FLORA MALESIANA SERIES I- SPERMATOPHYTA Flowering Plants Vol. 7, part 3 Aceraceae . Actinidiaceae s.str. Aizoaceae . Alismataceae Amaranthaceae . Ancistrocladaceae Aponogetonaceae Basellaceae Batidaceae. Betulaceae Bixaceae s. str. Burmanniaceae . Burseraceae Butomaceae Byblidaceae Callitrichaceae Campanulaceae . Cannabinaceae . Capparidaceae Caprifoliaceae Cardiopteridaceae Celastraceae Centrolepidaceae Ceratophyllaceae Chenopodiaceae Clethraceae Cochlospermaceae Combretaceae Connaraceae Convolvulaceae Corynocarpaceae Crassulaceae Cyperaceae Datiscaceae . Dichapetalaceae . Dilleniaceae Dioscoreaceae Dipsacaceae Droseraceae INDEX TO REVISED FAMILIES 42.3: 4: 4: . ae 4: 69, - 4: 4s 11542: " SE 5 5: * 4: 4: 13, 5: R 6: POSS ey 2 The G2-227- Ss 3 4: 4: 99, gE 4: 4: 5: 4: 338, Z 4: 4: als 4: 5: 4: 4: 4: 4: 592 37 267 317 593 8 213 300 414 207 239 592 209 598 93 389 421 41 594 139 61 533 495 599 262 197 435 382 305 141 293 290 377 Elatinaceae Epacridaceae Ericaceae . ; Erythroxylaceae . Fagaceae Ficoidaceae Flacourtiaceae Flagellariaceae Geraniaceae Gnetaceae . Gonystylaceae Goodeniaceae Haemodoraceae . Haloragaceae Hamamelidaceae Hippocrateaceae Hydrocaryaceae . Hydrocharitaceae Hydrophyllaceae Icacinaceae Juglandaceae Juncaceae. Juncaginaceae Lemnaceae Loganiaceae Lophopyxidaceae Malpighiaceae Martyniaceae Molluginaceae Moringaceae Myoporaceae Myricaceae Najadaceae Nyctaginaceae Nyssaceae . Ochnaceae. Oxalidaceae Papaveraceae Passifloraceae No Ww \o . — to | ~) eee N \o ANIARAADRPARAAMNAAIBRAAIAMAAMAMNMAARAA = Nn “A — | ~ aN oS a Pedaliaceae Pentaphragmataceae Pentaphylacaceae Philydraceae Phytolaccaceae . Pittosporaceae Plumbaginaceae . Podostemaceae . Polemoniaceae Pontederiaceae Portulacaceae Primulaceae Proteaceae. Punicaceae Restionaceae Rhizophoraceae . Salicaceae . Salvadoraceae Sarcospermaceae Saururaceae Scyphostegiaceae Simaroubaceae . Sonneratiaceae Sparganiaceae Sphenocleaceae . Stackhousiaceae . Staphyleaceae Stylidiaceae Styracaceae P Thymelaeaceae . Trigoniaceae Turneraceae Typhaceae Umbelliferae Valerianaceae Violaceae . Xyridaceae Zygophyllaceae . 4: 216 4: 517 5: 121 4:75 4: 229 5: 345 4: 107 4: 65 4: 195 4: 255 7: 121 6: 173 5: 147 4: 226 5: 416 5: 429 5: 107 4: 225 4: 32 4: 47 5: 297 ‘ 6: 193 4: 280, 513 4 4 6 4 4 aS \o . 49 4: 349, 6: 1 4: 59 , 4: 243 4: 113, 595 4: 253 7: 179 4: 366, 598 4: 64 CYPERACEAE (J. H. Kern, Leyden) Annual or perennial, often grass-like herbs, only the monotypic African genus Microdracoides tree-like; the perennial spp. with short- or long-creeping, mostly sympodial rhizome not rarely emitting stolons. Stems solid, exceptionally hollow, sometimes septate, often trigonous, more rarely 2-sided or terete, or 4-, 5-, or multangular, usually nodeless below the inflorescence. Leaves often 3- ranked, more rarely distichous or polystichous, basal and/or cauline, usually sheathing at the base, the sheaths closed (in Mal.), very rarely open, the blades as a rule sessile, linear (grass-like) or setaceous, rarely lanceolate and petioled, rarely much reduced or even absent; sheath and blade whether or not separated by a rim of short hairs or by a membranous ligule almost completely fused to the upper surface of the blade. F/owers simple, inconspicuous, each subtended by a bract (g/ume), arranged in small spiciform units (spike/ets), in subfam. Caricoideae strictly unisexual, in subfam. Cyperoideae tribe Hypolytreae composed of mon- androus lateral ‘flowers’ and a terminal ovary, in tribe Cypereae reduced to bisexual synanthia, a few of which may be functionally male or female by abortion of the other sex. Spikelets often (always?) cymose (“‘pseudo-spikelets’), (1-) few- to many-flowered. Inflorescence paniculate, anthelate, capitate, or spicate, with few to many spikelets, rarely reduced to a single spikelet, often subtended by 1-several leafy involucral bracts. Perianth consisting of bristles, hairs, or scales, but often absent. Stamens often 3, not rarely reduced to 2 or 1, very rarely more than 3 to numerous; filaments ligulate, free, only in a few Carex spp. connate, sometimes strongly elongating after anthesis; anthers basifixed, introrse, opening lengthwise by a slit. Ovary solitary, superior, usually 2- or 3-carpellate, unilocular; style not rarely thickened at the base, the thickened part whether or not articulated with the ovary; stigmas 2 or 3 (rarely more), only in a few spp. style unbranched; ovule solitary, erect from the base of the ovary, anatropous. Fruit indehiscent, a nut (often termed achene), sessile, or seated on a disk, free, or surrounded by a modified prophyll (perigynium, utricle). Seed erect, with thin testa not adhering to the pericarp; embryo small, at least partly surrounded by abundant mealy or fleshy endosperm. Distribution. About 70-80 genera with probably some 4000 spp., throughout the world. Among the genera occurring in Malesia only Capitularina is endemic (in New Guinea and the Solomons) and Paramapania (throughout Malesia) is almost endemic, with one Mal. sp. extending to the West Pacific. Malesian Cyperaceous genera show geographical relations in several directions: some have marginally westwards extending areas, for example Oreobolus, which genus is largely distributed in the southern half of the Pacific, reaching its most western station on the ancient high mountains of North Sumatra. Fig. 81. Another example is the very similarly distributed genus Uncinia of which the most western stations are found on Mt Kinabalu in North Borneo and in the Philippines. Another type of marginal occurrence is that of Kobresia (incl. Schoenoxiphium), a genus which is distributed over Africa and the northern hemisphere and shows up with one species in the ancient high mountains of North Sumatra. Several genera possess a worldwide range, sometimes restricted to the tropics and subtropics, e.g. Bulbostylis, Fuirena, Hypolytrum, Lipocarpha, and Mapania, sometimes however distributed (435) 436 FLORA MALESIANA [ser. I, volags over the warm and temperate zones, e.g. Carex, Cladium, Cyperus, Eleocharis, Fimbristylis, Machaer- ina, Rhynchospora, Schoenus, Scirpus, and Scleria. Among the latter Machaerina and Scleria have only a few representatives in the extra-tropical zones; reversely Carex is largely extra-tropical and in the tropics almost confined to montane stations. Cyperus and Fimbristylis occur predominantly in the tropical lowlands and hills, and rapidly diminish in number towards the temperate zones. The following genera are confined to the Old World: Carpha, Costularia, and Tetraria; Lepironia has a similar range, but its African stations are restricted to Madagascar, while Thoracostachyum occurs in the African region only in the Seychelles. Finally a number of genera occur only in the Old World, but are absent from Africa, viz Gahnia (fig. 96), Scirpodendron, Lepidosperma (fig. 76), and Tricostularia, among which the latter two are properly Australian-centred, extending with a single species through Malesia to Asia. Distribution of species. A fairly large number of species occurring in Malesia have (i) a worldwide or almost worldwide distribution, most of them in the warmer regions of the globe; for example: Bulbostylis barbata Eleocharis geniculata Carex curta nigrescens echinata parvula (not Australia) pseudocyperus retroflexa remota Fimbristylis complanata Cladium mariscus cymosa Cyperus brevifolius dichotoma compressus ferruginea cuspidatus hispidula cyperoides littoralis digitatus squarrosa halpan Rhynchospora corymbosa kyllingia gracillima (not America) odoratus triflora (not Australia) pedunculatus Scleria lithosperma polystachyos sesquiflorus unioloides It is not unexpected that these species are almost all plants which are indifferent to soil and belong to open lowland habitats, often inhabiting places with little competition, such as beaches and waste land, and are suitable to pioneering in disturbed and cultivated places. It is striking that there are no forest dwellers among them. It is not impossible that some may have attained their large area due to man in pre-historic or post-Columbian time. They are also almost all very common species. This is not always the case, as there are also some widely distributed species which show remarkable (ii) disjunct areas, e.g. Eleocharis variegata (Africa, Madagascar, Mauritius, then in North Sumatra and twice found in New Guinea), Carex michauxiana (NE. North America, China, Japan, and the highlands of New Guinea), Scleria mikawana (Africa, Ceylon, Japan, and in New Guinea), Scleria annularis (India to China, then New Guinea), and Sc/eria parvula (from Ceylon to Korea, then Luzon and New Guinea). Possibly future collections will fill the gaps, as many collectors have neglected to collect sedges. A range like that of Sc/eria parvula may show a quite natural pattern, as there are in New Guinea several plant species ranging from Japan, via Formosa and the Philippines to that island, although such cases mostly refer to mountain plants, e.g. several Carices. A few seemingly rather queer ranges have appeared to be due to misidentification or to curiously mis-localized collections for which is referred to Cyperus esculentus, Carex divulsa and Carex muri- cata (= C. pairaei). Besides the two categories mentioned, wor/dwide and disjunct, there are four others, three of which show ranges extending from the borders just into Malesia, viz from Asia, Australia, and from the Pacific, whilst the fourth category concerns the endemic species of Malesia. Of each of these groups examples will be given, not an exhaustive enumeration. 1974] Cy PERACEAE (Kern) 437 (iii) Asian representatives (mountain plants marked by an asterisk): Carex *duriuscula (Korea, New Guinea) *michauxiana (northern hemisphere, Guinea) Cyperus substramineus (Asia, Malaya) Eleocharis *acicularis (northern hemisphere, N. Sumatra, N. Luzon) *attenuata (Japan, China, New Guinea) Fimbristylis adenolepis (Thailand, Indo-China, Kangean Is.) *disticha (Burma to China, N. Sumatra) *nierotii (Himalaya to Japan, Luzon) New *thomsonii (Himalaya to Formosa, Malaya, N. Sumatra, Palawan) Machaerina *maingayi (Tonkin, Malaya) Rhynchospora malasica (Japan, China to W. Malesia) Scirpus wallichii (Japan to India, Malaya, Philip- pines) *wichurai (Himalaya, N. Sumatra) Scleria neesii (SE. Asia, Malaya) reticulata (SE. Asia, Malaya) thwaitesiana (Ceylon to Thailand, Malaya) (iv) Australian species just entering the borders of Malesia (mountain plants marked by an asterisk): Carpha *alpina (New Zealand, Tasmania, Aus- tralia, New Guinea) Cyperus angustatus (Australia, New Guinea) aquatilis (ditto) dietrichiae (Australia, New Britain) fulvus (Australia, New Guinea) */ucidus (ditto) pedunculosus (ditto) Eleocharis *acuta (New Zealand, Tasmania, Australia, New Guinea) *sphacelata (ditto) Fimbristylis furva {Australia; New Guinea: Wassi Kussa, Merauke; Aru Is.) recta (Australia; S. New Guinea: Kussa) schultzii (Australia, Bali, Sumba) Machaerina *gunnii (Australia, New Guinea) *teretifolia (New Zealand, Australia, W. New Guinea) Wassi Oreobolus *pumilio (Tasmania, Australia, New Guinea) Schoenus *melanostachys (Australia, Mindoro, Mt Kinabalu) *nitens (S. Chile, New Zealand, Australia, New Guinea) sparteus (Australia, New Guinea, Wetar) Scirpus *aucklandicus (New Zealand, Australia, New Guinea) *crassiusculus (ditto) *inundatus (temperate South America, New Zealand, Australia, New Guinea) *subtilissimus (New Zealand, Australia, New Guinea, W to Luzon, Mt Kinabalu) Scleria brownii (Australia, New Caledonia, Tonga, New Guinea) novae-hollandiae (Australia, Micronesia, New Guinea, Luzon) tricuspidata (Australia, S. Moluccas: Aru Is.) (v) The Pacific is botanically much poorer and has therefore only few species extending into East Malesia: Machaerina mariscoides (Polynesia, New Guinea) Scleria polycarpa (Fiji, Melanesia, New Guinea, Moluccas). Fig. 110. (vi) Of many genera endemic species occur in Malesia amongst which are many forest dwelling species of the Mapanieae (mountain plants marked with an asterisk): Capitularina involucrata (New Guinea) Carex *eremostachya (New Guinea & Solomons) *gajonum (N. Sumatra) */oheri (Philippines) malaccensis (Malaya) *merrillii (Philippines) nodiflora (Philippines) palawanensis (Philippines) *sarawaketensis (New Guinea) spathaceo-bracteata (New Guinea) Costularia pilisepala (Borneo, New Guinea) Cyperus cinereobrunneus (New Guinea) meistostylis (New Guinea) neoguineensis (New Guinea) pachycephalus (New Guinea) subpapuanus (New Guinea) Eleocharis *brevicollis (New Guinea) sundaica (Alor) Fimbristylis blepharolepis (New Guinea) caesia (Philippines, Java) calcicola (Malaya) capilliculmis (New Guinea) 438 FLORA MALESIANA [ser. I, vokage Fimbristylis celebica (Central Celebes) monostachya (Borneo) lineatisquama (Philippines) sessilis (W. Malesia) macassarensis (Luzon, SW. Celebes, Madura spadicea (Borneo) I.) squamata (W. Malesia) malayana (Malaya) wallichii (W. Malesia) subdura (Java) Oreobolus *kiikenthalii (N. Sumatra, Malaya) sumbaensis (Sumba I.) Paramapania flaccida (New Guinea) wetarensis (Wetar I.) gracillima (Philippines) Hypolytrum capitulatum (Borneo) longirostris (New Guinea) humile (West Java) radicans (Borneo) Kobresia *kobresioidea (N. Sumatra) rostrata (Philippines) Machaerina aspericaulis (Mt Kinabalu) simplex (New Guinea) *lamii (New Guinea) Schoenus *curvulus (New Guinea to Mt Kina- Mapania angustifolia (Borneo) balu) debilis (Borneo) delicatulus (Palawan, Mt Kinabalu) foxworthyi (Borneo) */ongibracteatus (New Guinea, Mt Kinabalu) graminea (Borneo) Scirpus *beccarii (Sumatra) holttumii (Malaya) *junghuhnii (N. Sumatra) latifolia (Borneo) Scleria cyathophora (W. Malesia) longiflora (Borneo, Malaya) densispicata (Luzon) lorea (W. Malesia) papuana (New Guinea) maschalina (Borneo) pygmaeopsis (Sumba) micropandanus (Malaya) Tetraria borneensis (Borneo) Ecology. Sedges occur in every formation and even in any biotope (except epiphytic) and not infrequently they form a substantial part of the herbaceous plant cover in grassland and savanna. A large part of them occurs in lowland eutrophic swamps and consequently in wet rice-fields; they abound also in swinging bogs on lake shores. However, some occur under distinct oligotrophic conditions, e.g. on kérangas,in peat swamps and high mountain marshes. Some are confined to the beach, and others to calcareous cliffs. Below an approximate survey is given of species which are characteristic of marked soil conditions or vegetation types. Going from the sea-shore inland we have then the following sequence: Mangrove. Along muddy sea-shores or in drier places in the mangrove are found Fimbristylis argentea and F. polytrichoides, the latter sometimes in quantity. Also Cyperus javanicus is largely confined to coastal swamps in brackish water. Sandy beach. The following species are characteristic: Cyperus bulbosus, C. dubius, C. hyalinus, C. pedunculatus (fig. 65), C. radians, C. stoloniferus (fig. 50), Fimbristylis cymosa, and F. sericea. Rhizomes or tubers are sometimes shielded by a tunica, for example in Fimbristylis sericea and Cyperus bulbosus respectively. All species produce runners for vegetative propagation, sometimes very long ones, as in Cyperus pedunculatus. Some are local or rare (Cyperus bulbosus, C. dubius, C. hyalinus), but others are very frequent (Fimbristylis cymosa) and may also occur somewhat inland behind the beach on low sandy flat coasts, as Bulbostylis barbata and B. puberula, which also may occur in abundance on the sandy beach. Saline habitats (“asinans’) occur here and there inland, especially in the seasonal districts. They are sometimes bound up with hot springs or mudwells. In such habitats several sedges are found which are in that district confined to such spots. Examples are Eleocharis parvula(E. Java, once), E. spiralis (often gregarious), Fimbristylis ferruginea, F. semarangensis, F . sieberiana, and Scirpus litoralis. Lowland swamp forests. The following species have been cited to be characteristic of or abundant in this vegetation type: Capitularina involucrata, Fimbristylis scaberrima, Mapania enodis (sometimes gregarious and dominant), M. lorea, M. macrocephala, M. palustris, Scirpodendron (sometimes forming pure stands), Scirpus ternatanus, and Thoracostachyum (2 spp.). Peat swamp forests. ANDERSON (Gard. Bull. Sing. 20, 1963, 216) mentioned two species, viz Tetraria borneensis, a tall sedge which is abundant in the centre of peat forest but also occurs in open heath forest (fig. 72), and Thoracostachyum bancanum which is found in all zones of the peat forest, but occurs also in mixed swamp forest. 1974] Cy PERACEAE (Kern) 439 Lowland swamps. No clear distinction could be made here between species of oligotroph and those of eutroph waters, nor between swamps with permanent water-level and those with fluctuating level. The first category (oligotroph waters) is rare and for example Lepironia belongs here; it can grow gregariously (fig. 6). Several other species obviously of eutroph waters can grow gregariously, for example Eleocharis dulcis (fig. 36), Fimbristylis dipsacea (locally in Bawean), F. pauciflora, Machaerina rubiginosa (fig. 94), Rhynchospora corymbosa, Scirpus mucronatus (big stands even in the mountains), and Scleria poaeformis. The following are characteristic, often common swamp and marsh sedges: Cladium mariscus (rare) Fimbristylis acuminata Cyperus babakan aestivalis digitatus aphylla distans complanata elatus ferruginea exaltatus (rare, often coastal) globulosa halpan griffithii holoschoenus (rare) intonsa (rare) imbricatus littoralis iria miliacea malaccensis perlaxa (rare) nutans tenuicula (rare) odoratus tetragona pilosus thomsonii (rare) platystylis (rare) Fuirena spp. polystachyos Lipocarpha chinensis procerus Machaerina disticha pulcherrimus rubiginosa pygmaeus (rare) Rhynchospora triflora (rare) sanguinolentus Scirpus articulatus (fig. 31) scariosus grossus (fig. 22) tenuispica Juncoides Eleocharis retroflexa lacustris spp. div. (fig. 35) lateriflorus Scleria rugosa Floating waterplants are few but characteristic, as they take part in the composition of floating islands of kumpai vegetation, often together with Eichhornia, Hanguana, Pistia, etc., and in that of swinging bogs. They are: Cyperus alopecuroides imbricatus cephalotes (fig. 61—62) ohwii elatus platystylis Scirpus confervoides takes a separate position in being a true submerged aquatic plant (fig. 25). Mixed dryland forests at lower altitudes have in primary condition almost only sedges of the tribe Mapanieae (Hypolytrum, Mapania, Paramapania) on the forest floor (fig. 13-14), but there are also a number of Cyperus spp. occupying that habitat viz C. diffusus, C. multispicatus, C. pedunculosus and allies (spp. 27-32), and Machaerina sinclairii (500-2000 m), M. glomerata (up to 1500 m) and M. mariscoides (up to 350 m). In the lowland and hill forest there are a number of Carices, most of which extend also to the colline and montane levels, e.g.: Carex cryptostachys (40—1500 m) rafflesiana (SO0—2400 m) horsfieldii (100-1100 m) satzumensis (400—2300 m) indica (up to 1000 m) speciosa (10-1500 m) maubertiana (400—2100 m) 440 FLORA MALESIANA [ser. I, vol. 7° Secondary thickets replacing primary forest harbour several sedges, amongst them a number of mostly tall Sc/eria species, which may form the major constituent, e.g. Scleria ciliaris, S. levis, S. lithosperma, S. oblata, S. purpurascens, S. scrobiculata, S. sumatrensis, S. terrestris; furthermore Fimbristylis dura and some species of Cyperus, notably C. cyperinus, C. cyperoides and C. diffusus. Podsolized soils. On kérangas soils some sedges are characteristic and confined to this oligotroph siliceous substratum, viz Fimbristylis fuscoides, Schoenus calostachyus (fig. 79), Tetraria borneensis (fig. 72), and Tricostularia undulata (fig. 78). Limestone hills are scarce in Malesia and their most characteristic development is found in the northern part of Malaya and in the Langkawi Is., as a southern extension of those hills in Thailand. They are remains of a geological formation which had in Tertiary time a much larger extension. The following sedges are confined to these hills on this specialized habitat: Carex malaccensis, Fimbristylis calcicola, F. malayana, F. trichophylla, Cyperus teneriffae (in Timor!), while C. hyalinus often occurs on limestone, but is not restricted to it. Craters and solfatara offer on the one hand little competition but on the other hand they are special- ized habitats because of the infertility of the ‘soil’, the presence of acids in the soil, and sulphureous and other poisonous gases in the air. Species which can stand this environment and are often even common here are Carex baccans, C. verticillata, and Gahnia javanica (fig. 97). None of them is bound to this ‘niche’. On volcanic screes and landslides some Carices may form large clumps, notably Carex baccans. Mid-mountain forest harbours proportionally few sedges and even here they prefer ridges or open spots near rocks and waterfalls. In such places one can find Carex turrita, C. verticillata(2000—3750 m), and Gahnia baniensis (sometimes gregarious, 900—2100 m). The sedges of the mountain forest mostly belong to Carex, e.g.: Carex anemocarya (1000—1200 m) lateralis (1400—2200 m) breviscapa (1000—1200 m) loheri (1300—2400 m) brunnea (1400-2100 m) longipes (1500—2200 m) commixta (800—1500 m) ramosii (medium altitude) dolichostachya (1200—1600 m) rhynchachaenium (800—2100 m) helferi (1100 m) satzumensis (400—2300 m) lamprochlamys (800-2700 m) Machaerina aspericaulis (1500—1600 m) Subalpine forest is again a vegetation type in which Cyperaceae are mainly represented by Carices: Carex brachyathera (3000—4000 m) perciliata (2400—4200 m) breviculmis (2200—3900 m) Phacelostachys (2200-3500 m) celebica (2400—3700 m) sarawaketensis (2700—3950 m) filicina (1400-3680 m) spathaceo-bracteata (3000—4000 m) finitima (2400-3900 m) verticillata (2000—3750 m) graeffeana (800—3800 m) Machaerina falcata (1900—3300 m) maculata (1800—3500 m) Uncinia riparia (2300—3680 m) myosurus (1700-3300 m) rupéstris (2900—3300 m) In Phillippine pine forest is found Fimbristylis pierotii (1550 m). Waterplants in the mountain streams are three species of Scirpus, viz S. beccarii (Sumatra, 2750— 3300 m; fig. 24), S. crassiusculus (New Guinea, 1800-3900 m), and S. fluitans (Java, 1600-3200 m). A characteristic river bank sedge is Carex teinogyna (Sumatra, 700-1150 m). Mountain swamps, bogs, marshes, and stream fringes abound with sedges, some of which may occur gregariously. The following have been selected: Bulbostylis densa (1000-3000 m) gajonum (3100—3300 m) Carex brownii (1500—2500 m) gaudichaudiana (1900-3700 m) capillacea (2000—4000 m) graeffeana (gregarious, 800—3800 m) celebica (2400—3700 m) jackiana (1350—2550 m) echinata (2400-3600 m) maculata (1800—3500 m) finitima (2400-3900 m) michauxiana (2250—2650 m) 1974] Cy PERACEAE (Kern) 44] montivaga (3200-3500 m) salbundia (900—2200 m) nubigena (1600—3000 m) Kobresia kobresioidea (gregarious, 2900—3400 m) oedorrhampha (1200—2400 m) Machaerina articulata (gregarious, 1900—2500 phacota (gregarious, 1500-2700 m) m) pruinosa (1560-2500 m) falcata (1900—3300 m). Fig. 90 pseudocyperus (1750-3225 m) gunnii (2250-3000 m) teres (2100-3500 m) lamii (3250—3520 m) Cyperus lucidus (gregarious, 2500 m) maingayi (900-2100 m) melanospermus (500-3000 m) rubiginosa (gregarious, 100-3225 m) sanguinolentus ssp. melanocephalus (gregar- teretifolia (2100 m) ious, 1000—3125 m) Rhynchospora rugosa (often gregarious, 0—2800 Eleocharis attenuata (1650—2800 m) m) congesta (50—2800 m) Scirpus inundatus (1750—2700 m) tetraquetra (650-2700 m) subcapitatus (1200—4000 m). Fig. 27—28 Fimbristylis consanguinea (1600—2300 m) subtilissimus (1600-3700 m) disticha (1200—1400 m) Mat-formation, stubs, and fairy rings. Many Cyperaceae show a tufted habit when rhizome inter- nodes are short, and finally big tufts may be formed, e.g. in several Carex, Cyperus and Gahnia species, and Rhynchospora rugosa (fig. 102). By the small species of the subalpine and alpine turf and heathland circular, low tufts are formed. Because of the rhizomes growing out in a centrifugal way such tufts may become fairly large clumps even leading to mats (fig. 84). In the central part of such roundish ‘Polster’ the oldest parts may die and decay, so that fairy rings are formed; this is a common feature observed in Oreobolus, similar to that found in tufts of Isachne pangerangensis, Centrolepis, etc. (fig. 83, 87). Though it is self-evident that through the growth mode an empty space is formed in the centre of the tuft, it is peculiar that this empty centre is not occupied by new young plants, it looks as if it were poisoned. It is not yet clarified whether the centre is exhausted of certain trace elements or that self-poisoning must be assumed. The latter assumption is not likely, because also other plant species do not occupy the bare centre of the fairy ring. In swamps liable to a seasonally oscillating water-level stubs can be formed which may attain quite some size, up to knee-high, as was observed on Tegal Pangonan, on Mt Diéng (Central Java) in Carex phacota. Such stubs are invaded by other marsh plants, but also by dryland herbs, in a sort of vertical succession. Subalpine heathland and meadows are generally dry, but in slight depressions and through seepage boggy places may be formed, so that this category is hardly distinct from the former. Moreover, such boggyness may be only temporary, so that the list below forms rather an addition than a class apart: Carex vesiculosa (1200—3500 m) Schoenus curvulus (1500-4000 m) Carpha alpina (3000—4200 m) longibracteatus (1500—3500 m) Eleocharis sphacelata (2200—2900 m) maschalinus (1800—3600 m) Gahnia javanica (1200-3560 m) melanostachys (1650-2400 m) Lepidosperma chinense (1500-3000 m). Fig. 77 nitens (3400 m) Machaerina gunnii (2250—3000 m) setiformis (2700-3750 m) Oreobolus ambiguus (2500—4000 m). Fig. 87 Scirpus aucklandicus (3300 m) kiikenthalii (1600-3460 m). Fig. 82—84 Junghuhnii (2200—3400 m) pumilio (2600—3800 m) In seasonal regions, with a long, distinct, annual period of drought, a large number of sedges occur in grasslands which may be swampy during the rainy season, usually in the period from November till March. In April and May sedges flower together with the grasses. This vegetation type is often subject to fire. It extends in West Java along the north coast east of Djakarta — Indramayu Residency has proved a very rich site — and then further eastwards through Central and East Java and the Lesser Sunda Islands to the southern border of New Guinea (Merauke to Wassi Kussa), along the Fly R. delta to Moresby and onwards to the east. Also parts of Celebes (mostly in the southern penin- sulas), spots in tne Moluccas (in Buru and Ceram), and the west sides of most Philippine Islands have 442 FLORA MALESIANA [ser. I, vol. 7° this type of climate. Some species are more common in the swampy places, others in permanently dry parts. The following are characteristic, some being rare or rarely collected: Carex stramentitia lanceolata tricephala macassarensis Cyperus alopecuroides microcarya angustatus recta bulbosus schultzii holoschoenus semarangensis nervulosus sieberiana paniceus subalata squarrosus subdura Fimbristylis adenolepis sumbaensis anisoclada wetarensis bisumbellata Gahnia aspera blepharolepis Lipocarpha microcephala dictyocolea Rhynchospora hookeri fimbristyloides Schoenus sparteus furva Scleria junghuhniana insignis psilorrhiza Savanna is of course hardly separable from the grassland category mentioned above, as these two vegetation types grade and are both liable to a dry season and annual burning. But savannas are generally dryland; the following sedges are characteristic: Fimbristylis cinnamometorum signata eragrostis wetarensis falcata Rhynchospora heterochaeta. Fig. 98 furva longisetis fusca rubra. Fig. 99 insignis subtenuifolia lanceolata wightiana ovata Schoenus falcatus recta punctatus Dispersal. Local extension by vegetative growth and even propagation is a very common fea- ture in Cyperaceae, on account of the frequent occurrence of rhizomes, runners and more rarely bulbs. This leads frequently to local almost pure stands (colonies), so characteristic of many swamp and highland species, and also to their rapid extension on waste land. It is characteristic in most sedges of the sandy beach, e.g. in Cyperus bulbosus, C. pedunculatus (fig. 65), C. stoloniferus (fig. 50), etc. It is of course equally common among inland species, e.g. in Cyperus rotundus (fig. 49), C. brevifolius (fig. 70), and other species of Cyperus sect. Kyllingia. This is one of the reasons that some sedges are difficult to eradicate, notably Cyperus rotundus, as parts of rhizomes or bulbs are in this way dispersed and act as vegetative diaspores. As in most large families there is quite an array of dispersal devices of the fruits. Unfortunately the number of actual observations in Malesia is very small and examples are largely derived from the northern hemisphere. They have also to be judged from the structure of the nuts and the habitat of the plants. Experiments in this field are also badly needed, as it is of course insufficient to find stomachs full of sedge nuts without checking latent germination power in those found in dung or droppings. Fruits (nuts) of Cyperaceae are generally small to minute, except those of Scirpodendron, a marsh plant, in which they measure 1—1/% by 1 cm (fig. 3). Mostly they have no device adapted to a special- ized kind of dispersal, but some have, for dispersal by water, by animals, by the sea, and a few by wind. Also vegetative dispersal can take place, by rhizomes or tubers. Vegetative dispersal by stolons is for example almost predominant in the very common Cyperus rotundus which very seldom sets fruit. 1974] Cy PERACEAE (Kern) 443 Most data in this paragraph are taken from RIDLEyY’s informative compilation (Dispersal of Plants, 1930). Sea-water. Some sedges which are characteristic of the sandy beach are certainly seaborne. These are Cyperus bulbosus, C. dubius, C. hyalinus, C. pedunculatus (Remirea) (fig. 65), C. radians, C. stoloni- ferus (fig. 50) and furthermore Fimbristylis cymosa and F. sericea. One might conclude that they would all be common species, like C. pedunculatus. They occupy indeed generally wide ranges, but although beaches abound in the archipelago, they are far from common, e.g. Cyperus bulbosus. For their scarcity one can assume either that some are selective for beach microniches or that there is some deficiency with their dispersal capacity. Such questions ask for experimental evidence. The most widespread is Cyperus pedunculatus and this has a specialized floating mechanism, a swollen corky rachilla internode enveloping the nut. Aquatic dispersal. As so many sedges inhabit marshy soil or swamps, stream- and river-banks, and man-made rice-fields, where they often occur in great quantity, fresh water must be a most important vector. Some of these marsh plants possess certain devices. Of some Carex species it is known that they derive buoyancy power by means of the inflated utricle enveloping the nut; Cyperus odoratus has swollen corky rachilla internodes remaining attached to the nut as in Cyperus pedunculatus ; Cyperus cephalotes (fig. 61-62) has a corky pericarp for buoyancy; nuts of Cyperus platystylis with narrow corky edges also float as found by CLARKE (J. Linn. Soc. 21, 1884, 28); Scirpodendron (fig. 3) has large fruits with a stony endocarp surrounded by a thick corky layer keeping them afloat for a long time; in Cladium mariscus the pericarp contains air-cells with thin walls and wide intercellular spaces and has been observed to float for 15 months; also Scirpus maritimus owes floating power to the pericarp structure. However, calculated in percentages the number of species possessing floating power are in distinct minority: normally nuts of Cyperus, Eleocharis, Scirpus, etc. sink in water. It is remarkable how few species have buoyant nuts, yet to observe how widespread and common, sometimes dominant, they occur in swampy places, even though the local dominance is not seldom due to vegetative reproduc- tion, e.g. by stolons in the perennial species. This discrepancy can in part be explained by arguing that even sinking seeds are of course dispersed by running water and floods and furthermore that whole plants or their debris are found in river drift. Cyperus cephalotes (fig. 62) is even a characteristic drifting plant, as described by CoeERT (Trop. Natuur 23, 1934, 13, fig. 8), as is Cyperus platystylis. Also vast numbers of non-buoyant nuts are found in drift and ‘sudd’, some having a means of attaching themselves, by their perianth bristles; even tubers of a Cyperus species have been found in such drift. Also in rice-fields nuts may be dispersed from one terrace to an other by irrigation methods. Though this accidental dispersal must be very common, it cannot well serve to explain dispersal of the majority of species and specimens found in swamps, permanent or temporary, which are not connected by streams or rivers. For this it is assumed that the major vector is the birds. Dispersal by birds. Examination of stomach contents of birds has revealed that especially ducks can really feed on sedge nuts. In one stomach 30.000 nuts were found of a Cyperus, in an other 64.000 of an Eleocharis, and in stomachs and crops also nuts were found of Carex, Cladium, Fimbristylis, Rhynchospora, Scirpus, Scleria, etc. Although, as remarked before, it has insufficiently been checked how far this food is digested, it can be assumed that some nuts will retain germination power and are dispersed with the excreta, a sort of accidental endozoic dispersal. In very few cases the nuts are attractive to birds by a display of contrasting colours, e.g. in Gahnia, where the nuts are yellow, or scarlet, and contrast vividly with the dark-brown to black inflorescences out of which they dangle from the filaments. In Sumatra this has been observed by JACOBSON with Pycnonotus bimaculatus. The nuts are very hard and whether they are digested is not clear. Carex baccans has fleshy red-coloured utricles which are equally attractive to birds. Epizoic dispersal by birds is assumed to be more important, as many sedges have very small seed which may adhere, with mud, to feet, beak and feathers of wading birds. In Europe this has been observed (KERNER) or supposed for species of Carex, Cyperus, Eleocharis, Rhynchospora, Scirpus, and Cladium mariscus. It may well be that bristles of certain species of Scirpus are favourable for epizoic dispersal. A specialized organ which certainly serves for this purpose is found in Uncinia, in which the rachilla protrudes from the utricle and is provided at the apex with a remarkable hook by which the utricles are adhesive to fur or feathers. 444 FLORA MALESIANA [ ser. I, volxas To which distances dispersal is effected by epizoic dispersal can of course not be observed, but in Europe it is assumed to cover several dozens of miles in e.g. Scirpus maritimus, in order to explain isolated localities in inland saline habitats through dispersal by wild-fowl. In East Java several halophilous sedges are found near salt (mud)wells which may be accounted for in this way. However, it remains quite uncertain whether accidental epizoic dispersal may also lead to effective long- distance dispersal, for example, to explain the total range of Scirpus maritimus, which occurs on the northern hemisphere and in Australia, but is extremely rare in Malesia, being a few times collected only in Luzon (in the mountains, but also at low altitude) and in New Guinea (in the high mountains). For, if long-distance transtropical dispersal is assumed to explain these huge disjunctions, short- distance dispersal in the same area must occur infinitely more frequently and this species should then not be so extremely rare in Malesia; this reasoning throws a grave doubt on the assumption of long- distance dispersal. Though in Uncinia effective epizoic dispersal can not be doubted, here also is doubt about its being effective over long distances (1000 km or more). Though on the one hand the range of Uncinia is, not reckoned oceanic disjunctions, fairly coherent and ‘natural’, it matches on the other hand that of many other plants which have quite different ways of dispersal or none (like Nothofagus). So a word of warning is in place against too hasty explanations based on the effectiveness of the specialized function of hooked fruits. Cattle is also assumed to contribute to dispersal of seed of sedges as has been observed in Europe (Carex, Scirpus). Nuts of Fimbristylis globulosa and F. littoralis do not yield to digestion of water buffaloes. This serves only for short distances and anyway such nuts are also dispersed in rice-field areas by water. Ants play a minor role in short-distance dispersal of sedges; ant dispersal has been observed in Carex in Europe where in some species the base of the utricule is provided with a sort of appendage with an oily body (SERNANDER). The role of man is certainly large but difficult to evaluate in extent and illustrate by examples. Traffic, transport, road-building, etc. must have contributed to dispersal of nuts and rhizomes, also sometimes overseas as for example Cyperus sphacelatus and C. aromaticus. In northern Italy several tropical and subtropical aquatic and marsh plant species are found in rice-fields of which the seed is assumed to have been carried as contamination of the rice seed supply. In Italy there is also the remarkable unique locality of Cyperus polystachyos in the isle of Ischia near Naples near the solfatara and hotwells as reported already by G. VON MARTENS (Flora 40, 1857, 346), where it can maintain itself by the grace of local, permanent higher temperature. This occurrence can only be accounted for by accidental dispersal by man from Asia Minor or Egypt, maybe dating back to surviving Crusaders who on return took to this famous holiday resort. A similar occurrence of Cyperaceae outside their natural area was observed by VAN STEENIS, who found Cyperus halpan, a common rice-field plant ascending to some 1650 m, on the summit of Mt Agung (Bali) at c. 3000 m near fumaroles together with a dozen other lowland or hill species. It is assumed that the diaspores of these plants were accidentally dispersed by man during the annual pilgrimage to the summit of this mountain where they can grow and maintain themselves in the subalpine permanent ‘open air hothouses’ near the fumaroles. Likewise R. VAN DER VEEN collected Cyperus cyperinus near hotwells in Lombok on Mt Rindjani at 2000 m, a species ascending normally to c. 1300 m. Wind plays a minor role in dispersal of seeds in the tropics, but it is assumed for the Australasian highland Carpha alpina in the Papuan Alps which possesses a plumose, persistent perianth of some 9 mm length. Here again long-distance dispersal is merely an assumption, because it immediately induces the question why then the genus Eriophorum which is so widely distributed over the northern hemisphere and occurs also in continental tropical-montane SE. Asia has not been capable to invade the West Malesian highlands. Wind has also incidentally been mentioned for Scirpus in which it has been observed that the glume may occasionally remain attached to the nut, acting as a wing. A similar mechanism has been men- tioned for certain Gahnia snecies where the lengthening stamens tear the nut loose and remain some- times entangled around it, but the nuts in Gahnia are heavy and are readily detached. For the various interesting mechanisms loosening the nut in this genus I refer to the discussion under the genus. Morphology. For a long time taxonomists have tried in Cyperaceae to derive the unisexual 1974] Cy PERACEAE (Kern) 445 flower from the bisexual one, viz that of Caricoideae from that of Cyperoideae, but there remained always an unbridgeable gap between these two types, as in the unisexual flowers there is never a vestige of a perianth nor of the other sex. If the opposite course is pursued, however, and we start our reasoning from the unisexual flower this gap can well be bridged. This idea was developed by MATTFELD (Proc. VI. Intern. Bot. Congr. 1936, 330-332) and independently by Ho_trum (Bot. Review 14, 1948, 525-541). It was further developed by me (Adv. Science 19, 1962, 141-148). Like in many other families the primitive structures are found in the tropical representatives. Instead of assessing insight in structure starting from a temperate species of Scirpus, we take Scir- podendron, a very coarse tropical sedge, with broad cutting leaves, large dense inflorescences, and drupaceous fruits attaining a centimetre in size. From the ground-plan of its spikelets with unisexual flowers the other types can easily be derived by (sometimes excessive) reduction, as is illustrated here by a number of diagrams in fig. 1. Starting from the primitive type of Scirpodendron, the other types MAPANIA PARAMAPANIA HYPOLYTRUM 5 CIRPODENDRON ea Bo 8 e 9 VE mein me e a, S5CIRPUS pp LIPOCARPHA CYPERUS ete CAREX Fig. 1. Floral diagrams in several genera of Cyperaceae, with axis, glume and scales all in black, the transverse keeled scales (blank) representing the prophyll (after KERN, 1962). (Mapania, Paramapania, etc.) can easily be derived, until we arrive at Hypolytrum, in which in the partial inflorescence (spikelet) there are no scales between stamens and ovary, the spikelet thus becoming indistinguishable from the diagram of the ‘flowers’ of some Scirpus species. This leads to the view that the ‘flower’ of Cyperoideae, which was so neatly supposed by CLARKE and others to represent a characteristic pentacyclic diagram as is usual in many Monocotyledonous families, though with reductions in the whorls, is really a pseudanthium or synanthium. An other most important feature is the interpretation of the structure of the spikelet and the role of the prophyll. In Cyperaceae as a rule the first leaf of every lateral branch is a 2-keeled prophyll, backing the axis from which the branch arises. KUNTH (in Wiegman’s Arch. Naturgeschichte 1, 2, 1835, 349-353, t. vi) definitely showed that the utricle in Carex, up till then generally taken for a perianth, is homologous with the prophyll, the margins of which are connate up to the top and so has become a bottle-shaped organ enclosing a female flower (later the nut). In Kobresia, as well as in the European genus El/yna and in the African genus Schoenoxiphium — which I all regard as congeneric (see my discussion in Acta Bot. Neerl. 7, 1958, 786-795) — the situation is more primitive than it is in Carex. The prophyll in these genera is either open or more or less connate at the margins, whereas the axis (rachilla) on which it is seated often bears some male flowers. In Carex these male flowers have disappeared and only in a few ~~ 446 FLORA MALESIANA [ser. I, vol. 7° species a reduced rachilla is still present. In Uncinia the vestige of the rachilla is at its apex transformed into a peculiar recurved hook which can be accepted to be homologous with an empty bract or reduced male flower; this hook is designated as a specialisation for epizoic dispersal. The idea of NELMES (Reinwardtia 1, 1951, 222-225; Kew Bull. 1951, 427—436) that the unispicate Carices from Europe and North America with a vestigial rachilla belong to the genealogy of Uncinia is untenable. According to the interpretation given above the morphological derivation of the structure of the spikelet and the essential role in it by the prophyll leads to the assumption that the most primitive state, with unisexual flowers, is still represented in the Mapanieae. Starting from the situation in Scirpodendron one can observe that in this tribe the prophyll (consisting of two transverse scales, which may be connate, in Hypolytrum sometimes even on both sides and thereby becoming more or less utriculiform) and flowers can be arranged in a series of successive reduction (vestigial to completely absent), leading finally in Cyperoideae to excessive reduction and the origin of pseudanthia. Another line of excessive reduction has led to the structure as is found in Caricoideae in which, however, the unisexuality of the flowers has been conserved. top-shaped top-shaped top-shaped broadly ellipsoid mushroom- top-shaped shaped Fig. 2. Embryo types in Cyperaceae. In the picture of Cyperus the various parts have been named in detail: col' is upper lip of coleoptyle, co/* lower lip of the same, cot is cotyledon, /' is primordium of first leaf, /? is primordium of second leaf, p is germination pore, r.c. is primordium of root-cap (simplified after VAN DER VEKEN). Coleoptyle Pax (in E. & P. Nat. Pfl. Fam. 2, 1887, 105) suggested that there would be a basic difference in Cyperaceae, advancing that the structure of the spikelets in Cyperoideae is monopodial and in tribe Rhynchosporeae sympodial. On this account ASCHERSON & GRAEBNER (Synopsis 2, 2, 1904, 339) raised the latter tribe to subfamily rank. This is, however, fallacious: the structure of the spikelets is in all Cyperaceae probably sympodial (KERN, 1962, /.c.). In passing it may be remarked that some genera were assigned to Rhynchosporeae by erroneous interpretation, e.g. Remirea which is simply a Cyperus in which a rachilla internode is transformed into a corky organ serving buoyancy (see the fuller account under the species). On the other hand Oreobolus is distinctly allied to Rhynchosporeae and must be placed in the Schoenoid affinity. Embryography. Interesting data on the embryography were published by P. VAN DER VEKEN (Bijdrage tot de systematische embryologie der Cyperaceae-Cyperoideae. Thesis, Leuven, 1964, pp. 230, illustr., in Dutch; Bull. Jard. Bot. Brux. 35, 1965, 285-354), who examined the embryo in 342 spp. of Cyperoideae. He found 6 main types which he named the Cyperus, Carex, Schoenus, Fim- bristylis, Bulbostylis, and Scirpus types. Some of these are depicted in fig. 2. It has appeared that in homogeneous taxa, e.g. in Cyperus sens. lat., always the same embryo type is found. This is also true for e.g. Schoenus, Bulbostylis, Eleocharis, and Fuirena, on the generic level. In our opinion a type may also be characteristic on infra-generic level, e.g. in Fimbristylis, where 1974] Cy PERACEAE (Kern) 447 sect. Abildgaardia and Actinoschoenus have embryotypes which differ from that in Fimbristylis. In Scirpus there are not less than 7 different embryo types. Conversely, one embryo type may occur in more than one genus, e.g. Lipocarpha has the Cyperus type of embryo. Whereas the homogeneity of Cyperus is supported by embryography, one may conclude that Scirpus is distinctly heterogeneous. This might be an argument for systematic heterogeneity involving a possibly polyphyletic assemblage; each of the natural sections in Scirpus possesses namely according to VAN DER VEKEN (1964, p. 140) only one embryo type. Though Scirpus confervoides (Websteria) has its own embryo type, similar to that of Eleocharis but somewhat more differentiated, I have kept it in Scirpus for flower-morphological reasons. As to Actinoschoenus, which has the Carex type of embryo combined with a Eucyperoid leaf-anatomy | have kept this in Fimbristylis, though VAN DER VEKEN (1964, p. 166) suggests it to belong to Rhynchos- poreae (see p. 591 under Fimbristylis thouarsii). Vegetative anatomy. Baas, Notes Jodrell Lab. 6 (1969) 1—20 (Hypolytreae); DUvVAL—JOUVE, Bull. Soc. Bot. Fr. 20(1873)91—95; GOVINDARAJALU, J. Linn. Soc. Bot. 59 (1966) 289-304 (Bulbostylis), ibid. 62 (1969) 27—40 (Fuirena), and ibid. 62 (1969) 41—58 (Cyperus); GREGORY & METCALFE, Notes Jodrell Lab. 5 (1967) 1—17 (bibliography); KAPHAHN, Beih. Bot. Centralbl. 18 (1904—05) 233-272 (silica); KoyAMA, Mem. N.Y. Bot. Gard. 15 (1966) 136—159 (Mapanieae = Hypolytreae) and ibid. 16 (1967) 46—70 (Sclerieae); KUKKONEN, Ann. Bot. 31 (1967) 523-544 (Uncinia); METCALFE, Am. J. Bot. 56 (1969) 782-790 (relationships and anatomy) and Anatomy of thé Monocotyledons V. Cyperaceae. Oxford (1971) 597 pp.; METCALFE & GreGory, Notes Jodrell Lab. 1 (1964) 1-11 (descriptive terms); PFEIFFER, Beih. Bot. Centralbl. 44 (1927) 90-176 (leaf anatomy of many genera) and Ber. Deut. Bot. Ges. 47 (1929) 78—82 (silica); RIKLI, Jahrb. Wiss. Bot. 27 (1895) 485—580 (leaf anatomy, bundle sheaths); SCHUYLER, Mitt. Bot. Staatssamml. Miinchen 10(1971)577—585 (Scirpeae); SHARMA, Curr. Sci. 41 (1972) 494—497 (Scirpus squarrosus); SHARMA & MEHRA, Res. Bull. Punjab Univ. n.s. 21 (1970) 119-128 (Kobresia) and Bot. Gaz. 133 (1972) 87—95 (Fimbristylis). METCALFE (1971, /.c.) provided a comprehensive survey of anatomical structures occurring within the family. His reference book also contains a very full bibliography and summaries by GREGORY of anatomical data collected in the past. Most of the following is taken from this book. Anatomical characters to be used for diagnostic purposes or for discussions of affinities are the following. Shape of lamina in transverse section taken midway the apex and the sheathing base. In the gener- ally occurring dorsiventral leaf type the lamina may be V-shaped (with flanged, medianly grooved and thick subtypes), corrugate, crescentiform (thin or thick), inversely W-shaped or triangular. The shapes of isobilateral leaves (Lepidosperma, Machaerina) range from subcruciform, winged fusiform, tetragonal, elliptical to constricted elliptical in transverse section. The pseudodorsiventral leaf type (restricted to Cladium) is V-shaped. Cylindrical leaves with a circular to subcircular shape of the transverse section occur in some species of Lipocarpha and Machaerina. METCALFE(1969, 1971) has suggested a derivation of the cylindrical, isobilateral and pseudodorsiventral leaf types from the dorsiventral ‘basic’ type. Prickle hairs are very common in the family and of little taxonomic interest. Other hair types are more infrequent and have some diagnostic value. They include adpressed hairs with tips directed towards the leaf apex (some spp. of Costularia, Scirpus and Scleria), unicellular flexible hairs (some spp. of Carex, Fimbristylis, Fuirena, Rhynchospora, Schoenus and Scleria), unicellular long and stiff hairs (some spp. of Carex, Fuirena, Rhynchospora and Schoenus), unicellular short and stiff hairs (some spp. of Carex, Fuirena, Gahnia, Lipocarpha, Oreobolus and Scleria) and finally lobed hairs in Schoenus (S.apogon). Papillae are present ina number of genera. Insome species of Carex, Cladium, Fimbristylis, Lepidosperma, Lepironia, Machaerina and Scleria the papillae are restricted to the cells around the stomata and overarch the latter. Epidermis. Stomata are generally paracytic but tend to be tetracytic in some genera. Silica bodies are of almost universal occurrence in the leaf epidermis of Cyperaceae and have proved to be of great taxonomic importance. They are absent or very doubtfully present in an unusual form in Hypolytrum and Lepironia, finely particulate silica has been noted in some species of Costularia, Fuirena, Oreobolus, Paramapania and Scleria. The bodies may be dome-shaped with their bases resting in the sinuations of anticlinal walls in Oreobolus or be present as solitary cones at the apices of sinuations of anticlinal walls (some spp. of Costularia and Scleria); cubical bodies associated with other types of silica bodies 448 FLORA MALESIANA [ser. I, vol. 7° occur in Paramapania and Rhynchospora; warty spherical to hemispherical bodies in Capitularina and Sc/eria; bridge-shaped bodies in Mapania and Thoracostachyum; wedge-shaped in Scirpodendron and Thoracostachyum. The most common type 1s the conical type with its base resting on the inner periclinal wall. This type intergrades with the nodular one. These types occur in the majority of genera and may be present in different numbers per cell, the number and arrangement of the bodies being of diagnostic value. The epidermis may include bulliform cells. In a number of Cyperaceae a parenchymatous or partly or wholly sclerenchymatous hypodermis is present. The presence or absence of bulliform cells and of a hypodermis is of restricted taxonomic value. Variations have been recorded below the species level. The distribution of sclerenchyma, mainly accompanying the vascular bundles shows an enormous range of variation within the family and the shapes of sclerenchyma girders, caps or strands may be of considerable diagnostic value at the species level. The mesophyll of the leaves may be virtually homogeneous, differentiated in palissade and spongy chlorenchyma or may contain radiate chlorenchyma (that is radiating from the vascular bundles). The latter type is conspicuously represented in most or all species of Bulbostylis, Cyperus, Fimbristylis and Lipocarpha, some or all species of Fimbristylis, Fuirena, Hypolytrum and Rhynchospora show this feature less markedly. Air cavities or areas with translucent cells, which give rise to such cavities occur in a great number of genera. The vascular bundles of the leaf are collateral and in most genera arranged in a single row as seen in transverse section. In cylindrical, isobilateral, pseudodorsiventral and a few dorsiventral leaves the arrangement differs. In cylindrical and isobilateral leaves they are arranged along the whole periph- ery of the transverse section. In the isobilateral leaves of Lepidosperma and Machaerina and the pseudodorsiventral leaves of Cladium they occur in opposite pairs with the xylem poles facing each other. This situation should not be confused with the arrangement of vascular bundles in two ranks in some dorsiventral leaves, each vascular bundle having a normal xylem and phloem arrangement. This occurs in species of Costularia, Cyperus, Lipocarpha, Mapania, Rhynchospora, Scirpodendron and Thoracostachyum. The vascular bundles are usually surrounded by a two-layered bundle sheath, with or without adaxial and/or abaxial merging with the accompanying sclerenchyma. Three basic types occur in Cyperaceae: 1) inner sheath sclerenchymatous, outer sheath parenchymatous (most common), 2) inner sheath parenchymatous, outer sheath sclerenchymatous (some species of Cyperus, Lipocarpha), and 3) bundle sheath three-layered, outer and inner sheath parenchymatous, middle layer sclerenchymatous (spp. of Bulbostylis, Carpha, Fimbristylis and Mapania). Some species show intermediate types of bundle sheaths. Culm anatomy also provides useful diagnostic features. These also include outline in transverse section (triangular, circular, hemicircular, quadrangular, pentagonal, hexagonal, polygonal, trape- zoid, winged fusiform, scutiform crescentiform and irregular), presence or absence and distribution of air cavities, arrangement of vascular bundles, and sclerenchyma. Roots and rhizomes are less well known anatomically, and the characters thought to be of diag- nostic value need further testing. Relationships with other families. Gramineae are different from Cyperaceae mainly in epidermal structure. In Gramineae the silica bodies are morphologically entirely different and the epidermal cells are Clearly differentiated in short and long cells. The latter feature, however, tends to be present in a few Cyperaceae as well. METCALFE (1971) has suggested that any phylogenetic relationship between the two families must be very remote. Restionaceae differ from most Cyperaceae in their culm anatomy. In Restionaceae there is a characteristic sequence of tissues: epidermis, chlorenchyma, parenchyma sheath, sclerenchyma sheath (enclosing peripheral vascular bundles) and medullary vascular bundles embedded in central parenchymatous ground tissue. In Cyperaceae some genera also possess a sclerenchyma ring in the culm, but the very regular zonation of tissues as in Restionaceae is always absent so that the resemblance of those Cyperaceae to Restionaceae remains superficial. Silica deposits occurring in some Restionaceae are either spheroidal nodular or are present as silica sand. Conical bodies, so characteristic of the majority of Cyperaceae are absent. Nodular bodies occur in a number of Cyperaceous genera but intergrade here with the conical type. According to METCALFE (1971) Cyperaceae are anatomically more similar to Juncaceae than to any other plant group. Juncaceae do not possess silica bodies, but this is also true for several genera of Cyperaceae. It may be noted in passing that most of the Cyperaceae lacking silica are amongst the genera belonging to the tribe 1974] CYPERACEAE (Kern) 449 Hypolytreae. This tribe is generally regarded as primitive within the family. It seems therefore justified to suggest that the plant group ancestral to Cyperaceae and Juncaceae also lacked silica bodies. The affinities of Cyperaceae with Juncaceae are also supported from other sources of evidence (cf. METCALFE, 1971, pp. 42 & 43). Anatomical affinities within the family. It is impossible to define the tribes and subfamilies anatomi- cally. All tribes contain such a range of anatomical variation and the overlap with other tribes is so extensive that only vague suggestions can be made. The tribe Hypolvtreae exhibits a great range of variation. The unusual bridge- and wedge-shaped silica bodies of some genera only occur outside this tribe in a few genera of the Rhynchosporeae. The absence of distinct silica bodies is moreover more or less confined to genera belonging to Hypolytreae. Several genera, notably Chrysithrix and Choriz- andra, show many resemblances with members of Rhynchosporeae (cf. BAAS, /.c.,and METCALFE, 1971, l.c.). All other tribes typically contain some sort of conical silica bodies, and Capitularina of the Hypolytreae has also been recorded to contain this type. In my opinion this genus is rather remote from the other Hypolytreae on other anatomical grounds as well (BAAS, /.c.). The tribe Cypereae does not exhibit any striking anatomical features which make them stand out within Cyperaceae. Anatomically there are no objections against including genera usually treated in the separate tribe Scirpeae in Cypereae. Rhynchosporeae vary considerably in their anatomy. Yet Oreobolus is rather isolated anatomically. Some genera resemble Hypolytreae (see above). The tendency in some non- Malesian genera to have wedge-shaped silica bodies also points to an affinity between the two tribes. Sclerieae usually possess silica bodies of deviating types in addition to the common conical ones. This seems to be the only outstanding anatomical character within the tribe. Cariceae are quite homogeneous anatomically. Most of the characters shared by the genera constituting this tribe are, however, also of widespread occurrence throughout the other tribes. Vegetative anatomy does not provide supporting evidence to distinguish two subfamilies Cyperoideae and Caricoideae. — P. BAAS. Palynology. The pollen grains in Cyperaceae are, as far as investigated, degraded tetrads, in which the wall of the pollen mother cell develops into an exine, while three of the four nuclei resulting from the tetrad division degenerate. SELLING (B. P. Bish. Mus. Spec. Bull. 38, 1947, 350—352) has created the term ‘pseudomonads’ for such grains (see the survey |_ CRANWELL, Bull. Auckl. Inst. Mus. 3, 1953, 42-47). ERDTMAN (Pollen Morph. & Pl. Taxon. 1952, 141-142), however, suspects that the pollen of Mapania may not belong to this category, although KoyAMa (cited in Cronquist, Evol. Class. Fl. Pl. 1968, 341—342) affirms its pseudomonad nature. Evidently careful studies are still necessary to solve such problems. Exine stratification is stated by ERDTMAN, /.c., to be similar to that found in other Angiospermous pollen grains. Size varies between 16—66 1, shape may be spheri- cal, pear-shaped or flattened triangular. Apertures are as a rule indistinct, variable in number (0—4) and in position, although one aperture may dominate. Due to lack of detailed studies, it is difficult to discuss the taxonomic significance of pollen characters in Cyperaceae. In Carex pollen morphology is known to be rather uniform and its pear-shaped, multiaperturate pollen type is also known from Bulbostylis, Costularia, Cyperus, Fimbristylis, Fuirena, Gahnia, Oreobolus, Scirpus, Scleria, Tetraria, and Uncinia. The pollen of the genera Mapania and Thoracostachyum is clearly monoporate and also differs in shape and, possibly, in wall structure from the Carex type. The formation of pseudomonads 1s unique for Cyperaceae. An analogous mode of pollen develop- ment occurs, however, in Juncaceae. Here all four nuclei develop normally, resulting in a tetrad in which the outer exine is also formed by the pollen mother cell wall, while the inner walls are thinner. This is generally taken to suggest that Cyperaceae are closely related to Juncaceae and may in fact have been derived from the latter family (cf. TAKHTAJAN, Evol. Angiosp. 