended by distinctly shorter glumes. Malesian specimens approaching this have only been found in N. Luzon. According to KoYAMA (1962) the greater part of the Malesian specimens belong to C. hattoriana NAKAI ex TUYAMA, occurring from Bonin and Formosa through Malesia to Australia. To him it is quite distinct from C. brunnea by the strikingly large utricles 4-4'/, mm long, the longer glumes, and the larger ¢ part of the spikelets. Carex spadiceo-vaginata Ouwi, from New Guinea, is said to differ from C. brunnea by its looser spikelets, sparsely setulose, 3 mm long utricles with longer beak. The type of C. buruensis NELMES is a very delicate plant with almost filiform leaves and reduced inflorescences, which may be due to the habitat. Essential differences with C. brunnea I cannot find. It is connected with broader-leaved specimens of C. brunnea by a collection from New Guinea, which to NELMES “represents a new variety or per- haps a new species”’. I share the opinion of Koyama that C. meyenii Nees (C. brunnea ssp. meyenii (NEES) KOYAMA) differing from typical C. brunnea by the lanceo- late, 3-4'/, mm long, long-beaked utricles setu- lose only on the margins, falls within the vari- ability of C. brunnea, but refrain from maintaining it as a subspecies. I also agree with KOYAMA in the reduction of C. kanehirae Onwt from Micronesia, to C. brunnea. | expect that several other ‘species’ described from Japan and the Pacific will have to be reduced to the polymorphic C. brunnea. Carex brunnea var. subteinogyna Kix. (non C. subteinogyna Oxnwti), from the Philippines was described as having looser spikelets, lighter coloured, longer, acuminate glumes 5 mm long, and very long-beaked and sparsely setulose utricles with longer stipe. NELMES (1950) altered this circumscription considerably so as to cover also specimens from Celebes and New Guinea, and described moreover a var. dolichocarpa NELMES with 5—53/, mm long utricles from Java. The latter variety is undoubtedly the same as C. megacarpa Koyama. I fail to see how to draw a line between the two varieties. The collection DE WILDE 14068 from the Gajolands often has 3 styles and accordingly trigonous (fertile) nuts, and the terminal spikelets are wholly 3. This might be a hybrid with 20. C. verticillata. — (Noot.) 58. Carex longipes D. Don, Trans. Linn. Soc. 14 (1825) 329; DresER, Symb. Caric. (1844) 24, t. 10; Mia. FI. Ind. Bat. 3 (1856) 347; Boorr, Ill. 4 (1867) 190; Boeck. Linnaea 40 (1876) 376; CLARKE, Fl. Br. Ind. 6 (1894) 704; J. Linn. Soc. Bot. 34 (1898) 108; ibid. 36 (1903) 295; Kuk. Pfil. R. Heft 38 (1909) 603; Bot. Jahrb. 70 (1940) 467, incl. var. ramosa KUK.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 61; NeLMeEs, Reinwardtia 1 (1951) 361; RAvM. Mém. Jard. Bot. Montréal n. 53 (1959) 58; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 493; STEEN. Mt. FI. Java (1972) pl. 14-6. — Fig. 123, 134. Rhizome very short. Stems tufted, slender, erect, trigonous, smooth except on the angles above, 15-80 cm by 1-1!/, mm, surrounded below the leaves by the fibrous, brownish remains of old leaf-sheaths. Leaves subbasal, rarely one higher up the stem, shorter to somewhat longer than the stem, flat, scabrid, 11/,-6 mm wide. Inflorescence very lax, with up to 7 spikelets, 10-40 cm long. Spikelets single at the nodes, erect or the lower cernuous, cylindric, simple or the lower 1-3 branched near the base, lax-flowered, androgynous (2 part much longer than the very short ¢ part), rarely wholly 9, 1—31/,(-41/,) cm by 4-6 mm, upper approximate, sessile, or subsessile on shortly exserted peduncles, lower distant on filiform, minutely scabrid, usually long-exserted peduncles, the lowest often near the base of the stem. Lower bracts foliaceous, shorter than inflorescence, long-sheathing, upper much reduced. Glumes 3-4 mm, shorter than the utricles, ovate-lanceolate to lanceolate, acute or subtruncate to bilobed- emarginate, pale ferrugineous with whitish hyaline margins, glabrous, with darker 3-nerved central’ stripe, the midnerve excurrent in a_ stoutish, straight, antrorsely scabrid, up to 5 mm long awn often overtopping the utricle. Utricles elliptic, compressed, biconvex, membranous, dorsally strongly 6—9-nerved, less distinctly nerved ventrally, glabrous and smooth or rarely the margins setulose, straight, obliquely erect, scarcely stipitate, pale green, fully ripe yellowish to light brown, shining, subabruptly long-beaked, 5-7 by 11/,-2 mm; beak sparsely hispid above, 2-3 mm long, deeply biden- tate (teeth '/,-*/, mm). Nut broadly elliptic to ovate, biconvex, stipitate, abruptly beaked, 2'/,-3 by 1'/,— 1979] “fd Ye TH a ® Fig. 134. Carex longipes D. Don. a. Habit, x '/3, 6. young fruit with utricle in axil of bract, c. fruit in utricle, d-e. fruits, all x 7 (VAN STEENIS 6788). Cy PERACEAE—II (Kern & Nooteboom) 175 1°/, mm. Style distinctly thickened at the base, subpersistent. Stigmas 2, shorter than the utricle. Distr. Nepal and India to China (Hupeh) and Indo-China (Tonkin); in Malesia very rare: Java (a few localities in W., Central and E.), Celebes (Menado, Poso, top of G. Lumut), and NE. New Guinea (Sattelberg). Ecol. In forests, along forest-trails, on swampy mountain meadows, 1500-2200 m. 59. Carex teinogyna Boortr, Ill. 1 (1858) 60, t. 158; BoeEck. Linnaea 39 (1875) 145; CLARKE, FI. Br. Ind. 6 (1894) 705; KUx. Pfl. R. Heft 38 (1909) 602, f. 102 F-H (‘teiogyna’), incl. var. scabriculmis KUx.; Bull. Jard. Bot. Btzg III, 16 (1940) 320, excl. pl. jav.; NELMES, Reinwardtia 2 (1954) 378; Mém. Mus. Hist. Nat. Paris n.s. B4 (1955) 146; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 59; Koyama, Bot. Mag. Tokyo 72 (1959) 307; J. Fac. Sc. Un. Tokyo III, 8 (1962) 162. — C. scabriculmis OuwI, Act. Phytotax. Geobot. 2 (1933) 27; Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 468; Ak1- YAMA, Car. Far East. Reg. Asia (1955) 102, t. 80; YOSHIKAWA, Ic. Jap. Carex 2 (1958) 246, t. 123. — C. brunnea (non. THUNB.) NELMES, Kew Bull. (1950) 200. — Fig. 123. Rhizome very short, woody. Stems tufted, slender, erect, triquetrous, smooth or scabrid on the angles above, 20-60 cm by !/,-1 mm, sur- rounded below the leaves by spadiceous sheaths ultimately fraying into persistent fibres. Leaves subbasal, shorter to slightly longer than the stems, flat to conduplicate, stiff, long-attenuate, asperous especially in the upper part, 2-3(-4) mm wide. Inflorescence with 3-6 fascicles of 1-3 spikelets, narrow, 10-20 cm long, upper fascicles approxi- mate, lower more distant. Spikelets erect, mostly simple, rarely branched at the base, lax-flowered, androgynous (2 part longer than the ¢ one), upper on scarcely, lower on shortly to rather long- exserted, smooth or scabrid peduncles, up to 3 cm long. Lower bracts foliaceous, longer than their fascicles but usually shorter than the in- florescence, long-sheathing, upper much reduced. Glumes about as long as_ utricles, oblong- lanceolate, acuminate, acute, translucent, glabrous, ferrugineous, sometimes with _ pale-hyaline margins above, muticous or with an awn up to 2 mm long, 3'/,-5 mm long. Urricles elliptic, plano-convex or compressed-biconvex, mem- branous, slenderly multinerved, — suberect, castaneous, whitish appressed-hispidulous, cuneate-stipitate, subabruptly beaked, 3'/,-4'/, (-5) by 1-1!/; mm; stipe 1/,;—'/, mm; beak 1'/,- 2 mm, bidentate. Nut elliptic to oblong-elliptic, compressed-biconvex, not stipitate, shortly beaked, brown, 2(-2!/3) by 1(-1/,) mm. Style-base slightly thickened. Stigmas 2, very long (7-12 mm), flexuous, persistent. Distr. Assam, Upper Burma, S. China (Hunan), Tonkin, Annam, Japan (Honshu, Shikoku, {ser, 1, vel.*9" 176 FLORA MALESIANA Kyushu), Korea (Quelpaert); in Malesia: N. Sumatra (Atjeh: Leuser; Gajolands: Mt Kemiri and Sangir Valley). Ecol. Rocky riverbanks, ravines, 200-1150 m. Note. Sometimes the glumes of the ¢ flowers cup-shaped, the margins connate in front. 26. Section Paludosae [Fries, Fl. Scan. (1835) 190, pro grege; TUCKERM. En. Meth. (1843) 14; O. F. LANG, Linnaea 24 (1851) 618]; BAILEY, Proc. Am. Ac. 22 (1886) 74, as group; KUK. Pfl. R. Heft 38 (1909) 730. — Subsect. Lacustres CAREY in Gray, Man. (1848) 561. — Sect. Tumidae MEINsH. Act. Hort. Petrop. 18 (1901) 283, 376. Type species: Carex paludosa GOODEN. Insufficiently known 60. Carex sp. Rhizome woody, emitting stout stolons covered with pale sheaths. Stem rather stout, triquetrous, smooth, leafy, c. 60 cm by 3 mm. Leaves basal, overtopping the stem, long-attenuate, flat or somewhat folded lengthwise, septate-nodulose, glaucous-green, subcoriaceous, c. 5 mm wide; ligule lanceolate, acute, c. 11/, cm long; margins smooth below, scabrid above; lower sheaths strongly septate-nodulose, not fibrous, stramineous or light brown. Inflorescence erect, consisting of 4 spikelets. Terminal spikelet g, c. 1!/, cm long, 2 mm wide, lateral spikelets 9, erect, cylindrical, densely flowered, the upper 2 approximate, sub- sessile, the lowest distant on a smooth, 1'/, cm long peduncle. Bracts foliaceous overtopping the inflorescence, not or scarcely sheathing. Glumes of the 2 flowers ovate, deeply emarginate at the top, pale with purplish margins, c. 3 mm long, the strong midrib excurrent in a firm, smooth or scabrid awn about as long as the glume; glumes of the 3 flowers similar, more oblong, purplish with pale centre. Utricles young, lanceolate, glabrous, many-nerved, 5—S!/, by 11/, mm, rather gradually narrowed into the stout, straight, bidentate beak with straight, 1/; mm long teeth. Style straight, ciliolate, c. 2 mm, not thickened at the base. Stigmas 3, longer than the style. Distr. Malesia: New Guinea: Western High- lands, Sirunki, swamp near Nanguris village, in fast flowing very deep water course (water depth approx. 230 cm), at c. 2500 m, 14 Sept. 1962: WALKER ANU 691 (CANB). Note. Only a single, immature specimen was collected. The plant is related to the Eurasian C. riparia CurtT., from which it differs by the narrower leaves, the small, single 3 spikelets, and the deeply incised g and 2 glumes, and to the Japanese C. rugulosa Kix. (not seen). II. Subgenus Vignea (BEAUV.) CLARKE, FI. Br. Ind. 6 (1894) 700; KUx. Bot. Jahrb. 27 (1899) 495; Pfi. R. Heft 38 (1909) 111; NeLMes, Reinwardtia 1 (1951) 431. — Vignea BEAUV. in Lestib. Ess. Fam. Cyp. (1819) 22. Type species: Carex arenaria L. 27. Section Divisae [CuRIsT, Bull. Soc. Bot. Belg. 24 (1885) 18, nomen]; (KUK.) KUx. Pfl. R. Heft 38 (1909) 119; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 234. — Sect. Capituligerae KUx. subsect. Divisae CHRIST ex KUK. Bot. Jahrb. 27 (1899) 500. Type species: Carex divisa Hups. 61. Carex duriuscula C. A. Mey. Mém. Ac. St. Pétersb. 1 (1831) 214, t. 8; Kunru, En. 2 (1837) 373; OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 234. — C. stenophylla WAHLENB. var. duriuscula Trauty. Act. Hort. Petrop. 10 (1887- 89) 537; KUK. Pfl. R. Heft 38 (1909) 121. — Carex sp. NELMES, Kew Bull. (1949) 386; Reinwardtia 1 (1951) 446. Rhizome creeping, slender, c. 1 mm thick, clothed with brown sheathing scales. Stems erect 1979 or curved, trigonous, smooth, or scaberulous below the inflorescence, 7-15 cm by c. !/; mm. Leaves subbasal, shorter than the stems, rigid, curved, circinnate at the apex, canaliculate to con- volute, smooth or scaberulous towards the apex, c. 1 mm wide. Spikelets 4-7, androgynous (¢ part about as long as the 9), sessile, approximate, elliptic-lanceolate (very young), subdense-flowered, 5-7 mm long, forming a more or less oblong head 1-11/, by c. !/, cm. Bracts glumiform, the lower long-awned. Glumes oblong-lanceolate or ovate- lanceolate, acutish, very thin, translucent, castaneous with broad whitish-hyaline margins, 3'/,4 mm long, the midrib not extending to the apex to very shortly excurrent. Utricles (very immature) elliptic, plano-convex, nerveless, gla- brous, scabrid on the margins in the upper half, 3-4 by c. 1 mn, tapering above into a short beak 177 with oblique mouth. Stigmas 2, about as long as the utricle. Distr. Siberia, N. Mongolia and Manchuria; in Malesia: New Guinea, Lake Habbema (Brass 9235) in W., Mt Victoria (LAE 61670) and Kondo, Mt Hagen (NGF 43535) in E. Ecol. In New Guinea in sunny bog and alpine grassland, 2400-3225 m. Note. NELMES, /.c., supposed this New Guinea collection to be close to C. arenicola F. SCHMIDT, but I agree with S. T. BLAKE, J. Arn. Arb. 28 (1947) 116, that it is evidently allied to C. stenophylla WAHLENB. of the northern hemisphere. The resem- blance to small, narrow-leaved specimens of the latter is surprising. I have seen extra-Malesian material of C. duriuscula; the Brass collection very well matches the collection I saw of this species, which is apparently very near to C. stenophylla. 28. Section Paniculatae [KUNTH, En. 2 (1837) 389, nomen]; MEINSH. Act. Hort. Petrop. 18 (1901) 281, 313; Kuk. Pfl. R. Heft 38 (1909) 174; NeLmes, Reinwardtia 1 (1951) 432. — Sect. Muricatae Friks subsect. Paniculatae KUNTH ex KUK. Bot. Jahrb. 27 (1899) 515. Type species: Carex paniculata L. 62. Carex appressa R. Br. Prod. (1810) 242; KunrTH, En. 2 (1837) 389; KUNZE, Suppl. Riedgr. (1840-50) 45, t. 11; Boorrt, Ill. 1 (1858) 46, t. 119, 120; Boeck. Linnaea 39 (1875) 99; Kix. Pfl. R. Heft 38 (1909) 178, f. 29 E-J; Bot. Jahrb. 69 (1938) 261; Onwt, Bot. Mag. Tokyo 56 (1942) 214; S. T. BLake, J. Arn. Arb. 28 (1947) 101; NELMEs, Kew Bull. (1949) 386, 392; Reinwardtia 1 (1951) 432; Kew Bull. 2 (1955) 318. — C. paniculata (non L.) BENTH. FI. Austr. 7 (1878) 440. — Fig. 123. Rhizome short, stout, woody, forming very dense clumps. Stems densely tufted, erect, triquet- rous (often very acutely so), rigid, scabrid on the angles above (see note), 30-180 cm by 2-4 mm, surrounded below the leaves by spadiceous, dark- nerved, bladeless sheaths and their fibrous remains. Leaves in the lower 1/,—"/, of the stem, shorter to longer than the stem, very stiff, flat to conduplicate, long-acuminate, with very scabrous margins, pale green, 3-10 mm wide. Inflorescence a slender, oblong-cylindric, contracted, spike-like panicle, 5-25 by 1-2 cm; branches numerous, erect, often appressed or even partly adnate to the stems, upper crowded, lower approximate or slightly distant. Bracts inconspicuous, setaceous, lower sometimes as long as the branches, upper reduced to glumes. Spikelets very numerous, sessile, androgynous, ovoid or ovoid-lanceolate, few- flowered (¢ and 2 parts about equal in length), 4-8 mm long. Glumes ovate, acute, thin and translucent, with ciliolate margins, otherwise glabrous, slenderly nerved, ferrugineous to casta- neous with hyaline margins, 2-3 mm long, the midrib often excurrent in a short mucro up to 1/, mm long. Utricles ovate, plano-convex, with obtuse margins, coriaceous, 6—-12-nerved dorsally, 3-6-nerved ventrally, glabrous, distinctly setulose- margined above, subpatent, stramineous to dark brown, rounded at the spongy-thickened base, shortly stipitate, subabruptly beaked, 2'/,-31/, by 1'/,-2 mm; beak '/,-1 mm long, bidentate, grooved on the back, with slightly oblique mouth. Nut ovate to obovate, compressed-biconvex, broadly stipitate, beakless or shortly beaked, 11,-13/, by c. 1 mm. Stigmas 2. Distr. Widespread in Australia, also in New Zealand and New Caledonia; in Malesia: New Guinea (Arfak Mts, Lake Habbema in W. and Western Highlands, Finisterre Range, Mt Sara- waket, in E.). Ecol. Open marshes, shores of lakes, alpine meadows, 1900-3225 m, on Mt Sarawaket as low as 900 m. Vern. Kwajare, Enga. Note. The stems are sometimes smooth or almost so: var. virgata (SOL. ex Boott) KUx. Pfi. R. Heft 38 (1909) 179 (C. virgata SoL. ex Boott in Hook. f. Fl. Nov. Zel. 1 (1853) 282; Ill. 1 (1858) 46, t. 121, 122; Boeck. Linnaea 39 (1875) 98.) — Only known from New Zealand and Tasmania. 178 FLORA MALESIANA [ser. I, vol. 91 29. Section Multiflorae (KUNTH, En. 2 (1837) 387, nomen]; (CAREY) BAILEY, Proc. Am. Ac. 22 (1886) 135, as group; KUK. Pfl. R. Heft 38 (1909) 142; NeLMgs, Reinwardtia 1 (1951) 433; RayM. Mém. Jard. Bot. Montréal n. 53 (1959) 19. — Sect. Vignea BEAUV. subsect. Multiflorae CAREY in Gray, Man. Bot. N. Un. St. (1848) 540. — Sect. Muricatae Fries subsect. Multiflorae KUK. Bot. Jahrb. 27 (1899) 514. Type species: Carex multiflora MUEHL. ex WILLD. 63. Carex nubigena D. Don, Trans. Linn. Soc. 14 Candollea 6 (1936) 430; BAcK. Bekn. Fl. Java (1825) 326; Nees in Wight, Contr. (1834) 120; (em. ed.) 10 (1949) fam. 246, p. 60; NELMEs, KuNTH, En. 2 (1837) 385; Boorrt, Ill. 1 (1858) 1, Reinwardtia 1 (1951) 434, excl. specim. Sumatrae; t. 2; Boeck. Linnaea 39 (1875) 90; CLARKE, Fl. KoyAMaA, Bot. Mag. Tokyo 72 (1959) 302; KERN in Br. Ind. 6 (1894) 702; J. Linn. Soc. Bot. 37 (1904) Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN. Mt 5, incl. var. fallax CLARKE; KUK. Pfl. R. Heft 38 FI. Java (1972) pl. 14-1. — C. fallax Steup. [in (1909) 145, incl. var. fallax KtxK.; in Hochr. Zoll. Syst. Verz. (1854) 60]; Syn. 2 (1855) 189; ~ Fig. 135. Carex nubigena D. Don in tufts around a shallow depression (sawahan), probably the site of an old silted-up crater, covered with a heavily deer-grazed, very short turf of herbs and grasses subject to frost in the dry season; background some scattered Casuarina junghuhniana Miq. Mixed with tussocks of Pennisetum alopecuroides (L.) Spr. East Java, Mt Jang, c. 2000 m altitude (VAN STEENIS, 1936). 1979] Mia. FI. Ind. Bat. 3 (1856) 347; Boeck. Linnaea 39 (1875) 57; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 247 (var. franchetiana Onwi). — Fig. 123, 135. Rhizome short, woody, forming dense tufts. Stems erect, slender but rigid, smooth or slightly scaberulous just below the inflorescence, obtusely trigonous, 20—60(-100) cm by 1-3 mm, clothed below the leaves by brownish to pale, bladeless sheaths and their fibrous remains. Leaves subbasal, shorter to longer than the stems, rigid, canaliculate to conduplicate, with scabrid margins, 1!/,-3 mm wide. Inflorescence pyramidal to oblong, head-like or spike-like, 1-3(—5) cm by 7-13 mm. Spikelets 5—10(-15), crowded, or lowest 1-2 sometimes slightly separated, ovoid to subglobose, sessile, androgynous (with very few ¢ flowers), 5-10 by 5-7 mm. Lower 2-3 bracts foliaceous, mem- branous-margined at the base, erect or curved, lowest much exceeding, other equalling to exceed- ing the inflorescence, upper much reduced to glume-like, none sheathing. Glumes ovate to oblong-lanceolate, acutish, very thin, translucent, ferrugineous to whitish hyaline, with brownish to greenish, 3-nerved central stripe, 2!/,—-3'/, mm long, with a mucro up to 1 mm long. Utricles ovate or Cy PERACEAE—II (Kern & Nooteboom) 179 ovate-lanceolate, plano-convex, membranous, strongly many-nerved on both faces, winged, glabrous, obliquely erect, greenish to brown, spongy-thickened at the base, shortly stipitate, subgradually beaked, 3'/,-4'/, by 1'/,-2 mm; beak c. 1'/, mm, serrulate-margined, dorsally grooved, bidentate. Nut elliptic to suborbicular, biconvex, broadly stipitate, shortly beaked, 1'/,-13/, by 4/--1 mm. Style-base not or scarcely thickened. Stigmas 2 (according to Boott occasionally 3). Distr. From Ceylon, S. India and the Himalaya to China (Hupeh, Yunnan), Formosa and Japan; in Malesia: Java (Central: Diéng Plateau; East: Mts Kawi, Tengger-Semeru & Jang). Distr. map in Bot. Mag. Tokyo 72 (1959) 302, f. 23 (the area in Malesia should be restricted to Central and East Java!). Ecol. Marshy places, along streams, according to BACKER 1600-3000 m, sometimes gregarious. On Mt Jang deer feed on this species of which the leaf- bases have a sweet taste, as in Gahnia javanica (VAN STEENIS, /.c.). Note. The immature collection LAE 65208 from New Guinea (W. slope of Mt Kenive, 9° 10’ S and 147° 45’ E) might possibly be referred to this species. — (Noot.) 30. Section Stellulatae KUNTH, En. 2 (1837) 399; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 253; NELMES, Reinwardtia 1 (1951) 441. — Sect. Elongatae KUNTH sensu KUK. Pfl. R. Heft 38 (1909) 226, p.p. Type species: Carex stellulata GOODEN. 64. Carex echinata Murr. Prod. Stirp. Gotting. (1770) 76; Boeck. Linnaea 39 (1875) 124; BENTH. Fl. Austr. 7 (1878) 439. — C. stellulata GOODEN. Trans. Linn. Soc. 2 (1794) 144; Kx. Pfl. R. Heft 38 (1909) 228. — C. nubigena (non Don) KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 314. — C. perileia S. T. Bake, J. Arn. Arb. 28 (1947) 102; NELMEs, Kew Bull. (1949) 386, 392; Reinwardtia 1 (1951) 441. — C. gajonum NELMES, Kew Bull. (1952) 84; Reinwardtia 2 (1954) 382. — C. omiana FRANCH. & SAv. var. perileia KoYAMA, Bot. Mag. Tokyo 69 (1956) 211. — Fig. 123, 131h. Rhizome short, forming dense tufts. Stems slender, erect, obtusely trigonous below, more acutely so above, smooth or slightly scaberulous just below the inflorescence, 15—90 cm by 1-2 mm. Leaves subbasa!, shorter to longer than the stems, canaliculate-conduplicate, long-attenuate, scabrid on the margins above, 1-3 mm wide, the lower ones reduced to pale to castaneous, bladeless sheaths. Inflorescence ovoid to oblong, head-like or spike- like, 11/,-3'/,(-4'/.) cm long. Spikelets 3-8, gynaecandrous (¢ flowers very few), ellipsoid, obovoid or ovoid to subglobose, sessile, dense- flowered, approximate or lowest somewhat distant, 5-10 by 5-8 mm, finally squarrose by the widely spreading utricles. Bracts glumiform, the lowest with a setaceous awn, others more shortly aristate, or indistinguishable from the glumes or the lower bracts foliaceous, overtopping the inflorescence. Glumes ovate, acute, thin, wholly pale or brownish with wide whitish-hyaline margins, 3-nerved in the centre, 2!/,—31/, mm long. Utricles elliptic or ovate- lanceolate to broadly ovate, rounded to cardate at the base, plano-convex, membranous, several- nerved on both faces, glabrous, smooth, straight or slightly recurved, widely patent to reflexed when mature, spongy-thickened at the base, scarcely stipitate, (3—)4—5(—5'/,) by (1-)1'/.-2 mm, yellowish green to brownish, subgradually nar- rowed into a bidenticulate beak more or less scabrid on the margins or rarely wholly smooth and with a dorsal split with brown overlapping margins. Nut oblong-ovoid, plano-convex, scarcely stipitate and beaked, brownish, 2-2'/, by 1-11/, mm. Style-base slightly thickened. Stigmas 2. Distr. N. America, Eurasia to Australia and New Zealand; in Malesia: N. Sumatra (Gajolands: 180 FLORA MALESIANA [ser. I, vol. 91 Mts Kemiri, Losir, Bandahara) and New Guinea (Arfak and Lake Habbema in W., and many mountains in E.). Ecol. Peat swamps, wet alpine grasslands, marshy lake shores, 1850-3600 m, locally often abundant. Vern. New Guinea: kisis, pemp, Papua, Mendi lang., koali, Enga lang., armemséna, kul, Mani- kiong lang. Notes. In the wide sense here accepted C. echi- nata is a widely spread species. The characters used for differentiating the numerous microspecies described in sect. Stellulatae are far from reliable; they mainly refer to the width of the leaves, the size of the utricles, the scabridity of their margins and the intensity of their nervation. For N. America MACKENzIE, N. Am. FI. 18 (1931) 99-114 recog- nized c. 20 spp. which can hardly be maintained. I have not seen Japanese materials of the section, but to judge from the descriptions and figures C. basilata Ouwi, Act. Phytotax. Geobot. 11 (1942) 258; YosHIKAWA, Ic. Jap. Carex 3 (1960) 296, t. 148 [C. muricata (non L.) OHwi1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 253; AktyAMa, Car. Far East. Reg. Asia (1955) 64, t. 31] is hardly different from the European plants, and C. omiana FRANCH. & SAv.; OHwI, /.c. 254; AKIYAMA, l.c. t. 32 with its lanceolate utricles less scabrid mar- gins must be very near to the New Guinean speci- mens. Carex perileia S. T. BLAKE was based on a specimen with a single fruiting culm, and distin- guished from C. echinata by its narrower leaves and its longer utricles with relatively larger beak deeply split on the back with entirely smooth margins. Several additional collections have shown that size and scabridity of the utricles are very variable. Sometimes the utricles are not longer and hardly less scabrid than in European materials. In typical C. echinata the uppermost spikelet is seemingly long-stalked by the relatively large number of 3 flowers at its base and sometimes almost wholly 3, in the New Guinean specimens also this spikelet is sessile as there are so few ¢ flowers that it has a wholly 2 appearance. This may be the same in some Australian forms of C. echinata, as according to BENTHAM, /.c., there are very few ¢ flowers at the base of the spikelets, sometimes even none. Whether C. perileia represents a special New Guinea race cannot be decided without a critical study of the whole section or of at least the Australasian and E. Asian representatives; in my opinion it is not specifically distinct. Carex gajonum NeELMeEs, from N. Sumatra (Gajolands: Mts Losir and Kemiri), was distin- guished because the lower bracts are foliaceous, much overtopping the inflorescence, and the utricles being broader, cordate at the base, and obliquely erect instead of widely patent when mature. In some Sumatran plants, however, the lower bracts are setaceous, in some New Guinean ones they are foliaceous, overtopping the inflores- cence. In the collection DE WILDE 13323, from Mt Bandahara, the ripe utricles are patent as in true C. echinata, making the spikelets squarrose. Furthermore I compared several utricles, and al- though they are generally broader in Sumatra, there is no constant difference. — (Noor.) 31. Section Elongatae KUNTH, En. 2 (1837) 402; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 256; Kuk. Pfl. R. Heft 38 (1909) 226, p.p.; NELMES, Reinwardtia 1 (1951) 438. Type species: Carex elongata L. 65. Carex remota LINNE, Amoen. 4 (1759) 293; Boeck. Linnaea 39 (1875) 129; KUx. Pfi. R. Heft 38 (1909) 233. The typical subspecies is widely distributed in Europe, extending to N. Africa and W. Asia; in Malesia two other subspecies occur: a. ssp. alta (BootT) KUx. Pfl. R. Heft 38 (1909) 234, incl. var. brizopyrum KwtKk.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 60; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 494; STEEN. Mt. Fl. Java (1972) pl. 14-4. — C. alta Boortrt, Proc. Linn. Soc. 1 (1845) 254; Trans. Linn. Soc. 20 (1846) 130; Ill. 1 (1858) 59, t. 153; Mia. FI. Ind. Bat. 3 (1855) 347; Boeck. Linnaea 39 (1875) 126; CLARKE, FI. Br. Ind. 6 (1894) 707; J. Linn. Soc. 37 (1904) 6; Netmes, Reinwardtia 1 (1951) 438. — C. brizopyrum KUNZE, Suppl. Riedgr. (1840-50) 169, t. 43. — C. remota var. rochebrunii CLARKE, J. Linn. Soc. 37 (1904) 6, p.p. (quoad ZOLLINGER 3192). — C. craspedotricha NELMES, Kew Bull. (1939) 657; ibid. (1946) 28; RAyM. Mém. Jard. Bot. Montréal n. 53 (1959) 19; Dansk Bot. Ark. 23 (1965) 252. — ? C. imbricata Kx. in Hand.-Mazz. Symb. Sin. 7 (1936) 1260, f. 38 n. 2. —? C. squamata Krecz. Not. Syst. Herb. Inst. Bot. Ac. Sc. URSS 9 (1946) 196. — Fig. 123. Rhizome very short, woody. Stems densely tufted, stiff to rather weak, erect or suberect, smooth, 15-80(—120) cm by 1—1!/, mm, surrounded at the base by brown, fibrous remains of old leaf- sheaths. Leaves in the lower 1/3 of the stem, shorter to longer than the stems, flat, scabrid on the margins especially towards the long-attenuated 1979 ] apex, 1'/,-4 mm wide; sheaths long, mouth concave in front. Spikelets 5—-18(—24) in an up to c. 15 cm long, spiciform inflorescence, sessile, obliquely erect, upper densely crowded, lower separated to their own length from one another, lowest 1-2 often distant, ellipsoid to ellipsoid- cylindric, densely flowered, gynaecandrous with only a few ¢ flowers, 5-15 by 3-5 mm. Lower bracts foliaceous, far exceeding the inflorescence, not sheathing, upper glumiform. Glumes ovate or oblong-ovate, acute, thin and translucent, finely nerved, whitish with greenish 3-nerved centre, 2-3 mm long, usually excurrent in a short, up to 1/, mm long mucro. Urricles ellipsoid or ovoid- ellipsoid (often in outline broadest above the middle because of the wings), obliquely erect, plano-convex, membranous, slenderly nerved on both sides in the lower centre, glabrous, winged in the upper */,—°/,, yellowish green to brownish, scarcely stipitate, subabruptly beaked, 2°/,-3 by 1-1'/, mm; wings varying in width, denticulate- ciliate; beak c. '/, mm long, bidentate with slender, straight teeth. Nut ellipsoid or ovoid, biconvex, shortly stipitate and beaked, 11/,-12/; by ?/,—5/; mm. Style slightly thickened at the base. Stigmas 2. Distr. Insufficiently known; according to literature from India (Himalaya) to Central China; C. craspedotricha NetMes from Thailand un- doubtedly belongs here. The type collection of both C. alta Boott and C. brizopyrum KUNZE are from Java; in Malesia only known from Java (from Mt Patuha in W. to Jang in E.). Ecol. In moist or swampy grasslands, along water-courses, damp forest-borders, 1500-2200 m; once collected at 1150 m. Cy PERACEAE—II (Kern & Nooteboom) 18] b. ssp. rochebrunii (FRANCH. & SAv.) Kix. Pfl. R. Heft 38 (1909) 234; Kern in Back. & Bakh. f. FI. Java 3 (1968) 494. — C. rochebrunii FRANCH. & Sav. En. Pl. Jap. 2 (1879) 126, 555; Netmes, Kew Bull. (1946) 29. — C. remota var. rochebrunii CLARKE, FI. Br. Ind. 6 (1894) 707. — C. monopleura KReECzZ. Not. Syst. Herb. Inst. Bot. Ac. Sc. URSS 7 (1937) 35; NeLMes, Kew Bull. (1950) 208; Rein- wardtia 1 (1951) 440. Usually slenderer than ssp. alta, with narrower leaves. Inflorescence up to 10 cm long, with 3-9 spikelets; upper spikelets not rarely sterile. Glumes */,-4 mm long. Utricles lanceolate, ventrally nerveless or nearly so, narrowly winged, 4—4!/, mm long. Nut elliptic-oblong, c. 2 by 1 mm. Distr. India (Sikkim), China, Japan, Formosa; in Malesia: Sumatra (Atjeh: G. Leuser, and W.: Mt Kerintji) and W. Java (Mt Papandajan). Ecol. Marshy places in forests, along rivers and streamlets, 1700-2700 m. Notes. Ssp. rochebrunii is very near to the European ssp. remota, mainly differing by the narrow, lanceolate, narrowly winged utricles and the oblong nuts. In my opinion the differences between C. rochebrunii (described from Japan) and C. monopleura (based on Sikkim material) are too slight to justify specific separation. In its extreme tall form, with relatively dense inflorescence and broadly winged utricles, ssp. alta gives the impression of being a separate species, but slender, weak plants occur (they were distinguished as var. brizopyrum by KUKENTHAL), and the width of the wings of the utricles is variable. KUNZE, /.c., in describing C. brizopyrum, stressed the importance of the bristly appendage of the con- nective, but thisisalso found in European C. remota. 32. Section Heleonastes KUNTH, En. 2 (1837) 393; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 261; NELMES, Reinwardtia 1 (1951) 443. Type species: Carex heleonastes EHRH. 66. Carex curta GOODEN. Trans. Linn. Soc. 2 (1794) 145; ScHkunr, Riedgr. 1 (1801) 43, t. C, f. 13; KuNTH, En. 2 (1837) 403; S. T. BLAKE, J. Arn. Arb. 28 (1947) 101; Netmes, Kew Bull. (1949) 386; Reinwardtia 1 (1951) 443; YosHIKAWA, Ic. Jap. Carex 1 (1957) 32, t. 16. — C. canescens (non L.) Boorrt, Ill. 4 (1867) 154, t. 496; Boeck. Linnaea 39 (1875) 122, excl. var. 8; BENTH. FI. Austr. 7 (1878) 439; CLARKE, FI. Br. Ind. 6 (1894) 706; KUxK. Pfl. R. Heft 38 (1909) 216, f. 35 C-E; OxwI, Mem. Coll. Sc. Kyoto Imp. Un. B11 (1936) 268; AKIYAMA, Car. Far East. Reg. Asia (1955) 60, t. 26. — Fig. 123. Rhizome short, forming dense tufts. Stems slender, erect, triquetrous, slightly scaberulous just below the inflorescence, 10-40(-60) cm by I- 11/, mm, surrounded below the leaves by light brown, withered leaf-bases. Leaves subbasal, shorter to longer than the stems, flat to condupli- cate, grey-green, scabrid on the margins above, long-attenuate to the triquetrous apex, 2—3(-5) mm wide. Inflorescence oblong, spike-like, 2—31/,(—6) cm long. Spikelets 4-8, gynaecandrous (¢ part few- flowered, inconspicuous), ovoid, ellipsoid, or sub- cylindric, sessile, dense-flowered, approximate or contiguous, 6-9 by 4-5 mm. Bracts glumiform, shortly aristate, the lowest rarely subulate to subherbaceous. G/lumes ovate, acute, very thin, whitish, 3-nerved in the centre, 2—2!/, mm long, the midrib sometimes slightly excurrent. Utricles ovate 182 FLORA MALESIANA [ser. I, vol. 9! or ovate-elliptic, biconvex, membranous, several- nerved on both faces, scarcely marginate, glabrous or slightly papillose at the apex, golden yellow, densely whitish puncticulate, straight, suberect, very shortly stipitate, 2—2!/, by 1-11/, mm, scarcely beaked; mouth minutely bidenticulate. Nut elliptic to broadly ovate, plano-convex or biconvex, scarcely stipitate, abruptly beaked, light brown, c. 1'/, by 1 mm. Style-base not thickened. Stigmas 2 Distr. Widely spread in N. America, extra- tropical S. America, Eurasia (also Kashmir and Japan), and SE. Australia (New South Wales, Victoria); in Malesia: New Guinea (Lake Habbema in W., Neon basin 15 km NNE of Woitape in E.). Ecol. Plentiful in open grassland on wet sandy soil and on marshy flats, 2800-c. 3225 m Note. For the correct name of this species, which has commonly been called C. canescens L., see E. S. MARSHALL, J. Bot. 45 (1907) 365; S. T. BLAKE, J. Arn. Arb. 28 (1947) 101; NeLmes, Reinwardtia 1 (1951) 444. Doubtful (Sect. Vigneastra) Carex subfilicina OHwi, Bot. Mag. Tokyo 56 (1942) 213; Netmes, Reinwardtia 1 (1951) 446 (under doubtful species). — I did not see any collections of this species. From the description it is possibly conspecific with either C. filicina NEES or C. cruciata WAHLENB. — (Noot.) Excluded Carex arnottiana NEES ex DREJER, Symb. Caric. (1844) 16, t. 5; Boeck. Linnaea 40 (1876) 436. — This species is endemic in Ceylon. BOECKELER, l.c., cited it also to occur in “Java, alt. 6000 ped. (Arnott). As the annotation for the type specimen is exactly matching this and ARNOTT never collected in Java, this record is based on a miswritten label. Carex cladostachya \WAHLENB. Vet.-Ak. Handl. 24 (1803) 149; Boeck. Linnaea 40 (1876) 361; KUK. Pfl. R. Heft 38 (1909) 267. — This Central Ameri- can species was recorded by BOECKELER, /.c., also from the Philippines on the strength of CUMING 625. This specimen must certainly belong to CuMING’s American collections, which were separately numbered from the Philippine ones. Carex dimorpholepis StEuD. Syn. 2 (1855) 214. — C. cernua Bootrt, Ill. 4 (1867) 171, t. 578, non J. F. Gmev. 1791. — The old specimens of this species in the Leyden Herbarium annotated “Arch. Ind. ? Leg. ?” in all probability do not originate from the Malayan Archipelago, as they would be the only Malesian collection extant. C. dimor- pholepis is known from India, Upper Burma, and China to Korea and Japan. See NELMES, Kew Bull. (1950) 208. Carex divulsa STOKES var. javanica NELMES, Kew Bull. (1950) 208; Reinwardtia 1 (1951) 436. — See below under C. pairaei. Carex erythrolepis Kix. Pfl. R. Heft 38 (1909) 628; BAcK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 65. — ‘Java, Hillebrand in herb. musei berol. sine indicatione loci.”” Wrongly localized; see Fl. Males. I, 1 (1950) 232. The name must be referred to the synonymy of C. wahuensis C. A. Mey., an endemic of the Hawaiian Islands. See NELMES, Kew Bull. (1950) 203. Carex haenkeana PrRESL, Rel. Haenk. 1 (1828) 205; SteuD. Syn. 2 (1855) 241; Mia. FI. Ind. Bat. 3 (1857) 354; F.-ViL_. Nov. App. (1882) 310. — “Habitat in insulis Philippinis.”’ This is C. pseudo- cyperus L. var. haenkeana (PRESL) KUxK. Bot. Jahrb. 27 (1899) 550; Pfl. R. Heft 38 (1909) 696, a S. American plant. The record attributed to the Philippines was based on an erroneously localized Malaspina Expedition specimen. See MERRILL, En. Philip. 1 (1923) 142. Carex lindleyana NEES in Wight, Contr. (1834) 121; Boeck. Linnaea 40 (1876) 362; KUK. Pfl. R. Heft 38 (1909) 280. — This is a species from the Deccan Peninsula and Ceylon. BOECKELER recorded it besides from the Philippines on the strength of an unnumbered collection of Cuminc. If correctly identified this certainly rests on an erroneous localisation; it might have been collected in Ceylon by CuMING himself. Carex pairaei F. SCHULTZ var. javanica NELMES, Kew Bull. (1950) 208; Reinwardtia 1 (1951) 437. Carex divulsa Stokes and C. muricata L. (= C. pairaei F. SCHULTZ) were recorded by Nemes from Java, the only ones from outside Europe, both based on a single specimen, that of C. divulsa even on a single culm. The scanty materials were alleged to have been collected by RIDLEY during his Java tour on Mt Papandajan. However, there is no doubt that the specimens were mislocalized, possibly by using drying paper to which remnants of rambles in Europe adhered. The Carex flora of the easily accessible Mt Papan- dajan is well known, e.g. by the intensive search for Carices by VAN STEENIS. Similar records opposing all rules of plant distri- bution are those of Elisma natans for Java, Linaria alpina for the Malay Peninsula, and Scheuchzeria palustris for Sumatra (see Taxon 5, 1956, 157). NELMES, in litt. Dec. 17, 1953, admitted that the records must be due to an error. Carex scabrifolia STEUD. Syn. 2 (1855) 237; Mia. FI. Ind. Bat. 3 (1857) 354. — ‘Carex nr 67 et 83. Herb. Zollinger. Java.”’ The type is not from Java, but from Japan, Decima in Nagasaki. See ZOLL. 1979 ] Cy PERACEAE—II (Kern & Nooteboom) 183 Syst. Verz. 1 (1854) 60; KUK. Pfl. R. Heft 38 (1909) Carex typhoides Bory; Hassk. Cat. Bog. 296; 737; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1962) Mua. FI. Ind. Bat. 3 (1857) 352. — This is C. bor- 248. — HUBERT WINKLER 2097, wrongly distributed bonica LAMK from the Mascarenes; certainly not as C. scabrifolia, belongs to C. baccans NEES. occurring in Malesia. 29. UNCINIA Pers. Syn. Pl. 2 (1807) 534; Boeck. Linnaea 41 (1877) 339; CLARKE, J. Linn. Soc. Bot. 20 (1883) 389; KUK. Pfl. R. Heft 38 (1909) 50; NeLMes, Kew Bull. (1949) 140; HAMLIN, Dom. Mus. Bull. 19 (1959) 1; BALGooy, Pac. Pl. Areas 3 (1975) 320, map 209; Noor. Blumea 24 (1979) 511. — Fig. 137a-d. Perennial, monoecious herbs, glabrous (or with hispid utricles). Stems central, tufted or approximate on a more or less creeping rhizome, erect or ascendent, sharply trigonous to subterete, striate, smooth, or scabrid below the inflorescence. Leaves narrowly linear, flat or involute, more or less scabrid on margins and nerves; basal sheaths bladeless, often disintegrating into fibres. Inflorescence a single, terminal spikelet; ¢ part above, shorter than the lower 2 part. Glumes spirally arranged, ovate to oblong, concave, persistent or caducous, all flower-bearing, the lowest often produced into a setaceous to foliaceous bract. — 3 Flowers naked, consisting of (1—)2—3 stamens with linear (or dilated, New World spp.) filaments and linear anthers; connective shortly produced. — 2 Flowers naked, enclosed in a bottle-shaped, obtusely trigonous organ (utricle, perigynium) which is closed up to the truncate top, glabrous (in all Mal. and Austr. spp.) or hispid; style incrassate at the base; stigmas 3, exserted from the utricle. Rachilla (see note) reduced to a rigid bristle below the nut and produced far beyond the mouth of the utricle, hooked at the top. Nut trigonous. Fig. 136. Range of the genus Uncinia Pers. (from VAN BALGooy, Pac. Pl. Areas 3, 1975, 320, map 209) 184 FLORA MALESIANA [ser. I, vol. 91 Distr. A genus of probably Antarctic origin, with wide southern distribution; from the extreme south of S. America including the Falkland Is., along the Andes to Mexico and Jamaica; islands in the southern parts of the Atlantic and of the Indian Ocean; from Tasmania through E. Australia northwards to Malesia, Mt Pulog in the Philippines being the most northern extension; highly developed in New Zealand and neighbouring islands, from there one species extending to Hawaii. Fig. 136. Note. Subdivision of the genus. KUKENTHAL divided Uncinia into two subgenera, Uncinia (‘Eu-Uncinia’) and Pseudocarex, the latter containing only U. kingii Bootr from Antarctic S. America, in which the rachilla is but shortly hooked. Subg. Uncinia is divided into two sections which are to some extent also geographically defined: sect. Uncinia, mainly Australian with few species in S. America, and sect. Platyan- dra which is exclusively American. KEY TO THE SPECIES 1. Glumes persistent. Stems sharply trigonous, scabrid beneath the inflorescence . . . 1. U. riparia 1. Glumes caducous, when young often an abscission layer visible. Stems obscurely trigonous, smooth 1. Uncinia riparia R. Br. Prod. (1810) 241; Boott in Hook. f. Fl. Tasmania 2 (1860) 102, t. 152 f. B; BENTH. FI. Austr. 7 (1878) 434; CLARKE, J. Linn. Soc. Bot. 20 (1883) 392, excl. var.; KUK. Pfi. R. Heft 38 (1909) 63, excl. var.; S. T. BLAKE, J. Arn. Arb. 35 (1954) 234; Noor. Blumea 24 (1979) 513. — Carex riparia (R. Br.) Por. in Lamk, Enc. Méth. Suppl. 3 (1813) 282. — U. sclero- phylla NELMES, Kew Bull. (1949) 143. — U. ohwiana KoyaMa, Bot. Mag. Tokyo 69 (1956) 214, f. 6. Rhizome more or less creeping. Stems approxi- mate on the rhizome, very slender, sharply trigo- nous, scabrid in the upper half, (10—)40-75 cm by 1/,-1 mm. Leaves from slightly shorter to slightly longer than the stems, rigid, flat or canaliculate, long attenuate, scabrous on margins and nerves in upper half, 1'/,-3(-?4) mm wide; basal sheaths bladeless or short-bladed, fuscous. Spikelet narrowly linear, often very loosely flowered, ebracteate or with a filiform bract usually not over- topping the inflorescence, 3—7(—15) by 2-5 mm, the 3 part few-flowered, 1—1!/,(-2) cm long. Glumes persistent, oblong-ovate, acute, muticous, rigid, stramineous with broad 3-nerved green centre and sometimes brown margins, (4—)5—6(—?8'/,) mm long, the midrib not reaching the apex. Utricles slightly exceeding the glumes, erect, linear-oblong or linear-lanceolate, compressed-trigonous, gla- brous and smooth, fine-nerved, stramineous, 6-7 by c. 1 mm, at the base subgradually narrowed into a c. 1'/, mm long stipe, at the apex into a c. 1'/, mm long, compressed-conical beak with narrow, hyaline mouth. Nut narrow ellipsoid. Distr. New Zealand, Tasmania, SE. Australia (Victoria, Upper Hume R. and Mt Kosciusko, according to BENTHAM, /.c.; no specimens seen); in Malesia: New Guinea (West Irian, summit of Mt Wilhelmina; Papua New Guinea). Ecol. In shaded places, usually between moss on the floor of the mossy or subalpine forest, 3000— 4100 m, above 4000 m also in grassland with shrubs. Fi. fr. Jan.—Dec. Note. Whether the glumes are persistent or caducous can often only be seen in old inflores- 2. U. compacta cences. It is not impossible that hybrids occur with U. compacta, thus giving more variability and plants in which the glumes fall very late. In these plants the sharply triquetrous culms become more or less rounded and smooth instead of scabrous (not found in Malesia). 2. Uncinia compacta R. Br. Prod. (1810) 241; Boott in Hook. f. Fl. Tasmania 2 (1860) 103; F.v.M. Fragm. 8 (1874) 152; BENTH. FI. Austr. 7 (1878) 434; CLARKE, J. Linn. Soc. Bot. 20 (1883) 395; CHEESEMAN, Man. New Zeal. FI. (1906) 800; KOK. Pfl. R. Heft 38 (1909) 65; CHEESEMAN, Man. New Zeal. Fl. ed. 2 (1925) 245; LourTEIc, Bull. Com. Nat. Fr. Rech. Antarct. (1968) 25; Noor. Blumea 24 (1979) 515. — Carex compacta Por. in Lamk, Enc. Méth. Suppl. 3 (1813) 282. — U. rupes- tris RAOUL, Ann. Sc. Nat. III, 2 (1844) 117; Bootr in Hook. f. Fl. Nov. Zei. 1 (1853) 286; CLARKE, J. Linn. Soc. Bot. 20 (1883) 392; Kix. Pfl. R. Heft 38 (1909) 64, incl. var. capillacea KUK.; HAMLIN, Dom. Mus. Bull. 19 (1959) 39. — U. filiformis Boott in Hook. f. Fl. Nov. Zel. 1 (1853) 286; HAMLIN, Dom. Mus. Bull. 19 (1959) 43. — U. ner- vosa Boott in Hook. f. Fl. Tasmania 2 (1860) 102. — U. riparia R. Br. var. stolonifera KUK. & STEEN. Bull. Jard. Bot. Btzg III, 13 (1934) 213. — U. riparia (non R. Br.) Ouwi, Bot. Mag. Tokyo 56 (1942) 213. — U. subtrigona NELMES, Kew Bull. (1949) 144. — Fig. 137a-d. Plant laxly to densely cespitose or with short rhizome and stems densely tufted, erect; sometimes stems decumbent, forming new tufts; culms slender, rarely more than 1 mm thick, obscurely trigonous, smooth, 5-45 cm. Leaves shorter or longer than stems, flat, involute, convolute or conduplicate, sometimes (var. nervosa) plano-convex and then often canaliculate, long attenuate, scabrous on margins and nerves at least in upper half, 1/,-3 mm wide, the tip mostly rather acute, triquetrous, rarely flat or plano-convex and blunt (var. nervosa); basal sheaths bladeless, brown. Spikelets nar- rowly oblong, loosely to very densely flowered, sometimes bracteate, (1-)11/,-5!/, cm by 2!/,- 1979 | Cy PERACEAE—II (Kern & Nooteboom) 185 Fig. 137. Uncinia compacta R. BR. var. nervosa CLARKE. a. Habit, nat. size, b. leaftip, x 7. — U. compacta R. Br. var. alpina Noor. c. Habit, nat. size, d. utricle, x 10. — Exocarya scleroides (F.v.M.) BENTH. e. Flower diagram (a-b ANU 7161, c-d ANU 7289). 186 FLORA MALESIANA [ser. I, vol. 9} 20 mm, the ¢ part 1/,-1'/, cm long. Glumes caducous, when young the abscission line often already visible, oblong-ovate or lanceolate, acute, muticous, at least the margins hyaline, greenish or brown, with 3-nerved central stripe but often with several more nerves, 4-6(—8 mm in the Antarctic Is.) by 1.6-3 mm. Utricles shorter to slightly longer than glumes, obliquely erect to patent, lanceolate elliptic or ovoid, with 2 conspicuous submarginal nerves, sometimes slenderly nerved towards the base or striate when these nerves are prolonged, 31/,-6(—7) by 1-2 mm, at the base contracted into a 1-1'/, mm long stipe, at the apex (gradually) narrowed into a 1-2 mm long beak. Nut ellipsoid. Distr. Australia (Victoria, New South Wales, Tasmania), New Zealand, Amsterdam I., Kergue- len, Crozet, Marion, Gough and Tristan da Cunha; in Malesia: Philippines (Luzon: Mts Pulog & Banahao; Mindanao: Mt Apo), N. Borneo (Mt Kinabalu), Celebes (Latimodjong Mts), New Guinea. Ecol. In the tropics in the high mountains, 2000-4300 m, outside the tropics in temperate and cold climates, in open places and in forest. Note. The second character in the key to the species differentiating U. compacta from U. riparia breaks partly down in some specimens from Vic- toria and New South Wales; these specimens have a sharply triquetrous and (interruptedly) scabrous stem. KEY TO THE VARIETIES 1. Spikelets c. 1 cm long. Leaves strongly circin- nate towards the apex. Usually small plants c. var. alpina 1. Spikelets 11/,-51/, cm. Leaves not or less circin- nate. Plants often larger. 