1959, 272). CRONQUIST, /.c., however, points to the resemblance between the distinctly monoporate Mapania pollen grains and those of Gramineae and Restionaceae and considers it unlikely that Cyperaceae inherited their pseudo- monads from some earlier group. — J. MULLER. Chemotaxonomy. The chemical characters of this huge family were summarized 10 years ago (HEGNAUER, Chemotaxonomie der Pflanzen 2, 1963, 124—133). Accumulation of silicic acid, which is deposited in highly characteristic, anatomically easily demonstrable (spodograms) patterns, usual absence of oxalate of lime, the relatively frequent occurrence of flavonoid (i.e. condensed) tannins probably derived from leucoanthocyanins or protoanthocyanidins and preponderance of starch as a carbohydrate reserve seem to be typical of Cyperaceous plants. Some taxa produce essential oils 450 FLORA MALESIANA [ser. I, vol. 7° which are deposited in more or less elongated oil cells. Cyanogenic compounds, alkaloids and sapo- nins seem to occur exceptionally only in this family. The scantiness of known chemical facts was stressed in 1963. In the meantime phytochemical research has not totally neglected Cyperaceae. Much new information about alkaloids, flavonoids, quinones and phenolic ketones of roots and rhizomes, and about the chemical composition of essential oils of several species of Cyperus became available in recent time. The alkaloids of Carex brevicollis, harman, brevicollin and brevicarin, were shown by Russian authors to be all B-carboline-type bases. Alkaloids were reported for several other Cyperaceae; their structures, however, have still to be established. The essential oils of rootstocks of Cyperus articulatus, C. rotundus and C. scariosus were studied intensively by several groups of workers; sesquiterpenes and sesquiterpenic alcohols and ketones are their main constituents. The taxonomically probably most interesting developments, however, concern polyphenolic compounds. E. C. BATE-SMITH published his survey of leaf phenolics of monocotyledonous plants (J. Linn. Soc. Bot. 60, 1968, 325). He noted a strong resemblance between Gramineae and Caricoideae and a tendency for Cyperoideae (Scirpoideae) to be “much more regular or ‘primitive’ in their phenolic pattern”; his sampling, however, was very poor with regard to Cyperaceae. H. T.CLIFFORD and J. B. HARBORNE (Phytochemistry 8, 1969, 123) studied the flavonoid pigments of inflorescences of sedges and arrived at the conclusions that grasses and sedges are radically different and that in wind-pollinated plants the correlation between flavonoid chemistry and taxonomy breaks down. The common anthocyanidins, flavonols and flavones seemed to be lacking in sedges; 3-deoxyanthocyanidins (carexidin), aurones (aureusidin), chalcones (okanin) and leuco-anthocyanidins were shown to occur in Cyperaceae, but to be erratically distributed in the family. Later HARBORNE (Phytochemistry 10, 1971, 1569) studied leaf pigments of 62 species representing 11 genera. This time a rather close resemblance between grasses and sedges and additionally palms was noted. Glycoflavones and the flavones luteolin and tricin are widespread in leaves of sedges and flavonols including rutin were found only in Eriophorum latifolium, Fuirena pubescens and in 7 of the 44 species of Carex investigated. The predominant occurrence of flavonols in Carex flava and related species suggests, according to HARBORNE, that these are the more primitive taxa within the genus. Evidently much more facts about the chemistry and distribution of phenolic compounds are needed before sound taxonomic conclusions become possible. Investigations aiming at extension of our knowledge were started by I. KUKKONEN (Mitt. Bot. Staatssamml. Miinchen 10, 1971, 622); many phenolics were demonstrated by him to occur in sedges; before a true evaluation of facts becomes possible, identification of these compounds is necessary. However, for the study of species aggregates chromatographic comparisons of leaf phenolics may be valuable without identification of the compounds concerned (R. I. Ep1Ger, Trans. Kansas Acad. Sc. 69, 1966, 152: Carex aggregata complex). Totally new compounds were detected in rootstocks of Australian Cyperaceae by R. J. ALLAN et al. (Tetrahedron Letters 1969, p. 4669 & 4673; ibid. 1970, p. 3945). These are the cyperaquinones and phenolic ketones, both probably derived from isopreny- lated and acylated phloroglucinols. The cyperaquinones were found to occur in species of Cyperus, Fimbristylis,and in Cyperus pedunculatus (R.BR.) KERN(Remirea maritima L.). Only the last mentioned species was investigated accurately for phenolic ketones; remirol, remiridiol, preemirol and isoevo- dionol occur in its rootstocks. These phloroglucinol-derived new constituents of sedges seem to be promising for the study of infra- and intergeneric relationships, but less so for the study of relation- ships between families. Summarizing it may be stated that even to-day our phytochemical knowledge of Cyperaceae is scanty. The facts available at present indicate that grasses and sedges share several striking features. Therefore the phytochemical data do not contradict the hypothesis postulating a common origin for these two large families of wind-pollinated Monocotyledons. — R. HEGNAUER. Taxonomical affinities. There is no unanimity of opinion about the affinities of Cyperaceae. As will be observed from the introductory chapters, the vegetative anatomy and palynology would suggest a remote tie with Juncaceae, chemotaxonomical evidence would point to Gramineae, while morphological structure would according to HoLTTuM point to ancient relationship with Pandanales, suiting the habit and drupaceous fruit of the primitive tropical members and the theory of the simple, unisexual primitive flower. 1974] CYPERACEAE (Kern) 451 Taxonomy and subdivision of the family. In spite of the much diversified structure of spikelets and flowers, Cyperaceae present in the customary circumscription, also adopted here, a natural unit. To GILLY (Iowa State Coll. J. Sc. 26, 1952, 210) the tribe Cariceae, comprising the large genus Carex and some closely allied smaller genera, is as nearly related to Gramineae as to Cyperaceae in the restric- ted sense, but his proposal to raise the tribe Cariceae to family rank as Kobresiaceae has found little support. According to BULLOCK (Taxon 7, 1958, 11), NELMES intended to raise some African genera to family rank as Coleochloaceae (Coleochloa, Trilepis, and Microdracoides) but he did not publish on this. These genera possess indeed some characters not found in other Cyperaceae, amongst others open sheaths of the cauline leaves, but they seem to me insufficient to warrant the distinction of a separate family. More important than the attempts to split Cyperaceae into smaller families is the tracing of the interrelationships within the family. Here the main problem is whether there is indeed an unbridge- able gap yawning between the monoclinous and diclinous members. The subdivision into subfamilies and tribes adopted below, and the arrangement of the Malesian genera of Cyperaceae, are mainly based on the ideas about relationships in this family as developed by MATTFEELD (Proc. 6th Int. Bot. Congr. A’dam 1, 1936, 330—332), HoLTTUM (Bot. Review 14, 1948, 525-541) and myself (Brit. Ass. Adv. of Sci. 19 (n. 78), 1962, 141—148) as explained in the chapter on morphology. Apart from some unimportant deviations they therefore agree with the system followed by SCHULTZE-MOTEL in the 18th edition of ENGLER’s Syllabus (1964). Diametrically opposed to it was the opinion prevalent among earlier cyperologists such as BOECK- ELER, CLARKE, PAX, and KUKENTHAL, and the system recently elaborated by the Japanese cyperol- ogist T. KOYAMA (J. Fac. Sc. Univ. Tokyo III, 8, 1961, 37-64). However different from each other their systems may be, they all start from the supposition that the strictly unisexual flowers in Cari- coideae (Carex, Scleria, and closely allied smaller genera) are reductions of the more primitive bisexual flowers in Cyperoideae (Scirpus etc.), which in their turn can be derived from the pentacyclic ground-plan of the monocotyledonous flower. According to this theory Cyperaceae with bisexual flowers must come first in the system, and Hypolytreae (Scirpodendron, Mapania, etc.) last, because their flowers are reduced to such an extent that they merely consist either of a single stamen or of an ovary. The structure in the axils of the glumes in Hypolytreae is considered a strongly reduced partial inflorescence. A serious objection to this theory is that it cannot bridge the gap between Cyperaceae with strictly unisexual flowers and those with bisexual ones, especially not because there has never been found a vestige of the other sex nor of hypogynous bristles or scales in the flowers of Caricoideae. Asearly as 1877 BENTHAM (J. Linn. Soc. Lond. 15, p. 506—512) defended the thesis that the so-called partial inflorescence in Hypolytreae is homologous with the bisexual Cyperaceous flower. In agree- ment with this view both MATTFELD and HOLTTUM take the flowers in Cyperoideae for synanthia, i.e. partial inflorescences ultimately reduced to functionally ‘bisexual flowers’. Several more or less advanced stages of reduction are found in Hypo/ytreae, which tribe therefore should be placed at the beginning of Cyperoideae, not at the end of Caricoideae. In the spikelets of the tropical Hypolytreae the glumes subtend a bisexual structure consisting of a terminal female flower and two or more lateral monandrous male ones, the latter seated in the axil of a scale. Two outer practically opposite scales are characterized by their being navicular and ciliate on the keel. They may be connate on the adaxial side (e.g. in several species of Hypolytrum) and the resulting 2-keeled bidentate scale cannot be dis- tinguished from the monocotyledonous prophyllum. Usually there is a varying number of flat glabrous scales between the ovary and the navicular scales, more or less distinctly arranged in one or two whorls and at least partly not subtending a stamen. Starting from the complex condition in Scirpodendron, in which there is a large but varying number of scales with stamens, the gradual reduc- tion can be followed through Lepironia, Chorizandra (Australian), Mapania, and Paramapania to the structures in Diplazia (South American), and Hypolytrum, where the scales surround the stamens which are arranged round the ovary, this being in effect the condition in several Cyperoideae. Such functionally bisexual ‘flowers’ are thus assumed to be synanthia, being derived from reduced partial inflorescences of unisexual flowers, as I have illustrated in 1962 (Adv. Sci. 19, July 1962, 141-148, 24 fig.). Compare fig. 1. In this way the gap apparently yawning between the unisexual flowers in Cari- coideae and the bisexual ones (synanthia) in Cyperoideae can be bridged. 452 FLORA MALESIANA [ser. I; vole FF The system here adopted is, as far as Malesian genera are concerned, as follows: A. SUBFAMILY CYPEROIDEAE I. Tribe Hypolytreae: 1. Scirpodendron 4. Thoracostachyum 6. Paramapania 2. Capitularina 5. Mapania 7. Hypolytrum 3. Lepironia II. Tribe Cypereae: 8. Scirpus 11. Eleocharis 13. Fimbristylis 9. Fuirena 12. Bulbostylis 14. Cyperus 10. Lipocarpha III. Tribe Rhynchosporeae: 15. Tetraria 19. Tricostularia 23. Machaerina 16. Costularia 20. Schoenus 24. Gahnia 17. Carpha 21. Oreobolus 25. Rhynchospora 18. Lepidosperma 22. Cladium B. SUBFAMILY CARICOIDEAE IV. Tribe Sclerieae: 26. Scleria V. Tribe Cariceae: 27. Kobresia 28. Carex 29. Uncinia SYNOPTICAL KEY TO THE MALESIAN GENERA (Reflecting the adopted system, not intended as a practical key) 1. Bisexual flowers (synanthia) present (by abortion of androecium or gynoecium part of the flowers maybe unisexual) . . . A. SUBFAM. CYPEROIDEAE 2. Outer 2 hypogynous scales (in H ypolytrum ‘the only o ones ‘present) folded, transverse and sharply keeled, ciliate or spinulose on the keel, free, or sometimes connate on the adaxial side . J/. Tribe Hypolytreae 3. Hypogynous scales numerous, indefinite in number. 4. Nut deeply grooved, 1—-I5 cmJong 6 <- 6) & eel eh be da ee sy 1 Sagi 4. Nut not grooved, much smaller. 5. Stems 5-angular. Inflorescence terminal, capitate . . rn Tener a ree A COn lsc 5. Stems terete. Inflorescence a single, pseudolateral spikelet 3. Hypogynous scales definite in number, up to 6. 6. Hypogynous scales 5 or 6. 7. Hypogynous scales 6. Stamens 3(—4). Outer scales ciliate on the keel. 3. Lepironia 8. Inflorescence paniculate. Exocarp thin, hard. . . 4. Thoracostachyum 8. Inflorescence capitate or consisting of a single spikelet. ‘Exocarp corky or r fleshy 3 5. Mapania 7. Hypogynous scales usually 5. Stamens 2(—3). Outer scales coarsely brown-spinulose on the keel. 6. Paramapania 6. Hypogynous scales 2 y ‘ 2. Hypogynous scales, when present, otherwise, or - perianth consisting of bristles. 9. Spikelets as a rule several- to many-flowered; only 1 or 2 lower glumes empty. . II. 10. Perianth represented by hypogynous bristles or scales. 11. Flowering stems leafy, at least at the base. Style continuous with the sa 12. Perianth consisting of bristles, exceptionally of 4 scales ey , 12. Perianth consisting of 2 or 3 scales. 13. Hypogynous scales 3, often with intervening bristles. 13. Hypogynous scales 2, median, hyaline 11. Stems leafless. Style articulated with the ovary, its base persistent on the nut 10. Perianth absent. 14. Nut crowned by the persistent style-base 14. Nut not crowned by the persistent style-base. 7. Hypolytrum Tribe Cypereae 8. Scirpus 9. Fuirena 10. Lipocarpha 11. Eleocharis 12. Bulbostylis 15 Style articulated with the ovary) 2. \Gouk) 2). 2 gee oul dee daa 15. Style continuous with the ovary. 16. Glumes distichous . 16. Glumes spiral; see 8. Scirpus. 9. Spikelets as a rule very few-flowered (mostly 1-2-flowered); usually several lower glumes 14. Cyperus empty. III. Tribe Rhynchosporeae 17. Style continuous with the ovary. 18. Stamens 6. Appendage of the connective nearly as long as the anther-cells; lower flower male. 15. Tetraria 1974] CYPERACEAE (Kern) 453 18. Stamens fewer (exceptionally 6 in some species of 24. Gahnia). 19. Perianth consisting of bristles, long-plumose throughout. 20. Style hispidulous. Inflorescence large, paniculate . . . . . . . +. +. 416. Costularia 20. Style glabrous. Inflorescence small, almost capitate . . oat ie ei eCarpha 19. Hypogynous scales or bristles, when present, not long- plumose. 21. Hypogynous scales incrassate after anthesis . . . . . 18. Lepidosperma 21. Hypogynous scales or bristles not incrassate after anthesis, or perianth absent. 22. Hypogynous scales densely white-hairy . . = SY Os Tricostalars 22. Hypogynous scales not densely white-hairy, or perianth absent. 23. Internodes of the rachilla very short between the lower (empty) glumes, elongate and prominently zigzag between the upper (flower-bearing) glumes. Glumes distichous . . . 20. Schoenus 23. Internodes of the rachilla not so markedly different in length. 24. Perianth consisting of 6hypogynous scales . . . . . . . +. +. +. 21. Oreobolus 24. Perianth otherwise, or absent. 25. Nut drupe-like, borne ona disk not falling off with the nut proper. Stems hollow 22. Cladium 25. Nut not borne on a disk. Stems ay 26. Glumes distichous. . . . So OT 2) gee Ee. oe Se AC HAGKING 26. Glumes spiral . . PES APL a, OE 24 Gahnia 17. Style articulated with the ovary, its base persistent ¢ onthe nut. . . 25. Rhynchospora 1. All flowers strictly unisexual . _ B. SUBFAM. CARICOIDEAE 27. Female flowers not enclosed by < a sac-like organ (a modified prophyll). IV. Tribe Sclerieae. 26. Scleria 27. Female flowers enclosed by a sac-like modified prophyll . . . . . . . . Vz. Tribe Cariceae 28. Modified prophyll more or less open on the abaxial side . . . . . . . +. +. 27. Kobresia 28. Modified prophyll closed throughout its length (utricle). 29. No needle-like hooked rachilla inside the utricle . i e5h2e-. Carex 29. A needle-like rachilla hooked at the top inside the utricle present. Inflorescence always a single spikelet. 29. Uncinia Useful & noxious plants. The following data are very concise and mostly taken from HEYNE (Nutt. Pl. 1927). See for more details in the text under the species. Various sedges are used for plaiting mats and thatching, the most commonly used being Cyperus malaccensis, Eleocharis dulcis, Lepironia, Scirpodendron, Scirpus grossus, S. lacustris, S. litoralis, S. mucronatus, and Thoracostachyum sumatranum. The stems of Eleocharis sphacelatus are in New Guinea used to make rush skirts for women. A peculiar use is made by fishermen of Cyperus malaccensis in Java (see p. 616 and fig. 53). The tubers of Eleocharis dulcis are edible and this species is sometimes cultivated for this purpose; possibly Malays brought it to the Northern Territory together with Egyptian Lotus. The apex of the stem and the leaf-bases of the sprouts of Gahnia javanica have a most agreeable sweet taste of nuts, but serve only as a titbit to the wandering naturalist. In Papua the rhizomes of Machaerina articulata are eaten. In north Central Java (near Pemalang) stem-pieces of Cyperus malaccensis are thickly inserted in ropes which are used to catch fry of the bandéng fish (VAN STEENIS, Trop. Natuur 29, 1940, 20, fig.). Noxious weeds are Cyperus rotundus etc. which by their subterranean tubers or rhizomes are difficult to eradicate. Several Cyperaceae occurring in great quantity in marshes and swamps invade the wet rice-fields. Most common paddy field sedges are Cyperus difformis, C. halpan, C. iria, Fimbristylis littoralis and Scirpus juncoides. They are of course weeded out, but when the rice is harvested there are still many which escaped weeding and are abundantly fruiting. As in many places the rice harvest is alternated with a dry farming crop, the seed (nuts) of the Cyperaceae must remain dormant in the soil because during the dry farming a quite different weed flora comes up. Such areas have hence a ‘double weed flora’, and consequently the seed or fruit of the dryland farming weeds must also be dormant during the wet period of the growing of rice. This is an interesting feature not yet fully disclosed, because we would like to know what factors are responsible for the dormancy of both categories, the secret of the factors inhibiting their germina- tion. Note. This revision will be published in two instalments due to the fact that the treatment of Carex and Uncinia is not yet finished. Acknowledgement. Since 1971 my failing eyesight prohibited actual research work and I 454 FLORA MALESIANA [ser. I, vol. 7° greatly appreciate the essential assistance of the general editor for composing the preliminary version of the introductory chapters, especially those on the ecology, dispersal, and plant geography, for my approval. KEY TO THE GENERA 1. Bisexual flowers always present (part of the flowers may be male or female by abortion of the other sex). 2. Flowers (the organs in the axils of the glumes of the spikelet) naked, i.e. not surrounded by a perianth consisting of hypogynous bristles and/or scales. 3. Nut crowned by the persistent style-base which is articulated with the nut proper. 4. Leaves reduced to bladeless sheaths. Inflorescence consisting of a single terminal spikelet. 5. Nut 114-2 mm long, seated on a conspicuous, /4—1 mm long gynophore. Spikelet 6-15 by 4-6 mm, plumes, 4—9 Mi lone): a Sao eu ee ee . . 13. Fimbristylis (tetragona) 5. Nut smaller, without gynophore. Spikelet and glumes smaller . . . . . 11. Eleocharis spp. 4. Leaf-blades well-developed. 6. Persistent style-base forming a minute dark button much narrower than the trigonous nut. Leaf- sheaths (if not too old) bearded in the throat with long white hairs. Stigmas 3. Annuals with capillary TEAWER. BAG 4 6 cde yas Bt & pit eds neni s Mal 2S. boars e481 dew dale 6. Persistent style-base not button-like. Leaf-sheaths not bearded. 7. Persistent style-base saddle-shaped, as broad as the biconvex nut. Stigmas 2. 25. Rhynchospora (gracillima) 7. Persistent style-base not saddle-shaped. Stigmas3 . . . . . 13. Fimbristylis (thomsonii) 3. Nut not crowned by the persistent style-base; when the nut is beaked then the beak continuous with the nut. 8. Style dilated at the base, articulated with the ovary, often fimbriate. Glumes spirally arranged, or more rarely distichous. Stigmas 2 or 3; nut 2-or3-sided. . . . . . . . ~~. 413. Fimbristylis spp. 8. Style continuous with the ovary. 9. Internodes of the rachilla very short between the lower (empty) glumes, elongate and prominently zigzag between the upper (flower-bearing) glumes; flowers (or fruits) in the hollows of the rachilla. Glumes distichous, frequently dark-coloured . . . . . . . 20. Schoenus spp. 9. Internodes of the rachilla not so markedly different in length. 10. Spikelets several- to many-flowered; as a rule only 1—2 lower glumes empty. LEGUMES IstichOUSIViaALrANSCOtet of) ecweeen aes el ae he Wane A ae 11. Glumes spirally arranged feu > aif «yew b-nmtgieeme unec Entube 0 A. i See 37. Female flower enclosed by a sac-like organ (a modified prophyll). Nut otherwise, without disk. 38. Modified prophyll (perigynium) more or less open on the abaxial side . 27. Kobresia (kobresiodea) 38. Perigynium closed throughout its length (a utricle). 456 FLORA MALESIANA [ser. 1, volar 39. Besides the ovary (or nut) the utricle contains a needle-like rachilla produced beyond the orifice of the utricle and strongly hooked at the top. Inflorescence always a single terminal spikelet. 29. Uncinia 39. Usually no rachilla inside the utricle; when there is a rachilla, it is not hooked at the top. Inflorescence paniculate, spicate, or a single spikelet guns 28. Carex 1. SCIRPODENDRON Zipp. ex Kurz, J. As. Soc. Beng. 38, 11 (1869) 84. — Ptychocarya R. BR. ex WALL. Cat. (1831) n. 3538, nom. nud. in synon.; LINDL. Veg. Kingd. (1846) 119, nom. nud.; VON Post & O.K. Lexic. Gen. Phan. (1904) 470, nom. nud. (Ptycho- caryum); H. PFEIFF. in Fedde, Rep. 21 (1925) 240. — Fig. 3. Stout perennial herbs, with woody, obliquely erect rhizome, in habit strongly resembling a small Pandanus. Stems relatively short, erect, triquetrous, leafy at the base only. Leaves 3-ranked, with short, open sheaths and very long linear blades. Inflorescence terminal, paniculate, branched in the lower half, the lowest branches ternate, sometimes branched again. Lowest 3 bracts pseudo- whorled, very long, leaf-like, the higher ones much smaller, the uppermost scale-like. Spikelets terete or more or less trigonous, many-flowered. Glumes subcoriaceous, spirally imbricate. Flowers hermaphrodite; terminal flower of each spikelet with a terminal ovary and several flat fascicled (probably spiral) scales each bearing a single stamen in its axil; lateral flowers strongly dorsiven- trally compressed, also with a terminal ovary, but with the 2 outer scales transversal, opposite, boat-shaped, sharply keeled, ciliate on the keel, usually connate on the adaxial side, the other scales variable in number (up to c. 10 in the lower flowers, often very reduced in the upper ones), flat, linear-lanceolate, acute, each with a single stamen in its axil (or uppermost scales sterile); arrange- ment in the lower fertile glumes of the spikelets usually more complex: two outer scales winged on the back, enclosing a normal central flower and two lateral more or less reduced ones. Style continuous with the ovary, not or hardly thickened at the base; stigmas 2 or 3 (in the same spikelet). Fruit drupaceous, coarsely ribbed; exocarp succulent, corky when dry; endocarp bony, black. Distr. Monotypic genus, extending from Ceylon to Australia and Polynesia. 1. Scirpodendron ghaeri (GAERTN.) Merr. Philip. J. Sc. 9 (1914) Bot. 268; Int. Rumph. (1917) 106; Brown, Min. Prod. Philip. For. | (1920) 352; En. Born. (1921) 64; En. Philip. 1 (1923) 131; HEyYNE, Nutt, Pl (1927), 3035S: i. BLAKE, Proc: Ro Soc: Queensl. 54 (1943) 73; RayM. Nat. Canad. 91 (1964) 131; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 454. — Chionanthus ghaeri GAERTN. Fruct. 1 (1788) 190, t. 39, f. 6a—e; Boeri. J. Linn. Soc. Bot. 31 (1895) 246. — Pandanus pumilus Moon, Cat. (1824) 67. — Pandanus caricosus [RUMPH. Herb. Amb. 4 (1743) 154] SpRENG. Syst. 3 (1826) 897; Mia. FI. Ind. Bat. 3 (1855) 163, non Kurz, 1867. — Scleria macrocarpa WALL. Cat. (1831) n. 3538, nom. nud. — Ptychocarya macrocarpa STEUD. Nomencl. 3 (1841) 416, nom. nud. — Hypolytrum costatum THw. En. Pl. Zeyl. (1864) 346. — S. costatum Kurz, J. As. Soc. Beng. 38, 11 (1869) 85; ibid. 39, 11 (1870) 85; SCHEFF. Nat. Tijd. N. I. 34 (1874) 91; Bentu. FI. Austr. 7 (1878) 341; GorBeL, Ann. Jard. Bot. Btzg 7 (1888) 122, t. 14, f. 1-11; CLARKE, FI. Br. Ind. 6 (1894) 684; Hook. f. in Trimen, Handb. Fl. Ceylon 5 (1900) 92, t. 97; RipL. J. Str. Br. R. As. Soc. n. 46 (1906) 227; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 106; MerR. Philip. J. Sc. 2 (1907) Bot. 422; CLARKE, Ill. Cyp. (1909) t. 116, f. 7-12; RipL. & WINKL. Bot. Jahrb. 44 (1910) 202; C. B. Ros. Philip. J. Sc. 6 (1911) Bot. 195; Koorpb. Exk. Fl. Java 1 (1911) 202; Atlas (1922) f. 266; RipL. Fl. Mal. Pen. 5 (1925) 175. — S. pandaniforme Zipp. ex Kurz, J. As. Soc. Beng. 38, 11 (1869) 85, in syn. — Pandanophyllum costatum Kurz l.c. in syn. — S. sulcatum Mia. Illustr. (1870) 65, t. 28, err. calam. — Pandanus acaulis MARTELLI, Webbia 4 (1913) 5.— Mapania macrocephala (non K. ScHuM.) Mere. En. Philip. 1 (1923) 132 p.p. (quoad ELMER 11978). — Ptychocaryum ghaeri H. PreirF. in Fedde, Rep. 21 (1925) 240; Bot. Arch. 12 (1925) 446; in Fedde, Rep. 28 (1930) 20; UITTIEN in Back. Bekn. Fl. Java. (em. ed.) 10 (1949) fam. 246, p. 53. — Fig. 3. CYPERACEAE (Kern) Fig. 3. Scirpodendron ghaeri (GAERTN.) MERR. a. Inflorescence, b. part of leaf and its section, x 24, c. spikelet, showing the glumes, d. flower, sustained by the glume, x 2, e. flower, with the two transversal scales, x 3, she pistil, g. partial infructescence, A. nut, i. ditto in CS, nat. size (after MIQUEL). 458 FLORA MALESIANA [ser. 1, vo Rhizome thick, forming large dense clumps, clothed with acute, imbricating scales and their fibrous remains. Stems smooth, 30-60cm by 5- 10 mm. Leaves numerous, coriaceous, drooping in the upper part, plicate, with 3 prominent nerves, aculeate-scabrous on the margins and the midrib beneath in the upper pat, rather gradually narrowed into a filiform, triquetrous, scabrous, 15-25 cm long tail, 1-4 m by 2—S cm; lower sheaths brown to dark castaneous. Panicle dense, oblong-ovoid, (5—)10—20 by up to 7cm when in fruit; branches short, thick, obliquely spreading. Lower bracts (30—)60—150 cm long, the upper ones broadly ovate with filiform scabrous tip. Spikelets in sessile or almost sessile clusters, ovoid. Glumes ovate, obtuse, many-nerved, stramineous or greenish, c. 1 cm long. Flowers slightly shorter than the glumes. Anthers linear, 3—4 mm long. Fruit conical-ellipsoid, acute, usually with c. 6 more or less tuberculate longitu- dinal ribs, dusky brown, 1-114 by c. 1 cm; ribs sometimes up to 10 and then often confluent at the top. Distr. Ceylon, Peninsular Thailand, North Queensland (Cook Distr., very rare), New Hebrides, Polynesia (Fiji, Samoa); in Malesia: Sumatra (and the adjacent islands Simalur, Mentawei, Banka), Malay Peninsula (also in P. Penang), West and Central Java (rare), Borneo (Sarawak, N. & E. Borneo), Philippines (Palawan, Luzon, Polillo, Leyte, Biliran, Mindanao), SE. Celebes (Kandari), Moluccas (Halmaheira, Ternate, Sulu Is., Buru, Ceram, Am- boina), New Guinea (Sorong). Ecol. Freshwater tidal areas on clayish soil, tidal swamp forests, transition forests behind the mangrove, along rivermouths, especially in places where during high tide or heavy rainfall the water is waist deep. Because of the cutting, pandan-like leaves often forming almost impenetrable pure stands, which on aerial photographs can easily be recognised amidst the small-crown swamp forests. According to RIDLEY, Disp. (1930) 240, 329, the large fruits readily float away when the water rises, and they are largely carried off by rats, which eat the corky exterior. Uses. In Sumatra, Leyte, and the Moluccas (also in Ceylon) the dried leaves are used for making mats and hats, in Fiji for thatching. In S. Sumatra (Djambi, Palembang) the species is sometimes culti- vated for this purpose. The material is apparently of inferior quality. In Samoa the fruits are eaten by the natives; this use is not reported from Malesia. Vern. Harashas, S, piés, rumbai latah, Lamp., garingging, Simalur, rumbai, séding ayér, Banka, sélingsing, Mal. Pen., pandan ayér, Mol., lasial, lasiatal, Alf. Amb., séwés, S. Halm. (Weda), sas- aréweén, sosaréwu, N. Halm. (Gal.), hehéwéhe, Tob., séa-séa, Ternate; Philip.: bilis, Sub., gaas, Bik., barongis, Bag. 2. CAPITULARINA KERN, nom. nov. — Capitularia VALCK. SUR. Nova Guinea 8 (1912) 711, non FLORKE, 1807 (quae est Cladonia), non RABENH. 1851 (quae est Uromyces). — Fig. 4. Perennial herbs with a woody, shortly creeping rhizome covered with ovate, many-nerved, fuscous to castaneous sheaths. Stems erect, quinquangular, hollow, inside with some indistinct septa not or hardly visible from the outside. Leaves basal, few, subcoriaceous, linear, folded lengthwise (at least when dry), complicate at the base, with 3 prominent nerves, or reduced to bladeless sheaths. Inflorescence terminal, capitate, consisting of 1-—several spikelets. Involucral bracts several, long, leafy, pseudo-whorled. Spikelets terete, many- flowered. Rachilla persistent. G/umes numerous, spirally imbricate, cartila- gineous, caducous, some lower ones empty. Flowers hermaphrodite, strongly dorsiventrally compressed; floral scales numerous, membranous, the lowest 2 transversal, opposite, free cucullate, with a narrow, minutely serrulate-ciliate wing on the acute keel, empty, the next 2 also empty, concave, posticous and anticous, the posticous one embracing the other; upper scales c. 8—15, flat or slightly concave, acute, each with a single stamen in its axil. Anthers linear, shortly apiculate. Style continuous with the ovary; stigmas 2, short. Nut conical-ovoid, many-ribbed, seated on a large, somewhat turgid, hollow stipe, therefore seemingly consisting of a fertile apical part and a sterile basal part; exocarp thin, not fleshy. Distr. Malesia: New Guinea and Solomon Is., probably monotypic. Note. Ripiey (Trans. Linn. Soc. II, Bot. 9, 1916, 244) united Capitularia with the Australian genus Chori- zandra R.Br. However, UITTIEN (Rec. Trav. Bot. Néerl. 33, 1936, 291) pointed out that Capitularia is well characterized by the quinquangular stems, not found in the related genera and very unusual in Cyperaceae, 1974] CYPERACEAE (Kern) 459 Fig. 4. Capitularina involucrata (VALCK. SUR.) KERN. a. Habit, x )4, a’. stem node in LS, x 3, b. inflorescence, x A, ¢. glume, d. transversal ciliate scale, e. inner scale, all x 214, f. flower, g. ditto, transversal scales removed, both x5, h. stamen with scale, i. nut on stipe, j. ditto, in LS showing hollow stipe, all x 2)4, k. pistil, x 5, /. floral diagram (a—c, f—g, k GJELLERUP 754, d, i—j BRASS 7064, e, h Brass 7003). 460 FLORA MALESIANA [ser. I, vol. 7° the terminal, capitate inflorescence subtended by several long leafy bracts, the four outer empty scales of the flower, the fertile inner ones, and the short stigmas. In Chorizandra the stems are terete, the inflorescence is pseudolateral because of the terete erect involucral bract as though continuing the stem, the outer scales are fertile, the inner ones empty, and the stigmas long. The peculiar nuts of Capitu/arina seated on a large sterile part are entirely different from those of Chorizandra. 1. Capitularina involucrata (VALCK. SUR.) KERN, comb. nov. — Capitularia involucrata VALCK. SUR. Nova Guinea 8 (1912) 711, t. 118. — Chorizandra involucrata RID. Trans. Linn. Soc. II, Bot. 9 (1916) 244. — Capitularia foliata var. archboldii UITTIEN, J. Arn. Arb. 20 (1939) 214. — Fig. 4. Stems borne single on the rhizome, acutely 5- angled (according to RIDLEY also 4-angled), smooth or slightly scaberulous just below the inflorescence, 35-65 cm by 2-4 mm, the base covered with blade- less, coriaceous, castaneous sheaths; upper sheaths scarious, lanceolate, 10-20 cm long. Leaf-blades 0-—S, gradually narrowed into a very acute to almost filiform scabrous point, scaberulous on the margins and the nerves on the under side at least in the upper part, glaucescent, 60-125 by 1-2 cm. Involucral bracts 5—8, similar to the leaves, erect, rigid, plicate, castaneous at the base, unequal, the lowest 20—75 by 1-2 cm. Spikelets half hidden in the bases of the involucral bracts, obovoid or obconical, 1—2 cm long and wide. Outer glumes obovate-cuneate, abruptly cuspidate, minutely serrulate-ciliate on the upper margin, pale at the base, castaneous at the apex, with a scabrous, up to 5 mm long awn, 7-11 by 5-7 mm; upper glumes gradually becoming lanceolate, pale, tipped with reddish brown. Flowers 5-11 mm long; outermost scales subspathulate, the wing on the keel up to 1 mm wide; posticous and anticous scales spathulate, obtuse, ciliolate at the top; fertile scales narrowly lanceolate, acute. Anthers linear, up to 6mm long. Style purplish, 5-10 mm long. Nut terete or slightly angular, very acute and acuminate or subacuminate, longitudinally striate- sulcate, stramineous with fuscescent base, 5—6 by 3—4 mm, the stipe (sterile basal part) fuscous, stri- olate, 3—4 by 2—3 mm. Distr. Malesia: New Guinea (W. New Guinea: Steenkool (Tembuni), Mt Carstensz (Canu Camp), Noord River near Alkmaar, Tor River, Utakwa River; Papua: Palmer River, W. Distr. in Kuinga Subdistr.). Ecol. In swampy parts of primary forests, at low altitude (up to 200 m?). Notes. Capitularia foliata U1TTIEN (Rec. Trav. Bot. Néerl. 33, 1936, 289) was based on Brass 3045 from the Solomon Islands, which collection I have not seen. According to UrTTIEN, C. foliata is quite similar to C. involucrata in stem, leaves, and spikelets, but more robust in all its parts, the stem provided with very long leaves at the base, the inflorescence consisting of several spikelets, the bracts much larger and awned. In 1939 UiTTIEN described C. foliata var. archboldii from Papua, differing from the typical form by the single spikelet and the shorter-awned glumes; C. foliata was then said to differ from C. involucrata by the leafy stem-bases. In my opinion it is impossible to distinguish between C. involucrata and C. foliata solely by the absence or presence of basal foliage-leaves, and I doubt whether C. foliata can be maintained as a species. In GJELLERUP 754 (L) from Tor River, the base of one of the two speci- mens bears scarious bladeless leaf-sheaths only, but in the other a well developed leaf is present. Both specimens certainly belong to C. involucrata; they were determined as such also by UITTIEN. In my opinion the few collections known from New Guinea belong to a single species which is very variable in the size of all its parts, even of its floral parts. A similar variability is found in several other members of the Mapanieae. 3. LEPIRONIA L. C. Ricu. in Pers. Syn 1 (1805) 70.— Chondrachne R. BR. Prod. (1810) 220. — Choricarpha Borck. Flora 41 (1858) 19. — Fig. 5—6. Perennial herbs with rush-like habit. Rhizome woody, horizontally creeping. Stems erect, terete, transversely septate within. Leaves reduced to bladeless sheaths open in front, the margins overlapping. Inflorescence consisting of a single spikelet, apparently lateral owing to the single erect involucral bract continuing the stem. Spikelet terete, many-flowered. Rachilla thick, spongious, conical, persistent. G/umes chartaceous, spirally arranged, very densely imbricate, caducous with the nut, some lower ones empty. Flowers strongly dorsiventrally compressed. Hypogynous scales numerous, (up to c. 152), the lowest 2 trans- versal, opposite, free, boat-shaped, ciliolate on the keel, the remaining ones fascicled (probably spiral), linear-lanceolate, acute, flat or nearly so. Stamens several (up to c. 102), one in the axil of each of the keeled scales, the others solitary in the axils of the outer flat scales; anthers linear, shortly apiculate. Ovary terminal; style continuous with the ovary, slightly incrassate at the base, the base persistent on the nut as a short beak; stigmas 2, long. Nut strongly dorsiventrally compressed, plano-convex, acutely keeled on the margins. 1974] CYPERACEAE (Kern) 461 ig: Lepironia articulata (RETZ.) DOMIN. a. Habit, x 14, b. spikelet, x 2, c. glume, d. flower with two trans- versal ciliate scales, enclosing inner scales and pistil, e. stamen, f. transversal scale, g. one inner scale, A. pistil, i. nut, j. ditto in CS; all x 5 (a—j KALKMAN 4021). 462 FLORA MALESIANA [ser. I, vol. 7° Distr. Usually considered a monotypic genus extending from Madagascar to Australia and Oceania. However, it is very likely that ENDLICHER (Gen. Pl. 1836, p. 116) was right in uniting Lepironia with Chori- zandra R.BR. The latter genus comprises 4 Australian spp. in habit very similar to Lepironia and with essentially the same flower-structure. The nut in Chorizandra possesses c. 8 very prominent longitudinal ribs. The species referred to Lepironia by MiQueL belong to Mapania and Thoracostachyum. 1. Lepironia articulata (RETZ.) Domin, Bibl. Bot. Heft 85 (1915) 486; Premrr. Bot. Arch. 12 (1925) 451, f. 25 & 46; FiscHER, Kew Bull. (1932) 70; Burk. Dict. 2 (1935) 1331; S. T. BLAkg, J. Proc. R. Soc. Queensl. 54 (1943) 71; J. Arn. Arb. 28 (1947) 213. — Restio articulatus Retz. Obs. 4 (1786) 14. — Scirpus coniferus Por. in Lamk, Enc. 6 (1804) 756; Suppl. 5 (1817) 90.— L. mucronata L. C. RIcu. in Pers. Syn. 1 (1805) 70; KuntH, En. 2 (1837) 366; STEUD. Fig. 6. Stand of Lepironia articulata (RETZ.) DOMIN on sandy soil with peaty water in the Tasek Bera swamp valley in Malaya; ground cover with Sphagnum (photogr. SOEPADMO). Syn. 2 (1855) 181: Mig. Fl. Ind. Bat. 3 (1856) 346: Sum. (1861) 262, 602: Kurz, Nat. Tijd. N. I. 27 (1864) 224. J. As. Soc. Beng. 38, ii (1869) 77: Flora 52 (1869) 438; Mia. Illustr. (1870) 60, t. 20: Borck. Linnaea 37 (1871) 140; ScHEFF. Nat. Tijd. N. I. 34 (1874) 89; BenTu. FI. Austr. 7 (1878) 342: GOEBEL, Ann. Jard. Bot. Btzg 7 (1888) 126, t. 14, f. 1213: CLaRKE, FI. Br. Ind. 6 (1894) 684; Rip. J. Str. Br. R. As. Soc. n. 46 (1906) 227; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 107; CrarkeE, Ill. Cyp. (1909) t. 118, f. 8-16; Koorp. Exk. Fl. Java 1 (1911) 202: Atl. (1922) f. 267 err. ‘mucronulata’: CAMUS, FI. Gén. I.-C. 7 (1912) 179, f. 25; MeRR. En. Born. (1921) 64; KUK. Bot. Jahrb. 50 (1924) 58; RipL. Fl. Mal. Pen. 5 (1925) 175; HEYNE, Nutt. Pl. (1927) 313: CHERMEZ. Fl. Madag., fam. 29 (1937) 244, t. 23, f. 5-8. — Chon- drachne articulata R. BR. Prod. (1810) 220. — Chori- carpha aphylla Boeck. Flora 41 (1858) 20. — L. conifera Druce, Rep. Bot. Exch. Club Br. Isl. 1916 (1917) 631. — Fig. 5-6. Madagascar, Ceylon, Thailand, Indo- China, S. China, coastal districts of N. and E. Australia, Carolines, New Caledonia, Fiji, in Male- sia: Sumatra (Tapanuli, Palembang, Lampongs), Banka, Lingga Arch., Malay Peninsula (Dindings, Pahang, Malacca, Johore, Singapore), Borneo, Cen- tral Celebes (Towuti Lake), Moluccas (Sula Is.: Taliabu; Buru), New Guinea. Ecol. In open swampy places, open marshes, swamps in savannah-forests, along quiet streams, often near the coast, in Sumatra up to 1000 m, in Trengganu (G. Padang) at 1200 m, in New Guinea (Wissel Lake Region) up to 1750 m. Often forming extensive communities. Vern. Tékor, tikér, tikuk, Lamp., purun, putjet, Banka, Mal. Pen., kérédjut, Lingga Arch., purum danau, Mal. Pen., Borneo. Uses. In S. Sumatra and Borneo mats for packing tobacco, rubber, and kapok, are made of the dried stems. The species is sometimes cultivated for this purpose. In New Guinea (Sepik Distr.) it is used by the natives for basket-making. 1974] CYPERACEAE (Kern) 463 Fig. 7. Thoracostachyum sumatranum (MiQ.) Kurz. a. Habit with inflorescence, x 2, b. floral diagram, c. spikelet, x 44, d. glume, e. flower with 2 outer transverse ciliate scales, both x 3, /. pistil, g. submature nut with style remains (after MIQUEL). 464 FLORA MALESIANA [ser. I, vol. 7° 4. THORACOSTACHYUM Kurz, J. As. Soc. Beng. 38, 11 (1869) 75; UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 133. — Mapania sect. Thoracostachyum (KURZ) KOYAMA, Mem. N.Y. Bot. Gard. 17 (1967) 50, in footnote. — Fig. 7-8. Perennial herbs, (in the Mal. spp.) with long-creeping stolons hardening into woody, creeping rhizomes covered with lanceolate acute scales. Stems arising from the centre of a basal tuft of normal leaves, erect, triquetrous. Leaves 3-ranked, equitant, subcoriaceous, linear, flat or somewhat folded lengthwise, very gradually narrowed into a very acute to filiform scabrous point, condupli- cate at the base (in 2 Indochinese spp. narrowed into a petiole); midrib (and often 2 of the lateral nerves) prominent underneath. Inflorescence terminal, paniculate, with divaricate rigid branches and several to many spikelets; lower bracts long, leaf-like, the upper ones much smaller. Spikelets broadly ovoid to oblong-ovoid, terete, solitary or more or less aggregated in twos or threes at the top of the ultimate branchlets, few- to many-flowered. Rachilla persistent. Glumes spirally imbricate, subcoriaceous, indistinctly 1—3-nerved, some lower ones empty. Flowers hermaphrodite, strongly dorsiventrally compressed, their structure as in Mapania: hypogynous scales 6, the lowest 2 transversal, opposite, free, boat-shaped, ciliate on the acute keel, each with a stamen in its axil, the upper 4 concave to almost flat, the third (anticous) one with a stamen in its axil, the remaining 3 empty (see note). Style continuous with the ovary; stigmas 3 (rarely in some flowers 2). Nut angular, hard, shining; endocarp stony, exocarp not spongy or fleshy; beak confluent with the nut proper. Distr. Small genus of about 7 closely related spp.; 1 in the Seychelles, 2 in Indo-China, 1 in the Carolines, 1 in the Fiji Islands, in Malesia 2 spp. I am not certain that they all deserve specific rank. Ecol. Usually in swampy forests, but also in open wet places, at low altitudes. Uses. T. sumatranum is sometimes used for making mats. Notes. The structure of the flowers is not as constant as it is usually assumed. Sometimes the lowest 4 (instead of 3) of the hypogynous scales bear stamens in their axils. UITTIEN (Bull. Bish. Mus. n. 141, 1936, p. 16) even observed 6 stamens in some of the flowers of T: vitiense UITTIEN all scales then being fertile. T. pacificum HosoKAwa is described as having 4 or 5 stamens. Thoracostachyum combines the habit of Hypolytrum with the flower-structure of Mapania. From Hypoly- trum the genus is sufficiently distinguished by the different number of hypogynous scales and the 3 stigmas. It is very closely related to Mapania, the only differences with Mapania sect. Cephaloscirpus lying in the paniculate (not capitate) inflorescence, and the hard, non-drupaceous fruit. KEY TO THE SPECIES 1. Stems smooth even at the top. Glumes ovate, 3-334 mm long. Flowers 3—3)4 mm long. Nut broadly obovoid, abruptly acuminate at the apex ._. . 1. T. sumatranum 1. Stems more or less scabrous at the top. Glumes broadly ovate, ee A mm long. Flowers 134-2 mm long. Nut ellipsoid, slightly acuminate, with 3 longitudinal grooves in the conical beak . 2. T. bancanum 1. Thoracostachyum sumatranum (MiQ.) Kurz, J. As. Soc. Beng. 38, ii (1869) 75; ScHEFr. Nat. Tijd. N. I. 34 (1874) 89; Urttien, Rec. Trav. Bot. Néerl. 33 (1936) 136, f. 1b & 2b; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 455. — Lepironia sumatrana Mia. Sum. (1861) 263, 604; Illustr. (1870) 62, t. 24. — Hypolytrum pandanophyllum F.v.M. Fragm. 9 (1875) 16. — Pandanophyllum hypolytroides F.v.M. L.c., in nota. — Mapania sumatrana BENTH. FI. Austr. 7 (1878) 341.— Mapania hypolytroides F.v.M. ex BENTH. FI. Austr. 7 (1878) 341. — Mapania panda- nophylla K. Scu. in K. Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 25; K. Scu. & Laur. Fl. Schutzgeb. (1900) 189; KoyAMa, Micronesica 1 (1964) 64. — T. hypolytroides CLARKE, FI. Br. Ind. 6 (1894) 680; J. Linn. Soc. Bot. 34 (1898) 94; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 102; CLARKE, Ill. Cyp. (1909) t. 108, f. 7-10; VALcK. Sur. Nova Guinea 8 (1912) 710; KOK. Bot. Jahrb. 59 (1924) 54; RipL. Fl. Mal. Pen. 5 (1925) 171. — T. dichromenoides RiDv. Bot. Jahrb. 44 (1910) 525; Merr. En. Born. (1921) 66. — T. pandanophyllum Domin, Bibl. Bot. Heft 85 (1915) 484; UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 138, f. lc & 2c; S. T. BLAKE, J. Arn. Arb. 28 (1947) 208; CYPERACEAE (Kern) 465 Fig. 8. Thoracostachyum sumatranum (MiQ.) KURZ. New Guinea, Cape Vogel Peninsula (Photogr. HOOGLAND; HOOGLAND 4646). UrTTIEN in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 51. — Mapania heyneana Back. Bull. Jard. Bot. Btzg III, 2 (1920) 328. — T. heyneanum UiTTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 200; ibid. 33 (1936) 139, f. 1d. — Fig. 7-8. Stems usually stout, smooth even at the top, (25—)60—150 cm by (2—)5—10 mm. Leaves green or pale green, distinctly reticulate when dry, up to 2 m long, (1—)2—3/4 cm wide; margins scabrous at least in the upper half; lowest sheaths bladeless, fuscous to blackish. Inflorescence globose or ovoid, often dense, (3—)10—20 cm in diam., with 10 to some hundred (according to CLARKE up to 800!) spikelets; lowest bract up to more than | m long, pale at the base; branches smooth or scabrous. Spikelets ovoid or oblong-ovoid, often nearly globular when in fruit, straw-coloured, many-flowered, 5-10 by 3- 5 mm, up to 7 mm wide when in fruit. Glumes ovate, obtuse, pale, with scarious margins, 3-334 by 2-234 mm. Flowers 3-314 mm long. Anthers linear, V4-1% mm long. Nut broadly obovoid, obtusely 3—S-angled, narrowed towards the base, abruptly acuminate at the apex, brown to castaneous, c. 3 by 2 mm (the c. 1 mm long beak included). Distr. Australia (NE. Queensland), Palau Islands (?), in Malesia: S. Sumatra (Palembang, Lampongs), Malay Peninsula (Malacca, Johore), W. Java (Danu swamp), Borneo, SE. Celebes, and New Guinea. Ecol. In swamps, marshy forests, ponds, on river- banks, on floating islands, locally often abundant; at low altitudes. Uses. In Sumatra (Palembang) and the Malay Peninsula the leaves are used for making mats. In Sumatra it is sometimes cultivated for this purpose in swamps and inundated fields. The mats are less durable than those made of the leaves of Pandanus Spp. Vern. M: umbai, rumput pandan biru, Mal. Pen., rumbai lilin, r. idju, Palemb., selingsing, Lamp., tigesangi, New Guinea (Trapi lang.). Notes. The inflorescence is very variable in size and density. UITTIEN treated T. sumatranum and T. pandanophyllum as specifically distinct on account of the large decompound inflorescence of the latter, and T. heyneanum on account of the less dense inflo- rescence and the longer spikelets. It is evident that it is impossible to trace dividing lines only with the aid of these characters. The details of flowers and fruits are the same in all three. In the herbaria T. sumatranum is often confused with Hypolytrum nemorum. It may be distinguished by the creeping stolons covered with long stramineous scales, the much firmer glumes, the 3 stigmas, and the nut abruptly narrowed into the beak. 2. Thoracostachyum bancanum (MiQ.) Kurz, J. As. Soc. Beng. 38, ii (1869) 76; ScHEFF. Nat. Tijd. N. I. 34 (1874) 89; CLARKE, FI. Br. Ind. 6 (1894) 680, incl. var. longispica; J. Linn. Soc. Bot. 34 (1898) 94; RIDL. J. Str. Br. R. As. Soc. n. 46 (1906) 226; Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 102; CLARKE, Ill. Cyp. (1909) t. 107, f. 12-20; Camus, Fl. Gén. I.-C. 7 (1912) 174; Mere. En. Born. (1921) 65; RipL. Fl. Mal. Pen. 5 (1925) 171; UittiEn, Rec. Trav. Bot. Néerl. 33 (1936) 136, f. la, 1f, 2a; OHwi1, Bot. Mag. Tokyo 56 (1942) 210; S. T. BLake, J. Arn. Arb. 28 (1947) 208. — Lepironia bancana MiQ. Sum. (1861) 263, 604; Illustr. (1870) 63. — Thoracostachys bancana Kurz, Nat. Tijd. N. I. 27 (1864) 224; Bot. Zeit. 23 (1865) 204, nom. nud.— Hypolytrum borneense Kurz, J. As. Soc. Beng. 38, ii (1869) 74; Mia. Illustr. (1870) 59; ScHeFF. Nat. Tijd. N. I. 34 (1874) 88. — Mapania bancana B. & H. ex Jacks. Ind. Kew. 2 (1895) 163. — T. ridleyi CLARKE in Ridl. J. Str. Br. R. As. Soc. n. 46 (1906) 226; Kew Bull. add. ser. 8 (1908) 53; Merr. En. Born. (1925) 66.— T. sub- capitatum VALCK. SuR. Nova Guinea 8 (1912) 710, (ig Wg Stems slender, more or less scabrous at the top, 15—80 cm by 1—3 mm. Leaves longer than the stems, usually glaucous, indistinctly reticulate when dry, scabrous on themargins,4—20 mm wide. Inflorescence loose, with scabrous branches, 2—7 cm in diam., with 5—60 spikelets; lower bracts usually not pale at the base, 15-40 cm long. Spikelets globular or ellipsoid, usually reddish at the top, few-flowered, 4—6 by 2—3 mm (c. 5 mm wide when in fruit). Glumes broadly ovate, obtuse, 134-2)4 by 14-2 mm. Flowers 124-2 mm long. Anthers oblong to linear- oblong, >4-1 mm long. Nui ellipsoid, slightly acuminate, with 3 longitudinal grooves in the conical beak, blackish brown at the base, olive- green at the top, 214-3 by 114-1'4(-2) mm (beak included). 466 Distr. Tonkin; in Malesia: Sumatra (also Men- tawei Is., Banka, and Riouw Arch.), Malay Peninsula (Perak, Pahang, Malacca, Johore, Singapore), Borneo, Moluccas (Buru), and New Guinea. Ecol. In swampy places in forests, in swamp forests and peat forests, often plentiful as a ground cover, also in wet open places, at low altitudes (in Buru at 1075 m). Vern. Mingsing, rumbai bajan, M, sending ajer, rumput séding, Banka, rumput senayan batu, r. FLORA MALESIANA [ser. I, vol. 7° séndérayan, séndayan, sératit, Mal. Pen., bélingi, rumput grinsing, r. rembog, W. Borneo. Notes. T. subcapitatum VALCK. SuR. is a form with elongated spikelets apparently due to the attacks of an Ustilago. This abnormality was also described as T. bancanum var. longispica CLARKE. It occurs with the normal form. A specimen from “E. Java, Tosari”’, leg. RIDLEY in the Kew Herbarium was obviously mislabelled (cf. Fl. Mal. I, 1, 1950, p. xxixa). 5. MAPANIA AUBLET, Hist. Pl. Gui. Frang. 1 (1775) 47; Bogck. Linnaea 37 (1871) 136; CLARKE, FI. Br. Ind. 6 (1894) 680; Kew Bull. add. ser. 8 (1908) 130; UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 145; l.c. 277. — Pandanophyllum HAssk. Tijd. Nat. Gesch. Phys. 10 (1843) 118, p.p. typ.; STEUD. Syn. 2 (1855) 134, p.p. — Fig. 9-16. Perennial, usually coarse herbs. Rhizome woody, erect, obliquely erect, or horizontal (sometimes up to 2 m long); long-creeping stolons covered with lanceolate, acute scales rarely present; roots usually thick, greyish brown to blackish, sometimes functioning as stilt roots. Flowering stems either central (arising from the centre of a basal tuft of leaves), erect, triquetrous, usually with 1—2 cauline leaves, or lateral (scapes), arising from the axils of the lower (rarely also higher) leaves or from below the leaves, trigonous or terete, the base clothed with some bladeless scale-like sheaths. Leaves 3-ranked, subcoriaceous to thickly coriaceous, linear, broadly linear, or lorate, slightly narrowed towards the conduplicate base, or rather abruptly narrowed into a petiole, the top (in almost all Mal. spp.) very gradually to abruptly narrowed into a filifc =m, triquetrous, scabrous tail; margins and midrib (rarely also 2 lateral nerves) aculeate-scabrous; midrib prominent beneath, 2 (rarely more) lateral nerves often prominent above. Inflorescence either consisting of a single spikelet or capitate (i.e. consisting of few to numerous sessile spikelets), in the latter case surrounded by some glume-like or foliaceous involucral bracts. Glumes spirally imbricate, coriaceous, chartaceous, or membranous, lower ones empty, upper ones flower-bearing. Flowers hermaphrodite, linear, strongly dorsiventrally compressed. Hypogynous scales (in the Mal. spp.) 6, thinly membranous, the lowest 2 transversal, opposite, free, rarely connate on the adaxial side, boat- shaped, ciliate on the acute or narrowly winged keel, each with a stamen in its axil, the third (anticous one) concave or 2-keeled, also with a stamen in its axil, the remaining 3 concave or flat, empty; in extra-Malesian spp. the number of scales and stamens less constant. Anthers linear; connective only shortly produced. Sty/e continuous with the ovary, not or hardly incrassate at the base; stigmas 3, more rarely 2. Nut terete or slightly angular, beaked, stipitate; exocarp thick, corky or fleshy; endocarp stony, black. Distr. About SO spp. in the tropical regions of South America and Africa (not in Madagascar!), and from Ceylon, Assam, Sylhet, Thailand and Indo-China to the W. Pacific and N. Queensland, in Malesia 25 spp. The centre of development in Malesia is Borneo, from where 19 spp. are known (10 endemic). In the Malay Peninsula 11 spp. (2 endemic), in Sumatra 7 spp., in W. Java only 3 spp. Fig. 10. The genus is not represented in E. Java and the Lesser Sunda Is. which have largely a seasonal climate, and but poorly in Celebes and the Moluccas. Mapania macrocephala, M. moseleyi, and M. baccifera are re- stricted to the eastern part of the Archipelago, M. macrocephala extending to N. Queensland, Samoa, the Solomons, and the Carolines. Only M. palustris and M. cuspidata are widely distributed. Ecol. All Malesian species occur in everwet primary rain-forests, preferably on moist, deep, muddy, 1974] CyPERACEAE (Kern) 467 Fig. 9. Mapania latifolia Uitt1EN. a—a’. Habit, x 4, db. floral diagram, c. spikelet, x 3, d. glume, e. flower, with two transverse scales, inner scales and stigmas, f. pistil, g—h. nut with style, i. nut in CS, all x 5 (a, g—i CLEMENS 27573, a’—f CLEMENS 30065). 468 FLORA MALESIANA [ser. I, volew® Fig. 10. Range of the genus Mapania AuBLET; above the hyphen the total number of species for each island or district, below it the number of endemic species. humous or peaty soils, often on stream-banks or in shallow pools at low and medium altitudes; in New Guinea M. macrocephala ascends to 2000 m. Taxon. Subdivision of the genus. The type species of Mapania is the South American M. sylvatica AUBL. Hist. Pl. Gui. Frang. 1 (1775) 47, t. 17. In sect. Mapania the leaves are as a rule all reduced to bladeless sheaths, and the inflorescence is sustained by an involucre of 3 large, oblong bracts. This section is not represented in Malesia. The Malesian species are generally classified into 3 sections, but I greatly doubt whether these sections can be upheld, the distinction between them not being sharp, especially that between sect. Pandanophyllum and sect. Macrolepironia. Some botanists have gone so far as to raise some sections to subgeneric or even generic rank, but in my opinion there is too much reticulate affinity to warrant that procedure. There are certainly groups of allied species in Malesia, but they cannot be rigidly classified into infrageneric taxa. Therefore I have decided to refrain from recognizing sections and I have arranged the species according to their supposed affinity. Moreover, there is a regrettable confusion as to the nomenclature. It has been overlooked that as early as 1870 MiQueEL used Pandanophyllum as a sectional name (in Lepironia) to cover the species with a capitate inflorescence. The type of this section is Pandanophyllum palustre Hassk. ex StEUD. Unfortunately CLARKE applied the name Pandanophyllum to the section with normally a single spikelet to the inflorescence, for which section MIQUEL coined the name Macrolepironia. This name must be reinstated. To those who want to recognize sections, the names to be used are as follows: Sect. 1. Cephaloscirpus (KURZ) B. & H. Gen. PI. 3 (1883) 1056. — Cephaloscirpus Kurz, J. As. Soc. Beng. 38, ii (1869) 83 (T: Hypaelyptum macrocephalum Gaupicu.).— Mapania subg. Cephaloscirpus CLARKE, Kew Bull. add. ser. 8 (1908) 131; Urrtten, Rec. Trav. Bot. Néerl. 32 (1935) 185. Flowering stems central. Involucral bracts foliaceous. Inflorescence capitate. Embraces the E. Malesian M. macrocephala and M. moseleyi. UrtTIEN referred also M. latifolia to this section, but this species shows strong relations to M. cuspidata and M. holttumii. Sect. 2. Pandanophyllum (Hassk.) B. & H. Gen. PI. 3 (1883) 1056. — Pandanophyllum Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 118, p.p. (T: Pandanophyllum palustre HAssk. ex STEUD.). — Lepironia sect. Pandano- phyllum Mia. Illustr. (1870) 60. — Mapania sect. Halostemma CLARKE, FI. Br. Ind. 6 (1894) 681; UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 284 (T: Mapania silhetensis CLARKE). — Mapania subg. Halostemma CLARKE, Kew Bull. add. ser. 8 (1908) 130; Uirtien, Rec. Trav. Bot. Néerl. 32 (1935) 185. Flowering stems lateral. Involucral bracts glume-like. Inflorescence always capitate. Sect. 3. Macrolepironia (MiQ.) KERN, Blumea 12 (1963) 25.— Lepironia sect. Macrolepironia MiqQ. Illustr. (1870) 60 (Lectotype: Lepironia enodis M1Q.). — Mapania sect. Pandanophyllum (non B. & H.) CLARKE, Fl. Br. Ind. 6 (1894) 682; Urrtien, Rec. Trav. Bot. Néerl. 33 (1936) 145. — Mapania subg. Pandanophyllum CLARKE, Kew Bull. add. ser. 8 (1908) 130; Urttien, Rec. Trav. Bot. Néerl. 32 (1935) 185. — Mapania subg. Pandanoscirpus UitTien, Rec. Trav. Bot. Néerl. 33 (1936) 278 (T: Mapania petiolata CLARKE). Flowering stems lateral. Inflorescence normally consisting of a single spikelet. Urrtien referred M. richardsii and M. longiflora to this section. However, their inflorescence is composed of several spikelets, exactly as in M. debilis, which he placed in the preceding section. In future the three might be treated as a separate section. 1974] CyPERACEAE (Kern) 469 In M. cuspidata, M. squamata, M. spadicea, and M. graminea there are not rarely some secondary spikelets branching from the axils of the lower (‘empty’) glumes; usually there is no distinct involucre, but sometimes the lower glumes assume the aspect of involucral bracts. Notes. Specific delimitation has proved to be difficult, this in contrast with the African species in which the structure of the fruit offers good characters. The flowers in many Malesian species vary considerably in size (partly due to age). Therefore differential characters have largely been based on vegetative characters and are less tangible than desirable. Herbarium material is often inadequate for critical study. Mapanias should be collected as whole plants, if possible both in flower and fruit. KEY TO THE SPECIES Texture and colour of the leaves are those of herbarium specimens ! 1. Leaves distinctly petioled. 2. Flowering stems central (arising from the centre of the shoot on which the normal leaves are borne), triquetrous, rough-hairy especially on the angles. Inflorescence capitate, with 8—SO spikelets, the lower 1—2 involucral bracts more or less foliaceous, tailed, overtopping the inflorescence . . 3. M. latifolia 2. Flowering stems (scapes) lateral (developing from shoots in the axils of the leaves or below the leaves), terete or trigonous, the base clothed with bladeless, scale-like sheaths. Inflorescence either a single spikelet or capitate, in the latter case involucral bracts not foliaceous, glume-like, not overtopping the inflorescence. 3. Inflorescence normally a single terminal spikelet, sometimes with 1—2(—4) sessile or peduncled spikelets in the axils of the lower aes Scapes glabrous and smooth, (}4—)1—2 mm thick. Leaves (4—)3—7 cm wide . 5. M. cuspidata 3. Inflorescence ‘always capitate, with 8-16 spikelets. Scapes. rough: hairy: (ultimately glabrescent), (2—)3-6 mm thick. Leaves 6-12 cm wide . . . . . M. holttumii 1. Leaves not petioled. 4. Flowering stems central (arising from the centre of the shoot on which the normal leaves are borne), triquetrous, nearly always with 1—2 cauline leaves. Inflorescence capitate, with several to numerous spike- lets, surrounded by foliaceous involucral bracts at least one of which much overtops the inflorescence. 5. Flowering stems stout to very stout, 4-8 mm thick. Leaves (2—)4—6 cm wide. Inflorescence with many to numerous spikelets. Spikelets 1/4— 3(— —4) cm long, flowers 10-15mm. . . . 1. M. macrocephala 5. Flowering stems rather slender, 14-2 mm thick. Leaves 0.6—1.1 cm wide. Inflorescence with 5-10 spikelets. Spikelets 1-114 cm long, flowersS5-7mm . . 2. M. moseleyi 4. Flowering stems (scapes) lateral (developing from shoots in the axils of the leaves or ‘below the leaves), terete or trigonous, the base clothed with bladeless scale-like sheaths; scapes sometimes very short to almost lacking. Inflorescence either a single spikelet or capitate, in the latter case involucral bracts not foliaceous, glume-like, not overtopping the inflorescence. 6. Apex of the leaves acute, but not caudate. Leaves with 3 prominent nerves, the latter all very scabrid on their raised surface almost from the base to the apex of the leaves . . . . 22. M. micropandanus 6. Apex of the leaves ending in a tail-like, triquetrous, scabrid acumen. Leaves with 1—several prominent nerves, but mostly only the midnerve distinctly scabrid on its raised surface. 7. Stem elongate, densely leafy, up to 50 cm long, producing in the leaf-axils solitary, subsessile spikelets. 23. M. maschalina 7. Stem short, not carrying a number of solitary, subsessile spikelets in successive leaf-axils. 8. Fruit with fleshy exocarp, berry-like, distinctly shrivelled when dried, yellow or orange (New Guinea). 25. M. baccifera 8. Fruit with dry exocarp, smooth to slightly rugulose, brown to blackish. 9. Inflorescence composed of several very densely packed spikelets at first sight hardly distinct from one another, and surrounded by some erect involucral bracts much firmer in texture than the pellucid, indistinctly nerved flower- -bearing glumes, the whole simulating a single spikelet. 10. Inflorescence (2—)2/4-4/4 cm long. Flowersc.20mmlong . . . . . . 7. M. longiflora 10. Inflorescence c. 1-14 cm long. Flowers 5—8 mm long. 11. Leaves broadly linear, 24-44 cm wide, abruptly contracted into the tail. Involucral bracts suborbicular . . . 6. M. richardsii 11. Leaves linear, 4- 1m wide, very gradually narrowed towards the apex. Involucral bracts elliptic or ovate... . . 8. M. debilis 9. Inflorescence either a single spikelet or composed of several to numerous distinct spikelets, in the latter case involucral bracts not strikingly different in texture from the coriaceous or chartaceous, distinctly nerved glumes. 12. Inflorescence always capitate, consisting of several to numerous distinct spikelets. 13. Bases of the leaves castaneous. 14. Leaves 1-3 cm wide. Involucral bracts 1-2 cm long. Glumes 9-12 mm long. Style usually bifid. Nutc.5by3mm. . . . 10. M. foxworthyi 14. Leaves 7-9 mm wide. Involucral bracts very short, 2-4 mm long. Glumes 34-4 mm long. Style trifid. Nut 3-3/4 by 144 mm_s Se ae hie op pms isa) ie cee ie een 13. Bases of the leaves stramineous to light brown. 470 FLORA MALESIANA [ser. I, vol. 7° 15. Glumes reddish brown, strongly nerved, almost ribbed when dry. Outermost flower-scales winged on the keel. Leaves with only the midnerve prominent 11. M. kurzii 15. Glumes stramineous, much less conspicuously nerved. Outermost flower-scales not winged. Leaves with 3 more or less prominent nerves . 9. M. palustris 12. Inflorescence normally consisting of a single terminal spikelet, ‘sometimes (on the same plant) some of them with 1—4 lateral sessile or peduncled spikelets added. 16. Bases of the leaves castaneous to blackish. 17. All spikelets sessile, triquetrous. Outer glumes sharply keeled. Flowers 15—30 mm a Nut c. 10 mm long, the 3—4 mm long conical beak included a i ek cnae M. sessilis 17. Spikelets at least partly peduncled. Outer glumes concave. 18. Leaves somewhat plicate lengthwise along several distinct lateral nerves . 18. Leaves with 1 or 3 prominent nerves. 19. M. tenuiscapa 19. Leaves very shining above, smooth, very densely crowded, strikingly 3-ranked, the rows very prominent and fan-like at the base; midnerve prominent beneath, 2 whitish lateral nerves somewhat prominent above. Scapes scabrous, or (at length?) glabrescent . 18. M. spadicea 19. Leaves when dry finely transversely wrinkled by numerous cross-nervules, not strikingly 3-ranked; midnerve prominent beneath, no prominent whitish lateral nerves. Scapes smooth. 17. M. lorea 16. Bases of the leaves stramineous or light brown, in M. graminea with castaneous margins. 20. Spikelets 6-10 mm long. Nut 2/4-2)4 mm long. Leaves 2—4 mm wide . 24. M. monostachya 20. Spikelets and nuts much longer. Leaves at least 5 mm wide, usually much wider. 21. Nut lageniform (subglobose in the centre, distinctly beaked, constricted in the lower part and then conspicuously widened into the truncate base). Leaves remotely scabrous on the margins. Scapes usually long, (10—)45—90 cm 16. M. enodis 21. Nut ellipsoid, obovoid, or pyriform. Leaves with densely serrate-scabrous margins (except in M. borneensis). Scapes often shorter. 22. Leaves conspicuously tessellate or with numerous prominent cross-nervules. 23. Leaves subcoriaceous, with 3 prominent nerves, conspicuously narrowed towards base and apex. Margins of the leaves scabrid only in the upper part 15. M. borneensis 23. Leaves mostly thickly coriaceous, with only the midnerve prominent, hardly narrowed towards the base, gradually narrowed upwards. Margins of the leaves serrate-scabrous throughout . 22. Leaves not tessellate. 17. M. lorea 24. Leaves very densely crowded, strikingly 3-ranked, very prominently equitant and fan-like at the base. Upper surface of the leaves scabrous just below the tail. Spikelets trigonous when young 21. M. graminea 24. Leaves not strikingly 4. ranked, not scabrous on the upper surface. Spikelets terete. 25. Leaves somewhat plicate lengthwise along several distinct lateral nerves. 25. Leaves with 1 or 3 prominent nerves. 19. M. tenuiscapa 26. Leaves dark green, 1-2 cm wide. Scapes (2—)10—40 cm long. Inflorescence always a single spikelet. Outermost flower-scales not winged 14. M. wallichii 26. Leaves pale glaucous, (2—)3/4—-5 cm wide. Scapes 2 2- 4(— 10) cm nm long. Inflorescence not rarely with 2—5 spikelets. Outermost flower-scales winged on the keel. 1. Mapania macrocephala (GAuUDICH.) K. SCH. in Warb. Bot. Jahrb. 13 (1891) 265; K. Scu. & Laut. Fl. Schutzgeb. (1900) 189; Merr. Philip. J. Sc. 2 (1907) Bot. 422; VALcK. Sur. Nova Guinea 8 (1912) 711; Mere. Int. Rumph. (1917) 107; En. Philip. | (1923) 132 p.p. (excl. ELMER 11978); KUxK. Bot. Jahrb. 59 (1924) 56; REHD. J. Arn. Arb. 14 (1933) 65; OuwI, Bot. Mag. Tokyo 56 (1942) 211; S. T. BLAKE, J. Arn. Arb. 28 (1947) 212; Proc. R. Soc. Queensl. 58 (1947) 48: Koyama, Micronesica 1 (1964) 66: KERN, Pac. Pl. Areas 2 (1966) map. 32. — Carex laevis major RuMPH. Herb. Amb. 6 (1750) 21. — Hypaelyptum macrocephalum GAuDICH. in Freycin. Voyage, Bot. (1829) 415.— Hypolytrum macroce- phalum KunTH, En. 2 (1837) 273; SteuD. Syn. 2 (1855) 133; Mia. Fl. Ind. Bat. 3 (1856) 334 (err. ‘macro- phyllum’). — Cephaloscirpus macrocephalus Kurz, J. As. Soc. Beng. 38, ii (1869) 84; ScHerr. Nat. Tijd. N. I. 34 (1874) 91. — Lepironia macrocephala Mia. Illustr. (1870) 64, t. 27. — Pandanophyllum macro- cephalum Boeck. Flora 58 (1875) 116 (not based on 13. M. squamata H ypaelyptum macrocephalum GAUDICH.). — Pandano- phyllum longifolium Boeck. I.c. 112. — M. longifolia CLARKE, Kew Bull. add. ser. 8 (1908) 54; Ill. Cyp. (1909) t. 115, f. 1-7; Premrr. Bot. Arch. 12 (1925) 450, f. 14, 39. — M. pandanacea RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 246; ? KUxk. Bot. Jahrb. 59 (1924) 57. — M. lactea KUkK. /.c. 56, ex descr. — M. moseleyi f. latifolia Uirr. J. Arn. Arb. 20(1939) 213. — M. geelvinkensis OHW1, Bot. Mag. Tokyo 56 (1942) 210, ex descr. — M. margaritae OHwWI, /.c. 211, ex descr. — M. palauensis HosoKAWA, Trans. Nat. Hist. Soc. Form. 32 (1942) 7, ex descr. — M. yapensis HOSOKAWA, /.c. 8, ex descr. Usually stout to very stout, resembling a small Pandanus, forming dense clumps. Stems central, triquetrous, smooth or somewhat scabrous at the top, much shorter than the leaves, 30-120 cm by 4-8 mm (up to c. 10mm at the incrassate top), with 1—2 well developed leaves about the middle, very rarely cauline leaves absent. Leaves coriaceous, broadly linear, with 3 prominent nerves, obscurely 1974] CYPERACEAE (Kern) 471 septate-nodulose, with serrulate-scabrous margins and the keel serrulate especially towards the apex, narrowed towards the conduplicate base (but not petioled), rather abruptly long-caudate, bright green or more or less glaucous, 1—3(—4!4) m by (2—)4—6 cm; lower sheaths bladeless, stramineous; cauline leaves with a 5—10 cm long sheath somewhat widened above. Inflorescence capitate, very dense, ovoid or globose, (3-)5-7/4(—9) cm in diam., with many to numerous spikelets. Bracts several, the lower 1—3 very long (up to 1/4 m), finally deflexed, similar to the leaves, the upper ones much shorter, ovate, caudate. Spikelets oblong, 1/4—3(—4) cm by c. | cm. Glumes chartaceous, oblong-lanceolate, obtuse, 3-nerved, 9-12 by c. 3mm, the lower ones empty, several- nerved. Flowers linear, longer than the glumes, 10-15 mm. Anthers up to 4mm long. Szyle 3-fid. Nut obovoid or pyriform, 5-7 by 3—5 mm, at the base narrowed into a slender 2—4 mm long stipe, greyish or purplish brown, the slender 2~3 mm long beak finally caducous. Distr. N. Queensland (the only Mapania here known), Samoa Is., Solomons, W. Carolines; in Malesia only in the eastern part: Philippines (Bohol, Balut), Moluccas (Talaud Is., Morotai, Halmahera, Batjan, Ceram, Ambon), throughout New Guinea (also in the Admiralty Is., Bismarck Arch., Louisiades, P. Rawak near Waigeu, and Aru Is.). Fig. 11. ‘ Po * b ie ~ % ee - Fey Oe st Feats eee “ee ee S % s es ‘ oe Fig. 11. Localities of Mapania_ macrocephala (GAUDICH.) K.SCH. Ecol. In dense swampy rain-forests, sago swamps, on shady banks of streams, usually 0—1000 m, but in the Arfak Mts (Kofo Anggi) at 2000 m. Vern. Sosaréwu ma dorou, Halm., gérre, kiki kella, New Guinea. Notes. RIDLEY mistook a specimen of M. palustris for M. macrocephala, and described a specimen of the latter species as M. pandanacea. I have not seen the type-collections of M. lactea KUux., M. geelvinkensis OHW1, M. margaritae OHWI, M. palauensis Hosokawa, and M._yapensis Hosokawa. To judge from the descriptions all fall under the circumscription of M. macrocephala as adopted above; see also Koyama, I.c. Like several other Mapaniae, M. macrocephala is extremely variable as to size, width of leaves, and dimensions of inflorescences. The fruits remain on the decaying inflorescence for a long time; when finally the greyish brown exocarp has rotted away the whitish outside of the endocarp becomes visible. In all probability M. lactea and M. margaritae were based on specimens with such old fruits. CLARKE based M. longifolia on Pandanophyllum longifolium BorEcK., but the specimen in the Kew Herbarium he took for M. longifolia (and figures as such in Ill. Cyp. t. 115) is undoubtedly syntype of Pandanophyllum macrocephalum BoECk. (‘‘spiculis . . . magnis conicis acutiusculis’’). For the rest, to me BOECKELER’s Pandanophyllum longifolium and P. macrocephalum are not specifically distinct. According to a field-note the flowers are fragrant. 2. Mapania moseleyi CLARKE, Kew Bull. add. ser. 8 (1908) 55; VALcK. SuR. Nova Guinea 8 (1912) 711; S. T. BLake, J. Arn. Arb. 28 (1947) 212.— M. ledermannii KUK. Bot. Jahrb. 59 (1924) 57; OHw1, Bot. Mag. Tokyo 56 (1942) 212. Stems central, slender but rigid, triquetrous, with scabrous angles at the top, much shorter than the leaves, 25—50 (—90) cm by 1/4—2 mm (up to 3 mm at the somewhat incrassate top), with 1 or 2 well developed cauline leaves. Leaves numerous, coria- ceous, linear, not petioled, with the midnerve prominent on the underside and several much less prominent lateral nerves, conduplicate at the base, with slightly revolute, spinulose-scabrous margins and the keel on the underside scabrous especially towards the apex, gradually tapering to a very long and very scabrous acumen, obscurely septate-nodu- lose, pale green, 6-11 mm wide, the lower sheaths bladeless or shortly laminate, pungent, stramineous to brownish. Inflorescence capitate, globose, with 5—6(-10) spikelets, 1/4 (finally 2/4-3) cm across. Bracts 3—5, the lower 1—2 very long, finally deflexed, similar to the leaves, the upper ones shorter, sub- erect. Spikelets ellipsoid, 1-1/4cm by 4-6 mm. Glumes chartaceous, ovate, rounded at the apex, with hyaline margins especially towards the apex, 3—5-nerved, brown or stramineous, c. 6 by 4mm; lower sterile glumes 5S—7-nerved. Flowers 5—7 mm long; outermost scales with narrowly winged keel; wings c. 4mm _ wide, densely ciliate. Anthers 2-2/4 mm long. Style 3-fid. Nut ellipsoid or obovoid, brown, 5—7 by 2)4—3 mm (beak included), the base narrowed into a broad c. 1 mm long stipe, the apex rostrate by the short conical style. Distr. Solomons (Bougainville), New Britain, Admiralty Is.; in Malesia: New Guinea (W. New Guinea: Japen-Biak, Nabire, Rouffaer R., Idenburg R., Cycloop Mts; NE. New Guinea: Sepik Distr.; Papua: Mt Musgrave, Mt Knutsford, Fergusson I., Normanby I.). Fig. 12. Ecol. In rain-forests, 200—1500 m. Fig. 12. Localities of Mapania moseleyi CLARKE. 472 FLORA MALESIANA [ser. I, vol. 7° 3. Mapania latifolia UiTTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 199; ibid. 33 (1936) 143. — M. montana RIDL. J. Str. Br. R. As. Soc. n. 44 (1905) 206; MERR. J. Strci Br: Re As “Soc. nn? 76'(1917). 79; En: Born: (1921) 65; non Laut. & K. Scu. 1900, nec UITTIEN, 1925. — Fig. 9. Stolons up to 15 cm long, covered with lanceolate, acute, many-nerved scales; brownish sheaths often present. Leaves subcoriaceous, lorate, with prominent midnerve and 2 of the lateral nerves more or less prominent, scabrous on margins and keel only at the very top, dark green above, paler beneath, 15-75 by 3-7 cm, the apex somewhat notched or suddenly contracted into a stiff, triquetrous, aculeate, 1-6 cm long tail, the base contracted into a con- duplicate ribbed petiole 15—SO cm long and 3—S mm wide; leaf-sheaths broad, purplish, the lower ones bladeless. Scapes terminal, triquetrous, gradually thickened towards the top, setulose-scabrous espe- cially on the angles, rusty when dry, 15—50 cm long, 2-3 mm thick at the base, 5—7 mm at the top, the base destitute of scales. Inflorescence capitate, with 8-50 densely crowded spikelets, (2—)3—5 cm across. Bracts c. 3, the outer 1-2 more or less leafy and overtopping the inflorescence, abruptly narrowed into a 3-7/4 cm long tail, the lowest sometimes slightly distant, similar to the leaves, shortly sheath- ing, up to 45 by 7cm. Spikelets oblong, acute or obtusish, 1-2 cm by 5—6 mm. Glumes oblong-ovate to oblong, obtuse, somewhat cucullate, prominently many-nerved, 7—9 mm long. Flowers about as long as iB» the glumes. Anthers linear, 3—4 mm. Style (2—)3- cleft. Nut ellipsoid or subpyriform, brown, 4 by 2-2/4 mm. Distr. Malesia: Borneo (Sarawak: Gat, Mt Dulit; N. Borneo: Mt Kinabalu, Kinabatangan, Mt Nunkok; E. Borneo: W. Kutei). Ecol. In primary forests, along streams, at low and medium altitudes, on Mt Kinabalu up to 1650 m. Note. In some spikelets I found several bifid styles among the trifid ones. 4. Mapania holttumii KERN, Blumea 9 (1958) 215. — M. insignis Hott. Gard. Bull. Sing. 11 (1947) 293, non SANDW. 1933. — Fig. 13. Very stout. Leaves subcoriaceous, lorate, drooping (the tip pointing downwards), with 3 more or less prominent nerves, scabrous on margins and keel only at the very top, dark green above, paler beneath, (30—)50—70 by 6-12 cm, the apex rather gradually narrowed into an up to 15 cm long aculeate tail, the base contracted into a conduplicate, ribbed petiole up to 60cm long and 6-10 mm wide; leaf-sheaths broad, purplish, up to 20 cm long, the lower ones bladeless. Scapes lateral, solitary in the lower leaf- axils or below the leaves, robust, obtusely trigonous, hardly incrassate at the top, very shortly and densely rough-hairy throughout, ultimately glabrescent, rusty when dried, 10—-35cm by (2—)3—6mm, the base clothed with some stiff, acute, up to 8 cm long, striate scales. Inflorescence capitate, consisting of Geo arn ZN % >. ~ Fig. 13. Mapania holttumii KERN in niapecann yi primary forest iphoieks: Counen; 1935; SF 30544). 1974] CYPERACEAE (Kern) 473 8-16 sessile spikelets, 3-6 cm across. Bracts ovate to lanceolate, acute, stiff, up to 3 cm long. Spikelets oblong, acute, 2—3 by lcm. Glumes narrowly oblong, with rounded apex, many-nerved, 9-11 mm long. Flowers about as long as the glumes; outermost scales ciliate on the keel near the apex only. Anthers linear, 3—5 mm. Style 3-cleft. Nut unknown. Distr. Malesia: Malay Peninsula (Trengganu: G. Padang; Pahang: G. Tahan, Karak For. Res.; Kemaman: Sungai Nipa). Ecol. In forests at low and medium altitudes, up to 1200 m. Note. Closely related to M. cuspidata, with the same vegetative habit, but much stouter. The very large leaves are scabrous at the very top only, the scapes thick and short-hairy (but finally glabrescent). The capitate inflorescence with several to many spikelets furnishes an easy character for discrimina- tion, but as M. cuspidata may sometimes have a few secondary spikelets in the axils of the basal glumes, the distinction between the two species on this ground is not sharp. According to a field-note of Corner the tufts are not stilted, unlike those of M. cuspidata. 5. Mapania cuspidata (MiQ.) UiTTieN, J. Arn. Arb. 20 (1939) 213; S. T. BLAKE, J. Arn. Arb. 28 (1947) 211; Hoxtr. Gard. Bull. Sing. 11 (1947) 293; UiTTIEN in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 49; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 454. — Lepironia cuspidata MiQ. Sum. (1861) 263, 603. — Pandanophyllum zippelianum Kurz, Nat. Tid. N. I. 27 (1864) 224, nom. nud. — Pandanophyllum humile (non STEUD.) OuDEM. Bot. Zeit. 23 (1865) 193; Kurz, J. As. Soc. Beng. 38, 11 (1869) 82; SCHEFF. Nat. Tijd. N.I. 34 (1874) 90. — Lepistachya praemorsa Zipp. ex Kurz, J. As. Soc. Beng. 38, 11 (1869) 82, in syn. — Lepironia humilis Mia. Illustr. (1870) 61, t. 23, quoad specim., haud Pandanophyllum humile SteuD. — M. humilis F.-VitL. Nov. App. (1882) 309, quoad specim., haud Pandanophyllum humile STEUD.; RIDL. J. Str. Br. R. As. Soc. n. 23 (1891) 14; CLARKE, FI. Br. Ind. 6 (1894) 683; Rip. J. Str. Br. R. As. Soc. n. 46 (1906) 227; CLARKE, Philip. J. Sc. 2 (1907) Bot. 109; Ripv. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105; Koorpb. Exk. Fl. Java 1 (1911) 203; Atlas (1922) f. 268; VALCK. SUR. Nova Guinea 8 (1912) 711; MERR. En. Born. (1921) 64; En. Philip. 1 (1923) 132; KUx. Bot. Jahrb. 59 (1924) 56; RipL. Fl. Mal. Pen. 5 (1925) 174; H. Preirr. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 174. — Pandanophyllum wendlandi Hort. ex Gard. Chron. 21 (1884) 711.— M. lucida N. E. Brown, Ill. Hort. 32 (1885) 77, t. 557. — M. triquetra RIDL. J. Str. Br. R. As. Soc. n. 41 (1904) 51; ibid. n. 46 (1906) 227; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 106; Rip_. & WINKL. Bot. Jahrb. 44 (1910) 525; Merr. \e = a Fig. 14. Mapauia cuspidata (M1Q.) U1TTIEN in Tawao F. R. (photogr. W. MeveR, 1960; SAN 19434). 474 FLORA MALESIANA [ser. I, vol. 7° En. Born. (1921) 65; Ripi. Fl. Mal. Pen. 5 (1925) 174. — M. petiolata CLARKE, Kew Bull. add. ser. 8 (1908) 54; Merr. En. Born. (1921) 65; UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 280; KUxK. Bot. Jahrb. 69 (1938) 261.— M. platyphylla MERR. Philip. J. Sc. 11 (1916) Bot. 54; En. Born. (1921) 65. — M. caudata KUx. in Fedde, Rep. 29 (1931) 201. — M. inopinata UntTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 151, f. 3. —M. stolonifera UitTIEN, l.c. 279. — Fig. 14. Leaves subcoriaceous, the lower ones reduced to bladeless sheaths, the upper ones very variable in shape and size, lorate to broadly linear, usually with 3 prominent nerves, scabrous on margins and keel beneath towards the apex, dark green above, paler beneath, pink to purplish when young, (3—)10—90 by (4-)3-7 cm, the apex not rarely notched and suddenly to rather gradually narrowed into an up to 10 cm long aculeate tail, the base contracted into a conduplicate ribbed petiole dilated into a scariously margined inflated sheath in the lower part. Scapes solitary in the leaf-axils or below the leaves, usually rather robust, obtusely trigonous or almost terete upwards, slightly incrassate at the top, smooth, (2—)5—35 cm by (}4-)1-2 mm, the base clothed with some hard, ovate-lanceolate, acute, striate scales scarious on the margins. Inflorescence normally consisting of a single terminal spikelet, sometimes with 1—2(—4) digitately arranged sessile or peduncled lateral spikelets in the axils of the lower glumes. Spikelet oblong when young, ovoid when mature, acute, (7—-)15—40 by (7-)10-20mm. Lower 4-7 glumes empty, firm, ovate or oblong-ovate, obtuse, many-nerved, with scarious margins, fuscescent, (6-)8—-9 by (3!4-)5-6 mm, upper ones gradually smaller and less firm, fertile. Flowers about as long as the glumes. Anthers linear, white, 3/4-5 mm. Stigmas 3. Nut ellipsoid or narrowly ellipsoid, rounded at the base, shortly stipitate or almost sessile, acuminate, greyish brown to dusky black, (4—)5—7 by (2—)3 mm. Distr. Thailand, Solomons (Bougainville), almost throughout Malesia, in Java only in the western part, lacking in the Lesser Sunda Is. Ecol. In damp localities in primary forests, on shady banks of streams, at low altitudes (up to 1500 m). Use. In the Malay Peninsula the leaves are used as a remedy against fever. Vern. Sélingsingan, pandan bangkuang, duhut bakkuang, bangkuang laju, simarbakkuang, M (Sum.), sohlenat, kigémbang, harassas, gébang, S, siak-siak rimba, pandan hutan, Mal. Pen., nanasaka, Talaud Is., héhéwéhé mabéeka, Morotai, kihadiéd, Ceram; Philip.: babakal, Sub., lubigan, P.Bis., maribari, S.L.Bis., malalubigan, Tag. Notes. In old stout specimens the old part of the stem has died off and the rosettes of leaves stand on thick stiff stilt roots. The flowers open late in the morning; the anthers hang on long filaments and fade rapidly. OUDEMANS (1865), who described a specimen cultivated in the Amsterdam Botanical Garden, was the first who applied HASsKARL’s nomen nudum Pandanophyllum humile to the species described above. Herein he was followed by most subsequent authors, but STEuDEL (1855) had given a quite different interpretation (see under Hypolytrum humile p. 494). STEUDEL’s interpretation has to be followed. The species is very variable, not only with regard to the shape and size of the leaf-blades, but also in the length and stoutness of the petioles and scapes, the size and number of spikelets, and even in the size of flowers and fruits. UITTIEN distinguished M. inopinata by the inflorescence consisting of (1—)3—5 spikelets, and referred it on this ground to sect. Halostemma. However, scapes bearing 2 or more spikelets (one or more in the axils of the lower normally empty glumes) are not rarely found in M. cuspidata, as was already observed by Kurz (1869), MiQuEL (1870), GoeBeL (Ann. Jard. Bot. Btzg 7, 1888, 129), and CLARKE (1908). M. stolonifera UITTIEN is said to differ from M. cuspidata by the horizontal stolons, but a creeping, often branched rhizome is always present in completely collected old specimens of M. cuspidata. The varieties UITTIEN distinguished were mainly based on the shape of the leaf-blades. They are connected by many intermediates. The intermediate form (var. petiolata) occurs throughout the area of the species, broad-leaved plants (var. cuspidata) have only been found in the western part, and narrow-leaved specimens (var. angustifolia) prevail in the Philippines and the Moluccas. Possibly they are only ecological forms. KEY TO THE VARIETIES 1. Leaf-blades small, 3—8(-12) by (4-)1-2 cm, usually only the midnerve prominent. Scapes very slender, almost filiform, c. 74 mm_ thick. Spikelets small, 7-10 mm long. Nut small, c. 4 yy 2mm ay. een ota pera eaees var. pumila 1. Leaf-blades larger, always with 3 prominent nerves. Scapes 1-2 mm thick. Spikelets (1/4—)2- 3)% cm long. Nut larger. 2. Leaf-blades 3—5 times as long as broad, up to 7cm broad, abruptly narrowed into the 114-3 mm thick petiole, the apex truncate to retuse; leaf-sheaths broadly ovate. var. cuspidata 2. Leaf-blades more than 5 times as long as broad. Petiole usually thicker. Leaf-sheaths narrower. 3. Leaf-blades 3!4-6cm broad, up to 90cm long, abruptly narrowed into the tail. var. petiolata 3. Leaf-blades (1—)2—3cm broad, shorter than in var. petiolata, rather gradually passing in the tail ae var. angustifolia var. cuspidata. — Pandoanophyllum _ wendlandi Hort. — M. lucida N. E. BRown. — M. triquetra Rip_t. — M. platyphylla Merr.— M._ inopinata UittT1en. — M. petiolata var. cuspidata UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 282. Distr. Malesia: Malay Peninsula, Sumatra, W. Java, Borneo. var. petiolata (CLARKE) UitTiEN, J. Arn. Arb. 20 (1939) 213; OHw1, Bot. Mag. Tokyo 56 (1942) 212; UrrTien in Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 49.—M. petiolata CLARKE. — M. humilis var. petiolata Rip. Fl. Mal. Pen. 5 (1925) 174; H. Preirr. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 174. Distr. As the species. 1974] CYPERACEAE (Kern) 475 var. angustifolia (UITTIEN) UiTTIEN, J. Arn. Arb. 20 (1939) 214. — M. petiolata var. angustifolia UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 282. Distr. Bougainville; Malesia: Philippines, Moluc- cas, New Guinea. var. pumila (UITTIEN) UiTTIEN, J. Arn. Arb. 20 (1939) 214.— M. petiolata var. pumila UrtTIENn, Rec. Trav. Bot. Néerl. 33 (1936) 283. — M. caudata KUK. in Fedde, Rep. 29 (1931) 201. — Probably a form of oligotrophic soil. — Fig. 15. Distr. Malesia: Malay Peninsula (Trengganu), Borneo, a few times collected. 6. Mapania richardsii UiTTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 195. Leaves subcoriaceous, broadly linear, with the midnerve prominent beneath and 2 of the lateral nerves prominent above, septate-nodulose by numer- ous cross-nervules, scabrous on the margins towards base and apex and on the midnerve near the top, gradually narrowed towards the conduplicate base but not petioled, abruptly contracted at the apex into a triquetrous, aculeate-scabrous, 1-8 cm long tail, 65-120 by 2)4-4)4 cm. Scapes lateral, slender, subterete, striate, smooth, slightly incrassate at the top, 13-17 cm by 1 mm, the lower 4—6 cm clothed with some loose, brownish, striate scales. Inflores- cence capitate, consisting of c. 6 very densely packed spikelets, simulating a single spikelet, broadly ellip- soid to globose, 9-11 mm long and wide. Bracts 3, coriaceous, erect, suborbicular, concave, many- ribbed, with scarious margins lacerate at the top. Spikelets strongly compressed, 7 mm long. Glumes thinly membranous, pellucid, narrowly oblong, obtuse, l-nerved or indistinctly 3-nerved, purplish puncticulate at the top, c. 7 by 2 mm. Flowers 6 mm long; outermost scales ciliate in the upper half. Anthers 3 mm long. Style 3-cleft. Nut unknown. Distr. Malesia: Borneo (Sarawak: Dulit trail: Belaga, Kapit Distr.). Ecol. In large tussocks in primary forests, all c. 800 m. Note. Ur1TieN placed this species next to M. cuspidata and M. borneensis, but in his system it should be referred to subg. Halostemma next to M. debilis and M. longiflora, which have the same type of inflorescence. 7. Mapania longiflora CLARKE, Kew Bull. add. ser. 8 (1908) 54; Ill. Cyp. (1909) t. 110, f. 7-13; MerR. En. Born. (1921) 64; Urttien, Rec. Trav. Bot. Néerl. 33 (1936) 146, excl. var. sessilis UITTIEN. Leaves coriaceous, linear, only slightly narrowed below, gradually narrowed upwards to the very slender, triquetrous, very scabrous tip, with 3 prominent nerves (the midnerve prominent beneath, scabrous, the lateral nerves prominent above), with aculeate-scabrous margins throughout their length, pale green or glaucous, 60-120 by 14-24% cm: sheaths dull stramineous to light brown. Scapes lateral, solitary in the leaf-axils or below the leaves, rather robust, trigonous upwards, slightly incrassate at the top, 15—30 cm by 2-4 mm, smooth, the base clothed with some hard, keeled, many-nerved scales up to 8cm long. Inflorescence capitate, seemingly consisting of a single spikelet because of the large, erect involucral bracts hiding the few to several densely packed spikelets inside, ellipsoid, obtuse, terete, (2-)2/4-44 by (1-)1)4-2 cm. Bracts 6-10, the lower ones slightly shorter than, the upper ones about as long as the inflorescence, coriaceous, concave or somewhat keeled, ovate, obtusish, many- striate, greenish, later on fuscescent, 2—3 cm long. Spikelets at first sight hardly distinct from one another, scarcely extending beyond the upper bracts, lanceolate, terete, many-flowered, stramineous, c. 2 by 4 cm. Glumes thinly membranous, pellucid, soon disintegrating into fibers, oblong-lanceolate, indistinctly 1—3-nerved, 1/4-2cm long. Flowers as long as the glumes, the lower ones often sterile or male, only the upper ones fructiferous; outermost scales not winged. Anthers linear, 5-6 mm. Style 3-cleft. Nut ellipsoid or subpyriform, smooth, greyish brown, stipitate (stipe 2-4 mm), acuminate, 5—6 by 3-3/4 mm (the 1-2 mm long beak included). Distr. Malesia: Malay Peninsula (Singapore: Bukit Timah), E. Borneo (Nunukan; E. Kutei: Sungei Susuk region; Banjermasin, Batulitjin). Ecol. In damp primary forests, along streams, at low altitudes. Vern. Sélingsing, SE. Borneo. Note. UrTIen placed M. longiflora next to M. enodis, M. squamata, etc. These are certainly not its nearest allies because of the quite different structure of the inflorescence. 8. Mapania debilis CLarkE in Ridl. J. Str. Br. R. As. Soc. n. 46 (1906) 227; Kew Bull. add. ser. 8 (1908) 53: Mere. En. Born. (1921) 64.— M. zeylanica var. RIDL. J. Str. Br. R. As. Soc. n. 46 (1906) 227. — M. gracilipes MERR. J. Str. Br. R. As. Soc. n. 85 (1922) 158.— M. flagellaris UttTieEN, Rec. Trav. Bot. Neéerl. 32 (1935) 198, f. 3. Short stolons covered with lanceolate, brown, striate scales sometimes present. Leaves subcoria- ceous or coriaceous, linear, slightly narrowed towards the conduplicate base, very gradually narrowed up- wards to the very slender, triquetrous, aculeate- scabrous tip, with the midnerve prominent beneath and some lateral nerves somewhat prominent above, with aculeate-scabrous margins throughout their length, glaucous, up to 90 cm by 5-10 mm; sheaths stramineous to brownish. Scapes lateral, from the caudex below the leaves, very slender, obtusely trigonous to terete, striate-sulcate, smooth, up to 20 cm by 1—2 mm, the base clothed with some loose, striate scales up to 5 cm long. Inflorescence capitate, when in flower seemingly consisting of a single spikelet, ovoid to subglobose, containing 3—5 very densely packed spikelets, 10-13 mm long. Bracts few, coriaceous, concave, ovate or elliptic, acutish, many-nerved, with broad, scarious, brownish punc- ticulate-lineolate margins, 9-13 mm long. Spikelets at first sight hardly distinct from one another, somewhat extending beyond the bracts, ellipsoid. stramineous, 5S—10 by 2-4 mm. Glumes thinly mem- branous, pellucid, oblong, obtuse, indistinctly 1- nerved, brownish puncticulate-lineolate at the top, 5—8 mm long. Flowers as long as the glumes: keel of the outermost scales minutely ciliolate at the top. Anthers c.4 mm. Style 3-cleft. Nut obovoid-pyriform, slightly rugulose, greyish brown, shortly stipitate (stipe /4 mm), acuminate, shortly beaked, c. 3 by 14-2 mm. Distr. Malesia: Borneo (Sarawak: Mt Poi, Mt Matang, Mt Dulit; N. Borneo: near Sandakan: 476 FLORA MALESIANA [ser. I, vol. 7° Fig. 15. Mapania cuspidata (MiQ.) UITTIEN var. pumila (UitTIEN) UrtTiEN. Habit, x )4 (ENDERT 3557). Wb dale: . : a ee ene ee eee ee 1974] CYPERACEAE (Kern) 477 E. Borneo: W. Kutei; also collected by TEYSMANN in W. Borneo, without definite locality). Ecol. In damp primary forests at low altitudes, up to 1200 m. Note. Both M. debilis and M. gracilipes are known from a single, rather poor collection only. M. flagellaris was separated from them on account of the coriaceous leaves with much longer acumen. As the length of the acumen in the specimens with thicker leaves is variable, and the leaves of M. gracilipes are intermediate in texture between those of M. debilis and M. flagellaris, 1 think the three should be united, at least provisionally, until more material will be available. 9. Mapania palustris (HASSK. ex STEUD.) F.-VILL. Nov. App. (1882) 309; CLARKE, FI. Br. Ind. 6 (1894) 681; J. Linn. Soc. Bot. 34 (1894) 95; Ill. Cyp. (1909) PaOo- RIDE. Jo Str. Bro Re As. Soc: n223)(1891) 15; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 103; Koorp. Exk. Fl. Java 1 (1911) 203; Atlas (1922) f. 269; MERR. Philip. J. Sc. 9 (1914) Bot. 267; En. Philip. 1 (1923) 132; PremrF. Bot. Arch. 12 (1925) 449, f. 7, 34, 51; Rip. Fl. Mal. Pen. 5 (1925) 172; UmTien, Rec. Trav. Bot. Néerl. 32 (1935) 194; ibid. 33 (1936) 285; in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 49; Kern in Back. & Bakh. f. Fl. Java 3 (1968) 455. — Pandanophyllum palustre [? Hassk. Tijd. Nat. Gesch. Phys. 10 (1843) 119; Cat. Hort. Bog. (1844) 297; Mor. Syst. Verz. (1846) 99; ZoLL. Syst. Verz. 1 (1854) 61, nom, nud.] ex SteUD. Syn. 2 (1855) 134; Mia. Fl. Ind. Bat. 3 (1856) 334; Kurz, J. As. Soc. Beng. 38, ii (1869) 78, incl. var. malesica KURZ & var. silhetana KURZ, excl. WALLICH 3541; BOECK. Linnaea 37 (1871) 138; ScHerr. Nat. Tijd. N. I. 34 (1874) 89.— Pandanophyllum zeylanicum (non THWAITES) KuRzZ, J. As. Soc. Beng. 38, ii (1869) 80, p-p., quoad specim. andam. — Lepironia palustris Mig. Illustr. (1870) 63, t. 25.— M. andamanica CLARKE, FI. Br. Ind. 6 (1894) 681; J. Linn. Soc. Bot. 34 (1898) 95; UitTreNn, Rec. Trav. Bot. Néerl. 33 (1936) 288.— M. silhetensis CLARKE, FI. Br. Ind. 6 (1894) 681; J. Linn. Soc. Bot. 34 (1898) 94. — M. kurzii (non CLARKE) MerR. Philip. J. Sc. 2 (1907) Bot. 422; C. B. Ros. ibid. 6 (1911) Bot. 194; MerRR. En. Philip. 1 (1923) 132; E_m. Leafl. Philip. Bot. 10 (1938) 3537. — M. albescens CLARKE, Kew Bull. add. ser. 8 (1908) 54; Rip. Fl. Mal. Pen. 5 (1925) 172; Urrtien, Rec. Trav. Bot. Neéerl. 33 (1936) 288. — M. banahaensis ELM. Leafi. Philip. Bot. 2 (1909) 574; UiTiEN, Rec. Trav. Bot. Néerl. 33 (1936) 288. — M. macrocephala (non K. SCH.) RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 245. — M. affinis MERR. J. Str. Br. R. As. Soc. n. 85 (1922) 157; En. Philip. 1 (1923) 131; UrrTiEN, Rec. Trav. Bot. Néerl. 33 (1936) 287. — M. grandiceps KUxk. Bot. Jahrb. 59 (1924) 55, ex descr.; UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 287; non OHwW1, Bot. Mag. Tokyo 56 (1942) 211.— M. longiflora (non CLARKE) PFEIFF. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 173.— M. javana UrTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 287; in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 50. Leaves coriaceous, linear to broadly linear, with the midnerve prominent beneath and 2 of the lateral nerves more or less prominent above, slightly narrowed towards the conduplicate base, gradually to rather abruptly narrowed to the up to 10cm long, aculeate-scabrous, triquetrous tail, aculeate-scabrous on the margins and on the keel beneath, up to 2/4m by (2-)3-6cm; sheaths brownish. Scapes lateral, trigonous to almost terete, densely furfuraceous to almost smooth, (10—)30- 70 cm by (2—-)3—6mm, up to 15 mm thick at the incrassate top, sheathed at the base with some loose, ovate to lanceolate, striate, up to 15 cm long scales. Inflorescence capitate, semiglobose to globose, with few to numerous spikelets, (1/4-)4-6(-8) cm across. Bracts 4—8, coriaceous, ovate to lanceolate, somewhat shorter than to about as long as the inflorescence, many-ribbed. Spikelets usually 10-50, sometimes only 3—5, not rarely up to 100, very densely packed, ellipsoid, 114-2 by 5-10 mm. Glumes subcoriaceous, oblong or lanceolate, often lacerate at the top. Flowers about as long as the glumes, (7—)10—15 mm long. Outermost flower-scales ciliate on the keel, the third (anterior) scale ciliate on both keels. Stamens 4-8 mm long. Style 3-cleft. Nut obovoid, shortly stipitate, with small recurved beak, greyish brown, c. 4 by 2-2/4 mm. Distr. SE. Asia (from Assam to Indo-China and S. China); in Malesia: Sumatra, Riouw Arch., Malay Peninsula, W. Java, Borneo, Natuna_Is., Philippines, New Guinea. Ecol. In wet or muddy places in damp primary forests, along streams, on wet rocks, from few m altitude up to 1500 m. Use. In the Malay Peninsula the leaves are used for making mats and baskets. Vern. Harashas, bangkonoh, S, ménkuang tédong, ménkuang lobo, Mal. Pen. (from its resemblance to the méngkuan, Pandanus furcatus ROXB.), assingsing, Sum. E. C.; Philip.: kulibang, blas, Sub. Notes. The most variable species of the Malesian Mapaniae. However different in appearance the extremes may be, it seems impossible to treat them as separate species. The characters used by UITTIEN (Rec. Trav. Bot. Néerl. 33, 1936, 285, spp. 4—10) are unfit for practical use, as all the ‘species’ are con- nected by numerous intermediates. M. andamanica CLARKE and M. banahaensis ELM. are more slender and have smaller — often much smaller — spikelets and flowers than typical M. palustris. As was already stated by UrtTiEN, M. albescens CLARKE differs from M. andamanica only by the pale colour of the dried specimens. M. affinis MERR. was separated on account of the larger inflorescences and the shorter glabrous scapes. However, still larger inflorescences and glabrous scapes are often found in M. palustris. M. grandiceps KUK. is also a stout form, but with scabrid scapes. I fail to see any difference whatever between M. silhetensis CLARKE and M. javana UtITTIEN. Both have inflorescences with few large spikelets. This form may have some taxonomical value; KURZ treated it as a variety. MIQUEL (1870) very properly remarked that the size of the spikelets diminishes when their number increases. CLARKE, who saw but a few specimens of typical M. palustris, thought M. silhetensis were also characterized by the smooth scapes in contrast to the scabrous scapes of M. palustris, and UITTIEN characterized M. javana also by the leaf-bases rounded on the back. These charac- ters do not hold good. In CLaRKeE’s flower diagram (1909; also in PFEIFFER, |.c., whose diagrams are obviously copied 478 FLORA MALESIANA [ser. I, vol. 7° from those of CLARKE’s) the outermost flower scales are figured as being connate. I always find them completely free. The specimen from “East Java, PLOEM” cited by UnrTIEN (1936, p. 287) was collected in W. Java. No Mapania is known to occur in E. Java. Stout specimens of M. palustris are often confused in the herbaria with M. macrocephala, but the latter is readily recognizable by the central flowering stems and the very long leaf-like bracts of the inflorescence. 10. Mapania foxworthyi Mere. Philip. J. Sc. 11 (1916) Bot. 53; En. Born. (1921) 64; UrtTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 286. Leaves coriaceous, linear, with only the midnerve prominent, hardly narrowed towards the condupli- cate base, gradually attenuate towards the very slender, triquetrous, aculeate-scabrous tip, densely aculeate-scabrous throughout the margins and on the upper part of the midnerve on the lower side, dark green, up to 170 by 1—3 cm; sheaths castaneous. Scapes lateral, trigonous, slightly thickened at the apex, smooth, 30-80 cm by 3—4 mm (up to 6 mm at the top), the base clothed with some lanceolate acute sheaths scabrous at the top and up to 15 cm long. Inflorescence capitate, globose or semi-globose, composed of c. 20 densely crowded spikelets, 3-4 cm across. Bracts ovate to lanceolate, many- ribbed, with scarious margins, scabrid at the top, 1-2 cm long. Spikelets ovoid to ellipsoid, 1)4-2 cm by 8-10 mm. Glumes coriaceous, oblong, obtuse, strongly many-nerved, reddish brown, 9-12 by 5—6 mm, the flower-bearing ones much less firm, 3-nerved. Flowers about as long as the glumes, 9-14 mm long; outermost scales ciliate in the upper half, not or hardly winged. Anthers linear, 4-5 mm. Style bifid, rarely some trifid. Nut obovoid, brown, c. 5 by 3 mm, the very short beak included. Distr. Malesia: Borneo (Sarawak: Lundu, Mt Poi). Ecol. In forests, up to 1700 m. Note. Very near to M. kurzii, but apparently satisfactorily characterized by the castaneous leaf- sheaths, the larger flowers and anthers, the oblong glumes, and the usually bifid style. It was collected in the type locality (Mt Poi) as early as 1866 by BECCARI (PB2401). 11. Mapania kurzii CLARKE, Fl. Br. Ind. 6 (1894) 681: J. Linn. Soc. Bot. 34 (1898) 95; RipL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 104; Fl. Mal. Pen. 5 (1925) 172; UitTiEN, Rec. Trav. Bot. Néerl. 33 (1936) 288. — M. multispicata CLARKE [ex RIDL. J. Str. Br. R. As. Soc. n. 23 (1891) 15, nom. nud.] Fl. Br. Ind. 6 (1894) 682, p.p.; J. Linn. Soc. Bot. 34 (1898) 95, p.p., excl. specim. jav. — M. longispica RIDL. J. Str. Br. R. As. Soc. n. 44 (1905) 205; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 104; Fl. Mal. Pen. 5 (1925) 172. — M. valida Rip. J. Str. Br. R. As. Soc. n. 44 (1905) 205, excl. RIDLEY 1714; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 104; Fl. Mal. Pen. 5 (1925) 172. Leaves coriaceous, linear, with only the midnerve prominent beneath, and several very indistinct lateral nerves, hardly narrowed towards the con- duplicate base, gradually attenuate to the very slender, triquetrous, aculeate-scabrous tip, densely aculeate-scabrous throughout the margins and on the upper part of the keel on the lower side, glaucous, up to 120 by (1—)2(—3) cm; sheaths stramineous to light brown. Scapes lateral, trigonous, slightly thick- ened at the apex, smooth, 10-40 cm by 1-3 mm, the base clothed with some lanceolate, acute sheaths up to 15 cm long. Inflorescence capitate, with 5—15 spikelets, 2-S cm long and wide. Bracts ovate to lanceolate, many-ribbed, with scarious margins, 1—2 cm long. Spikelets ovoid or ellipsoid to cylindri- cal, 1-5 cm by 6-8 mm. Glumes coriaceous, ovate, obtuse, strongly many-nerved, almost ribbed when dry, reddish brown, 7—9 by 5—6 mm, the upper ones less firm, 3—5-nerved. Flowers about as long as the glumes or slightly longer; outermost scales with a narrow, membranous, ciliate wing on the keel. Anthers linear, 2-3 mm. Style 3-fid. Nut ellipsoid or pyriform, shortly stipitate, brown, 474-5 by 2- 2/4 mm, the 1-1)4 mm long, recurved beak in- cluded. Distr. Thailand; in Malesia: Central Sumatra (W. Indragiri: Taluk region), widely distributed in the Malay Peninsula (Perak, Selangor, Negri Sem- bilan, Malacca, P. Penang, Singapore). Ecol. In primary forests, at low altitudes, up to 950 m. Notes. Closely allied to M. palustris, but well characterized by the 1-nerved leaves, brown inflo- rescence, the ribbed bracts and glumes, and the narrow wings of the outermost flower scales. Very variable in width of the leaves, thickness and length of the scapes, and number and length of the spikelets. On the whole Perak specimens have long cylindrical spikelets, but some have them short. Penang speci- mens have mostly very slender scapes and few, rather short spikelets. The specimens with long cylindrical spikelets were separated as M. longispica RIDL., but there is no other difference with typical M. kurzii. The name M. multispicata was intended by CLARKE as a new name for Pandanophyllum humile HAssk. ex STEUD., but published with a description drawn up from a specimen of M. kurzii (see Hypoly- trum humile, p. 494). 12. Mapania angustifolia U1TTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 197, f. 2. Leaves coriaceous, linear, with prominent mid- nerve beneath and 2 of the lateral nerves more or less prominent above, hardly narrowed at the conduplicate base, gradually narrowed at the top into a long aculeate-scabrous acumen, serrulate- scabrous throughout the margins and on the midnerve in the upper part, glaucous, up to 80 cm by 7-9 mm; sheaths shining castaneous. Scapes lateral, obtus- angular, striate-sulcate, scaberulous especially to- wards the top, 18-30 cm by 1 mm, the base clothed with some membranous, brown, acute, striate sheaths up to 5 cm long. Inflorescence capitate, consisting of 3-5 spikelets, c. 1 by 1/4-2/% cm. Involucral bracts c. 3, very small, ovate or triangular, 2-4 mm long. Spikelets narrowly ellipsoid or lanceolate, acute, 7-13 by 3-4mm. Glumes coriaceous, ovate to oblong, rounded at the apex, indistinctly 3-nerved, fuscous, with scarious margins, 3/4—4 by c. 2mm. Flowers as long as the glumes; outermost scales minutely ciliolate on the keel. Style 3-fid. Nut ovoid- ellipsoid, acuminate at both ends, stipitate (stipe c. % mm), rugulose, brown or greyish brown, 3-3% by 1/4 mm, the short, conical, recurved beak included. 1974] CYPERACEAE (Kern) 479 Distr. Malesia: Borneo (Sarawak: Mt Dulit). Ecol. In tussocks on steep stony slope in primary forest, under 300 m. 13. Mapania squamata (KURZ) CLARKE, Kew Bull. add. ser. 8 (1908) 53; Pretrr. Bot. Arch. 12 (1925) 449, f. 9: UitTiEN, Rec. Trav. Bot. Néerl. 33 (1936) 149; in Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 50; KERN in Back. & Bakh. /. Fl. Java 3 (1968) 455. — Pandanophyllum squamatum Kurz, J. As. Soc. Beng. 38, ii (1869) 80; ScHEFF. Nat. Tijd. N. I. 34 (1874) 90.— Lepironia squamata Mig. Illustr. (1870) 64, t. 26. — M. wallichii (non CLARKE) RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105, p.p., quoad RipLey 11475; Fl. Mal. Pen. 5 (1925) 173, p.p. — M. heterocephala Merr. J. Str. Br. R. As. Soc. n. 85 (1922) 158. Leaves subcoriaceous, broadly linear, with the midnerve prominent beneath and 2 of the lateral nerves prominent above, densely serrulate-scabrous throughout the margins and on the upper part of the midnerve on the lower surface, pale glaucous, slightly narrowed towards the conduplicate base, rather suddenly narrowed upwards into a 7-14 cm long, triquetrous, very scabrous tail, 50—150(—180) by (2-)3)/4-5cm; sheaths stramineous to light brown. Scapes lateral, from the caudex below the leaves, obtusely trigonous to almost terete, smooth, usually very short (2-4 cm), rarely up to 10cm, 2-3 mm thick, clothed with ovate-oblong, acute, concave, striate scales usually almost to the top. Inflorescence consisting of a single spikelet or composed of 2-5 fasciculately arranged spikelets (both types often occurring on the same plant). Spikelets ovate or lanceolate in outline, terete, acute, 2-3 by 1-2)4cm. Lower sterile glumes coriaceous, oblong-elliptic, obtuse or lacerate at the top, many-nerved, brown with scarious margins, c. 14 cm long, the upper fertile ones chartaceous, less nerved. Flowers (11—)15(-17)mm long; keel of the outermost scales distinctly winged, the wing c. 4mm wide, brown, ciliate at the top. Anthers linear, c. 6 mm. Style 3-cleft. Nut ovoid, indistinctly 3-ribbed, slightly rugulose, greyish brown, shortly stipitate (stipe c. 1 mm), 6-7 by 2)4-3)4 mm (the recurved beak included). Distr. Malesia: Malay Peninsula (Selangor: Batu Tiga; Johore: Gunong Panti; Singapore: Bukit Timah), W. Java (very rare), Borneo (Sarawak: Matang; N. Borneo: Sandakan, Mt Kinabalu; E. Borneo: Nunukan, W. Kutei). Ecol. In damp primary forests, along streams, at low and medium altitudes, up to 1500 m. Note. Not known from Sumatra and Central Java; the records ““Palembang” and “Sumatra” (UITTIEN, l.c.) refer to leaves of Pandanaceae; WALLICH 3541, also cited under M. squamata, is the type collection of M. wallichii. 14. Mapania wallichii CLARKE, Fl. Br. Ind. 6 (1894) 682; J. Linn. Soc. Bot. 34 (1898) 95; Ill. Cyp. (1909) t. 110, f. 1-6; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105, p.p., excl. RipLey 11475; CLARKE, Kew Bull. add. ser. 8 (1908) 130 (‘wallichiana’); Merr. En. Born. (1921) 65; RipL. Fl. Mal. Pen. 5 (1925) 173, f. 220, p.p.; Uirtien, Rec. Trav. Bot. Néerl. 33 (1936) 148. — Pandanophyllum palustre (non Hassk. ex STEUD.) Kurz, J. As. Soc. Beng. 38, ii (1869) 79, p.p., quoad WALLICH 3541. Leaves coriaceous, linear, with only the midnerve prominent beneath and sometimes 2 of the lateral nerves somewhat prominent above, densely serrulate- scabrous throughout the margins and on the upper part of the midnerve on the lower surface, dark green, slightly narrowed towards the conduplicate base, very gradually narrowed into a long, aculeate- scabrous, triquetrous acumen, shining above, dull beneath, up to 150 by 1—2 cm; sheaths stramineous to light brown. Scapes lateral, from the caudex below the leaves, obtusely trigonous to terete, smooth, striate, very variable in length, usually much longer than in M. squamata, (2—)10—40 cm by 2-3 mm, hardly incrassate at the top, the base clothed with some ovate to oblong, striate scales up to 4cm long. Inflorescence always a single spikelet, ovoid-conical, terete, acute, 2-4 by 1/4-2 cm. Sterile glumes numerous (up to 50), the lower ones coriaceous, broadly ovate or ovate, obtuse, the upper ones oblong, chartaceous; fertile glumes 15-20 mm long, less nerved. Flowers about as long as the glumes. Anthers linear, c. 8 mm. Style 3-cleft. Keel of the outermost scales ciliate in the upper half or narrowly winged and ciliate. Nut (immature) ellip- soid or obovoid, shortly beaked, greyish brown to blackish, c. 5 by 3 mm; stipe 1 mm. Distr. Malesia: P. Musala (W. of Sumatra), Banka, P. Lingga, Malay Peninsula (Singapore, P. Batam), Borneo. Ecol. In primary forests, marshy coastal forests, along streams, at low altitudes (up to 500 m). Vern. Séding babi, Banka. Note. M. wallichii is so near to M. squamata that I very much doubt whether the two are specifically distinct. As their distribution is practically the same, they cannot represent geographical races. In general M. wallichii is characterized by the stiff, narrow, dark green leaves, the inflorescence always consisting of a single spikelet, the usually much longer scapes, and the quite or almost wingless outermost flower- scales, but throughout the genus these characters are not very satisfactory. The question should be studied in the field, e.g. on Bukit Timah (Singapore), where both occur. 15. Mapania borneensis Merr. J. Str. Br. R. As. Soc. n. 76 (1917) 78; En. Born. (1921) 64. — M. dictyo- phlebia S. T. BLake, J. Arn. Arb. 28 (1947) 211, t. 2. — Cf. M. alpina E_o., p. 484. Leaves subcoriaceous, broadly linear, with 3 prominent nerves, septate-nodulose by numerous cross-nervules, conspicuously narrowed towards the conduplicate base (but not petioled), rather gradually to abruptly contracted into a long, triquetrous, aculeate-scabrous acumen, spinulose-scabrous on the margins in the upper part, 30—65 cm by (10—)15- 25 mm. Scapes lateral, from the caudex below the leaves, slender, terete or subtrigonous, striate, smooth, 6-20 cm by c. 1 mm, the base clothed with some pale brown, striate, loose scales up to 5 cm long. Inflorescence consisting of a single spikelet, ovoid or oblong-ovoid, acutish, terete, finally 114-2 by 1/4 cm. Glumes subcoriaceous, ovate-oblong, obtuse, many-nerved, with scarious margins, 8- 10 mm long. Flowers about as long as the glumes: outermost scales ciliolate on the keel in the upper half. Style 3-cleft. Nut obovoid-pyriform, terete, slightly rugulose, brown, purplish spotted, shortly 480 FLORA MALESIANA [ ser. 1, vole stipitate (stipe 4-74 mm), c. 4 by 2-244 mm, the short recurved beak included. Distr. Solomons (Bougainville); in Malesia: N. Borneo (Mt Kinabalu, Mt Nungkok), W. New Guinea (Idenburg R.). Ecol. In damp primary forests, along streams, on forested hillsides, 850—1400 m altitude. Note. In habit very similar to M. cuspidata, but the leaves not contracted into a petiole and without broad sheaths at the base. 16. Mapania enodis (MiQ.) CLARKE, Kew Bull. add. ser. 8 (1908) 53; UitTIEN, Rec. Trav. Bot. Neéerl. 33 (1936) 149. — Lepironia enodis MiQ. Sum. (1861) 263, 603; Illustr. (1870) 60, t. 21; Boeck. Linnaea 37 (1871) 141. — Lepironia foliosa Miq. Sum. (1861) 263, 603. — Pandanophyllum miquelianum Kurz, J. As. Soc. Beng. 38, ii (1869) 81; ScHEFF. Nat. Tijd. N.I. 34 (1874) 90. — M. longa [Rip-. J. Str. Br. R. As. Soc. n. 23 (1891) 14, nom. nud.] ex CLARKE, FI. Br. Ind. 6 (1894) 683, p.p.; J. Linn. Soc. Bot. 34 (1898) 95, p.p., quoad specim. Singap.; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105; Fl. Mal. Pen. 5 (1925) 173; non Camus, FI. Gén. I.-C. 7 (1912) 178 (quae est Carex vietnamica RAyYM.).— Pandanophyllum an- gustifolium KURZ ex CLARKE, FI. Br. Ind. 6 (1894) 683, nom. nud., in syn. — M. zeylanica (non CLARKE) RIDL. & WINKL. Bot. Jahrb. 44 (1910) 525; MerRrR. En. Born. (1921) 65, p.p., quoad WINKLER 2585, fide Urttien. — M. tumida Urrt1en, Gard. Bull. S. S. 10 (1939) 182. Leaves coriaceous, linear, with the midnerve prominent beneath and 2 of the lateral nerves promi- nent above, rather gradually attenuate towards the slender, triquetrous, aculeate-scabrous tip, remotely aculeate-scabrous on the margins and the upper part of the midnerve on the lower side, hardly narrowed towards the conduplicate base, septate-nodulose by numerous cross-nervules, glaucous, paler beneath, (45 cm—)1—-1)4 m by 1-2/4 cm; sheaths stramineous. Scapes lateral, rather robust, much shorter than the leaves, obtusely trigonous, thickened at the apex, smooth, glaucous, (10—)45—90 cm by 14-2 mm (up to 4mm at the top), the base clothed with some lanceolate, acute, striate sheaths, the upper sheaths widened upwards, with scarious brown margins, up to 15 cm. Inflorescence consisting of a single spikelet (very rarely | or 2 lateral spikelets in the axils of the lower glumes), conical, acute, 2—3/4(-4) by 1- 1/A(-2) cm. Glumes coriaceous, oblong-lanceolate, obtuse, greenish with narrow scarious brown margins, the lower empty ones many-nerved, 8—10 by 4-5 mm, the fertile upper ones less firm, oblong, less nerved, 8-9 by 244-3 mm. Flowers about as long as the glumes; outermost scales ciliate on the keel in the upper half. Anthers not seen. Style 3-cleft, sometimes some of them 2-cleft. Nut lageniform, subglobose in the centre, distinctly beaked, constricted towards the base and then conspicuously widened, truncate at the very base, shortly stipitate, with 3 longitudinal ribs, brown, 5—7 by 24-3 mm. Distr. Peninsular Thailand; in Malesia: Sumatra (Djambi, Palembang), Malay Peninsula (Johore, Singapore), W. Borneo. Ecol. In swampy forests, along streams, at low altitudes, sometimes dominant. Vern. Rumput sélingsing, M, térissi, Borneo. Notes. The type collection of Lepironia enodis MiQ. consists of fruiting scapes only. MIQUEL placed this species next to Lepironia articulata (RETZ.) DOMIN (= L mucronata L. C. RICH.) as he supposed it to be leafless. Immature complete specimens from the same locality were described as Lepironia foliosa Mig. In 1870, when aware of his mistake, MIQUEL united the two under the binomial Lepironia enodis, which choice must be followed. M. longa CLARKE was based on two collections, RIDLEY 169 from Singapore and Mor ey 1190 from Borneo (see J. Linn. Soc. Bot. 34, 1898, 95 and Ill. Cyp. 1909, t. 110). In the Flora of British India CLARKE placed the species in his section Pandano- phyllum, remarking that perhaps it might better be referred to sect. Halostemma. This remark is due to the fact that RIDLEy’s specimen is M. enodis (belong- ing in CLARKE’s sect. Pandanophyllum), whereas MOTLEY’s collection, from which the description was drawn up, belongs in his sect. Halostemma. 17. Mapania lorea UITTIEN, Rec. Trav. Bot. Néerl. 33 (1926) 150. — M. valida Riwu. J. Str. Br. R. As. Soc. n. 44 (1905) 205, p.p., quoad RIDLEY 1714; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 104, p.p.; Fl. Mal. Pen. 5 (1925) 172, p.p. — M. tenuiscapa (non CLARKE) RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105, p.p., quoad RIDLEY 11501; Fl. Mal. Pen. 5 (1925) 173, p.p. — M. zeylanica (non CLARKE) MerRR. En. Born. (1921) 65, p.p., quoad Nat. Coll. 350. Leaves thickly coriaceous, linear, with the midnerve prominent beneath, and several indistinct lateral nerves, finely transversely wrinkled by very numerous cross-nervules connecting the nerves, slightly narrowed towards the conduplicate base, gradually attenuate to the very slender, triquetrous, aculeate- scabrous tip, densely serrate-scabrous throughout the margins and on the upper half of the keel on the lower side, also aculeate-scabrous on the upper surface in the upper part, glaucous, up to 1m by 1—2 cm; sheathing base dark castaneous, more rarely shining light brown. Scapes lateral, from the axils of the lower leaves or below the leaves, terete or com- pressed, slightly thickened at the apex, smooth, 5-30 cm by 1—2 mm, the base clothed with some light brown scales up to 5 cm. Inflorescence always a single spikelet, subglobose to cylindrical, terete, very obtuse when in fruit, 1-4 by 1-1/4 cm. Glumes coriaceous, ovate, obtuse, strongly many-nerved, pale brown, with scarious margins, densely purplish puncticulate in the upper half, 7-9 mm long, the upper ones less firm, oblong, several-nerved. Flowers about as long as the glumes; keel of outermost scales sparingly ciliolate. Anthers c. 4mm. Style 3-cleft. Nut ellipsoid or pyriform, shortly stipitate, fulvous, 5-6 by 244-344 mm, the short recurved beak included. Distr. Malesia: Sumatra (Siak R. region and P. Rupat), Enggano, Malay Peninsula (Perak, Johore, Singapore), Borneo (Sarawak, once collected; E. Borneo: Long Tubus; N. Borneo: Kilagan). Ecol. In swamps and swampy forests at low altitudes, in P. Enggano characteristic of the open forest on tertiary red soil. Vern. Sélingsing, séling sélingan, prupd, M. Notes. Distinguishable from M. tenuiscapa by the much thicker, leathery, transversely wrinkled leaves with only one distinct nerve, the more robust scapes, and the leaves scabrous also on the upper side just 1974] CYPERACEAE (Kern) 481 below the tail. In appearance it is similar to M. kurzii, but this species has always a capitate inflo- rescence, with several dark reddish spikelets, almost ribbed glumes, smooth upper side of the leaves, and distinctly winged outermost flower-scales. The base of the leaf is usually dark brown to almost black, but in the Enggano specimens of a much lighter colour. 18. Mapania spadicea UitTieN, Rec. Trav. Bot. Néerl. 33 (1936) 147. Leaves very densely crowded, strikingly 3-ranked, the rows prominent and fan-like at the base (almost as in M. graminea), coriaceous, linear, with the midnerve prominent beneath and 2 whitish lateral nerves somewhat prominent above, slightly narrowed towards the conduplicate base, gradually narrowed into a long, flagelliform, triquetrous, very scabrous acumen, densely serrate-scabrous throughout the margins and on the midnerve beneath in the upper part, glaucous, very shining above, dull beneath, up to 170 by 9-14 mm; sheathing bases shining dark brown to blackish. Scapes lateral, from the caudex below the leaves, stout, subterete or compressed, scabrous or (at length?) almost smooth, (2—)10— 30 cm by 3—4 mm, the base clothed with stiff, acute, dark brown scales. Inflorescence usually a single spikelet, sometimes consisting of 2—3 fasciculately arranged spikelets (both types occurring on the same plant). Spikelets lanceolate or narrowly lanceolate in outline, terete, acute, 2/44-5cm by 10-15 mm. Lower glumes coriaceous, lanceolate, acutish, cas- taneous, with scarious margins, many-nerved, flower-bearing glumes lanceolate, thinner and less nerved, 12—15 mm long. Flowers about as long as the glumes; keel of outermost scales ciliate in the upper part. Anthers c. 8mm. Style 3-cleft. Nut ellipsoid, subtrigonous, acuminate, shortly stipitate (stipe c. 1 mm), brown, 4—5 by 2/4 mm. Distr. Malesia: Borneo (W. Borneo: Sanggau; E. Borneo: W. Kutei). Ecol. In primary forests, along streams, at low altitudes. Vern. Méringsing, W. Kutei. 