2. Leaves flat (always in New Guinea), involute, convolute or conduplicate with acute, tri- gonous or triquetrous tip. . a. var. compacta 2. Leaves plano-convex with flat, or plano- convex blunt tip. b. var. nervosa a. var. compacta. Spikelets 11/,-5'/, cm. Leaves flat, involute, convolute or conduplicate. Leaf tip acute, trigo- nous or triquetrous, undulate or (somewhat) circinnate. Distr. As the species. Ecol. In forest, 2000-3700 m. Fi. fr. Jan._Dec. b. var. neryosa CLARKE, J. Linn. Soc. Bot. 20 (1883) 395. — U. nervosa Bootr in Hook. f. Fl. Tasmania 2 (1860) 102; HAMLIN, Dom. Mus. Bull. 19 (1959) 50. — Fig. 137a-b. Spikelets 1'/,-3 cm. Leaves plano-convex, usually canaliculate, sometimes some of the leaves convolute. Leaf tip flat, or plano-convex, blunt. Distr. Australia (New South Wales, Kosciusko area and Tasmania), New Zealand; in Malesia: New Guinea. Ecol. In open places, mostly in grassland, 3000-4026 m. FI. fr. June—Nov. c. var. alpina Noor. Blumea 24 (1979) 519. — Fig. 137c—d. Spikelets c. 1 cm long. 2 Flowers c. 5; utricles 3!/,4 mm. Leaves involute, strongly circinnate towards the triquetrous or trigonous apex. Distr. Malesia: New Guinea (Mt Wilhelm and Mt Giluwe). Ecol. In exposed places, often temporarily covered by snow, also on solifiuction terraces, 3770-4350 m. Fi. fr. Nov.—April. ADDENDA 7: 452 Add to A. SUBFAMILY CyPEROIDEAE I. Tribe Hypolytreae after 6. Paramapania: 6a. Exocarya. Change in the SYNOPTICAL KEY TO THE MALESIAN GENERA fork 6, second lead, into: 6. Hypogynous scales 2 or 4. 6; ciyporynousiscales, 275. oa econte ck 6’. Hypogynous scales4. . . . . 7: 454 Change in the KEY TO THE GENERA fork 19, first lead, into: 19. Hypogynous scales 2 or 4. Stigmas 2. Inflorescence paniculate. 19’. Hypogynous scales 2, either of them with a stamen in its axil . 19’. Hypogynous scales 4, stamens 3 6a. EXOCARYA BENTH. in Hook. Ic. Pl. 3 (1877) pl. 1206. — Fig. 137e. Leaves situated throughout the stem. Inflorescence an umbellate panicle, the lower 2-3 bracts long, foliaceous. Spikelets small, the upper | or 2 flowers bisexual, the other 2-3 flowers male. Glumes imbricate, the lower ones often empty. Flowers 1979] Cy PERACEAE—II (Kern & Nooteboom) 187 compressed, the 2 outer hypogynous scales subopposite, folded, transverse and sharply keeled, the 2 inner ones flat or concave, parallel with the glumes. Stamens 3. Style broadened towards the base, with 2 filiform stigmas. Nut exserted from the glumes, crowned by the persistent style-base. Distr. Monotypic. Australia (Queensland, New South Wales); in Malesia: E. New Guinea (Ferguson I.). 1. Exocarya scleroides (F.v.M.) BENTH. in Hook. Ic. Pl. 3 (1877) pl. 1206; F. M. BatLey, Queensl. Fl. 6 (1902) 1777; S. T. BLaKe, Proc. R. Soc. Queensl. 54 (1943) 72. — Cladium scleroides F.v.M. Fragm. 9 (1875) 12. — Scleria ustulata F. M. Battey, 3rd Suppl. Syn. Queensl. Fl. (1890) 81. — E. montivaga Domi, Bibl. Bot. XX, Heft 85 (1915) 484. — Fig. 137e. Stems from a creeping rhizome, to c. 1 m high, foliaceous, 3-angled. Leaves flat, to 6!/, mm broad, tapering into long subulate points; sheaths close, with minutely fimbriate mouth. /nflorescence an umbellate panicle, varying in size, but often very large, to 20 cm @, the longest rays up to 15 cm, pedicels filiform. Involucral bracts several, folia- ceous, the longest scarcely as long as the inflores- cence. Spikelets dark brown, narrowly obovate, c. 4 mm long. Lower glumes empty, outer ones very short, c. 1 mm, gradually passing into the c. 2 mm long flowering ones. Hypogynous scales as long as the glumes. Nut much exserted, ovoid- oblong (or globose: ‘E. montivaga’), c. 4 mm long, smooth, the remains of the spikelet forming a small tuft at its base. Style-base black, larger than the ovary at the time of flowering but not enlarged afterwards. Distr. E. Australia (New South Wales, Queens- land); in Malesia: Papua New Guinea (once collected in Ferguson I.). Ecol. In forest, 720 m. Fi. fr. Nov. Although sometimes rather abundant, its occurrence in Australia is very sporadic over its rather extensive geographical range. It seems to be usually a constituent of some of the less densely closed forests. A notable feature is that only a small propoftion of the spikelets produce mature nuts (BLAKE, /.c.). Note. This plant belongs to the tribe Hypoly- treae and is obviously intermediate between Paramapania and Hypolytrum, differing from the former in the presence of only 2 interior, flat hypogynous scales, while Hypolytrum lacks those scales. The stomata are tetracytic, as in Lepironia and Scirpodendron. Excluded Eriophorum comosum (WALL. in Roxb.) NEEs in Wight, Contr. (1837) 110; Mia. Fl. Ind. Bat. 3 (1856) 330; STEEN. Bull. Jard. Bot. Btzg III, 13 (1933) 200. MIQUEL cited this to occur in Malaya (Penang I.); on what grounds is uncertain. It must be a mystifica- tion or mislocalisation of specimens. CLARKE (1893) did not mention this locality in Fl. Br. Ind. In Herbarium Bogoriense VAN STEENIS, /.c., found specimens of E. comosum, said to have been collected in Karimata I. (off W. Borneo), mixed with specimens of Machaerina rubiginosa. This was interpreted as an unintentional mixture; E. comosum certainly does not occur in the low Karimata I. Eriophorum filamentosum Boeck. Bot. Jahrb. 5 (1884) 506 was based on a GRIFFITH collection credited to have been collected in Malaya. CLARKE (FI. Br. Ind. 6, 1893, 664) reduced this to Xerotes leucocephala R. Br., of which Xerotes filamentosa A. CUNN. msc. is a synonym. This is not a Cyperaceous plant, but belongs to Lomandra of the Liliaceae sens. lat., a genus which occurs outside Australia only in New Guinea. It is hard to believe that BOECKELER made such an error. This may be a slip of the pen by CLARKE. The identity of BOECKELER’s type should still be checked; but no Eriophorum has ever been found in Malaya. LILIACEAE—I (J. P. Jessop, Adelaide)! Herbs, usually glabrous, with perennial underground stems (corms, bulbs, tubers, or rhizomes) in all Mal. spp. Aerial stems usually herbaceous and annual, erect or climbing. Leaves simple, caespitose and basal, sometimes distichous, if cauline usually alternate, generally linear to lanceolate or oblanceolate especially when basal, but sometimes shorter and broader (to ovate) when cauline, usually sessile (in Asparagus and Petrosavia reduced to non-photosynthetic scales), usually with parallel venation. Stipules 0. Jnflorescence terminal or axillary, usually racemose (less often at least partly umbellate) or flowers solitary, usually bracteate. Flowers bisexual (except, in Mal., Asparagus cochinchinensis and Astelia alpina), usually actinomorphic. Perianth segments almost invariably 6 in two more or less similar or less often distinctly dissimilar whorls of 3, petaloid, connate or free, the outer whorl sometimes saccate at the base. Stamens 6, inserted on receptacle or perianth; filaments connate or free, rarely forming a corona-like ring attached to the perianth; anthers basifixed or dorsifixed, rarely sessile, usually 2-celled, extrorse to introrse or rarely dehiscing by an apical pore. Ovary usually superior, of 3 (usually fused) carpels; styles 1 or 3, simple or 3-branched; locules usually 3 (1 in Trica- listra); ovules 1 to numerous, placentation axile, rarely basal or parietal, usually in 2 rows. Fruit usually a loculicidal or septicidal capsule or berry, rarely the ovary wall ruptured by the developing seed which develops unprotected by a fruit, perianth caducous or persistent. Seeds with copious fleshy or cartilaginous endo- sperm. Distribution. About 180 genera with approximately 3500 spp., distributed all over the world, especially in the temperate regions of Asia, Australasia and Africa, but relatively poorly repre- sented in South America (13 genera). In Malesia 22 genera, with a total of 31 spp., no genus being represented by more than two species. The only genus endemic to the region is the Malayan genus Tricalistra whose separation from Tupistra is, however, somewhat uncertain. Most other genera are represented in Malesia by a minority of their species, exceptions being Gloriosa and Peliosanthes, which are probably both monotypic, and Petrosavia, which consists probably of two species. The genera can roughly be arranged into three geographical groups. Old World genera are Asparagus, Chlorophytum, Dianella, Gloriosa and Iphigenia, among which Chlorophytum is mainly Africa-centred, and Dianella mostly Australasian. Northern hemisphere genera, especially from the Far East, Sino-Himalayan, are the following: Aletris, Disporopsis*, Disporum, Lilium*, Liriope*, Ophiopogon, Peliosanthes, Petrosavia, Tricyrtis*, and Tupistra. Of these four, provided with an asterisk, are found only in Malesia in the Philippines, and Tupistra only in Malaya and Sumatra. All of them are absent from East Malesia. Most of their species occur in the montane zone, testimony of their subtemperate ecology. Australasia-derived genera are Arthropodium, Astelia, Caesia, Schelhammera, Thysanotus, and Tricoryne. Their occurrence in Malesia is confined to New Guinea, except for Thysanotus chinensis BtH. which is found through Malesia as far as Thailand and S. China. Their ranges are sometimes wider in Austral regions, as Arthropodium and Caesia occur also in the Malagasy area, and Caesia also in South Africa, while Astelia ranges widely from Mauritius to the southern Pacific islands and the Falkland Islands. Except fer Aste/ia, which occurs in Malesia only in the (1) Gratefully acknowledging a stipend from C.S.I.R., Pretoria, for work on this revision during the period July 1973 to June 1974, at the Rijksherbarium, Leyden, when on long-leave from Rhodes Univer- sity, Grahamstown, South Africa. With bibliographic co-operation of the General Editor. (189) 190 FLORA MALESIANA [ser. I, vol. 9! alpine zone of New Guinea, all the species of the genera of this southern group are bound to lowland drought habitats. Ecology. Of the 22 genera 13 are integrated in lowland to montane everwet-forest conditions. Astelia alpina is a high-altitude cushion plant which sometimes plays a significant role in the alpine bogs of New Guinea. The eight other genera, Arthropodium, Asparagus, Caesia, Gloriosa, Iphigenia, Schelhammera, Thysanotus, and Tricoryne, are constituents of areas subject to a seasonal climate. Consequently, the ranges of Asparagus racemosus, Gloriosa superba, Iphigenia indica and Thysanotus chinensis show in Malesia the usual disjunctions of drought-preferring plant species. They are predominantly grassland or open savannah plants at low altitudes. Most of them are Australasia-centred. Except for Astelia papuana, species of these Liliaceae sensu stricto do not form major con- stituents or natural communities. Dispersal. The great majority of Liliaceae spread and reproduce vegetatively by the branching of their subterranean axes. In most species this appears to be a slow process, with the branches often not extending more than a few centimetres in a year. It may, however, result in fairly dense monospecific stands, for example, in Astelia and Liriope. Fruits are generally capsules or berries. In the former, dispersal mechanisms do not usually result in the removal of seeds to any great distance, although wind and water can contribute significantly. Birds are probably the most efficient vectors over longer distances. Several genera have fleshy fruits and two (Ophiopogon and Peliosanthes) have a fleshy coating to the seeds, which are exposed through rupture of the ovary wall. Mammals may also disperse the seeds by eating the fruits. Liliaceae seeds in other areas are known to be carried by ants if there is a substance attractive to ants (often oil bodies) in the testa or fruit. Specific data on the Malesian species have not, however, been found. For a study of the structure and relationships of the seeds, see HuBER, Die Samenmerkmale und Verwandtschaftsverhaltnisse der Liliifloren, Mitt. Bot. Staatssamml. Miinch. 8 (1969) 219-538. Cytology. Liliaceae, because of the usually large size of their chromosomes, because of the ease with which material can often be obtained at the stages of division required for study, and because many species are in cultivation, have been fairly well studied cytologically. Chromosome numbers, and even basic chromosome numbers vary widely sometimes within, as well as between, genera. At least six different somatic numbers have, for example, been reported for Disporum, based on x = 6, 7, 8, 9 and 11. Other genera, for example, Asparagus (x = 10) and Dianella (x = 8), have relatively stable basic numbers, although polyploidy may be common. Taxonomy. The family Liliaceae, in the sense of BENTHAM & HooKER and of KRAUSE in E. & P. Nat. Pfl. Fam. is a very large and rather heterogeneous one including possibly as many as 3500 species. Many more recent authors have attempted to distribute these species over a larger number of families. In this treatment the family delimitation of HUTCHINSON (Families of flower- ing plants, ed. 3, 1973) has been adopted, with two modifications: the inclusion of Petrosavia which HuTCHINSON placed in its own family, and of the naturalised Nothoscordum which HUTCHINSON placed in the Amaryllidaceae. It is very doubtful if the family is more naturally defined by excluding a number of genera represented in Malesia, as HUTCHINSON has done, but this has been followed here as much for the convenience of dealing with smaller families as for any conviction that these families have any botanical significance. Table 1 indicates the genera retained in the Liliaceae and the families to which other genera, sometimes included in the Liliaceae, were ascribed by HUTCHINSON. These families have also been included in the key to the genera of Malesian Liliaceae. Much work remains to be done on the relationships of Liliaceous genera and HUTCHINSON’S work, although the most recent, is probably no better than KRAUSE’s. HUTCHINSON’S placing of Ophiopogon and Peliosanthes in separate tribes is, for example, almost certainly unjustified. Uses. Several species of Liliaceae native to Malesia have been taken into cultivation as garden ornamentals, for example of the genera Dianella, Gloriosa, Lilium, Liriope, and Ophio- pogon. Other uses are, however, rather few. Several genera, for example, Asparagus, Gloriosa, Ophiopogon, have been used in traditional medicines but they have not contributed to modern 1979] LILIACEAE—I (Jessop) 191 Table 1. Malesian Liliaceous genera in the classification by KRAUusE (1930), first column, and by HUTCHIN- SON (1973), third column. Subfamily I. Melanthioideae Is 2. Petrosavieae Petrosavia (Petrosaviaceae) le 6. Uvularieae Schelhammera . Glavioxa Uvularieae it: 7. Tricyrteae Tricyrtis Tricyrtideae il. 8. Anguillarieae Iphigenia Iphigenieae Subfamily III. | Asphodeloideae IIl. 11. Asphodeleae Chlorophytum Thysanotus Asphodeleae Ill. lla. Asphodelinae Arthropodium Asphodeleae Tricoryne Johnsonieae Caesia Asphodeleae Ill. 11g. Dianellinae a } Dieiehe III. 17. Lomandreae Lomandra Rommnalaa (Xanthorrhoeaceae) Subfamily IV. Allioideae Nothoscordum (Amaryllidaceae) (introduced) Subfamily V. Lilioideae Lilium Tulipeae Subfamily VII. Dracaenoideae VII. 27. Dracaeneae Cordyline Dracaena (Agavaceae) Astelia Milliganieae Subfamily VIII. Asparagoideae VI. 28. Asparageae Asparagus Asparageae VIII. 29. Polygonatae seals } Peligonaine isporopsis VIII. 30. Convallarieae Tupistra Asiairene Tricalistra Subfamily IX. | Mondoideae Liriope SF eee Ophiopogon PRESS Peliosanthes Peliosantheae Subfamily X. Aletroideae Aletris Narthecieae Subfamily XI. Luzuriagoideae Luzuriaga Geitonoplesium (Philesiaceae) Eustrephus Subfamily XII. Smilacoideae Smilax Rhipogonum Heterosmilax (Smilacaceae) 192 FLORA MALESIANA [ser. I, vol. 9! medicine. Others have edible underground parts (e.g. Arthropodium) or fruits (e.g. Astelia), but none is probably of great significance. Notes. A very large number of exotic Liliaceae are recorded to be or have been cultivated in gardens in Malesia. They have been treated elaborately, with keys for their identification by C. A. BACKER in his ‘Handboek voor de Flora van Java’, part 3 (1924), in Dutch, and by C. A. BACKER & R. C. BAKHUIZEN VAN DEN BRINK Jr in their ‘Flora of Java’, volume 3 (1968), in English. Only one exotic, Nothoscordum inodorum, has been introduced and has run wild in West Java; this has been incorporated in the treatment. Almost all drawings were made by Mr. L. DuTKtEwicz, Adelaide. KEY TO THE GENERA including the families sometimes segregated from Liliaceae sensu lato 2 Leaves reduced to small, non-photosynthetic scales. . Stem! branched: bearing green'cladodes 475-7 4 = = = =) Se OR ASDArACUS 2. Stem simple; Small echlorophyllous'saprophyte” =. > = = 2 5) 4 7) eee 4. Petrosavia Leaves usually well-developed, always green (sometimes absent when the plant i is in flower). a Anthers 1-celled. Plants pag woody and often prickly climbers (Heterosmilax, Rhipogonum, Smilax): 1.008.569 2 ERS Saw Me) ee: SO Smilacaceae 3. Anthers 2- celled. Plants erect or - herbaceous, not prickly, rarely woody climbers. 4. Plants dioecious. 5. Panicle glabrous or scaberulous. Flowers clustered at all or most nodes (Lomandra) Xanthorrhoeaceae 5. Panicle silvery-scaly. Flowers solitary ateachnode ........ 1 ach eu CEC eA Stella 4. Flowers bisexual. 6. Foliage leaves caespitose, usually basal or radical. 7., Inner perianth'segments'fringed . . 9. 2 ee <3). 2 2. ee ee Oe ySanotus 7. Perianth segments not fringed. 8. Inflorescence a simple umbel. Bulbous . . . .... . . 23. Nothoscordum 8. Inflorescence a raceme, spike or panicle. Plant rhizomatous. 9. Flowers sessile. Anthers sessile or subsessile on the perianth. 10:. Fruita capsules ssc. seep ich apedise 7 e e ke ees ry EIS 10. Fruit fleshy. 11. Style distinct. Stigma simple or indistinctly lobed. . . ..... =... . 17. Tupistra 11. Style absent. Stigmas distinctly3 . 5b) oo, Ghee, Sune Gomi ricalistral 9. Flowers pedicelled. Filaments usually well- ‘developed. 12. Seeds fleshy and exposed soon after the commencement of their development. 13: Filaments free of one’ another; Corona absent: = = =). eee 18. Liriope 13. Filaments connate or anthers sessile on a staminal corona. 14. Anthers borne on a distinct staminal corona. . ....... ... . 20. Peliosanthes 14. Anthers borne on connate filaments ............. .. 19. Ophiopogon 12. Seeds retained in the fruit until mature. 15. Anthers dorsifixed. 16. Ovary half-inferior . . 3 ee ew ts os tee one gu oe eS 16. Ovary fully superior. 17. Fruit a capsule. Stems to 5 cm long and 2-3 mm diameter (Romnalda) . Xanthorrhoeaceae 17. Fruit a berry. Stems woody and usually long (Dracaena, incl. Pleomele) . . Agavaceae 15. Anthers basifixed. 7 18: Fruita berry . 5 20 ee a 2 ence 18. Fruit a capsule. 19. Base of anthers with a papillose appendage. Pedicels 12-20 mm. Outer perianth segments distinctly broader than inner ones . . . . . 6. Arthropodium 19. Base of anthers without appendages. Pedicels 3- 12 mm. Perianth segments equal. 20. Perianth spirally twisting after anthesis, blue ........... » one caesia 20. Perianth segments persistent, but not twisted after flowering, white to green 8. Chlorophytum 6. Foliage leaves distributed at intervals along the stem. 21. Plant with scaly bulb. Flowers 10-25cmlong ............... . 12. Lilium 1979] LILIACEAE—I (Jessop) 193 21. Plant with a rhizome, corm or tuber. Flowers up to at most 9 cm. 22. Stem(s) woody. Venation reticulate (Eustrephus, Geitonoplesium). . . . . . . Philesiaceae 22. Stem(s) herbaceous. Venation parallel. 23. Style simple, or if 3 lobed the filaments forming a corona. 2Aeruita berry, Inflorescenceia panicle ... 2... wi. i =: - . . . 9. Dianella 24. Fruit consisting of 1-3 nutlets. Inflorescence consisting of umbels each ¢ on a winged peduncle 11. Tricoryne 23. Style branched, sometimes to the base. 25. Flowers large, 5-9 cm. At least some leaves ending in a coiled tendril . . . 1. Gloriosa 25. Flowers smaller, up to 3 cm. Leaves never ending in a tendril. 26. Filaments expanded to form acorona. Fruita berry ..... . . . . 15. Disporopsis 26. Filaments free, sometimes connivent. DAL EMUI tia) DELLY 2 0h eae) even oe ns: ee eee, ces, et ee nl Soe DIS DOFUIN 27. Fruit a capsule. 28. Pedicels not articulated. Anthers dorsifixed. 29. Rootstock a rhizome. Style branches bifid .............. 2 Tricyrtis 25 mRootstocksa comm- style branchesstmple” 2 2) 39)... . 24. = = 3. Iphigenia oRmPedicelsaruculated) Anthers) basifixed, <2. 2 9-9. . 3 =. - = - 5. Schelhammera 1.GLORIOSA LinnE, Sp. Pl. (1753) 305; Gen. Pl. ed. 5 (1754) 144; Baker, J. Linn. Soc. Bot. 17 (1879) 457; Krause in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 266; HutcH. Fam. FI. Pl. ed. 2, 2 (1959) 606; FiELD, Kew Bull. 25 (1971) 243; Lilies and other Liliaceae 1973 (1972) 93. — Methonica TourRN. ex CRANTZ, Inst. rei herb. 1 (1766) 474. — Fig. 1, 2. Climbing, or less often erect, glabrous herbs. Rhizome perennial, tuberous, horizontal; roots fibrous. Aerial stem annual, moderately branched; the branches spreading or erect-spreading at the base. Leaves cauline, alternate, opposite or in whorls of 3 (4), flat, with many veins and a strong midrib, sessile, entire, lanceolate to ovate, slightly amplexicaul, obtuse at the base, narrowing gradually to an apical, coiled tendril (sometimes lacking tendrils in erect forms); basal leaves lacking a lamina or tendril, ensheathing the stem. Pedicels solitary, axillary in the axils of vegetative leaves, not articulated, cernuous. Perianth segments connate at the base, many-veined, subequal, reflexed or rarely spreading with a longitudinal papillose furrow in the basal + third of the adaxial surface. Filaments filiform, attached to the receptacle; anthers dorsifixed, linear-oblong, extrorse. Ovary superior, sessile, oblong-ovoid to oblong-obovoid, 3-celled, 1 cell slightly shorter than the other 2; ovules axile, numerous; style filiform with 3 stigmatic branches, reflexed or spread- ing from the attachment to the ovary. Capsule coriaceous, septicidal ; seeds globose; perianth persistent but withering as the capsule enlarges. Distr. Widespread in tropical and southern Africa, Madagascar, India, Burma and SE. Asia, as far as West Malesia. Ecol. Usually climbing in bushes, in habitats ranging from savanna to forest. Note. Stated by FreLp (1971, 1972) to be monotypic. In Malesia only a single indigenous species has ever been recognized, although other species have been described elsewhere. 1. Gloriosa superba Linné, Sp. Pl. (1753) 305; Nutt. Pl. (1927) 436; Back. Onkr. Suiker. (1928) Baker, Fl. Cap. 6 (1897) 525; BACK. Trop. Natuur 189, Atlas (1933) t. 201; GAGNep. Fl. Gén. I.-C. 6 3 (1914) 117, tab. col.; Ri. Fl. Mal. Pen. 4 (1924) (1934) 807; H. Perrier, Fl. Madag. fam. 40 (1938) 338; BACK. Handb. Fl. Java 3 (1924) 50; PercH, 135; Suva, Ceyl. J. Sc. sect. A, 12 (1945) 155; Ann. R. Bot. Gard. Perad. 9 (1925) 243; HEYNE, SANTAPAU, J. Bomb. Nat. Hist. Soc. 46 (1946) 202; 194 FLORA MALESIANA [ser. I, vol. 91 Fig. 1. Gloriosa superba L. Botanic Garden Singapore, Febr. 1952 (Photogr. M. R. HENDERSON). 1979] HEND. Mal. Gard. Pl. 4 (1951) 10; Mal. Wild Flow. Monoc. (1954) 178, f. 104; Hottr. MAHA Mag. 15 (1958) 75 (hybrids); BACK. & BAKH. f. FI. Java 3 (1968) 85; HutcH. & Datz. Fl. W. Trop. Afr. ed. 2, 3 (1968) 106; FieLp, Kew Bull. 25 (1971) 243. — Methonica superba (L.) CRANTZ, Inst. rei herb. 1 (1766) 474; ZoLv. Syst. Verz. 1 (1854) 66; Nat. Tijd. N. I. 14 (1857) 149; Mia. Fl. Ind. Bat. 3 (1859) 550. — G. virescens LINDL. Bot. Mag. (1825) t. 2539. — Fig. 1, 2. Stems usually climbing to c. 2 m (rarely to 6 m), less often erect; green. Leaves (including the tendril) 8-17!/,(-25) by (14/4-)1'/.4(4'/.) cm; tendril usually less than 1 cm long. Pedicels 4'/,-19 cm. Perianth segments narrowly elliptic, with undulate or crisped margins, 5—7(-9) by 3/,-1'/,(-3) cm broad, yellow or red, often (perhaps always in Asia) yellow or green towards the base at first but becoming red throughout later. Filaments spread- ing, 2'/.-5 cm long; anthers 7-10 mm long. Ovary 8-15 mm long; style including the filiform, 3-7 (-12) mm long style branches 31/,-5!/, cm long. Capsule 4-10 by c. 1'/,-2 cm. Seeds vivid-red or orange-red, with a fleshy testa, c. 5 mm @, tardily falling. Distr. Tropical and southern Africa, Madagas- car, India to Indo-China; in Malesia: Java (also Madura and Kangean Is.), S. Celebes, and all Lesser Sunda Is. Ecol. Brushwood, hedges, teak-forest, only in regions subject to a strong dry season, from near the beach and dunes up to c. 300 m altitude (very rarely 600 m), locally common. It is not native in Sumatra and Borneo, and probably not in con- tinental Malaya. This disjunction in its range between continental SE. Asia and Central South Malesia is clearly caused by its drought preferring ecology. Fi. fr. Jan.—Dec. Uses. Commonly grown asa garden ornamental. The tuber is said to be poisonous (through colchicin), but only slightly so as was tested by BoorsMA (BACKER, 1914; HEYNE, 1927). Fig. 2. Vern. Klimlelie, D, flame lily, superb lily, E; Java: kembang djonggrang, k. kuku matjan, k. sung- LILIACEAE—I (Jessop) 195 sang, M, dongkel sungsang, mandalika, pa(n)tjing towo, J, katongkat, S, mand(h)alika, Md, Balin.; Lesser Sunda Is.: enatha, sikal, Dawan lang., Timor. Note. In sterile state easily distinguished from another climbing monocot with coiled apical leaf- tendrils, Flagellaria indica L., by absence of a leaf-sheath. Fig. 2. Gloriosa superba L. Old rhizome with scar, the apex with a new tuber emitting roots and a vertical shoot. The two triangular elongations of the new tuber will grow later into new rhizomes; x 1/,. Botanic Garden, Bandung, 1952. Dug up by L. VAN DER PIL. 2. TRICYRTIS WALL. Tent. FI. Nap. (1826) 61, t. 46; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, I5a (1930) 269; HutcH. Fam. FI. Pl. ed. 2, 2 (1959) 608, nom. gen. cons. — Compsoa D. Don, Prod. FI. Nep. (1825) 50. — Fig. 3. Erect puberulous or glabrescent herbs. Rhizome short, creeping. Aerial stem annual, simple or branched. Leaves cauline, alternate, flat, with several veins and a strong midrib, sessile, entire, lanceolate to ovate, with a sheathing base. Jnflores- cence terminal, racemose, simple or branched, or flowers in the axil of vegetative leaves. Pedicels solitary, not articulated. Perianth segments free or very shortly united, equal or subequal but the outer three saccate at the base, erect to spreading. 196 FLORA MALESIANA [ser. I, vol. 9} Fig. 3. Tricyrtis imeldae GUTIERREZ. a. Habit, x 1/4, b. flower, nat. size, c. outer perianth segment, d. inner perianth segment, e. gynoecium, all x 1'/,, f. fruit, nat. size, g. seed, x 12 (Redrawn from GUTIER- REZ, Philip. J. Sc. 103, 1974, 3, fig. 1). LILIACEAE—I (Jessop) 197 Filaments flattened, more or less connivent, free of the perianth; anthers dorsifixed, versatile, oblong, extrorse. Ovary superior, sessile, oblong, 3-celled, 3-angled; ovules axile, numerous; style columnar; with 3 spreading or recurved bifid branches. Capsule septicidal. Seeds oblong or ovoid. Distr. Possibly c. 20 spp., largely in Japan, also in Manchuria, Korea, throughout China to the Hima- layas, Taiwan, and North Malesia: Philippines. Notes. HUTCHINSON placed this genus in the tribe Tricyrtideae possibly with a South African genus Sandersonia as the only other member. KRAUSE placed it in the Tricyrteae with the closely related, possibly congeneric, Brachycyrtis. The genus appears to be taxonomically rather isolated, but probably closest to Gloriosa of the Malesian genera. A thorough revision of the species is badly needed. 1. Tricyrtis imeldae GUTIERREZ, Philip. J. Sc. 103 (1974) 171, f. 1. — Fig. 3. Stems erect to 70 cm high, unbranched, puberu- lous at first. Leaves thick and fleshy when fresh but becoming membranous when dried, acute, the lower narrowly elliptic-lanceolate, with cuneate base, the upper broadly lanceolate to elliptic, with cordate base, (6—-)12-16 by (3—-)4-5 cm, glabrous except for the main veins beneath. Jnflorescence a terminal bifurcate raceme c. 18-20 cm, puberulous; pedicels 3-5 mm, puberulous. Flowers greenish- white with purple spots inside, to over 3 cm long, infundibuliform, glabrous, segments linear-oblong to oblong spathulate. Filaments 16-18 mm long; anthers 3 mm long, yellowish-brown. Ovary 10 mm long; style 8 mm, its branches 8 mm long, purple, spreading, tuberculate on the inner surface. Fruit c. 25-30 by 4-6 mm. Seeds flat, oblong, c. 2 mm long. Distr. Malesia: S. Philippines (Mindanao: Tasaday, Cotabato), reported to be rather rare; only known from the type. Ecol. Primary forest, along stream at c. 1300 m. Fl. Aug. Vern. Philippines: amutmagiso, Tasaday. Notes. I have only seen the type and have not been able to add to the description by GUTIERREZ. He compared the species with the Formosan T. stolonifera, from which it chiefly differs in the shape and colouring of the perianth segments. Close study of more material and variability is needed to check the specific difference. 3. IPHIGENIA KuNTH, En. Pl. 4 (1843) 213; BAKER, J. Linn. Soc. Bot. 17 (1880) 450; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 272; H.Perrier, Fl. Madag. fam. 40 (1938) 136; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 612; OBERMEYER, Kirkia | (1961) 84, nom. gen. cons. — Aphoma Rar. FI. Tellur. 2 (1836) 31, nom. rejic. — Fig. 4. Erect, glabrous herbs. Corm enclosed in dry leaf bases; roots fibrous. Aerial stem annual, unbranched. Leaves cauline, with tubular, ensheathing bases; the lowest | or 2 often with a poorly developed lamina; the others decreasing in size from the lowest towards the uppermost, the uppermost often approaching the lowest bracts in size, c. 4-7; lamina flat, glabrous, sessile, entire, linear to lanceolate, acute. Flowers solitary, in a few-flowered cluster or in a raceme terminating the stem. Pedicels solitary, erect-spreading, not articulated. Perianth segments free, + equal, spreading or reflexed, few-veined. Filaments flattened, attached to the receptacle; anthers dorsifixed or basifixed, extrorse. Ovary superior, sessile, ovoid, oblong or obovoid, 3-celled; ovules ov, axile; styles 3, (free or) fused at the base only. Capsule globose, cylindrical or ellipsoid, loculicidal. Seeds globose or angular; perianth deciduous. Distr. Southern and tropical Africa (5 spp.), Madagascar (2 spp.), SE. Asia (4 spp.), of which one species extends through Malesia to Australia, and New Zealand. Ecol. Usually in open grassland, sometimes in damp areas. 198 FLORA MALESIANA [ser. I, vol. 91 1. Iphigenia indica (L.) A. GRAY ex KUNTH, En. Pl. 4 (1843) 213; Mia. Fl. Ind. Bat. 3 (1859) 552; Bru. Fl. Austr. 7 (1878) 31; BAKER, J. Linn. Soc. Bot. 17 (1880) 450; E.v.M. Descr. Not. 6 (1885) 18; Hook. f. Fl. Br. Ind. 6 (1892) 357; BAILEY, Queensl. Fl. 5 (1902) 1641; Laut. Bot. Jahrb. 50 (1913) 292; Merr. En. Philip. 1 (1922) 202; Back. Handb. FI. Java 3 (1924) 51; Krause, Bot. Jahrb. 59 (1925) 548; Back. & Baku. f. Fl. Java 3 (1968) 85. — Melanthium indicum LinnE, Mant. 2 (1771) 226. — Anguillaria indica (L.) R. Br. Prod. (1810) 273; WALL. PI. As. Rar. 3 (1832) 37, t. 259. — Fig. 4. Corm usually + globose, 5-10 mm @. Leaves linear-lanceolate, often with a single conspicuous vein and 2-8 rather inconspicuous ones, the longest c. 10-40 cm long, up to 6 mm wide, the shortest often less than a quarter of the length of the longest; lower leaf bases sometimes pubescent, glandular pubescent or scabrid, especially on the veins. Flowers 1-3. Pedicels erect or erect-spreading 5-40 mm. Perianth segments + _ spreading, narrowly oblanceolate (inner whorl sometimes narrower than the outer), 5-9 by up to 2 mm, dark- brown or red-brown, reddish, purplish or white, described by BACKER (1924) as having a green keel and apex. Filaments linear, up to half as long as the perianth, green basally, brown distally; anthers dorsifixed, c. '/.-1 mm long. Ovary obovoid to ovoid, c. 11/,-21/, mm long; styles recurved, c. 1 mm. Capsule c. 10-20 mm long; seeds c. 1'/, mm @. Distr. Ceylon, India to Thailand, S. China, in Malesia rare: N. Sumatra (Lake Toba), Java (Indramayu in W, Surabaja and Madura I. in E), the Lesser Sunda Is. (Timor and Wetar), Philip- pines (Luzon, Mindanao), New Guinea, and Australia (W. Australia, Northern Territory and Queensland). Ecol. A rather uncommon species of open, often poor grassland, always under seasonal climatic conditions. In Java only in the lowland but else- where also in the hills, in N. Sumatra at c. 1000 m. Flowers are reported by BACKER (1924) to occur for a short period during the rainy period (Dec.— Jan.) in Java, with fruit ripening in March, after which the aerial parts soon wither and disappear. Elsewhere other flowering dates have been noted from July to August, N of the equator, where seasons are reversed. Fig. 4. Iphigenia indica (L.) A. GRAY ex KUNTH. a. Habit, nat. size, b. flower, c. fruit, both x 2 (Partly after WALLICH, PI. As. Rar. 3, 1832, t. 259). 4. PETROSAVIA Becc. Nuov. Giorn. Bot. Ital. 3 (1871) 7, t. 1; Rip. J. Str. Br. R. As. Soc. n. 24 (1891) 170; Krause in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 256; HuTcH. Kew Bull. (1933) 156; STEEN. Trop. Natuur 23 (1934) 52; NAKAI, J. Jap. Bot. 17 (1941) 191; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 546. — Protolirion RipL. Ann. Bot. 9 (1895) 45; Groom, /.c.; RIDL. Fl. Mal. Pen. 4 (1924) 322; KRAUSE in E. & P. Nat. 1979] LILIACEAE—I (Jessop) 199 Pfl. Fam. ed. 2, 15a (1930) 257, f. 87; NAKAI, J. Jap. Bot. 17 (1941) 191. — Miyoshia MAKINO, Bot. Mag. Tokyo 17 (1903) 144; NAKAI, J. Jap. Bot. 17 (1941) 191. — Fig. 5. Erect, glabrous, saprophytic, pale yellow or cream coloured, herbs lacking chlorophyll. Rhizome slender, simply or sparsely branched, bearing alternate, often imbricate scale-leaves. Aerial stems | or less often up to 3 or more, un- branched. Leaves scale-like, spiral, sessile, entire; the base usually partly embracing the stem, rather well-spaced. Raceme terminal, usually simple, sometimes corym- bose. Pedicels solitary, with 0-2 alternate bracteoles near the centre or in the lower half (often concealed by the subtending bract), not articulated, in the axils of bracts resembling the scale-leaves. Perianth segments erect-spreading, with a single vein, cream-coloured to yellow; the outer 3 distinctly inserted outside the inner 3, even in the open flower, and narrower and shorter. Filaments sublinear attached to the receptacle or to the base of the perianth segments; anthers basifixed, introrse or dehiscing laterally. Ovary superior or semi-inferior; the 3 carpels free above the receptacle; styles 1 on each carpel, capitate or subcapitate; ovules attached to ven- tral surface, numerous, in 3—c. 6 rows. Fruit dry, dehiscing along the ventral suture; the 3 segments spreading. Seeds numerous, ovoid, with longitudinal ridges; perianth persistent. Distr. Japan (Prov. Mino), China (Kwangsi and Taiwan), Indo-China (Tonkin), in Malesia: Malaya, N. half of Sumatra, Borneo (Sabah, Sarawak), and Central Celebes. In addition to the two species from Malesia and Japan, a third has been described from Tonkin and S. China (Kwangsi): Petrosavia sinii (KRAUSE) KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 257; GAGNEP. FI. Gén. I.-C. 6 (1934) 802, f. 78 (10-13); (Anonymous) Icon. Corm. Sin. 5 (1976) 424, f. 7677. — Protolirion sinii KRAUSE, Notizbl. Berl.-Dahl. 10 (1929) 806. It is highly probable that this is a synonym of P. sakuraii and would thus neatly fill a gap in the range of that species. Ecol. Saprophytes on the forest flora, in Malesia in the hills at 1000-2000 m. Notes. This genus has been placed in the Liliaceae (Liliales) by KRAUSE (1930), the Petrosaviaceae (Alismatales) by HUTCHINSON (1959) and the Miyoshiaceae (Miyoshiales) by NAKAI (1941). Both in being saprophytic and in having 2- or multi-seriate ovules it is anomalous in either the Liliales (sensu HUTCHIN- SON) or the Alismatales. ERDTMAN (Pollen Morph. & Pl. Taxon., Angiosp., 1952, 235) described the pollen as 1-sulcoidate which is unknown in the Scheuchzeriaceae or Alismataceae (Alismatales) but present, although uncommon, in the Liliaceae. M. Y. STANT (Bot. J. Linn. Soc. 63, 1970, Suppl. 1, 147) investigated the anatomy of P. stellaris and found it to be indistinguishable from that in the (saprophytic) Triuridaceae (Triuridales). Although here retained in the Liliaceae it is admitted that further investigation may show it to be better placed elsewhere. The roots lack root-hairs but contain an endotrophic mycorrhiza (Groom, 1895). KEY TO THE SPECIES 1. Inflorescence corymbose; pedicels all arising close to the peduncle apex, usually 10 mm or more long 1. P. stellaris 1. Inflorescence racemose; pedicels spaced along the peduncle, usually 8 mm or less long 2. P. sakuraii 1. Petrosavia stellaris BEcc. Nuov. Giorn. Bot. Ital. 3 (1871) 8, t. 1; Rm. J. Str. Br. R. As. Soc. n. 24 (1891) 171; Groom, Ann. Bot. 6 (1892) 380; HutcH. Kew Bull. (1933) 156; STEEN. Trop. Natuur 23 (1934) 52, f. 12 right; NAKat, J. Jap. Bot. 17 (1941) 191; Henp. Mal. Wild Flow. Monoc. (1954) 178, f. 103; STANT, Bot. J. Linn. Soc. 63 (1970) Suppl. 1, 147, anat. — Protolirion paradoxum Riwv. Ann. Bot. 9 (1895) 56; Groom, l.c. 45, pl. 3; RipL. Mat. Fl. Mal. Pen. Monoc. 2 (1907) 87; Fl. Mal. Pen. 4 (1924) 322, f. 195; Nakal, J. Jap. Bot. 17 (1941) 191. — Fig. 5a. Aerial stems (4-)6-11 cm high. Scale-/eaves of rhizome ovate, c. 2-5 mm long, their base amplexi- caul, often forming a closed sheath. Scale-leaves of aerial stem narrowing gradually to the acute apex, 3-6 mm long, their base partly embracing the stem. Raceme simple, corymbose, with 1-12 flowers. FLORA MALESIANA fser. I, vol. 9! Fig. 5. Petrosavia stellaris Becc. a. Habit, nat. size (Redrawn from HUTCHINSON, Fam. FI. Pl. 2, 1959, fig. 347). — P. sakurati (MAKINO) J. J. SMITH ex STEEN. b. Habit, nat. size, c. flower, d. flower in fruit, both x 10 (Redrawn from MAKkINo, Bot. Mag. Tokyo 17, 1903, pl. 5). Pedicels all arising close to the apex of the stem, usually less than 1 mm apart, (5—)10-16(—20) mm. Outer perianth segments ovate to lanceolate, acute, 1-2 by c. 1/, mm. Inner perianth segments ovate, + acute, 2!/,-31/, by 11/.-2 mm. Filaments c.2 mm; anthers less than 1/, mm long. Ovary + superior; styles up to 1 mm long. Capsule segments 3-4 mm long. Seeds c. 3/, mm long. Distr. Malesia: Sumatra (West Coast Res.), Malay Peninsula, Borneo (Sarawak, Sabah), and Central Celebes (Masamba: Mt Kambuno). Ecol. Recorded by RmDLey (1924) “in dry woods at the foot of Dacrydiums” and by EYMA (EYMA 1305, L) from ‘forest, rather dark, without undergrowth’’. Other records refer to sandy forest, mossy forest and among bamboos. Recorded between 100 and 1000 m. F/. usually Febr.—Sept. 2. Petrosavia sakuraii (MAKINO) J. J. SMITH ex STEEN. Trop. Natuur 23 (1934) 52. — Miyoshia sakuraii MAKINO, Bot. Mag. Tokyo 17 (1903) 144, pl. 5; /.c. 208. — Protolirion miyoshia-sakuraii MAKINO, Bot. Mag. Tokyo 17 (1903) 208, nomen err. et provis., illeg.; PILG. in E. & P. Nat. Pfl. Fam. Nachtr. 3 (1908) 44, f. 8, ditto. — Protolirion sakuraii (MAKINO) Danby, J. Bot. 69 (1931) 53. — Fig. 5b-d. Aerial stems (5—)10—21(-27) cm high. Scale- leaves of rhizome ovate, c. 2-5 mm long, their base amplexicaul, often forming a closed sheath. Scale- leaves of aerial stem narrowing gradually to the acute apex, 4-6 mm long, their base partly embrac- ing the stem or rarely completely amplexicaul. Raceme simple or with few-flowered branches towards the base, elongate, with (3—)6—25(-30) flowers. Pedicels of mature flowers usually at least 2 mm apart, 3-8 mm. Outer perianth segments ovate to lanceolate, acute, 11/,-2 by 1/,-1 mm. Inner perianth segments ovate, + acute, 2-3 by 11/,-13/, mm. Filaments c. 2 mm; anthers less than 1/, mm long. Ovary superior to semi-inferior; styles up to 1 mm. Capsule segments 2-3 mm long. Seeds c. 3/, mm long. Distr. Japan (Prov. Mino), Formosa, Burma and Malesia: northern half of Sumatra (Gajo- lands; Westcoast Res.). Ecol. Along forest paths and on flat forest ridges, 1000-2000 m. F/. March-July, Nov. 5. SCHELHAMMERA R. Br. Prod. (1810) 273; Bru. FI. Austr. 7 (1878) 31; BAKER, J. Linn. Soc. Bot. 17 (1879) 466; BAILEY, Queensl. FI. 5 (1902) 1642; MAIDEN & BETCHE, Cens. N.S.W. Pl. (1916) 40, as Schellhammera; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 266; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 606, as Schelhammeria; nom. gen. cons. — Fig. 6. 1979] LILIACEAE—I (Jessop) 201 annual, simple or with 1-3 branches, often slightly swollen at the nodes; branches erect. Leaves cauline, alternate, flat, with many veins and usually a strong midrib, sessile or shortly petioled, lanceolate to ovate, often at least partly amplexicaul, obtuse or cuneate at the base, acute at apex. Leaves on rhizome and lower parts of aerial stems and branches scale-like. /nflorescence terminal, consisting of a sessile or peduncled umbel or of a single flower. Pedicels straight, articulated or not. Perianth segments free, equal, spreading to shallowly campanulate, with several usually inconspicuous veins. Filaments flattened, tapering from the base, attached to the base of the perianth segments; anthers basifixed, linear-oblong, extrorse. Ovary superior, sessile, obovoid, globose or oblong, often fairly deeply 3-lobed, 3-celled; ovules axile, few (c. 4-12) per locule; style filiform, with 3 deeply divided branches. Capsule somewhat fleshy. Distr. Three spp. in eastern Australia, one of which also in East Malesia: New Guinea. Ecol. Most records suggest a preference for rain-forest, but also recorded in scrub and on open slopes in the lowland and hills. 1. Schelhammera multiflora R. Br. Prod. (1810) 274; F.v.M. Descr. Not. 4 (1876) 73; Bru. FI. Austr. 7 (1878) 32; Laut. Bot. Jahrb. 50 (1913) 292; HALL. f. Nova Guinea 8 (1914) 989; KRAUSE, Bot. Jahrb. 59 (1925) 548. — Fig. 6. Stems 10-40 cm, rarely minutely and sparsely pubescent. Leaves 4-8 by 1-3 cm, usually minutely ciliate on the margin and veins towards the base; petiole usually absent, rarely up to 5 mm. Scale- leaves ovate to lanceolate, dry, sessile, amplexicaul, usually 5-10 mm long. Pedicels usually 5-30, articulated at base of flower, 1-3 cm. Perianth segments obovate, acute, with the sides turned up to form a gutter-shaped structure in which the anthers are partly enclosed, swollen on the abaxial surface at the base, 4'/,-8 by c. 2 mm, white. Filaments c. 3 mm; anthers c. 2 mm long, brown or black. Ovary obovoid, 6-lobed, c. 11/; mm long; style 3-4 mm, divided for at least half its length; branches adhering to one another rather firmly at first, later recurving; ovules few (c. 2-4) per locule. Fruit and seeds not seen. Distr. Australia (Queensland) and E. Malesia: S. New Guinea (Merauke area: Okaba; Fly R. area). Ecol. In lowland grassland on open slopes, sub- ject to a strong or distinct dry season, in Queens- land also in open forest, obviously a rare species, below 400 m. F/. March—Sept. Note. The two further Australian species are S. undulata, which has larger solitary flowers, and Fig. 6. Schelhammera multiflora R. Br. a. Habit, S. pedunculata, which has peduncled umbels. x 1/,, b. flower, c. capsule, both x 2. 202 FLORA MALESIANA [ser. I, volt 91 6. ARTHROPODIUM R. Br. Prod. (1810) 276; Baker, J. Linn. Soc. Bot. 15 (1876) 351; Bru. Fl. Austr. 7 (1878) 55; BAILEY, Queensl. Fl. 5 (1902) 1629; Ewart, FI. Vict. (1930) 292; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 286; BLAcK, FI. S. Austr. 1 (1943) 193; PAYENS, Nova Guinea n.s. 8 (1957) 388; SCHLITTLER, Mitt. Bot. Mus. Un. Ziirich 207 (1957) 6, map; HuTcu. Fam. FI. Pl. ed. 2, 2 (1959) 598. — Dichopogon KUNTH, En. Pl. 4 (1843) 622; BAKER, J. Linn. Soc. Bot. 15 (1876) 318; Bru. Fl. Austr. 7 (1878) 58; BAILEY, Queensl. FI. 5 (1902) 1631; Ewart, Fl. Vict. (1930) 291; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 285; BLAck, FI. S. Austr. 