19. Mapania tenuiscapa CLARKE, Fl. Br. Ind. 6 (1894) 683; J. Linn. Soc. Bot. 34 (1898) 95; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 105, p.p.; FI. Mal. Pen. 5 (1925) 173, p.p.; UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 150.— M. versicolor BECc. For. Born. (1902) 517, nom. nud.; MeRR. En. Born. (1921) 65.— M. archboldii Uittien, J. Arn. Arb. 20 (1939) 214; S. T. BLAKE, ibid. 28 (1947) 211. Leaves densely crowded, subcoriaceous, linear, with the midnerve prominent beneath, somewhat plicate lengthwise along several distinct lateral nerves, slightly narrowed towards the conduplicate base, very gradually narrowed into a long, flagelliform, triquetrous, aculeate-scabrous acumen, densely aculeate-scabrous throughout the margins and on the midnerve beneath, pale or bright green, up to 1m by 7-10 mm; sheaths ferrugineous to brown. Scapes lateral, from the caudex below the leaves, very slender, much shorter than the leaves, striate, 7-30cm by 4-1 mm, up to 2mm thick at the somewhat incrassate apex, the base clothed with some stramineous or brown, loose scales up to 5 cm long. Inflorescence consisting of a single spikelet, ovoid when in fruit, terete, acute, 10-15 by 5—6(—10 when in fruit)mm. Glumes subcoriaceous, with scarious margins, ovate-lanceolate, many-nerved, 8-10 mm long, the upper ones less firm, oblong, obtuse or lacerate, brown-puncticulate at the top, 5—7-nerved. Flowers about as long as the glumes; outermost scales ciliate on the hardly or narrowly winged keel. Style 3-cleft. Nut fusiform or pyriform, terete, shortly stipitate (stipe 4-1 mm), greyish or blackish brown, 4-7 by 2-3 mm, the short recurved beak included. Distr. Thailand, Indo-China; in Malesia: Malay Peninsula (Kedah, Perak, Pahang, Malacca, Johore, P. Langkawi), Borneo (Sarawak), New Guinea (W. New Guinea: Japen-Biak; Papua: Palmer River). Ecol. In primary forests, on river banks, locally abundant, at low altitudes. Vern. Sendayan, Mal. Pen. Notes. CLARKE says that the spikelets are “more or less compound (i.e. basal partial spikes containing more than one spikelet)’’. I always find them simple. In the New Guinea specimens (M. archboldii UrTien) the fruits are 6—7 mm long, in the other specimens examined 4—6 mm. Otherwise M. arch- boldii agrees in details with M. tenuiscapa. 20. Mapania sessilis MerR. J. Str. Br. R. As. Soc. n. 85 (1922) 156. — M. longiflora var. sessilis UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 146. Leaves coriaceous, linear, slightly narrowed below, gradually narrowed upward to the very slender, triquetrous, aculeate-scabrous tip, with the midnerve prominent beneath and 2 lateral nerves somewhat prominent above, the margins serrulate-scabrous throughout except near the base, and the keel scabrous on the lower surface in the upper part, glaucous, up to 14m by 1-14 cm; sheathing bases shining castaneous. Scapes lateral, along the caudex below the leaves, very short and inflorescences therefore practically sessile. Inflorescence always a single spikelet, lanceolate in outline, acute, tri- quetrous, up to 3-4 by 1-1/4 cm. Sterile glumes coriaceous, ovate to oblong-ovate, acute, sharply keeled at least in the upper half, shining castaneous, 2-3 by 1-14 cm; fertile glumes membranous, oblong, white, 3—5-nerved. Flowers 15-30 mm long; outermost scales with a distinct, pellucid, white wing 4-24 mm wide, sparsely and minutely ciliolate at the top. Anthers 7-15 mm. Nut oblong-ellipsoid, terete or slightly compressed, stipitate (stipe 1-2 mm), brown, c. 10 by 3/4 mm (the long, straight, conical, 3—4 mm long beak included). Distr. Malesia: Sumatra (E. Coast Res.), Banka, N. Borneo (near Sandakan). Ecol. In damp forests, along small streams, at low altitudes. Vern. Séding rimba, Banka. Notes. The number of stigmas is unknown. Only a few times collected, probably owing to the fact that the spikelets, all sessile at the very base of the plant, are easily overlooked. UnTiEn reduced this species to a variety of M. longiflora scarcely deserving recognition. However, it differs from the latter species not only by the lack of a scape, but also by the different structure of the inflorescence, the castaneous base of the leaves, the keeled outer glumes, the winged outermost flower- scales, and the larger nuts. 482 FLORA MALESIANA [ser. I, voli F 21. Mapania graminea UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 147, incl. var. stipitata UITTIEN. Rhizome woody, erect or ascending, 1-1/4 cm thick. Leaves very densely crowded, strikingly 3- ranked (the rows very prominent and fan-like at the base, the base of the foliage in cross-section therefore Y-shaped), coriaceous, linear, with the midnerve prominent beneath, slightly narrowed towards the conduplicate base, very gradually narrowed into a long, flagelliform, triquetrous, aculeate-scabrous acu- men, densely serrate-scabrous throughout the mar- gins and on the midnerve beneath in the upper part. also aculeate-scabrous on the upper surface in the upper part, glaucous, up to 144m by 5-20 mm; margins of the sheaths dark brown or purplish. Scapes lateral, from the caudex below the leaves, very variable in length, subtrigonous or compressed, striate, smooth, 1-15 cm by 1-2/4 mm, the base (when very short wholly) covered by some ovate to lanceolate, pale brown sheaths up to 4 cm long. Inflo- rescence consisting of a single spikelet, rarely of 2—3 fasciculately arranged spikelets. Spike/et ellipsoid or lanceolate in outline, trigonous when young, acute, 2-3 by 1-1/4 cm. Lower (sterile) glumes coriaceous, with scarious margins, lanceolate, acutish, many- nerved, fertile ones thinner, oblong, less nerved, 11-14 mm long. Flowers about as long as the glumes; outermost scales ciliate on the keel in the upper half. Anthers 6-6/4 mm. Style 3-cleft. Nut obovoid- pyriform, subtrigonous, shortly beaked, shortly stipitate (stipe 4-74 mm), brown or greyish brown, 4-5 by 244 mm. Distr. Malesia: Borneo (Sungei magne, Bukit Kasian, Nunukan; E. Borneo: W. Kutei, Mt Palimasan; N. Borneo). Ecol. In primary forests at low altitudes. Note. Almost sessile and distinctly pedunculate spikelets sometimes occur on the same plant; it seems superfluous to distinguish specimens with elongated scapes as var. stipitata. 22. Mapania micropandanus Hoitr. Gard. Bull. Sing. 11 (1947) 294. Rhizome /4-1 cm thick, woody, erect or finally becoming almost horizontal, the older part dying off and then the leafy end standing upon long, greyish brown stilt roots 15—30cm above the ground. Leaves herbaceous, linear, pale green, with denti- culate-aculeate margins throughout their length, 20-40 cm by 8-13mm, the apex rather abruptly narrowed into the acute tip, not caudate, the base slightly narrowed, conduplicate, the midrib prom- inent on the underside and 2 of the lateral nerves prominent on the upper side, all 3 main nerves scabrous on their raised surface almost from base to apex of the leaf. Scapes lateral, hidden in the leaf- axils, very short, the solitary spikelet almost sessile. Spikelet lanceolate in outline, trigonous, acute, 15—20 by 5-6 mm. Glumes lanceolate, many-nerved, almost ribbed when dry, acute, pale, with scarious margins, c. 15mm long, the lower sterile ones shorter and broader, ovate to broadly ovate, firmer, c. 10mm long. Flowers as long as the subtending glume (c. 15 mm), linear; outermost scales narrowly winged on the keel, sparsely ciliate at the top only. Stamens 3; anthers linear, c. 8 mm. Style 3-fid. Nut unknown. Distr. Malesia: Malay Peninsula (SE. Johore: Sungai Sedili basin; Pulau Tioman). Ecol. In dry Dryobalanops forests and in the drier parts of freshwater swampy forests, at low altitudes. Note. This remarkable species, like a miniature Pandanus, is distinct from all other species by the non-caudate leaves toothed on the margins and throughout the three main veins. Its nearest ally may be M. graminea UITTIEN from Borneo, in which the spikelets are often also subsessile and trigonous, the texture and nervature of the glumes very similar, and the outermost flower scales also winged. 23. Mapania maschalina KERN, Blumea 12 (1963) 23, f. 1. — Fig. 16. Rhizome woody, ascendent. Stems elongated, erect, sustained by 2-3 mm thick terete roots, densely leafy, c. 50cm tall. Leaves coriaceous, narrowly linear, rigid, dull green, glaucescent, stra- mineous towards the conduplicate base, gradually narrowed into a triquetrous, very scabrous tail, up to 50 cm long, 5—7 mm wide, the upper part densely serrulate-scabrous on the margins and the prominent midnerve beneath, the lateral nerves scarcely distinct. Scapes lateral, very short, hidden in the axils of the cauline leaves, each with a solitary spikelet. Spikelets subsessile, ovoid, acute, c. 15 by 7-10 mm. Sterile glumes coriaceous, ovate, acute, many-nerved, fuscous, with scarious margins, 8—9 mm long; fertile glumes lanceolate, less firm, 9-10 mm long. Flowers linear, 8-10 mm long; scales membranous, the transversal 2 ciliolate on the keel. Anthers linear, 5—6 mm long. Style 3-fid. Nut subpyriform, narrowed into a short, slightly curved beak, shortly stipitate (stipe 14 mm), rugulose, greyish brown, 6 by 3 mm (beak included). Distr. Malesia: N. Borneo (Distr. Sandakan: Malubok Kinabatangan). Only a single collection known of this remarkable species. Ecol. In swamp forest. 24. Mapania monostachya UITTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 194, f. 1; RicHaRDs, J. Ecol. 24 (1936) 15. Leaves subcoriaceous, linear, with the midnerve very prominent on the underside and 2 of the lateral nerves more or less prominent above, serrulate- scabrous on margins and midnerve at least in the upper part, the apex gradually narrowed into a filiform, triquetrous, aculeate-scabrous acumen, hardly narrowed at the conduplicate base, brilliant metallic blue-green, 20—40 cm by 2—4 mm; sheaths stramineous. Scapes lateral, somewhat angular, very slender, much shorter than the leaves, (4—)8—15 cm by 4-74 mm, the base clothed with some greenish or stramineous, striate, up to 15 mm long scales. Inflorescence consisting of a single spikelet, 6-10 by 2-3 mm. Bracts 2-3, oblong-lanceolate, obtusish, many-nerved, densely fuscous-puncticulate, 5—7 mm long, the lowest 1-3 mm distant. Glumes hardly differing from the bracts but somewhat narrower, with less conspicuous and less numerous nerves. Flowers 4—6mm long; outermost scales sparsely ciliolate on the keel in the upper part. Stamens 3; anthers linear, white, 2-3 mm. Sty/e 2- or 3-fid. Nut pyriform, terete, narrowed at the base, acuminate at the apex, dusky brown, darker spotted, slightly rugulose, 24-24% by 114-1)4 mm. 1974] 484 FLORA MALESIANA [ser. I, vol. 7° Distr. Malesia: Borneo (Sarawak: Upper Rejang R., Mt Dulit; N. Borneo: Distr. Temburong, Kuala Belalong; W. Borneo: Sungei Simun and Sanggau). Ecol. In primary forests, along streams, from low altitudes up to 1100 m. Note. The smallest species of the genus, very similar in habit to Paramapania gracillima from the Philippines (see p. 488). Readily distinguishable from this by the longer flowers and glumes, the number of stamens, the linear much longer anthers, the flower- scales not spinulose on the keel, and the different shape of the nut. 25. Mapania baccifera CLARKE, Kew Bull. add. ser. 8 (1908) 53; Ill. Cyp. (1909) t. 115, f. 8-9; UiTTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 278, 279: S. T. BLAKE, J. Arn. Arb. 28 (1947) 211.— M. radulosa RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 245. — M. papuana RIDL. I.c. 246; UirTien, Rec. Trav. Bot. Néerl. 33 (1936) 146, 151, 278, 279; S. T. BLAKE, J. Arn. Arb. 28 (1947) 210. — M. grandiceps (non KUxK.) OHWI1, Bot. Mag. Tokyo 56 (1942) 211. Leaves coriaceous, linear, slightly narrowed to- wards the conduplicate base, gradually to rather suddenly narrowed upward to the slender, triquet- rous, aculeate-scabrous tip, with the midnerve prominent beneath and several rather prominent lateral nerves, the margins finely scabrous to almost smooth and the midnerve scabrous on the lower surface in the upper part, glaucous, up to 1/4 m by c. 1)4-4cm:; sheathing bases stramineous. Scapes lateral, from the axils of the lower leaves or below the leaves, trigonous to almost terete, incrassate at the top, scabrous by very short, stiff hairs, at length glabrescent, up to 60(—90?) cm by 1/4-3 mm. Inflo- rescence either consisting of a single spikelet or capitate, with up to 4 spikelets surrounded by a few coriaceous, lanceolate, acute, many-nerved, up to 3cm long bracts. Spikelets oblong-ovoid, acute, terete, 2-4 cm long. Lower glumes coriaceous, ovate- lanceolate, acute, many-nerved, with scarious margins, 10-15 mm long: flower-bearing glumes thinner, oblong, several-nerved. Flowers 10-15 mm long: outermost scales ciliate on the keel. Style 3-cleft. Nut obovoid-pyriform, terete, berry-like by the succulent exocarp, stipitate (stipe 1—2 mm), shortly beaked, strongly wrinkled when dry, yellow or orange, 6-7 by 3/4—4 mm (beak included). Distr. Solomons; in Malesia: ?Moluccas (Moro- tai); W. New Guinea (Bume R. near Nabire, Mt Carstensz, Idenburg R.), Japen, Biak. Ecol. In dense rain-forest, in mossy forest under- growth, at low altitudes, up to 900 m. Note. Very insufficiently known. Provisionally I have united all Malesian Mapaniae with succulent yellowish fruits, for the material available at the present is inadequate for critical study. The type collections of M. baccifera, M. papuana, and M. radulosa consist of leaf fragments and a single inflorescence or a few detached ones. In the type of M. baccifera from the Solomons the inflorescence contains 4 overmature spikelets surrounded by some involucral bracts, in the specimens collected in those islands by Brass and referred to M. baccifera by both UitTren and BLAKE the inflorescence is a single spikelet. The type of M. papuana is still more fragmentary than that of M. baccifera. The leaves are c. 2cm wide, the inflorescence a very young spikelet. In the specimens UITTIEN referred to M. papuana the leaves are much wider and in BRAss 13079 (referred to M. papuana by BLAKE) the inflo- rescence is apparently compound. The latter col- lection is very similar to the type of M. radulosa, the original description of which is inaccurate: the leaves are 2 (not 1-1/4) cm wide and the involucral bracts 3 (not 8) cm long. Doubtful Mapania alpina Ev. Leafl. Philip. Bot. 10 (1938) 3535. — Based on ELmMer 14864 from Irosin (Mt Bulusan), Luzon, in the woods at 3500 ft altitude. — I have not seen this collection; judging from the description it might belong to M. borneensis MERR. — MERRILL, En. Philip. 1 (1923) 132, referred it to M. cuspidata (M1Q.) UITTIEN (‘M. humilis’), but ELMER properly observes that it is distinct from this species. 6. PARAMAPANIA UiTTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 186; ibid. 33 (1936) 141; KERN, Blumea 9 (1958) 215. — Fig. 17. Perennial herbs with short, woody, obliquely erect rhizome covered with the fibrous remains of decayed leaf-sheaths. Leaves 3—ranked, equitant, conduplicate at the base, subcoriaceous, either linear, gradually narrowed into a tail-like, aculeate-scabrous acumen, or, more rarely, narrowly oblong, petiolate, abruptly acuminate and caudate; midnerve prominent on the under side; ligule absent. Flowering scapes slender, usually much shorter than the leaves, arising laterally, either from the axils of the lower leaves, or below the leaves, with some basal bladeless sheaths, not rarely also with 1—3 bladeless sheaths higher up. Inflorescence terminal, corymbose, capituliform, or consisting of a single spikelet; bracts very short. Spikelets small, (narrowly) ellipsoid, ovoid, or obovoid, terete, many-flowered, usually brown. Glumes spirally imbricate, subcoriaceous, brown, with paler midnerve, nerveless sides, and hyaline margins, some lower ones empty. Flowers compressed, bisexual; hypogynous scales 1974] CYPERACEAE (Kern) 485 Fig. 17. Paramapania parvibractea (CLARKE) UITTIEN. a. Habit, x 4, b. floral diagram, c. spikelet, x 5, d. glume, e. flower with two transverse ciliate scales, inner scales each with apex of filament, and stigmas, f. three inner scales, g. pistil, 4. mature nut, all x 15 (a—h SULIT 6308). 486 FLORA MALESIANA [ser. I, vol. 7° (squamellae) usually 5, the lowest 2 transversal, opposite, free, boat-shaped, coarsely brown-spinulose on the acute keel, the 3rd (anticous) scale as a rule absent, the upper 3 concave, empty; rarely scales 6, the anticous one present, but smaller than the upper 3. Stamens 2, very rarely 3, in the axils of the lower scales; anthers oblong-linear, scarcely appendaged. Style continuous with the ovary; stigmas 3, rarely 4. Nut subterete or angular, turgid in the middle, narrowed towards the base, the apex with a cupuliform, conical-pyramidal, or elongated beak often confluent with the nut proper; exocarp hard, thin. Distr. Small genus (7 spp.), almost entirely confined to Malesia; P. parvibractea extending to the western Pacific. Ecol. All the species inhabit very damp rain-forests at low or medium altitudes. Notes. The species of Paramapania are mutually closely related, and form a natural group. Nevertheless it is difficult to characterize the genus in a satisfactory way. In structure the flowers are very similar to those of Mapania and Thoracostachyum, but generally more reduced, as the third hypogynous scale is usually lacking and the number of stamens is reduced to 2. When the third scale is present it is smaller than the upper ones, and as a rule empty. Occasionally there may be a stamen in its axil, but if so that stamen is less developed than the other two. The relatively coarse, brown spinules on the keel of the two lowest (lateral) scales are characteristic of Paramapania; the corresponding scales in Mapania and Thoracostachyum are softly ciliate on the back. Paramapania can readily be distinguished from Mapania sect. Cephaloscirpus and Thoracostachyum by the lateral, leafless scapes. It is, however, difficult or practically impossible to distinguish between Para- mapania and the scapigerous species of Mapania and Hypolytrum by their habit alone. Especially some African Hypolytrum spp. (e.g. H. nudicaule from Madagascar) bear a close resemblance to P. parvibractea, but in them the number of scales is reduced to 2. As a rule the spikelets and flowers in Mapania are much larger than those in Paramapania, and the fruits drupaceous by the thick, fleshy or spongy exocarp. Corymbose inflorescences are not found in Mapania. Specific delimitation is difficult as the greater part of the species (P. gracillima, P. flaccida, P. rostrata, P. longirostris, and P. simplex) are only known from a few collections, and the more widely distributed species are highly variable. Several of the characters used by UITTIEN are apparently taxonomically less important than he supposed them to be. KEY TO THE SPECIES 1. Flowering scapes strongly compressed, ancipitous. Leaves abruptly acuminate, shortly caudate, the margins scabrous only at the very top J oe Me Pate, eae fe Demis, sie 1. Flowering scapes terete or trigonous. 2. Stigmas 4, or in a few flowers 3. Inflorescence consisting of a single spikelet 2. Stigmas 3, or in a few flowers 4. 3. Nut lageniform, with a hexagonal, conical, obtuse, densely verruculose beak, 2mm long. Glumes distinctly spinulose-ciliolate on the upper pe Inflorescence dense, head-like, or reduced to a single spikelet 2. P. radians 3. Nut with a triquetrous, acute, smooth beak, or beak indistinct. Glumes not or scarcely (not spinulose) ciliolate. 4. Nut indistinctly beaked, 14-274 mm long. Inflorescence usually corymbose, bb well-developed rays and many to numerous spikelets . : . I. P. parvibractea 4. Nut with a distinct, triquetrous, 14-3 mm long beak. Inflorescence either heat of a single spikelet or head-like, with up to 5 spikelets. 5. Leaves abruptly acuminate, caudate, 8-12 mm wide, their margins scabrous vi at the Bey top, the base narrowed into a 1-5 cm long petiole . . P. simplex 5. Leaves very gradually narrowed into a long point, 4-8 mm wide, scabrous on the Be at least in the upper half, conduplicate at the base, but not distinctly petioled. 6. Nuts stellately spreading, their upper part dark brown, suddenly narrowed into the pale, obconical lower part. Beak of the nut firm, straight or but slightly curved, hence spikelets echinate. 5. P. rostrata 6. Nuts obliquely erect, brown, their body ellipsoid, rather ee attenuate at both ends. Beak of the nut slender, curved; spikelets not echinate ‘ 6. P. longirostris 3. P. gracillima 1. Paramapania parvibractea (CLARKE) UITTIEN, Rec. Bull. add. ser. 8 (1908) 51; VALCK. Sur. Nova Trav. Bot. Néerl. 33 (1936) 143; S. T. BLake, J. Arn. Arb. 28 (1947) 209; KERN, Blumea 9 (1958) 217, excl. pl. Philip. — Hypolytrum parvibractea CLARKE, Kew Bull. (1899) 114. — Mapania montana LAuT. & K. Scu. in K. Sch. & Laut. Fl. Schutzgeb. (1900) 189. — Hypolytrum parvibracteatum CLARKE, Kew Guinea 8 (1912) 709; RipL. Trans. Linn. Soc. II, Bot. 9 (1916) 243.— Mapania luchanensis ELMER, Leafl. Philip. Bot. 2 (1909) 573. — Hypolytrum parvibracteatum var. quadriglumatum VALCK. SUR. Nova Guinea 8 (1912) 709, t. 116, nom. inval. — Hypolytrum quadriglumatum VALCK. Sur. l.c., nom. 1974] CYPERACEAE (Kern) 487 inval. — Thoracostachyum montanum VALCK. SUR. I.c. 710; KUK. Bot. Jahrb. 59 (1924) 54; OHw1, Bot. Mag. Tokyo 56 (1942) 209.— Thoracostachyum lucbanense KUx. ex Mere. Philip. J. Sc. 11 (1916) Bot. 258; En. Philip. 1 (1923) 132, p.p. — Thoraco- stachyum longistylum KOK. Bot. Jahrb. 58 (1924) 54, incl. f. parvum KtUK.— Hypolytrum radians RIDL. Fl. Mal. Pen. 5 (1925) 170. p.p. (quoad specim. Johor.), non Mapania radians CLARKE. — P. luc- banensis UiTTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 190, quoad basion. — P. johorensis UirTieEN, l.c. 191; ibid. 33 (1936) 141. — P. amboinensis UITTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 191, f. 4. — P. montana UnTien, I.c. 200, nom. provis.— P. longistyla UiiTien, Rec. Trav. Bot. Neéerl. 33 (1936) 141. — Thoracostachyum parvibractea KUx. Bot. Jahrb. 69 (1938) 261.— ? Thoracostachyum macilentulum Ouwil, Bot. Mag. Tokyo 56 (1942) 210, e descr. — Mapania parvibractea KoyAMA, Micronesica | (1964) 66. — Fig. 17. Forming compact, 3-angled clumps. Leaves linear, flat or somewhat folded lengthwise, very gradually or (in broad-leaved specimens) rather suddenly narrowed into a subfiliform, scabrous point, antrorsely aculeate- scabrous on the margins and on the midnerve beneath in the upper part, up to 70(-100) by %— 1/4(-2) cm, the base strongly conduplicate, with reddish brown or purplish, scarious margins. Scapes trigonous, smooth, usually reddish brown, up to 45 cm by 1-2 mm, in or about the middle with 1—2 tubular, 3—5 cm long sheaths split up at the top, the base clothed with some loose, lanceolate, fuscous sheaths. Inflorescence corymbiform, simple or com- pound, with few to many spikelets, rarely in some of the scapes reduced to a single spikelet, up to 3/4 by 7cm, the lower branches divaricate, up to 3 cm long, the upper ones suberect, very short. Fruiting spikelets subglobose, ellipsoid, or oblong-ovoid, 3—6(—10) by 3—4 mm. Glumes ovate, obtuse, minutely ciliolate on the upper margin, 2—3 by i4- 2mm. Flowers 134-2(-3)mm_ long; hypogynous scales 5(—6), the upper 3(—4) lanceolate or narrowly ovate. Stamens 2, very rarely 3; anthers oblong-linear, A- 1mm. Style triquetrous, more or less scabrid; stigmas 3, or in a few flowers 4. Nut obliquely erect, fusiform or subpyriform, subterete or slightly com- pressed, acuminate at both ends, shortly stipitate, rounded at the apex, brown or greyish black, with 3 longitudinal pale stripes, 1/4-2)4 by 4-1/4 mm; beak indistinct, confluent with the nut proper. Distr. W. Carolines, Solomons, Samoa, Fiji; in Malesia: W. Sumatra, Lingga Archipelago, Malay Peninsula (Kelantan, Johore), Borneo (Distr. Sanda- kan), Philippines (Luzon, Samar, Mindanao), Celebes (E. Peninsula), Moluccas (Talaud Is., Morotai, Amboina, Ceram, P. Gebe), New Guinea and adjacent islands. Ecol. In rain-forests, along forest-trails, on seep- ages, in the lowlands and on the lower mountain slopes, up to 1200 m. Vern. Philip.: pavong-payong, tingol, Sub. (these names may refer also to P. rostrata); New Guinea: masibu, Kutubu lang. Notes. The type collection (GIULIANETTI s.n., K) is said to be collected on Mt Scratchley at 12.200 ft, i.e. in the subalpine zone. See, however, VAN STEENIS, Fl. Mal. I, 1 (1950) xxi. The type collection of P. lucbanensis (ELM.) UrtTIEN is Ev_mer 9116 from Lucban, prov. of Tayabas, Luzon (E). It belongs undoubtedly to P. parvibractea. UITTIEN, who did not see the type, took Ramos 23642, distributed as Thoracostachyum luc- banense, but which belongs to P. rostrata, for P. lucbanensis. See also under P. rostrata. Originally UrrTIEN distinguished between P. johorensis from the Malay Peninsula and P. parvi- bractea (= P. amboinensis Uirr.) by the number of flower-scales (6 in P. johorensis, 5 in P. parvibractea), and the smaller number of spikelets to the inflo- rescence in the former. Later on he referred specimens from P. Lingga, with 5 scales and inflorescences bearing up to 20 spikelets, also to P. johorensis on account of the globose, stipitate nut. As I fail to see any essential difference in the nuts, and the number of scales is variable also in the eastern part of the area, I refer P. johorensis to the synonymy of P. parvi- bractea. More deviating from typical P. parvibractea is P. longistyla (KUK.) UlTTIEN, only known from two New Guinea collections: LEDERMANN 19586 (L, P!) and 19728 (f. parva KUx., not seen). The leaves are up to 2cm wide, rather suddenly acuminate, the flowers about 3 mm long, with 6 scales and 3 stamens, the style is distinctly scabrid on the angles, and the nut somewhat larger. In P. parvibractea, however, the width of the leaves is very variable, 3mm long flowers with 6 scales and 3 stamens are also found in narrow-leaved specimens, and the style is more or less scabrid. Provisionally I consider P. longistyla a local race of P. parvibractea, the polymorphism of which needs further study. 2. Paramapania radians (CLARKE) UITTIEN, Rec. Tiav. Bot. Néerl. 32 (1935) 188, f. 1; 200; KERN, Blumea 9 (1958) 217, p.p. (quoad pl. Born.). — Mapania radians CLARKE in Ridl. J. Str. Br. R. As. Soc. n. 46 (1906) 226; Kew Bull. add. ser. 8 (1908) 53; Merr. En. Born. (1921) 65. — Hypolytrum radians RipL. Fl. Mal. Pen. 5 (1925) 170, excl. specim. Malacc. Near to the preceding species. Leaves linear, flat or somewhat folded lengthwise, very gradually narrowed into a subfiliform, aculeate-scabrous point, antrorsely scabrous on the margins and on the midnerve beneath in the upper part, pale green or glaucescent, up to 80cm by 3-8mm; sheaths conduplicate, fuscescent, with reddish brown, scar- ious margins. Scapes trigonous or subterete, smooth, often dark-puncticulate, 4-16 cm by 4-1 mm, the base clothed with a few lanceolate, fuscous sheaths, otherwise without sheaths, nodeless. Inflorescence simple, very dense, head-like, subglobose, with 2-10 spikelets, up to 1/4 cm across, not rarely in some of the scapes reduced to a single spikelet. Fruiting spikelets ovoid-ellipsoid, brown, 7-10 by 4-5 mm. Glumes ovate, obtuse, spinulose- ciliolate on the upper margin, 2 by 14-14 mm. Flowers 114-124 mm long; hypogynous scales 5, the upper 3 lanceolate or narrowly ovate. Stamens 2; anthers 14-% mm long. Stigmas 3, rarely in a few flowers 4. Nuts obliquely erect, lageniform, terete or obscurely trigonous, with 6 longitudinal ribs, smooth, shining, dark brown, 2 by 34-1 mm (beak included); beak clearly distinct from the nut proper, pyramidal- conical, obtuse, hexagonal by 6 ribs, densely verru- culose, c. 4 mm long. 488 FLORA MALESIANA [ser. I, volar Distr. Malesia: Borneo (Sarawak; Brunei; Sabah; NE. Borneo: Nunukan near Tarakan; E. Borneo: W. Kutei). Ecol. In primary Dipterocarpaceous and Agathis forests, up to 600 m. 3. Paramapania gracillima (KUK. & MerRR.) UITTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 190, f. 3. — Mapania gracillima KOK. & MerR. Philip. J. Sc. 9 (1914) Bot. 267; MerrR. En. Philip. 1 (1932) 132. Leaves linear, flat, gradually attenuate at both ends, glaucescent, aculeate-scabrous on the margins and especially at the filiform, triquetrous top, up to 30 cm by (2—)4—5 mm. Scapes very slender, filiform, terete or obtusangular, slightly asperous-furfura- ceous, 5-10 cm by 4—/)4 mn, in the middle often with a single, tubular, 4-2 cm long sheath split up at the top, the base clothed with some lanceolate, fuscous, bladeless sheaths. Inflorescence consisting of a single spikelet. Spikelet small, at first linear- ellipsoid, when in fruit obovoid or ellipsoid, 4—7 by 3-4mm. Glumes ovate, obtuse, 2 by les mm. Flowers c. 134 mm long; hypogynous scales (5—)6, the anticous one present in the majority of flowers, but smaller than the upper scales. Stamens 2; anthers elliptic-oblong, 4-74 mm long. Stigmas 4 (or in a few flowers 3). Nut fusiform-sublageniform, narrowed at both ends, subterete, slightly angular by 4 longi- tudinal ribs, shortly stipitate, with short, obtuse beak scarcely distinct from the nut proper, smooth, stramineous to brown, 2 by 1 mm. Distr. Malesia: Philippines (Luzon: Prov. of Tayabas, Mt Dingalan; Mindanao, Distr. Zam- boanga). Ecol. On forested ridges, at about 1100 m. 4. Paramapania flaccida UiTTIEN, Rec. Trav. Bot. Neéerl. 33 (1936) 142. Leaves linear, flat, rather abruptly acuminate, shortly caudate (the aculeate-scabrous tail 1—2 cm long), gradually narrowed towards the strongly conduplicate base, scabrous, on the margins only at the very top, pale green, fuscescent at the base, 10—20 cm by 7-14 mm. Scapes strongly compressed, ancipitous, scabrid on the edges only at the top, 5-18 cm by 1mm, the base clothed with some lanceolate, fuscous sheaths. Inflorescence consisting of 1(—3) spikelets; branches when present short, 74-1 cm long. Spikelets oblong-obovoid, narrowed at the base, brown, 7-10 by 4-6 mm when in fruit. Glumes oblong, obtuse, brown, 3-3/4 by 1/4 mm. Flowers oblong, c. 3 mm long; hypogynous scales 6, the anticous one small, 2 mm long, the upper ones 3 mm. Stamens 2. Style triquetrous, smooth; stigmas 3. Nut ellipsoid-fusiform, obtusely trigonous, slightly dorsiventrally compressed, acuminate at both ends, dark brown, 224-3 by 14-14 mm, with obtuse beak almost confluent with the nut proper. Distr. Malesia: W. New Guinea (along Rouffaer River). Ecol. In lowland rain-forest, 250 m. Notes. Only known from the type collection, DOCTERS VAN LEEUWEN 10413. In appearance very similar to P. simplex, but distinguishable by the ancipitous scapes and the almost erostrate nuts. The irrelevant specific epithet was obviously intended to refer to the compressed (Dutch ‘vlakke’), but not flaccid scapes. 5. Paramapania rostrata UITTIEN, Rec. Trav. Bot. Neéerl. 32 (1936) 189.— Mapania rostrata ELM., nomen in sched. ad Phil. Isl. Pl. 16150. — P. longi- rostris UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 142, p.p. (quoad specim. Philip.). — Thoracostachyum luc- banense MeErRR. En. Philip. 1 (1923) 132, p.p., non Mapania lucbanensis ELM. — P. lucbanensis UITTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 190, quoad specim. cit., non Mapania lucbanensis ELM. — P. radians (non UITTIEN) KERN, Blumea 9 (1958) 216, 217, p.p. (quoad pl. Philip.). Leaves linear, flat or with revolute margins, very gradually narrowed into a flagelliform, aculeate- scabrous point, also aculeate-scabrous on the margins and the midnerve beneath, glaucescent, up to 65 cm by 5—8 mm, the base strongly conduplicate, pinkish or purplish. Scapes filiform, minutely scabrid, 3- 20cm by 4-4 mm, the base clothed with a few, brown, bladeless or short-bladed sheaths, the longer ones also witha sheath about the middle. Inflorescence simple, often consisting of a single spikelet, not rarely head-like with up to 5 spikelets; branches when present up to 5mm. Spikelets at first ellipsoid or oblong-ellipsoid, dark brown, 6—14 by 2 mm, finally subglobose, echinate, up to 8 mm thick. Glumes ovate, brown with scarious margins, 2-24 mm long. Flowers 114-2)4 mm long; hypogynous scales 5, oblong-elliptic. Stamens 2. Style triquetrous; stigmas 3. Nuts stellately spreading, stipitate, dis- tinctly rostrate, in cross-section hexagonal by 6 ribs, 3-5/4 by 1)4 mm (beak included), the brown upper part of the body abruptly narrowed into the pale, obconical, narrowly winged lower part, rather grad- ually narrowed into the firm, triquetrous, straight or but slightly curved, smooth, pale, 1/4—3 mm long beak. Distr. Malesia: Philippines (Luzon: Prov. Sor- sogon, Mt Bulusan; Lake Polog and Sierra Madre, NNE of Dingalan; Alabat Island). Ecol. On Mt Bulusan in loose, humus covered ground among shrubberies at 1750 m, in the Sierra Madre in primary Dipterocarpaceous forest at 300—400 m. Notes. In Blumea, l.c., I treated P. rostrata as being conspecific with the Bornean P. radians, but after having seen recent Philippine collections with ripe fruits I am convinced that the two are well distinct. In 1936 UrtTIEN referred P. rostrata to the sy- nonymy of the New Guinean P. longirostris. The two are certainly closely related, but the shape of their fruits differs considerably, so that I prefer to keep them apart. Some additional minor differences corroborate this view. In both P. rostrata and P. longirostris the beak of the fruit is variable in length. Though in RAMos 23642 from Lake Polog and in JAcosBs 7666 from Sierra Madre the beak is shorter than in ELMER 16150 from Mt Bulusan, the type collection of P. rostrata, | refer them to that species on account of the remaining characters. See also under P. parvi- bractea. 6. Paramapania longirostris (KUK.) UITTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 142, p.p. (quoad specim. Nov. Guin.); KERN, Blumea 9 (1958) 216. — Mapania longirostris KOK. Bot. Jahrb. 59 (1924) 55; Ouw1, Bot. Mag. Tokyo 56 (1942) 211.— P. attenuata S. T. BLAKE, J. Arn. Arb. 28 (1947) 210, t. 1. 1974] CYPERACEAE (Kern) 489 Leaves narrowly linear, flat, very gradually narrowed into a flagelliform, curved to circinnate, aculeate-scabrous point, scabrous on the margins and keel at least in the upper half, pale green or glaucescent, 30-65 cm by 4—-6mm; sheaths con- duplicate, sharply keeled, purplish or fuscous. Scapes obscurely trigonous, smooth, (1—)3—10(—20) cm by Y%-Y% mm, the base clothed with a few bladeless, lanceolate, fuscous sheaths. Inflorescence simple, with up to 4 spikelets, usually consisting of a single spikelet; branches when present up to WA cm long. Spikelets at first oblong-ellipsoid, when in fruit ovoid or obovoid, brown, 5-15 by 5—-6mm. Glumes oblong-ovate, obtuse, 3 by 14-134 mm. Flowers 2/4-3 mm long; hypogynous scales 5, the upper 3 lanceolate or narrowly ovate. Stamens 2. Stigmas 3. Nut obliquely erect, lanceolate in outline, stipitate, 6-ribbed, trigonous because 3 of the ribs are more prominent and end in the beak, smooth, shining, brown, 3/4—-5 by 1-1/4 mm (beak included); body ellipsoid, rather gradually attenuate at both ends; beak gradually long-attenuate, triquetrous, more or less curved, 1/4—2 mm long. Distr. Malesia: New Guinea (W. New Guinea: Sorong; Idenburg R.; Dallman, Nabire; NE. New Guinea: Sepik region). Ecol. In very damp rain-forests, on seepages, up to 1200 m. Notes. Very similar in habit to P. gracillima, but readily distinguishable by the quite different shape of the nut. P. attenuata S. T. BLAKE is said to differ from P. longirostris by the twice as long glumes, the flowers slightly shorter than the glumes, and the indistinctly 3-angled (not 6-angled) nut passing gradually (not abruptly) into the relatively shorter beak. However, “glumae ...1,5 mm longae” in UITTIEN’s description of P. rostrata UittiEN (1935, l.c.) is obviously an error for 2.5 mm. The young nuts in the type collec- tion of P. attenuata (BRASS 12930) do not differ from those of P. longirostris. 7. Paramapania simplex (RIDL.) UITTIEN, Rec. Trav. Bot. Néerl. 32 (1935) 190; S. T. BLAKE, J. Arn. Arb. 28 (1947) 209.— Thoracostachyum simplex RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 244. Very near to P. longirostris. Leaves broader, narrowly oblong to linear, 5-30 cm by 8-12 mm, abruptly acuminate, caudate (the tail 1-3 cm long), the base narrowed into a 1—5S cm long petiole, the margins smooth almost throughout, scabrous only at the very top just below the aculeate-scabrous tail. Inflorescence consisting of a single spikelet, 5-8 by 4-5 mm. Glumes 234-34 by 1°%4-2){ mm. Upper sterile scales narrower and slightly longer than in P. longirostris, very narrowly elliptic to oblong-linear. Distr. Malesia: New Guinea (W. New Guinea: Ramoi, Vogelkop, Otakwa R., Idenburg R.; Papua: Fly R., Palmer R.). Ecol. In rain-forests, on shaded river-banks, at low alt., up to 900 m. Note. However different in habit tall specimens of P. longirostris with narrowly linear leaves very gradually narrowing towards the apex, may be from dwarf specimens of P. simplex with oblong, petiolate and caudate leaves, it is questionable whether they are specifically distinct. Provisionally they are kept apart because the difference in facies seems to correlate with one in the shape of the upper hypo- gynous scales, and there is also a marked difference in the scabridity of the leaf-margins. On the other hand I fail to find any essential difference in the nuts. Whether the characters indicated above hold true, can only be decided when the variability of both species will better be known. 7. HYPOLYTRUM L. C. RIcu. in Pers. Syn. 1 (1805) 70. — Hypaelyptum VAHL, En. 2 (1806) 283, p.p. — Beera Lest1B. Essai Fam. Cyp. (1819) 43. — Tunga Roxs. FI. Ind. i (1820) 187, p.p. — Albikia PResL, Rel. Haenk. 1 (1828) 184. — Pandanophyllum HASSK. Tijd. Nat. Gesch. Phys. 10 (1843) 118, p.p.; SrEUD. Syn. 2 (1855) 134, p.p. — Fig. 18-20. Perennial herbs with short, rather stout rhizomes; long-creeping stolons absent (in Mal. spp.). Flowering stems centrally arising from a tuft of normal leaves, erect, trigonous, usually leafy, or from the axils of the lower leaves or below the leaves and then scapiform. Leaves 3—ranked, basal and usually one to several higher on the stem, subcoriaceous or herbaceous, conduplicate at the base, very acute, with 3 more prominent nerves, linear to lanceolate; cauline leaves sheathing the stem, in the scapigerous species reduced to bladeless sheaths. Inflorescence terminal, paniculate, more or less corymbose, sometimes contracted into a single capituliform cluster. Bracts long, leaf-like, not sheathing at the base, in the scapigerous species small, almost glume-like. Spikelets few to very numerous, ovoid to oblong-ellipsoid, often almost globose when in fruit, terete, several- to many-flowered. Glumes membranous to chartaceous, spirally imbricate, glabrous, smooth, 1—nerved, muticous or with slightly excurrent midrib, some lower ones empty. Flowers hermaphrodite, strongly 490 FLORA MALESIANA [ser. I, vol. 7° dorsiventrally compressed. Hypogynous scales 2, hyaline, transversal, opposite, boat-shaped, ciliate on the acute keel, often (always in Mal. spp.?) delicately connate on the adaxial side, but readily rupturing when the fruit develops (in some extra-Mal. spp. also connate on the abaxial side and then forming a tube); no inner flat scales present. Stamens 2, in the axils of the scales (in some extra- Mal. spp. stamens 3). Ovary terminal. Style continuous with the ovary, not or hardly incrassate at the base; stigmas 2, long. Nut biconvex, ovate, elliptic, or almost orbicular, hard, smooth or irregularly wrinkled, with long spongy beak confluent with the nut proper. Distr. Pantropic. In absence of a monograph the number of good species is unknown; it has usually been estimated at c. 50, but this seems much too high, judging from the numerous reductions which I felt compelled to make for the Malesian species. Taxon. Originally Hypolytrum L. C. Ricu. as well as Hypaelyptum VAHL (in all probability a mutilation of RICHARD’s name) and Tunga ROxB. comprised species partly belonging to Hypolytrum partly to Lipocarpha in the present sense. No generic distinction was made between Hypolytrum and Lipocarpha as in both genera two hypogynous scales are found. However, the median scales in Lipocarpha are morphologically quite different from the transversal, sharply keeled, usually connate, ciliate scales in Hypolytrum, the latter in my opinion homologous with the cladoprophyll generally present at the base of the branches in Cyperaceae. KEY TO THE SPECIES AND’ SECTIONS 1. Flowering stems solitary, arising from the centre of a tuft of normal basal leaves; usually also 1-several cauline leaves with well developed leaves present. Lower bracts of the inflorescence foliaceous, the lowest overtopping the inflorescence. 1. Sect. Foliigera CLARKE. 2. Inflorescence paniculate, with well developed rays. 3. Glumes obtuse, not mucronulate, 1’4—2!4 mm long. Anthers pies A % mm long. Nut 1/4-3 mm fone sa . 1. H. nemorum 3. Glumes acute, mucronulate by the slightly excurrent mid- -nerve, 4-5 mm long. Anthers linear, c.UAZAmm long. Nut3—4mmlong . : . 2. H. compactum 2. Inflorescence densely contracted, capituliform or capitate. 4. Stems rather robust, 2-3 mm o. Glumes chartaceous, acutish, with narrow scarious margins, 4-5 mm long. Anthers c. 1/4 mm long. Nut with a narrow acute beak. 2. H. compactum 4. Stems very slender, 1-1)4 mm g. Glumes thinly membranous, obtusish, 3— “34 mm long. Anthers 14-% mm long. Nut with a broad obtusish beak .. . 3. H. capitulatum 1. Flowering stems several, scapiform, arising from the axils of the lower leaves or below the leaves; cauline leaves when present reduced to sheaths. Bracts of the inflorescence small, not foliaceous, none of them overtopping the inflorescence. 2. Sect. Scaposa CLARKE 4. H. humile 1. Section Foliigera CLARKE, Fl. Trop. Afr. 8 (1902) 486. — Sect. Latifolia CHERMEZ. Fl. Madag. fam. 29 (1937) 242. The Malesian species of this section are all very closely related. 1. Hypolytrum nemorum (VAHL) SPRENG. Syst. | 108; (1825) 233; KERN, Blumea 9 (1958) 218; in Back. & Bakh. f. Fl. Java 3 (1968) 456. — Carex laevis Rip_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 100; CLARKE, Ill. Cyp. (1909) t. 106, f. 1-12; Koorpb. Exk. Fl. Java 1 (1911) 184; Atlas (1922) f. 192; minor RUMPH. Herb. Amb. 6 (1750) 21. — Scirpus anomalus Retz. Obs. 5 (1789) 15, non H. anomalum STEUD. (1855). — Schoenus nemorum VAHL, Symb. Bot. 3 (1794) 8; En. 2 (1806) 227. — Hypaelyptum nemorum BEAUV. Fl. Owar. 2 (1810) 13.—H. latifolium L. C. Ricu. in Pers. Syn. 1 (1805) 70; KUNTH, En. 2 (1837) 271; StEUD. Syn. 2 (1855) 132; Mia. Fl. Ind. Bat. 3 (1856) 333: Kurz, J. As. Soc. Beng. 38, ii (1869) 72; Mia. Illustr. (1870) 58; SCHEFF. Nat. Tijd. N. I. 34 (1874) 87; Hook. f. Bot. Mag. 103 (1877) t. 6282; BenTH. Fl. Austr. 7 (1878) 339; F.-VILL. Nov. App. (1882) 309; F.v.M. Descr. Not. 7 (1886) 34; CLARKE, FI. Br. Ind. 6 (1894) 678; J. Linn. Soc. Bot. 34 (1898) 93; K. Scu. & Laut. FI. Schutzgeb. (1900) 191; Ripv. J. Str. Br. R. As. Soc. n. 46 (1906) 225; CLARKE, Philip. J. Sc. 2 (1907) Bot. VALCK. SuR. Nova Guinea 8 (1912) 709; Camus, FI. Gén. I.-C. 7 (1912) 171, f. 22, 2-8; Merr. Int. Rumph. (1917) 106; En. Born. (1921) 54; En. Philip. 1 (1923) 103; KUx. Bot. Jahrb. 59 (1924) 53; ibid. 69 (1938) 260; RipL. Fl. Mal. Pen. 5 (1925) 170, incl. var. penangense; OWI, Bot. Mag. Tokyo 56 (1942) 209, incl. var. depauperatum; S. T. BLAKE, Proc. R. Soc. Queensl. 54 (1943) 71; J. Arn. Arb. 28 (1947) 207; UitTIEN, in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 52. — Tunga diandra Rox. FI. Ind. 1 (1820) 188. — Albikia schoenoides PRESL, Rel. Haenk. 1 (1828) 185, t. 34. — Alhikia scirpoides PRESL, l.c., t. 35. — H. diandrum Dietr. Sp. Pl. 2 (1833) 365. — H. giganteum WALL. ex NEES in Wight, Contr. Bot. Ind. (1834) 93; Boeck. Linnaea 37 (1871) 131. — H. schoenoides Nees, Linnaea 9 (1934) 288; 1974] CYPERACEAE (Kern) 49] Monk. Syst. Verz. (1846) 97; ZOLL. Syst. Verz. 1 (1854) 61.— H. trinervium KUNTH, En. 2 (1837) 272: STEUD. Syn. 2 (1855) 132; Mia. FI. Ind. Bat. 3 (1856) 332; Illustr. (1870) 59; F.-ViLL. Nov. App. (1882) 308; CLARKE, FI. Br. Ind. 6 (1894) 679: Camus, FI. Gén. I.-C. 7 (1912) 173. — H. myrianthum Mia. FI. Ind. Bat. 3 (1856) 333. — H. giganteum « normale O.K. Rev. Gen. Pl. 2 (1891) 751. — H. penangense CLARKE, FI. Br. Ind. 6 (1894) 679; J. Linn. Soc. Bot. 34 (1898) 94: RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 101. — H. costatonux CLARKE in Ridl. J. Str. Br. R. As. Soc. n. 46 (1906) 225: Kew Bull. add. ser. 8 (1908) 52; Merr. En. Born. (1921) 53.— H. viridinux CLARKE, Philip. J. Sc. 2 (1907) Bot. 109: RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 244: Merr. En. Born. (1921) 54.—H. philippense CLARKE, Philip. J. Sc. 2 (1907) Bot. 109; KUK. Bot. Jahrb. 59 (1924) 53; ibid. 69 (1938) 261. — H. amplectens VALCK. Sur. Nova Guinea 8 (1912) 708. — H. anomalum Domin, Bibl. Bot. 85 (1915) 484: H. PrFeIFF. Bot. Arch. 12 (1925) 450, f. 21-22: non STEUD. 1855. — H. scirpoides MeERR. En. Philip. 1 (1923) 103; Etmer, Leafl. Philip. Bot. 10 (1938) 3534. — H. scabrum UittTien, J. Arn. Arb. 20 (1939) 215; S. T. BLAKE, J. Arn. Arb. 28 (1947) 207. — H. microcarpum S. T. BLAKE, J. Arn. Arb. 35 (1954) 235. — Fig. 18-19. KEY TO) THE VARIETIES 1. Spikelets brown. Glumes not or hardly scarious- margined, not Jacerate. Usually stout plants. var. nemorum 1. Spikelets whitish. Glumes with broad white scarious margins, soon lacerate. Usually small slender plants : var. proliferum var. nemorum. Stems usually stout (but slender specimens often occur), rigid, triquetrous upwards, smooth or sca- berulous at the top, with 1—2 well developed leaves above the base, 60-120 cm by (2—)3—6 mm. Leaves longer than the stems, subcoriaceous or chartaceous, flat, more or less scabrous on the margins and mid- nerve beneath, 1-3/4 cm wide; basal sheaths blade- less, keeled, ferrugineous or brownish. Inflorescence paniculate, open to very dense, more or less corym- bose, compound or decompound, with few to very numerous spikelets, S—15 cm across; branches rigid, divaricate, smooth or more or less scaberulous. Bracts 2—4, the lowest similar to the leaves, much overtopping the inflorescence. Spikelets oblong- ovoid or ellipsoid when in flower, 2-6 by 1-2 mm, ovoid to subglobose and 3—6mm wide when in fruit, brown. Glumes membranous, ovate to almost orbicular, concave, obtuse, not or hardly scarious- margined, entire, muticous, 1/4-2)4 by 14-14 mm. Flowers slightly shorter than the glumes. Anthers oblong, 4-% mm. Nut compressed-ovoid to sub- globose, rugulose, sometimes almost smooth, brown or chestnut, 114-3 by 1-2 mm (the conical paler beak included). Distr. From India to Formosa, NE. Queensland and Polynesia, widely distributed in Malesia, not yet collected in the Lesser Sunda Is. BENTHAM /.c. says that it is common in tropical Asia and Africa, and closely allied to if not identical with an American species. KUKENTHAL (1924, l.c.) also gives the distribution as pantropical. It may be Fig. In flower, SPRENG. 1961 18. Hypolytrum nemorum (VAHL) Sanggau, W. Borneo, Sept. (photogr. Father E. ELSENER). better indeed to consider the very closely related African and American ‘species’ as at most racially distinct. Ecol. In swampy places in forests and clearings, at low and medium altitudes, up to 1200 m. Vern. Harassa lalaki, papajungan, S, ielat, M, giring-giring, rumput séndéreian, r. supiding, r. susat bélukar, sélinsing, M (Mal. Pen.), sepédjam, Palemb.. pandan sinhit, Djambi, pandan oweng, Simalur: Philippines: baliis, Tag., bélas, bubugo, salagsalag, Sub., marakibkib, root, Neg., paraibiau, Sul., alisahis, Buk.; New Guinea: kikisa, Orne lang. Notes. VAHL’s description of Schoenus nemorum applies very well to the variety described above, to which the two specimens in the Copenhagen Herbar- ium marked ‘“Schoenus nemorum’ by VAHL certainly belong. CLARKE and others referred the name to an African Hypolytrum, but it is based on an Indian plant. NELMES (Kew Bull. 1955, 71) wrongly referred it to H. wightianum Boeck. (Linnaea 37, 1871, 130: CLARKE, FI. Br. Ind. 6, 1894, 678: J. Linn. Soc. Bot. 34, 1898, 93; Ill. Cyp. 1909, t. 106, f. 13-14), according to CLARKE endemic in Malabar and the Nicobars, with pale, usually glandular-punctate nut. In my opinion H. wightianum cannot be separated specifically from H. nemorum. In H. nemorum in the sense here accepted certainly several races are involved. Its polymorphism has lead to the segregation of numerous so-called species, which I fail to distinguish after having studied a rich material. Specimens with slenderer stems more or less scabrous at the top, and also the axis and branches of the inflorescence scabrid, with copious panicle, small few-flowered spikelets, and small (1’4—2 mm long) nuts have been described as 492 FLORA MALESIANA [ser. I, vol. 7° 19. Hypolytrum nemorum (VAHL) SPRENG. In fruit, same place as fig. 18 (photogr. Father E. ELSENER). Fig. Albikia scirpoides PRESL, H. costatonux CLARKE, H. viridinux CLARKE, H. philippense CLARKE, H. amplectens VALCK. SUR., H. scabrum UrtT., and H. microcarpum §. T. BLAKE. However different the specimens from the Philippines and New Guinea may be from typical H. nemorum, none of the char- acters indicated are reliable, and there are too many intermediates to consider them even as a well-marked subspecies. H. myrianthum Mia. with copious panicle and small nuts was already withdrawn by MIQUEL himself. In the type of H. costatonux CLARKE the flowering stem arises from the centre of the tuft of normal leaves, not laterally as was indicated by CLARKE; longitudinally ribbed fruits often occur in mature specimens of H. nemorum. H. amplectens VALCK. Sur. is said to differ also by the hypogynous scales connate on one side. These scales are always connate on the adaxial side in H. nemorum during anthesis (see also Note under H. capitulatum). According to UITTIEN H. scabrum is amongst others characterized by the leaves beautifully spotted with reddish brown. The same mottling I observed in otherwise typical specimens of H. nemorum, in the related African H. purpurascens CHERMEZ., and in the American H. sylvaticum Porpp. & KUNTH (see CHERMEZON, Bull. Soc. Bot. Fr. 80, 1933, 508: NELMES, Kew Bull. 1955, 69). H. penangense CLARKE differs from normal H. nemorum only by the longer, cylindrical spikelets. Already RIDLEY (1907) supposed it to be abnormal H. latifolium. Obviously the abnormal shape of the spikelets is due to the attack of a fungus, like in Thoracostachyum bancanum ‘var. longispica CLARKE. Remarkable specimens of H. nemorum were col- lected by FORSTEN in 1841 in the Moluccas and by WINKLER (n. 2107) in Borneo. Besides the central stem they possess lateral leafless scapes with very short bracts. var. proliferum (BOECK.) KERN, stat. nov. — H. proliferum Boeck. Linnaea 37 (1871) 126; CLARKE, Fl. Br. Ind. 6 (1894) 679: J. Linn. Soc. Bot. 34 (1898) 94; Rip. J. Str. Br. R. As. Soc. n. 46 (1906) 225; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 101; CLARKE, Ill. Cyp. (1909) t. 104, f. 1-5; Camus, Fl. Gén. I.-C. 7 (1912) 173; Merr. En. Born. (1921) 54; Rip. FI. Mal. Pen. 5 (1925) 170; H. Preirr. Bot. Arch. 12 (1925) 451, f. 19. Stems usually slender, (20—)30—60(—90)cm_ by 1—2(—4) mm. Leaves herbaceous or subcoriaceous, (3—)6—8(—20) mm wide. Inflorescence very variable in size, with few (rarely numerous) spikelets. Spikelets glistening white; upper margin of the glumes con- spicuously white-scarious, pellucid, soon lacerate. Beak of the nut more acute than in the typical variety. Distr. Tonkin (CAMus, I.c.); in Malesia: Banka, Riouw & Lingga Arch., Malay Peninsula (P. Penang, Pahang, Johore, Singapore, Karimon Is.), Borneo. Ecol. In damp woods, on margins of lakes and along streams, up to 800 m. Note. In its typical form readily recognizable by its slender weak habit, narrow leaves, small inflo- rescence, and the white margins of the glumes almost completely covering the brown bases. Some stout specimens I have referred to this variety on account of the white spikelets. As there is nothing proliferous in the plant. BOECKELER’s epithet is inappropriate. 2. Hypolytrum compactum Nees & Mey. [{ Linnaea 9 (1834/5) 288, nom. nud.| ex KUNTH, En. 2 (1837) 271; Nees, Nov. Act. Ac. Caes. Leop.-Car. 16, suppl. 2 (= 19, suppl. 1) (1843) 73; StEuD. Syn. 2 (1855) 132; Mig. Fl. Ind. Bat. 3 (1856) 333: Kurz, J. As. Soc. Beng. 38, ti (1869) 75; Boeck. Linnaea 37 (1871) 127; F.-ViLL. Nov. App. (1882) 309; Merr. Philip. J. Sc. 1 (1906) Suppl. 29; CLarke, Philip. J. Se. 2 (1907) Bot. 109; WINKL. Bot. Jahrb. 44 (1910) 523?; Merr. En. Philip. 1 (1923) 102; KUk. Bot. Jahrb. 59 (1924) 53; ibid. 69 (1938) 261; MerRR. En. Born. (1921) 53?; UrrtTieN, Rec. Trav. Bot. Neéerl. 33 (1936) 155; Onw1, Bot. Mag. Tokyo 56 (1942) 209; S. T. BLAKE, J. Arn. Arb. 28 (1947) 207. —H. xerocarpum CLARKE, Kew Bull. add. ser. 8 (1908) 52; VALCK. Sur. Nova Guinea 8 (1912) 709. — Mapania thoreliana Camus, Not. Syst. 1 (1910) 250; Fl. Gén. I.-C. 7 (1912) 178. Stems rather stout, rigid, subtriquetrous, smooth, sometimes slightly scaberulous just below the inflo- rescence, with a well-developed leaf in or above the middle, 30—60 cm by 2—3 mm. Leaves about as long as or longer than the stem, subcoriaceous, flat, more or less scabrous on the margins in the upper part, pale green, 7-14 mm wide, two lateral nerves very 1974] Prominent beneath; basal sheaths bladeless, acute, keeled, ferrugineous or brownish. Inflorescence com- pound, with several to many spikelets, contracted into a very dense, ovoid to globose mass, 2-5 by 14-4 cm, the lowest cluster often somewhat remote, rarely the whole inflorescence somewhat open. Bracts 4—7, the lowest similar to the leaves, up to 60cm long. Spikelets oblong to linear-oblong, ovoid when in fruit, many-flowered, ferrugineous- brown, 7-9 by 1)4-3 mm, 5-6 mm wide when in fruit. Glumes chartaceous, narrowly ovate to lanceo- late, acutish, minutely mucronulate by the slightly excurrent midnerve, with narrow scarious margins, 4-5 by 134-2 mm. Flowers slightly shorter than the glumes, narrowly elliptic; scales ciliate-setulose on the keels, the setulae more or less confluent. Anthers linear c. 114mm. Nut compressed, ovate, with keeled margins, irregularly rugulose, acuminate into the acute pale beak, 3-4 by 174-2 mm. Distr. Andamans (CLARKE, I.c.), Indo-China, Queensland (Cape York Peninsula); in Malesia: Borneo, Philippines (Palawan, Mindoro, Luzon), Celebes (Kendari), Moluccas (Ceram), New Guinea (incl. Japen, Waigeo, and Aru Is.). Ecol. In rain-forests at low altitude, up to 750 m. Vern. Nomahagino, Sorong, gafaat, Waigeo; Philip.: tuhog-dalag, Tag. 3. Hypolytrum capitulatum VALCK. SuR. ex CLARKE, Kew Bull. add. ser. 8 (1908) 51; UitTIEN, Rec. Trav. Bot. Néerl. 33 (1936) 155. — Fig. 20. Stems very slender, subtriquetrous, smooth, (10—)40—60 cm by 1-1/4 mm, with 1-2 well developed leaves in or above the middle, rarely in small speci- mens cauline leaves absent. Leaves about as long as the stems, herbaceous, flat, 5-8 mm wide, two lateral nerves rather prominent, margins scaberulous towards the top; basal sheaths bladeless, keeled, ferrugineous or brownish. Inflorescence simple, densely contracted, capitate or almost so, with 3—6 spikelets, 1-2 cm across. Bracts 2-3, the lowest similar to the leaves, S—15 cm long, the upper ones much shorter. Spikelets ovoid, many-flowered, ru- fescent, (S—)8—10 by 5—7 mm when in fruit. Glumes thinly membranous, ovate-elliptic, obtusish, with rather broad pellucid margins, purplish lineolate, 3-3)% by 1)4-134 mm, the inconspicuous midnerve ending somewhat below the apex. Flowers slightly shorter than the glumes, narrowly elliptic, 24-3 by 1 mm; scales minutely ciliolate on the keel. Anthers linear, 4-24 mm. Nut compressed, elliptic, longitu- dinally rugose, rufous or brown, 3-4 by 1}4-2 mm (beak included), the beak obtusish, slightly longer than the nut proper. Distr. Malesia: W. Borneo (Bukit Singkadjang, Sendabai Lakes), only 3 collections. ‘ge Margins of lakes, clearings, at low alti- tude. Note. Closely allied to H. compactum; differing from it in the much slenderer habit, the thinly membranous obtusish glumes with pellucid margins, the smaller anthers, the different shape of the beak of the nut, and the minutely ciliolate scales. Glumes with pellucid margins are also found in H. nemorum var. proliferum, but here the inflorescence is not capitate, the spikelets are smaller, and the glumes and nuts much smaller. CLaRKE, I.c., calls the hypogynous scales connate only at the base. However, they are fused on the CYPERACEAE (Kern) 493 Fig. 20. Hypolytrum capitulatum VALCK. SuR. ex CuarKE. a. Habit, x 4, b. floral diagram, c. glume, x 6, d—e. flower, x 10, f-g. nut enveloped by scale, frontal and dorsal views, h. ditto, scale removed, both x 6. 494 adaxial side almost to the top, which is probably the case in all Malesian spp. They are torn up quite or FLORA MALESIANA [ser. I, vol. 7° almost to the base by the ripening nut, which is enclosed by the persistent scales. 2. Section Scaposa CLARKE, Fl. Trop. Afr. 8 (1902) 487. — Sect. Africana CHERMEZ. Fl. Madag. fam. 29 (1937) 240. 4. Hypolytrum humile (STEUD.) Boeck. Linnaea 37 (1871) 128; ScHerrF. Nat. Tijd. N. I. 34 (1874) 58; UriTien, Rec. Trav. Bot. Néerl. 33 (1936) 153; in Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 52; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 456. — Pandanophyllum humile Hassk. [ex Mor. Syst. Verz. (1846) 98; ZoLL. Syst. Verz. 1 (1854) 61, nom. nud.| ex STEUD. Syn. 2 (1855) 134; Mia. FI. Ind. Bat. 3 (1856) 334. — Mapania multispicata (non RIDL. J. Str. Br. R. As. Soc. n. 23, 1891, 15) CLARKE, Fl. Br. Ind. 6 (1894) 682 p.p., quoad specim. jav. et. synon.; J. Linn. Soc. Bot. 34 (1898) 95, p.p. (quoad specim. jav.); KoorpD. Exk. Fl. Java 1 (1911) 203; H. PreirF. Bot. Arch. 12 (1925) 449. Scapes lateral, obtusely trigonous to almost terete, more than 1 m by 10—20 mm, margins and midnerve scabrous at the top. Inflorescence globose, obovoid, or ellipsoid, consisting of 10—60 densely congested spikelets, 114-274 cm long. Bracts 3-4, similar to the basal sheaths of the scape, subspathaceous, shorter than to as long as the inflorescence, the lowest often somewhat remote. Spikelets (when in flower) oblong, acute, many-flowered, 5—8 by c. 2mm. Glumes ovate to oblong-ovate, obtuse, with membranous-pellucid apex and margins, 2)4-3 by 134-2 mm. Flowers about as long as the glumes. Anthers linear, 1-114 mm long. Nut slightly flattened, ellipsoid or obovoid, acuminate at both ends, apiculate, longitudinally and subreticulately rugose, 24-3 by 134-2 mm, fuscescent. leafless, smooth, 10—35 cm by 1—2 mm, at the base with a few acute ferrugineous sheaths up to 4cm long, often also a bladeless sheath in or about the middle. Leaves subcoriaceous, linear, acute, up to Distr. Malesia: West Java. Ecol. In damp forests, 400—1000 m. Vern. Irateun hahajaman, S. 8. SCIRPUS LINNE, Sp. Pl. 1 (1753) 47; Gen. Pl. ed. 5 (1754) 26; KUNTH, En. 2 (1837) 157; STEUD. Syn. 2 (1855) 83; Boeck. Linnaea 36 (1869-70) 475; B. & H. Gen. Pl. 3 (1883) 1049; Pax in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 111; CLARKE, Kew Bull. add. ser. 8 (1908) 111; BEETLE, Am. J. Bot. 27 (1940) 63-64; ibid. 28 (1941) 469-476, 691-700; ibid. 29 (1942) 82-88, 653-656; ibid. 30 (1943) 395—401; ibid. 31 (1944) 261-265; ibid. 33 (1946) 660—666. — Fig. 21-31. For the numerous generic names here referred to Scirpus, see under the synonymy of the sections. Annual or perennial herbs of very various habit, stonoliferous or cespitose, glabrous or only minutely hairy in the inflorescence. Stems erect or procumbent, sometimes floating or submerged, terete to acutely angled, usually smooth. Leaves either with more or less developed blades, or reduced to bladeless sheaths; ligule membranous or wanting. Inflorescence terminal, but often pseudolateral, capitate or anthelate, or consisting of a single spikelet. Involucral bracts 1—several, in pseudolateral inflorescences seemingly a continuation of the stem, in 1-spiculate inflorescences usually reduced and glume-like. Spikelets solitary or clustered, terete or angular, usually many-flowered. Rachilla persistent, not winged. Glumes spirally arranged, acropetally caducous, the lower 1—3 often empty. Flowers hermaphrodite, the upper ones often tabescent. Perianth consisting of 1—6(—11) setaceous (rarely flattened, scale-like) bristles, not rarely absent. Stamens 1—3; anthers linear or oblong, with shortly produced, smooth or bristly connective. Style caducous, continuous with the ovary (neither separated by a constriction nor different in texture, but passing insensibly into the top of the ovary), glabrous or very rarely sparsely fimbriate, hardly or not dilated at the base; stigmas 2—3. Nut 2- or 3-sided, obovate or elliptic to oblong 1974] CYPERACEAE (Kern) 495 Fig. 21. Scirpus junghuhnii Mig. a. Habit, x 4, b. spikelet, x 4, c. glume, d. flower, e. nut, all x 8 (a—b, d VAN STEENIS 8484, c, e JUNGHUHN 477). 496 FLORA MALESIANA [ser./1, valigF in outline, often apiculate, smooth or rugulose; epidermal cells hexangular or vertically oblong to linear. Distr. A large genus (possibly 200 spp.) of worldwide distribution, in Malesia 21 spp. Several Malesian species are mountain plants with disjunct areas. Scirpus beccarii and S. junghuhnii are apparently endemics of Sumatra. Ecol. Most species prefer open, wet localities: swamps, pools, ditches, wet rice-fields, lake-shores, and stream banks: S. confervoides grows always, S. fluitans often submerged. In Malesia prolificating inflorescences have been observed in S. mucronatus, S. inundatus, and S. subtilissimus. Uses. Only a few species have some economic importance. The stems of S. grossus and S. mucronatus are often used for making cheap but durable mats and bags; S. juncoides is sometimes used as a green food for cattle. Notes. The genus is in comparison with other genera of the family heterogeneous. It comprises all the species of subfamily Cyperoideae left after the more or less homogeneous genera have been taken out. The only characteristics of Scirpus are the spirally arranged glumes and the style not articulated with the ovary. Several attempts have been made to split it up into smaller genera, or to divide it into subgenera. These efforts have failed in my opinion, as the resulting genera or subgenera are just as well heterogeneous, and often agglomerates of very dissimilar species. The only way out is obviously to distinguish a number of sections each comprising a set of probably allied species, a procedure also employed by CHERMEZON in the Fl. Madag. The Malesian species are tentatively arranged in such sections, to which a separate key has been added. Specific delimitation offers several difficulties, as many species appear to be racially differentiated, especially when isolated populations have been formed in mountainous country. A good example of this is the S. subcapitatus complex. It is remarkable that certain characters, generally accepted as important in Cyperaceae, as for example the structure of the perianth bristles, are less constant in such complexes, which are doubtless genetically coherent. In some cases specific limits turn out to be less distinct than was assumed formerly; for instance S. mucronatus ssp. clemensii from New Guinea distinctly diminishes the demarcation between S. mucronatus and S. juncoides. KEY TO THE SECTIONS REPRESENTED IN MALESIA 1. Spikelets not squarrose. Perennials, or annuals with nuts longer than /4 mm. 2. Involucral bracts several, foliaceous, flat. 3. Hypogynous bristles inconspicuous, shorter than the glumes, or absent. 4. Stems nodeless:Leaves basaly Spel (2 >. .755 fr teas . = =. . I. Sect. Actnsemes 4. Stems noded, leafy. 5. Spikelets usually not numerous, large. Glumes S—7 mm long. Spp.2-3 . . . 2. Sect. Maritimi 5. Spikelets very numerous in a decompound umbelliform inflorescence, small. Glumes much smaller. Sp Aaeeeee ote ee eee lie ote | Sa ee dey a 3. Sect. Scirpus 3. Hypogynous bristles strongly elongating after anthesis, at maturity greatly exceeding the glumes, strongly curled and entangled. Sp. 5 OS, ea eee Gt 4. Sect. Trichophorum 2. Involucral bract non-foliaceous, single, either looking like a continuation of the stem, or glume-like, or absent. 6. Stems branched, procumbent, floating, or submerged. 7. Hypogynous bristles absent. Spikelet several-flowered. Spp.6-8 . . . . . 5. Sect. Eleogiton 7. Hypogynous bristles present. Spikelet 1(—2)-flowered.Sp.9. . . . . . 6. Sect. Confervoidei 6. Stems not branched, erect. 8. Inflorescence terminal, subtended by the outer caducous, often mucronate glume. Sp. 10. 7. Sect. Baeothryon 8. Inflorescence pseudolateral, subtended by a persistent bract looking like a continuation of the stem. 9. Stems tall, usually stout. Bristles usually present. Nut (1—-)1/4-2/4 mm long, smooth or transversely ridged. Spp. 11-17 tinted aE hatte ters Fee rrp eee ee 9. Stems low, setaceous. Bristles absent. Nut smooth, 4-14 mm long. Spp. 18-20 . 9. Sect. Isolepis 1. Spikelets squarrose by the recurved mucros of the glumes. Small tufted annuals with nuts c. J4 mm long. Sask. MUI TAI Le) OD WAU Wie oe eT Ee KEY TO THE SPECIES 1. Spikelets strongly squarrose by the recurved mucros of the glumes. Small tufted annual with nuts c. 4 mm ot DOV. 3) Ey eae ee Tp ee ear rm. eee ae Ree ae | 21. S. squarrosus 1. Spikelets not squarrose. Perennials, or annuals with nuts longer than 4 mm. ps pe i several, flat, leaf-like, not continuous with the stem. Stout perennials with well developed leaf- lades. 3. Leaves all basal. Stem nodeless. Stigmas 3. Nut trigonous. . . . . . . . . I. S. grossus 3. Stems noded, leafy. 4. Glumes 1/4-2)4 mm long. Nut 4-1)4 mm long. 5. Perianth-bristles strongly elongating after anthesis, finally greatly exceeding the glumes, curled and entatisied. stiomas 3. Nutirieonops |; 2-6. + oe ee th a ee 5. Perianth-bristles inconspicuous, shorter than the glumes, straight, or absent. Stigmas 2 (rarely in some flowers 3). Nut planoconvex or unequally biconvex shes Alo. 1 4S Steraianins 4. Glumes 4—7 mm long. Nut larger. 6. Inflorescence terminal, umbelliform. Glumes pubescent outside. Bristles straight, retrorsely scabrous. 2. S. maritimus 1974] CYPERACEAE (Kern) 497 6. Inflorescence consisting of up to 4 distant corymbiform partial inflorescences. Glumes papillose or hispidulous at the top. Bristles flexuous, antrorsely scabrous. . . 3. S. junghuhnii 2. Inflorescence subtended by a single bract similar to and continuous with the stem (sometimes a very small patent second bract present), or bracts absent or glume-like. 7. Inflorescence pseudolateral because of the bract looking like a continuation of the stem. 8. Glumes 5—7 mm long, pubescent onthe back . . epee - 2+ SaMlanitimus 8. Glumes at most 4 mm long, glabrous (but sometimes with ciliate margins). 9. Stem and bract transversely septate, the bract longer than the stem proper (hence inflorescence seemingly inserted in the lower half of the stem)... . . 16. S. articulatus 9. Stem and bract not septate, the bract shorter than the stem (hence inflorescence seemingly inserted in the upper half of the stem). 10. Perianth absent. 11. Nut conspicuously transversely wavy-ridged, black when mature. Inflorescence sometimes capitate, but usually with one of the rays I(—4) cm long... . . 17. S. lateriflorus 11. Nut smooth, stramineous to light brown when mature. Inflorescence capitate or consisting of a single spikelet, sometimes proliferous. 12. Stems branched. Style 2-cleft, incompletely 3-cleft, and 3-cleft on the same specimen. Nuts planoconvex and triquetrous on the same specimen, 1%-2mmlong. . . . 7. S. beccarii 12. Stems unbranched. Style 3-cleft, very rarely in some flowers 2-cleft. Nuts triquetrous, usually c. | mm long, rarely up to 114 mm. 13. Stamens 3. Anthers “4-1 mm long, with distinctly produced connective. Inflorescence partly hidden by the dilated base of the bract . . . 18. S. aucklandicus 13. Stamens 1—2(—3). Anthers 74 mm long, with hardly produced connective. Inflorescence not hidden by the base of the bract. 14. Rhizome filiform, shortly creeping, forming mats. Stems filiform, terete, 4—)4 mm thick. Leaf-blades well developed. Spikelets solitary or in clusters of 2-3 . . 19. S. subtilissimus 14. Without definite rhizome. Stems tufted, strongly compressed, 14-1 mm wide. Leaves usually all reduced to their sheaths, only the upper one often with a very short, rarely + elongated blade. Spikelets (except in depauperate specimens) in clusters of 3-12 . . 20. S. inundatus 10. Perianth present, consisting of setaceous bristles or ligulate-spatulate scales. 15. Inflorescence anthelate, open. Glumes often notched at the apex, mucronate. Connective of the anthers with a fimbriate appendage. 16. Hypogynous bristles 5—6, setaceous, retrorsely scabrous. Glumes distinctly ciliate. 11. S. lacustris 16. Hypogynous scales usually 4 (3—S), ligulate-spatulate, plumosely fringed with antrorse hairs. Glumes only microspically ciliolate. . . . . 12. S. litoralis 15. Inflorescence capitate. Glumes not emarginate, apiculate. Connective of the anthers with smooth appendage. 17. Stems sharply triquetrous, usually rather stout (up to 8 mmthick) . . . 13. S. mucronatus 17. Stems terete or more or less 4—S-angular, but not sharply triquetrous, usually slender. 18. Involucral bract very short (1—2 cm), hardly overtopping the inflorescence. Glumes distinctly many-nerved (especially when dry). Stems usually rather coarse. 13. S. mucronatus ssp. clemensii 18. Involucral bract slender, 5—15 cm long, much overtopping the inflorescence. Glumes faintly nerved (only the midnerve distinct). Stems slender. 19. Bristles shorter to slightly longer than the nut, the longest 2-24 mm. Nut biconvex (only low-convex on the ventral side). Anthers 1-1/4 mm long. . . . 14. S. juncoides 19. Bristles all distinctly longer than the nut. Nut planoconvex (the ventral side flat). Anthers Y4-%mmiong. . . . . . IS. S. wallichii 7. Inflorescence terminal, or stems and branches terminated by a single spikelet. 20. Spikelets 1(—2)- flowered. Submerged very delicate aquatic, the stem-nodes with numerous thread-like sterile stems not to be confused with leaves .. . 9. S. confervoides 20. Spikelets more-flowered. Habit quite different, if an aquatic each node with one true leaf. 21. Leaf-blades well developed. Perianth-bristles absent. Stigmas 2; nut planoconvex or biconvex. Stems procumbent or floating, branched, the branches bearing a single terminal spikelet. 22. Nut planoconvex, sharply angled, 1/4— 1% by c. 1 mm. Spikelets 3-6 by 2-3 mm .__ 6. S. fluitans 22. Nut slightly biconvex in the centre, thinner towards the a and there thickened into an obtuse edge, 174-2 by c. 1 mm. Spikelets 5-8 by3-4mm . . . 8. S. crassiusculus 21. Leaf-blades reduced to an up to 2 cm long (usually much shorter) mucro. Perianth-bristles 6. Stigmas 3; nut trigonous. Stems erect, not branched. Inflorescence consisting of 1—5 spikelets. 10. S. subcapitatus 1. Section Actinoscirpus OxHwI, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 98. — Hymenochaeta BEAUV. ex Lestis. Ess. Fam. Cyp. (1819) 43?; emend. Nees, Edinb. New Phil. J. 17, n. 34 (1834) 264; Linnaea 9 (1834) 293; in Wight, Contr. Bot. Ind. (1834) 498 FLORA MALESIANA [ser. I, vol. 7° 110. — Sect. Hymenochaeta BEETLE, Am. J. Bot. 33 (1946) 661. ype species: S. grossus Lf. 1. Scirpus grossus LINNE f. Suppl. (1781) 104; PRESL, Rel. Haenk. 1 (1828) 195; SteEuD. Syn. 2 (1855) 87; Mia. FI. Ind. Bat. 3 (1856) 307 (f. minor); BOoEcK. Linnaea 36 (1870) 723; CLARKE, FI. Br. Ind. 6 (1893) 659; Ill. Cyp. (1909) t. 49, f. 12; VALcK. Sur. Nova Guinea 8 (1912) 705; Camus, Fl. Gén. I.-C. 7 (1912) 136; Mere. En. Philip. 1 (1923) 117; Ripi. Fl. Mal. Pen. 5 (1925) 162; BAck. Onkr. Suiker. (1928) 151, t. 153; OHwi1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 122; BeeTLte, Am. J. Bot. 33 (1946) 661; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p: 10:°S: I BEAKE,, JArne Arb: 35° (1954) 2205; RAYMOND, Nat. Canad. 84 (1957) 117; KERN in Back. & Bakh. /f. Fl. Java 3 (1968) 458. — S. kysoor (non ROxB.) LLANOsS, Fragm. Pl. Filip. (1851) 20 (‘kisoor’); F.-ViLL. & Naves in Blanco, FI. Filip. ed. 3, 4' (1880) 14 (‘kisoor’). — Cyperus difformis (non L.) BLANCO, FI. Filip. (1837) 32; ed. 2 (1845) 22; ed. 3, 1 (1877) 41. — S. aemulans Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 87. — S. maritimus var. aemulans Mia. FI. Ind. Bat. 3 (1856) 306. — S. grossus var. kysoor (non CLARKE, Fl. Br. Ind. 6, 1893, 660) CLARKE, Philip. J. Sc. 2 (1907) Bot. 100. — Schoenoplectus grossus PALLA, Allg. Bot. Zeitschr. 17 (1941) Beil. 3. — Fig. 22. Stout perennial, with rather long stolons ending in small tubers. Stems erect, sharply triquetrous with concave sides, septate-nodulose, smooth, or slightly scaberulous at the top, 80-200 cm tall, up to 2)4 (at the top 4-1) cm thick. Leaves basal, 50-180 by up to 3 cm, in the lower half keeled, triquetrous, in the upper half almost flat, acute, very gradually acuminate, strongly septate-nodulose, with more or less scabrous margins; sheaths spongy, stramineous. Inflorescence terminal, large, corymbiform, decom- pound to supradecompound. Involucral bracts sev- eral, patent, flat, at least 2 of them far overtopping the inflorescence, 15-70 cm. Primary rays several, unequal, spreading, scaberulous. Spikelets numerous, solitary, sessile and peduncled, ovoid to oblong- ovoid, acutish, densely many-flowered, 4-10 by 3/4-4 mm. Glumes membranous, appressed, con- cave, ovate to broadly ovate, obtuse, muticous or very shortly mucronulate, with strong green midnerve and ferrugineous to reddish brown nerveless sides, more or less puberulous, glabrescent, minutely ciliolate, 244-3 by 174-2 mm. Bristles 4—6, retrorsely scabrous, incurved at the top, as long as to somewhat longer than the nut. Stamens 3; anthers linear, 1/4- 14 mm. Style glabrous; stigmas 3. Nut trigonous, Fig. 22. Scirpus grossus L.f.in Rawa Tembaga, a lowland swamp near Bekassi(E of Djakarta) with Nymphoides indica (L.) O.K. (photogr. VAN STEENIS, 1941). 1974] CYPERACEAE (Kern) 499 obovate, apiculate, smooth, brown, 14-13% by c. | mm. Distr. Throughout India (except the NW), SE. Asia, S. China, Bonin Is., tropical Australia (N. Territory, N. Queensland), probably throughout Malesia, but not yet collected in the Lesser Sunda Is. and the Moluccas. Ecol. In swampy or inundated localities, pools, ditches, rice-fields, often abundant, 0-850 m. Uses. Often used for making sleeping mats, bags, and baskets; the stems are dried after removal of one of the ribs, then flattened, and bleached in the sun. Vern. Méndarong, ménsiang (mansiang, mésiang, musiang, masiang), ménurong, murong, M, beba- wangan, walingi, S, lingi, wlingén, wlingi, wlingian, J, balingeh, Md, basiang, Bat., mansiro daun, Menangk., réduk, Palembang, bundung, W. Borneo, kaingas, kawasar, tinorong, Alf. Minah.; Philip.: agds, bang- kuang, ragiudiu, Bik., bagads, baki-baki, balangot. P.Bis.. balakbak, Pang., tikug, Mbo, tikiu, titiu, Tag. Note. In var. kysoor (ROXB.) CLARKE, FI. Br. Ind. 6 (1893) 660 (S. kysoor Roxs. Hort. Beng. 1814, 6: Fl. Ind. 1, 1820, 235. — S. grossus f. kysoor BEETLE, Am. J. Bot. 33, 1946, 661) from India the glumes are more distinctly mucronate, the weaker bristles villous to almost plumose by longer weaker hairs. and the stems usually scabrous at the top. According to CLARKE the hairs of the bristles are multicellular (see also Ill. Cyp. t. 49, f. 11), which is incorrect. CLARKE (1907) recorded this variety from the Philippines and “‘many examples from the Malayan Peninsula and Archipelago’, but I have not seen any Malesian specimen. It is obviously restricted to India. 2. Section Maritimi CHERM. Fl. Madag. fam. 29 (1937) 155, descr. gall. — Scirpus b. Euscirpus «. Phyllanteli BEURL. Pl. vasc. Scand. (1859) 55. — Sect. Euscirpus B. & H. Gen. Pl. 3 (1883) 1051, p.p. — Bolboschoenus PALLA in Koch, Syn. Deut. u. Schw. Fl. ed. 3, 3 (1904) 2531. — Sect. Bolboschoenus BEETLE, Am. J. Bot. 29 (1942) 82. Typespecies: S. maritimus L. 2. Scirpus maritimus LINNE, Sp. Pl. 1 (1753) 51: KuNTH, En. 2 (1837) 167; Boeck. Linnaea 36 (1870) 722; BENTH. Fl. Austr. 7 (1878) 335; CLARKE, FI. Br. Ind. 6 (1893) 658. Perennial, with horizontally creeping rhizome forming hard, ovoid tubers at the nodes. Stems erect, approximate or solitary from a tuberous base, slender to stout, triquetrous, striate, smooth, or scaberulous just below the inflorescence, 15—180 cm tall and 1-15 mm thick towards the base. Leaves cauline, stiff, flat, with revolute margins when dry, scabrid on the margins, very gradually narrowed into the long, triquetrous tip, (1—-)2-12 mm wide, the upper ones overtopping the inflorescence; ligule absent; sheaths tight, smooth, with slightly emar- ginate, truncate or somewhat produced mouth, the basal ones greyish brown. Inflorescence very variable in development, umbelliform with several unequal, smooth, up to 10cm long primary rays and short secondary rays, to capitate, sometimes even reduced to a single spikelet. Bracts 1-several, foliaceous, dilated at the base, not sheathing, overtopping the inflorescence, the longest usually erect, the remainder oblique. Spikelets solitary or in clusters of 3-10, ovoid to oblong-ovoid, terete, densely many-flowered. acutish, ferrugineous to castaneous, 1—2(—4) cm long. Glumes membranous, appressed, not keeled, ovate or elliptic, emarginate or lacerate at the top, pubescent outside, 4-7 mm long, 1(—3)-nerved, the strong midnerve produced into an antrorsely scabrid awn. Bristles (0O—)3—6, retrorsely scabrous, caducous. Stamens 3; anthers linear, 2—5 mm long, with bristly appendage of the connective. Style long, glabrous: stigmas 2 or 3 (the number often differing in flowers of the same spikelet). Nut two-sided or trigonous, obovate, brown to blackish, 2/4—4 mm long. Distr. Scirpus maritimus in the wide sense here accepted is widely distributed over the tropical and temperate regions of the whole world: from Malesia only 4 collections are known: Philippines (Luzon: Laguna), New Guinea (NE. New Guinea: Western Highlands and Mt Sarawaket; Papua: Lower Fly River). Ecol. In lowland rice-field (Laguna), in tall reed swamp (Western Highlands, c. 2500 m), on loose sand on open foreshores (Lower Fly R.), in moist localities at c. 1150 m (Mt Sarawaket). Notes. Extremely polymorphous. The type is from Europe. In European specimens the style is trifid as a rule, but usually there are bifid styles in some basal flowers of each spikelet. Specimens in which digynous flowers prevail are very rare in Europe (var. digynus Gopr. Fl. Lorr. 3, 1844, 91). The Malesian plants are strictly or predominantly digynous. S. maritimus var. fluviatilis Torr. Ann. Lyc. Nat. Hist. N.Y. 3 (1836) 325, mainly characterized by its constantly trigonous nuts and 6 persistent hypogynous bristles, may be a good species [S. fluviatilis (TorR.) A. Gray, Man. Bot. ed. 1 (1848) 527; KoyAMA, J. Fac. Sc. Un. Tokyo III, 7 (1958) 334]. It is widely distributed (N. America, Japan, Korea, Manchuria, China, and Indo-China), but not known from Malesia. The specimens from the Western Highlands in New Guinea are stout, c. 180 cm tall, with pale green leaves up to 12 mm wide, a compound inflorescence with c. 8 unequal, up to 10 cm long rays, solitary, 10-15 mm long spikelets, c. 4mm long glumes, 3-5 caducous bristles, 3—3)4 mm long anthers, and 2 or 3 stigmas in flowers of the same spikelet. They may come nearest to a taxon described from New Zealand as S. medianus Cook, Trans. Proc. R. Soc. New Zeal. 76 (1947) 569, also with bifid and trifid styles in the same spikelet, but with a simple inflo- rescence and probably a different nut. The Philippine plants may belong to the E. Asian form known as S. planiculmis FR.SCHM., Reisen 500 FLORA MALESIANA [ser. I, vol. 7° Amurl. (1868) 190; Koyama, J. Fac. Sc. Un. Tokyo III, 7 (1958) 330. The stems are up to 55cm tall, 2-3 mm thick, the capitate inflorescence consists of 3-4 spikelets, the glumes are c. 6mm long, the anthers 4 mm, and there are c. 4 caducous bristles; no fruits present; style 2-fid. The Lower Fly River collection consists of very slender, narrow-leaved specimens (stems 30—60 cm by 1-14 mm; leaves 1-1/4 mm wide; inflorescence a single, pseudolateral spikelet; glumes 5—7 mm; bristles 2—3; nut lenticular, strongly compressed, slightly concave on the inner side, 374-4 mm long). I take them for a somewhat depauperated form of S. planiculmis. S. T. BLake, J. Arn. Arb. 35 (1954) 204, referred them with some doubt to S. affinis RoTH in R. & S. Syst. Veg. 2 (1817) 140 [S. strobilinus Roxs. FI. Ind. 1 (1820) 222] from Central, S. and SE. Asia, which has large, straw-coloured spikelets, and smaller nuts. In my opinion neither S. planiculmis nor S. affinis can be separated specifically from S. maritimus. 3. Scirpus junghuhnii Mig. Fl. Ind. Bat. 3 (1856) 307; Suppl. 1 (1861) 261; KUxK. Bull. Jard. Bot. Btzg 16 (1940) 301, incl. var. minor KUK.; in Fedde, Rep. 53 (1944) 102. — Fig. 21, 23. Perennial with very short rhizome. Stems erect, usually stout, trigonous, smooth, up to 150 by 1 cm. Leaves basal and cauline, coriaceous, flat, gradually tapering into a long scabrous tip, 10-25 mm wide; margins scabrous in the upper part; lower sheaths shining fuscous; ligule a dark, minutely ciliolate rim. Fig. 23. Scirpus junghuhnii Mig. in a mountain heath (blang) on Mt Goh Lembuh (Gajo Lands, N. Sumatra), at c. 3000 m (photogr. VAN STEENIS). 1974] CYPERACEAE (Kern) 501 Inflorescence large, decompound, up to 70 by 20 cm, consisting of (1—)2—4 distant corymbiform partial inflorescences with hispidulous rachis; peduncles erect, rigid, trigonous, smooth, subtended by long leafy bracts, the lowest up to 25cm; secondary branches divaricate, up to 3 cm, tertiary ones slender, up to 2 cm, with 3—7 spikelets. Spikelets approximate, at right angles to the rachis, easily caducous, shortly peduncled, often curved upwards, oblong-lanceolate, somewhat angular, acute, loosely 8—10-flowered, 10-15 by 2/4-3 mm. Glumes rather firm, slightly keeled, oblong to narrowly lanceolate, acute, shortly mucronate (mucro c. }4 mm), with strong midnerve, papillose or hispidulous at the top, auricled at the base, rufous to dark brown, 5-6 by 2/4—-3 mm, the margins more or less inrolled when dry. Bristles (4—)6, delicate, flexuous, antrorsely scabrous, S— 6mm. Stamens 3; filaments strongly elongated after anthesis; anthers linear, 2)}4-3)4 mm. Style c. 5 mm long, fimbriate at the top, slightly dilated at the base; stigmas 2 and 3 in the same spikelet, somewhat shorter than to almost as long as the style. Nut biconvex or trigonous, elliptic, shortly annulate at the top, densely puncticulate, brown, 124-2 by 1 mm. Distr. Malesia: N. & Central Sumatra (Atjeh: Gajo Lands; Lubuk Raja, Mt Malintang, Mt Kerintji). Ecol. In brushwood, on river banks, on open heaths, at high altitudes (2200-3400 m), often gre- garious. Notes. KUKENTHAL’s var. minor is merely a less robust form of very high altitudes, without systematic value. KUKENTHAL placed this remarkable species in the affinity of S. maritimus; however, it probably repre- sents a separate section. 3. Section Scirpus Seidlia Opi1z, Beitr. Naturgesch. 11 (1826) 349. — Sect. Euscirpus B. & H. Gen. Pl. 3 (1883) 1051, p.p. — Sect. Phylloscirpus Pax in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 112. — Sect. Sylvatici CLARKE, Fl. Br. Ind. 6 (1893) 661, p.p. — Sect. Seidlia CLARKE, Kew Bull. add. ser. 8 (1908) 113, p.p. — Sect. Taphrogeton Rcus. Fl. Germ. Excurs. 1 (1830) 79; BEETLE, N. Am. FI. 18 (1947) 486. Type species: S. sylvaticus L. 4. Scirpus ternatanus REINW. ex Mig. FI. Ind. Bat. 3 (1856) 307; in De Vriese, Pl. Ind. Bat. Or. (1857) 140 (‘ternatensis’); CLARKE, J. Linn. Soc. 34 (1898) 83; Philip. J. Sc. 2 (1907) Bot. 100; Ill. Cyp. (1909) t. 52, f. 1-4 (‘ternatensis’); MERR. En. Philip. 1 (1923) 118; KOK. Bot. Jahrb. 69 (1938) 259; Bull. Jard. Bot. Btzg III, 16 (1940) 301; OHwi, Bot. Mag. Tokyo 56 (1942) 204; Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 108; KUx. in Fedde, Rep. 53 (1944) 101; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 10; S. T. Biake, J. Arn. Arb. 35 (1954) 203; RAYMOND, Nat. Canad. 84 (1957) 115; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 458. — S. chinensis Munro in Seem. Bot. Voy. Herald (1857) 423; CLARKE, Fl. Br. Ind. 6 (1893) 662; VALCK. SurR. Nova Guinea 8 (1912) 705; Rip. Trans. Linn. Soc. II, Bot. 9 (1916) 242, non OsBeck, 1757. — Hypolj- trum minus RIDL. l.c. 244, p.p. Perennial. Stems erect, rather stout to stout, trigonous, triquetrous just below the inflorescence, leafy, smooth, several-noded, 50-200 cm by 4-8 mm. Leaves rigid, flat, gradually acuminate, the cauline ones often overtopping the stem, %—2)4 cm wide; margins scaberulous; ligule a membranous rim, or absent in the upper leaves; lower sheaths shining fuscous to castaneous. Inflorescence terminal, um- belliform, copiously branched, compound to supra- decompound, usually dense, 10-25 cm_ across. Involucral bracts 3-5, leaf-like, overtopping the inflorescence, the lowest up to 50 cm. Primary rays several, divaricate, very unequal, smooth, up to 10 cm. Spikelets in clusters of 4-10, ovoid to oblong- ovoid, terete, obtuse, rarely acutish, very densely many-flowered, brown, 2/4-6 by 2-3 mm. Glumes membranous, tightly appressed, hardly keeled, ovate to almost orbicular, obtuse, submucronulate, 1- nerved, 1/4-2% by 1-14 mm. Bristles 0—3(-6), delicate, sparsely antrorsely scaberulous in the upper half. Stamens 2-3; anthers oblong-linear, %%-1 mm. Style glabrous; stigmas 2, rarely in some flowers 3. Nut plano-convex or unequally biconvex, strongly dorsiventrally compressed, obovate, mi- nutely apiculate, smooth, pale brown, 34-1 by “-% mm. Distr. From the NW. Himalaya through S. Asia to S. China, Formosa, the Ryu Kyu Is. and Bonin Is.; in Malesia: throughout Sumatra, W. Java (Mt Gedeh-Pangerango; near Talun between Mt Papan- dajan and Mt Kantjana), Philippines (Luzon, Min- danao), Borneo (Mt Kinabalu; Sabah: Mt Trusmadi), Celebes, Moluccas (Ternate), New Guinea. Ecol. In open wet places, in thickets and forests, on banks of streams, often in large clumps, chiefly between 800 and 2000 m; in New Guinea up to 2650 m, but also collected at 30 m (PULLE 152). Vern. Riat séla, Minahasa; Philip.: dadtan, Tag., sagari, Bag., sanganga, Klg., siilak, Ig. Note. Very variable as to size of spikelets and glumes, and number of bristles. According to CLARKE the plants are often stoloniferous when growing in swamps, and their leaf-sheaths occasion- ally perforated by descending aerial stolons (like in S. radicans SCHK.). 502 FLORA MALESIANA [ser. I, vol. 7° 4. Section Trichophorum (Pers.) A. Gray, Man. Bot. ed. 5 (1867) 565. — Trichophorum Pers. Syn. 1 (1805) 69; Nes, Linnaea 9 (1834) 293. — Sect. Sylvatici CLARKE, FI. Br. Ind. 6 (1893) 661, p.p. — Sect. Seidlia CLARKE, Kew Bull. add. ser. 8 (1908) 113, p.~. Type species: S. cyperinus (L.) KUNTH. 5. Scirpus wichurai Boeck. Linnaea 36 (1870) 729: Ouwi, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 107; RAYMOND, Nat. Canad. 84 (1957) 115.—S. eriophorum (non MicHx) CLARKE, Fl. Br. Ind. 6 (1893) 661. — S. asiaticus BEETLE, Am. J. Bot. 33 (1946) 662. Perennial: rhizome short, without stolons. Stems erect, rather stout to stout, rigid, obtusely trigonous to subterete, leafy, smooth, 5—9-noded with brownish nodes, 100-150 cm by 5—8 mm. Leaves shorter than the stems, basal and cauline, rather rigid, flat, gradually acuminate, scabrous on the margins, 5-20 mm wide; ligule a membranous rim; lower sheaths brownish. Inflorescence consisting of 1—4 anthelas, the terminal one large, decompound or supradecompound, up to 25 by 20cm, the lateral ones when present axillary, much smaller. Involucral bracts 2—3, similar to the leaves, often overtopping the inflorescence. Primary rays scaberulous at the top, up to 15 cm, secondary ones and peduncles of the spikelets very scabrous. Spikelets very numerous, solitary or partly in clusters of 2-4, ellipsoid to oblong-ellipsoid, terete, obtuse, rather densely many- flowered, 3-6 by 2—3mm. Glumes membranous, appressed, not keeled, ovate, rufous, acute or minutely mucronulate, 1-nerved, 1/4—2 by 1-14 mm. Bristles 6, delicate, strongly elongated, curled and entangled after anthesis, smooth below, antrorsely scaberulous near the top. Stamens 1—2; anthers linear, c. 1 mm. Style glabrous, c. mm; stigmas 3, papillose. Nut compressed-trigonous, elliptic or slightly obo- vate, with a distinct c. 4 mm long beak, smooth, stramineous to pale brown, 1—1/4 (beak included) by 0.6 mm. Distr. India (E. Himalaya, Khasia, Assam), Annam, S. China, Korea, Japan, in Malesia only in Sumatra: Atjeh (Peuet Sago, Bur ni Gérédong), West Coast Res. (Mt Kerintji). Ecol. Swampy places in forests, on river banks, 1700-2200 m. Note. The Malesian specimens perfectly agree with the type collection and additional materials from India. Scirpus wichurai belongs to an intricate group of mutually closely related taxa, which has another centre of development in North America. It comes near to the American S. cyperinus (L.) KUNTH and S. rubricosus FERN. (= S. eriophorum Micux, nom. illeg.). It differs by the ovate (not oblong-ovate) glumes, the shorter, somewhat firmer bristles scabrid in the upper part, and the larger nuts. Possibly these differences are too trifling to warrant specific distinctness, the more so as some Japanese specimens show a clear approach to the American taxa in having narrower glumes, and bristles only denticulate at the very top. Also OHwi refers the Japanese plants to S. wichurai. The question whether all the species distinguished can be upheld, or should be considered racial differentiations of one linneont can only be solved by comparative study of extensive materials from the whole area of the group. 5. Section Eleogiton (Link) Pax in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 111. — Eleogiton Linx, Hort. Berol. 1 (1827) 284. — Sect. Monostachyi *Fluitantes CLARKE, Kew Bull. add. ser. 8 (1908) 111. — Sect. Fluitantes CHERM. Fl. Madag. fam. 29 (1937) 142, descr. gall. Type species: S. fluitans L. 6. Scirpus fluitans LINNE, Sp. Pl. 1 (1753) 48; BOoECK. Linnaea 36 (1870) 485; BENTH. FI. Austr. 7 (1878) 325; CLARKE, FI. Br. Ind. 6 (1893) 653; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 9; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 458; STEEN. Mt FI. Java (1972) t. 14:16.— Isolepis fluitans R.BR. Prod. (1810) 221. — Eleogiton fluitans Link, Hort. Berol. 1 (1827) 284. — Eleogiton curvulus Nees in Wight, Contr. Bot. Ind. (1834) 110. — Isolepis curvula KUNTH, En. 2 (1837) 189; ZOLL. Syst. Verz. 1 (1854) 62 (‘curvata’ err. typ.). — Eleo- giton fluitans var. fasciculata MiQ. Fl. Ind. Bat. 3 (1856) 313, incl. f. curvula et f. tenerior. Perennial. Stems weak, filiform, fascicled, Ae 1 mm thick, sympodially branched (each superposed branch terminated by a spikelet), floating and then up to 40cm long, or in terrestrial forms much shorter, creeping or procumbent, and often forming dense cushion-like tufts. Leaves narrowly linear, almost setaceous, 3—4-nerved, rather acute, 1-7 cm by 4-1 mm. Peduncles pseudolateral, up to 12 cm long, each bearing a single, terminal, ebracteate, ovoid to oblong, 5—10-flowered, 3-6 mm _ long, 2-3 mm wide spikelet. Glumes membranous, ap- pressed, concave, ovate to broadly ovate, obtusish, muticous, faintly many-nerved, 2-2/4 by 1/4 mm; margins hyaline; sides often purplish. Bristles none. Stamens 2—3; anthers oblong, 4-1 mm. Stigmas 2. Nut planoconvex, elliptic or slightly obovate, sharply angled, smooth, pale brown, shortly apiculate, Iy-14 by 0.9-1 mm. Distr. Widely distributed throughout the Old 1974] CYPERACEAE (Kern) 503 World, from Europe to Australia, in Malesia: W. Java (Mts Gedeh-Pangerango and Papandajan), Central Java (Diéng plateau), E. Java (Mts Lawu, Argopuro, Tengger, and Smeru, Jang plateau). Ecol. In swampy localities, along streams, or submerged in shallows, often abundant, 1600 3200 m. Vern. Reubeu, Md. Note. In the Javan plants the number of stamens is usually 2, only some flowers are triandrous. The European plants are constantly triandrous; they have usually still narrower, 3-nerved leaves, with less distinct cross-nerves. 7. Scirpus beccarii Boeck. Bot. Jahrb. 7 (1886) 275; KERN, Blumea 8 (1955) 161. — Fig. 24. Very near to S. fluitans. Spikelets solitary or 2 together, 6—7 by 3 mm, subtended by an erect bract as though continuing the peduncle, as long as or somewhat longer than the spikelet(s), up to 1/4 cm, auricled at the base. Stamens 3. Style 2-cleft, incom- pletely 3-cleft, or 3-cleft (on the same specimen). Nut planoconvex, or with a raised dorsal angle, or sharply triquetrous, slightly larger than in S. fluitans (134-2 by 1-1, mm), and longer-beaked. Distr. Malesia: Sumatra (Atjeh: Mt Losir, Mt Kemiri; Ophir Distr.: Mt Talamau; Mt Singgalang). Ecol. In tufts along the shore of lakes, in and along brooklets, 2750—3300 m. Note. BOECKELER placed the species in the affinity of S. varians BoEcK. (according to Ind. Kew. = S. inundatus Por.) and S. supinus L., presumably on account of the pseudolateral inflorescence. Its true relationship, however, is clearly with S. fluitans L. In Sumatra S. fluitans is apparently absent and replaced by S. beccarii. 8. Scirpus crassiusculus (HOOK. /.) BENTH. Fl. Austr. 7 (1878) 326; KUK. Bot. Jahrb. 69 (1938) 258; S. T. BLAKE, J. Arn. Arb. 35 (1954) 205.— Isolepis crassiuscula HooK. f. Fl. Tasm. 2 (1860) 86, t. 143. — S. fluitans (non L.) CLARKE, Kew Bull. (1899) 113°: VALCK. Sur. Nova Guinea 8 (1912) 704?; HOoGL. Blumea, Suppl. 4 (1958) 235.— S. pseudo-fluitans MAKINO, Bot. Mag. Tokyo 19 (1905) 28; Onw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 100. — S. fluitans L. ssp. pseudo-fluitans KOYAMA, J. Fac. Sc. Un. Tokyo III, 7 (1958) 326. Perennial. Stems tufted, erect, or creeping and rooting at the nodes, or submerged and elongated, much coarser than in the closely related S$. fluitans. Flowering branches 5—15cm long. Leaves linear, longer and usually broader (1-2 mm) than in S. fluitans. Spikelet ovoid, terete, acute, c. 15-flowered., 5-8 by 3-4mm. G/umes membranous, appressed, with somewhat prominent keel, oblong-ovate to oblong, obtusish, muticous, many-nerved, with pur- plish margins, 34-4 by 1/4-2 mm. Bristles none. Stamens 3; anthers linear, c. 1mm; connective shortly produced, smooth. Style c. 1/4 mm; stigmas 2, about as long as the style. Nut strongly compressed, slightly biconvex in the centre, thinner towards the margin and there thickened into an obtuse edge. elliptic, distinctly apiculate, brown, 134-2 by c. 1 mm. Distr. SE. Australia, Tasmania, New Zealand; also known from a single locality in Japan (Honshu: Iwaki, Kanayama); in Malesia: New Guinea (W. New Guinea: Lake Habbema, Mt Wilhelmina; NE. New Guinea: Morobe Distr., Mt Sarawaket; Papua: Central Div., Mt Albert Edward; Eastern Highlands: Mt Wilhelm; Milne Bay Distr.: Duiri; Mt Giluwe, Sugarloaf complex). Ecol. On sand bars or submerged in slow-moving streams, in shallows of alpine tarns, on wet peaty soil, in dense patches, 1800-3900 m. Vern. Iwarud, kasump, Mindi lang. Fig. 24. Scirpus beccarii BOECK. a. Habit, x }4, b. spikelet, c. bract, both x 5, d. glume, e. flower, f. style, g. nut, all x 10 (a—~g DocTeRS VAN LEEUWEN 3977). 504 FLORA MALESIANA [ser. I, vol. 7° 6. Section Confervoidei CHERM. FI. Madag. fam. 29 (1937) 143. — Websteria S. H. WriGurt, Bull. Torr. Bot. Club 14 (1887) 135. Type species: S. confervoides Por. B.C Stone del. 971 Fig. 25. Scirpus confervoides Poir. a. Habit, x %, b. spikelet, c. glume, d. deflorate flower, all x 5 (after STONE). 1974] CYPERACEAE (Kern) 505 9. Scirpus confervoides Pork. in Lamk, Encycl. 6 (1804) 755; KUNTH, En. 2 (1837) 173, p.p.; BOECK. Linnaea 36 (1870) 487; CHERM. Fl. Madag. fam. 29 (1937) 143; BEETLE, N. Am. FI. 18 (1947) 496; Stone, Mal. Nat. J. 24 (1971) 91, f. 1. — Baeothryon confervoides A. DietR. Sp. Pl. 2 (1833) 94. — Schoenus confervoides WILLD. ex KUNTH, En. 2 (1837) 173, in syn. — Eleocharis? confervoides Mig. FI. Ind. Bat. 3 (1856) 303, guoad bhasion.— S. ruppioides Tuw. ex Sauv. Fl. Cub. (1871) 80, in obs. —S. submersus SAUV. Anal. Acad. Ci. Habana 8 (1871) 79: Fl. Cub. (1873) 175; CLARKE, FI. Br. Ind. 6 (1893) 653; in Dur. & Schinz, Consp. FI. Afr. 5 (1895) 631; in Urban, Symb. Antill. 2 (1900) 91. — Rhynchospora ruppioides BENTH. in Hook. Ic. Pl. 14 (1881) 31, t. 1344. — Websteria limnophila S. H. WriGuHT, Bull. Torr. Bot. Club 14 (1887) 135; TRIMEN in Hook. f. Handb. Fl. Ceylon 5 (1900) 78.— Websteria sub- mersa BRITTEN, Bull. Torr. Bot. Club 15 (1888) 99; SUESSENG. Bot. Jahrb. 73 (1943) 122; LEON, FI. Cuba | (1946) 204; EiTen, Phytologia 20 (1970) 276. — Fig. 25. Submerged aquatic rooting in the mud, flaccid, tassel-like when lifted out of the water, with filiform branched stems bearing at the nodes pseudo-whorled fascicles of numerous sterile, leaf-like side-stems. Stems terete, smooth, less than 14 mm thick; lower internodes elongate, upper shorter; side-stems ultra- capillary, 244-10 cm long, with a tubular, short, hyaline sheath at the base. Spikelets solitary, on filiform peduncles from amongst the sterile stems, oblong-lanceolate, 1(—2)-flowered, 6-12 mm long; peduncles emergent from the water, smooth, 4- fistular, 114-30 cm long, the base often rooting and enclosed in a hyaline sheath. G/umes 2, membranous, erect, narrowly lanceolate, acute, green with hyaline margins, 7-10 mm long, the lower one 3-nerved, empty, the upper one longer, 1-nerved, with a bisexual flower. Bristles 6-11, very delicate, re- trorsely barbed, yellow. Stamens 3; anthers linear, with shortly produced connective. Style very slender, halfway bifid, the stigmas filiform. Nut biconvex, broadly obovate, smooth, long-beaked, 3-3/4 mm long (beak included), pale brown. Distr. Subtropical and tropical America (Florida, Guatamala, Cuba, Guyana, Paraguay), tropical Africa, Madagascar, Ceylon; in Malesia: Malay Peninsula (Trengganu: Kp. Jambu _ Bongkok; Pahang: Tasek Bera); generally local and rare. Not known from Java; the records for Java and Surinam refer respectively to Eleocharis retroflexa (see p. 534) and Eleocharis flaccida URB. (cf. UITTIEN in Fl. Surinam 1, 1934, 111). Ecol. Oligotroph lakes in lowland swamps, in Malaya associated with Pandanus helicopus and Lepironia articulata. 7. Section Baeothryon (A. Dietr.) B. & H. Gen. Pl. 3 (1883) 1050. — Baeothryon A. Dietr. Sp. PI. 2 (1833) 89. — Sect. Monostachyi **Caespitosi CLARKE, Kew Bull. add. ser. 8 (1908) 111, p.p. — Sect. Anthelophorum Ouw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 95. — Sect. Paucispicati BEETLE, Am. J. Bot. 33 (1946) 664. Type species: S. cespitosus L. 10. Scirpus subcapitatus THwaites, En. Pl. Zeyl. (1864) 351; Boeck. Linnaea 36 (1870) 704; CLARKE, Fl. Br. Ind. 6 (1893) 661; Ill. Cyp. (1909) t. 51, f. 1-2; KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 301. incl. f. rigidus KOK. et var. triangularis KUK.; in Fedde, Rep. 53 (1944) 101; BEETLE, Am. J. Bot. 33 (1946) 665; Kern, Reinwardtia 4 (1956) 89; RAYMOND. Nat. Canad. 84 (1957) 142. — S. cespitosus (non L.) F.v.M. Trans. R. Soc. Vict. | (1889) 35. — S. clarkei Starr, Trans. Linn. Soc. Bot. 4 (1894) 244; RIDL. J. Str. Br. R. As. Soc. n. 46 (1906) 224; J. Fed. Mal. St. Mus. 6 (1915) 192: Starr in Gibbs, J. Linn. Soc. Bot. 42 (1914) 174; BeeTLe, Am. J. Bot. 33 (1946) 664; S. T. BLAKE, J. Arn. Arb. 35 (1954) 206. — S. pakapakensis STAPF in Gibbs, J. Linn. Soc. Bot. 42 (1914) 174. — S. clarkei var. pakapakensis BEETLE, Am. J. Bot. 33 (1946) 665. — Fig. 26-28. Perennial; glabrous. Rhizome woody, very shortly creeping. Stems slender, erect, rigid, densely to very densely tufted, terete to more or less distinctly triangular, grooved, smooth, or obscurely scaberul- ous just below the inflorescence, 10—75cm_ by Y-l% mm, the base clothed with tight brown sheaths bearing an up to 2 cm long (usually much shorter) smooth or scabrid mucro; orifice of the sheaths obliquely truncate, with a hyaline border. Inflorescence terminal, consisting of 1—5 spikelets, when |-spiculate often ebracteate, otherwise the first spikelet sessile, the others on short smooth peduncles, and with an involucral bract sheathing the stem, 5—10 mm long, with a smooth or scabrid, up to 4 mm long mucro. Spikelets oblong-lanceolate, terete, acute, 5—10-flowered, 5-15 by c. 2mm. Glumes membranous, oblong-lanceolate, slightly keeled, the outermost obtusish, the inner more acuminate, ferrugineous-brown, (2/4—)314—5 by L4- 24% mm, with 3 yellowish nerves and hyaline margins. Bristles 6, very slender, flexuous, scabrid near the top, much longer than the nut, (3—)5—7 mm. Stamens 3; anthers 1-3 mm long, apiculate. Sry/e glabrous; stigmas 3, papillose. Nut compressed-trigonous, elliptic to oblong, apiculate, smooth, fuscous, 134- 2/4 by 0.6—1 mm; epidermal cells minute, rounded to hexagonal. Distr. India, Thailand, Ceylon, Annam, S. China, Formosa, in Malesia: N. and Central Sumatra, Malay Peninsula (Trengganu: Padang; Pahang: Mt Tahan), N. Borneo (Mt Kinabalu), New Guinea (Papua: Mt Albert Edward, Owen Stanley Range, Mt Wilhelm, Mt Knutsford; Sugarloaf complex: HOOGLAND & SCHODDE 7108); two subspecies in Luzon and Celebes. Ecol. In open boggy, sandy or rocky places (grassland slopes, forest glades, banks of streams, mountain heaths), often dominant, 1200—4000 m. Fig. 27-28. [ser. I, vall@e FLORA MALESIANA 506 , x %, b. involuc nut with style and stigmas (a—e BUNNEMEUJER 2908). ral bract, c. glume, d. perianth bristle, e. immature rpus subcapitatus THw. a. Habit Fig. 26. Sci CYPERACEAE (Kern) 507 Fig. 27. Scirpus subcapitatus tufts (dark) in mountain heaths (b/ang) on Mt Kemiri at c. 3300 m (Gajo Lands, N. Sumatra) (photogr. VAN STEENIS, 1937). Notes. I fail to trace dividing lines between S. subcapitatus s. str., S. clarkei (said to differ by the very slender habit, the more advanced development of the lamina of the leaves, the solitary spikelets, and the more distinctly barbellate perianth-bristles), and S. pakapakensis (differing from S. subcapitatus by the loose inflorescence and the more distinctly barbellate bristles). More distinct geographical races are: ssp. pulogensis (MERR.) KERN, Reinwardtia 4 (1956) 90. — S. pulogensis MERR. Philip. J. Sc. 5 (1910) Bot. 333; En. Philip. 1 (1923) 118. Stems very slender, |-spiculate. Perianth-bristles smooth, about as long as the nut. Distr. Malesia: Philippines: Luzon (Benguet, Mt Pulog). ~ Ecol. In very damp ravines and on open grassy slopes, 2400-2700 m. ssp. celebicus KERN, Reinwardtia 4 (1956) 90, f. |. — Fig. 29. Stems very slender, l-spiculate. Spikelet very small, 3-5 by 1-2 mm. Glumes ovate or broadly ovate, 134-2 by 14-124 mm. Perianth-bristles distinctly papillose, 2-3 mm long, unequal, the outer ones flat, straight or slightly flexuous, the inner ones filiform, strongly flexuous. Nut 114-1)4 by 0.75- 0.9 mm. Distr. Malesia: SW. Celebes, Subdiv. Enrekang (Latimodjong Mts; B. Rante Mario, Masimbollong, Pokapindjang). Ecol. Open moist localities, 2700—3000 m. 508 FLORA MALESIANA [ser. I, vol. 7° 8. Section Schoenoplectus Rcup. Ic. Fl. Germ. 8 (1846) 40 ex B. & H. Gen. Pl. 3 (1883) 1051. — Pterolepis SCHRAD. Gott. Gel. Anz. (1821) 2071.— Schoenoplectus PALLA, Sitz. Ber. Zool. Bot. Ges. Wien 37 (1888) 49; Bot. Jahrb. 10 (1888) 298. — Sect. Scirpus (proper) CLARKE, FI. Br. Ind. 6 (1893) 655, p.p. — Sect. Euscirpus CLARKE, Kew Bull. add. ser. 8 (1908) 112, p.p. — Sect. Littorales CHERM. Fl. Madag. fam. 29 (1937) 155, descr. gall. — Sect. Mucronati CHERM. l.c. 151, descr. gall. — Sect. Lacustres BEETLE, Am. J. Bot. 28 (1941) 691. — Sect. Pterolepis BEETLE, N. Am. FI. 18 (1947) 502. — Sect. Actaeogeton RCHB. Fl. Germ. Excurs. 1 (1830) 78; BEETLE, N. Am. FI. 18 (1947) 498. Type species: S. lacustris L. Fig. 28. Tuft of Centrolepis fascicularis LABILL. invaded by Scirpus subcapitatus THw. and Gentiana sp.(one flower) on Mt Losir(Gajo Lands, N. Sumatra), c. 3250 m (photogr. VAN STEENIS, 1937). 11. Scirpus lacustris LINNE, Sp. Pl. 1 (1753) 48. In Malesia only: ssp. validus (VAHL) KoyAMaA, Can. J. Bot. 40 (1962) 927. — S. validus VAHL, En. 2 (1806) 268; BEETLE, Am. J. Bot. 28 (1941) 693 (quoad specim. philipp. p.p. min.; MERRILL 1792); S. T. BLAKE, Trans. R. Soc. S. Austr. 67 (1943) 54; J. Arn. Arb. 35 (1954) 205. — S. triqueter var. segregatus CLARKE, FI. Br. Ind. 6 (1893) 658, saltem quoad specim. nov.-guin.; VaALCK. SuR. Nova Guinea 8 (1912) 704.—S. litoralis (non SCHRAD.) K.ScuH. & LAuT. Fl. Schutzgeb. (1901) 195, p.p. — S. lacustris (non L.) MerR. Philip. J. Sc. 5 (1910) Bot. 172; En. Philip..1 (1923). p.p. — S. tabernaemontani (non GMEL.) KUK. Bot. Jahrb. 59 (1924) 51; OHwi, Bot. Mag. Tokyo 56 (1942) 203; RAYMOND, Nat. Canad. 84 (1957) 139. — S. lacustris var. validus KOK. in Fedde, Rep. 23 (1926) 200. Perennial. Rhizome stout, horizontally creeping, scaly. Stems rather approximate, stout, erect, terete or nearly so, soft and easily compressed, smooth, 44-2 m by /4-2 cm at the base. Leaves reduced to the sheathing bases, rarely the uppermost with an up to 10 cm long blade. Inflorescence pseudolateral, compound or subdecompound, consisting of many spikelets, 5—10(—15) cm long. Involucral bract erect, shorter than the inflorescence, 2—S cm. Primary rays rather stiff, scabrous on the edges, in the upper part up to 5cm; secondary rays scabrous, up to 2 cm. Spikelets solitary or partly in clusters of 2—3, ovoid to oblong-ovoid, rather acute, densely many- flowered, fulvous to deep brown, 5—10 by 4—5 mm. Glumes scarious, appressed, ovate to oblong-ovate, with prominent midrib excurrent into a short mucro, more or less notched at the top, more or less ciliate on the margins, 3-4 by 2-2/4 mm. Bristles 5—6, slightly longer than the nut, retrorsely scabrous. Stamens 3; anthers linear, 1/4—2 mm, the connective produced into a triangular-ovate bearded appendage. Style bifid. Nut planoconvex (with low rounded back), obovate, apiculate, smooth, greyish black, c. 2 by 1)4-1)4 mm. Distr. Widely distributed in the countries border- ing the Pacific Ocean, in Malesia: Philippines (Luzon: Cagayan, Zambales, Benguet), New Guinea. Ecol. In open marshes, open sandy foreshores, freshwater swamps, 0—1900 m. Uses. In N. Luzon used for weaving mats. Vern. Philip.: tikér, Ilk. Note. Scirpus lacustris in the wider sense is a highly polymorphous cosmopolitan species. The 1974] Cy PERACEAE (Kern) Jeo ie A h Mi I i ) NYA, DE Ni // bit, «34,6. spikelet, x 6, c—e. lower empty Fig. 29. Scirpu ssp. celebicus KERN. a. Habit, x 4 - fl h th bristles, all x 12, i. perianth bristles, enlarged (a—i ie s subcapitatus THwW. glumes, f. fertile glume, g. deflorate flower, 510 typical subspecies is restricted to Europe. The circum-Pacific ssp. validus differs from it by the constantly bifid styles (trifid in ssp. lacustris) and some minor very inconstant characters. Because of its bifid styles ssp. validus approaches the European- W. Asian ssp. glaucus (SMITH) HART. (= S. taber- naemontani GMEL.), to which KUKENTHAL and OHwI refer the Malesian plants. Typical ssp. glaucus has smaller stems (but very stout specimens occur!) and densely gland-dotted glumes. According to BLAKE (1943) the rays of the inflorescence in ssp. validus are more distinctly scabrous than in ssp. glaucus, but this is incorrect. On account of the many intergrading specimens BEETLE (1941) treated ssp. glaucus as a variety of S. lacustris, viz S. lacustris var. tabernaemontani (GMEL.) DOELL, whereas in his opinion S. validus is a clearly distinct species. In this connection it may be remarked that in Europe ssp. lacustris and ssp. glaucus frequently occur together and hybrids of the two are formed, whereas ssp. validus inhabits an entirely separated area. The differentiating characters between ssp. lacustris and ssp. validus are not more important than those between ssp. lacustris and ssp. glaucus. They are too trifling to justify specific distinction. 12. Scirpus litoralis SCHRAD. Fl. Germ. 1 (1806) 142, t. 5, f. 7; KUNTH, En. 2 (1837) 166; STEUD. Syn. 2 (1855) 86; BENTH. FI. Austr. 7 (1878) 334; K.ScH. & HOLtr. Fl. Kais. Wilhelm Land (1889) 25; CLARKE, Fl. Br. Ind. 6 (1893) 659; Ill. Cyp. (1909) t. 50; VALCK. SuR. Nova Guinea 8 (1912) 705; BACK. Onkr. Suiker. (1928) 150, t. 152; STEN. Arch. Hydrobiol. Suppl. 11 (1932) 289, f. 73; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 10; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 459. — S. subulatus VAHL, En. 2 (1806) 268; KUNTH, En. 2 (1837) 165; Mia. FI. Ind. Bat. 3 (1856) 306; Boeck. Linnaea 36 (1870) 715; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 122.—S. thermalis Tras. in Batt. & Trab. Fl. Alg. (Monoc.) ed. 2 (1895) 99. — S. lacustris (non L.) Mere. En. Phil. 1 (1923) 117, p.p.—S. validus (non VAHL) BEETLE, Am. J. Bot. 28 (1941) 693, p.p. (quoad CURRAN 17290 & McGREGOR 41439). Perennial. Rhizome short; sometimes slender stolons present. Stems rather stout to stout, erect, terete but obtusely trigonous just below the inflo- rescence (see note), glaucous, 60—150 cm by 3-10 mm. Leaves reduced to bladeless or shortly laminate sheaths. Inflorescence pseudolateral, simple or com- pound, few- to many-spiculate, 2-8 cm long; rays slender, smooth, or scaberulous at the top. Involucral bract erect, continuous with the stem, rigid, chan- nelled along the inner side to triquetrous, 2- 5(—10) cm. Spikelets solitary, ovoid to oblong-ovoid, terete, acutish, densely many-flowered, rusty or brown, 8-15 by 3—4 mm. Glumes scarious, appressed, elliptic to oblong, obtuse or slightly notched, minutely ciliolate at the apex, otherwise glabrous, 3)4-4 by c. 2 mm; midrib prominent, excurrent into a short but distinct 4-4 mm long mucro; sides nerveless. H ypogynous scales usually 4 (3—5), ligulate- spatulate, plumosely fringed in the upper part with antrorse moniliform hairs, ferrugineous, about as long as the nut. Stamens (2—)3; anthers 114-2 mm, the connective with a fimbriate appendage. Style deeply bifid. Nut unequally biconvex, strongly FLORA MALESIANA [ser. I, vol. 7° dorsiventrally compressed, elliptic to obovate, api- culate, smooth, castaneous to blackish, c. 2 by V4A-14% mm. Distr. From the Mediterranean region through S. Asia to Australia, Africa, in Malesia very rare: Java (here and there near the sea, in W. Java also inland near Garut), Madura, Kangean, Lesser Sunda Is. (Bali: Batur Lake), Philippines (Luzon: Panga- sinan Province, MCGreGcor BS 41439, CLEMENS 18104), NE. New Guinea. Ecol. In brackish swampy places, saline pools near the sea, sometimes abundant; at Tarogong (Mt Guntur) at 800m in water originating from hot springs, along Batur Lake (Bali) at 1000 m. Uses. In W. Java (Indramaju) used for making mats. Vern. Endong, péndjalinan, J. Notes. In appearance very similar to S. lacustris, but usually with looser, less compound inflorescence, and easily recognized by the flat, plumose hypo- gynous scales. In the Mediterranean plants and also in those from the Middle East, China and Mongolia (S. litoralis s.s.) the stems are sharply triangular throughout; however, in Scirpus this character does not warrant specific separation. S. subulatus was based on speci- mens from the Nicobars identical with the Malesian ones, to which the above description only refers. It may be distinguished as S. litoralis var. subulatus (VAHL) CuHiov. (Distr.: Tropical Africa, S. Africa, Madagascar, India, Malesia, Micronesia). See also TOWNSEND, Kew Bull. 15 (1961) 415-417. Also the African S. pterolepis KUNTH with more copiously branched inflorescence and longer, oblong spikelets is possibly not specifically distinct from S. litoralis. 13. Scirpus mucronatus LINNE, Sp. Pl. 1 (1753) 50; DecngE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 361; Descr. Herb. Timor. (1835) 33, excl. var. «; KUNTH, En. 2 (1837) 161; Mia. Fl. Ind. Bat. 3 (1856) 304; Boeck. Linnaea 36 (1870) 703; BENTH. FI. Austr. 7 (1878) 332; F.-VILL. Nov. App. (1882) 308; CLARKE, FI. Br. Ind. 6 (1893) 657; Philip. J. Sc. 2 (1907) Bot. 100; RipL. Mat. Fl. Mal. Pen. 3 (1907) 79; CaMus, Fl. Gén. I.-C. 7 (1912) 134; VALCK. Sur. Nova Guinea 8 (1912) 704; Merr. En. Philip. 1 (1923) 118; KUk. Bot. Jahrb. 59 (1924) 51; ibid. 69 (1938) 259; RipL. Fl. Mal. Pen. 5 (1925) 161; BACK. Onkr. Suiker. (1928) 149, t. 151; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 10; S. T. BLAKE, J. Arn. Arb. 35 (1954) 204; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 59, incl. var. planoconvexus KOYAMA; RAYMOND, Nat. Canad. 84 (1957) 129; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 459; STEEN. Mt FI. Java (1972) t. 14:17. — S. triangulatus Roxs. Fl. Ind. 1 (1820) 219; ed. Carey 1 (1832) 217; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 60.— S. acutus PRESL, Rel. Haenk. 1 (1828) 192, non MUHL. — S. preslii Dietr. Sp. Pl. 2 (1833) 175 (‘presslii’); KUNTH, En. 2 (1837) 161; Mig. FI. Ind. Bat. 3 (1856) 305; F.-ViLL. Nov. App. (1882) 308 (‘preslei’). — S. javanus NEES in Wight, Contr. Bot. Ind. (1834) 112; KUNTH, En. 2 (1837) 161; Mor. Syst. Verz. (1846) 97; Mig. Fl. Ind. Bat. 3 (1856) 305. — S. sundanus Mia. FI. Ind. Bat. 3 (1856) 304; in De Vriese, Pl. Ind. Bat. Or. (1857) 140. — Fig. 30. Perennial. Rhizome very short. Stems rather stout, stiff, erect, tufted, triquetrous with more or less 1974] Cy PERACEAE (Kern) 511 Fig. 30. Scirpus mucronatus L. a. Habit, x 74, b. spikelet, x 3, c. glume, d. deflorate flower, e. young stamen, f. style, all x 13 (a—f LORZING 6723). 542 concave sides, smooth, Y-1 m by (2—)3—8 mm. Leaves reduced to 1-2 membranous, stramineous to brownish, bladeless sheaths. Inflorescence pseudo- lateral, capitate (sometimes proliferous, with some slender branches bearing one or more spikelets), with (2—)4—-25 spikelets, up to 4cm across. Involucral bract at first erect, looking like a continuation of the stem, finally often patent to reflexed, triquetrous like the stem, 1-10 cm. Spikelets sessile, ovoid to oblong- ovoid, terete, rather acute, densely many-flowered, stramineous to brownish at maturity, 10-20 by 4-6 mm. Glumes firm, tightly appressed, concave, but slightly keeled near the apex, ovate or broadly ovate, acutish, shortly mucronulate, 3/4-4 by c. 2/4mm; midrib prominent, sides many-nerved, yellowish brown, margins minutely ciliolate at the top. Bristles 5—6, stout, unequal, slightly to distinctly longer than the nut, retrorsely scabrid. Stamens 3; anthers linear, 1-174 mm; connective distinctly produced. Style 2—2)4 mm, sparsely fimbriate or papillose; stigmas 3, but often in some flowers 2. Nut strongly dorsiventrally compressed, planocon- vex, or trigonous but the dorsal angle indistinct, broadly obovate, shortly apiculate, scarcely rugulose to smooth, shining black at maturity, 174-2)4 by 14-1 mm. Distr. Warmer parts of the Old World, from S. Europe to Japan, and through S. Asia to Australia, rare in tropical Africa; probably throughout Malesia, but not yet seen from the Moluccas. Ecol. In open wet places, swamps, ditches, pools, margins of lakes, often dominant, or codominant with Leersia hexandra, sometimes in inundated rice-fields, 0—c. 2100 m. Uses. The dried and flattened stems are often used for making cheap but durable mats and bags, and for strings. In the Padang Uplands (Sumatra West Coast) the species is cultivated for this purpose. Vern. Djadjaruman, walini, S, méndongan, J, rumput kumbah, M, baion login, login djau, Sum. E. C., buku ajér, kértjut, kumbuh, mansiang agam, Sum. W. C., abulbugul, Sibil, kumbuéh, mansiro hitam, Minangk., ma’siang, Simalur, bubundelan, Banten, andung, Borneo (Mal.), maufles, ututu, Timor, pusu, Alf. Minah., bimpolu, SW. Celebes; Philip.: bibiran, kanubsuban, Tag., parapipit, Ilk., pulutapit, pupuegan, Bon.; New Guinea: ticomba, Orne lang. Note. The above description refers only to the Malesian plants; S. mucronatus, with its wide range, is very polymorphous. In the European form the style and anthers are shorter and the nuts distinctly rugulose, but the Malesian specimens cannot be separated specifically from them. Very remarkable is: ssp. clemensii KUK. Bot. Jahrb. 69 (1938) 259. — S. clemensii OHW1, Bot. Mag. Tokyo 56 (1942) 203; KUx. Mitt. Thiir. Bot. Ver., N.F. 50 (1943) 13 (‘clemensiae’); S. T. BLAKE, J. Arn. Arb. 35 (1954) 204. Stems terete or very obtusely trigonous. Glumes fuscous to purplish. Distr. Malesia: New Guinea (W. New Guinea: Arfak Mts, Wissel Lake region, Lake Habbema; NE. New Guinea: Morobe Distr., Mt Sarawaket; Mt Sugarloaf; Mt Giluwe). According to Koyama (Willdenowia 5, 1969, 493) S. fohaiensis TANG & WANG (FI. Reip. Pop. Sinic. FLORA MALESIANA [ser. I, vol. 7° 11, 1961, 23, 222, t. 10, f. 6-10) from Yunnan should belong to the same taxon. Ecol. In open swamps, sandy marginal shallows of lakes, tarns, locally often abundant, 1750-3225 m. Note. As additional characters distinguishing this taxon from S. mucronatus s. str., KOKENTHAL (1943) indicated the creeping rhizome, the non- triquetrous bracts, and the obovate nut. I fail to see differences in the rhizome and in the nut; the in- volucral bract is obtusely trigonous to almost terete like the stem (in S. mucronatus s. str. both are tri- quetrous). The colour of the glumes is very variable in S. mucronatus s. str. For these reasons I prefer to maintain the original subspecific rank. Both KUKENTHAL and OHwlI indicate 3 stigmas, but just as in S. mucronatus s. str. digynous flowers often occur with trigynous ones, even in one spikelet. Slender specimens of this subspecies strongly resemble S. juncoides. They may be distinguished by the characters given in the key on p.497. 14. Scirpus juncoides Roxs. Fl. Ind. 1 (1820) 218; ed. Carey 1 (1832) 216; KunTH, En. 2 (1837) 160; ZOLL. Syst. Verz. 2 (1854) 62; Mig. Fl. Ind. Bat. 3 (1856) 303; CHERM. Fl. Madag. fam. 29 (1937) 152; S. T. BLAKE, Proc. R. Soc. Queensl. 62 (1952) 88; RAYMOND, Nat. Canad. 84 (1957) 134; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 57; J. Fac. Sc. Un. Tokyo III, 7 (1958) 310, incl. var. ohwianus (KOYAMA) KoyaMa, I.c. 311; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 459. — S. luzonensis PRESL, Rel. Haenk. 1 (1828) 193; Nees in Wight, Contr. Bot. Ind. (1834) 112; Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 361; Descr. Herb. Timor. (1835) 33; Mig. Fl. Ind. Bat. 3 (1856) 304. — S. junciformis Ness in Wight, Contr. Bot. Ind. (1834) 112.—S. mucronatus var. « DECNE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 361; Descr. Herb. Timor. (1835) 33. — S. timorensis KUNTH, En. 2 (1837) 162; Mia. Fl. Ind. Bat. 3 (1856) 305 (‘timoriensis’). — S. supinus var. elatior Boeck. Linnaea 36 (1870) 700.—S. erectus (non PoiR.) CLARKE, Fl. Br. Ind. 6 (1893) 656; Philip. J. Sc. 2 (1907) Bot. 99; RipL. Mat. FI. Mal. Pen. 3 (1907) 79; CLARKE, Ill. Cyp. (1909) t. 48, f. 11-12; Camus, Fl. Gén. I.-C. 7 (1912) 136; Me_rr. En. Philip. 1 (1923) 117; RipL. Fl. Mal. Pen. 5 (1925) 161; Back. Onkr. Suiker. (1928) 148, t. 149; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 11.