1 (1943) 193; Hurcu. Fam. FI. Pl. ed. 2, 2 (1959) 598. — Fig. 7. Stemless herbs. Rhizome very short. Roots fibrous or somewhat fleshy, often producing well-defined, distant tubers. Leaves basal, linear to lanceolate, gradually expanding towards the base to form (apparently dry and membranous) open sheaths, sometimes ciliate or with bristles along the margins; veins of the sheaths sometimes persisting on an outer zone of fibres. Jnflorescence a raceme or panicle. Pedicels \—3(—5)-nate, in the axils of bracts and usually associated with a few smaller bracts also apparently in the axil of the principal bract, articulated usually in the distal half, rather long and filiform. Perianth segments usually free, the inner whorl broader, with 3 or less often 5 veins, spreading. Filaments linear-filiform, usually attached to the base of the perianth segments; anthers basifixed, oblong to linear, often strongly arcuate, dehiscing laterally or introrsely; papillose or pubes- cent appendages always present, usually adnate to the filaments and often also to the anthers. Ovary superior, sessile, subglobose to ellipsoid, 3-celled; ovules axile, 2-10 in each locule; style filiform, simple, minutely capitate. Capsule dehiscing loculicidally; perianth segments adhering, marcescent, not twisting after flowering. Seeds angular. Distr. Madagascar (1 sp.), New Zealand (2 spp.), New Caledonia (1 sp.), and c. 9 spp. in Australia, of which one extends into Malesia: New Guinea (South Papua). Ecol. Usually in open grassland or open woodland, often at rather low altitudes (below 250 m). Species have been described as having chocolate or vanilla scents. Notes. PAYENS (1957) made a strong case for combining Dichopogon with Arthropodium. Nevertheless, authors (e.g. BURBIDGE and GRAY, FI. A.C.T., 1970) have continued to separate these genera principally on whether the papillose or pubescent staminal appendages are adnate to the filaments (in Arthropodium) or to the anthers (in Dichopogon). An examination of species represented at Leyden and Adelaide has convinced me that these two genera cannot be separated. A particularly significant form is A. neo- caledonicum in which the appendages are attached to both filaments and anthers such that it would be difficult to assign this species to a genus. In most of the material of Dichopogon the appendages are also distinctly adnate to the filaments as well as to the anthers. Dichopogon strictus and Arthropodium capillipes, although placed in separate genera, have almost indistinguishable stamens. 1. Arthropodium strictum R. Br. Prod. (1810) 276; Roots bearing distant tubers c. 1—-1'/, cm long. F.v.M. Descr. Not. 6 (1885) 17; Laur. Bot. Jahrb. 50 (1913) 292; Krause, ibid. 59 (1925) 548; PAYENS, Nova Guinea n.s. 8 (1957) 390; EICHLER, Suppl. Fl. S. Austr. (1965) 83. — Dichopogon strictus (R. Br.) BAKER, J. Linn. Soc. Bot. 15 (1876) 319; Bru. Fl. Austr. 7 (1878) 58; BAILEy, Queensl. FI. 5 (1902) 1631; Ewart, Fl. Vict. (1930) 291; GARDNER, En. Pl. Austr. Occ. (1931) 18; BLAck, FI. S. Austr. 1 (1943) 194. — Fig. 7. Leaves 3-12, suberect, sublinear but narrowing gradually towards the apex and sometimes also towards the basal sheath where they again become broader, glabrous or minutely ciliate on the mar- gins, 20-45 cm by 3-7 mm; veins of leaves not forming fibrous sheaths at base of plant. Peduncle simple or more often with 1-4 ascending branches, (20-)30-60(—110) cm high. Bracts rarely up to 6 cm long at the base of the lowest branch, but usually 1979] 5-15 mm long, ovate to lanceolate, narrowing gradually to the apex, partly scarious. Pedicels solitary or less often 2- or 3-nate, erect-spreading, 12-20 mm. Perianth segments purple, 9-14 by c. 3-3'/, mm. Filaments c. 1'/,-2 mm; anthers 3-5 mm long; appendages less than 1 mm long, free of or only shortly adnate to filaments. Ovary globose-ellipsoid, 11/,.-4 mm long; ovules 8-10 in each locule; style 6-7 mm. Capsule subglobose, c. 5mm @, with several seeds; perianth marcescent or rarely persistent. Distr. Australia (all states but not yet from the Northern Territory) and Malesia: SE. New Guinea (Port Moresby area). Ecol. Open grassland and open woodland at low altitude, subject to a long dry season. Vern. Chocolate lily, E (Australia). Note. PAYeENsS (1957) stated that the perianth segments are connate for 1 mm. This was not con- firmed by me for material he had examined. LILIACEAE—I (Jessop) 203 ae <) \ \ j y J J : ot J Ry} 2 - 7] ." Ft ~ D e \; aS {Fe ly CEH TRA Sa c Fig. 7. Arthropodium strictum R. Br. a. Habit, x 1/,, b. flower, X 2, c. withered flower, x 3, d. gynoecium, X 7, e. anther, x 10(C. R. ALcock 2875, S. Australia). 7. CAESIA R. Br. Prod. (1810) 277; BAKER, J. Linn. Soc. Bot. 15 (1876) 357; Brn. Fl. Austr. 7 (1878) 46; BAKER, FI. Cap. 6 (1897) 400; BAILEY, Queens]. Fl. 5 (1902) 1632; Ewart, FI. Vict. (1930) 289; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 288; BLACK, FI. S. Austr. 1 (1943) 192; PHiLuips, Gen. S. Afr. Fl. Pl. ed. 2 (1951) 204 FLORA MALESIANA [ser. I, vol. 9! 1002; PAYENS, Nova Guinea n.s. 8 (1957) 383; SCHLITTLER, Mitt. Bot. Mus. Un. Ziirich 207 (1957) 6, map; HuTcH. Fam. FI. Pl. ed. 2, 2 (1959) 596; OBERMEYER, Bothalia 11 (1973) 122. — Fig. 8. Stemless herbs. Rhizome very short. Roots fibrous, fleshy or tuberous. Leaves basal, subulate to linear, expanded at the base to form a sheath. Inflorescence a raceme or more often a panicle. Pedice/s 1—3-nate, in the axils of bracts and usually associated with a few smaller bracts also apparently in the axil of the principal bract, articulated usually in the distal half. Perianth segments free or shortly con- nate, subequal, 3-veined, spreading. Filaments filiform or linear, glabrous, attached to the receptacle or to the base of the perianth segments; anthers basifixed, oblong, dehiscing introrsely. Ovary superior, sessile, globose or subglobose, 3-celled; ovules axile, 2 in each locule; style filiform, simple, minutely capitate. Capsule dehiscing loculicidally; perianth segments adhering, marcescent, twisting spirally after flowering. Seeds angular or globose, usually solitary in each locule. Distr. Madagascar (1 sp.), South Africa (2 spp.; Cape Province) and Australia (7-9 spp.), of which one species also occurs in Malesia: S. Papua New Guinea (W. Distr.: Wassi Kussa area). Ecol. Usually in the open, but also in savanna. Note. Generically this group is regarded as well-defined by the spirally-twisting perianth and 2-ovuled locules. The South African monotypic genus Nanolirion Btu. (1883) was formerly distinguished by having a 1-3-flowered inflorescence (PHILLIPS, 1951), but OBERMEYER (1973) placed it under Caesia. 1. Caesia setifera BAKER, J. Linn. Soc. Bot. 15 (1876) 359; Bru. Fl. Austr. 7 (1878) 47; Ewart & Davies, Fl. North. Terr. (1917) 71; PAYENS, Nova Guinea n.s. 8 (1957) 384, f. 1. — Fig. 8. Roots bearing distant spindle-shaped tubers c. 1-1'/, em long. Leaves 2-4, suberect, filiform, glabrous, c. 10-25 cm long, c. 1 mm broad; veins of leaves forming fibrous sheaths at base of plant. Peduncle thin and wiry, usually with | or 2 ascend- ing branches, (17—)25-45 cm high. Bracts bearing branches or pedicels in their axils, rarely up to 5mm long, ovate to lanceolate, partly scarious. Pedicels 1-6-nate, usually erect-spreading, (3-)5-10 mm. Perianth segments blue, linear-oblong, 6-8 by 1-1'/, mm. Filaments 3-4 mm (the outer up to 1 mm longer than the inner); anthers yellow, c. 1/.- 3/,mm long. Ovary c. 3/,-1 mm long; style 4-5 mm. Capsule subglobose, deeply 3-lobed, c. 3 mm @. Distr. Australia (Queensland and Northern Territory); in Malesia: S. Papua New Guinea: Western District, Wassi Kussa area: Morehead, Weam, Arufi, Tarara. Ecol. In the Wassi Kussa area scattered on open, grass-sedge plains on thin sand over clay, in maintained savanna grassland, in savanna with Melaleuca and Acacia, on wet flats in savanna forest, on sour grey soils, at very low altitudes sub- ject to a long dry season. F/. July-Aug., Dec.—Jan. A field note recorded that usually one flower is opening in sequence. Fig. 8. Caesia setifera BAKER. a. Flower after anthesis, with persistent twisted perianth, b. sepal inside, c. stamen, all x 6, d-e. anther, dorsal and lateral, x 15 (a BRAss 8599, b-e BRAss 8560). 1979] LILIACEAE— I (Jessop) 205 8. CHLOROPHYTUM KER-GAWL. Bot. Mag. 27 (1808) t. 1071; BAKER, J. Linn. Soc. Bot. 15 (1876) 321; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 284; PoELLN. Ber. Deut. Bot. Ges. 61 (1943) 126; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 598; Dress, Baileya 9 (1961) 29; OBERMEYER, Bothalia 7 (1962) 690; BACK. & BAKH. f. Fl. Java 3 (1968) 86; CHAROENPHOL, Thai For. Bull. 7 (1973) 67; PANIGRAHI, Kew Bull. 30 (1975) 563; MARAIS & REILLY, Kew Bull. 32 (1978) 653. Perennial stemless herbs. Rhizome horizontal, often very short. Roots fibrous, or fleshy or tuberous. Leaves basal, usually rosulate, linear to lanceolate, often with a fimbriate margin, expanding towards the base to form a sheath. Inflorescence a raceme or a panicle. Pedice/ls 1—6-nate, in the axils of bracts and associated with a few small bracts also apparently in the axil of the principal bract, articulated usually near or below the middle. Perianth segments free, subequal, with 3 or 5 veins, most frequently spreading or reflexed. Filaments filiform, glabrous or papillate, attached to the receptacle; anthers basifixed, linear-oblong, introrse. Ovary superior, sessile or shortly stipitate, globose or obovoid, 3-lobed, 3-celled; ovules axile, 2 or more in each locule; style filiform, simple, minutely capitate. Capsule dehiscing loculicidally; perianth segments adhering, marcescent, not twisting after flowering. Seeds flat, suborbicular. Distr. Especially in Africa (mainly southern and tropical), Madagascar and Asia (especially India) to Australia (2 spp.); in West Malesia two non-endemic species. The total number of species is estimated by KRAUuSE (1930) at c. 100 and by OBERMEYER (1962) at nearly 300. Ecol. Species of Chlorophytum occur in a wide range of habitats from coastal to montane regions. They grow in many soil types and in rock crevices, and in open grassland and dense forest. Notes. OBERMEYER (1962) recorded that in species with fascicled flowers plants do occasionally occur in which the flowers are borne spirally on a distinct but very short lateral branch of the inflorescence. This supports the theory that fascicled flowers and their associated bracts represent an abbreviated lateral shoot. She also discussed the difficulty encountered in separating Chlorophytum from Anthericum L. (1753). She was only able to find a single character on which these genera could always be separated: the seeds of Chlorophytum are flat, but of Anthericum angular and smaller. Several other characters were found to be generally, but not universally, of value in separating them. KEY TO THE SPECIES 1. Perianth segments 3-5 mm long. Anthers less than 1 mm long. Leaves usually 4-8 mm broad 1. C. laxum 1. Perianth segments 7-12 mm long. Anthers 4-5 mm long. Leaves usually 10-50 mm broad 2. C. malayense Roots often bearing tubers c. 1-5 cm long (absent in material I have seen from Malesia and stated by 1. Chlorophytum laxum R. Br. Prod. (1810) 277; Hook. f. Fl. Br. Ind. 6 (1892) 336; BAck. Handb. Fl. Java 3 (1924) 52, incl. f. javanicum (HASsK.) BACK.; GAGNEP. FI. Gén. I.-C. 6 (1934) 804; BAck. & Baku. f. Fl. Java 3 (1968) 86; Hutcu. & Daz. Fl. W. Trop. Afr. ed. 2, 3 (1968) 100. — C. laxi- florum Baker, J. Linn. Soc. Bot. 15 (1876) 328, nom. illeg. — Nolina javanica Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 120; Mia. FI. Ind. Bat. 3 (1859) 554. BACKER & BAKHUIZEN VAN DEN BRINK f/f. to be absent in Javanese plants). Leaves 4-12 or rarely more, linear to lanceolate, usually channelled especially towards the base, with a rather promi- nent midrib; suberect, glabrous, 10—30(—50) cm by 4-8(-12) mm, rarely forming fibrous sheaths round the base of the plants; the outer arcuate and recurved; the inner often straight. Peduncle straight 206 FLORA MALESIANA [ser. I, vol. 91 or flexuous, unbranched or rarely with a single branch, (5—)15—35(—60) cm. Lower bracts 3-12 mm long, usually lanceolate-acuminate, partly scarious. Pedicels solitary or, less often, 2-nate, often erect- spreading but recurving in fruit, 3-12 mm. Perianth segments white or ‘greenish white’, linear- oblong, 3-5 by c. 1 mm. Filaments (1'/,-)2-3 (4) mm long (the outer often longer than the inner); anthers up to !/, mm long. Ovary c. 1 mm long, obovoid or globose; style less than 2 mm; ovules 2 per locule. Capsule usually obovoid, less often globose or depressed-globose, 3-lobed, 5-10 mm long. Distr. Tropical Africa (e.g. Senegal, Liberia, Ghana, Nigeria, Zambia, Ethiopia), S. Asia (e.g. India, Indo-China, Thailand), China (incl. Hainan); in Malesia: N. Sumatra (Eastcoast Res.), Malay Peninsula, W. and Central Java, and SE. Borneo (Bandjermasin), extending to N. Australia (Queensland and Northern Territory). Ecol. Recorded from a variety of localities including rock crevices and in sandy soils, but usually a species of shady places (including bamboo forest, deciduous forest and a coconut grove), usually below 1000 m. F/. Jan.—Dec. Notes. BACKER (1924) and BACKER & BAK- HUIZEN VAN DEN BRINK f. (1968) treat the Javanese plants as belonging to forma javanicum (HASSK.) BACK. They give no reason for this decision and in the absence of a monograph of this species I would prefer not to uphold this form. Extra-Malesian synonyms are omitted from the references. 2. Chlorophytum malayense RIDL. Fl. Mal. Pen. 5 (1925) 341; PANIGRAHI, Kew Bull. 30 (1975) 565. — C. orchidastrum (non LINDL. Bot. Reg. 10 (1824) t. 813) RipL. Mat. Fl. Mal. Pen. Monoc. 2 (1907) 92; Fl. Mal. Pen. 4 (1924) 327; GaGnep. Fl. Gén. I.-C. 6 (1934) 806; CHAROENPHOL, Thai For. Bull. 7 (1973) 67. Roots sometimes bearing tubers 3-4 mm or more long. Leaves 3-10, sublinear or indistinctly petioled and with a lanceolate lamina, channelled towards the base, with a rather prominent midrib, suberect, glabrous, (10—)25-60 by (*/,-)1-5(-10) cm, never forming fibrous sheaths round the base of the plant. Peduncle glabrous or less often glandular pubes- cent, with 0-~ branches (30—)40—50(?—90) cm high. Lower bracts up to 9 cm long, usually narrowly lanceolate-acuminate, usually partly scarious; the lowest usually sterile. Pedicels usually 2-3-nate, less often solitary, erect-spreading, not recurving in fruit, 3-10 mm. Perianth segments white, elliptic to linear-oblong, 7-12 by 2-3 mm. Filaments occasionally minutely pubescent, 4-5 mm (not always similar in length but neither whorl regularly longer than the other); anthers 4-5 mm long. Ovary c. 11/,-2 mm long, obovoid or globose; style 3-4(-8, GAGNEPAIN, 1934) mm; ovules 4-6 per locule. Capsule globose, strongly emarginate and very deeply 3-lobed, c. 5-8 mm @. Distr. Indo-China, Thailand and Malesia: Malay Peninsula (Perak and Perlis). Ecol. This is usually a forest (including bamboo, oak, pine) species, often associated with limestone, 50-1500 m. F/. Jan.—Dec. Notes. GARRETT (in sched.) recorded that the plant is night-flowering in Thailand. RIDLEY (1925) gave the new name C. malayense to what he had previously identified as C. orchid- astrum LINDL. However, he gave no characters by which these species could be separated. In the neighbouring territories of Thailand and Indo-China CHAROENPHOL (1973) and GAGNEPAIN (1934) have continued to recognize C. orchidastrum, but Iam following PANIGRAHI (1975) who considers that C. orchidastrum sensu stricto is confined to Africa, that the Indian material should be referred to C. nimmonii and C. glaucum, and that the SE. Asian material is C. malayense. Our species can, according to PANIGRAHI, be recognized inter alia by drying greenish brown or glaucous, in the leaves not being distinctly petioled and 3-5 cm broad, in the scape being up to 50 cm long and shorter than the leaves and in the bracts being up to 9 cm long. Few collections have been made of C. malayense and further field work is needed toconfirm its status. C. longissimum Rw. (J. Str. Br. R. As. Soc. n. 49, 1907, 209) was described from Trang, Peninsular Thailand, close to the Malesian border. It closely resembles C. malayense. CHAROENPHOL distinguished these species on whether the inflores- cence is erect and sometimes branched (C. malay- ense) or trailing on the ground and never branched (C. longissimum). The type of C. longissimum has not been seen, but specimens at Kew identified as such, and agreeing with the type description, are possibly sufficiently characterized by these inflores- cence characters to retain it as a distinct taxon. Whether the differences are sufficient for specific rank must be left to future examination; it is provisionally kept distinct. It was not discussed by PANIGRAHI. 9. DIANELLA LAMK [Encycl. 2 (1786) 276, nom. inval.] ex Juss. Gen. Pl. (1789) 47; BAKER, J. Linn. Soc. Bot. 14 (1875) 574; Bru. Fl. Austr. 7 (1878) 13; BAck. Handb. FI. Java 3 (1924) 53; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 1979] LILIACEAE—I (Jessop) 207 295; SCHLITTLER, Mitt. Bot. Mus. Un. Ziirich 163 (1940) 256; Blumea 6 (1948) 200; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 606. — Rhuacophila BL. En. Pl. Jav. (1827) 13. — Fig. 9. Erect or climbing glabrous herbs. Rhizome short to stoloniferous. Roots fibrous. Leaves usually cauline and distichous or basally rosulate, linear, distally dorsiven- tral, in the lower parts sometimes laterally compressed and keeled and often form- ing a closed sheath at the base. /nflorescence a panicle. Pedicels solitary or few, usually in the axils of bracts, articulated immediately below the flower. Perianth segments free, 3-7-nerved, spreading or recurved, equal or subequal. Filaments filiform or linear, often swollen in the distal half, glabrous, attached to the recep- tacle or the inner whorl attached to the perianth segments; anthers basifixed, linear to oblong, dehiscing by an apical pore or by a slit which is initiated in an apical pore. Ovary superior, more or less sessile, globose, 3-celled; ovules axile, 4-8 in each locule; style filiform, simple, minutely capitate. Fruit a berry, usually shiny blue; perianth segments adhering, marcescent, not twisting after flowering; base of style persistent. Seeds globose or angled. Distr. Continental Africa (one record), Madagascar, through India into China, throughout Malesia and Australia to New Zealand and the Pacific Islands (New Caledonia, Sandwich Is., Norfolk and Fiji). Estimates of the number of species vary widely; there are possibly 20-30 spp. in all. Ecol. Both in forest and in more open localities. KEY TO THE SPECIES 1. Leaf bases strongly compressed and keeled. Leaves usually with minute teeth or prickles along the margin and abaxial surface of the midrib. Fertile bracts differing considerably in size from even the smaller leaves. Filaments strongly thickened at the apex under the anther. . . . . 1. D. ensifolia 1. Leaf bases obtuse in section, not keeled. Leaves always smooth. Fertile bracts grading into the leaves. Filaments widened about the middle 1. Dianella ensifolia (L.) DC. in Redoute, Liliac. 1 (1802) t. 1, cf. Coop in Bosser c.s. Fl. Mascar. Lil. (1978) 32; Mia. FI. Ind. Bat. 3 (1859) 560; HEmsL. Rep. Chall. Bot. 1, 3 (1884) 201; RipL. Mat. FI. Mal. Pen. Monoc. 2 (1907) 92; Laur. Bot. Jahrb. 50 (1913) 293; HALL. f. Nova Guinea 8 (1914) 995; Merr. En. Born. (1921) 114; En. Philip. 1 (1922) 203; Rip. Fl. Mal. Pen. 4 (1924) 329; KRAUSE, Bot. Jahrb. 59 (1925) 551; Merr. Contr. Arn. Arb. 8 (1934) 18; Witp, Kew Bull. 8 (1953) 251; M. R. HENDERSON, Mal. Wild Flow. Monoc. (1954) 186, f. 110; Back. & Baku. f. Fl. Java 3 (1968) 87; STEEN. Mt. Fl. Java (1972) t. 28-1. — Gladiolus odoratus indicus RumMpH. Herb. Amb. 5 (1747) 185, t. 73. — Dracaena ensifolia LINNE, Mant. (1767) 63. — Dianella nemorosa LAMK, Encycl. 2 (1786) 276, nom. inval.; Tabl. Enc. 2 (1792) 388, t. 250, nom. illeg.; SCHLITTLER, Mitt. Bot. Mus. Un. Ziirich 163 (1940) 256; Blumea 6 (1948) 209, incl. many forms, /.c. 216-223. — Dracaena ensata THUNB. Diss. Bot. Drac. (1808) 4. — Dianella montana BL. En. Pl. Jav. 1 (1827) 12; Hassk. Tijd. Nat. Gesch. Phys. 11 (1844) 180; PI. Jav. Rar. (1848) 114; Mia. Fl. Ind. Bat. 3 (1859) 560; Back. Handb. Fl. Java 3 (1924) 54. — 2. D. javanica D. odorata [RumPpH.] BL. En. PI. Jav. 1 (1827) 13; HALL. f. Nova Guinea 8 (1914) 996; Merr. Int. Rumph. (1917) 136; Krause, Bot. Jahrb. 59 (1925) 550; SCHLITTLER, Mitt. Bot. Mus. Un. Zurich 163 (1940) 258. — D. revoluta (non R. BR.) SCHAUER, Nov. Act. Ac. Nat. Cur. 19 (1843) Suppl. 1: 445; Laut. Bot. Jahrb. 50 (1913) 293; Merr. En. Philip. 1 (1922) 203. — D. bancana Mia. FI. Ind. Bat. Suppl. (1861) 610; BAck. Handb. FI. Java 3 (1924) 54. — D. caerulea (non Sims, Bot. Mag. 15, 1801, t. 505) Merr. Philip. J. Sc. 2 (1907) Bot. 266; ibid. 5 (1910) Bot. 337; HALL. f. Nova Guinea 8 (1914) 993; Gipss, Arfak (1917) 100; Merr. En. Philip. 1 (1922) 202; Krause, Bot. Jahrb. 59 (1925) 552; HoLtuuts, Blumea 5 (1942) 167. — D. robusta Evmer, Leafli. Philip. Bot. 5 (1913) 806. — D. bam- busifolia HALL. f. Nova Guinea 8 (1914) 995, t. 182; KRAUSE, Bot. Jahrb. 59 (1925) 550. — D. flabellata Hatt. f. lc. 997, t. 183; Krause, /.c. 549. — D. carinata HALL. f. l.c. 999, t. 186; KRAUSE, /.c. 550. — D. parviflora Zipp. ex HALL. f. l.c. 998, t. 184; Krause, /.c. 551. — C. albiflora HALL. f. l.c. 998, t. 185; KRausE, /.c. 551. — D. monophylla Hatt. f. lc. 1000, t. 188; Krause, /.c. 551. — D. serrulata HALL. f. l.c. 1000, t. 187; KRAUSE, FLORA MALESIANA i ' ‘ 8 ae 2 = ; 1 2 ‘ ; ' [ser. I, vol. 91 Fig. 9. Dianella javanica (BL.) KUNTH, on Mt Kaba, S. Sumatra (Photogr. DE Voocp). lic. 549. — D. parviflora Ripv. J. Fed. Mal. St. Mus. 6 (1915) 186. — D. pullei KRAUSE, Nova Guinea 14 (1924) 175; Bot. Jahrb. 59 (1925) 553. — D. ledermannii KRAUSE, Bot. Jahrb. 59 (1925) 549. — PD. monticola Krause, I.c. 553. — ? D. levis (non R. Br.) C. T. Wuirte, Proc. Linn. Soc. N.S.W. 51 (1926) 298. — D. sparsiflora SCHLITTLER, Mitt. Bot. Mus. Un. Zurich 163 (1940) 262. — D. ensata (THUNB.) R. J. HENDERSON, Taxon 26 (1977) 136. Stem 0-1 m high, unbranched, rarely with a few branches. Rhizome horizontal, moderately branched. Leaves basal, scattered along the stem or in a terminal rosette, distichous, with a sheathing lower part, (25—)30—60(-100) cm by (4—)8-30 mm; above the base keeled and with the sides of the lamina becoming firmly appressed to one another and fused to form an isobilateral portion; distally with a dorsiventral linear or linear-lancelate 1979] lamina which is sometimes absent from the lower leaves, almost always with minute serrations or prickles along the midrib on the lower surface, with a conspicuous midrib and numerous smaller veins. Inflorescence exceeding the leaves, lax or with short terminal branches often c. 1-2 cm long, bearing up to 20 pedicels. Lower bracts usually narrowly linear-lanceolate and bilaterally com- pressed above the basal sheath like the leaves; bracts subtending pedicels 1-4(-7) mm long or rarely absent. Pedicels 4-15(-22) mm. Perianth segments blue, white, lilac or yellow, spreading, (4-)5—8(-9) mm long. Filaments often more than half as long as the perianth segments, filiform or narrowly linear, white or yellow with a yellow or orange, glabrous swelling below the anther. Ovary green, 1'/,-2 mm long; ovules 4 in each locule; style green, white or blue, 4-6 mm long. Fruit shiny blue, 6—-8(—11) mm @. Seeds several, 3-4 mm long. Distr. Continental Africa (WiLp, 1953), Madagascar, continental Asia (Himalayas, Burma, Thailand, Indo-China) to southern China (Yunnan, Hainan), Japan and Formosa, throughout Malesia to Australia (Northern Territory, Queensland, New South Wales), Tasmania, New Zealand and many Pacific islands. Ecol. A highly adaptable species, occurring in habitats ranging from open grassland to primary forest, from sea-level to over 3000 m. F/. Jan.— Dec: Vern. Malaya: benjuang, satagit, senjuang, siak-siak jantan; Sumatra: (akar) tu(n)daun, mentuntil, tengari, ukop, Banka, siak-siak, Riouw, sieuh, Djambi, sitanggit, Batak, sesiah, Pasemah, sitangie, Indragiri; Java: djamaka, d. putih, suliga, S, tegari, J; Borneo: labeh-labeh, Dusun Penam- pang dial., angkup-angkup, Bokan dial., tembalong tipoh, Dusun dial.; Philippines: abldas, Bag., bariu- bariu, oyon-dyon, P.Bis., dungau, Ig., hogangan, If.; Moluccas: mariuii, Talaud; New Guinea: suruma; bururl, tirambaramp, Mendi, buru-buru, Biak dial., moalengen, Aiome, tanglenu, Wigote, Wapi lang., sinda, Mumuni, Orokaiva lang., tsiri kande kande, Hagen Subdistr., bonkaige, Sinasina lang., Nimai dial., pfifiriki, Kutubu lang., kili-kili, Bembi, kikipatia, Koroba Subdistr., sabetari, Rawa, kilina, Kaigorin, sarpeim, Miwaute, Wapi lang., pengeh-pengeh, Maipa, Mekeo lang., baibigehgi, Utukap, Miniafi lang. Notes. BACKER (1924, 1968) recognized two species, D. montana and D. bancana, to include the material from Java, which he distinguished mainly on flower colour, venation of petals, and inflores- cence form. However, the large number of inter- mediates makes this separation impracticable. SCHLITTLER (1940) recognized 3 spp. in Malesia (D. odorata, D. nemorosa and D. sparsiflora), but in 1948 he reduced these to a single one under the name D. nemorosa, with 24 forms in Malesia. Being a common species over a large area, LILIACEAE—I (Jessop) 209 D. ensifolia is exceptionally well represented in herbaria and, despite its variability, I doubt that even with intensive field work distinct subspecific taxa can be defined. 2. Dianella javanica (BL.) KUNTH, En. Pl. 5 (1850) 52; Mia. FI. Ind. Bat. 3 (1859) 561; HALL. f. Nova Guinea 8 (1914) 995; Merr. En. Born. (1921) 114; En. Philip. 1 (1922) 203; Rip. Fl. Mal. Pen. 4 (1924) 328; Back. Handb. FI. Java 3 (1924) 53; KRAUSE, Bot. Jahrb. 59 (1925) 549; Merr. Contr. Arn. Arb. 8 (1934) 19; ScHLITTLER, Blumea 6 (1948) 206, incl. f. stenophylla, alba et rubra SCHLITTLER, /.c. 208; Mitt. Bot. Mus. Un. Zirich 207 (1957) 6, 11, map, 25; BAcK. & BAKH. f. FI. Java 3 (1968) 87; STEEN. Mt. Fl. Java (1972) t. 28-2. — Rhuacophila javanica BL. En. Pl. Jav. (1827) 14; RIDL. J. Linn. Soc. Bot. 42 (1914) 166; J. Fed. Mal. St. Mus. 6 (1915) 186. — Rhuacophila celebica BL. En. Pl. Jav. (1827) 14. — Eustrephus javanicus (BL.) D. Dretr. Syn. Pl. 2 (1840) 1117. — Eustre- Phus celebicus (BL.) D. Dietr. l.c. — D. celebica (BL.) KuntuH, En. Pl. 5 (1850) 45; Mia. Fl. Ind. Bat. 3 (1859) 561. — D. austro-caledonica SEEM. FI. Vit. (1868) 312; Laur. Bot. Jahrb. 50 (1913) 294. — Fig. 9. Stem always present, up to 2 m high, unbranched or branched. Rhizome ‘horizontal, moderately branched. Leaves concentrated towards the ends of the branches, distichous, 12—35(-40) by 3/,— 2'/, cm, with a sheathing but not closed base which is continuous with the lamina, lacking a compressed and fused intermediate portion, lacking serrations or prickles; midrib usually more conspicuous than the many other veins. Jnflorescence lax, exceeding the leaves. Bracts subtending branches of the inflorescences grading gradually in size into the foliage leaves, up to 25 by 3 cm, ovate to lanceolate not bilaterally compressed; bracts subtending the pedicels 1/,-3(-5) mm long or absent. Pedicels 6-20 mm. Perianth segments blue (usually pale), white to yellow or lilac, spreading, (6-)8-12 mm long; outer whorl strongly cucullate at the apex. Filaments usually less than half as long as the perianth segments, white or yellow, filiform, usually with a distinct papillose fusiform swelling about the middle. Ovary green, c. 2 mm long; ovules c. 8-10 in each locule; style white or green 2-5 mm. Fruit green or yellow at first, sometimes (at least) becoming black, ellipsoid, 8-15 mm long, with up to 10 seeds. Seeds c. 11/, mm @. Distr. Throughout Malesia, also in New Cale- donia and Ile des Pins, and Fiji Is. (Viti Levu). Ecol. Grows in a wide variety of habitats, including mossy forests, thickets, forest borders, on narrow open ridge-crests, in debris of craters, among rocks, on stream banks and in exposed places, locally common, (1000—)1500-3000 m. F7. Jan.—Dec. Vern. Java: hadjéra, S; Philippines: kallawad, If., apilug, sapilau, urdya, \g., talobatub, Bon. 210 FLORA MALESIANA [ser. I, vol. 9" Fig. 10. Thysanotus tuberosus R. Br. a. Habit, x 1/4, b. flower, x 5, c. sexual organs, x 10, d. fruit in persistent perianth, x 5. — T. chinensis Btn. e. Habit, x '/, (a-d BRASS 6517, e VAN ROYEN & SLEUMER 5632). 1979] LILIACEAE—I (Jessop) 10. THYSANOTUS R. Br. Prod. (1810) 282; Baker, J. Linn. Soc. Bot. 15 (1876) 334; Bru. Fl. Austr. 7 (1878) 36; Rip. Fl. Mal. Pen. 4 (1924) 328; KRAUusE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 285; Ewart, FI. Vict. (1930) 288; BLACK, FI. S. Austr. 1 (1943) 190; PAYENS, Nova Guinea n.s. 8 (1957) 386; HuTCH. Fam. FI. Pl. ed. 2, 2 (1959) 598, nom. gen. cons. — Chlamysporum SAuIsB. Parad. Lond. (1808) t. 103. — Halongia JEANPLONG, Act. Bot. Ac. Sc. Hung. 16 (1970) 293, f. 1-6. — Fig. 10. Stemless herbs. Rhizome horizontal, very short. Roots fibrous or tuberous. Leaves basal, rosulate, filiform or linear, glabrous, expanding towards the base to form a sheath. Inflorescence a raceme, panicle or umbellate, erect or twining. Pedicels 1—7-nate, in the axils of bracts and associated with a few small bracts also apparently in the axil of the principal bract, or umbellate, articulated often in the median third. Perianth segments free, 3—5-nerved, spreading; outer whorl green; inner whorl usually blue to purple or violet, fringed. Stamens usually 6, rarely 3. Filaments linear, glabrous, attached to the perianth segments, bent over to one side of the ovary (BLACK, 1943); 3 often shorter; anthers basifixed, linear, curved de- hiscing introrsely. Ovary superior, sessile, oblong to globose, 3-celled; ovules axile, 2 in each locule; style filiform, simple, minutely capitate. Capsule dehiscing locu- licidally; perianth segments adhering, marcescent, twisting after flowering; style persistent. Seeds angled, with a fleshy white or orange strophiole. Distr. Up to 30 spp., all in Australia (all states, but especially Western Australia); in Malesia: 2 of these spp., one in New Guinea and another throughout Malesia as far as Thailand, Indo-China and southern China. Ecol. A wide range of mostly open habitats from grasslands and sandy heaths to open forests, including both Eucalyptus savanna and pine forest. Notes. The fleshy appendage of the seed has been referred to as a caruncle (e.g. BLACK, 1943), but it appears to be derived from the funicle rather than from the testa and I am, therefore, using the term strophiole as done by PAYENS (1957). The term ‘cluster’ is sometimes used in preference to umbel as in some material the inflorescence appears umbellate but in other material the pedicels arise from a short but distinct axis. KEY TO THE SPECIES . 1. T. chinensis . 2. T. tuberosus 1. Roots lacking tubers. Inflorescence a simple umbel_. ; 1. Roots with tubers. Inflorescence a panicle, the branches terminated by umbels 1. Thysanotus chinensis Btu. Fl. Hongk. (1861) 372; BAKER, J. Linn. Soc. Bot. 15 (1876) 337; HALL. /. Nova Guinea 8 (1914) 994; Merr. En. Philip. 1 (1922) 202; Krause, Bot. Jahrb. 59 (1925) 548; GARDN. En. Pl. Austr. Occid. (1931) 18; SCHLIT- TLER, Mitt. Bot. Mus. Un. Zurich 207 (1957) 6, map; PAYENS, Nova Guinea n.s. 8 (1957) 386; STEEN. Blumea 20 (1972) 433. — T. chrysantherus F.v.M. [in Bru. Fl. Hongk. (1861) 372, nomen] Fragm. 5 (1866) 202; Brn. FI. Austr. 7 (1878) 40; Naves, Nov. App. (1880) 266; VIDAL, Phan. Cuming. (1885) 153; BAILEY, Queensl. Fl. 5 (1902) 1629. — Chlamysporum chrysantherum (F.v.M.) O. K. Rev. Gen. Pl. 2 (1891) 708. — T. siamensis Row . J. Str. Br. R. As. Soc. n. 59 (1911) 209; FI. Mal. Pen. 4 (1924) 328. — Halongia purpurea JEANPLONG, Act. Bot. Ac. Sc. Hung. 16 (1970) 296, f. 1-6. — Fig. 10e. Roots fibrous. Leaves c. 5 to numerous, erect, often shallowly channelled, usually 15-40 cm, up to c. 1 mm wide. Peduncle about as long as the leaves, straight, unbranched, terminating in a 2-6(-12)-flowered cluster. Bracts with scarious margins, 2-4 mm long. Pedicels 1-2 cm, articulated in the basal third, erect in bud, usually spreading or recurving in fruit. Outer perianth segments (2 always) green, with a scarious, white margin, 6-9 by 2-3 mm; inner segments pale blue to purple or light violet, with inturned usually fringed margins, 6-9 by 3-5 mm. Filaments 1'/,-2 mm, 252 FLORA MALESIANA [ser. I, vol. 91 Fig. 11. Tricoryne platyptera RcuB. a. Habit, x 1/4, b. flower, x 2, c. withered flower, x 4, d. stamen, x 20, e. fruit, x 5 (NGF 38690, PULLEN 7090). 1979 ] the outer whorl shorter than the inner; anthers 1!/,-3 mm long, the outer shorter than the inner. Ovary c. 1 mm long; style c. 3-4 mm. Capsule oblong, 4-5 mm long. Seeds c. 1'/, mm long. Distr. Australia (Western Australia and Nor- thern Territory); in Malesia: New Guinea (West New Guinea and Papua), SE. Moluccas (Aru Is.: Tranggan), Celebes (Masamba), Philippines (Luzon, Mindanao), Lesser Sunda Is. (Flores), onto continental SE. Asia: S. Peninsular Thailand (Setul), Indo-China (Tonkin), S. China (incl. Hong Kong). Ecol. Open grassland, in grassy pine forest, most frequently in open savanna, in New Guinea of Melaleuca, etc., in places subject to a moderate to strong dry season, from close to sea-level up to 1600 m. F/. Dec.—Aug. 2. Thysanotus tuberosus R. Br. Prod. (1810) 282; BAKER, J. Linn. Soc. Bot. 15 (1876) 335; Bru. FI. Austr. 7 (1878) 41; Ewart, Fl. Vict. (1930) 289; GARDN. En. Pl. Austr. Occid. (1931) 18; Brass, J. Arn. Arb. 19 (1938) 190; BLAck, FI. S. Austr. 1 (1943) 191; PaveNs, Nova Guinea n.s. 8 (1957) 387; Burs. & Gray, FI. A.C.T. (1970) 102. — Chlamysporum tuberosum (R. Br.) O. K. Rev. Gen. PI. (1891) 708. — Fig. 10a-d. Roots partly fibrous, bearing spindle-shaped distant tubers 10-25 mm long. Leaves 2-6, erect, linear to terete, 20-50 cm by up to c. 1 mm broad. Peduncle about as long as the leaves or up to 50% longer, usually with 1-6 branches; main axis LILIACEAE—I (Jessop) 213 and branches each terminating in a 2-6(-20)- flowered cluster; 1-—3(—8) flowered fascicles of flowers usually also present in the axils of bracts along the main axis and branches. Bracts usually with scarious margins, the lowest 3-—60(-80) mm long; the upper shorter. Pedicels 7-17 mm, articula- tion usually below the centre, erect to spreading. Outer perianth segments purple with a scarious pale margin, 7-14 by 2-3 mm; inner segments purple with inturning fringed margin, 7-15 by 2!/,-5 mm. Filaments 1-2 mm, those of the outer whorl often shorter than the inner; anthers 24 mm long; outer often shorter than the inner. Ovary 1—1'/, mm long; style (2-)3-5 mm. Capsule oblong, c. 5-6 mm long. Seeds up to 2 mm long. Distr. Australia (all states except Tasmania) and S. Papua New Guinea (Western District: Wassi Kussa; Mabaduan). Ecol. Common grass associate in lowland savanna forest on poorly drained flats, subject to a distinct dry season. F/. Dec.—April. In Australia found in exposed localities up to 500 m. Notes. VAN RoyeN recorded in Queensland that the outside of the flowers was white with a green midrib and the anthers dark purplish green. PAYENS (1957) recognized two varieties separable on flower size. Both occur in Australia but only var. parviflora BTH. in Papua. The material avail- able is insufficient to justify recognition of these varieties. Six other synonyms based on extra-Malesian material are omitted from the synonymy. 11. TRICORYNE R. Br. Prod. (1810) 278; BAKER, J. Linn. Soc. Bot. 15 (1876) 361; Bru. Fl. Austr. 7 (1878) 50; BatLey, Queensl. FI. 5 (1902) 1636; Ewart, Fl. Vict. (1930) 287; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 287; BLAck, FI. S. Austr. 1 (1943) 192; Hutcu. Fam. FI. Pl. ed. 2, 2 (1959) 602. — Fig. 11. Erect, glabrous or variously hairy herbs; stems green, terete, angled or flattened and leaf-like. Rhizome horizontal or erect, often very short; roots thick or fibrous. Leaves basal and/or cauline, more or less linear or reduced to scales, alternate, often amplexicaul at the base. Inflorescence of terminal umbels surrounded by small scarious bracts and outer larger, sometimes leaf-like bracts. Pedice/s articu- lated just below the flower. Perianth segments free or shortly connate, equal or sub- equal, with 3 or 5 (less often 7) prominent veins, spreading. Filaments filiform, with a dense tuft of hairs in the distal part, attached to the receptacle; anthers basifixed, linear to oblong; introrse. Ovary superior, sessile, deeply 3-lobed, 3-celled; ovules basal, 2 in each locule; style filiform, simple, minutely capitate. Fruit consisting of (1-)3 indehiscent nutlets; perianth segments adhering, marcescent, twisting spirally after flowering. Seeds subglobose. Distr. In Australia (all states) 6 spp., of which one in Malesia: S. Papua New Guinea. Ecol. Low altitude plant. BAILEY recorded most of the Queensland species from ‘sandy shores’ and specimen annotations suggest a preference for sandy soils in grassland or savanna. 214 FLORA MALESIANA [ser. I, vol. 9! Note. The terms ‘stem’ and ‘leaves’ give some difficulty as the aerial shoot might be better regarded as an inflorescence often bearing leaf-like bracts rather than as a vegetative shoot with leaves. 1. Tricoryne platyptera Rcnp. Syst. Pflanzenk. (1871) 72; Bru. Fl. Austr. 7 (1878) 51; BAtLey, Queensl. Fl. 5 (1902) 1636; Compr. Cat. Queens]. Pl. (1913) 559, f. 539; Brass, J. Arn. Arb. 19 (1938) 190; PAYENS, Nova Guinea n.s. 8 (1957) 385. — T. pterocaulon BAKER, J. Linn. Soc. Bot. 15 (1876) 363. — Fig. 11. Virgate, erect or subscandent herb. Stems and branches strongly flattened (winged), terete towards the base of the plant, glabrous, 25-80 cm high; flattened portions with a prominent midrib, c. 2-8 mm broad. Rhizome short; roots rather thick. Leaves cauline, narrowly triangular, scale-like, up to 12(-25) mm long. Bracts 1-4 mm long. Pedicels Perianth segments yellow, oblong or elliptic- oblong, with a scarious margin, 6-8(—-10) by up to c. 3 mm; those of the outer whorl usually with (3-)5(-7) veins; those of the inner whorl slightly narrower, with 3 veins. Filaments 2'/,-3 mm, yellow; anthers c. 1-11/, mm long. Ovary c. 1/, mm long; style 31/,-4 mm. Nutlets green. Distr. Australia (tropical Queensland) and Malesia: 8. Papua New Guinea (Western District:. Wassi Kussa area), Thursday I. Ecol. In New Guinea in savanna with Melaleuca and Acacia, in lowland savanna-woodland on sour grey soil and in grass of creek flats, subject to a strong dry season. F/. Dec.—Jan., July—Aug. usually 5-12(-15) in number, (2-)4-12 mm. In N. Queensland found up to 950 m. 12. LILIUM LINNE, Sp. Pl. (1753) 302; Gen. Pl. ed. 5 (1754) 143; Baksr, J. Linn. Soc. Bot. 14 (1875) 225; ELwes, Monogr. genus Lilium (1880); WiLson, Lilies of E. Asia (1925); KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 329; DRYSDALE Woopcock & STEARN, Lilies of the World (1950); SEALY, Kew Bull. 5 (1950) 273; HutTcu. Fam. FI. Pl. ed. 2, 2 (1959) 609. Erect, usually glabrous herbs. Bulb scaly; roots thick, produced from below the bulb and in some species from the base of the aerial stem. Aerial stem annual, erect, usually unbranched. Leaves cauline, alternate or verticillate, linear or broadly flattened, usually sessile, sometimes with bulbils in their axils. Flowers solitary and terminal, or in a racemose inflorescence of solitary flowers in the axils of the often leaf-like bracts. Pedicels erect to cernuous, not articulated. Perianth segments free, + equal, infundibuliform or campanulate, sometimes clawed, sometimes adhering and forming a tube, spreading or recurving distally, with a nectariferous longitudinal furrow towards the base. Filaments filiform or subulate, attached to the receptacle or to the base of the segments; anthers dorsifixed, linear, introrse, versatile. Ovary superior, sessile, oblong to obovoid, 3-celled; ovules axile, co; style terete sometimes clavate, 3-lobed. Capsule coriaceous, loculicidal, perianth deciduous. Seeds flat. Distr. Widespread with possibly 80 spp. in the temperate regions of North America, Europe and Asia (also Taiwan); in Malesia: Philippines. Ecol. The species occupy a wide range of habitats, including open areas and woods. Note. Many species are of horticultural importance. KEY TO THE SPECIES . 1. L. longiflorum 2. L. philippinense 1. Leaves 6-15 mm broad. Nectariferous furrow on perianth segments glabrous 1. Leaves 2-4 mm broad. Nectariferous furrow on perianth segments papillose 1. Lilium longiflorum THuns. Trans. Linn. Soc. 2 (1925) 23; DrysDALE Woopcock & STEARN, (1794) 333; ELwes, Monogr. genus Lilium (1880) t. 7; MeRR. En. Philip. 1 (1922) 204; Back. Handb. Fl. Java 3 (1924) 62; Witson, Lilies of E. Asia Lilies of the World (1950) 253; Hatrus. Mem. Fac. Agric. Kagoshima Un. 5 (1966) 62, err. longifolium; BAcK. & BAKH. f. Fl. Java 3 (1968) 90. 1979] Bulb usually subglobose, often yellowish, up to 6'/, cm @; scales closely imbricate. Stem 30-90 (-100) cm high, smooth, green, producing roots above the bulb. Leaves alternate, 20—40(—60), sessile, lanceolate or oblong-lanceolate, attenuate, with up to 20 or more veins of which up to 7 are usually more conspicuous, the largest on each plant up to 15 by 1'/, cm, usually much smaller distally. Flowers often solitary, but up to at least 7, white, horizontal, (12—)15—20 cm long, infundibuliform, the tube (in dried material) 9-12 mm @. Perianth segments not clawed, reflexed distally; nectari- ferous furrow glabrous. Filaments filiform above, linear below, in dried material 9-14 cm; anthers 7-12 mm long; pollen yellow. Style 8-12 cm; stigma deeply 3-lobed. Capsule c. 4-6 cm long. Distr. China, Japan and Taiwan; in Malesia: Philippines: Batan Is. and Y’ami (N of Luzon), cf. MERRILL and HATUSIMA. Native country not exactly known, possibly naturalized over part of its range. Ecol. Open grassy slopes at low altitude. Vern. Easter lily, E; teppo-yuri, Japan; Philip- pines: vonitan, Iv. Note. Stated by DrysDALE Woopcock & STEARN to be commercially the most important species of Lilium, with numerous cultivated forms. 2. Lilium philippinense BAKER, Gard. Chron. (1873) 1141, f. 243; J. Linn. Soc. Bot. 14 (1875) 228 LILIACEAE—I (Jessop) 215 (‘philippense’); Curtis, Bot. Mag. III, 32 (1876) t. 6250; ELwes, Monogr. genus Lilium (1880) t. 3; Merk. Philip. J. Sc. 5 (1910) Bot. 337; En. Philip. 1 (1922) 204; WiLson, Lilies of E. Asia (1925) 20; DRYSDALE Woopcock & STEARN, Lilies of the World (1950) 311. Bulb subglobose, whitish, c. 3'/,-4 cm @; scales unknown. Stem 30-90 cm high, smooth, green or mottled with purple, producing roots above the bulb. Leaves alternate, 30-40, linear, attenuate, with up to 7 veins of which 1 or 3 are more con- spicuous, 8-17 cm by 2-4 mm. Flowers 1 or 2, white, with green and reddish outside towards the base, horizontal, (10—)14—25 cm long, infundibuli- form, the tube (in dried material) 8-12 mm @. Perianth segments not clawed, spreading distally; nectariferous furrow papillose. Filaments linear in dried material, 13-17 cm; anthers 5-15 mm long; pollen yellow. Style 10'/,-16 cm; stigma deeply 3-lobed. Capsule c. 5 cm long. Distr. Taiwan and Malesia: Philippines: North Luzon (Bontoc, Benguet and Pangasinan Prov.). Ecol. Open grassy slopes in the pine region, 1100-2300 m. FI. May, fr. Oct. Vern. Philippines: kanyon, Ilk., soyasoi, tubtubkau, \g., tuktukpdu, Bon. Note. Except for the presence of papillae on the nectariferous furrow hard to distinguish in flower from L. longiflorum. luplipak, 13. ASPARAGUS LINNE, Sp. Pl. (1753) 313; Gen. Pl. ed. 5 (1754) 147; Baker, J. Linn. Soc. Bot. 14 (1875) 594; Bru. Fl. Austr. 7 (1878) 17; BAKerR, FI. Trop. Afr. 7 (1898) 425; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 362; Hutcn. Fam. FI. Pl. ed. 2, 2 (1959) 608; Jessop, Bothalia 9 (1966) 31; BACK. & BAKH. f. Fl. Java 3 (1968) 92. — Asparagopsis KUNTH, Abh. K. Ak. Wiss. Berlin (1842) 35. Climbing or erect, glabrous or pubescent, softly woody plants usually with bisexual flowers, rarely dioecious. Rhizome perennial; roots mostly thick and often tuberous. Aerial stems annual or perennial, usually much-branched; branches sometimes similar to the main stems and sometimes morphologically distinct. Leaves cauline, alternate, scale-like, usually brown and at least partially scarious, often with a spine from the abaxial surface; leaves of the rhizome scale-like, lacking a spine. Leaf-like structures (cladodes) solitary or fascicled, flat, angled or terete, arising in the axils of cauline leaves but sometimes absent from flower-bearing branchlets. Pedicels solitary or fascicled in the axils of the cauline leaves, articu- lated. Perianth segments free or minutely connate, equal or subequal, 1-veined, white or nearly so, often with a green longitudinal band on the abaxial surface, spreading or less often reflexed. Filaments flattened, attached to the perianth seg- ments; anthers dorsifixed, oblong, introrse. Ovary superior, + sessile, 3-celled; ovules axile, 2 to few; style usually filiform with 3 short branches, less often divided 216 FLORA MALESIANA [ser. I; vol9? nearly to the base. Fruit usually a red 1- to few-seeded berry; perianth not usually persistent. Seeds globose or partly angled. Distr. Widespread in Europe, Africa, Madagascar, and Asia; in Malesia 2 spp., one of which is wide- spread in the Old World and the single one known from Australia. There are probably fewer than 100 spp., but the taxonomy of the genus is poorly understood. Ecol. Species have a preference for arid areas in the open and for savanna, but some occur in damp forests. Notes. The morphology of the spines has been discussed by CusseT & TRAN (Bull. Soc. Bot. Fr. 113, 1966, 151). The nature of the leaf-like organs (cladodes) is controversial. They are most frequently treated as axillary structures, i.e. modified branches (e.g. KAUSSMANN, Bot. Stud. 3, 1955), and my own work confirms this. However, ARBER (Monocotyledons: a morphological study, 1925) believed that in a few species they were in fact leaves, while SCHLITTLER (Bot. Jahrb. 79, 1959, 428) concluded that they are leaves in all species. KEY TO THE SPECIES 1. Flowers unisexual, borne at normal vegetative nodes. Cladodes usually flat, less often triquetrous 1. Flowers bisexual, usually on branches lacking cladodes. Cladodes triquetrous . . 1. Asparagus cochinchinensis (LouR.) MERR. Philip. J. Sc. 15 (1919) Bot. 230; En. Philip. 1 (1922) 206; GAGNEP. FI. Gén. I.