—-S. ohwianus Koyama, Bot. Mag. Tokyo 69 (1956) 212, f. 4. Annual. Stems rather slender, erect, tufted, usually subterete, not rarely more or less angular, ribbed, smooth, 15—75(—120)cm by 1-—2(—3)mm. Leaves reduced to 2-3 membranous sheaths obliquely truncate at the top, often with a very small rudimen- tary blade. Inflorescence pseudolateral, capitate, con- sisting of (1—)2—7 spikelets. Involucral bract erect, looking like a continuation of the stem, grooved on the inner side, with a callous tip, always much shorter than the stem, 5-15 cm. Spikelets sessile, ovoid to oblong-ovoid, terete, obtusish, densely many-flowered, stramineous to brownish at maturity, 7-18 (sometimes lengthening to 30) by 5—6mm. Glumes firm, tightly appressed, concave but slightly keeled near the apex, broadly ovate, obtuse, shortly mucronulate, with strong midrib, faintly many- nerved purplish lineolate sides, and margins minutely ciliolate at the top, 34-4 by 3-34 mm. Bristles 1974] CYPERACEAE (Kern) 513 5—6, stout, unequal, shorter than or the longer ones slightly exceeding the nut, retrorsely scabrid, up to 2(-24%) mm. Stamens (2-)3; anthers linear, 1- 1% mm. Style glabrous, 2 mm; stigmas 2, sometimes a short third stigma present. Nut strongly dorsi- ventrally compressed, unequally biconvex (only low- convex on the ventral side), broadly obovate, shortly apiculate, more or less pitted, shining black, c.2 by 14-14 mm. Distr. Madagascar; from India, China, and Japan to Hawaii and tropical Australia; the North American S. purshianus FERN. (= S. debilis PURSH, non LAM), curiously omitted by BEETLE in Fl. N. Am., is hardly different and in all probability not specifically distinct; in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Is. (Lombok, Timor), Borneo, Philippines (Mindoro, Luzon, Polillo, Leyte, Biliran, Mindanao), Celebes, W. New Guinea (Biak Res.: Arupa). Rather common in the western part of the region, much rarer in the eastern part, not yet collected in the Moluccas. Ecol. In open wet localities: swamps, river banks, and particularly in inundated rice-fields, only in fresh water, both in regions with a strong and a feeble dry monsoon, 0—1200 m, rarely up to 2000 m. Uses. Sometimes used as a green-food for cattle; feeding-value rather high. Vern. Kambo-mantjik, M, bawang kladi, njon- joran, J, babawangan, kutjaja, wawalingian, S, paro- paro, Minangk., luluwang, Alas, siwuntuh, Sum. W.C., rumput prut tikus, Mal. Pen., adas-adasan, Banten: Philip.: bitubituinan, gumi, Tag., paratupit, Ikk.; New Guinea: nunu, Kapauku. Note. S. ohwianus KOYAMA according to KOYAMA occurring from W. Japan to S. China and Luzon. I fail to see a demarcation between S. juncoides and S. ohwianus. var. triangulatus (HONDA) OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 115. — S. erectus var. triangulatus HONDA, Bot. Mag. Tokyo 45 (1931) 45. Stems distinctly (3—)5-angled. Distr. Japan, in Malesia: West Java. Note. I have not seen the type of this variety. Ouwi described it as more robust in all its parts, with anthers |.7—2 mm long, and suspected hybrid origin (S. juncoides x preslii?). The Javan specimens are rather robust, distinctly 5-angled, with anthers c. 1’%4 mm long. In herbarium specimens it is often difficult to decide whether in vivo the stems were terete or angular. 15. Scirpus wallichii Nees in Wight, Contr. Bot. Ind. (1834) 112; KuNTH, En. 2 (1837) 160; Steub. Syn. 2 (1855) 84; Onwi, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 113; KERN, Reinwardtia 4 (1956) 90; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 60; RAYMOND, Nat. Canad. 84 (1957) 136.—S. erectus var. wallichii BEETLE, Am. J. Bot. 29 (1942) 654. Closely related to S. juncoides. Stems very slender, 4—5-angular, 10-40 cm_ by 34-1 mm. Inflorescence with 1—2(—3) spikelets. Spikelets oblong-ovoid, acute, greenish, 3—4 mm wide. G/umes membranous, ovate to elliptic, acute, shortly mucronate, finally greyish green, 34-4 by c. 244 mm. Bristles 4—5, distinctly longer than the nut, the outer ones somewhat shorter than the inner ones, the latter c. 3mm. Anthers oblong-linear, 4-34 mm long. Style always bifid. Nut planoconvex (the ventral side flat), somewhat smaller than in S. juncoides, c. 134 by 1)4 mm. Distr. India, Annam, Tonkin, Mergui Arch., Japan, Korea, Formosa, in Malesia apparently very rare: Malay Peninsula (Kedah: Kepala Batas; Malacca: Batu Berendam, Sawar); Philippines (Luzon: Prov. of Bulacan, Manila, Rio Tansa). Ecol. Insufficiently known; the habitats are probably similar to those of S. juncoides as it was often collected together with that species (open wet localities, especially wet rice-fields). 16. Scirpus articulatus LINNE, Sp. Pl. 1 (1753) 47; Boeck. Linnaea 36 (1870) 702; BENTH. FI. Austr. 7 (1878) 331; CLARKE, FI. Br. Ind. 6 (1893) 656; Philip. J. Sc. 2 (1907) Bot. 99; Ill. Cyp. (1909) t. 49, f. 1-2; Camus, Fl. Gén. I.-C. 7 (1912) 133; Merr. En. Philip. 1 (1923) 116; BAck. Onkr. Suiker. (1928) 149, t. 150; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 11; RAYMOND, Nat. Canad. 84 (1957) 132; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 458. — S. incur- vatus ROXB. FI. Ind. | (1820) 217; LLANOs ex F.-VILL. & NAVESs in Blanco, FI. Filip. ed. 3, 4' (1880) 105. Isolepis prolongata Nees in Wight, Contr. Bot. Ind. (1834) 108; KunTH, En. 2 (1837) 199. — Carex glomerata (non THUNB.) BLANCO, FI. Filip. ed. 2 (1845) 24; ed. 3, 1 (1877) 45. — Fig. 31. Fig. 31. Scirpus articulatus L. on swampy ground amidst matted Marsilea crenata in Papua, Central Distr., Kairuku Subdistr.. c. 5 miles NW of Hisiu village (DARBYSHIRE 826). 514 Annual or perennial. Stems erect, densely tufted, terete, fistulose, more or less distinctly transversely septate, smooth, (4—)10—35 cm by 3-6 mm. Leaves reduced to 1—2 bladeless, scarious, obliquely truncate, stramineous to brownish sheaths. Inflorescence pseu- dolateral, capitate, globose, consisting of numerous (up to 30) spikelets, 1-3 cm across. Involucral bract erect, similar to and continuous with the stem, terete, transversely septate, somewhat to much longer than the stem proper, up to 60 cm. Spikelets sessile, ovoid to oblong-ovoid, terete, acutish, densely many- flowered, 8-17 by 4-5 mm. Glumes firm, appressed, concave, with scarcely prominent keel, broadly ovate to suborbicular, rather acute, apiculate, many- nerved, rufous-testaceous, 3/4—5 mm long and wide. Bristles none. Stamens 3; anthers oblong, 4-1 mm. Style 14-2 mm; stigmas 3, shorter than the style. Nut triquetrous with slightly concave sides, obovate, narrowed towards the base, shortly apiculate, con- spicuously to obsoletely transversely wavy-ridged, black, c. 4 by | mm. Distr. From the Mediterranean through Africa and S. Asia to Australia (N. Australia, Queensland), in Malesia: scattered through Java, Bawean I., Lesser Sunda Is. (Bali, Lombok, Timor), Philippines (Luzon, Cebu, Mindoro), SW. Celebes; once col- lected in New Guinea (Papua, Central Distr., Kairuku Subdistr; fig. 31). Ecol. In regions with a pronounced dry season, in open swampy or inundated places, shallow pools, rice-fields, locally sometimes abundant, at low altitude, in Java below 50 m, in Bawean I. at 200 m, in Timor at 450 m. Vern. Méndong, J, bawang-bawang, pakitan, Ba- wean; Philip.: apurau, Pang. Note. The above description refers only to the Malesian specimens of this widely distributed poly- morphous species. NEES distinguished between Iso- lepis prolongata (Potr.) Nees (S. prolongatus Pore. Enc. 6, 1804, 764), with an involucral bract 3—4 times as long as the obsoletely articulated stem, broadly ovate glumes, and transversely rugulose nuts, and /. articulata (L.) NEES, with an involucral bract twice as long as the distinctly articulated stem, ovate glumes, and smooth nuts. Apparently only the former occurs in Malesia. The Indian S. articulatus sensu NEES (S. articulatus var. major Boeck. Linnaea 36, 1870, 702) has often stouter stems and larger nuts (c. 1% by 1/4 mm), but very stout specimens are not rare in Malesia. African plants with slender stems only 2—3 mm thick, obtuse small spikelets (3 by 4mm), small glumes (3 by 2mm), and somewhat smaller (114 by | mm), transversely wavy-ridged nuts repre- sent Isolepis senegalensis HOCHST. ex STEUD. Syn. 2 (1855) 96. I think these three taxa do not deserve specific rank. 17. Scirpus lateriflorus GmeEL. Syst. Veg. 1 (1791) 127; S. T. BLAKE, Proc. R. Soc. Queens]. 62 (1952) 87; KoyaMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 59; KERN, Blumea, Suppl. 4 (1958) 164; in Back. & Bakh. f. Fl. Java 3 (1968) 459. — S. lateralis Retz. Obs. 4 (1786) 12; ibid. 5 (1789) 16; RAYMOND, Nat. Canad. 84 (1957) 132; non Forsk. 1775. — Eleocharis tristachyos Mor. Syst. Verz. (1846) 97, non Scirpus tristachyos Rotts. — Isolepis ambigua STeuD. in Zoll. Syst. Verz. 2 (1854) 62, nom. nud., non STEUD. Syn. 2 (1855) 91. — Isolepis oryzetorum FLORA MALESIANA [ser. I, volive (err. typ. oryectorum) STEUD. Syn. 2 (1855) 96. — S. tristachyos ZOLL. ex STEUD. I.c., in syn., non ROTTB. — Isolepis uninodis (non DELILE) Mia. FI. Ind. Bat. 3 (1856) 308. — Isolepis? juncoides Mia. FI. Ind. Bat. 3 (1856) 312. — S. supinus (non L.) F.—VILL. Nov. App. (1882) 308; CLARKE, Philip. J. Sc. 2 (1907) Bot. 99; RIDL. Mat. Fl. Mal. Pen. 3 (1907) 79; Camus, FI. Gén. I.-C. 7 (1912)) 135, an-f. 17, 67> MERRe Ene Philip. | (1923) 118; RipL. Fl. Mal. Pen. 5 (1925) 161; BAcK. Onkr. Suiker. (1928) 148, t. 148; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 12. —S. supinus var. uninodis CLARKE, FI. Br. Ind. 6 (1893) 656; Philip. J. Sc. 2 (1907) Bot. 99, non Isolepis uninodis DELILE. — S. oryzetorum OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 112. Annual. Stems slender, weak, densely tufted, obtusangular to subterete, smooth, with a node a short distance above the base, 10-35 cm by 4-1 mm. Leaves reduced to 2-3 sheaths slightly widened upwards, obliquely truncate at the apex, often with a small rudimentary, rarely + elongate blade. Axils of the basal sheaths often with a solitary female flower with very long, exserted, trifid style. Terminal inflorescence pseudolateral, with 4—10(—15) spikelets, sometimes capitate, usually with one of the rays developed 1(—4)cm long. Involucral bracts 1-2, the lower erect, similar to and continuous with the stem, always distinctly shorter than the latter, up to 15 cm, the second when present patent, much shorter, up to 5 cm. Spikelets oblong-ovoid, angular, acutish, densely many-flowered, 5— 10(-15) by c. 244 mm. Glumes membranous, appressed, distinctly keeled nearly throughout, ovate, stramineous to ferrugin- eous, often reddish variegated, 2—2'4 by 1%- 2mm; midnerve prominent, sides nerveless, margins hya- line, microscopically ciliolate at the apex. Bristles none. Stamens 3; anthers oblong, 0.4—0.6 mm; connective produced, papillose-bristly at the top. Style 4-34 mm; stigmas 3, about as long as the style. Nut compressed-trigonous, obovate, shortly apiculate, conspicuously transversely wavy-ridged, black, !—1.2 by 0.9-1 mm. Distr. SE. Asia (from India to S. China and Formosa), Australia, in Malesia: Malay Peninsula (Kedah, Wellesley, Kelantan, Trengganu, P. Lang- kawi), W.-E. Java, Kangean Arch., Madura, P. Bawean, Lesser Sunda Is. (Bali, Lombok), Philip- pines (Luzon, Panay). Ecol. In open wet places, edges of swamps, rice- fields, only in fresh water, sometimes abundant, at low altitude (0Q—300 m, rarely up to 1300 m). Vern. Domdoman, sriwit, weko, J. Notes. S. /ateriflorus belongs to a group of closely related species which is greatly in need of revision. The type of S. /ateralis Retz. is from Ceylon (leg. KONIG), ReTzIUs erroneously ascribed a bifid style to it. The species was renamed by GMELIN because RETZIUS’s name is a later homonym of S. lateralis FORSK., a nomen confusum of which the type got lost (“Jn herbario Forskdlaei non aderat” VAHL 1806). S. T. BLAKE (1952) considered S. erectus Por. = Isolepis uninodis DELILE synonymous with S. lateri- florus. However, S. erectus is a clearly distinct African species, with larger glumes, more distinctly bristly connective of the anthers, bifid styles, and larger biconvex nuts (see CHERMEZON, Arch. Bot. 4, Mem. 7, 1931, 26). BLAKE wrongly placed Isolepis? 1974] Cy PERACEAE (Kern) 545 juncoides Mig. in the synonymy of S. juncoides Roxs. It was not based on the latter binomial; MIQUEL’s description and the type-specimen in the Leyden Herbarium prove that it belongs to S. lateriflorus. The latter is often united with the Eurasian S. supinus L., which can be distinguished by its less slender habit, the nodeless stems, the absence of basal flowers, the always solitary in- volucral bract, the always capitate inflorescence, the many-striate glumes, and the smooth appendage of the connective. 9. Section Isolepis (R.BR.) GriseB. Spicil. Fl. Rumel. Bith. 2 (1844) 417; K.ScH. in Engler, Pfi. Welt Ost-Afrikas C (1895) 125.— Isolepis R.Br. Prod. (1810) 221; Linx, Hort. Berol. 1 (1827) 285. — Sect. Cernui CHERM. Fl. Madag. fam. 29 (1937) 145, descr. gall. — Sect. Setacei CHERM. I.c. 144. Type species: S. setaceus L. 18. Scirpus aucklandicus (HooK. f.) Boeck. Linnaea 36 (1870) 491; S. T. BLAKE, Proc. R. Soc. Queensl. 58 (1947) 39; Contr. Queensl. Herb. 8 (1969) 18. — Isolepis aucklandica Hook. f. Fl. Antarct. | (1844) 88, t. 50; SreuD. Syn. 2 (1855) 92; Hook. f. Handb. New Zeal. Fl. (1867) 302. Perennial; forming loose mats. Rhizome very slender, creeping, branched. Stems slender, erect, setaceous, ribbed, smooth, 5-15 cm by 14-14 mm. Leaves narrowly linear, from somewhat shorter than to often overtopping the stems, semiterete, often somewhat canaliculate by the inrolled margins, 3-nerved, obtuse, smooth, c. /4 mm wide; lower sheaths ferrugineous. Inflorescence pseudolateral, consisting of 1(—2) sessile spikelets. Involucral bract similar to the leaves, erect, continuous with the stem, strongly dilated at the base, 1-1/4 cm. Spikelet ovoid, terete, obtusish, several-flowered, c. 3 by 2 mm, partly hidden by the base of the bract. Glumes membranous, ovate, keeled, obtuse, muticous, fer- rugineous, the lowermost often + bract-like but fertile, cucullate at the top, c. 3 mm long, the remain- ing ones 1/£—2 mm long; midnerve distinct, green; sides finely nerved. Brist/les none. Stamens 3; anthers oblong-linear, *4-1 mm; connective distinctly pro- duced. Style 4-14 mm; stigmas 3, much longer than the style. Nut triquetrous with somewhat concave sides and rib-like angles, elliptic-ovate, slightly dorsiventrally compressed, apiculate, smooth, stra- mineous, 1-1/4 by 4-7) mm. Distr. SE. Australia (New South Wales, Victoria, Tasmania), New Zealand and its outlying islands, Amsterdam I.,in Malesia: NE. New Guinea (Western Highlands: Mt Hagen; Eastern Highlands: Mt Wilhelm). Ecol. In tussock grassland at the edge of an alpine swamp, c. 3300 m. 19. Scirpus subtilissimus (Borck.) S. T. BLAKE, Contr. Queens]. Herb. 8 (1969) 20. — Isolepis sub- tilissima BOECK. Flora 41 (1858) 416. — S. inundatus (non Poir.) STAPF, Trans. Linn. Soc. Bot. 4 (1894) 244; Ripv. J. Str. Br. As. Soc. n. 46 (1906) 224: CLARKE, Philip. J. Sc. 2 (1907) Bot. 99. — Cyperus tenellus (non L. f.) CLARKE, /.c. 81. — Schoenoplectus merrillii PALLA in Kneucker, Cyp. et Junc. exsicc. 8 (1911) n. 223; Allg. Bot. Zeitschr. 17 (1912) Beil. 3. — S. merrillii KUK. ex MERR. En. Philip. 1 (1923) 117;S.T. BLAKE, Proc. R. Soc. Queensl. 58 (1947) 38: J. Arn. Arb. 35 (1954) 206. Perennial; forming loose mats. Rhizome filiform, creeping, branched, forming small, irregular, 2—6 mm long tubers. Stems very slender, erect, filiform. terete, smooth, 2-10cm by 4A mm. Leaves filiform, from somewhat shorter than to often overtopping the stems, flat or more or less canalicu- late, 3-nerved (lateral nerves submarginal), with rounded apex, smooth, 4—-)4 mm _ wide; lower sheaths bladeless, transversely truncate, ferrugineous. Inflorescence pseudolateral, capitate, consisting of [—2(—3) sessile spikelets, not rarely proliferous. Involucral bract similar to the leaves, erect, continu- ous with the stem, 4-1/4 cm, at the base with scarious purplish auricles. Spikelets ovoid, terete, acutish, several-flowered, 1/4-3 by 1-14 mm. Glumes membranous, concave, very broadly ovate, obtuse or slightly notched, muticous or scarcely apiculate, 1-1/4 by 34-1 mm; midnerve distinct, green; sides finely nerved, often purplish; keel curved. Bristles none. Stamens |(—2); anthers oblong, YA-)4 mm; connective hardly produced. Style 34-1 mm (the 3 stigmas included; stigmas exceptionally in some of the flowers 2). Nut triquetrous with rib-like angles, elliptic, apiculate, nearly as long as the sub- tending glume, smooth, yellowish to brownish, 0.8—1 by 0.6—0.75 mm. Distr. New Zealand, Australia (Tasmania, Vic- toria, New South Wales, Queensland), in Malesia: N. Borneo (Mt Kinabalu), Philippines (Luzon, Negros, Mindanao), SW. Celebes, New Guinea. Ecol. In marshes, shallow swamps, tussock grassland, on wet banks of alpine streams, on open seepages, associated with mosses, Haloragis mi- crantha R.BR. and Isachne clementis MERR. var. vulcanica (MERR.) JANSEN, 1600-3700 m. Vern. Philip.: sa/aisoi, Bag.; New Guinea: noma, Mendi lang. 20. Scirpus inundatus (R.BR.) Pore. Enc. Suppl. 5 (1817) 103; SpRENG. Syst. | (1825) 207; BENTH. FI. Austr. 7 (1878) 329; ?K UK. Bot. Jahrb. 69 (1938) 258; OHwI, Bot. Mag. Tokyo 56 (1942) 203. — Isolepis inundata R. Br. Prod. (1810) 222. — S. setaceus (non L.) RENDLE in Gibbs, Arfak (1917) 90: an K.ScH. & Laut. Fl. Schutzgeb. (1901) 195?. Perennial; rhizome not creeping, not forming 516 mats. Stems usually coarser than in S. subtilissimus, tufted, slender, erect, strongly compressed, striate, smooth, 5-30 cm by )4—1 mm wide. Leaves usually all reduced to their sheaths, the upper one often with a very short, rarely + elongated blade; lower sheaths purplish. Inflorescence pseudolateral, capitate, con- sisting of 3-12 (in depauperate specimens sometimes 1 or 2) spikelets, frequently proliferous. Involucral bract at first erect, finally pushed aside by the mature inflorescence, obtuse, at the base with whitish or more or less purplish hyaline auricles, very short, as long as or slightly longer than the inflorescence, 4-10 mm. Spikelets ovoid to oblong-ovoid, some- what angled, acute, (2—)4-6 mm long. Glumes mem- branous, ovate or broadly ovate, concave, somewhat keeled, obtuse or minutely mucronulate, not emar- ginate, 1/4-134 by 1-1)4 mm; midnerve distinct, green, sides finely 3—4-nerved, striate with purplish FLORA MALESIANA [ser. I, vol. 7° streaks, rarely pale throughout. Brist/es none. Sta- mens 1-2; anthers oblong, c. 4 mm; connective shortly produced. Sty/e very short; stigmas 3, much longer than the style. Nut triquetrous with narrowly ribbed angles, elliptic-ovate, slightly dorsiventrally compressed, apiculate, smooth, stramineous, %%—1 by %-% mm. Distr. Widely distributed in the eastern Australian States, Tasmania, New Zealand, also in temperate South America, in Malesia: New Guinea (W. New Guinea: Arfak Mts, Wissel Lake region; NE. New Guinea: Western Highlands, Hagen Distr.; Mt Sarawaket, according to KUKENTHAL, /.c., not seen; Papua: Southern Highlands: Mt Giluwe; Eastern Highlands: Mt Wilhelm, Brass 30421. Ecol. In open marshes and peat swamps, !1750- 2700 m. Vern. New Guinea: tambugo, Mendi lang. 10. Section Micranthi C. B. CLARKE, FI. Br. Ind. 6 (1893) 663. — Sect. Microstyli CLARKE, Kew Bull. add. ser. 8 (1908) 113. — Sect. Squarrosi CHERM. Fl. Madag. fam. 29 (1937) 141, descr. gall. Type species: S. squarrosus L. 21. Scirpus squarrosus LINNE, Mant. 2 (1771) 181; Rotts. Descr. & Ic. (1773) 49, t. 17, f. 5; VAHL, En. 2 (1806) 259; Boeck. Linnaea 36 (1870) 734, quoad Specim. asiat.; CLARKE, Fl. Br. Ind. 6 (1893) 663; Ill. Cyp. (1909) t. 52, f. 11-13; Camus, Fl. Gén. I.-C. 7 (1912) 134; KERN, Reinwardtia 4 (1956) 93.— Isolepis squarrosa R.& S. Syst. 2 (1817) 111; NEEs in Wight, Contr. Bot. Ind. (1837) 106. — Ascolepis tenuior STEUD. Syn. 2 (1855) 105. — S. chinensis (non OsB.) RAYMOND, Nat. Canad. 84 (1957) 123. Annual; glabrous. Stems erect, tufted, setaceous, obtusangular, smooth, 5-20cm by /4-)4 mm. Leaves shorter than the stems, filiform, smooth or slightly antrorsely scaberulous at the top, c. | mm wide; ligule absent; lower sheaths ferrugineous, purplish striate, bladeless. Inflorescence capitate, usually pseudolateral, consisting of (1—)2—4 spikelets. Involucral bracts |—3, similar to the leaves, dilated at the base, the lowest usually erect, continuous with the stem, !-6cm, the others when present much shorter, patent to reflexed. Spikelets sessile (at length, when the lower glumes and nuts have fallen off, seemingly peduncled by the naked base of the rachilla), ovoid to subcylindrical, terete, obtuse, very densely many-flowered, greenish or brownish, 3—7 by c. 3 mm. G/umes thinly membranous, squarrose, easily caducous, obovate-rhomboid, 3-—5-nerved (midnerve prominent, side-nerves delicate), suddenly narrowed into a c. 14 mm long recurved mucro, 4-4 mm (mucro included) by c. 14 mm. Bristles none. Stamens \(—2); anther oblong, c. 14 mm. Style hardly any; stigmas 3, short, much recurved. Nut trigonous, obovate or oblong-obovate, not or hardly apiculate, smooth, ultimately castaneous to black, 0.5—0.6 by 0.3—0.35 mm; epidermal cells hexangular. Distr. S. Asia, in Malesia very rare: Malay Penin- sula (P. Penang, Johore, Singapore). In FI. Br. Ind. 6 (1893) 663 also recorded for Java. Ecol. In cultivated land (P. Penang), sandy open places, grassy fields, at low altitude. Note. S. squarrosus is often confused with Lipocarpha microcephala (R.BR.) KUNTH, which -it deceivingly resembles in habit. The latter species is easily recognized by the narrow c. | mm long nut enveloped by 2 thinly membranous hypogynous scales, and by the widely patent or reflexed involucral bracts. Excluded or Doubtful Scirpus borneensis H. Preirr. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 172. — West Borneo, lower course of Serawei R., HANS WINKLER 258, 1258. The type collection got lost during the war. From the description it seems doubtful whether it actually belonged to Scirpus. It is incomprehensible why PFEIFFER supposed ““S. macrothyrsus MiQ.” in De Clercq, Nieuw Plantk. Woordenb. Ned. Ind. (1907) might belong here, for this name is an error for Carex macrothyrsa Miq. 9. FUIRENA Rotts. Descr. & Ic. (1773) 70; KUNTH, En. 2 (1837) 180; Borck. Linnaea 37 (1871) 98; B. & H. Gen. Pl. 3 (1883) 1053; CLARKE, Kew Bull. add. ser. 8 (1908) 115. — Fig. 32. 1974] CYPERACEAE (Kern) St7 Fig. 32. Fuirena ciliaris (L.) Roxs. a. Habit, x 74, b. spikelet, x 6, c—d. glume, dorsal and lateral views, x 12, e. deflorate flower, f. scaly perianth segment and two bristle-like ones, g. nut, all x 24 (a—g CorrtT 51). Annual or perennial herbs. Stems erect, noded, leafy throughout their length. Leaves usually with elongate, linear or lanceolate blades and closed sheaths. Ligule membranous. Inflorescence paniculate, elongate, consisting of a terminal partial inflorescence and 1-several axillary ones, but often much reduced. Spikelets clustered, terete, usually greyish green or lead-coloured. Rachilla persistent, not winged, with lozenge-shaped scars. Glumes spirally imbricate, acropetally caducous, not keeled, nearly always aristate, 3-nerved, hairy on the back at least in the upper half, the lower 1-2 empty. Flowers hermaphrodite, the upper ones tabescent. Perianth (in sect. Fuirena) consisting of 3 outer bristles opposite the angles of the nut (sometimes reduced or wanting), and 3 inner scales opposite the faces of the nut, or (in extra-Malesian sections) of 2 series of 3 518 FLORA MALESIANA [ser. I, vol. 7° (sometimes reduced or wanting) bristles. Stamens (2—)3. Style not or hardly dilated at the base, continuous with the ovary, glabrous; stigmas 3. Nut small, triquetrous, obovate or ovate, more or less stipitate, smooth or trabeculate, beaked, falling off enclosed by the hypogynous scales. Distr. About 30 spp. in the warm regions of the globe, most of them in tropical Africa and tropical America; in Malesia only 2 spp. Ecol. The Malesian spp. grow in open wet localities at low and medium altitudes. Notes. Both Malesian spp. belong to sect. Fuirena (sect. Fuirena ‘proper’ CLARKE, FI. Br. Ind. 6, 1893, 666; sect. Eu-Fuirena CLARKE, Fl. Trop. Afr. 8, 1902, 463; sect. Genuinae CLARKE, Kew Bull. add. ser. 8, 1908, 116.—Type species: F. umbellata ROTTB.). This section is characterized by the 3 broad, petal-shaped hypogynous scales often alternating with 3 hypogynous bristles. The most acceptable interpretation of these bristles and scales is that they represent the perianth. This was already accepted by LesTIBOUDOIS (Essai sur la famille des Cypéracées, 1819, 13, 34), and since BENTHAM (J. Linn. Soc. Bot. 15, 1877, 510) advocated it, by most modern authors. NEEs (Linnaea 9, 1834, 5, 278, t. 4, f. 2-3), assuming that the stamens were inserted below bristles and scales, took the latter two for staminodes, whereas BOECKELER, !.c., was of the opinion that only the scales should be interpreted as such. The delimitation of the genus against Scirpus is very difficult, as in several extra-Malesian spp. the perianth- segments are either all bristly or lacking, like in Scirpus; on the other hand they are flat and petal-like in Scirpus litoralis, recalling the scales of Fuirena sect. Fuirena. The noded, leafy stems and the dark, aristate, hairy glumes, in some way also characteristic of Fuirena, are likewise found in some species of Scirpus. KEY TO THE SPECIES 1. Hypogynous scales hardly clawed, almost sessile, obovate, narrowed at the base, usually with a minute curled mucro at the truncate apex; bristles usually wanting. Mostly rather stout perennial; a glabrous or ciliate only at the base, very rarely pubescent, (5—)8—15(—25) mm wide . F. umbellata 1. Hypogynous scales distinctly clawed, subquadrate, cordate at the base, tridentate at As apex; bristles present (though usually short). Slender annual; leaves pubescent or at least ciliate throughout the margins, 3—8 mm wide 1. Fuirena umbellata Rotts. Descr. & Ic. (1773) 70, t.19 (i.e. t. 18 altera), f. 3; VAHL, En. 2 (1806) 383; KUNTH, En. 2 (1837) 185; Mor. Syst. Verz. (1846) 97; ZOLL. Syst. Verz. 2 (1854) 61; SteuD. Syn. 2 (1855) 126; Mig. FI. Ind. Bat. 3 (1856) 328; Boeck. Linnaea 37 (1871) 110, incl. var. pentagona; BENTH. FI. Austr. 7 (1878) 337; F.-VILL. Nov. App. (1882) 308; VIDAL, Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 284; CLARKE, FI. Br. Ind. 6 (1893) 666; K. Scu.& Laut. Fl. Schutzgeb. (1900) 194; Rip-. J. Str. Br. R. As. Soc. n. 46 (1906) 224; Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 80; CLARKE, Philip. J. Sc. 2 (1907) Bot. 101; Ill. Cyp. (1909) t. 59, f. 9; Camus, FI. Gén. I.-C. 7 (1912) 141, f. 18, 6; VALcK. Sur. Nova Guinea 8 (1912) 706; Merr. En. Borneo (1912) 58; En. Philip. 1 (1923) 116; KUx. Bot. Jahrb. 59 (1924) 52; RipL. Fl. Mal. Pen. 5 (1925) 162; BAck. Onkr. Suiker. (1928) 146, t. 147;S.T. BLAKE, J. Arn. Arb. 28 (1947) 228; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 16; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 457. — Scirpoides paradoxus Rotts. Progr. (1772) 27, nom. provis. — F. pentagona' W.& A. ex Nees in Wight, Contr. Bot. Ind. (1834) 93; in Hook. J. Bot. Kew Misc. 6 (1854) 28; Mia. FI. Ind. Bat. 3 (1856) 329; Kurz, Nat. Tijd. N. I. 27 (1864) 224, non SCHUM. 1827. — F.quinquangularis Hassk. Tijd. Nat. Gesch. Phys. 9 (1842) 128; Flora 25 (1842) Beibl. 3; Pl. Jav. Rar. (1848) 58; Steup. Syn. 2 (1855) 127. — F. uncinata (non KUNTH) CAMUS, FI. Gén. I.-C. 7 (1912) 141. — F. philippinensis GaND. Bull. Soc. Bot. Fr. 66 (1920) 297. 2. F. ciliaris Perennial with short thick rhizome. Stems tufted, erect, slender to rather stout, acutely 4—5-angular, pubescent in the inflorescence or very rarely through- out, 30-110cm by up to 8mm at the base, the lowest internode not rarely bulbously thickened. , Leaves much shorter than the stems, rather rigid, flat, acute, glabrous or ciliate at the base, very rarely pubescent, with 5 prominent nerves, (5—)8—15(—25) mm wide, the lower ones very short; sheaths shorter than the internodes, usually glabrous. Inflorescence usually with a terminal partial inflorescence and several axillary ones, the latter single or binate, on pubescent peduncles. Lower bracts similar to the leaves, as long as or slightly overtopping the inflo- rescence, obliquely erect to patent, sheathing at the base; upper ones much shorter, not or hardly sheathing. Spikelets in very dense clusters, ovoid to oblong-ovoid, acute, finally more or less squarrose, brownish or greyish to blackish green, 4—10 by 2)4-3 mm. G/umes membranous, ovate or obovate, rounded at the apex, short-hairy, pilose at the apex, 2-24 by 114-14 mm, somewhat below the apex with a recurved, often pilose, 4-1/4 mm long awn. Perianth usually uniseriate; bristles usually absent, very rarely present, but always much shorter than the nut; scales about as long as the nut, subsessile (with a very short curved claw), obovate, narrowed at the base, truncate at the somewhat thickened top, the midnerve usually excurrent into a short curled mucro, minutely ciliolate at the apex, 3-nerved, brown. Stamens (2—)3; anthers linear-oblong, 4—- A ' Illegitimate name, antedated by F. pentagona ScHuM. Guin. PI. (1827) 42, which is conspecific with F. pentagona NEES, but based on a different type. All belong to the same species. 1974] Cy PERACEAE (Kern) 519 mm long. Nut elliptic to obovate, triquetrous, narrowed into a stipe and beak, smooth, shining stramineous to fuscous, 1-114 by 0.6-0.7 mm. _ Distr. Widely distributed in the tropical and subtropical, not too dry regions of the whole world, presumably common throughout Malesia, but not yet collected in the Lesser Sunda Is. and the Moluc- cas; according to BACKER (1928, I.c.) in Java less common than F. ciliaris. Ecol. In open wet lands: swamps, pools, ditches, marshy shores of lakes, swampy grasslands, rice- fields, sometimes in secondary forests, at low and medium altitudes, up to 1800 m. Vern. Djukut, wawalingian, pépéntul, S, kasabon, J, rumput buku buloh, r. lidah ménkarong, r. kélulut, M (Mal. Pen.), blidang ajér, M (Palembang), rumput paro-paro, M (Djambi), tihe galbus, Asahan, anggal- bur, Toba, tikang bélangkas, S.& E. Borneo, rumput ragi, r. kuluwing, Minahassa, hehe, W. New Guinea, sinbora, Orokaiva lang., Papua. Note. Usually stouter and more glabrous than F. ciliaris, with broader leaves and much more compound inflorescence, but these characters are all highly variable. Specimens with strongly pilose stems, leaf-blades and sheaths are very rare. The two spp. can always be distinguished with certainty by the quite different shape of the hypogynous scales. 2. Fuirena ciliaris (L.) Rox. [ Hort. Beng. (1814) 81] Fl. Ind. 1 (1820) 184; Merr. Fl. Manila (1912) 119; En. Philip. | (1923) 116; Back. Onkr. Suiker. (1928) 146, t. 146; Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 16. — Scirpus ciliaris LINNE, Mant. 2 (1771) Loe Rome a Descr. & Ice. (1773) 55, t: 17, f. 1. cf. CLARKE, J. Linn. Soc. Bot. 30 (1894) 314. — F. glomerata LAMK, Tabl. Enc. Méth. Bot. | (1791) 150: VAHL, En. 2 (1806) 386; Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 360: Descr. Herb. Timor. (1835) 32; KUNTH, En. 2 (1837) 184; SteuD. Syn. 2 (1855) 127; Mia. FI. Ind. Bat. 3 (1856) 328: Boeck. Linnaea 37 (1871) 107; BENTH. FI. Austr. 7 (1878) 338; F.-VILL. Nov. App. (1882) 309; CLARKE, FI. Br. Ind. 6 (1893) 666; Philip. J. Sc. 2 (1907) Bot. 101; Rip_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 81: CLARKE, Ill. Cyp. (1909) t. 59, f. 6; Camus, Fl. Gen. I.-C. 7 (1912) 140; Rip. Fl. Mal. Pen. 5 (1925) 163. — F. rotboellii NEES in Wight, Contr. Bot. Ind. (1834) 94; ZOLL. Syst. Verz. 2 (1854) 61; Mia. FI. Ind. Bat. 3 (1856) 329.—F. striata LLANOs, Fragm. PI. Filip. (1851) 21; repr. by F.-Vit_. & NAves in Blanco, FI. Filip. ed. 3, 4! (1880) 14. — F. wallichiana (non KUNTH) Camus, FI. Gen. I.-C. 7 (1912) 141. — Fig. 32. Annual. Stems slender, obtusangular, striate-sul- cate, pubescent at the top or sometimes throughout, (S—)10—40 cm by |—2 mm, not incrassate at the base. Leaves rather weak, more or less pubescent or at least ciliate on the margins, with 3—5 more prominent nerves, light green, 3-8 mm wide; sheaths pubescent. Inflorescence sometimes a single terminal cluster, but usually also with 1—2 clusters on short pubescent peduncles in the upper axils. G/lumes obovate to oblong-obovate, short-hairy almost throughout the back, pilose in the centre, 1%—13% by c. 1/4 mm (the c. |!mm long awn excluded). Perianth biseriate: bristles scaberulous or almost smooth, sometimes as long as the nut, usually much shorter; scales distinctly clawed, subquadrate, cordate or hastate at the base, with 3 minute teeth at the distinctly thickened top (central tooth largest), glabrous or minutely hairy at the apex. Anthers oblong, c. 14mm long. Nut slightly smaller than in F. umbellata, 34-1 by 0.5—0.6 mm. Otherwise as F. umbellata. Distr. In the tropics of the Old World: tropical Africa, throughout SE. Asia (extending to S. China, Japan, and to tropical Australia), in Malesia: N. Sumatra (Atjeh), Central Sumatra, Malay Peninsula (Kedah, Wellesley, Perak, Kelantan, P. Langkawi, P. Penang), throughout Java, Borneo (Sarawak), Lesser Sunda Islands (Sumbawa, Timor), throughout the Philippines, S. Celebes, W. New Guinea (WICHMANN M14 in L). Ecol. In open wet lands: in swampy grassland, on riverbanks, and especially in wet rice-fields, at low altitudes (in Java up to 800 m, in Atjeh collected at 1180 m), usually scattered but sometimes abundant. Vern. Kapohan, J, rumput halia, Mal. Pen., rébu osap, Sumbawa; Philip.: pugdpak, Tag. Excluded Fuirena glabra ECKL. ex KUNTH, En. 2 (1837) 182; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 17. — Only known from S. Africa. A specimen of this species in the Herbarium of the Amsterdam University is labelled ““Lamongan, Java, Nov. 1904, P. MOLHUYSEN’’. As MOLHUYSEN’s collections include several plants from Africa, which are certainly not native to Java, this record is apparently based on an erroneously localized specimen. See Fl. Mal. I, | (1950) 366. 10. LIPOCARPHA R.Br. in Tuckey, Narr. Exped. Congo, App. (1818) 459, nom. cons.; KUNTH, En. 2 (1837) 266; Steup. Syn. 2 (1855) 129; Bogeck. Linnaea 37 (1871) 114; CLARKE, Kew Bull. add. ser. 8 (1908) 116. — Hypaelyptum VAHL, En. 2 (1806) 283; R.BR. Prod. (1810) 219, p.p.; K.ScH. in Engl. Pflanzenw. Ost. Afr. C (1895) 126. — Tunga Roxs. FI. Ind. 1 (1820) 187, p.p. — Fig. 33. Annual or perennial glabrous herbs. Stems tufted, erect, smooth, leafy only at the base. Leaves elongate, linear; sheaths of the basal ones open, of the cauline ones closed at the base. Ligule absent. Inflorescence terminal, capitate, with (1—)2—8 spikelets, subtended by some involucral bracts similar to the leaves. 520 FLORA MALESIANA [ser. 1, VoL? Fig. 33. Lipocarpha chinensis (OsB.) KERN. a. Habit, x 7%, 5. floral diagram, c. glume, d. glume with flower, e—f. the two transparant inner scales, g. nut with style, all x 18 (a—g RAHMAT Si BOEEA 4999). 1974] Cy PERACEAE (Kern) 521 Spikelets small, sessile (finally seemingly peduncled by the rachilla naked at the base because of the caducous glumes), terete, very densely many-flowered. Rachilla persistent, not winged, marked by lozenge-shaped scars. Glumes spirally imbricate, acropetally caducous, not keeled, the lower 1—2 empty. Flowers hermaphrodite, the upper ones tabescent. Perianth consisting of 2 (readily overlooked!) hyaline, nerved scales placed in the median plane (i.e. fore and aft, parallel with the subtending glume, the posticous scale embracing the anticous one). Stamens 1—2; anthers small, oblong to linear, with shortly produced smooth connective. Style small, not dilated at the base, continuous with the ovary, glabrous; stigmas 2—3. Nut small, dorsiventrally compressed, trigonous or planoconvex, oblong-obovate to narrowly oblong, smooth, reticulate, often slightly curved, enclosed by the hypogynous scales; epidermal cells isodiametric. Distr. About 15 spp.. in tropical America, tropical and South Africa, and from continental S. and SE. Asia through Malesia to Australia, apparently absent from the Pacific islands; most of the species in Africa; in Malesia only 2 spp. Ecol. The Malesian species inhabit open (permanently or temporarily) marshy localities at low and medium altitudes; L. chinensis was once found in N>w Guinea at 2800 m. L. microcephala is almost restricted to regions with a pronounced dry monsoon. Notes. The most important character of the genus lies in the two median hypogynous scales, which are difficult to discern as they are very thin and closely adhere to the nut. They are interpreted in various ways (see CLARKE, FI. Trop. Afr. 8, 1902, 468). The most acceptable interpretation to me is that they represent the perianth, a view already advanced by BENTHAM (J. Linn. Soc. Bot. 15, 1877, 509, 510, 518), and followed by HOoLtTtTuM (Bot. Rev. 14, 1948, 525-541). They take the scales for homologous with the bristles in Scirpus and Eleocharis. Then the genus has to be placed next to Scirpus sect. Micranthi, to which it shows affinities in several respects. According to VON GOEBEL (1887) each stamen of L. chinensis has two thecae, but each theca has only one pollen chamber (checked by Dr. W. VAN HEEL, 1972). CHERMEZON divided the genus into 2 sections: Sect. Acutae: spikelets squarrose by the mucronate glumes. Sect. Obtusae (correct name at present sect. Lipocarpha): spikelets not squarrose; glumes not mucronate. Of the Malesian species L. chinensis belongs to sect. Lipocarpha, L. microcephala to sect. Acutae. KEY TO) DHE SPECLES 1. Spikelets not squarrose, whitish, 4-5 mm wide; glumes not mucronate, but with a short erect or slightly incurved triangular tip. Hypogynous scales distinctly longer than the nut, c.2 mm. cancer er: %-1 mm long. Nut )4—!4 mm wide. Rather firm plant with leaves 2-4 mm wide one . L. chinensis 1. Spikelets squarrose by the distinct, finally recurved mucros of the glumes, greenish, - 3mm wide. Hypogynous scales but slightly longer than the nut, c. 1 mm. Anthers Bote “~ 7% mm long. Nut YY, mm wide. Very slender, weak plant with leaves 1-2 mm wide... 2. L. microcephala 1. Lipocarpha chinensis (OsB.) KERN, Blumea Suppl. 4 (1958) 167; in Back. & Bakh. f. Fl. Java 3 (1968) 457. — Scirpus chinensis Oss. Dagb. Ostind. Resa (1757) 220. — Scirpus senegalensis LAMK, Tabl. Enc. Meth. Bot. | (1791) 140. — Hypolytrum senegalense RIcu. in Pers. Syn. | (1805) 70, non CLARKE, 1902. — Hypaelyptum argenteum VAHL, En. 2 (1806) 283. — Hypolytrum argenteum KUNTH in H.B.K. Nova Gen. & Sp. Pl. 2 (1816) 218, guoad basion. — Tunga laevigata Roxs. FI. Ind. 1 (1820) 188. — L. laevigata Nees ex WIGHT, Cat. Pl. (1834) 110; in Wight, Contr. Bot. Ind. (1834) 92; in Hook. J. Bot. Kew Misc. 6 (1854) 28.— L. argentea R.BR. ex NEES, Linnaea 9 (1835) 287; KuNTH, En. 2 (1837) 266, p.p. (excl. pl. am.); StEUD. Syn. 2 (1855) 129; Mia. FI. Ind. Bat. 3 (1856) 331; Boeck. Linnaea 37 (1871) 114, p.p.; BENTH. FI. Austr. 7 (1878) 336; F.-VILL. Nov. App. (1882) 308; CLARKE, FI. Br. Ind. 6 (1893) 667; K.ScuH. & Laut. Fl. Schutzgeb. (1900) 197: Rip-. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 81; CLARKE, Philip:) 3: ‘Se? 2) ~(1907)" ‘Bot. 109; “ly Gyp. (1909) t. 60, f. 4-7; Koorp. Exk. Fl. Java 1 (1911) 183; Atlas f. 191; Camus, Fl. Gén. I.-C. 7 (1912) 143, t. 1, f. A, 1,2; VALCK. Sur. Nova Guinea 8 (1912) 706; Merr. En. Born. (1921) 53; En. Philip. | (1923) 102; KUx. Bot. Jahrb. 59 (1924) 51; ibid. 69 (1938) 259: Rip. Fl. Mal. Pen. 5 (1925) 163: BAcK. Onkr. Suiker. (1928) 124, t. 118. — Kyllingia albes- cens STEUD. Syn. 2 (1855) 68; Mig. Fl. Ind. Bat. 3 (1856) 294: F.-ViLL. Nov. App. (1882) 300. — L. senegalensis TH. & HEL. DURAND, Syll. Fl. Congol. (1909) 619; Danby, J. Bot. 70 (1932) 331; Ouw1, Bot. Mag. Tokyo 56 (1942) 204; Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1943) 166; S. T. BLAKE, J. Arn. Arb. 28 (1947) 229; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 28.—L. triceps (non NEES) Camus, Fl. Gén. I.-C. 7 (1912) 144. — L. debilis RIpL. Trans. Linn. Soc. II, Bot. 9 (1916) 243; ef. KERN, Reinw. 4 (1956) 96.— Cyperus lipocarpha Koyama, Bot. Mag. Tokyo 73 (1960) 438. — Fig. 33. 522 FLORA Annual or perennial. Rhizome hardly any. Stems tufted, erect, rather stiff, obtusely trigonous, striate- sulcate, smooth, (10—)20—S0(—70) cm by 1-2 mm. Leaves usually much shorter than the stems, rather rigid, flat or involute, gradually attenuate, obtusish, practically smooth (only minutely scaberulous at the very top), 2-4 mm wide. Inflorescence 1—-1)4(—2) em across. Involucral bracts 2—3(—5), much overtopping the inflorescence, finally horizontally spreading to reflexed, dilated at the base, the longest up to LO{—15) cm. Spikelets 3—6(—10), ovoid to oblong-ovoid, terete, obtuse, whitish, 5-8 by 4-5 mm. Glumes membranous, spathulate to oblong-obovate, sub- truncate at the apex with obtuse triangular tip, concave, with strong midnerve and faintly 1|—2- nerved sides, often purplish lineolate, 134-27 by 1-l)4mm. Hypogynous scales oblong, 5—7- nerved, much longer than the nut, 134-2 mm long. Stamens \(—2); anthers linear, 74-1 mm. Style (2—)3- fid. Nut oblong-obovate to oblong, straight or slightly curved, minutely apiculate, stramineous to fuscous, 1-114 by 4-4 mm. Distr. Tropical and S. Africa, Madagascar, Mas- carene Is., S. and SE. Asia (to Formosa and the Ryu Kyu Is.), tropical Australia, in Malesia: throughout Sumatra, Banka, Lingga Arch., Malay Peninsula (common all over the peninsula), Central Java (only once collected in the Rawah Pening near Ambarawa, 1941), Madura, Borneo, Anambas & Natuna Is., Philippines (Luzon, Mindanao), Celebes, Moluccas, and New Guinea. In view of the wide distribution in the Malesian region it is very remarkable that there is only a single record for Java, and none for the Lesser Sunda Is. Ecol. In open marshy places; swamps, marshes, along lakes and pools, on riverbanks, in moist grassfields, wet rice-fields, clearings in forests, locally often abundant, at low and medium altitudes (usually between 0 and 1800 m, in W. New Guinea once at 2800 m: Brass 10741). Vern. Rumput lida angsa, r. rotan, Mal. Pen., imbulu tano, tihe-tihe, Sum. E. C., pohun, Lingga Arch.: Philip.: balabak, Buk.; New Guinea: kurumb, Mendi lang. 2. Lipocarpha microcephala (R.BR.) KUNTH, En. 2 (1837) 268; SteubD. Syn. 2 (1855) 130; Mia. FI. Ind. Bat. 3 (1856) 331; Boeck. Linnaea 37 (1871) 118; BENTH. FI. Austr. 7 (1878) 337; F.v.M. Pap. Pl. 7 (1886) 34; CLarkeE, FI. Br. Ind. 6 (1893) 668; Koorpb. Minah. (1898) 285; RipL_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 82, p.p.; CLARKE, Philip. J. Sc. 2 (1907) Bot. 110; Koorp. Exk. Fl. Java | (1911) 183: CAMUS, FI. Gén. I.-C. 7 (1912) 144; Merr. En. Philip. 1 (1923) 102; KUk. Bot. Jahrb. 59 (1924) 51; RIDL. Fl. Mal. Pen. 5 (1925) 163, p.p.; OHw1, Bot. Mag. MALESIANA [ser. I, vol. 7° Tokyo 56 (1942) 204; Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 167; S. T. BLake, J. Arn. Arb. 28 (1947) 228: Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 27; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 457. — Hypaelvptum microcephalum R.Br. Prod. (1810) 220. — Ascolepis kyvllingioides STEUD. [in Zoll. Syst. Verz. 2 (1854) 63, nom. nud.] Syn. 2 (1855) 105; Mig. FI. Ind. Bat. 3 (1856) 313: Koorp. Exk. Fl. Java 1 (1911) 184. — Kvillingia squarrosa STEUD. Syn. 2 (1855) 68; Mig. Fl. Ind. Bat. 3 (1856) 294. — L. zollingeriana Boeck. Flora 42 (1859) 100. — Scirpus squarrosus (non L.) K.ScH. & Laut. Fl. Schutzgeb. (1900) 195; VaLcK. Sur. Nova Guinea 8 (1912) 705.— Cyperus zollingerianus KOYAMA, Bot. Mag. Tokyo 73 (1960) 438. Annual. Stems tufted, erect, very slender, often filiform, obtusely trigonous, striate-sulcate, smooth, 5—30(-40) cm by 14-1 mm. Leaves usually much shorter than the stems, weak. flat or involute, gradually acuminate, acute, smooth, or minutely scaberulous at the very top, 1-2 mm wide. Inflores- cence '4—1 cm across. Involucral bracts 2—3, much overtopping the inflorescence, finally horizontally spreading to reflexed, dilated at the base, the longest up to 10cm. Spikelets (1—)3(—4), globose-ovoid to oblong-ovoid, terete, squarrose, obtuse, greenish, 3—5 (finally to 8) by 2-3 mm. G/umes thinly membra- nous, oblong-obovate, concave, with strong mid- nerve and nerveless sides, often purplish lineolate, without the mucro 1-1/4 by 7%-?4 mm; mucro finally recurved, smooth, 4-34 mm. Hypogynous scales linear, 3—S-nerved, slightly longer than the nut, |—1!4 mm long. Stamens 1(—2); anthers oblong, 4-24 mm long. Style 2(—3)-fid. Nut oblong-linear, slightly curved, minutely apiculate, stramineous to fuscous, 0.9-1 by Z-'4 mm. Distr. SE. and E. Asia (Indo-China, S. China, Japan, Korea, Ryu Kyu Is., Formosa), all mainland Australian States, in Malesia: Malay Peninsula (Kelantan: Kota Bharu; P. Penang, n.v.; Singapore), Java (W. Java: Indramaju, Cheribon; Central Java: Kedu, Borobudur; E. Java: Kediri), Madura, Philip- pines (Luzon: Apayao, Ifugao, Bulacan, Rizal), Celebes (Minahassa, Makassar), Moluccas (Talaud Is., Buru, Ceram, Ambon), New Guinea (also on P. Japen and Daru I.), New Britain. Ecol. In open wet localities (wet grassfields, rice- and sugar-cane-fields, on damp soil of savannah- forests) at low altitude, very local and absent over large tracts of Malesia, apparently largely bound to areas subject to a dry season. Vern. Kukus imbola, Minahassa; New Guinea: famdi, Wapi lang. Note. Very similar in habit to Scirpus squarrosus L. and often confused with it; for the differences see p. 516. 11. ELEOCHARIS R. Br. Prod. (1810) 224: KuntTH, En. 2 (1837) 139; Steupb. Syn. 2 (1855) 75; Boeck. Linnaea 36 (1869-70) 418; CLARKE, Kew Bull add. ser. 8 (1908) 105; SvENS. Rhodora 31 (1929) 121-135; 152-163; 167-191; 199-219; 224-242; ibid. 34 (1932) 193-203; 215-227: ibid. 36 (1934) 377—389; ibid. 39 (1937) 210-231; 236-272: ibid. 41 (1939) 1-19; 43-77; 90-110:S.T. BLAKE, Proc. R. Soc. Queensl. 1974 | CYPERACEAE (Kern) 523 50 (1939) 88-132; KoyAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 84-99. — Heleo- charis Lest1B. Ess. Fam. Cyp. (1819) 41. — Chaetocyperus Nees, Linnaea 9 (1835) 289. — Chlorocharis RiKL1, Jahrb. Wiss. Bot. 27 (1895) 564. — Fig. 34—36. Annual or perennial herbs, in the latter case often stoloniferous. Stems usually tufted, erect or arcuate, terete to acutely angled. Leaves reduced to bladeless tubular sheaths often bearing a short erect mucro (vestige of the blade) at the apex. Inflorescence a terminal, ebracteate, single spikelet, sometimes proliferous. Spikelet terete or angular, rarely compressed, few- to many- flowered. Rachilla persistent. Glumes spirally imbricate, sometimes subdistichous, caducous, the lowest 1—2 usually empty and more persistent than the fertile ones (in ser. Aciculares all glumes fertile). Flowers hermaphrodite. Perianth normally consisting of 6—10 bristles, not rarely reduced, very rarely wanting. Stamens 1—3; anthers ovate to linear, the connective produced into a small, rarely subulate appendage. Style usually glabrous, articulated with the nut, the dilated base persistent on the nut, in ser. Pauciflorae not sharply differentiated; stigmas 2—3. Nut 3- or 2-sided, obovate, orbicular, or pyriform, crowned by the enlarged style-base; epidermal cells hexagonal, transversely oblong to linear, or vertically oblong. Distr. According to SVENSON c. 150 spp., from the tropics to the polar regions of both hemispheres with the greatest concentration in the New World, in Malesia 19 spp. Ecol. Semi-aquatic and marsh plants inhabiting wet muddy places along streams, or swamps and shallow waters of lakes and ponds. Nearly all Malesian spp. are lowland plants ascending to medium altitudes; E. attenuata ascends to 2800 m, the northern E. acicularis was found at 900-1450 m; E. tetraquetra is almost restricted to altitudes above 1000 m. Uses. Only a few species are of economic value. Some are used for matting (E. ochrostachys, E. dulcis). The tubers of E. dulcis are edible and the species is therefore cultivated. Notes. Eleocharis isa natural genus nearest to Scirpus, Fimbristylis,and Bulbostylis; it ismainly characterized by the dilated style-base articulated with the ovary and persistent on the fruit, and the bladeless leaf-sheaths. The embryo, a variant of the Fimbristylis-type, is characterized by the first leaf more or less protruding from the germination-pore (VAN DER VEKEN, Bull. Jard. Bot. Brux. 35, 1965, 330). Only the members of ser. Pauciflorae (represented in Malesia by E. parvula) cause some difficulty as to the delimitation of the genus. Many authors consider them to belong to Scirpus. Their style-base, though confluent with the nut, is clearly different in texture and colour; therefore their natural place is obviously in Eleocharis; the type species of the series (see p. 537) has a typical Eleocharis embryo. Rik, Jahrb. Wiss. Bot. 27 (1895) 564, 574, Aared the genus into Eleocharis s.s. with eucyperoid anatomical structure, and Chlorocharis with chlorocyperoid anatomy. Both Scirpus and Fimbristylis lack the persistent style-base of Eleocharis and Bulbostylis. The leaf-sheaths of Eleocharis are at most mucronate, but in Scirpus a comparatively well developed blade is usually present. In Fimbristylis and Bulbostylis the perianth is always lacking; in the former the style, falling off as a whole, leaves no button on the nut. The etymologically more correct spelling ‘Heleocharis’ of LestriBouDo!s was followed by many authors. In the present revision the two spellings have not been considered as different in citing synonymy. In general SVENSON’s arrangement, somewhat modified by S. T. BLAKE, has been followed: however, the circumscription of several species has been broadened. KEY TO THE SERIES AND SUBSERIES represented in Malesia 1. Style-base sharply differentiated from and articulated with the nut proper (see E. spiralis). 2. Glumes indurated, cartilaginous on the back, with a distinct midnerve and finely many-nerved sides, not keeled or only obscurely so. Mostly coarse perennials with the stem as wide or almost as wide as the spikelet. Style 2-fid or 3-fid. Nut lenticular, stramineous or brown, 114-2 mm long. Spp. 1-8. 1. ser. Mutatae 2. Glumes membranous, with a distinct midnerve and nerveless sides, often keeled. Stem much narrower than the spikelet. 3. Style 3-fid. 4. Nut triquetrous, trigonous, or lenticular, smooth or variously sculptured (finely reticulate, or cancellate i.e. with deep honeycomb reticulation), but not trabeculate, the epidermal cells inconspicuous, mostly shortly vertically oblong. 5. Coarse or rather coarse perennials (excepts for small annual forms of E. congesta). Nut obtusely trigonous, not costate on the rounded angles, usually 1 mm or more long. Uppermost leaf-sheath distinctly mucronate at the mouth (muticous or almost so in E. congesta). 524 FLORA MALESIANA [ser. I, vol. 7° 6%) NutinigonoussSpp)9=li ems. be Oe nS A 2 gser se Vintec, 6. Nutlenticular.Sp.12 . . . 3. ser. Acutae 5. Dwarf, tufted annuals with capillary stems. Nut (in Mal. spp.) sharply triquetrous with costate angles, small, rarely up to 1 mm long. Uppermost leaf-sheath not mucronate. Spp. 13-14. 4. ser. Tenuissimae 7. Nut smooth or only finely reticulate under high magnification. Sp. 13. 4a. subser. Leiocarpeae 7. Nut coarsely cancellate.Sp.14 . . . 4b. subser. Chaetarieae 4. Nut obscurely trigonous or subterete, trabeculate (i. e. with some longitudinal ribs connected by numerous cross-ridges), the epidermal cells transversely linear or oblong. Lowest glume flower- bearing. Sp. 15. eal eid Se ee I Sandee ene MaRS OMSL AY 275-0. GCE TEI ES 3. Style 2-fid. Nut biconvex, smooth. 8. Rather coarse, usually stoloniferous perennials. Style-base spongy. Nut more than 1 mm long, yellow or brown. Sp.16 . . . 6. ser. Palustriformes subser. Palustres 8. Small annuals, without stolons. Style- -base not spongy. Nut less than 1 mm long, purplish or black. Spp.17-18 . . . 7. ser. Maculosae subser. Rigidae 1. Style-base confluent with the apex of the nut, not constricted at the base. Nut trigonous, usually reticulate under high magnification. Style 3-fid.Sp.19 . . . . . . . . +. +. +. +. #8. ser. Pauciflorae KEY TO THE SPECIES The length of the nut is given without the persistent style-base crowning the nut 1. Stems transversely septate, i.e. with a single large central cavity divided into several chambers by distinct transverse septa visible (at least when dry) from the outside. 2. Stems tufted on a short rhizome with elongated stolons sometimes bearing subglobose tubers. Spikelets 3—6 mm wide. Glumes greyish, 4— 6/4 mm long. Bristles 6-8. Nut 14-2 mm long ._ . 7. E. dulcis . Stems in a close linear series on a very stout, shortly creeping rhizome never producing tubers. Spikelets 8-9 mm wide. Glumes light brown, 7/4—9 mm long. Bristles 8-10. Nut 2)4—2/4 mm long. 8. E. sphacelata 1. Stems not transversely septate. 3. Glumes cartilaginous, not or hardly keeled, finely many-nerved, usually with prominent midnerve. Nut lenticular, stramineous to brown, 1/4—2/4 mm long. Style 2-fid or 3-fid. Spikelets hardly (if at all) wider than the stem (E. ochrostachys often excepted). 4. Nut constricted below the apex into a short but definite neck, the apex expanded. Stems acutangular. 5. Epidermal cells of the nut for the greater part transversely oblong to linear, shallowly pitted. Glumes appressed, not in 4 rows. Spikelet terete. Stems triquetrous, usually 3—4 mm wide. 1. E. acutangula 5. Epidermal cells of the nut for the greater part isodiametric, hexagonal, deeply pitted. Glumes finally subsquarrose, almost in 4 rows. Spikelet angular. Stems acutely 4—5-angled, c. 2 mm wide. 2. E. philippinensis 4. Nut not constricted below the apex, without neck. 6. Glumes very obtuse to truncate, very densely imbricate (the exposed part broader than long, only 14-1 mm long). 7. Stems triquetrous at least in the upper part. Style-base c. 4 as long and /4 as wide as the nut. Epidermal cells of the nut transversely On Rs linear to linear, in c. 20 vertical series oneitherface . . ; . 3. E. spiralis 7. Stems terete. Style- base c. Ki as long and almost as wide as the nut. Epidermal cells of the nut transversely oblong, in c. 30 vertical series on either face. . . . . 4. E. sundaica 6. Glumes obtuse, not particularly densely imbricate (the exposed part longer than broad, 2-3 mm long). 8. Epidermal cells of the nut small, not inflated, transversely oblong, in 15—20 vertical rows on either face. Nut with prominent longitudinal italien) . 5. E. ochrostachys 8. Epidermal cells of the nut large, strongly inflated, isodiametric or slightly transversely elongated, in c. 12(—15) vertical rows on either face. Nut not longitudinally ridged. 6. E. variegata 3. Glumes thinly membranous, often keeled, with distinct midnerve but nerveless sides. Nut triquetrous, trigonous, or lenticular. Spikelet much wider than the stem. 9. Style 3-fid. 10. Nut lenticular, plano-convex or biconvex. Uppermost leaf-sheath prominently mucronate. 12. E. acuta 10. Nut trigonous, subterete or triquetrous. 11. Nut trabeculate (with some longitudinal ribs connected by numerous cross-ridges) or cancellate (with deep, regular honey-comb reticulation). Stems capillary. 12. Nut obovoid, sharply triquetrous, almost truncate at the apex, cancellate. Style-base about as wide as the nut, 3-lobed, the lobes decurrent on the angles of the nut. 14. E. retroflexa 1974] 12. Nut obovoid-oblong, obscurely trigonous or subterete, minute, at most /% as wide as the nut, not decurrent Cy PERACEAE (Kern) 325 trabeculate. Style-base 15. E. acicularis 11. Nut smooth, neither trabeculate nor cancellate, only finely reticulate under high magnification. 13. Style-base not constricted at the base, confluent with the apex of the nut and seemingly a continuation of it, but of different colour and texture. Dwarfish (stems 1—7 cm by YA mm), often with filiform stolons ending in small fusiform tubers. 19. E. parvula 13. Style-base constricted at the base, sharply differentiated from and articulated with the nut. 14. Nut triquetrous with costulate angles, very small ( (in Malesian specimens) (c. 4mm long). Bristles absent E. nigrescens 14. Nut trigonous, not costulate, ae ym mm long. Bristles 6, as long as or longer than the nut. 15. Stems acutely quadrangular. Bristles densely scabrous to barbellate- subplumose. Nut 1/4—1/4 mm long, crowned by a large, cap-like style-base. 9. E. tetraquetra 15. Stems terete, more or less angular-ribbed but not quadrangular. Bristles more or less scabrous, not subplumose. Nut %—1)4 mm long with short- pyramidal style-base. 16. Style-base 3% to almost as wide as the nut. Glumes 2—2)4(—3) mm long. Uppermost leaf-sheath truncate or slightly oblique, distinctly mucronate. Nut 1-17 /Z mm long 10. E. attenuata 16. Style-base /4—/4 as wide as the nut. Glumes 14 2 mm long. Uppermost leaf-sheath usually oblique, not or but minutely mucronate. Nut 74-1 mm long 9. Style 2-fid. Nut lenticular. 11. E. congesta 17. Rather coarse, stoloniferous perennial. Nut 1/4— 1A mm long, yellowish or pale brown. Glumes c. 44 mm long 16. E. brevicollis 17. Small annuals without stolons. Nut less than 1 mm long, blackish when ripe. Glumes at most 2 mm long. 18. Spikelets c. 3 mm thick. Glumes scarcely keeled, 124-2 mm long. Bristles 6-8, slightly longer than the nut, ferrugineous to brown. Nut 34-24, mm long 7. E. geniculata 18. Spikelets 114-2 mm thick. Glumes keeled, up to VA mm long. Bristles 3— —4(—6), shorter than to as long as the nut, whitish, translucent, sometimes absent. Nut 14— “a mm long. 1. Series Mutatae SvENS. Rhodora 31 (1929) 127. — Limnochloa NEEs in Wight, Contr. Bot. Ind. (1834) 71; Linnaea 9 (1834) 294, non BEAUV. ex LeEsTIB. Ess. Fam. Cyp. (1819) 41? — Subg. Limnochloa C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 105.— Type species: E. mutata (L.)R. & S. 1. Eleocharis acutangula (ROxB.) SCHULT. in R.& S. Mant. 2 (1824) 91; Troupin, Fl. Sperm. Pare Nat. Garamba (1956) 108; ANDREws, Flow. PI. Sudan 3 (1956) 359; KERN in Back. & Bakh. f. Fl Java 3 (1968) 461. — Scirpus fistulosus Poir. Enc. 6 (1804) 749, non Forsk. 1775. — Scirpus acutangulus Roxs. FI. Ind. | (1820) 216. — E. fistulosa SCHULT. in R. & S. Mant. 2 (1824) 89; KunTH, En. 2 (1837) 155; Mor. Syst. Verz. (1846) 97; SteuD. Syn. 2 (1855) 80; Mia. Fl. Ind. Bat. 3 (1856) 302; Boeck. Linnaea 36 (1870) 472; NAves, Nov. App. (1882) 306; CLARKE, FI. Br. Ind. 6 (1893) 626; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 76; CLARKE, Ill. Cyp. (1909) t. 35, f. 1-4; Koor». Exk. Fl. Java | (1911) 197; Camus, Fl. Gén. I.-C. 7 (1912) 84, f. 13, 3 & 4; Merr. En. Philip. | (1923) 120; Rip. Fl. Mal. Pen. 5 (1925) 151; Fern. Rhodora 27 (1925) 39, t. 149, f. 5-10; Svens. Rhodora 31 (1929) 152, t. 188, £3; ibid. 41 (1939) 4;S.T. BLAKE, 18. E. atropurpurea Proc. R. Soc. Queens!. 50 (1939) 97, t. 7, f. 1-3; Ouwl, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 31; S. T. BLAKE, J. Arn. Arb. 28 (1947) 226; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 15, p-p.; KOYAMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 36; J. Fac. Sc. Un. Tokyo III, 8 (1961) 98. — E. planiculmis Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 80'; Naves, Nov. App (1882) 306; Mia. FI. Ind. Bat. 3 (1856) 301. — PE variegata (nec PRESL, nec KUNTH) BOECK. Linnaea 36 (1870) 470, p.p.; BACK. Onkr. Suiker. (1928) 153, Pp. Perennial, with short rhizome and long stolons. Stems erect, tufted, rather stout, somewhat spongy. triquetrous, smooth, not transversely septate, 30—70 cm by (1—)3—4mm. Sheaths membranous, rather loose, with oblique mouth, pointed at the summit, often rubescent. Spike/et cylindrical, scarcely if at all broader than the stem, terete, acute, many-flowered, dusky green, 2—3)4.cm by 3-4mm. Glumes firm, rather loosely imbricate, closely appressed, ovate or oblong-ovate, obtuse, not keeled, many-nerved with prominent midnerve, dirty stramineous, with rather broad scarious margins, 414—4'4(—-5) by 2)4-3)4 mm. Bristles 6-7, rather stout, one very short, one about as long as the nut, the others subequal, usually as long as or slightly surpassing the style- base, rusty brown. Stamens 2-3; anthers linear, ' The type is erroneously cited as ZOLLINGER 284; should be 281. 526 FLORA MALESIANA [ser. I, volLy? shortly apiculate, 1’4—2 mm. Style 3-fid. Nut turgidly and unequally biconvex, with scarcely ribbed mar- gins, obovate, distinctly constricted below the apex into a short neck c. 24 as wide, the expanded apex c. 4 as wide as the body of the nut, glistening stramineous, 14-2 by 14-14 mm; epidermal cells conspicuous, transversely oblong, in c. 15 vertical rows on either face, shallowly pitted. Style-base ovate-deltoid, flat, dark brown, c. 1% as long and /% as wide as the nut. Distr. Widely distributed but apparently un- common species in the tropics both of the Old and the New World: West Indies, tropical S. America, tropical Africa, Madagascar, S. and E. Asia (to Formosa and Japan), tropical Australia (Queens- land), in Malesia: Sumatra, Malay Peninsula (Kedah, Perak, Malacca, P. Langkawi, Singapore), W. and Central Java (also in Kangean, Madura), Lesser Sunda Islands (Timor, Alor, Wetar), N. Borneo, Philippines (Luzon, Mindanao), Celebes, New Guinea (Papua, NE. New Guinea). Distribution map in Rhodora 41 (1939) 10, f. 28. Ecol. In open wet places (swamps, rice-fields) at low alt., up to 800 m. Vern. Bhan-talobhanan, Md., komlak, Alor. Note. After TRouPIN, I.c., had pointed out that the correct name of this species is E. acutangula, not E. fistulosa, KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 98, accepted again the latter combination in ascribing it to Link, Jahrb. Gewachsk. 3 (1820) 78. However, the combination was not definitely made there (“Scirpus fistulosus . . . ist ebenfalls eine Eleocharis’’). 2. Eleocharis philippinensis SVENS. Rhodora 31 (1929) 155, f. 9: S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 98, t. 7, f. 7-10; KERN, Reinw. 4 (1956) 94; in Back. & Bakh. f. Fl. Java 3 (1968) 462. — E. fistulosa (non SCHULT.) BENTH. Fl. Austr. 7 (1878) 293. — E. variegata var. laxiflora (non CLARKE, FI. Br. Ind. 6, 1893, 626) CLARKE, Philip. J. Sc. 2 (1907) Bot. 90; Mer. Fl. Manila (1912) 114; En. Philip. | (1923) 121, p.p. — E. nuda (non CLARKE) SveNS. Rhodora 41 (1939) 8, p.p. — Fig. 34. Perennial, with short rhizome and long, slender stolons. Stems erect, tufted, slender, sharply and unequally 4—5-angled, smooth, not transversely septate, (10-)30-50 cm by c. 2mm. Sheaths thin, rather loose at the apex, purple at the base. Spike/et linear, scarcely if at all broader than the stem, angular, acute, many-flowered, dusky green, 2-6 cm by 2-3 mm. Glumes firm, almost in 4 rows, rather loosely imbricate, subsquarrose, more or less con- volute when dry, oblong-ovate, obtuse, rather prom- inently keeled, many-nerved, more or less distinctly brown-zonate within the narrow scarious margins, 4-5 by 2-2)4mm. Bristles 6—7, rather coarse, unequal (the inner ones distinctly longer than the outer ones), from less than half as long as the nut proper to slightly longer than the nut together with the style-base, retrorsely scabrous, brown. Stamens 3; anthers linear, 1-1/4 mm. Stigmas 2—3. Nut turgidly biconvex with slightly ribbed margins, obovate, somewhat pyriform, slightly narrowed into a distinct neck which is upwards outwardly curved into a prominent annulus, stramineous to fuscous, 1/4-174 by c. 114mm; epidermal cells in 15-20 regular vertical series on either face, hexagonal, deeply pitted. Sryle-base deltoid, flattened, c. 4-7 as long and 74-%% as wide as the nut. Fig. 34. Eleocharis philippinensis SVENS. a. Habit, x %, b. spikelet, x 11%, c. glume, x4, d. deflorate flower, x 8 (a—d RAmos 1461). 1974] CYPERACEAE (Kern) 527 Distr. Tropical Australia, New Caledonia, Thai- land (Bangkok, Kerr 11105), Indo-China (Bien Hoa), in Malesia: Malay Peninsula (Kedah), W. Java (near Djakarta), Madura, Lesser Sunda Is. (Sumba: Waik- abubak; Tainimbar Is.: P. Jamdena), Philippines (Luzon), New Guinea (tributaries of Goragatabu Creek 17 miles NE. of Port Moresby). Ecol. On open marshy ground, in wet rice-fields, in P. Jamdena in Melaleuca-forest surrounded by primary forest; at low altitude, up to 400 m. Vern. The local name rebbha lob-tolobhan, Md, is reported from Madura. It may also refer to other Eleocharis species (cf. E. acutangula). Notes. In 1939 referred by SVENSON to E. nuda C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 21. I have followed BLAKE, who keeps the two apart, dis- tinguishing E. nuda from E. philippinensis by the terete or trigonous stems, the relatively broader appressed glumes, the distinctly pyriform nut with shallower pitting, the strongly upcurved annulus and the caducous bristles. The specimens of E. nuda in the Kew Herbarium show that EF. nuda is clearly distinct. E. philippinensis is also related to E. plicarhachis (GRISEB.) SVENS. of the West Indies and South America, from which it differs in the slender, elongate spikelet with somewhat spreading glumes, and the larger nuts with more deeply pitted, almost iso- diametric epidermal cells and a wider annulus. E. ochrostachys differs in the terete or obscurely angular stems, the short spikelet with appressed glumes, the different shape and the transversely linear marking of the nut. As BLAKE pointed out, the outer cells of the nut in E. philippinensis occasionally show a tendency to lengthen transversely, but not to such an extent as to alter the typical character of the nut. In HeYLIGERS 1316 (CANB), the only collection known from New Guinea, the bristles are very short. This form is also known from Queensland; see S. T. BLAKE, /.c. The plants from the other parts of Malesia and those from continental Asia have long bristles. 3. Eleocharis spiralis (RoTTB.) R. & S. Syst. Veg. 2 (1817) 155; Kuntu, En. 2 (1837) 155; StEup. Syn. 2 (1855) 81; Boeck. Linnaea 36 (1870) 473; BENTH. FI. Austr. 7 (1878) 292, p.p.; CLARKE, FI. Br. Ind. 6 (1893) 627; Ill. Cyp. (1909) t. 35, f. 5-7; Camus, FI. Gén. I.-C. 7 (1912) 85, f. 11, 6-7; MerRR. En. Philip. 1 (1923) 121; Back. Onkr. Suiker. (1928) 153, t. 155: SvENS. Rhodora 31 (1929) 135, t. 188, f. 12: ibid. 41 (1939) 9; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 97, t. 7, f. 4-6; J. Arn. Arb. 28 (1947) 227; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 15; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461. — Scirpus spiralis Rotts. Descr. & Ic. (1773) 45, t. 15, f. 1. — E. variegata (non PReEsL, nec KUNTH) Merk. En. Borneo (1921) 60, p.p. Perennial, with short rhizome and creeping stol- ons. Stems erect, tufted, rather robust, triquetrous at least in the upper part, smooth (somewhat rugulose when dry), not transversely septate, glaucous, 25—60 cm by 2-4 mm. Sheaths thin, rather loose at the apex, pale green to brownish, more or less tinged with purple, obliquely truncate, with a setaceous, up to 4mm long point at the apex. Spike/et cylindrical, somewhat broader than the stem, terete, obtuse, very densely many-flowered, yellowish, 114-314 cm by 3—5 mm. G/umes rather firm, strikingly spirally arranged, very closely packed, (the exposed portion broader than long), tightly appressed, broadly cuneate-obovate to almost quadrate, subtruncate at the apex, faintly many-nerved (the midnerve not very prominent), stramineous to light brown, more or less distinctly brown-zonate within the narrow scarious margins, 234-334 mm long and about as wide. Bristles 4—6, delicate, from much shorter than to about as long as the nut, with weak both antrorse and retrorse teeth, ferrugineous. Stamens 3; anthers linear, shortly apiculate, 1/4-2 mm. Style 2—3-fid. Nut turgidly biconvex (the margins not ribbed), obovate, without neck and apical annulus (or latter indistinct), shining, at first. stramineous, finally deep brown, 14-134 by c. 114mm; epidermal cells transversely oblong-linear to linear, in c. 20 vertical series on either face, not pitted. Stv/e-base triangular, almost confluent with the nut, c. 14 as long and /4 as wide as the nut. Distr. Tropical Africa, Madagascar (here per- haps introduced), Mauritius, Ceylon, S. India, S. China, N. Australia, Queensland, New Caledonia, in Malesia apparently very rare: Malay Peninsula (Singapore), W.-E. Java, in a few localities; Madura; N. Borneo; Philippines (Luzon, Rizal); New Guinea (Papua, Central Distr., Kairuka Subdistr.). Distri- bution map in Rhodora 41 (1939) 5, f. 17. Ecol. In open wet places, pools, swamps, often on clayish soil, at low altitudes (up to 100 m?); almost restricted to brackish or salt localities, sometimes forming extensive, practically pure stands; far inland along salt mudwell near Kesongo (Java, res. Rem- bang). Uses. In Indramaju (W. Java) used for making mats. Vern. Boroslanang (endong), J (Indramaju). Note. Eleocharis spiralis is very near to E. mutata (L.) R. & S. (Scirpus mutatus LINNE, Amoen. Ac. 5, 1759, 391) from America and tropical Africa, and might perhaps better be treated as a subspecies of a pantropic species. The two were united by BOECKELER, Linnaea 36 (1870) 473. E. spiralis differs from E. mutata by the firmer and more sharply truncate glumes, the shorter and thicker spikelets, the smoother nuts with finer markings, and the more irregularly toothed, slenderer bristles. 4. Eleocharis sundaica KERN, Blumea 9 (1958) 220, fale Perennial, with short rhizome and long-creeping stolons ending (always?) in tubers. Stems erect, tufted, rather robust, terete, smooth (rugulose when dry), not transversely septate, 65-85 cm by 3—6 mm. Sheaths loose, membranous, pale green to stramin- eous, often darker at the base, obliquely truncate, muticous. Spike/et cylindrical, terete, subacute, very densely many-flowered, 1/4-2cm by 4-6 mm. Glumes rather firm, very closely packed (the exposed portion broader than long), broadly ovate or sub- orbicular, rounded at the apex, not keeled, faintly many-nerved, stramineous, brown-zonate within the broad scarious margin, c. 4 by (3—)}4 mm, the lowermost empty, embracing the stem. Bristles 5—6, firm, retrorsely scabrous, 2—3 reaching the top of the persistent style-base, other ones slightly shorter to slightly longer than the nut. Stamens 3. Style 2-fid 528 (always?). Nut biconvex, broadly elliptic or broadly obovate, subtruncate, with slightly ribbed angles, slightly reticulate, shining, castaneous when ripe, c. 2 by 124-134 mm; epidermal cells transversely oblong, in c. 30 vertical series on either face. Style-base triangular, flattened, spongy, >4 as long and almost as wide as the nut. Distr. Malesia: Lesser Sunda Is. (Alor, near Naumang). Ecol. Abundant along a lake, at 450 m. Vern. Moliuta. Note. Closely related to E. spiralis, from which it differs by the terete stems, the larger, not almost truncate glumes, the larger nut with the external cells oblong, the much stouter style-base, and the longer and firmer bristles. Apparently also close to E. brassii 8. T. BLAKE, Proc. R. Soc. Queensl. 49 (1938) 154: ibid. 50 (1939) 100, t. 7, f. 16-19, from N. Australia and Queensland. Not seen; according to description and figure mainly differing by the much narrower, ovate glumes, the length of the bristles (4 about as long as the nut, 2 very small or absent), and the hexagonal epidermal cells of the nut in c. 20 vertical rows on either face. 5. Eleocharis ochrostachys SteuD. [in Zoll. Syst. Verz. | (1854) 62, nom. nud.| Syn. 2 (1855) 80; Mia. Fl. Ind. Bat. 3 (1856) 301; Boeck. Linnaea 36 (1870) 452; CLARKE, FI. Br. Ind. 6 (1893) 626; RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 76; Fl. Mal. Pen. 5 (1925) 151; Onwi1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 32; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461. — Scirpus laxiflorus THw. En. Pl. Zeyl. (1864) 435. — E. variegata (PorR.) PREsL var. laxi- flora CLARKE, FI. Br. Ind. 6 (1893) 626; MerR. En. Philip. | (1923) 121, p.p.; KUK. Bot. Jahrb. 59 (1924) 47; ibid. 69 (1938) 257; SveNs. Rhodora 31 (1929) 156, f. 10. — E. subulata Boeck. Flora 41 (1858) 412; Koorp. Exk. Fl. Java 1 (1911) 197. — Scirpus ochrostachys O.K. Rev. Gen. Pl. 2 (1891) 758. — E. laxiflora H. Pretrr. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 169; Svens. Rhodora 41 (1939) 8, t. 537, f. 3; S. T. BLAKE, J. Arn. Arb. 28 (1947) 227. — E. varie- gata (non PRESL, nec KUNTH) Boeck. Linnaea 36 (1870) 470, p.p. (excl. svn.); Koorb. Exk. Fl. Java | (1911) 196; Back. Onkr. Suiker. (1928) 153, p.p. — E. fistulosa (non LINK) BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 15, p.p. — Fig. 35. Perennial, with short rhizome and long stolons. Stems erect, tufted, slender, rigid, terete or obscurely angular, smooth, not transversely septate, 35-60 cm by (34-)2-3(—5) mm. Sheaths membranous, closely appressed, purplish at the base, obliquely truncate, acute or rounded at the apex. Spikelet cylindrical, as broad as to distinctly broader than the stem, somewhat angular, acute, few- to several-flowered, pale green, (S—)10—-20 by 3—4mm. G/umes firm, loosely imbricate, appressed, oblong-ovate, obtuse, somewhat convolute when dry, many-nerved with prominent midnerve, usually brown-zonate within the broad scarious margins, 4-5 by 2-3/4 mm. Bristles 5—7, coarse, up to 2—3 times as long as the nut, retrorsely scabrous in the upper half, yellowish or light brown. Stamens 2-3; anthers linear, 1-1/4 mm, shortly apiculate. Sty/e 2—3-fid. Nut turgidly and unequally biconvex, ribbed on the margins, obovate to broadly obovate, not constricted but with an annular prominence c. 14-34 as wide as the nut at FLORA MALESIANA [ser. I, vol. 7° the apex, shining stramineous to greyish, 1)4-2 by 1-1/4 mm; epidermal cells small, transversely ob- long, in 15—20 vertical series on either face, not pitted, the longitudinal ridges prominent. Sty/e-base deltoid, flat, dark brown, 4-34 mm high. Distr. From India and Ceylon to Thailand, Indo-China, Formosa and the Ryu Kyu Islands, Micronesia (Carolines) and Melanesia (Solomon Is., Fiji), almost throughout Malesia, but often very local; in Java only in the western part, in the Philip- pines only known from Luzon and Mindanao, in the Moluccas only from Halmahera; not yet collected in the Lesser Sunda Is. Distribution map in Rhodora Al(1939) 5, fel Se Ecol. In open wet places, swamps, on margins of lakes, floating islands efc., at low and medium altitudes (up to 1400 m), sometimes dominant. Uses. Sometimes used for making bags (Sumatra E.C., W. Borneo). Vern. Purun, purun tikus, purun dama, rumput sikat, M, ngunu boba (Halmahera); these names may partly refer to E. acutangula. Note. Except for some minor points the collection KALKMAN 4003 from W. New Guinea, Sibil valley, very well answers the description of E. difformis S. T. BLAKE, Proc. R. Soc. Queens}. 50 (1939) 99, of which I have not seen the type. Similar slender specimens with stems not fully | mm wide I have seen from the Malay Peninsula and Borneo. In my opinion they are not specifically distinct from E. ochrostachys, and I doubt whether E. difformis can be upheld. 6. Eleocharis variegata (Poir.) PRESL in Oken, Isis 21 (1828) 269: KunTH, En. 2 (1837) 153; Boeck. Linnaea 36 (1870) 470, p.p.; SVENS. Rhodora 41 (1939) 8, t. 537, f. 4: KERN, Reinw. 4 (1956) 95. — Scirpus variegatus Por. Enc. 6 (1804) 749. Closely related to E. ochrostachys and agreeing with this species in almost every detail, except for the quite different marking of the nut. G/umes often reddish-zonate within the broad, lacerate, scarious margins. Nut glossy, stramineous to orange, without prominent longitudinal ridges, the epidermal cells large, strongly inflated, isodiametric or somewhat horizontally elongated, in 12—15 vertical series on either face. Distr. Tropical Africa, Madagascar, Mauritius, Seychelles, in Malesia only a few times collected: N. Sumatra (S. foot of Mt Piso-piso, NW. of Toba Lake, LORZING 9374) and NE. New Guinea (Western Highlands). Distribution map in Rhodora 41 (1937) Det elite Ecol. In swamps, !400—1500 m. Note. Though in habit very similar to E. ochro- stachys readily recognized by the large inflated outer cells of the nut, according to SVENSON approached nowhere else in the genus except in the Caribbean E. cellulosa Torr. In the Malesian specimens the stems are more or less quadrangular (with one of the sides narrower than the other ones), especially in the upper part. Therefore they agree very well with the description of E. calocarpa CHERM. (Arch. Bot. Caen 4, Mém. 7, 1931, 41) from tropical Africa, of which I have not seen the type. In SVENSON’s opinion E. variegata and E. calocarpa are very closely related, the chief distinction lying in the 4-angled stems of E. calocarpa as compared with the cylindrical stems of E. variegata, which are sometimes trigonous below 1974] Cy PERACEAE (Kern) 529 Fig. 35. Eleocharis ochrostachys STEUD. in foreground gregarious in front of drifting mats of Leersia hexandra and Isachne miliacea and behind open water with Nymphoides indica. Rawa Tembaga near Bekassi(E of Djakar- ta) (photogr. VAN STEENIS, 1941). the inflorescence. It is somewhat surprising that one of the collections cited by SVENSON under E. calocarpa is annotated ‘culms terete’. As in my opinion angular stems alone are an unsatisfactory character for specific separation, I referred the Malesian speci- mens to E. variegata. 7. Eleocharis dulcis (BURM. f/f.) HENSCHEL, Vita Rumph. (1833) 186; Merr. Int. Rumph. (1917) 104; Sp. Blanc. (1918) 82; Brown, Min. Prod. Philip. For. 2 (1921) 250, t. 8; En. Philip. 1 (1923) 119; Back. Onkr. Suiker. (1928) 152, t. 154: SveNs. Rhodora 31 (1929) 158, t. 188, f. 16; ibid. 41 (1939) 11; STEEN. Arch. Hydrobiol. Suppl. 11 (1932) 285, f. 15, 36, 53; OcusE, Veg. Dutch E. Ind. (1931) 219, f. 132; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 103, t. 8, f. 6-9; J. Arn. Arb. 28 (1947) 227; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 12; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 35; J. Fac. Sc. Un. Tokyo III, 8 (1961) 97; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 460. — Cyperus dulcis RuMPH. Herb. Amb. 6 (1750) 7, t. 3, f. 1; cf. TRIN. Clav. Agr. (1822) 120. — Andropogon dulce BuRM. f. Fl. Ind. (1768) 219. — Scirpus plantaginoides Rotts. Descr. & Ic. (1773) 45, t. 15, f. 2. — Scirpus plantagineus Retz. Obs. 5 (1789) 14. — Hippuris indica Lour. Fl. Cochinch. (1790) 16. — E. planta- ginea R. & S. Syst. 2 (1817) 150; Mia. FI. Ind. Bat. 3 (1856) 302; Naves, Nov. App. (1882) 306; BoEck. Linnaea 36 (1870) 474, p.p.; CLARKE, FI. Br. Ind. 6 (1893) 625; Ill. Cyp. (1909) t. 33, f. 1-5; t. 34, f. 7; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 75; Koorp. Exk. Fl. Java 1 (1911) 196; ibid. 4 (1922) f. 243; Camus, Fl. Gén. I.-C. 7 (1912) 82; Rip. FI. Mal. Pen. 5 (1925) 150. — Scirpus tuberosus ROXB. Fl. Ind. 1 (1820) 213.— Scirpus tumidus Roxs. Fl. Ind. | (1820) 215. — E. tuberosa R.& S. Mant. 2 (1824) 86; Mia. FI. Ind. Bat. 3 (1856) 302; NaAves, Nov. App. (1882) 306. — E. tumida R. & S. Mant. 2 (1824) 86; Naves, Nov. App. (1882) 306. — E. equisetina PREsL, Rel. Haenk. | (1828) 195; Steub. Syn. 2 (1855) 82; Mig. Fl. Ind. Bat. 3 (1856) 302: 530 Naves, Nov. App. (1882) 306; CLARKE, FI. Br. Ind. 6 (1893) 626; Philip. J. Sc. 2 (1907) Bot. 89; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 75; Merr. Fl. Manila (1912) 114; Camus, Fl. Gén. I.-C. 7 (1912) 83; MerR. En. Philip. 1 (1923) 120; Rip. Fl. Mal. Pen. 5 (1925) 150; SveNs. Rhodora 31 (1929) 161, t. 188, f. 14; KUk. Bot. Jahrb. 69 (1938) 257; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 104, t. 8, f. 10-13. — Carex tuberosa (non DEGL.) BLANCO, FI. Filip. (1837) 35; ed. 2 (1845) 24; ed. 3, | (1877) 45, t. 15. — E. sphacel- ata (non R.Br.) BENTH. FI. Austr. 7 (1878) 292, p.p.— E. plantaginoidea W. F. WiGutT, Contr. U.S. Nat. Herb. 9 (1905) 267. — E. plantaginoides Domin, Bibl. Bot., Heft 85 (1915) 445. — E. indica DRucE, Rep. Bot. Exch. Club Br. Isl. 4 (1917) 621. — Fig. 36. Perennial; rhizome short with elongated stolons sometimes bearing subglobose, brownish to blackish, zoned tubers c. | cm o(up to 4 cm in cultivated races). Stems erect, tufted, slender, terete, finely longitudi- nally striate, conspicuously transversely septate. smooth, greyish to shining dark green, 40—200 cm by (1-)3-10 mm. Sheaths membranous, oblique at the apex, purplish. Spike/et cylindrical, as broad as or somewhat broader than the stem, terete, rather obtuse to acute, many-flowered, 114-6 cm by 3-6 mm. Glumes numerous, firm, rather densely im- bricate, appressed, oblong-obovate to oblong, obtuse to truncate, finely many-nerved with distinct mid- nerve, greyish green or stramineous, with a narrow scarious margin at the apex and a faint brown zone within it, 4-64 by 2-3mm. Bristles 6-8, from about as long to twice as long as the nut, retrorsely Fig. 36. Eleocharis dulcis (BURM. f.) HENSCHEL grown in a muddy pool near Wapanamanda (Papua), used for women’s skirts (photogr. HOOGLAND). FLORA MALESIANA [ser. I, vol scabrous, light brown. Stamens 3; anthers linear, 2-3 mm, the connective with a distinct appendage. Style 2—3-fid. Nut biconvex, obovate to broadly obovate, not costate on the margins, hardly con- stricted at the summit but with an inconspicuous annular thickening about 14 as wide as the nut, usually with a shallow longitudinal furrow on the anticous side, shining brown, 1)4-2 by L4-L4% mm: epidermal cells very small, hexagonal to oblong- hexagonal. Style-base triangular, flat, dark brown, c. % as wide and /¥ as long as the nut, the remainder of the long style often persistent. Distr. Tropical West Africa, Madagascar, SE. Asia (from India to S. China, Formosa, the Ryu Kyu Is. and Japan), Australia (N. Australia, Queensland), Micronesia (Marianas, Carolines), Melanesia (New Caledonia, Fiji, Samoa), in Malesia: Sumatra, Banka, Riouw Arch., Malay Peninsula, Java, Kan- gean, Lesser Sunda Is. (Bali, Sawu, Alor, Timor), Philippines (Luzon, Negros, Mindanao), Celebes, Moluccas (Key & Aru Is.), and New Guinea (W. New Guinea, Papua). Ecol. In open wet places, both in salt or brackish and in fresh water swamps, in pools, ponds, rice- fields; often forming pure stands surrounding the open water; 0—-1350 m; on Mt Diéng (Central Java) at 2000 m (not flowering); in most parts of the area of local occurrence. Uses. The boiled tubers of the wild form are sold in large quantities at the Djakarta and Manila markets in the months of August to December (kulub, S). They are usually made into chips, kripik (émping téki). See OcHsE, /.c. In the warmer parts of China the species has long been cultivated and developed into a strain yielding tubers which are superior in size and sweetness to those produced by the wild plants and esteemed as a nutritious delicacy in Chinese cookery. They are also extensively eaten raw as a substitute for fresh fruits because of the crisp apple-like flesh. The much larger tubers of this cultivated form (‘matai’ = horse’s hoof; E. tuberosa R. & S. s.s.) are imported in Indonesia. Recently there is vivid interest in the establishment of this species as a new crop in the United States of America (see HopGE in Econ. Bot. 10, 1956, 49-65, with bibliography). In Sumatra (Padang Highlands) and N. Celebes the species is also cultivated; here the stems are used for making sleeping mats (this use is also reported from Halmahera). By some Papuan tribes they are used for making skirts for the women. Vern. Chinese waterchestnut, E; M; begau, biga, Sum. W. C., mansiro buluk, m. bolong, m. kuning, Menangk., putjut, Banka, pepérétan, Djakarta; baba- wangan beureum, lembang, S; bélané péret, titimbihen, Minah., tingkal, Halmahera, /orio, Biagi, Papua; Philippines: cabezas de negrito, Sp., apulid, Tag., Bik., kalangub, C. Bis., potok, Tag.; special names for the tuberiferous form (according to HEyNE): M: tiké, Djakarta, tjikai, Palemb.; dékéng, téki-tiké, J, pangoké, tékéré, Mak., goro, pagoro, péru-péru lénrong, Bug. Notes. Originally SvENSON (1929) distinguished between E. dulcis s.s. and E. equisetina PRESL, the latter characterized by the punctulate (not reticulate) nuts; however, he considered them conspecific in 1939. BLAKE (1939, 1947) distinguished E. equisetina by stolons never tuberiferous, harder stems usually 1974] less flattened in herbarium specimens, broader, shorter, subtruncate glumes somewhat incurved when dry, long-apiculate anthers, bright brown nut with acutely costulate margins and more evenly arranged external cells, weaker, usually much shorter hypogynous bristles which are quite free from another (not conspicuously connate at the base), and non- prominent receptacle. Apart from the difficulties encountered in naming the numerous incomplete collections (without rhizomes, or in flower only), I fail to divide satis- factorily the Malesian materials into two groups, on account of the many intermediates. Undoubtedly several races are involved in this widely distributed and extensively cultivated, extremely polymorphic species; possibly additional collections of whole plants with ripe fruits will enable to distinguish between them. The stems of the cultivated form are much more robust than those of the wild plants; cytological study must show whether polyploidy is involved. E. kuroguwai Ouwl1, J. Jap. Bot. 12 (1936) 654, from Japan, characterized by the narrowly oblong glumes and the linear appendage of the connective, apparently also belongs here; according to Koyama (1961, /.c.) it is a good species, char- acterized by the less imbricate, rounded glumes, the conspicuously annulate nuts with narrower style- base, and the spikelets gradually narrowed towards the apex. 8. Eleocharis sphacelata R. Br. Prod. (1810) 224: Boeck. Linnaea 36 (1870) 475; BENTH. FI. Austr. 7 (1878) 292, p.p.; CLARKE, III. Cyp. (1909) t. 34, f. 1-6; Svens. Rhodora 31 (1929) 160, t. 188, f. 15; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 102, t. 8, f. 1-5. Very closely allied to E. dulcis. Rhizome woody, very stout, shortly creeping, up to | cm thick, never producing tubers. Stems in a close linear series, up to 2m tall and 12 mm thick. Spike/et acute, 8-9 mm wide. Glumes densely packed, light brown, 7!4-9 mm long. Bristles 8-10, at least partly reaching the top of the style-base or overtopping it. Anthers 344-4 mm long. Nut broadly obovate to orbicular, with somewhat costulate margins and a definite longitudinal furrow on the anticous side, light yellowish to tawny or pale brown, 2)4-2)4 by 1%- 2% mm. Style-base from 4 to fully as long as the nut. Distr. Australia (not in the Western part), Tasmania, New Zealand, in Malesia: NE. New Guinea, Western Highlands District (Sirunke, Wabag area, Kandep valley and northern slopes of Sugarloaf complex). Ecol. In swamps, c. 2200—2900 m. Uses. Used by native women for making rush skirts. Vern. Kur, Enga language. Notes. Distinguishable from E. dulcis by its stout horizontal rhizome without tuber-bearing stolons, broader spikelets, longer glumes, and slightly larger nuts. The stems are always stout, but equally stout stems occasionally occur in E. dulcis. As only two Malesian collections are known, I have followed BLAKE’s monograph of the Australian CYPERACEAE (Kern) 531 Eleocharis species in treating it as a separate species extending from the Australian mainland to Tas- mania, New Zealand, and New Guinea, though it probably may better be treated as a subspecies of E. dulcis. 2. Series Multicaules Svens. Rhodora 41 (1939) 4, descr. angl.; ex KERN‘. — Sect. Leiocarpeae C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 106, p.p. — Ser. Tuberculosae & ser. Inter- mediae SVENS. Rhodora 31 (1929) 129, p.p. — Type species: E. multicaulis (SM.) SM. 9. Eleocharis tetraquetra NEES in Wight, Contr. Bot. Ind. (1834) 113; KUNTH, En. 2 (1837) 150; BoEck. Linnaea 36 (1870) 447; BENTH. FI. Austr. 7 (1878) 294; Naves, Nov. App. (1882) 307; CLARKE, FI. Br. Ind. 6 (1893) 630; Ill. Cyp. (1909) t. 37, f. 17-20; Koorpb. Exk. Fl. Java | (1911) 197; ibid. 4 (1922) f. 247; Merr. En Philip. | (1923) 121; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 111, t. 9, f. 12-15; SvensS. Rhodora 41 (1939) 99; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 13; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 36; KoyAMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 38; J. Fac. Sc. Un. Tokyo III, 8 (1961) 91; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461; STEEN. Mt FI. Java (1972) t. 14: 13. — E. erythrochlamys Mia. FI. Ind. Bat. 3 (1856) 300. — Scirpus tetraqueter THWAITES, En. PI. Zeyl. (1864) 434; O.K. Rev. Gen. Pl. 2 (1891) 757 (‘tetraquetrus’). — E. tetraquetra var. micranthera Oxuwt, Bot. Mag. Tokyo 56 (1942) 201. Perennial, with short rhizome; long slender stolons sometimes present. Stems erect, tufted, slender, rigid, acutely 4-angled, with usually prominently ribbed angles and finely longitudinally striate or rather prominently ribbed sides, smooth, (15—)30—70 cm by 1-1)4 mm. Sheaths herbaceous, closely appressed, purplish below, the upper one slightly thickened and brownish at the mouth, truncate or somewhat oblique, shortly mucronate just below the apex. Spikelet ovoid to oblong-ellipsoid, broader than the stem, often somewhat inclined, terete, acute, densely many-flowered, ferrugineous to brown, 10-20 by 3-5 mm. Glumes rather firm, rather densely imbricate, closely appressed when young, somewhat spreading with age, elliptic to oblong- elliptic or lanceolate, obtuse, not or only faintly keeled, 1-nerved, ferrugineous with paler keel and narrow scarious margins, 3—4 by c. 2 mm. Brist/es 6, coarse, flat, about as long as or slightly exceeding the nut including the style-base, very densely retrorsely scabrous or subplumose, rufous. Style 3-fid. Stamens 2-3; anthers linear, minutely apiculate, 1-1/4 mm. Nut obtusely trigonous, compressed, obovoid, not constricted below the apex, shining, yellowish or olivaceous to brown, obsoletely reticulate by the small, vertically oblong epidermal cells, 114-1/4 by %o-1% mm. Style-base large, deltoid to submitri- form, compressed above, somewhat pyramidal below, spongious, fuscous, c. 14-34 as long and 4-% as wide as the nut. Distr. From Ceylon, India, China, Formosa, the Ryu Kyu Is. and Japan to Australia (Queensland, ' Culmi vulgo subvalidi. Vagina summa ore truncata vel oblique secta, saepe mucronata. Stylus trifidus. Nux obtuse trigona, angulis ecostatis. 532 FLORA MALESIANA N. S. Wales), in Malesia: Sumatra, Java, Lesser Sunda Is. (Flores), Philippines (Luzon, Mindanao), SW. Celebes, New Guinea. Distribution map in Rhodora 41 (1939) 75, f. 46. Ecol. Open swampy places, at medium and high altitudes (in Java between 1000 and 2200 m, in the Philippines between 650 and 1800 m, in Sumatra Eastcoast Res. descending to 450 m), rarely ascend- ing to 2700 m (W. and NE. New Guinea). Vern. Djukut djarum, S, tihe-tihe, tihe alk, si gonde langet, si martihe-tihe, Sum. E.C.; New Guinea: Kuli, Enga lang., patso, Mendi lang. Note. In VAN STEENIS 4058 (in BO), mentioned by BACKER (1949, /.c.) as being similar in habit to E. tetraquetra but differing from it by the distinctly reticulate nuts, I fail to see any difference with the other specimens of Javan. E. tetraquetra. 10. Eleocharis attenuata (FRANCH. & SAV.) PALLA, Monde d. Pl. 12 (1910) 40; Hara, J. Jap. Bot. 19 (1943) 153; ibid. 20 (1944) 333; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 91; TANG & WANG, FI. Reip. Pop. Sinic. 1! (1961) 58.— Scirpus attenuatus FRANCH. & Sav. En. Pl. Jap. 2 (1877) 110; (1879) 543. — E. major Hara, J. Jap. Bot. 11 (1935) 820, f. 24. — E. laeviseta NAKAI var. major HaRA, J. Jap. Bot. 14 (1938) 521; OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 42.— E. pellucida PRESL f. attenuata Ounwt, I.c. 40, quoad basion. — E. pellucida (non PRESL) S. T. BLAKE, J. Arn. Arb. 28 (1947) 227. Perennial (always?). Rhizome short, without sto- lons. Stems densely tufted, erect or obliquely erect, slender, terete, longitudinally striate-costate, smooth, 20-60 cm by 14-1 mm. Lower sheaths obliquely truncate, reddish brown, uppermost truncate or somewhat oblique, distinctly mucronate. Spikelet ovoid or narrowly ovoid, obtusish, terete, densely many-flowered, 5-10 by 3—5mm. Glumes mem- branous, appressed, obovate, obtuse with rounded apex, convex (scarcely keeled), !-nerved, greenish white often tinged with purple, with ay or green midnerve and scarious margins, 2—2!4(—3) mm long. Bristles 6, slender, distinctly to obscurely retrorsely scabrous [smooth in f. laeviseta (NAKAI) HARA (E. laeviseta NAKAI in Fedde, Rep. 13 (1914) 246), not known from Malesia], as long as or slightly longer than the nut, but not overtopping the style-base. Stamens 2-3; anthers short-linear, c. #4 mm, shortly apiculate. Style 3-fid. Nut obtusely trigonous, obo- void, rounded at the apex, somewhat attenuate towards the base, smooth, shining olivaceous to brown, 1-1/% by 4 44 mm. Style-base depressed- deltoid-pyramidal, 3% to almost as wide as the nut. Distr. Japan, Ryu Kyu Is., S. Korea, Quelpaert, E. and NW. China, in Malesia: New Guinea (W. New Guinea: Lake Habbema; Territory of N.G.: W. and E. Highlands). Ecol. In swampy ground: along pools, swampy places in forests and grassy fields, on creek-banks, 1650-2800 m. Note. Very closely related to E. congesta and possibly only a geographical (E. Asian) race of it. 11. Eleocharis congesta D. Don, Prod. Fl. Nepal. (1825) 41; KuNTH, En. 2 (1837) 152; CLARKE, FI. Br. Ind. 6 (1893) 630; Camus, Fl. Gén. I.-C. 7 (1912) 88; SveNS. Rhodora 41 (1939) 102, t. 546 f. 2; Koyama, [ser. I, vol. 7° Contr. Inst. Bot. Un. Montréal n. 70 (1957) 37; J. Fac. Sc. Un. Tokyo III, 8 (1961) 89; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461. — E. pellucida PRESL, Rel. Haenk. | (1828) 196; Mia. FI. Ind. Bat. 3 (1856) 301; Naves, Nov. App. (1882) 307; Svens. Rhodora 41 (1939) 101; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 37. — E. afflata Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.] Syn. 2 (1855) 76; Mig. Fl. Ind. Bat. 3 (1856) 299; Naves, Nov. App. (1882) 307; CLARKE, FI. Br. Ind. 6 (1893) 629; Philip. J. Sc. 2 (1907) Bot. 90; Camus, Fl. Gén. I.-C. 7 (1912) 87, f. 13; 5; Murr. En. Philip. 1 (1923) 119; Back. Onkr. Suiker. (1928) 154, t. 157; Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 14. — E. sub- prolifera Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 80; Mig. Fl. Ind. Bat. 3 (1856) 300; Boeck. Linnaea 36 (1870) 426; Koorp. Exk. Fl. Java | (1911) 197; Atlas (1922) f. 246. — E. subvivipara Boeck. Linnaea 36 (1870) 424; CLARKE, Fl. Br. Ind. 6 (1893) 629; Ill. Cyp. (1909) t. 37, f. 13- 16. — Scirpus afflatus BENTH. Fl. Hongk. (1861) 394; O.K. Rev. Gen. Pl. 2 (1891) 757. — E. japonica Mig. Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 142. — Scirpus zollingeri O.K. Rev. Gen. Pl. 2 (1891) 758. — Scirpus subviviparus O.K. l.c. Annual of perennial, without stolons. Stems tufted, erect or oblique, slender, often capillary, striate or somewhat angular- ribbed, smooth, bright green, 5-40cm by 4-1 mm. Sheaths appressed, often purplish at the base, muticous or with a minute mucro at the oblique or very oblique mouth. Spikelet ovoid or lanceolate, acutish, terete, densely several- to many-flowered, 3-9 by 1/4-3 mm, often pro- liferous at the base. Glumes membranous, appressed, oblong-ovate to lanceolate, obtuse, scarcely keeled, I-nerved, pale ferrugineous, often tinged with red, with ig midnerve and whitish, scarious margins, 14-2 mm long. Bristles 6, slender, retrorsely scab- rous, slightly to distinctly longer than the nut, at first whitish, finally ferrugineous. Stamens 2(-—3); anthers linear, shortly apiculate, 7{9—/4 mm long. Style 3-fid. Nut trigonous, obovoid, smooth or obsoletely longitudinally striolate, shining, yellowish green to olivaceous, %—-1 by 4- 34 mm: epidermal cells longitudinally oblong. Sty/e-base shortly pyra- midal, triangular, as long as or slightly longer than wide, ye 4, as broad as the nut. Distr. SE. and E. Asia (from India to China and Japan), in Malesia: N. and Central Sumatra, Malay Peninsula (P. Penang; Selangor: Kuala Lumpur, here introduced ?), Java (in Central and E. Java less common than in W.), Lesser Sunda Is. (Flores), N. Borneo (according to MERRILL, En. Born. p. 59; not seen), Philippines (Luzon, Babuyan Is.), SW. and Central Celebes. Ecol. In shallow water, swampy places, ditches, wet rice-fields, 5|0—2800 m. Vern. Bulu mata munding, S, dongdoman, J; Philippines: hallopnot, If. Notes. E. congesta and E. pellucida are treated as specifically distinct by SvENSON, but distinguished only by the thicker, rigid stems of the former. KoyaMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 89, assigns varietal rank to E. pellucida [E. congesta var. japonica (MiQ.) KoyAMA], distinguishing it from E. congesta var. congesta by the capillary stems, the small spikelets up to7 mm by 14% mm, and the small, less than 11% mm long glumes. 1974] CYPERACEAE (Kern) 533 However different in habit the small, annual plants of the wet rice-fields in the lowlands (E. pellucida) are from the much coarser, apparently perennial ones occurring in more natural habitats at medium and high altitudes (E. congesta s.s.), | fail to trace a dividing line between them. CLARKE’s suggestion, E. congesta might be a western variety of E. afflata (= E. pellucida), cannot be maintained, as E. congesta s.s. is widely distributed in Malesia, although of much rarer occurrence than E. pellucida. The group of spp. 9-11 is a very difficult one. F. attenuata is not always easily distinguishable from E. congesta. Also E. tetraquetra is sometimes identifiable only with difficulty, as the quadrangular nature of its stems is not always clearly pronounced, and the hypogynous bristles vary from densely subplumose to scabrous. The length of the nut may serve as an auxiliary character. 3. Series Acutae S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 114, descr. angl.; ex KERN'.— Type species: E. acuta R.Br. 12. Eleocharis acuta R. Br. Prod. (1810) 224; STEUD. Syn. 2 (1855) 82; BENTH. Fl. Austr. 7 (1878) 294, excl. var.; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 114, t. 10 f. 1-7; SveNs. Rhodora 41 (1939) 102, t. 538 f. 2. — E. mucronulata Nees, Ann. Nat. Hist. I, 6 (1841) 46; Steup. Syn. 2 (1855) 82; Boeck. Linnaea 36 (1870) 466. Perennial, with slender, creeping stolons. Stems erect, in distant tufts, rigid, terete, longitudinally striate-sulcate, not transversely septate, up to 90 cm by 1—2(-—3)mm. Sheaths herbaceous or rigidly membranous, appressed, more or less purplish at the base, the uppermost truncate, somewhat thickened at the often dark brown mouth, prominently mucro- nate. Spikelet oblong-ovoid to linear, broader than the stem, terete, acute, rather densely many-flowered, dark brown or variegated with brown, 15—30 by 3-7 mm. Glumes membranous, rather densely im- bricate, closely appressed when young, somewhat spreading with age, ovate-lanceolate or oblong-ovate, subobtuse to very acute, faintly keeled, l-nerved, deep brown or reddish brown with greenish back and whitish, scarious margins, 3!4-5’4 mm long, the two lowermost firmer, shorter, rounded, empty. Bristles 6—7(—9), rather slender, somewhat unequal, about as long as or slightly exceeding the style-base, strongly retrorsely scabrous, occasionally ill-devel- oped. Stamens 3; anthers linear, 114—2)4 mm incl. the long, prominent appendage of the connective. Stv/e 3-fid. Nut (not present in the only Malesian collec- tion) plano-convex or biconvex, broadly obovate, not costate on the margins, shining yellowish to brown, 174-1% by 1-1/4 mm, the epidermal cells minute, shortly vertically oblong. Sty/e-base ovate or triangular, compressed, cellular, whitish or finally discoloured, 7~—*% by 4-2% mm. Distr. Australia, Tasmania, New Zealand, Nor- folk Island, in Malesia: New Guinea (Territory of New Guinea, Western Highlands, Yobobos grass- land area, source of Lagaip R., Lagaiam Subdistr.). Ecol. On sandy bank along river, c. 2500 m. Vern. Guli, Enga language, Kepilam. Note. The determination remains somewhat doubtful as long as no fruits of the New Guinean plants are known. According to S. T. BLAKE, l.c., the most variable of the Australasian species. The Malesian plants differ from the few Australian ones I have seen in the slightly longer glumes and the c. 14 mm long, more or less bristly appendage of the connective. In the Australian specimens the glumes are up to 414 mm long and the c. 4mm long appendage of the connective is smooth or almost so. 4. Series Tenuissimae (CLARKE) SVENS. Rhodora 31 (1929) 129. — Sect. Leiocarpeae a. Tenuissimae C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 106. — Type species: E. tenuissima CLARKE — Distribution map in Rhodora 39 (1937) PAN he 4a. Subseries Leiocarpeae (CLARKE) SVENS. /.c. — Sect. Leiocarpeae C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 106. — Type species: E. nigrescens (NEES) STEUD. 13. Eleocharis nigrescens (NEES) STEUD. Syn. 2 (1855) 77; CLARKE, Ill. Cyp. (1909) t. 38, f. 1-4; SvENS. Rhodora 39 (1937) 223, t. 462, f. 5-7; S. T. BLAKE, Proc. R. Soc. Queens]. 50 (1939) 108, t. 9 f. 1-4. — Scirpidium nigrescens Nees [Linnaea 9 (1834) 293, nom. nud.] in Mart. Fl. Bras. 2, 1 (1842) 97. — ? E. schweinfurthiana Boeck. Flora 62 (1879) 562; SvENS. Rhodora 39 (1937) 252 t. 461, f. 13, saltem quoad specim. philipp. — E. microcarpa (non Torr.) CLARKE, Philip. J. Sc. 2 (1907) Bot. 91. — ? E. perrieri CHERM. Bull. Soc. Bot. Fr. 73 (1926) 554; Fl. Madag. fam. 29 (1937) 213, t. 19, f. 10—12; SvENs. Rhodora 39 (1937) t. 461, f. 14, in syn. Annual (or perennial?). Stems erect or obliquely erect, tufted, filiform, obscurely quadrangular, sul- cate, punctate, 3—10(—20) cm by c. 14 mm. Sheaths thinly herbaceous, appressed, obliquely truncate, readily disintegrating, stramineous or reddish. Spike- let ovoid, obtusish, densely many-flowered, 2—3(—5) by 1-114 mm. Glumes thinly membranous, spreading in fruit, ovate-elliptic, obtuse to slightly emarginate, keeled (the keel curved in profile), 1- nerved, yellow- ish, purplish puncticulate (or brownish), 8 1 by 4% mm. Bristles wanting (or half as long as to nearly equalling the nut, light brown, obscurely toothed). Stamens 1(—2); anther oblong, )4(—)4) mm long. Style 3-fid. Nut triquetrous with costulate angles, obovoid, smooth, obscurely reticulate, shining, stra- mineous to light brown, %4-7%, by 4-% mm. Style-base very short, pyramidal or depressed, apiculate, somewhat narrower than the nut. Distr. Southern United States to Mexico and Brazil, tropical Africa and Madagascar, Australia (N. Australia, Queensland, S. Australia); in Malesia: Philippines (Central Luzon, about 1900, LOHER 5193). Map in Rhodora 39 (1937) 224. Notes. Originally the LOHER specimens in the Kew Herbarium were determined by CLARKE as E. microcarpa TorRR.; the name E£. schweinfurthiana BOECK. was given in synonymy. The N. American E. microcarpa, though closely related, seems to be Culmi rigidi, subvalidi. Vagina summa ore truncata, distincte mucronata. Stylus trifidus. Nux lenticularis, angulis ecostatis, cellulis extimis minutis oblongis. 534 FLORA MALESIANA [ser. I, vol. 7% clearly distinct by the non-costulate nuts. CLARKE’s remark (1907, /.c.) ““No 5193 LOHER may be taken to be a depauperated state of the common Eleocharis afflata Strub.” [= E. congesta D. Don] is certainly wrong. In 1932 SvENSON annotated the specimens as follows: “Identical with PERRIER DE LA BATHIE 17947 and 17282 from Madagascar. It is very close to E. nigrescens KUNTH, from which it differs in stouter growth and in having an achene which is smooth or at most with traces of black striolation. E. nigrescens has a clearly marked reticulation on the achene.” In Rhodora 39 (1937) 224 SveNson referred E. perrieri with a question mark to E. nigrescens (PERRIER DE LA BATHIE 17947), on p. 252 it is cited as a synonym of E. schweinfurthiana (same collector 17282) and LOHER’s specimens considered identical in outward aspect and in minute details of the nut with the type collection of E£. schweinfurthiana. How- ever, in SCHWEINFURTH 1949 (type collection of E. schweinfurthiana) the bristles are about half as long as the nut, in LOHER 5193 they are absent. In SVEN- SON’s key (/.c. 214) absence of bristles leads to E. nigrescens. In the Australian plants bristles are also wanting (S. T. BLAKE, /.c.). According to SVENSON E. schweinfurthiana stands between E. nigrescens and E. anceps RIDL., the latter characterized by the somewhat broader stems (‘4-14 mm wide) and the slightly larger nuts (% mm long), whereas E. helenae Busc. & MUSCHL. is intermediate between E. schweinfurthiana and E. anceps. NELMES (in /itt.) saw specimens of an Eleocharis from N. Rhodesia agreeing with E. schweinfurthiana in every way, except that they had no bristles in some flowers, but up to 3 short white bristles in other flowers. Therefore he took the Philippine collection for E. schweinfurthiana. As I fail to see the distinguishing characters between all these ‘species’, I provisionally refer the Philippine plants to E. nigrescens, of which I have not seen the type. In the above description the data between brackets refer to extra-Malesian specimens. 4b. Subseries Chaetarieae (CLARKE) SVENS. Rhodora 31 (1929) 129. — Chaeto- cyperus NEES in Wight, Contr. Bot. Ind. (1834) 95; Linnaea 9 (1834) 289, p.p.— Sect. Chaetarieae C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 106, p.p. — Type species: E. chaetaria R. & S. 14. Eleocharis retroflexa (Porr.) Urs. Symb. Ant. 2 (1900) 165; Merr. En. Philip. 1 (1923) 120; BAcK. Onkr. Suiker. (1928) 155, t. 160; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 14; SveNs. Rhodora 39 (1937) 236, t. 461, f. 11; KERN in Back. & Bakh. f. FI. Java 3 (1968) 460. — Cyperus setaceus Retz. Obs. 5 (1789) 10; VAHL, En. 2 (1806) 305, non E. setacea R. Br. Prod. (1810) 225. — Scirpus pygmaeus LAMK, Ill. 1 (1791) 139, non E. pygmaea Torr. (1836). — Scirpus retroflexus Pork. in Lamk, Enc. 6 (1804) 753. — E. chaetaria R. & S. Syst. Veg. 2 (1817) 154; KunTH, En. 2 (1837) 140; Mor. Syst. Verz. (1846) 96 (var. subbiflora); Zoi. Syst. Verz. 1 (1854) 62; Boeck. Linnaea 36 (1870) 428, p.p.; NAves, Nov. App. (1882) 307; CLarRKE, FI. Br. Ind. 6 (1893) 629; Philip. J. Sc. 2 (1907) Bot. 90; Rmp_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 77; Koorp. Exk. Fl. Java (1911) 197; Atlas (1922) f. 245; Camus, Fl. Gén. I.-C. 7 (1912) 87, f. 12, 7-8; Ripe. Fl. Mal. Pen. 5 (1925) 151; Svens. Rhodora 39 (1937) 250, t. 461, f. 10; S. T. BLAKE, Proc. R. Soc. Queens}. 58 (1947) 42; KOYAMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 37; J. Fac. Sc. Un. Tokyo III, 8 (1961) 88.— Chaetocyperus setaceus Nees, Linnaea 9 (1834) 289: Mia. FI. Ind. Bat. 3 (1856) 298; non Kurz, Nat. Tijd. N. I. 27 (1864) 223. — Chaetocyperus limnocharis NEES in Wight, Contr. Bot. Ind. (1834) 96. — Chaetocyperus arenicola STEUD. Syn. 2 (1855) 73. — E.? confer- voides Mia. FI. Ind. Bat. 3 (1856) 303, quoad specim. jav., non STEUD. Syn. 2 (1855) 32, nec Scirpus con- fervoides Porr. (basionym). — E. setacea [non R. BR. Prod. (1810) 225] Merr. En. Born. (1921) 59." Annual. Stems very slender, tufted, often curved, angular, 4—5-ribbed, puncticulate, light green, 5—20 cm by 4-lK mm. Sheaths reddish at the base, whitish-scarious at the apex. Spikelet ovoid, obtuse, more or less compressed, loosely few-(3—10)-flowered, 2-5 by 1'4-2/4 mm. G/umes membranous, spreading in fruit, ovate, obtuse, keeled, with distinct greenish midnerve, purplish sides, and scarious margins, 24-34% by 14-2 mm, the lower ones almost distichous. Bristles 6, rather delicate, unequal, the longest usually longer than the nut, retrorsely scabrous, whitish to yellowish, occasionally reduced to obsolete. Stamens 3; anthers linear, minutely apiculate, 14-34 mm. Style 3-fid. Nut equilaterally triquetrous, urceolate, truncate or slightly convex at the apex, conspicuously cancellate, costulate on the angles, dull stramineous to fumose, % —1 by Ao-Ao mm; epidermal cells conspicuous, deeply pitted, hexagonal or roundish, in 6—10 vertical series on each face. Style-base pyramidal, in the Malesian specimens usually much depressed and blunt at the apex, c. )4 as high as and equal in width to the nut, 3-lobed, the lobes decurrent on the angles of the nut. Distr. Probably pantropic: tropical America, Asia, and Australia (Queensland); common through- out Malesia. The records for tropical Africa (CLARKE, Fl. Trop. Afr. 8, 1902, 408) SVENSON partly refers to E. brainii Svens. Rhodora 39 (1937) 251, remarking that it is questionable whether E. chaetaria actually occurs in tropical Africa. Map of E. retroflexa s.s. in Rhodora 39 (1937) 237. Ecol. In wet muddy places: along streams, along humid road-sides, in shallow pools, in rice-fields, 0—1600 m. Vern. Rumput kakamatan, M, djadjaruman, djukut bulu mata, S, duhut tilam, Asahan, lumut, Borneo, Minahassa, djukut kambing, bulu babi, bulu idung, E. Borneo, tamomo oé, Celebes. Notes. In running water with hairlike stems and proliferous spikelets (E. confervoides of MIQUEL). In shallow, stagnant water often propagating by re- curving or decumbent stems developing young plants in the axils of the glumes. 1 MERRILL I.c. cited “(ReTz.) R. BR. Prodr. (1810) 224”. Here BRown did not make a combination based on Cyperus setaceus RETZ.; E. setacea R. Br. Prod. (1810) 225 is a quite different species. 1974] SvENSON thinks E. retroflexa of the New World and E. chaetaria of the Old World are clearly distinct, E. chaetaria having a much lower and blunter style-base, and larger and deeper markings on the nut. Apart from the question whether these trifling characters are sufficient to justify specific separation, it may be remarked that in the Malesian specimens the style-base is variable in shape and sometimes much approaches that of American ones. Apparently this is also the case in the Australian specimens (see S. T. BLAKE, /.c.). Already KUNTH (1837) remarked that E. retroflexa differs from E. chaetaria only in the longer, more acute style-base, and in its native country. Agreeing with his ‘species vix servanda’ | think it desirable to unite the two. 5. Series Aciculares (CLARKE) SvENS. Rhodora 31 (1929) 128. — Sect. Aciculares C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 105. — Type species: E. acicularis (L.) R. & S. — Distribution map in Rhodora 41 (1939) 2. 15. Eleocharis acicularis (L.) R. & S. Syst. Veg. 2 (1817) 154; KuntH, En. 2 (1837) 141; Boeck. Linnaea 36 (1870) 431; Merr. Philip. J. Sc. 9 (1914) Bot. 264; En. Philip. 1 (1923) 119; STEEN. Arch. Hydrobiol. Suppl. 11 (1932) 284; Svens. Rhodora 31 (1929) 184; ibid. 41 (1939) 18, t. 59, f. 1, 9 b; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 35; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 36; J. Fac. Sc. Un. Tokyo III, 8 (1961) 87.— Scirpus acicularis LINNE, Sp. PI. 1 (1753) 48. Perennial, with capillary stolons usually forming dense mats. Stems erect, capillary, angular-sulcate (usually 4-angled), smooth, 3—10(—30)cm by )%- YA(-4) mm, often much elongated in deep or running water. Sheaths thinly membranous, loose, oblique at the mouth, often reddish at the base, the apex scarious and often somewhat inflated. Spikelet ovate to lanceolate, more or less flattened, acute, (3—)5—8(—15)-flowered, 2-4(-7) by 1-14 mm. Glumes thinly membranous, suberect, oblong-ovate, obtusish, l-nerved, pale green with scarious margins and often reddish brown sides, 14-2 by 1-114 mm, the lower ones subdistichous. Brist/es up to 4, very slender, equalling or exceeding the nut, often reduced or absent (see note). Stamens 2—3; anthers linear, 34-1 mm; connective shortly produced. Style 3-fid. Nut very obtusely trigonous, almost terete, obovate- oblong, on each face with 2—4 distinct longitudinal ribs connected by numerous transverse cross-ridges, whitish to light brown, 4-1 by 4-14 mm; epidermal cells transversely linear. Sty/e-base minute, depressed pyramidal, )4— Yas wide as the nut. Distr. Widely distributed in North America, throughout Europe and northern Asia, Japan, Korea, extending southward to the Ryu Kyu Is., Yunnan, Annam, and Formosa, in Malesia: N. Sumatra (Toba Lake), Philippines (N. Luzon; Ben- guet Subprov., Baguio). A distinct northern element in the Malesian flora. Ecol. In open damp places; around Toba Lake rather common on very wet sandy soil at c. 900 m; in Luzon on seepage slopes about rice paddies at c. 1450 m. Notes. The Malesian plants (and with a few exceptions all East Asiatic ones) belong to: CYPERACEAE (Kern) 335 var. longiseta SveNsS. Rhodora 31 (1929) 189; Ouw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 36. — Scirpus yokoscensis FRANCH. & Say. En. Pl. Jap. 2 (1879) 109, 543. — E. svensonii ZINSERL. Fl. U.R.S:S. 3 (1935) 71, 580. — E. vokoscensis TANG & WANG, Fl. Reip. Pop. Sinic. 11 (1961) 54, t. 20 f. 10-13. — Bristles 3—4, conspicuously longer than the nut. The chromosome number (2n = 20) differs from that of typical E. acicularis (2n = 30-58; 50-70). In the majority of the European plants the bristles are lacking or greatly reduced, very rarely equalling the nut, in the North American specimens they are either very slender and equalling the nut, or absent. The South American E. costulata NEES & MEY. ex KUNTH (En. 2, 1837, 142), mainly differing by the pale green glumes and the 14 mm long, distinctly apiculate anthers, is very near to E. acicularis, and is probably only racially distinct. Also E. pusilla R. BR. Prod (1810) 225; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 106. t. 8 f. 14-19; Svens. Rhodora 41 (1939) 19, t. 539 f. 2, widely spread in extra-tropical eastern Australia and New Zealand, is obviously very near to E. acicu/aris. For the trifling differences see S. T. BLAKE, I.c. 107. According to SVENsON, !.c. 95, in nota, E. acicularis has been introduced into Australia, and between it and E. pusilla all inter- mediate stages are found. 6. Series Palustriformes SvENS. Rhodora 31 (1929) 128. — Type species: E. palustris (L.) R. & S. 6a. Subseries Palustres (CLARKE) SvVENS. Rhodora 31 (1929) 128. — Subg. Eleogenus c. Palustres C. B. CLARKE, Kew Bull. add. ser. 8 (1908) 105.— Type species: E. palustris (L.) Ro &s. 16. Eleocharis brevicollis KERN, Blumea 13 (1965) I ges oud Perennial, with woody, creeping rhizome 2—3 mm thick. Stems erect, in tufts, slender, subterete, longitudinally ribbed, smooth, pale green, (30—)60—90 cm by 1-2 mm. Sheaths herbaceous, shining, dark purplish at the base, the uppermost scarious at the very oblique, not mucronate mouth. Spikelet erect, cylindric, obtuse, densely many-flowered, pale to dark brown, 12-18 by c. 3mm. Glumes thinly membranous, appressed, concave (not keeled), ob- long-ovate, acutish, |-nerved, with narrow, green, median streak, otherwise brown, purplish zonate within the broad, whitish, scarious margins, c. 414 mm long, the lowest firmer, obtuse, embracing the spikelet, sterile. Bristles (3—)4, firm, retrorsely scab- rous, reaching as high as the style-base or slightly overtopping it, pale. Stamens (2—)3; anthers linear, c.2 mm long, with ovate appendage of the connective. Style 2-fid, occasionally 3-fid. Nut biconvex, turgid, elliptic, constricted at the top into a short, but distinct neck, not costate on the angles, shining, yellow or pale brown, 1!4—-174 by 1/4 mm, obsoletely reticulate by the subisodiametric epidermal cells. Style-base triangular, flattened, cellular-spongious, c. % as wide as the nut. 536 FLORA MALESIANA [ser. I, vol. 7° Distr. Malesia: Territory of New Guinea, West- ern Highlands (Sirunki and Lake Inim), Southern Highlands (Tari Gap). Ecol. In swampy, mixed Acorus and sedge sward, rooting in fine detritus clay, at Lake Inim forming ring round edge of marsh, c. 2500 m. Note. The only Malesian member of subser. Palustres, a very difficult group of chiefly Holarctic distribution. 7. Series Maculosae Svens. Rhodora 31 (1929) 128.—Type species: E. maculosa (VAHL) R. & S. 7a. Subseries Rigidae Svens. /.c. — Type species: E. atropurpurea (RETZ.) PRESL. 17. Eleocharis geniculata {L.) R. & S. Syst. Veg. 2 (1817) 150: FurtTaApDo, Gard. Bull. S. S. 9 (1937) 293, 298: Svens. Rhodora 41 (1939) 50;S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 124, t. 10, f, 29-31; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 93; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461. — Scirpus geniculatus LINNE, Sp. Pl. 1 (1753) 48.— Scirpus caribaeus Rotts. Descr. Pl. Rar. Progr. (17712) 242 Deserv& Te: (17713); 46, t2 155.1 35. capitata R. BR. Prod. (1810) 225; Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 361; Herb. Timor. Descr. (1835) 33; KUNTH, En. 2 (1837) 150; Mia. FI. Ind. Bat. 3 (1856) 299: Boeck. Linnaea 36 (1870) 461; BENTH. FI. Austr. 7 (1878) 296; Naves, Nov. App. (1882) 307; CLarKE, Fl. Br. Ind. 6 (1893) 627; Philip. J. Sc. 2 (1907) Bot. 90; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 77; Koorp. Exk. Fl. Java | (1911) 197; Atlas (1922) f. 248; Merr. Fl. Manila (1912) 114: Camus, Fl. Gén. I.-C. 7 (1912) 85; MerRR. En. Borneo (1921) 59; Rip. Fl. Mal. Pen. 5 (1925) 151, non Scirpus capitatus L.— E. atropurpurea Pres_, Rel. Haenk. | (1828) 196, guoad specim. cit., excl. syn. — Scirpus retroflexus (non Potr.) LLANOS, Fragm. Pl. Filip. (1851) 19; F.-Vitt. & NAves in Blanco, FI. Filip. ed. 3, 4' (1880) 13. — E. caribaea S. F. BLake, Rhodora 20 (1918) 24: Merr. Sp. Blanc. (1918) 83; En. Philip. 1 (1923) 119; Back. Onkr. Suiker. (1928) 155, t. 159; SveNs. Rhodora 31 (1929) 225, f. 48: BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 13. Annual. Stems tufted, slender but rather rigid, erect or oblique, angular-striate, smooth, S—40 cm by 4-1 mm. Sheaths appressed, purplish at the base, the uppermost with an oblique and more or less attenuate mouth. Spikelets globose to oblong-ovoid, very obtuse, terete, densely many-flowered, 4-8 by 3-34 mm. Glumes membranous, appressed or obliquely ascending, broadly elliptic, broadly ovate or suborbicular, very obtuse, scarcely keeled, l-nerved, ferrugineous with greenish keel and sides more or less tinged with purple, 174-2 by 14-1)4 mm. Bristles 6-8, rather coarse, ferrugineous to brown, somewhat longer than the nut. Stamens 2—3; anthers linear-oblong, minutely apiculate, c. 14 mm. Style 2-fid. Nut biconvex, obovate, smooth, shining black, YAH. by K-RKmm; epidermal cells very in- conspicuous, subquadrate. Stvle-base conical, usu- ally much depressed, about 14 as wide as the nut. Distr. The most widely distributed Eleocharis, throughout the warmer parts of both the Old and the New World, in Malesia rather rare, but locally sometimes abundant: N. and Central Sumatra, Malay Peninsula (Perak, Trengganu, Pahang, Jo- hore, Singapore), W.-E. Java, Bawean, Madura, Lesser Sunda Is. (Timor), N. Borneo (not seen), Philippines (Luzon, Bohol, Cebu), Celebes. Ecol. In open wet lands (swamps, swampy grasslands, wet rice-fields, sometimes on brackish clay), from sea-level up to 900 m. Vern. Djangga tembe, S, sriwit, sudjinan, djémbut, 18. Eleocharis atropurpurea (RETZ.) PReEsL, Rel. Haenk. | (1828) 196, excl. specim. cit.; KUNTH, En. 2 (1837) 151; Gay, Flora 25 (1842) 641; Borck. Linnaea 36 (1870) 458, excl. varr. B & 7; BENTH. FI. Austr. 7 (1878) 296, p.p.; CLARKE, Fl. Br. Ind. 6 (1893) 627; Philip. J. Sc. 2 (1907) Bot. 90; Ill. Cyp. (1909) t. 36, f. 6-9; Merr. En. Philip. 1 (1923) 119; Back. Onkr. Suiker. (1928) 154, t. 158; SvENs. Rhodora 31 (1929) 227, t. 191, f. 49; S. T. BLAKE, Proc. R. Soc. Queensl. 50 (1939) 125, t. 10, f. 32-33; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 13; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 94; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 461. — Scirpus atropurpureus Retz. Obs. 5 (1789) 14. — E. ochreata (non NEES, nec STEUD.) CLARKE, Philip. J. Sc. 2 (1907) Bot. 90, non al.; MERR. En. Philip. | (1923) 120. Dwarf annual. Stems tufted, erect or curved, capil- lary, angular-sulcate, smooth, 4-15 cm by 4-4 mm. Sheaths appressed, purplish or brownish at the base, the apex of the uppermost oblique, often attenuate. Spikelet oblong-ovoid, obtusish, somewhat angular, densely many-flowered, 2—4 (finally lengthening to 8) by 1/4-2 mm. G/umes membranous, at length more or less spreading, elliptic, obtuse, keeled, 1-nerved, with green keel and pale or more or less purplish sides, %{,-l% by 4-, mm. Bristles 3-4-6), slender, minutely retrorsely scabrous to almost smooth, shorter than to about as long as the nut, whitish, translucent, not rarely vestigial or absent. Stamens 1—2; anthers linear or oblong-linear, mi- nutely apiculate, 4-4 mm long. Style 2-fid. Nut biconvex, obovate, smooth, shining, black, 4-74 by 4-24 mm; epidermal cells very inconspicuous. Style-base minute, flattened, strongly depressed, disciform or depressed-conical, 14-14 as wide as the nut. Distr. Scattered through the tropics of both the Old and New World, also in the U.S.A., and in Europe in Italy and Switzerland; in S. and E. Asia to China, Formosa, and Japan, in Australia in W. and N. Australia, and Queensland, in Malesia very rare: Central Sumatra, W. Java, Madura, Philippines (Luzon), New Guinea (near Port Moresby). The record ‘Central Java’ in Backer, 1949 /.c., is based on a young specimen of E. geniculata. Ecol. In wet rice-fields, wet grassy places, 90— 250 m. Note. Usually readily distinguishable from the more common E. geniculata by its smaller size. From small specimens of the latter it may be distinguished 1974] by the smaller nuts, the much more delicate bristles which are glistening white and translucent, and by the more prominently keeled glumes giving the spikelet an angular appearance. 