-C. 6 (1934) 780; Merr. Comm. Lour. (1935) 108; MaxkIno, Ill. Fl. Japan (1954) 735. — Melanthium cochinchinense Lour. FI. Coch. (1790) 216. — A. lucidus LINDL. Bot. Reg. 30 (1844) Misc. 29; Baker, J. Linn. Soc. Bot. 14 (1875) 605; MerR. & ROLFE, Philip. J. Sc. 3 (1908) Bot. 96, incl. var. dolichocladus MERR. & ROLFE. Plant dioecious. Stems glabrous, erect, procum- bent or climbing, shallowly ridged; branches numerous, uSually simple, arising singly, less than 10(-30) cm long, spreading or erect-spreading, often straight. Roots with distant, elongate tubers. Scale-leaves with scarious portion less than 5(-8) mm long; spines absent on branches, poorly developed or up to 5 mm long on main axes. Cladodes present on branches and towards the ends of stems, flat, or 3-angled, 1—3-nate, linear-arcuate, 5—15(-40) by '/,-11/, mm. Pedicels 1- or 2(-3)-nate, arising from the axils of scale-leaves subtending cladodes, articulated near or above the middle, 2-3(-6) mm long. Perianth segments yellow-green, pale green or white, spreading similar, oblong- elliptic, c. 2-31/, mm long. — ¢ Flowers: filaments shorter than the perianth segments, cuneate from a broad base; anthers nearly 1 mm long (one specimen); ovary rudimentary. — @ Flowers: filaments c. half the length of the perianth seg- ments; anthers rudimentary; ovary obovoid, 1'/,-2 mm long; style c. 1/, mm, with 3 stigmatic ridges; ovules 2 per locule. Berry green when ripe, c. 4-7 mm @. Seeds 1-4, globose or angled, c. 2-3 mm @. Distr. Korea, Japan, Ryu Kyu Is., S. China, Indo-China, Taiwan; in Malesia: Philippines (N. Luzon: Benguet Prov.), one record (LOHER 1928), probably from above 1200 m. Ecol. There are several records from within the spray zone on coral or limestone substrates; also in 1. A. cochinchinensis 2. A. racemosus bush up to at least 200 m, but no ecological details known from the Philippines. Notes. The recorded flower colours may be mis- leading. It is possible that the segments are white with a green or yellow-green band. GAGNEPAIN recorded that the flowers were uni- sexual. I have seen insufficient material to determine whether the plants are always dioecious or to con- firm that the flowers are always functionally unisexual, but in all flowers I examined one sex appeared to be rudimentary. 2. Asparagus racemosus WILLD. Sp. Pl. 2 (1799) 152; BAKER, J. Linn. Soc. Bot. 14 (1875) 623; Bru. Fl. Austr. 7 (1878) 17; ENGL. Bot. Jahrb. 7 (1886) 448; Ripi. Fl. Mal. Pen. 4 (1924) 331; Back. Handb. Fl. Java 3 (1924) 72; HeyNne, Nutt. Pl. (1927) 444; H. Perrier, Fl. Madag. fam. 40 (1938) 21; STEEN. Atlas Trop. Nederl. (1938) map 7*; Jessop, Bothalia 9 (1966) 72; HutcH. & DALz. Fl. W. Trop. Afr. ed. 2, 3 (1968) 93; Back. & BAKH. f. Fl. Java 3 (1968) 93. — A. dubius DECNE, Nouv. Ann. Mus. Paris 3 (1834) 363; Herb. Timor. Descr. (1835) 35; SPAN. Linnaea 15 (1941) 476, added A. penduliflorus ZipP., nomen, in syn. — Asparagopsis decaisnei KUNTH, En. Pl. 5 (1850) 103, nom. illeg.; ZOLL. Syst. Verz. 1 (1854) 67; Mia. FI. Ind. Bat. 3 (1859) 562; RmpL. in Forbes, Wand. (1885) 520. — Asparagopsis schoberioides KUNTH, En. Pl. 5 (1850) 70; IL’mn, Fl. U.S.S.R. 4 (1968) 328. — Asparagopsis javanica KUNTH, En. Pl. 5 (1850) 100; ZoLL. Syst. Verz. 1 (1854) 67; Mia. FI. Ind. Bat. 3 (1859) 562. Plant with bisexual flowers. Stems glabrous, usually climbing, up to 2-3 m high or more, smooth or grooved, in their lower part un- branched; branches numerous and _ branched; solitary; final branches usually 1—4-nate, up to 5(-10) cm long, spreading or ascending, straight. Roots with distant, elongate tubers. Scale-/eaves 1979] with scarious portion up to 5(-10) mm long and spine usually absent on final branches, up to 5(-10) mm long. Cladodes present mainly on branches and towards the ends of stems, triquet- rous, linear-crenate, 1-—3(-4)-nate, (7—)10-25 (40) mm long, rarely over 1 mm broad. Pedicels 1- or 2-nate, usually on branches 2-6 mm long lacking cladodes, less often on normal branches, articulated usually near the middle, 3—S mm long. Flowers bisexual, scented. Perianth segments white with a green band, spreading similar oblong to obovate-oblong, 2-3(-4) mm long. Filaments shorter than or about the same length as the perianth segments; anthers c. 0.2-0.3 mm long. Ovary obovoid, c. 11/, mm long; style c. '/, mm, with 3 stigmatic ridges or 3 short branches; ovules 2 per locule. Berry red when ripe, c. 4-6 mm @. Seeds 1-3, globose or angled, c. 2-3 mm @. Distr. Widespread in Africa including the southern Cape, Guinea and Madagascar, through southern Asia into China, in South Malesia and the northern states of Australia; in Malesia: Malay Peninsula (Langkawi, on limestone rocks), Java (in the western half only on the Northcoast, in E on both sides; also Madura and Kangean Is.), Lesser Sunda Is. (Lombok, Sumba, Sumbawa, Flores, Timor), and SE. Moluccas (Tenimber Is.). The range in Malesia is distinctly disjunct on both ends which is due to its drought preference; LILIACEAE—I (Jessop) 217 it is absent from the Sundaland rain-forest core. Ecol. In Malesia a distinct drought-loving plant and bound to the seasonal areas, in Java to the zone with at most 20 rainy days in the 4 driest consecutive months of the year, mostly in coastal areas, in sunny thickets and on dunes, in monsoon forest with Bambusa spinosa, Acacia leucophloea, etc., predominantly in the lowland, but ascending to c. 1150 m (BACKER). F/. mostly Aug.—Jan. Vern. Christusdoorn, D; bek bun, Chinese; Java: sangga langit, J; Timor: niesie saub, nénésan; Tenimber Is.: skikirie, Saumlak. Notes. There are many synonyms from Africa, Asia and Australia, but none are known to have been used for the Malesian area. A. racemosus is closely allied to A. cochinchinen- sis and can only be distinguished by a combination of characters. In Malesia they are geographically separated but until a revision of the genus, at least in Asia, is undertaken the taxonomy of this group must remain uncertain. Excluded According to MERRILL (En. Philip. 1, 1922, 206) Naves (Nov. App. 1880, 264) has credited A. decli- natus L. and A. racemosus WILLD. to the Philippine flora, but both were apparently admitted on erroneous identifications. 14. DISPORUM SALIsB. Trans. Hort. Soc. 1 (1812) 331; D. Don, Trans. Linn. Soc. Lond. 18 (1841) 513; BAKER, J. Linn. Soc. Bot. 14 (1875) 588; Hook. f. Fl. Br. Ind. 6 (1894) 359; BAcK. Handb. FI. Java 3 (1924) 73; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 368. — Lethea Norona, Verh. Bat. Gen. ed. 1, 5 (1790) Art. 4, 2, nomen. — Drapiezia Bu. En. P|. Jav. (1827) 8. — Fig. 12. Glabrous rhizomatous herbs. Aerial stems annual, erect, branched. Leaves cauline; the lower brown and scale-like with a sheathing base; the others alternate becoming opposite distally, flat, with many veins, petioled, usually ovate, obtuse at base. Peduncles terminating vegetative branches, not articulated, bearing 1-6 nutant flowers in an umbel. Perianth segments free, equal or subequal, erect or distally recurved, several-veined, usually saccate or spurred at the base. Filaments linear to lanceolate, attached to the receptacle or the base of the perianth; anthers dorsifixed, linear-oblong, extrorse. Ovary superior, sessile, ovoid to obovoid, 3-celled; ovules axile, 2 per cell; style filiform, with 3 recurved stigmatic branches. Fruit a 1-3-seeded berry. Distr. Approximately 30 spp. currently recognized in the western U.S.A. and Canada, Japan, China, northern Deccan Peninsula, through Thailand to West Malesia: Malay Peninsula, Sumatra, Java, and Bali. In my opinion it is highly unlikely that more than half of these will be recognized after critical revision. Ecol. Typical forest plants. Note. BACKER /.c. pointed to the peculiar sympodial structure of the stem. 218 FLORA MALESIANA [ser. Dvor 1. Disporum cantoniense (LouR.) MERR. Philip. J. Sc. 15 (1919) 229; MerR. Comm. Lour. (1935) 109. — Fritillaria cantoniensis Lour. Fl. Coch. (1790) 206. — Uvularia chinensis KER-GAWL. Bot. Mag. (1806) t. 916. — D. pullum Sautss. Trans. Hort. Soc. 1 (1812) 331; Hassk. Pl. Jav. Rar. (1848) 105; Mia. FI. Ind. Bat. 3 (1859) 552; BAker, J. Linn. Soc. Bot. 14 (1875) 589; Hook. f. Fl. Br. Ind. 6 (1892) 360; Rip. J. Fed. Mal. St. Mus. 4 (1909) 82, incl. var. multiflorum RIDL.; Koorp. Fl. Tjib. 1 (1922) 47; Ripv. FI. Mal. Pen. 4 (1924) 338; HEND. Mal. Wild Flow. Monoc. (1954) 185, f. 109; CHAROENPHOL, Thai For. Bull. 8 (1974) 89. — Drapiezia multiflora BL. En. Pl. Jav. (1827) 8; (1974)89. — Drapiezia multiflora Bv. En. Pl. Jav. (1827) 8; JUNGH. Java ed. 2 (neerl.) 1 (1853) 522; Zou. Syst. Verz. 1 (1854) 66, incl. var. albiflora Zoi. — D. leschenaultianum D. Don, Proc. Linn. Soc. 1 (1839) 45; Trans. Linn. Soc. 18 (1841) 518; Merr. Contr. Arn. Arb. 8 (1934) 19. — D. hors- fieldii D. Don, Proc. Linn. Soc. 1 (1839) 45 (WALLICH 5088D). — Streptopus multiflorus (BL.) Fig. 12. Disporum cantoniense (LOUR.) MERR. a. Habit, x 3/4, b. flower of the spurred form, c. ditto of the non-spurred form, both x 2!/,, d. gynoecium, e. stamen, both x 5, /f. fruit, x 11/, (drawn from various collections). 1979] D. Dretr. Syn. PI. 2 (1840) 1121. — D. multiflorum (BL.) D. Don, Trans. Linn. Soc. 18 (1841) 518; Mia. FI. Ind. Bat. 3 (1859) 552. — D. calcaratum D. Don, Trans. Linn. Soc. 18 (1841) 516; BAKER, J. Linn. Soc. Bot. 14 (1875) 588; Hook. f. Fl. Br. Ind. 6 (1892) 359; CHAROENPHOL, Thai For. Bull. 8 (1974) 89. — Uvularia multiflora (BL.) KUNTH, En. Pl. 4 (1843) 207. — D. chinense (KER-GAWL.) O. K. Rev. Gen. Pl. 2 (1891) 708; BAck. Handb. Fl. Java 3 (1924) 73; Docr. v. LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. II, 31 (1933) 147; BACK. & BAKH. f. Fl. Java 3 (1968) 94. — Fig. 12. Stems erect, often 45-80 cm high, up to 2!/, m, branched only in the upper half; branches erect- spreading, becoming flexuose distally. Roots fibrous or slightly thickened but not tuberous. Leaves usually ovate, less often oblong or lanceo- late, rarely suborbicular, acuminate, 6'/,-17 by 2'/,-9 cm, with 5-9 veins prominent on the lower surface and numerous finer veins; petiole to 8 mm. Inflorescence an umbel of 2-7 flowers terminating a short branch which arises opposite a leaf and bears a single sometimes somewhat reduced leaf; peduncle 0-3!/, cm; pedicels 1-4 cm. Perianth segments lanceolate to oblanceolate, acute, keeled below and usually saccate or less often with a spur to 5 mm long at the base, white, greenish to purple, 1—2(-2'/,) cm long. Filaments linear to lanceolate, often less than half as long as the LILIACEAE—I (Jessop) 219 perianth; anthers bright yellow, 2'/,-4 mm long. Berry bluish black, 7-10 mm @. Seeds c. 3 mm @. Distr. SE. Asia from the northern Deccan Peninsula to southern China and Japan; in Malesia: Malay Peninsula (Perak, Pahang) and throughout Sumatra, Java, and Bali. Ecol. In both primary and secondary forest, rarely in the open, (700—)1000-2550 m; especially common in W. Java. F/. Jan._Dec. DOCTERS VAN LEEUWEN /.c. observed the flowers to be proto- gynous; self-pollination is possible but bumblebees also regularly visit the flowers. Vern. Java: baradja_ lintang, kamalakian, kibeunteur areuj, (ki)tamiang, lili leaweung, radja lintang, tangkal milon, S, égér égér, glinggangan, lenguk, srintil, tombagan, J; N. Sumatra: si- demsapo, sumbul sumbul, ‘Karo-Batak, kayu (si-mar)soma-soma, S. Notes. Variation in the length of the spur has been used in segregating species, but BACKER recognized the continuity of this gradation in Javanese material. Some Javanese plants have spurs as long as those in typical “‘D. calcaratum’. Branching is partly sympodial. Short terminal axes bear the umbel. Continuation of growth of the aerial shoot is by a branch arising in the axil of the second leaf below the umbel. The node separating two leaves associated with the inflorescence is sometimes so short that the leaves appear opposite. 15. DISPOROPSIS HANCcE, J. Bot. 21 (1883) 278; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 370; Nakal, J. Jap. Bot. 12 (1936) 147; CHANG & Hsu, Taiwania 19 (1974) 64. — Fig. 13. Glabrous rhizomatous herbs. Aerial stems annual, erect, unbranched. Leaves cauline, alternate, flat, with many veins, petioled, entire, ovate to oblong, obtuse or subcuneate at the base; brown scale-like leaves on the rhizome, less often also at the base of the aerial stems. Flowers solitary in the axils of the leaves, nutant. Pedicels articulated. Perianth campanulate; segments 3-5, fused, equal, viscid, saccate at the base. Filaments expanded to form a corona attached to the perianth; anthers dorsifixed, sagittate, introrse. Ovary superior, sessile, ovoid, 3-celled ; ovules axile, 4-6 per cell; style short and thick, with simple or 3-lobed stigma. Fruit a 1—5-seeded berry. Distr. Probably 4 or 5 spp. from Thailand, Indo-China, southern China, Taiwan; in Malesia: Philip- pines. Ecol. Forests, usually in the mountains. 1. Disporopsis fusco-picta HANCE, J. Bot. 21 (1883) 278. — Disporum pullum (non SA.isB.) MERR. Philip. J. Sc. 1 (1906) Suppl. 182. — Disporum luzoniense MeERR. Philip. J. Sc. 5 (1910) Bot. 338; En. Philip. 1 (1922) 206. — Fig. 13. Stems erect, 25-45 cm. Roots fibrous or slightly thickened but not tuberous. Leaves 6-9, ovate to oblong, obtuse or subcuneate at the base, acumi- nate, 6-12!/, by 2!/,-5 cm, with usually 7 veins prominent on the lower surface and numerous finer veins; petiole (3—)5—12 mm. Flowers in the axils of the lower leaves. Pedicels 1-2'/, cm. Perianth seg- 220 [ser. I; vols 9? FLORA MALESIANA Fig. 13. Disporopsis fusco-picta HANCE. a. Habit, x 1/3, b. flower, c. ditto, laid open showing corona and stamens, both x 2!/,, d. fruit, x 2 (a—c STEINER 2147, d PNH 7471). ments fused below middle, 13-15 by 2-3 mm, lan- ceolate, obtuse, shallowly saccate at the base, white with (always?) dull purple inside. Corona attached close to perianth sinus, c. 4 mm long, its lobes emarginate. Anthers sessile on the corona, c. 1 mm long, very shallowly sagittate. Berry becoming purple, blue or black, c. 1 cm @. Seeds c.4 mm @. Distr. Southern China, Taiwan, in Malesia: Philippines (N. Luzon: Lepanto, Bontoc and Benguet Prov.). Ecol. Mossy forest, 1700-2500 m, with one record of association with (the secondary pyro- genous savanna of) Pinus insularis. Fl. rarely recorded: May-June, or later. 16. TRICALISTRA RIDL. J. Fed. Mal. St. Mus. 4 (1909) 83; Fl. Mal. Pen. 4 (1924) 330. — Tupistra sensu HUTCH. Fam. FI. Pl. ed. 3 (1973) 749, in part. — Fig. 14d. Stemless herbs. Rhizome horizontal, woody; roots thick and fleshy. Leaves large, basal, caespitose, petioled, expanding towards the base to form a sheath, with a strong main vein and numerous finer veins. Jnflorescence a spike. Flowers numerous, each in the axil of a bract. Perianth segments 6, fleshy, fused for about half their length, campanulate, equal. Stamens 6, attached to the perianth; anthers sub- sessile, dorsifixed, oblong, dehiscing introrse-laterally. Ovary superior, sessile, subcylindrical, 3-lobed, 1-celled, containing (2 or) 4 discord ovules but with traces of two further carpels; stigmas 3, hippocrepiform, sessile. Fruit a drupe, green when unripe, globose, 1|-seeded. Distr. Monotypic. Malesia: Malay Peninsula. Note. Regarded as a synonym of Tupistra by HUTCHINSON and recognized, but with some doubt, by Airy SHAW (Willis Dict. ed. 8, 1973). The only consistent difference is that Tricalistra has no style, con- sistently 6 stamens, and a 3-lobed stigma, a set of characters of equal standing as those separating other genera in the Aspidistreae. 1979] 1. Tricalistra ochracea RIDL. J. Fed. Mal. St. Mus. 4 (1909) 83; Fl. Mal. Pen. 4 (1924) 331. — Fig. 14d. Scale-like leaves several, sessile, lanceolate to 5 cm long. Foliage /eaves few, broadly lanceolate to oblanceolate, acuminate, cuneate at the base; dark green, chartaceous, 30-40 cm long when in flower, lengthening in fruit, 8-11 cm broad; petiole poorly differentiated, sometimes winged, 12-20 cm. Inflorescence 12-15 cm long when in flower, lengthening in fruit, with 25-35 flowers. Bracts ovate, obtuse, caducous, to 3 mm long. Perianth fleshy, c. 5 mm long, lobes as long as the tube, recurved, ovate, acute, dull ochreous yellow. LILIACEAE—I (Jessop) 221 Anthers united below the mouth of the perianth tube, thecae divaricate at base, less than 1 mm long. Ovary c. 3 mm long. Fruit to c. 2 cm long. Distr. Malesia: Malay Peninsula (Pahang: Cameron Highlands). Ecol. One collection annotated ‘on rocks in open places’, 1000 m. F/. Nov. (one record), fr. April. Note. Syntypes collected by RrmpLey (13692) are in SING and K. RIDLey referred to the leaf having 6 veins. However, leaves on both type specimens have at least 100 of which about 13 are more conspicuous than the others. The only other collec- tion (Nur SF 32725) agrees with the type. 17. TUPISTRA KER-GAWL. Bot. Mag. 39 (1814) t. 1655; BL. Tijd. Nat. Gesch. Phys. 1 (1834) 67; BAKER, J. Linn. Soc. Bot. 14 (1875) 580; Hook. f. Fl. Br. Ind. 6 (1892) 324; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 372. — Fig. 14a-—c. Stemless herbs. Rhizome tuberous or horizontal, thick and woody; roots thick and fleshy. Leaves large, basal, scattered or distichous, usually distinctly petioled, expanding towards the base to form a sheath, with a strong main vein and numer- ous finer veins. Inflorescence a spike. Flowers numerous, each in the axil of a bract. Perianth segments 6 or 8, fleshy, fused, campanulate, equal. Anthers sessile or sub- sessile, inserted in tube, dorsifixed, oblong or ovoid, introrse. Ovary superior, sessile, subglobose or not externally differentiated from style, 3(—4)-celled; ovules 2 in each locule; style cylindrical; stigma large, peltate or capitate, entire or variously lobed. Fruit a globose berry, usually 1-seeded; seeds turgid; perianth persisting below the fruit. Distr. Eastern Himalayas to southern China; in Malesia: Malay Peninsula and Sumatra. The type of 7. squalida (the type species) was described, undoubtedly in error, from Amboina. About 25 spp. have been described, but many of these should probably be reduced to synonymy. Ecol. Most frequently recorded from dry evergreen forest, often in deep shade and often from near streams. Syst. It appears to me that in the Aspidistreae too many small genera are distinguished on insignificant characters. Note. The Malesian species have been insufficiently collected for convincing taxonomic judgements to be made. KEY TO THE SPECIES 1. Spike to 17 cm long; style and ovary up to 7 mm long; stigma usually 1'/,--3 mm @ . 1. T. violacea 1. Spike at least 20 cm long; style and ovary at least 7'/, mm long; stigma at least 4"/, mm @ 1. Tupistra violacea Rip-. J. Str. Br. R. As. Soc. n. 41 (1904) 35; Mat. Fl. Mal. Pen. Monoc. 2 (1907) 93; Fl. Mal. Pen. 4 (1924) 330. Leaves few, elliptic to oblanceolate, acute or acuminate, cuneate at the base; lamina 50-70 by 7T'/,-13 cm; petiole well-defined, to 40 cm. Jnflores- cence ascending to 17 cm long when in flower, lengthening in fruit, with 30-40 flowers. Bracts somewhat amplexicaul ovate, c. 5 by 3-8 mm. 2. T. grandis Perianth segments violet; the tube 5-6 mm long, 41/,-6 mm wide; lobes darker than tube, oblong to ovate, 4—5!/, by 3-4 mm. Anthers sessile, attached in tube or throat, c. 1—-1'1/, mm long. Style and ovary white, spotted violet, 3'/,-7 mm long; stigma 11/,-4 mm @, obscurely lobed. Distr. Malesia: Malay Peninsula (Perak: Bujong Malacca; Penang Highlands); probably also in Thailand. 222 FLORA MALESIANA [ser. I, vol. 91 Fig. 14. Tupistra grandis Rw. a. Habit, < '/,, b. flower from outside, c. flower laid open, both x 2. — Tricalistra ochracea RiDL. d. Flower, halved, showing gynoecium, 3 tepals, to which stamens are attached, x 10 (a—c LORZING 8753, d RIDLEY 13692). 1979] Ecol. Mountain forest, apparently rare and no collections made since 1901 have been seen. Fi. March, Dec. Note. This species closely resembles 7. squalida, the type species of the genus, from the Himalayas. Further study is needed to confirm that they should be kept separate. In the absence of recent collec- tions much of the information here is taken from Rim.ey (1924). The scale-like leaves recorded for T. gracilis are likely to occur in 7. violacea, but the basal part of the plant has not been preserved. 2. Tupistra grandis RIDL. J. Bot. (1900) 73; Mat. Fl. Mal. Pen. Monoc. 2 (1907) 93; Fl. Mal. Pen. 4 (1924) 330; B. M. ALLEN, Mal. Nat. J. 19 (1966) 303. — T. perakensis Nicnots. Ill. Dict. Gard., Cent. Suppl. (1901) 722. — Fig. 14a-c. Leaves few, surrounded by several sessile lanceo- late scale-like leaves to 25(—40) cm long, elliptic or sublinear to oblanceolate, acute or acuminate, cuneate at the base; lamina (50—)70-150 by 6— 14 cm; petiole often winged and poorly defined, to 40 cm. Inflorescence erect, 20-50 cm long when in flower, lengthening slightly in fruit, with up to 100 or more flowers, with a strong smell. Bracts somewhat amplexicaul, ovate, 5-10 by 2-6 mm. Perianth segments violet to dark purple; tube 5-11 mm long, 5-10 mm wide; lobes darker than tube, oblong to ovate, 4-14 by 3-5 mm. Anthers sessile, attached near top of tube, 1!/,-2 mm long. Style and ovary white, 7!/,-16 mm long; stigma 4'/,-12'/, mm @, peltate, flat to biconvex, some- times rugulose or irregularly lobed. Distr. Malesia: Malay Peninsula (Perak, Kelantan, Pahang, Selangor, Langkawi) and N. Sumatra (Tapanuli Res.: Karo Highlands); probably also in Thailand. Ecol. In Malaya records suggest that it usually occurs in wet rocky places on limestone. In Sumatra it grows in forest at 600-1225 m, but in LILIACEAE—I (Jessop) 223 Malaya there is a record of 150 m in Perak. FI. Jan.—Dec. Vern. N. Sumatra: singkut antu, Karo-Batak. Note. The Sumatran specimens tend to have longer spikes and larger flowers than the Malayan material. No characters have, however, been found on which to base taxonomic separation. Excluded Tupistra singapureana [WALL. Cat. n. 5195]; BAKER, J. Linn. Soc. Bot. 14 (1875) 581; Hook. f. Fl. Br. Ind. 6 (1892) 325, was omitted from the genus by S. Kurz (J. As. Soc. Beng. 44, ii, 1875, 199) and has indeed appeared not to belong to Liliaceae. It was referred by RIDLEY (Mat. Fl. Mal. Pen. Monoc. 1, 1907, 232) to Neuwiedia curtisii ROLFe and by ROLFE (Kew Bull. 1907, 412) to Neuwiedia singapureana (BAKER) ROLFE. According to DE VoGEL (Blumea 17, 1969, 331) = Neuwiedia zollingeri RCHB. var. singapureana (BAKER) DE VOGEL (Orchidaceae). Tupistra squalida Ker-GAwL. Bot. Mag. 39 (1814) t. 1655; E>warps, Bot. Reg. (1823) t. 704; LoppiGcEs, Bot. Cab. 6 (1821) t. 515; BL. Tijd. Nat. Gesch. Phys. 1 (1834) 67, t. IIIC; Mia. FI. Ind. Bat. 3 (1859) 569; Baker, J. Linn. Soc. Bot. 14 (1875) 580; cf. Hook. f. Fl. Br. Ind. 6 (1892) 324, in nota sub T. nutans. — Rhodea tupistra SCHULT. Syst. 7, 2 (1829) 173, nom. illeg. The provenance of this species, the type of the genus, was given as ‘Amboyna’. This is certainly erroneous. It was described from a cultivated plant in the nurseries of LoppIGEs, and was obviously in the former century cultivated in several botanic gardens. The more curious it is that its proper identity and native country remains more or less uncertain. BAKER /.c. reduced 7. nutans WALL. (Bot. Reg. t. 1333) from India to this species, but Hooker f. kept these two entities apart. 18. LIRIOPE Lour. Fl. Coch. (1790) 200; L. H. Bamey, Gent. Herb. 2 (1929) 3; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 376; WANG & TANG, Act. Phytotax. 1 (1951) 331; Hume, Baileya 9 (1961) 135. — Ophiopogon (non KER-GAWL.) KUNTH, En. Pl. 5 (1850) 297, in part. — Fig. 16d. Stemless or shortly caulescent, rhizomatous herbs. Roots thick, sometimes with tuberous swellings. Leaves usually basal, petioled or subpetioled linear to oblanceo- late or lanceolate, many-nerved. Inflorescence a spike or raceme. Flowers solitary or fascicled in the axil of each bract. Perianth segments fleshy, campanulate, free or with a broad tube and short lobes, equal, with 1 vein, violet or white. Anthers 6-8, subsessile or pedicels short, attached near base of perianth, dorsifixed, in- trorse. Ovary superior, sessile, ovoid to subglobose, 3—4-celled; ovules axile, 2 in [ser. I, vol. 9! 224 FLORA MALESIANA SS \ \? FF SSS TT — = “= =— S —- : as : AS =—_ = SSS ——— — Se, q's “(Ce a e AN Gr’ SF — = = —— iS = . i Fi \ \ \ = In forest above mountain garden Tjibodas, Febr. 1936. Fig. 15. Ophiopogon caulescens (BL.) BACK. Habit, with seed, x 1/4. 1979 | 225 LILIACEAE—I (Jessop) each locule; style short and thick; stigma capitate or peltate. Ovary wall rupturing early in the growth of the seeds which are therefore exposed during most of their development. Seeds black, globose or slightly elongate, with fleshy testa; perianth persistent. Distr. About 5 spp., in Japan, China, Indo-China and North Malesia: N. Philippines. Note. Some species are widely grown as ornamentals. 1. Liriope graminifolia (L.) BAKER, J. Linn. Soc. Bot. 17 (1879) 499; Merr. En. Philip. 1 (1922) 207; Hume, Baileya 9 (1961) 150. — Asparagus gramini- folius LINNE, Sp. Pl. ed. 2 (1762) 450. — Dracaena graminifolia (L.) LINNE, Syst. Nat. ed. 12 (1767) 275..— ? L. spicata Lour. FI. Coch. (1790) 201; L. H. BaiLey, Gent. Herb. 2 (1929) 33; MerR. Comm. Lour. (1935) 109; Hume, Baileya 9 (1961) 150, 152, 158. — ? Ophiopogon spicatus (LouR.) Ker-GAWL. Bot. Reg. 7 (1821) t. 593; Naves, Nov. App. (1880) 264. — Mondo graminifolia (L.) Komwz. Tokyo Bot. Mag. 40 (1926) 333. — L. muscari [non (DECNE) L. H. BAILEY] HaAtus. Mem. Fac. Sc. Kagoshima Un. 5, 3 (1966) 62. — Fig. 16d. Rhizome horizontal, slender, moderately woody. Roots bearing distant tubers. Leaves basal, caespi- tose, linear to narrowly linear-oblanceolate, minutely denticulate on the margins, the central vein sometimes conspicuously larger than the others, expanded to form membranous wings towards the base, 25-90 cm long, 2-9 mm broad. Peduncle erect, (12—)30-50 cm long, shorter than leaves. Bracts deltoid, to 4 mm long. Pedicels 2—5-nate, 2-12 mm long, articulated at the base of the flower. Leafy shoots occasionally produced from axil of bracts on inflorescence. Perianth segments free, 3'/.-4 mm long, violet. Filaments to 2 mm long; anthers c. 1 mm long. Seeds oblong (perhaps only when young) or globose, c. 5 mm long. Distr. Japan, China and North Malesia: Philippines (Batan Is.; Mindoro and Luzon). The paucity of collections suggests that this species is rare in the Philippines. Ecol. Open slopes at c. 1400 m (MERRILL, /.c.), but obviously in the Batan Is. at low altitude. Notes. The differences between L. graminifolia and L. spicata are not clear. HUME depended for their separation largely on the conspicuous mem- branous basal wings to the leaves of the former, associated with quantitative characters of the leaves and inflorescence. L. graminifolia is likely to remain the correct name for the Philippine species whether L. spicata is treated as a synonym or not. L. muscari (DECNE) L. H. BAILey (Gent. Herb. 2, 1929, 35) differs according to HUME by caespitose habit and stiffer and wider leaves (8-26 mm). 19. OCPHIOPOGON KER-GAWL. Bot. Mag. 27 (1807) t. 1063; Hook. f. Fl. Br. Ind. 6 (1892) 267; RIDL. Fl. Mal. Pen. 4 (1924) 326; RODRIGUEZ, Bull. Soc. Bot. Fr. 75 (1928) 997; FI. Gén. I.-C. 6 (1934) 655; BACK. & BAKH. f. Fl. Java 3 (1968) 95, nom. gen. cons. — Mondo ADANS. Fam. 2 (1763) 496; FARWELL, Amer. Midland Nat. 7 (1921) 41; L. H. BAILey, Gentes Herb. 2 (1929) 17; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 377; OHwi in Fedde, Rep. 36 (1934) 45. — Flueggea Ricu. Neues J. Bot. 2 (1807) 8 (‘Fluggea’); BAKER, J. Linn. Soc. Bot. 17 (1879) 500. — Chloopsis BL. En. Pl. Jav. (1827) 14; HAssk. Flora 34 (1851) 481. — Fig. 15, 16a—c. Stemless herbs. Rhizome sometimes very short; roots fibrous or fleshy, sometimes tuberous. Leaves basal, linear or petioled with a broad lamina, expanded at the base to form a conspicuous scarious sheath. Inflorescence a raceme. Pedicels 1—several- nate, in the axils of bracts and usually associated with a few smaller bracts also apparently in the axil of the principal bract, articulated usually in the distal half. Perianth segments free, equal, with 1 vein, spreading or campanulate, white or violet. Filaments often connate, short, thick, glabrous, attached to the base of the perianth segments or to the receptacle; anthers basifixed, linear-oblong, dehiscing introrsely. Ovary superior to inferior, 3-celled; ovules basal, 2(—6) in each locule; 226 FLORA MALESIANA [ser. I, vol. 9! style ooniditing: sitely 3-lobed. Ovary wall rupturing early in the growth of the seeds which are therefore exposed during most of their development. Seeds blue, often globose, with fleshy testa; perianth often wholly or partly persistent. Distr. India through to southern China to Thailand, Indo-China, Japan and Taiwan; in Malesia: Malay Peninsula, Sumatra, Java, Borneo, and Philippines. About 70 spp. of Ophiopogon have been described, but I doubt whether more than a third of these should be recognized. Ecol. In forest. Notes. RIDLEY recognized O. malayanus, O. intermedius and O. prolifera from the Malay Peninsula. No fertile material has been seen from this area and the only specimens seen (CORNER SF 37872, K & L, and SozEPADMO & MAHOMUD 1214, K) do not differ significantly from O. caulescens. No Malayan material identified as O. intermedius or O. prolifera has been seen. Riptey characterized O. prolifera by having the filaments connate, and distinguished O. intermedius from O. malayanus on its larger flowers (12 mm wide). He did not give a comparable figure for O. malaya- nus but recorded that the segments were 3 mm long. The descriptions could all be of varieties of O. caulescens, falling within the morphological range known for that species from Java, except that the shortest perianth recorded for Java material is 4 mm long (almost 50% greater than the length recorded for O. malayanus). O. malayanus has also been recorded from the Philippines and Borneo (MERRILL, 1922). Material from continental Asia of O. intermedius resembles O. japonicus, but no material has been seen from the Malay Peninsula. KEY TO THE SPECIES 1. The broadest leaves on each plant more than 4 mm broad. Rhizome well-developed. Lowest bracts 12-25 mm long . 1. O. caulescens 1. The broadest leaves one each plant less than 3 mm 1 broad. Rhizome poorly developed. Lowest bracts 5-10 mm long 1. Ophiopogon caulescens (BL.) BACK. Handb. FI. Java 3 (1924) 74; VAN HELTEN, Med. Alg. Proefsta- tion Landb. n. 16 (1924) 49 (‘gauliscens’); RIDL. J. Bot. 43 (1925) Suppl. 122; Doct. v. LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. II, 31 (1933) 252; BAcK. & BAKH. f. Fl. Java 3 (1968) 95. — ? O. inter- medius D. Don, Prod. Fl. Nep. (1825) 48; RipL. J. Fed. Mal. St. Mus. 4 (1909) 81, incl. var. macran- thum RIDL.; Fl. Mal. Pen. 4 (1924) 327. — Chloop- sis caulescens BL. En. Pl. Jav. (1827) 14; Zo. Syst. Verz. 1 (1854) 67; Mia. Fl. Ind. Bat. 3 (1859) 553. — Chloopsis acaulis Bu. En. P|. Jav. (1827) 14; Mia. Fl. Ind. Bat. 3 (1859) 553. — ? O. prolifera LINDL. J. Hort. Soc. 1 (1846) 76; Mia. FI. Ind. Bat. 3 (1859) 568; RipL. Mat. Fl. Mal. Pen. Monoc. 2 (1907) 91. — Fluggea wallichiana KuntH, En. PI. 5 (1850) 303. — O. wallichianus (KUNTH) Hook. f. Fl. Br. Ind. 6 (1892) 268; Rip. J. Fed. Mal. St. Mus. 8 (1917) 118. — O. malayanus Ri . J. Str. Br. R. As. Soc. n. 41 (1904) 34; Mat. Fl. Mal. Pen. Monoc. 2 (1907) 91; Merr. En. Born. (1921) 115; En. Philip. 1 (1922) 207; Contr. Arn. Arb. 8 (1934) 19. — Mondo malayanum (RiIDL.) FARWELL, Amer. Midland Nat. 7 (1921) 42. — O. japonicus (non (L. f.) KER-GAWL.) Koorpb. FI. Tjib. 1 (1922) 47. — O. acaulis (BL.) Rw. J. Bot. 63 (1925) Suppl. 122. — Fig. 15, 16b. Rhizome rather woody and well-developed, sometimes supported by several thick prop-roots; roots thick but not bearing tubers. Leaves numer- 2. O. japonicus ous, linear to linear-oblanceolate, often slightly arcuate, subacute or narrowly obtuse, often with a prominent midrib, glaucous (waxy), or with glaucous stripes, on the lower surface, (15-)25- 55(-65) cm by 3-10(—22) mm. Peduncle filattened, 10-35(-46) cm long. Flowers 3-12, + secund. Lower bracts 12-25 mm long. Pedicels solitary or, less often, 2-nate, spreading or erect-spreading, articulated often near the centre, 3-8 mm. Perianth segments white or violet, oblong or ovate- or elliptic-oblong, usually obtuse, the inner often slightly smaller than the outer, free segments (5-)7—7'/, by 2-3 mm. Anthers (2-)4—5!/, mm long; filaments connate at the base, c. 1/.-2 mm, up to 1 mm broad at the base. Ovary inferior or semi- inferior; style terete, linear-obconic, (41/,-)5— 7(-9) mm, simple or minutely trifid; ovules 2 per locule. Seeds up to 6, glossy blue, globose or slightly ovoid or ellipsoid, 4-8 mm long. _ Distr. Continental SE. Asia and West Malesia: Malay Peninsula (also Langkawi), Sumatra, all over Java, Sabah (Kinabalu area), S. Philippines (Sulu Is.: Jolo). Ecol. Generally a forest species reported from ‘rain forest’ and ‘primary forest’. Also recorded from screes, (650—)1000-2000 m. Fi. Jan.—Dec. DOCTERS VAN LEEUWEN /.c. stated that self-pollina- tion is the rule. Vern. Java: Murut dial. suket alank, J; Sabah: ryran, 1979] Note. If it is shown that O. intermedius is synonymous with O. caulescens, it will become the correct name. 2. Ophiopogon japonicus (L. f/f.) KER-GAWL. Bot. Mag. 27 (1807) t. 1063; Merr. Philip. J. Sc. 1 (1906) Suppl. 35; ibid. 5 (1910) Bot. 338; En. Philip. 1 (1923) 207; Hume, Baileya 9 (1961) 142. — Convallaria japonica LInnE f. Suppl. (1781) 204. — Mondo japonicum (L. f.) FARWELL, Amer. Mid- land Nat. 7 (1921) 42. — O. merrillii MAsAm. Bull. Soc. Bot. Fr. 84 (1937) 90. — Fig. 16a. Rhizome poorly developed; roots fibrous with distinct tubers usually less than 1'/, cm long and 4 mm broad (not present in all herbarium material even where roots are well represented). Leaves numerous, linear, usually more or less straight, acute or subacute, usually minutely denticulate on vein and margin, usually with a distinct midrib, glaucous (waxy) on the lower surface, 18-60 cm by 1!/,-3(—31/,) mm. Peduncle (10-)12-35 cm. Lower LILIACEAE—I (Jessop) 227 bracts 5-1Q mm long. Pedicels solitary, spreading or erect-spreading, articulated near or below the middle, 2'/,-3 mm. Perianth segments white or violet, oblong-elliptic or ovate, obtuse, the inner slightly smaller than the outer, free segments, 3'/,-4 by 2-3 mm. Anthers 2-2'/, mm long; filaments connate at the base, c. '/, mm long and up to '/, mm broad at the base. Ovary inferior or semi-inferior; style terete, sublinear, c. 3 mm, simple; ovules 2 per locule. Seeds up to 6, glossy blue, globose or slightly ellipsoid, c. 4 mm long. Distr. Japan and North Malesia: Philippines (Luzon: Benguet Prov.; Mindanao: Mt Apo). Ecol. Chiefly in mossy forest, rather common in the Mountain Province, 850—2400(—2900) m. Fi. May-July, fr. Dec.—Jan. Vern. Philippines: Jangigit, Ig., takadu, Bon., uli-uli, Bag. Note. A commonly cultivated garden plant in Java and other places in Malesia, especially for lining borders, but never flowering in the lowland. 20. PELIOSANTHES ANDR. Bot. Repos. 10 (1810) t. 605; HAssk. Tijd. Nat. Gesch. Phys. 10 (1843) 121 (‘Piliosanthes’); BAKER, J. Linn. Soc. Bot. 17 (1879) 503; Hook. f. Fl. Br. India 6 (1892) 265; RipDL. Fl. Mal. Pen. 4 (1924) 323; RopRIGUEZ, FI. Gén. I.-C. 6 (1934) 668; Jessop, Blumea 24 (1976) 141. — Teta Roxs. [Hort. Beng. 1814, 24, nomen] Fl. Ind. ed. Carey 2 (1832) 165. — Bulbisperma REINW. ex BL. Cat. (1823) 59, nomen. — Bulbospermum BL. En. Pl. Jav. (1827) 15. — Lourya BAILL. Bull. Soc. Linn. Paris 1 (1888) 743. — Neolourya RODRIGUEZ, Bull. Mus. Hist. Nat. Paris II, 6 (1934) 96. — Fig. 16e-¢. Erect, stemless, perennial herbs. Rhizome usually very short and horizontal; roots thick. Leaves basal, usually distinctly petioled, the blade sublinear to ovate or obovate, many-nerved. Peduncles flattened, at least at the base, erect, usually soli- tary. Pedicels and inflorescence surrounded at the base by scarious scale-like leaves. Inflorescence a simple raceme. Pedicels articulated. Flowers 1—6-nate in the axils of each bract. Perianth segments fleshy, campanulate or subglobose, fused below, equal, l-veined, white, green, blue, violet or purple. Anthers sessile, attached to a short annular tube (corona) arising from the perianth, introrse. Ovary superior to inferior, 3-celled; ovules basal, 2—5 in each locule; style simple, conical to cylindri- cal; stigma capitate or undifferentiated. Ovary wall rupturing early in the growth of the seeds which are therefore exposed during most of their development. Seeds blue, ellipsoid to pyriform, with fleshy testa; perianth often persistent. Distr. Monotypic, widespread in continental SE. Asia, from the southern Deccan to NE. India and southern China, in West Malesia: Malay Peninsula, Sumatra, Java, Lesser Sunda Is. (Sumbawa) and Borneo. Ecol. Usually in forest, from the lowland to the mountains. Note. In the past far too many species were described. For an account of the taxonomic problems on specific delimitation, see Jessop (1976). 228 FLORA MALESIANA [ser. I, vol. 9! Fig. 16. Ophiopogon japonicus (L. f.) KER-GAWL. a. Habit, x 1/4. — O. caulescens (BL.) BACK. b. Flower, x 2.— Ophiopogon sp. c. Mature seeds, x 2. — Liriope graminifolia (L.) BAKER. d. Flower, x 4. — Peliosanthes teta ANDR. ssp. humilis (ANDR.) Jessop. e. Habit, x */,, f. flower, perianth shown reflexed to reveal corona, x 3, g. LS of flower to show attachment of corona and position of ovary, x 7 (a, c SWINBURNE S.n. ex Hort. Adelaide, b after STEEN. Mt. FI. Java pl. 28: 3b, d largely after E. & P. Nat. Pfl. Fam. ed. 2, 15a, fig. 153 ‘O. spicata’, e-g SORENSEN c.s. 2960). 1. Peliosanthes teta ANDR. Bot. Repos. 10 (1810) t. 605; BAKER, J. Linn. Soc. Bot. 17 (1879) 505; Hook. f. Fl. Br. Ind. 6 (1892) 265; Rip. Mat. FI. Mal. Pen. Monoc. 2 (1907) 88; Fl. Mal. Pen. 4 (1924) 323; RopriGcugz, Fl. Gén. I.-C. 6 (1934) 669; Jessop, Blumea 24 (1976) 154. For synonyms see under the subspecies. Leaves (2-)4~-8(-12); leaf-blades (7'/,-)12'/,- 47/2 by 1*/,-8/,(-11'/.) cm; petioles (4~)7!/,- 50 cm long, slightly compressed. Peduncles to 35(-75) cm high; lower sterile bracts 0-4(-15), to 15(—40) mm long; fertile bracts to 15(—30) mm long, smaller towards the apex of the raceme. Flowers 1-6-nate. Pedicels 1-6(-10) mm long, enlarging after flowering; articulation usually close to flower. Perianth segments suborbicular to linear, 11/,- 6(—8) mm long. Corona forming a disk c. 3-4 mm 2, entire or 6-toothed. Anthers usually rather closely adpressed to the style, c. 1/2 mm long. Ovary most frequently semi-inferior; style often 3- or 6-ridged or fluted, */,-1(-2) mm long. Seeds up to 10-12 mm long. Distr. Tropical SE. Asia; in Malesia: Malay Peninsula, Sumatra, Java, Lesser Sunda Is. (Sumbawa), Borneo. Ecol. Primarily in wet evergreen forest, from 0-—3000 m above sea-level. A few records indicate that dry areas are also occasionally occupied, possibly i in wet enclaves. The subspecies eysal to grow in similar habitats. Note. Grown as a garden ornamental or pot plant. a. ssp. teta. — Cf. Jessop, Blumea 24 (1976) 155. — Teta viridiflora Roxs. Fl. Ind. ed. Carey 2 (1832) 165. — P. teta var. mantegazziana PAMP. Nuovo Giorn. Bot. Ital. n.s. 11 (1904) 151; Bull. R. Soc. Toscana Ortic. III, 10 (1905) 50, f. 11. — P. mante- gazziana (PAMP.) PAMp. Nuovo Giorn. Bot. Ital. n.s. 13 (1906) 138; Merr. & Quis. Philip. J. Sc. 82 LILIACEAE—I (Jessop) 229 Fig. 17. Range of Peliosanthes teta ANDR. ssp. teta (broken line), and ssp. humilis (ANDR.) Jessop (even line). (1953) 323. — P. graminea Riwe. J. Str. Br. R. As. Soc. n. 59 (1911) 207. — P. teta var. angusti- folia Rw. /.c. — P. tonkinensis WANG & TANG, Bull. Fan Mem. Inst. Biol. Peiping, Bot. 7 (1936) 83. Pedicels 2—6-nate. Leaf index 2—24(—34). Flowers usually green, rarely blue. Anthers c. 0.5-0.6 mm long. Distr. India to southern China; in Malesia: Malay Peninsula (Pahang, Selangor, Penang and Langkawi Is.). Fig. 17. b. ssp. humilis (ANDR.) Jessop, Blumea 24 (1976) 155. — P. humilis ANpR. Bot. Repos. 10 (1811) t. 634; Mia. Fl. Ind. Bat. 3 (1859) 568. — Bulbi- sperma ovigera REINW. ex BL. Cat. (1823) 59, nomen. — Bulbospermum javanicum Bu. En. PI. Jav. (1827) 15; Zo. Syst. Verz. 1 (1854) 68. — P. javanica (BL.) Dieter. Syn. Pl. 2 (1840) 1123; Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 121; Pl. Jav. Rar. (1848) 116; Mig. FI. Ind. Bat. 3 (1859) 568; Back. Handb. FI. Java 3 (1924) 75; BAcK. & BAKH. f. Fl. Java 3 (1968) 95; STEEN. Mt. Fl. Java (1972) t. 28-4. — P. violacea WALL. ex Baker, J. Linn. Soc. Bot. 17 (1879) 504; HEND. Mal. Wild Flow. Monoc. (1954) 184, f. 108. — Lourya campanulata BAIL. Bull. Soc. Linn. Paris 1 (1888) 743. — P. albida BAKER, Bot. Mag. 116 (1890) t. 7110; Hook. f. Fl. Br. Ind. 6 (1892) 267; RipL. Mat. Fi. Mal. Pen. Monoc. 2 (1907) 90; Merr. En. Born. (1921) 115; FiscHerR, Kew Bull. (1932) 183. — P. viridis Ripv. J. Str. Br. R. As. Soc. n. 31 (1898) 95. — P. lurida Riv. lI.c. 95. — P. grandifolia Rw. I.c. 97. — P. stellaris RIL. l.c. 97. — P. parviflora RI . ibid. n. 61 (1912) 61. — P. sumatrensis Riv. J. Fed. Mal. St. Mus. 8 (1917) 118. — P. sessiliflora Riv. l.c. 118. — P. hypogyna Rw. ibid. 10 (1920) 121. — P. monti- cola Rip. /I.c. 155. — P. campanulata (BAILL.) RODRIGUEZ, Bull. Mus. Hist. Paris II, 6 (1934) 96. — Fig. 16e-g. Pedicels solitary in the axil of each bract. Leaf index 2-10. Flowers sometimes green, but often white, blue, violet or purple. Anthers c. 1/,-2 mm long. Distr. India to southern China; in Malesia: Malay Peninsula (throughout, incl. the Langkawi, Penang, Singapore and Tioman Is.), Sumatra (throughout, incl. Simalur and Billiton Is.), Java (throughout), Lesser Sunda Is. (Sumbawa) and Borneo (Sarawak, Southeast, Sabah, incl. the Anambas and Karimata Is.). Fig. 17. 230 FLORA MALESIANA [ser. I, vol. 91 21. ALETRIS LINNE, Sp. Pl. (1753) 319; Gen. Pl. ed. 5 (1754) 149; Amoen. Acad. 3 (1756) 11 (‘Alethris’); Hook. f. Fl. Br. Ind. 6 (1892) 264; FRANCHET in Morot, J. de Bot. 10 (1896) 178, 195; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 378; Hara, J. Jap. Bot. 42 (1967) 312. — Metanarthecium Max. Bull. Ac. St. Pétersb. 11 (1867) 438; KRAUSE in E. & P. Nat. Pfl. Fam. ed. 2, 15a (1930) 260. — Meta-aletris Masao. Trans. Nat. Hist. Soc. Formosa 28 (1938) 46. — Fig. 18, 19. Erect, stemless, rhizomatous herbs. Roots fibrous. Leaves basal, linear or lanceo- late, sessile, the veins of the decayed bases persisting as fibres at the base of the plant. Inflorescence a raceme or spike. Flowers solitary in the axils of the bracts, with a single bracteole. Pedice/s not articulated. Perianth segments connate at the base, equal, with three (often indistinct) veins, ascending or reflexing, glabrous or pubescent, white or pink. Filaments subulate, shorter than and attached to perianth; anthers dorsifixed, oblong to ovoid, dehiscing introrsely. Ovary half-inferior, 3-celled; ovules axile, numerous; style simple or minutely 3-lobed. Fruit a capsule; perianth persistent. Seeds oblong. Distr. North America (few spp.) and eastern Asia, from Japan to southern China (also Taiwan) and the Himalayas, in Malesia: only on the high mountains of the Philippines (N. Luzon), Sabah (Mt Kina- balu) and N. Sumatra (Gajolands and Westcoast Res.). There are possibly fewer than 15 spp., mostly in Asia, but HARA /.c. estimates the number at c. 30. Ecol. Elfin forest and mossy mountain forest, but mostly in open, sometimes damp situations in sedge- or grasslands, crevices of rocks, and mountain heaths, locally often common, between (1000—)1600 and 3250 m. KEY TO THE SPECIES Pe: 1. A. foliolosa 1. Perianth glabrous. Leaves mostly spreading, usually at least 4 mm broad 1. Perianth glandular-pubescent. Leaves usually erect, less than 4 mm broad 1. Aletris foliolosa STAPF, Trans. Linn. Soc. II, 4 (1894) 240; Doct. v. LEEUWEN, Trop. Natuur 9 (1920) 98, fig.; Merr. En. Born. (1921) 115; Docr. v. LEEUWEN, Verh. Kon. Ak. Wet. A’dam sect. II, 31 (1933) 251, f. 66 (‘foliosa’). — A. rigida STAPF, Trans. Linn. Soc. II, 4 (1894) 241; Gress, J. Linn. Soc. Bot. 42 (1914) 164; Merr. En. Born. (1921) 115. — ? Liriope brachyphylla MeErr. Philip. J. Sc. 2 (1907) Bot. 266; En. Philip. 1 (1922) 206. — ? Metanarthecium brachyphyllum (MERR.) MASAM. Bull. Soc. Bot. Fr. 84 (1937) 18. — A. sumatrana MasAM. /.c. 18. — Meta-aletris sumatrana (MASAM.) Masao. Trans. Nat. Hist. Soc. Formosa 28 (1938) 46. — Meta-aletris rigida (STAPF) MASAM. I.c. 46. — Fig. 18a-d, 19. Leaves glabrous, ascending at first but usually spreading when mature and finally recurved, to 6(-10) cm by 4-7(—10) mm; veins several, close to one another. Peduncles 1 or more, 6-65 cm high, glabrous, rigid, with 1-6 narrow-lanceolate, sterile bracts to 2 cm long; fertile bracts 4-20, lanceolate, 5-10 mm long; bracteoles similar to bracts but shorter. Pedicels 0-5 mm long in flower, sometimes over | cm in fruit, usually expanding gradually to apse i] a » ie kl * al} =_ a iat on . ioe ae 7 - t¢ f ¥ | ; > | 4 | uty “6 a ee | 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 REQUESTS AND RELATED CORRESPONDENCE SHOULD BE DIRECTED EXCLUSIVELY TO SISTHOFF & NOORDHOFF INTERNATIONAL PUBLISHERS Volume 1. Volume 2. Volume 3. Volume 4. Volume 5. Volume 6, Volume 7. Volume 8. Volume 9. P.O. Box 4, Alphen aan den Rijn, The Netherlands For sale only - Not for exchange Series I - Spermatophyta (Flowering Plants) Cyclopaedia of collectors & collections. 1950. pp. clii + 639. Out of print. Available in microfiche. Malesian vegetation (In preparation) Malesian plant geography (In preparation) General chapters and revisions. 1948-1954. pp. ccix + 631 | Bibliography, specific delimitation & revisions. 1955-1958. pp. cccxlii + 596 Systematic revisions, Parts 1-6. 1960-1972. PP. 20 + 1023. Out of print. Available in microfiche. Systematic revisions. Parts 1-4. 1971-1976. pp. 18 + 876 Cyclopaedia of Collectors, Suppl. 2. Systematic revisions. Parts 1-3. 1974-1978. pp. 19 + cxv + 577 Systematic revisions. Part 1. 1979. SIJSTHOFF & NOORDHOFF INTERNATIONAL PUBLISHERS, - ALPHEN AAN DEN RIJN, THE NETHERLANDS PRINTED IN THE NETHERLANDS ISBN 90 286 0629 7