8. Series Pauciflorae Svens. Rhodora 31 (1929) 127. — Type species: E. pauciflora (LIGHTF.) LINK [correct name: E. quin- queflora (HARTM.) SCHWARZ |. 19. Eleocharis parvula (R. & S.) LINK ex BLUFF, Nees & SCHAUER in Bluff & Fingerh. Comp. Fl. Germ. ed. 2, 1' (1836) 93; Svens. Rhodora 31 (1929) 168, t. 189, f. 18; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 34; KERN, Reinw. 4 (1956) 94; Blumea 10 (1960) 643; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 87; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 460. — Scirpus pusillus VAHL, En. 2 (1806) 246, non E. pusilla R. Br. (1810). — Scirpus nanus (non Por.) SPRENG. Pugill. | (1813) 4. — Scirpus parvulus R.&S. Syst. Veg. 2 (1817) 124; Boeck. Linnaea 36 (1869-70) 477. Dwarf perennial, densely tufted, forming mats, and propagating by small, fusiform, brown or purplish, 3—6 mm long tubers at the end of filiform stolons. Stems erect, weak, capillary, terete, striate when dry, smooth, pale green, 1-7 cm by 4-14 mm. Sheaths membranous. Spikelet ovoid, obtuse, compressed, few-(2—9)-flowered, 2-4 by 1)4-2 mm. Glumes sub- distichous, membranous, obliquely spreading in fruit, ovate, obtuse, scarcely keeled, with distinct green midnerve and pale green or stramineous, sometimes dull brown sides, 114-134 by c. | mm. Bristles 4—6, delicate, retrorsely scabrous, yellowish, unequal, somewhat shorter to slightly longer than the nut, sometimes reduced. Stamens 3; anthers linear, c. 1 mm long, the connective with a distinct reddish appendage. Style 3-fid. Nut equilaterally trigonous, with prominent angles, obovate, smooth, shining stramineous, c. | by 34 mm; epidermal cells inconspicuous, oblong. Style-base minute, pyra- midal, confluent with the apex of the nut and seem- ingly a continuation of it, but of different texture, greenish. Distr. Along the Atlantic coast of North America from New Foundland to the West Indies and the Pacific coast from N. California to British Columbia, rarely inland; in South America in Brazil; Mediter- Cy PERACEAE (Kern) 537 ranean coast of Europe and N. Africa, Atlantic coast of Europe north to Norway, rarely inland; Japan (Kiushiu); Cochinchina, once collected; in Malesia: E. Java (Bangil, S of Surabaya). Ecol. Salt marshes and brackish mud along the seacoast, sometimes inland in salt lakes, at Bangil near the salt iodine-wells not far from the coast. Note. Often treated as a species of Scirpus. As the style-base, though small and confluent with the nut, clearly differs in texture from the latter, it stands more naturally in Eleocharis (see p. 523). According to SVENSON, it may perhaps be considered the most primitive member of ser. Pauciflorae. Doubtful Eleocharis alta BoOECK. Cyp. Nov. 1 (1888) 17. — “Java, Vulcan Gede, alt. 2400m”. No collector mentioned. The description is insufficient. The plant is stated to be very close to E. variegata KUNTH. It was placed in the synonymy of E. tetraquetra NEES by CLARKE, Fl. Br. Ind. 6 (1893) 630. Eleocharis obtusa (WILLD.) SCHULT. Mant. 2 (1824) 89; SveNs. Rhodora 31 (1929) 214; ibid. 55 (1953) 1.— Scirpus obtusus WILLD. En Hort. Berol. | (1809) 76. — The specimens in the Kew Herbarium, cited as “E. ovata R. BR.” by CLARKE in J. Bot. 25 (1887) 269 from ‘Java, ZOLLINGER’ and in J. Linn. Soc. Bot. 34 (1898) 50 from ‘Java’ (see also SVENS. Rhodora 31, 1929, 213) are in fact E. obtusa. Origi- nally they belonged to the HOOKER herbarium; they are provided only with the small printed part of a ZOLLINGER-label “Plantae javanicae cl. ZOLLING- ERO lectae’. This supposed ZOLLINGER collection is not represented in any of the other Eleocharis collections I could study, therefore it is very unlikely that the plants were actually collected in Java. E. obtusa is known from the eastern United States, the Pacific region from northern California to British Columbia, and the Hawaiian Islands. By its annual, cespitose habit, the bifid style, and the biconvex nut E. obtusa is similar to E. geniculata and £. atropurpurea. It belongs in ser. Ovatae characterized by the flat lamelliform style-base, which is here nearly or quite as broad as the nut. In the closely related, and possibly not specifically distinct E. soloniensis (DuBoIs) HARA = E. ovata (ROTH) R. & S. the style-base is much narrower than the nut. 12. BULBOSTYLIS KUNTH, En. 2 (1837) 205, nom. cons., nec STEV. 1813, nec DC. 1836. — Steno- phyllus RAF. Neog. (1825) 4. — Oncostylis NEES in Mart. Fl. Bras. 2, 1 (1842) 80. — Scirpus sect. Oncostylis BoECK. Linnaea 36 (1870) 736. — Fimbristylis sect. Oncostylis BENTH. & HooK. Gen. PI. 3 (1883) 1049. — Fig. 37. Perennial or (in Mal.) annual herbs. Stems tufted, erect, very slender, angular, striate or sulcate, leafy only at the base. Leaves very narrow, nearly always capillary; sheaths generally bearded in the throat with long white hairs sometimes disappearing with age. Inflorescence terminal, subtended by foliaceous involucral bracts, capitate or anthelate, sometimes reduced to a 538 FLORA MALESIANA [ser. I, vol. 7° single spikelet. Spikelets usually not compressed, angular, several- to many- flowered. Rachilla persistent, narrowly winged. Glumes spiral, acropetally caducous, with strong midnerve; lower 1—2 empty. Flowers achlamydeous (hypogynous bristles or scales absent), bisexual, the uppermost often male or barren. Stamens 1—3; anthers oblong or linear, with shortly produced connective. Style articulate with the ovary, slender, glabrous; stigmas 3, very rarely 2 (not in Mal.); style-base incrassate, bulbiform, persistent on the apex of the nut as a minute, darker coloured button. Nut trigonous or triquetrous, very rarely biconvex (not in Mal.), obovate, obtuse, scarcely stipitate. Distr. Probably c. 100 spp. in the tropical and subtropical regions of the world; centres of development are tropical Africa and America; in Malesia only 3 spp., none of them common. Ecol. B. barbata and B. puberula prefer rather dry sandy localities at low altitudes, B. densa occurs in wet or swampy localities at medium and high altitudes. Notes. The Malesian spp. are easily recognized by the capillary leaves with needle-like white hairs at the orifice of the sheaths, as well as by the peculiar button crowning the nut (like in Eleocharis, which is to Scirpus as Bulbostylis is to the closely related genus Fimbristylis). SVENSON (N. Am. FI. 18, 1957, 540) attaches much value to the shape of the epidermal cells of the nut as a generic character (longitudinally elongated in Bulbostylis, isodiametric to horizontally elongated in Fimbristylis). This would, however, place the closely related B. barbata and B. puberula in different genera and Fimbristylis hispidula (p. 560, lacking the button on the nut!) in Bulbostylis, with which I do not agree. It is somewhat doubtful whether the name Bulbostylis was validly published by KUNTH, who proceeded to describe the members of the genus as species of /solepis, though he proposed Bulbostylis as a separate genus intermediate between Isolepis and Fimbristylis. KEY TO THE CSRECIES 1. Glumes muticous, acute. Inflorescence anthelate, open, rarely reduced to a single spikelet. Nut minutely granular, obsoletely transversely wrinkled .. . ee esl is” cuore Sp pat ee 1. Glumes mucronulate, the mucro finally excurved. Inflorescence capitate or, if anthelate, congested. Nut either smooth or distinctly transversely wrinkled. 2. Nut smooth, minutely reticulate by the isodiametric epidermal cells, 4-74 mm long. Glumes glabrous, or but slightly pubescent... 2. B. barbata 2. Nut distinctly transversely wrinkled, with longitudinally oblong-linear epidermal cells, 4-1 mm long. Glumes rather densely pubescent . 1. Bulbostylis densa (WALL. in RoxB.) HAND.-MAZz. in Karsten & Schenk, Vegetationsb. 20, 7 (1930) 16; Hara, J. Jap. Bot. 18 (1942) 467; OHwi1, Mem. Coll. Sc. Kyoto B 18 (1944) 50; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 468. — Scirpus densus WALL. in Roxs. FI. Ind. 1 (1820) 231. — Isolepis trifida NEES in Wight, Contr. Bot. Ind. (1834) 108. — Isolepis capillaris, formae indicae KUNTH, En. 2 (1837) 212; Mio. FI. Ind. Bat. 3 (1856) 312. — Isolepis trichocolea Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.] Syn. 2 (1855) 96 (‘trichokolea’); Mia. Fl. Ind. Bat. 3 (1856) 308.— Fimbristylis capillacea HOCHST. ex Steup. [in Zoll. Syst. Verz. 1 (1854) 61, nom. nud.] Syn. 2 (1855) 111; Mig. Fl. Ind. Bat. 3 (1856) 320. — Scirpus capillaris (non L.) Boeck. Linnaea 36 (1870) 759, p.p. — B. capillaris var. trifida CLARKE, FI. Br. Ind. 6 (1893) 652; Philip. J. Sc. 2 (1907) Bot. 98. — Stenophyllus capillaris var. trifidus DoMIN, Bibl. Bot. 20, Heft, 85 (1915) 464. — B. capillaris (non CLARKE) Camus, FI. Gén. I.-C. 7 (1912) 127. — Stenophyllus capillaris B. trifidus Preirr. Bot. Arch. 6 (1924) 188. — B. trichokolea BEETLE, Leafl. West. Bot. 4 (1944) 45. — Stenophyllus capillaris (non Britt.) BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 26. — B. trifida Nemes, Kew Bull. (1950) 209. — Fimbristylis densa KOYAMA & CHUANG, Quat. J. Taiwan Mus. 13 (1960) 229. Stems setaceous, glabrous and smooth, (1—)5—40 3. B. puberula cm by 14-14 mm. Leaves much shorter than the stems, capillary, glabrous (except for the long white hairs at the orifice of the sheaths), slightly scaberulous at the top, 4-)K mm wide. Inflorescence very variable, simple or subcompound, with 1—many spikelets, in Mal. usually very loose, up to 5 cm long. Involucral bracts usually very short, glume-like, or the lower 1—2 setaceous with dilated scarious base, sometimes up to 7 cm. Rays 0-7, filiform, glabrous, smooth, up to 4cm. Spikelets solitary, oblong-ovate to oblong, acute, rather densely flowered, 3-6 by 134-2 mm. Glumes membranous, ovate to broadly ovate, acute, muticous, strongly keeled, with nerveless sides and ciliolate margins, brownish to castaneous, 134-2 mm long. Stamens 2; anthers oblong, 4—/4 mm. Style 34-1 mm; stigmas 3, shorter than the style. Nut triquetrous, obovate to broadly obovate, densely granular-puncticulate, stramineous to fumose, 0.7— 0.9 by 0.5-0.8 mm; epidermal cells minute, iso- diametric to oblong. Distr. Widely distributed in the Old World tropics and subtropics: S. and E. Asia to China and Japan, tropical Africa and Australia, in Malesia: N. Sumatra (Atjeh, E. Coast Res.), Central and E. Java, Lesser Sunda Islands (Bali, Lombok, Wetar, Alor), Philippines (Luzon: Cagayan, Pampanga, Lepanto, Bontoc, Benguet), SW. Celebes, and New Guinea. Ecol. In open wet places, locally often abundant, 1974] CYPERACEAE (Kern) a9 at medium and high altitudes, usually 1000—3000 m; ELBERT 4551 from Wetar was collected between 425 and 500 m. Vern. Nomai, poudan, New Guinea (Enga lang.). Notes. B. densa is very near to the American B. capillaris (L.) CLARKE, differing by the more acute, glabrous or only slightly pubescent glumes, and the granular, not or hardly transversely rugulose nuts. Remarkable specimens of this species were col- lected on Mt Pulog in Luzon (FB 16143: MERRILL Philip. Pl. 541): the inflorescence is reduced to 1—3 spikelets, the latter are larger than usual, c. 3 mm wide; the glumes c. 3 mm long; the nuts broadly obovate to turbinate, 1.1 by 0.95 mm: the anthers somewhat longer than usual, c. 4 mm. The cited collections may represent a local race of this ex- tremely variable species. Isolepis trichocolea STEUD. was based on ZOL- LINGER 613. The sheet in STEUDEL’s herbarium (P) contains two specimens, one of which is B. densa, the other B. barbata. Hence STEUDEL’s description partly refers to the former species, partly to the latter. In accordance with CLARKE (FI. Br. Ind. 6, 1893, 652), the specimen of B. densa is selected as the type. 2. Bulbostylis barbata (RoTTB.) CLARKE, FI. Br. Ind. 6 (1893) 651; Philip. J. Sc. 2 (1907) Bot. 98; RIDL. Mat. Fl. Mal. Pen. 3 (1907) 77; Camus, Fl. Gén, I.-C. 7 (1912) 126; Merr. En. Philip. 1 (1923) 127; RIDL. Fl. Mal. Pen. 5 (1925) 160; KUx. Bot. Jahrb. 70 (1940) 463, incl. f. paupercula KUK.; KERN in Back.& Bakh. f. Fl. Java 3 (1968) 468. — Scirpus barbatus Rotts. Progr. (1772) 27; Descr. & Ic. (1773) 52, t. 17 f. 4; Boeck. Linnaea 36 (1870) 751, excl. var. B; VIDAL, Phan. Cuming. Philip. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 284. — Isolepis barbata R. BR. Prod. (1810) 222; Prest, Rel. Haenk. | (1828) 187; KUNTH, En. 2 (1837) 208; DECNE, Herb. Tim. Descr. (1835) 32; Mia. Fl. Ind. Bat. 3 (1856) 310. — Onco- stylis barbata Nees in Hook. J. Bot. Kew Misc. 6 (1854) 29. — Isolepis involucellata StEup. [in Zoll. Syst. Verz. | (1854) 62, nom. nud.] Syn. 2 (1855) 101; Mia. FI. Ind. Bat. 3 (1856) 311. — Isolepis cumingii STEUD. Syn. 2 (1855) 101; Mig. Fl. Ind. Bat. 3 (1856) 310. — Isolepis armerioides Mig. Fl. Ind. Bat. 3 (1856) 310, incl. var. B; Kurz, Nat. Tijd. N. I. 27 (1864) 223. — Fimbristylis barbata BENTH. F1. Austr. 7 (1878) 321; F.-VILL. Nov. App. (1882) 308; MERR. Fl. Manila (1912) 116; Back. Onkr. Suiker. (1928) 157, t. 161. — Fimbristylis cumingii F.-VILL. Nov. App. (1882) 308. — Iriha barbata O.K. Rev. Gen. PI. 2 (1891) 753. — Stenophyllus barbatus Cooxe, FI. Bombay 2 (1908) 887; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 27. — Fimbristylis actino- schoenus (non CLARKE) Camus, FI. Gén. I.-C. 7 (1912) 124, excl. descr. et fig. — Fimbristylis armerioides BEETLE, Leafl. West. Bot. 4 (1944) 45. — Scirpus mindorensis BEETLE, l.c. 46. Stems setaceous, glabrous and smooth, 5—30 cm by 4—/4 mm. Leaves much shorter than the stems, capillary, glabrous (except for the long white hairs at the orifice of the sheaths), smooth or slightly scaberulous at the top, 4-14 mm wide. Inflorescence capitate, hemispherical, with (1—)3—20 spikelets, 5—15 mm across. Involucral bracts 1-3, somewhat shorter to much longer than the inflorescence, filiform, with dilated scarious base, 5—20 mm. Fig. 37. Bulbostylis puberula(Poir.) CLARKE. a. Habit, x A, b. spikelet, x5, c. deflorate flower, d. nut, both x 15 (a—~d BUNNEMEUER 6002). 540 FLORA MALESIANA [ser. 1, volove Spikelets sessile, ovate to oblong, acute, 3-8 by 1— 1/4mm. Glumes membranous, ovate, mucronulate (mucro finally excurved), strongly keeled, with nerve- less sides and minutely ciliolate margins, otherwise glabrous or but slightly puberulous, rusty brown with green keel, 1)4-2)4 by UA4-UAZ mm. Stamen 1; anther oblong-linear, c. /4mm. Style 74-1 mm; stigmas 3, shorter than the style. Nut triquetrous, broadly obovate, smooth, finely reticulate by the minute isodiametric epidermal cells, whitish to stramineous, 0.5—0.75 by 0.4—0.6 mm. Distr. Widely distributed over the warm parts of the Old World, also in the southern U.S.A. (see SVENSON, N. Am. FI. 18, 1957, 544), throughout Malesia, but nowhere common. Ecol. In open dry sandy places, locally sometimes abundant, often near the sea: dunes, fields, savan- nahs, sometimes on rocks, usually at low altitudes (e.g. in Java up to 250m, in Flores at 1000 m, in Papua up to 450 m); according to MERRILL in the Philippines ascending to 1500 m; a specimen from Luzon (Benguet Subprov., MEARNS BS 4257) was collected at 2100 m. Vern. Mapu, Sumba, papoa, Sula Is., bese baroho, korompau nguas, Tanimbar; Philip.: humot, Iv., kulilis, Tag., tirtiris, Ik. Note. A variable species. A form known from Ceylon and India, with larger glumes, 3 stamens with larger anthers, and slightly larger nuts is sometimes treated as a separate species. 3. Bulbostylis puberula (Porr.) CLARKE, FI. Br. Ind. 6 (1893) 652; RipL. Mat. Fl. Mal. Pen. 3 (1907) 78; Camus, Bl. Gen; L-C.>7 (1912): 128, «f)-17, 1=2'; RIDL. Fl. Mal. Pen. 5 (1925) 160; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 468. — Scirpus puberulus Pore. in Lamk, Enc. 6 (1804) 767; Boeck. Linnaea 36 (1870) 767, excl. syn. Isolepis armerioides. —— Isolepis puberula KUNTH, En. 2 (1837) 213; STeuD. Syn. 2 (1855) 103. — Scirpus setaceus (non L.) Camus, FI. Gen. I.-C. 7 (1912) 132. — Stenophyllus puberulus Kituip, J. Siam Soc. Nat. Hist. Suppl. 7 (1927) 57; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 27.— Fimbristylis puberula Back. ex STEEN. Arch. Hydrobiol. 3, Suppl. Band 11 (1932) 236. — Fig. 37. Stems setaceous, glabrous or pubescent, some- times hispid just below the inflorescence, 5—40 cm by Y%-% mm. Leaves much shorter than the stems, capillary, short-pubescent, /4—/4 mm wide; sheaths stramineous to brownish, pubescent. Inflor escence simple or subcompound, anthelate with up to 5 short rays c. /4 cm long, often congested to almost capitate, sometimes reduced to a single spikelet, up to 14cm long and wide. Involucral bracts 2-4, filiform, scarious at the dilated base, the lowest erect, often as though continuing the stem, usually longer than the inflorescence, up to 4 cm. Spikelets solitary, stalked (except in very small plants), but often Closely packed, ovate to oblong, acute, 4-8 by c. 14 mm. Glumes membranous, broadly ovate, mucronulate (mucro finally excurved), strongly keeled with nerve- less sides, rather densely pubescent, greenish to brown, 134- 24 by 1)4-2 mm. Stamen 1; anther oblong-linear, 4-34 mm. Style 4-1 mm; stigmas 3, shorter than the style. Nut triquetrous, obovate to broadly obovate, transversely wavy-wrinkled, whit- ish to stramineous, 0.75—1 by 0.6—0.75 mm; epi- dermal cells longitudinally oblong-linear. Distr. Tropical Africa and Asia, in Malesia: Sumatra, Riouw Arch., Malay Peninsula, W.-E. Java, Madura, W. & N. Borneo, Anambas & Natuna Is.; everywhere rare. Ecol. In sandy localities at low altitudes, often near the sea, locally sometimes abundant. Uses. According to a label in SING the plant is (was?) cultivated by the Chinese in Singapore and exported to China, where it is used as a diuretic. Vern. Rumput hulu hidang, M (Mal. Pen.). Note. The habit is much like that of B. barbata, but the texture of the pericarp is quite different. The nut character is the most distinctive; sessile or stalked spikelets is also a good distinction, except in very small plants. The hairiness of the stem is variable. 13. FIMBRISTYLIS VAHL, En. 2 (1806) 285, nom. cons.; KUNTH, En. 2 (1837) 220; STEUD. Syn. 2 (1855) 106; Mia. Fl. Ind. Bat. 3 (1856) 314; Bogcx. Linnaea 37 (1871) 2; ibid. 38 (1874) 384; B. & H. Gen. Pl. 3 (1883) 1048, excl. sect. Oncostylis; PAX in E. & P. Pfl. Fam. 2, 2 (1887) 113; CLARKE, Kew Bull. add. ser. 8 (1908) 107; Onwt, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 52; KERN, Blumea 8 (1955) 10; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 99, excl. subg. Bulbostylis. — Cyperus subg. Iria L. C. Ricu. ex PERS. Syn. 1 (1805) 65. — Iria HEDw. Gen. Pl. (1806) 360. — Gussonea PRresL, Rel. Haenk. 1 (1828) 183. — Iriha O.K. Rev. Gen. PI. 2 (1891) 751. — Trachystylis S. T. BLAKE, Proc. R. Soc. Queensl. 48 (1937) 89. — Tylocarya NELMES, Kew Bull. (1949) 139. — Fig. 38—48. For the other generic names referred to Fimbristylis in the present treatment see under the synonymy of the sections. Annuals, or perennial herbs with short, rarely creeping rhizome. Stems usually tufted, erect or obliquely erect, rarely decumbent, solid, more or less angular or subterete, striate or sulcate. Leaves linear, all basal, or also a few in the lower part of the stems, often cellular-reticulate above, those of the 1974] CYPERACEAE (Kern) 541 Fig. 38. Fimbristylis subdura OHW1. a. Habit, x /4, a’. leaf tip, b. spikelet, both x 5, c. glume, d. anther, e. deflorate flower, f. nut, all x 10 (a—f BACKER 6525). 542 FLORA MALESIANA [ser. I, vole flowering stems not rarely reduced to bladeless or short-bladed sheaths; ligule a fringe of short hairs or a membranous projection between sheath and blade, or absent. Inflorescence terminal (very rarely pseudolateral because of the erect lowest bract seemingly continuing the stem), often anthelate (terminal spikelet or cluster overtopped by the lateral ones), simple or more or less com- pound, not rarely subcapitate or capitate by suppression of the rays, or con- sisting of a single spikelet. Rays subtended by foliaceous but often much reduced bracts, the base enclosed in a tubular prophyll (cladoprophyll). Spikelets solitary or in clusters, terete, angular (because of the acutely keeled glumes), or strongly laterally compressed, few- to many-flowered; axis (rachilla) as a rule persistent after the glumes and fruits have fallen off, often winged by the persistent basal part of the glumes, exceptionally spikelets falling off as a whole. Glumes usually spirally arranged, more rarely subdistichous or distichous, acropetally deciduous, only in a few spp. persistent on the rachilla, some (up to 6) lower ones empty. Flowers bisexual (the uppermost of the spikelet often tabescent), naked (hypo- gynous bristles or scales absent). Stamens 1—3; anthers usually with shortly produced connective, rarely the latter produced in a conspicuous, smooth or bristly appendage. Style articulated with the ovary, deciduous with its dilated base, leaving no button on the nut (in a few spp. style-base subpersistent), ciliate or glabrous, triquetrous or flat; stigmas 2 or 3 (number sometimes varying in the same specimen). Nut trigonous or lenticular, from orbicular or broadly obovate to oblong-linear, smooth, verrucose, or tuberculate, reticulate by the hexagonal or roundish epidermal cells, or lineolate when those cells are linear, or trabeculate (with longitudinal ribs connected by cross-bars), or cancel- late (latticed). Distr. Large genus with some hundreds of species, chiefly in the tropics of both hemispheres, but with some species extending to the warmer parts of the temperate regions. In Malesia 78 spp.; unlike in Cyperus only a small number of them (8) is pantropical. The number of endemic spp. is relatively high (17), but their distribution is usually inadequately known. The vast majority of the Malesian spp. is widely distributed in S. and SE. Asia, several extending to tropical Australia. Australian elements extending to E. Malesia are F. schultzii, F. recta, F. furva, F. lanceolata, F. signata, and F. acicularis. Very disjunct areas are shown by F. dictyocolea (Thailand and Indo-China — New Guinea), F. adenolepis (Thailand and Indo-China — Kangean), F. intonsa (Bengal — N. Sumatra — New Guinea), and F. semarangensis (Indo-China — Java). Future collections may fill some of these gaps. Ecol. Most spp. prefer wet localities: swamps, wet rice-fields, margins of lakes, river-banks. Relatively few occur in forests, savannahs, or savannah forests, or are characteristic of the sandy or muddy sea-shore. Three spp. (F. trichophylla, F. calcicola, and F. malayana) are restricted to open, dry places on the limestone hills of the Malay Peninsula. F. dichotoma and F. littoralis are sometimes troublesome weeds. Nearly all the Malesian spp. are lowland plants ascending to medium altitudes. Only F. aestivalis, F. aphylla, F. consanguinea, F . fusca, F . pierotii,and F.. salbundia occur above 1500 m, F. consanguinea ascending to 2300 m. Uses. The genus has but little economic value. Some species are used for matting; F. globulosa is sometimes cultivated for that purpose. The foliage of F. dichotoma and F. littoralis furnishes a rather good cattle-fodder. Notes. VAHL, the founder of the genus, segregated from Scirpus only those species with spirally arranged glumes, and flat, ciliate, distigmatic, deciduous style with enlarged base. For the species with the same flower- structure but distichous glumes he created the genus Abildgaardia; the tristigmatic species were left in Scirpus. Already ROBERT BROWN recognised the deciduous style articulated with the nut as the essential character of the genus, and added several tristigmatic Australian species. This extension of the genus was carried through by KuntH for the extra-Australian species. NEES tried to revive LesTIBOUDOIS’ genus Trichelostylis comprising the tristigmatic species, but he was never followed. Asa GRAY, BENTHAM, and recently KoyAMA, merged Bulbostylis into Fimbristylis. To me Bulbostylis is morphologically as well circumscribed as many other Cyperaceous genera. VAN DER VEKEN, Bull. Jard. Bot. Brux. 35 (1965) 323-328 and 333-336, who investigated the embryos of 45 Fimbristylis spp. and 19 Bulbostylis spp., found that, with a few exceptions in Fimbristylis, the embryos of the two genera are of a different type, another reason for keeping Bulbostylis apart. In a few Fimbristylis spp. the embryo is a variant of the Bulbostylis type, and of those species the systematic place is indeed disputable also on other grounds (see F. hispidula). The embryo of F. thouarsii differs essentially from both the Fimbristylis and Bulbostylis types. As also the eucyperoid anatomical structure and the articulated rachilla are not found elsewhere in the genus, it may be 1974] Cy PERACEAE (Kern) — 543 better to reinstate the genus Actinoschoenus, probably also comprising F. yunnanensis, of which anatomical structure and embryo-type are as yet unknown. The species with a single terminal spikelet, and especially F. tetragona with greatly reduced leaves, are often taken for Eleocharis spp. The genus Eleocharis differs from Fimbristylis by the presence of hypogynous bristles and by the persistent style-base forming a button on the nut. The spikelets of Fimbristylis spp. with distichous glumes strongly resemble those of Cyperus. In the latter genus the style is not articulated with the ovary. Probably due to the attack ofa parasite the spikelets of F. dura, F. globulosa, and F .dichotoma are sometimes comose by the strongly elongated, sterile upper glumes (see MiQueEL, Fl. Ind. Bat. 3, p. 318 under F. efoliata). F. germainii CAMUS was based on such abnormal specimens of F. dichotoma. In outline the subdivision of the genus accepted below agrees with that of OHwi in J. Jap. Bot. 14 (1938) 571. A natural classification has not been attained. In some of the tristigmatic species the number of stigmas is often reduced to two (F. cymosa, F. globulosa, F. sericea, and to a lesser extent F. pauciflora and F. schultzii). Distigmatic F. scaberrima has been placed next to tristigmatic F. dura, to which it shows affinity in almost every respect. The arrangement of the glumes in sect. Abildgaardia and sect. Fuscae is sometimes not clearly distichous. KEY TO THE SECTIONS represented in Malesia 1. Rachilla of the spikelets persistent. Glumes acropetally deciduous. 2. Nut obovate, oblong-obovate, orbicular, pyriform, or turbinate. 3. Stigmas 3; style triquetrous, usually glabrous. Nut trigonous or triquetrous (see F. globulosa, F. scaberrima, and F. cymosa!). 4. Glumes spirally arranged. 5. Nut smooth or verruculose. 6. Inflorescence anthelate or capitate. 7. Ligule a fringe of short hairs. Spp.1-6. . . : . . . . . =. . UJ. Sect. Trichelostylis 7. Ligule.absent. 8. Cauline leaves reduced to bladeless sheaths. Spp. 7-13. . . . . . . 2. Sect. Miliaceae 8. Cauline leaves with well developed blades. 9. Glumes not or only microscopically ciliolate. 1OwRather stout perennials. Spp: 14-20) 8S es i 232 Sect.\Cymosae 10. Small annuals. Spp. 21-22. . . Rube 5.4 Wits eth 2 Chose sel pe 4 WS cGhaencrae 9. Glumes long-ciliate. Spp. 23-28 . ee ee ee SS. Sect, Leptocladac 6. Inflorescence consisting of a single spikelet. Spp. 29-30 . . 6. Sect. Heleocharoides 5. Nut coarsely tuberculate, pyriform. Cauline leaves reduced to bladeless sheaths. Inflorescence anthelate.Sp.31 . . sibs Abra. Mbt Roe be Secrsignatae 4. Glumes distichously arranged (see F. eragr ostis!). 11. Nut large, 2-3 mm long, usually coarsely tuberculate. Inflorescence usually consisting of a single Spikelet®Sp.32... . 8. Sect. Abildgaardia 11. Nut much smaller, smooth or verruculose. Inflorescence with a few to numerous spikelets. Spp. 33—44. 9. Sect. Fuscae 3. Stigmas 2; style flat, usually ciliate. Nut biconvex or planoconvex. Glumes spirally arranged. 12. Nut not coarsely rugulose. 13. Glumes ovate or orbicular, less than twice as long as broad. Inflorescence usually anthelate. 14. Ligule a fringe of short hairs, very rarely membranous. iS Nat smooth. Spp.45=50: ©... 4 ee oe EE BE I Seek: Dichelosivlis i> Nut trabeculate. Spp. 51-582. os eee te OS OL SE Sect aBinmbstylis 14. Ligule absent. 16. Spikelets terete or subterete, 2-4 mm wide. Spp.59-62 . . . . . . 12. Sect. Rigidulae 16. Spikelets angular, 1-1/4 mm wide. Spp.63-67. . . . . 13. Sect. Pogonostylis 13. Glumes oblong-ovate or oblong-lanceolate, twice or more than twice as long as broad. Inflorescence usually consisting of a single spikelet. Spp.68-71 . . . 14. Sect. Neodichelostylis 12. Nut coarsely rugulose by transverse ridges. Cauline Jeaves reduced to bladeless sheaths. Inflorescence consisting of a single spikelet. Spp.72-74 . . . . . . . . . . . 15. Sect. Nutantes 2. Nut subcylindric, with almost parallel edges. 17. Inflorescence consisting of a single spikelet. Cauline leaves reduced to bladeless sheaths. Nut without clavate appendages. Sp.75.. . . 16. Sect. Mischospora 17. Inflorescence anthelate. Cauline leaves with well developed blades. Nut ornamented with rows of clavate appendages. Sp. 76. . . 17. Sect. Dipsaceae 1. Rachilla deciduous, spikelets falling off entire. Glumes distichous. Cauline leaves reduced to bladeless sheaths. Inflorescence Capitate. Spu7 1) «sins Be ewe cues 6 Sali ee te ee lO. SeCie ACENOSCHOCHNS KEY TO THE SPECIES Ripe fruits are indispensable 1. Glumes (at least in young spikelets) exactly distichous, the distichous arrangement in mature spikelets sometimes less pronounced by torsion of the rachilla. Spikelets strongly laterally compressed, much resembling those in Cyperus. 544 FLORA MALESIANA [ser. I, vol. 7° 2 Stiemas 2. Nut bicenvex. Seésub 56:2 Ge. 6He Gonke es Row ee eo 0S BS sealieeae 7 Stigmas 3. Nut trigonous or triquetrous. 3. Inflorescence a dense globose head of sessile, stellately spreading spikelets, the latter when mature falling offasawhole. . . . 77. F. thouarsii 3. Inflorescence anthelate, with distinct rays, or consisting of a ‘single terminal spikelet, rarely almost capitate. Glumes acropetally deciduous from the persistent rachilla. 4. Nut 2-3 by 1)4-2 mm, coarsely tuberculate, rarely almost smooth. Inflorescence usually consisting of a single terminal spikelet, more rarely | or 2 peduncled lateral spikelets added . . 32. F. ovata 4. Nut much smaller, smooth or verruculose. Inflorescence usually with several to numerous spikelets. 5. Glumes densely reddish gland-dotted. Leaves very narrow, setaceous or almost so, )4—1 mm wide. Spikelets few-flowered. 6. Nut smooth, inconspicuously reticulate by the isodiametric epidermal cells. Anthers with a subulate, 4% mm long ‘appendage of the connective . _ 35. F. fuscoides 6. Nut verruculose, transversely lineolate by the linear or - oblong- linear epidermal cells. Connective of the anthers but shortly produced. 7. Perennial with creeping, woody rhizome. Inflorescence compound or decompound, with many to numerous spikelets. Glumes 2?4—4 mm long. Stamens 3. Style shenty hairy at the base, 3-314 mm long. . . 33. F. cinnamometorum 7. Delicate, tufted eet vith marae Eee ieceeonce simple or Salmpst so, with (1—)3—5 spikelets. Glumes 114-2 mm long. Stamen 1. Style glabrous, 1-1%,mmlong. . . . 34. F. adenolepis 5. Glumes not gland-dotted, either glabrous or hairy. 8. Outer leaf-sheaths coriaceous, shining purplish or fuscous. Leaves long-acuminate, with bristle-like, readily caducous tip . . . 36. F. vanoverberghii 8. Outer leaf-sheaths stramineous or light brown. Leaves with rounded, apiculate top. 9. Nut finely longitudinally striate and transversely lineolate by the transversely linear epidermal cells in 3—4 vertical rows on each face. . . . . 44. F. calcicola 9. Marking of the nut otherwise; epidermal cells isodiametric or almost so. 10. Base of the pyriform nut abenpy truncate, conspicuously broader than the short but distinct stipes. c =. disc) heals Sabsin latent. diya sie oeO en) fm DN eyes 10. Nut not truncate at the base. 11. Nut perfectly smooth. Glumes long-acuminate, minutely puberulous at the top, 334-4 mm long. 42. F. malayana 11. Nut verruculose or tuberculate. 12. Glumes hairy. 13. Glumes very broadly ovate, 174-2 mm long. ee 1-14 mm hone: Anthers oblong-linear, 4 mm long . . 41. F. intonsa 13. Glumes lanceolate or ovate- lanceolate, 4- 64 mm long. Style 4-6 mm long. Anthers linear, 174-2 mm long. 14. Leaves much shorter than the stems, often scarcely // as long, (1—)2—4 mm wide. Inflorescence compound to supradecompound, with several to numerous spikelets. Glumes scabrid by very short, stiffish hairs, with broad, whitish hyaline, glabrous margins . . 38. F. fusca 14. Leaves longer, up to 35 cm, 1-2 mm wide. Inflorescence simple, rarely one of the rays with a short secondary ray, with 3—7 spikelets. Glumes pubescent Ks soft hairs, not or scarcely hyaline-margined, ciliolateinthe upper part . . . . . . . 39. F. fulvescens 12. Glumes glabrous. 15. Perennial with short, woody rhizome. Leaves (2—)3—5 mm wide. Spikelets 2-4 mm wide. Glumes 3—5 mm long, distinctly mucronate, chartaceous . i yoles ERY eragrostis 15. Annual with fibrous roots. Leaves 1-2 mm wide. Spikelets yn mm wide. Glumes VA4-2(-244) mm long, minutely apiculate just below the apex, membranous. . . . 43. F. disticha 1. Glumes spirally arranged. Spikelets terete or angular, not strongly compressed. 16. Nut subcylindrical, oblong-linear in outline (with almost parallel edges). 17. Inflorescence consisting of a single terminal spikelet, 6-15 by 4-6 mm. Glumes muticous, 3-5 mm long. Nut trabeculate, seated on a conspicuous, up to 1 mm long gynophore, 1’4—2 mm long. 75. F. tetragona 17. Inflorescence anthelate, with up to 15 spikelets, rarely reduced to a single one. Spikelets 3—6 by 2-3 mm. Glumes mucronate, the blade about 1 mm long. Nut subsessile, ornamented on the edges with a row of clavate appendages, Y%-Y%Ammiong . 9? S25) ea Ose dinsacen 16. Nut obovate, oblong-obovate, orbicular, pyriform, or turbinate. 18. Stigmas 3, rarely i in some of the flowers 2. Nut trigonous or triquetrous, when dorsiventrally compressed with a raised dorsal angle. 19. Glumes with long-ciliate margins. 20. Nut truncate or depressed at the top, transversely wrinkled by the prominent walls of the longi- tudinally elongate, linear epidermal cells .. . 24. F. hispidula 20. Nut neither truncate or depressed, nor transversely wrinkled, ‘the epidermal cells isodiametric or transversely linear. 21. Upper side of leaves asperulous by short, bulbous-based hairs. Nut trabeculate, with transversely linear epidermal cells. Ligule a dense fringe ofhairs. . 56.27 blepharolepis 21. Leaves without bulbous-based hairs. Nut not trabeculate, its epidermal cells not linear. Ligule absent. 1974] Cy PERACEAE (Kern) 545 22. Inflorescence consisting of a single terminal spikelet 5-6 mm wide. Glumes 5—7 mm long. Nut 124-2 mm long. FO ais Woon meets 22. Inflorescence usually with ‘several spikelets. Spikelets much narrower. Glumes and nuts shorter. 23. Glumes densely gland-dotted. 24. Spikelets aggregated into small clusters. Glumes veg small, at most 114 mm long. Anthers with smooth connective. . . 23. F. leptoclada 24. Spikelets not clustered. Glumes larger, 2 ay mm long. Connective of anthers with setulose top. 25. Spikelets linear, 10-25 mm long. Glumes 4-4/4 mm long. Style 3-374 mm. Nut 74, mm long. 27. F. lanceolata 2S Spikelets ovoid or ee -ovoid, 5-10 mm long. Glumes 2—3 mm long. Style 1-2 mm. Nut s7Ammiong. . Ae i ee nee ce eee ee et ee nA CASSALCNSIS 23. Glumes not gland-dotted. 26. Inflorescence anthelate. . > 2 © =26; Fo furva 26. Inflorescence a small, hemispherical or 1 subglobose head. See under 50. . 22. F. schultzii 19. Glumes with glabrous or only microscopically ciliolate margins. 27. Nut truncate and quasi-tridentate at the apex. Inflorescence usually consisting of a single spikelet, pseudolateral because of the erect bract as though continuing the stem . . . 78. F. fenestrata 27. Nut neither truncate nor quasi-tridentate at the apex. Inflorescence terminal. 28. Lowest (empty) glume more than half as long as the spikelet. Inflorescence always consisting of a single spikelet. 29. Sheaths of the cauline leaves disintegrating in front into fine, reticulate fibres. Spikelet 7—9 by 3 mm. Glumes I-nerved. Style 3-5 mmlong. . . 29. F. dictyocolea 29. Leaf-sheaths not disintegrating into reticulate fibres. Spikelets aly by Ly mm. Glumes several- nerved. Style 114-2)4 mm long. . . 30. F. pauciflora 28. Lowest glume much shorter Ban half the spikelet. ‘Inflorescence consisting of several to numerous spikelets, but rarely reduced to a single one. 30. Leaves ligulate (blade and sheath separated from each other by a fringe of short hairs). 31. Stems obtusely trigonous or quadrangular, only slightly compressed. Spikelets 2-2’4 mm wide. Glumes 7—9-nerved. Nut 114-174 mm long. Style abruptly thickened at the base. 1. F. thomsonii 31. Stems strongly compressed, ancipitous. Spikelets 1-2 mm ce Glumes with 3-nerved keel. Nut usually shorter than 1 mm (in F. consanguinea 1-1}£ mm). 32. Spikelets very small, 1 mm wide. Glumes 1 mm long. Style’/4mm . . . 5. F. microcarya 32. Spikelets larger, we 2/4 mm wide. Glumes 2—3 mm long Style 1-2 mm. 33. Stems and leaves very narrow, )4- Zan wide. Margins of the glumes scarcely hyaline. Anthers 4 mmlong . . 4. F. capilliculmis 33. Stems 1-4 mm wide, leaves (te ae 5 mm. Margins of the glumes whitish hyaline. Anthers 1-1/4 mm long. 34. Inflorescence subsimple, with few spikelets. Stems 1-1/4 mm wide. Glumes (2/4-)3 mm long. Nut smooth, 1-1), mmlong . . 3. F. consanguinea 34. Inflorescence compound or decompound, with many spikelets. Stems 2-4 mm wide. Glumes 2-2/4 mm long. Nut verruculose, {,.-% mmlong. . . 2. F. complanata 30. Leaves eligulate (sheaths on the inner side gradually passing into the blades). 35. Spikelets solitary, or only a few of them more or less clustered. 36. Cauline leaves (at least the upper one) reduced to bladeless, tubular sheaths. 37. Glumes 334-4 mm long, the midnerve excurrent in a scabrid, 1-114 mm long awn. Nut coarsely tuberculate, about 144 mmlong . . . 31. F. signata 37. Glumes much shorter, muticous or shortly apiculate. Nut smooth or verruculose, smaller. 38. Basal leaves strongly twisted to the left. Inflorescence simple or almost so, with few (rarely more than 4) spikelets . . . 21. F. obtusata 38. Basal leaves when present not arising i in a left-hand spiral. Inflorescence usually compound to supradecompound, with many or numerous spikelets. 39. Basal leaves equitant, laterally compressed, without prominent midnerve, the outer margin thin, the inner grooved . . . . 8. F. littoralis 39. Basal leaves when present not equitant, dorsiventrally compressed, with prominent midnerve and thickened, rib-like margins. 40. Stems obtusangular or subterete, not deeply grooved. 41. Spikelets globose, ovoid, or ellipsoid, obtuse, 3-4 mm wide. Glumes muticous, the midnerve ending somewhat below the ape cae . 7. F. globulosa 41. Spikelets lanceolate, acute, 1/4 mm wide. Glumes apiculate or mucronulate by the more or less excurrent midnerve ~ ckinaiaal stuart Siok & tase Nee ee 40. Stems acutely 4—5-angled, deeply grooved. 42. Nut obovoid, densely verruculose, its epidermal cells transversely linear in 4—6 vertical rows on each face. 43. Annual with fibrous roots. Spikelets angular, 1— 1% mm wide. Glumes distinctly keeled, apiculate by the shortly excurrent midnerve, \(~ 14)mmlong. . . 10. F. miliacea 43. Rhizomatous perennial. Spikelets terete, 114-2 mm wide. Glumes ares keeled, muticous,1'44mmlong . . . . pete Oa eres cos aphylla 546 FLORA MALESIANA [ser. I, vol. 7° 42. Nut broadly obovoid, smooth or very sparsely verruculose, its epidermal cells transversely elliptic or oblong in 9—13 vertical rows on each face, or nut obscurely reticulate. 44. Inflorescence 2—4 (rarely up to 7) cm long. Involucral bracts very short, up to 1/4 cm. Spikelets, 3-5 by 2mm. Glumes 2-2/4 mm long. Stamens (2-)3. Style 1 mm long. Nut %-%) mm long ._.. . . II. F. salbundia 44. Inflorescence 12—20 cm long, ‘rarely ‘shorter, much interrupted. Lowest bract 4—8 cm long. Spikelets 2— 3 by 1-114 mm. Glumes wy VA mm Lone Stamens 1-2. Style 4-34 mm long. Nut 4-4 mm long . . 12. F. anisoclada 36. Cauline leaves with well developed blades, ¢ or all the leaves basal. 45. Spikelets 2-4 mm wide. Glumes 4)4—6 mm long. Style 2/4—5 mm long. 46. Glumes mucronulate. Nut obovoid-pyriform, truncate or somewhat depressed at the top. Rhizome not creeping. . . . 17. F. insignis 46. Glumes muticous. Nut broadly obovoid, rounded at the top. Rhizome creeping. 16. F. pierotii 45. Spikelets 1-2 mm wide. Glumes 114-2/4 mm long. Style 4-1 mm long. 47. Stems strongly compressed, ancipitous in the upper half. Leaves 3-8 mm wide. Lowest bract overtopping the inflorescence. . . 14. F. dura 47. Stems not or but slightly compressed. Leaves at most 3 mm wide. Lowest bract much shorter than the inflorescence. 48. Perennial with stiff, coriaceous leaves and rigid, 1-2 mm thick stems. Leaves not twisted. 19. F. cymosa 48. Small annual with herbaceous, strongly falcate leaves twisted to the left, and setaceous stems. 21. F. obtusata 35. Spikelets in clusters, some solitary ones often added. 49. Leaves and glumes glabrous. Rhizome not creeping, or absent. 50. Small annual with setaceous, 10-25 cm tall stems and 1-1/4 mm wide leaves. Inflorescence a single, hemispherical or subglobose head. Glumes mucronate from the sinus of the bilobed apexoer . 22. F. schultzii 50. Perennials with stouter stems and broader leaves. ‘Inflorescence usually with well developed rays and several clusters of spikelets, rarely subcapitate. Glumes muticous, or mucronulate from the apex. 51. Glumes distinctly mucronulate, 3-5 mm long, irregularly subspiral. Spikelets Gee )3-4 mm Widees ta . 37. F. eragrostis 51. Glumes muticous or ‘apiculate, pss 2% mm 1 long, regularly spiral. Spikelets 14-2 mm wide. 52. Nut whitish or stramineous, *#4—1 mm long. Connective of the anthers distinctly produced, bristly at the top . . 18. F. falcata 52. Nut blackish, 74- 44 mm. ‘Connective but shortly produced, smooth . . 19. F. cymosa 49. Leaves densely silky-pubescent beneath. Glumes pubescent. Rhizome creeping. 20. F. sericea 18. Stigmas 2. Nut strongly dorsiventrally compressed, biconvex or planoconvex. 53. Nut coarsely rugulose by transverse wavy ridges. Inflorescence consisting of a single terminal spikelet. 54. Spikelet more or less nodding, broadly ovoid or ovoid, 3—S mm wide. Style strap- eee Y4-Ko mm wide. Nut with 3—S(—6) ridges... . 73. F. nutans 54. Spikelet erect, lanceolate or oblong- -ovoid, {2 3% m mm wide. Style less than YY mm wide. Nut with 5-8 ridges. 555 Spikelet 6-12 by 2-314 mm. Nut broadly obovate or orbicular, 14-174 mm long and wide, stramineous or brownish. . : . 72. F. acuminata 55. Spikelet 4-6 by 1- VA(-2) mm. Nut ‘obovate, Y- Aol- 1) by ve Al 4) mm, white or pale stramineous. . . 74. F. acicularis 53. Nut smooth or verruculose, but not ‘coarsely rugulose. 56. Stems strongly flattened, ancipitous. Rachilla broadly winged . . . . . 415. F. scaberrima 56. Stems not ancipitous. 57. Glumes oblong-ovate or oblong-lanceolate, twice or more than twice as long as broad. Stems usually terminated by a single spikelet. 58. Flower-bearing glumes muticous. Inflorescence with 1—3 spikelets, often subtended by a short- bladed bract. Nut reticulate by the small epidermal cells in about 30 vertical rows on either face. 68. F. polytrichoides 58. All the glumes shortly but distinctly mucronulate. Stems always with a single spikelet. 59. Leaf-sheaths and sometimes also the blades hairy. Glumes obtuse, 2—2’4 mm long. Stamen 1. Nut reticulate by the transversely oblong epidermal cells in about 15 vertical rows on either face. 69. F. tenuicula 59. Glabrous. Glumes acute, 3 mm long. Stamens (1—)2-3. Nut smooth. . . . 70. F. celebica 57. Glumes ovate or orbicular, less than twice as long as broad. 60. Stem-base clothed with bladeless, subinflated sheaths. Spikelets when young globose. Glumes muticous. Nut verruculose, its epidermal cells in 12—15 vertical rows on either face. 7. F. globulosa 60. Blades of the cauline leaves well developed, rarely the upper one short, or all the leaves basal. Other characters not united. 61. Leaves ligulate (sheaths and blades separated from each other by a fringe of short hairs or a membranous projection). 1974] CYPERACEAE (Kern) 547 62. Glumes hairy, at least in the apical part. 63. Lower sheaths coriaceous, shining brown or castaneous. Blades of the cauline leaves rather short, up to 10 cm by }4-1/4 mm. Involucral bracts usually shorter than the inflorescence. Spikelets acute. Glumes pubescent, ferrugineous . wet Vey Se 45. F. ferruginea 63. Lower sheaths herbaceous, stramineous or ferrugineous. Blades of the cauline leaves long, up to 30 cm by 14-2 mm. Lowest involucral bract usually overtopping the inflorescence. Spikelets obtusish. Glumes densely tomentose, usually dark brown . . . . +. +46. F. sieberiana 62. Glumes glabrous. 64. Nut smooth, distinctly stipitate by the conspicuous, 4-24 mm long gynophore. 65. Apex of the nut broadly emarginate by the 14-4 mm wide style-scar. 66. Stems tufted. Spikelets 1-3 to the stem, 3—4 mm wide. Glumes 4)4—5 mm long, anthers ca lummestyleves2mmiitw ike cos ee OS Mere aie tre 40 RR caesia 66. Stems approximate on the short-creeping rhizome. Inflorescence consisting of a single, 4—5 mm wide spikelet. Glumes 6—7 mm long, anthers 2/4—3 mm, style 314-4 mm. 50. F. subalata 65. Apex of the nut rounded or umbonulate, not emarginate. 67. Inflorescence with 1—2(—3) spikelets which are 3-4 mm wide. Glumes 2)4-3 mm long. Anthets, min long. se eerie whos MEY ES erie) Ve Bu Sorters Baas R" <—34 by 4-% mm. Distr. Lower Bengal, Lower Burma, Peninsular Thailand; in Malesia: Malay Peninsula(P. Langkawi, Pahang, Malacca, Johore, P. Penang, Singapore), Borneo (Sarawak: Kuching; W. Borneo: Banjer- masin, Sanggau; N. Borneo: Jesselton, Mt Kinabalu); Celebes: Pare-Pare; specimens in the Leyden Herb. “Sumatra, leg. WAITZ”’ may be mislabelled. Ecol. Grass-fields, grassy road-sides, at low alti- tude; on Mt Kinabalu collected at 900 m. Note. Clearly distinct from F. tenera R. & S., of which CLARKE considered it a variety. Mainly characterized by the flat, relatively broad, obtuse leaves and the muticous, glabrous, glumes. By the leaves arising in a left hand spiral and the tendency to distichous arrangement of the glumes it shows affinity to the species of sect. Fuscae (e.g. F. fimbri- stvloides). 22. Fimbristylis schultzii Borck. Linnaea 38 (1874) 391; BENTH. FI. Austr. 7 (1878) 320; S. T. BLAKE, Proc. R. Soc. Queens]. 58 (1947) 45; KERN, Blumea 8 (1955) 116. — F. platystachys Boeck. Linnaea 38 (1874) 390; Doin, Bibl. Bot., Heft 85 (1915) 464, incl. var. schultzii DomiN. — Iriha platystachys O.K. Rev. Gen. Pl. 2 (1891) 753. Glabrous annual with fibrous roots. Stems very slender, setaceous, tufted, quadrangular, smooth or "The other part belongs to F. onchnidiocarpa KERN. minutely scabrid at the top, leafy at the base, 10- 25 cm by 4-4 mm. Leaves radical and 1-2 some- what higher on the stem, much shorter than the stems, subfalcate, flat or with inrolled margins, acute, scabrid on the margins in the upper part, 1-114 mm wide; ligule absent. Inflorescence capitate, hemi- spherical or subglobose, with 5—12 spikelets, 5— 10 mm across. Involucral bracts 4—6, spreading or reflexed, dilated at the base, the lower 1—2 somewhat longer than the inflorescence, 1-174 cm long. Spike- lets sessile, stellately spreading, oblong-ovoid, angu- lar, acute, few- to several-flowered, 3—8 by 114-2 mm; rachilla winged. G/umes spiral, membranous, erect, ovate, keeled, with strong midrib and nerveless, stramineous to ferrugineous sides, glabrous or denticulate-ciliolate at the upper edge, mucronulate to strongly mucronate from the incised apex, 1/4-3 mm long; mucros almost straight to strongly recurved, 4-174 mm long. Stamens 2-3; anthers oblong or linear, 4-1/4 mm. Sryle triquetrous, pyramidally thickened at the base, glabrous, (?4—)1- 2 mm long; stigmas 3 (or in a few flowers 2), about as long as the style. Nut obtusely trigonous, obovoid or obovoid-pyriform, shortly stipitate, obtuse, minutely umbonulate, smooth and subcancellate by the slightly impressed epidermal cells in c. 9 vertical rows on each face, or densely verruculose, whitish or shining dark brown, 4—34(-1) by 14-24 mm. Distr. Australia (Northern Territory, W. Aus- tralia, Queensland); in Malesia only twice collected in the Lesser Sunda Is. (W. Bali: Gilimanuk; Sumba: Waikelo). Ecol. In Bali in grassy fields at low altitude. Notes. BENTHAM, /.c., is of opinion that BOECKELER’S separation of F. schultzii and F. platystachys is apparently justified by the specimens in the Berlin herbarium, but as those of the Kew herbarium differ much less from each other, and F. MUELLER’S specimens are intermediate in most respects, he finds himself unable to give tangible characters even for two distinct varieties. S. T. BLAKE also considered F. schultzii and F. platystachys as one species. I find the two SCHULTZ collections in the Kew herbarium, though young, fairly distinct, but having seen only a few collections I have provisionally followed BENTHAM and BLAKE. The plants from Sumba (IBoeT 358a in BO) agree very well with the type collection of F. schultzii (SCHULTZ 96, according to BOECKELER from Adelaide, but probably from Port Darwin). The glumes are 1/4-2 mm long with a short, but slightly spreading awnlet }4—)4 mm long, not ciliate, the anthers ’-¥% mm long, the style 34-1 mm, the nut rather pyriform than obovoid, not verruculose, and brown. The Bali plants (VAN STEENIS 7586), though similar in habit and certainly closely related, differ considerably in the spikelets, which are strongly squarrose by the stout, recurved, long awns, in the denticulate- ciliolate upper margins of the glumes, and in the nuts, which are obovoid, densely verruculose, and white. By these characters they come near to the type collection of F. platystachys (SCHULTZ 792), in which, however, the glumes are larger (c. 3 mm without the up to 134 mm long awn), and the anthers c. 1144 mm long (in the Bali plants respectively 114-2 mm and c. 4% mm). S. T. BLAKE, Proc. R. Soc. Queensl. 58 (1947) 46, described S. stellata, “‘very similar to F. schultzii in 560 FLORA habit, foliage, anthers, style and shape of glumes”’, but differing “in the broader, more squarrose spike- lets, the more coarsely and obtusely keeled glumes more densely hairy at the margins and with longer stouter awnlets, and in the white (not black) simply obovoid (neither pyriform nor obcordate) somewhat larger nut which also appears to be more densely and MALESIANA [ser. I, vol. 7? evenly verrucose”’. Ina duplicate of the type collection I fail to see in what way it differs from F. platystachys. I am inclined to consider F. schultzii and F. platystachys well-founded taxa, F. stellata synon- ymous with the latter, and the Bali plants a local race of it. 5. Section Leptocladae Onwil, J. Jap. Bot. 14 (1938) 572. — Fimbristylis ser. Leptocladae OHwW1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 54. F.. leptoclada BENTH. Type species: 23. Fimbristylis leptoclada BENTH. Fl. Hongk. (1861) 393, non FI. Austr. 7 (1878) 314; CLARKE, FI. Br. Ind. 6 (1893) 647; RipL_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 96; Camus, Fl. Gén. I.-C. 7 (1912) 118; MerR. En. Borneo (1921) 61; En. Philip. 1 (1923) 124; Rip. Fl. Mal. Pen. 5 (1925) 159. — F. retusa THWAITES, En. Pl. Zeyl. (1864) 349; Boeck. Linnaea 37 (1871) 46. — Iriha leptoclada O.K. Rev. Gen. PI. 2 (1891) 753. —Iriha retusa O.K. l.c.— F. disticha (non Boeck.) Camus, FI. Gén. I.-C. 7 (1912) 125, p.p. Annual (?), glabrous except for the glumes. Stems densely tufted, very slender, often setaceous, angular, smooth, or scabrid at the top, 15-65cm by %4- 1(—2) mm. Leaves basal, usually much shorter than the stems, flat but narrow, rather abruptly acuminate, with scabrid margins, 1—2?4 mm wide; ligule absent; lower sheaths stramineous or ferrugineous. Inflo- rescence simple, sometimes head-like, consisting of 1-7 small clusters of spikelets, 1—-2cm across. Involucral bracts (1—)2—3, very short, the longest 4-1(-2) cm. Rays erect to patent, often curved, compressed, smooth, up to 2cm long. Spikelets 3-8 to each cluster, sessile, globose to ovoid or ellipsoid, slightly angular, obtuse, 10—15-flowered, dark brown, 2-3 by 1/4 mm; rachilla narrowly winged. G/umes spiral, membranous, ovate or sub- orbicular, very obtuse, more or less emarginate- bilobed at the apex, mucronulate from the sinus, keeled, with prominent midrib, nerveless, gland- dotted sides, and densely ciliate margins, 1/4-1/4 by 1-144 mm. Stamens 1(-—2); anthers oblong- linear, Y- ¥, mm long. Style triquetrous, abruptly bulbously or pyramidally thickened in the lower )4-'%, glabrous, 4-34 mm long; stigmas 3, about as long as the style. Nut very obtusely trigonous, obovoid, shortly stipitate, inconspicuously umbonulate, verru- culose (see Note), whitish to stramineous, 49-7 by -Ko mm; epidermal cells isodiametric, roundish or hexagonal. Distr. Ceylon, Peninsular Thailand, Indo-China, S. China; in Malesia: Sumatra (E. Coast and W. Coast Res.), Malay Peninsula (Setul, Pahang, Malacca, P. Penang, Singapore), Lesser Sunda Is. (Flores), N. Borneo and W. Borneo, Philippines (Luzon, Culion, Palawan). The collection from the Moluccas (Amboina, ROBINSON 1901 in US) may be mislabelled as all other specimens of this number examined belong to F. dichotoma. Ecol. In dry open sandy soil at low altitudes, in P. Penang on rocks dripping with water. Vern. Imbulu tano, Sum. E.C Note. The only known New Guinean collection differs from the typical variety (var. leptoclada) and may be distinguished as: var. etuberculata KERN, Blumea 10 (1960) 637. Nut smooth, not verruculose. Distr. W. New Guinea: aerodrome. Tanah Merah, on 24. Fimbristylis hispidula (VAHL) KUNTH, En. 2 (1837) 227; Steup. Syn. 2 (1855) 112; Boeck. Linnaea 37 (1871) 27; KERN, Blumea 8 (1955) 120; Rein- wardtia 4 (1956) ispi 2 (1806) 276. — Isolepis exilis KUNTHin H.B.K. Nov. Gen. 1 (1815) 224. — F. exilis R. & S. Syst. Veg. 2 (1817) 98. — Abildgaardia pubescens PRESL, Rel. Haenk. 1 (1828) 180, non F. pubescens Link. — F. preslii KUNTH, En. 2 (1837) 228; SteupD. Syn. 2 (1855) 108; F.-VILL. Nov. App. (1882) 308; Merr. En. Philip. 1 (1923) 127.— F. clavinux CLARKE, Il. Cyp. (1909) t. 41 f. 1-3. — Bulbostylis pubescens Svens. N. Amer. Fl. 18 (1957) 542. — Fig. 40. Annual (always ?). Stems densely tufted, setaceous, angular, smooth or hispidulous-scabrid just below the inflorescence, 5-30cm by 4-4 mm. Leaves basal, much shorter than the stems, filiform, erect, hispidulous, /4—)4 mm wide; ligule absent; sheaths stramineous, hairy. Inflorescence consisting of 1—5 (sometimes more) spikelets, the rays when present 1-1/4 cm long. Involucral bracts 1-2, glume-like, mucronate, or filiform and up to 2 cm long. Spikelets solitary, erect, lanceolate, subterete, acute, 4—12- flowered, 4-10 by 2-214(-3)mm _ wide; rachilla winged. G/umes spiral, membranous, ovate-lanceo- late, acute, muticous or minutely mucronulate, densely short-pubescent, with strong midnerve on both sides bordered by a yellowish stripe, ferrugi- neous or brownish often sanguineous-variegated sides and densely ciliate margins, 3—4 mm long. Stamens 1-3; anthers linear, 1-2 mm long. Style triquetrous, pyramidally thickened at the base, glabrous, or sparsely spinulose-ciliolate at the top, 134-214 mm long; stigmas 3, about as long as the style. Nut trigonous with prominent angles, turbinate, usually slightly depressed at the apex, minutely stipitate, umbonulate in the apical depression, not tubercled, with up to 10 prominent transverse wrinkles on each side, brownish or fumose, 1-114 mm long and wide; epidermal cells linear, longitudinally elongate. Distr. Very common in tropical Africa, rare in tropical America (Mexico, Columbia); very rare in Malesia: E. Java (near Situbondo), Lesser Sunda Is. 1974] Cy PERACEAE (Kern) 561 Ee eoas nesses) Fig. 40. Fimbristylis hispidula (VAHL) KUNTH. a. Habit, x 74, b. spikelet, x 6, c. glume, d. anther, e. nut with style and stigmas, all x 13 (a—e RAMos BS 32797). (Wetar, Flores), Philippines (Luzon: Burgos, Ilocos Norte Prov.). Ecol. In Java in grassy fields, in Wetar in Euca- lyptus forest and swampy places, 425-500 m. Notes. Abildgaardia pubescens PRESL was based on specimens collected by HAENKE “in Mexico et in Luzon”. MERRILL, /.c., thought it was probably not Philippine. However, in July 1918 specimens were collected near Burgos in Luzon which undoubtedly represent PRESL’s species. They cannot be separated specifically from the widely distributed and poly- morphous F. hispidula. The inflorescence usually consists of a single terminal spikelet, sometimes with 1—2 lateral ones added, the glumes are 4 mm long, the style is sparsely ciliolate and c. 2 mm long. The very slender specimens from Java, Flores and Wetar perfectly agree with some of the African specimens I have seen. The inflorescence is anthelate consisting of 1—5 spikelets and subtended by small setaceous bracts, the glumes are 3 mm long, the style is glabrous and c. 134 mm long. Nuts with longitudinally elongate epidermal cells are very rare in Fimbristylis, whereas they frequently occur in the closely related genus Bulbostylis, to which F. hispidulashows affinity also in other respects. VAN DER VEKEN, Bull. Jard. Bot. Brux. 35 (1965) 327, found that its embryo is of the Bulbostylis type. According to Svenson, N. Amer. FI. 18 (1957) 540 & 550, the cells are always longitudinally elongate in Bulbostylis, but never so in Fimbristylis. Consequently he transferred Abildgaardia pubescens PRESL to Bulbostylis. However, in B. capillaris and B. barbata, also to him true Bulbostylis species, the epidermal cells are isodiametric. Nowadays for F. hispidula mostly the name F. exilis (KUNTH) R. & S., based on Isolepis exilis KUNTH, is accepted. KUNTH deliberately rejected his epithet exi/is in favour of hispidula, but he did not cite Scirpus hispidulus in synonymy. In VAHL’s Enumeratio 94 spp. of Scirpus are enumerated, two of them without specific epithet. Of the remaining 92 species 91 are meticulously accounted for in KUNTH’s Enumeratio, only Scirpus hispidulus VAHL is missing. There is, therefore, ample circumstantial evidence that KUNTH based his name on that of VAHL, the more so as VAHL’s type locality (Guinea) is cited. 25. Fimbristylis recta F. M. BAILEY, 3rd Suppl. Syn. Queensl. Fl. (1890) 80; Queens]. Fl. 6 (1902) 1762: S. T. BLAKE, Proc. R. Soc. Queens]. 58 (1947) 44; J. Arn. Arb. 35 (1954) 211; KERN, Blumea 8 (1955) 120.— F. xyridis R.Br. var. rigidula BENTH. FI. Austr. 7 (1878) 307.— F. macgillivrayi CLARKE, Kew Bull. add. ser. 8 (1908) 24. — F. stricticulmis Domin, Bibl. Bot., Heft 85 (1915) 452. Perennial with short rhizome. Stems densely tufted, rigid, angular-sulcate, somewhat incrassate just below the inflorescence, antrorsely scabrous, 30-80 cm by c. 1 mm. Leaves basal, shorter than the stems, up to 30 cm long, erect, flat, rigid, with obtuse or rounded apex, glabrous, with scabrous margins, greyish green or glaucous, 2-3 mm wide; ligule absent; sheaths white-scarious in front. Inflorescence 562 FLORA MALESIANA [ser 1, vor consisting of a single terminal spikelet. Spikelet erect, ellipsoid or slightly obovoid, terete, acute, densely many-flowered, 8—20 by 5—6 mm; rachilla narrowly winged. Glumes spiral, membranous, ob- long-ovate, very obtuse, minutely apiculate, not keeled, with strong midnerve, nerveless sides, and densely ciliate upper margin, reddish brown in the apical half, much paler in the basal half, 5-7 by 34%-4 mm; lower empty glumes much smaller, manifestly mucronate. Stamens 3; anthers linear, 3-4 mm long. Style triquetrous, pyramidally thick- ened at the base, glabrous, 4—5 mm long; stigmas 3, somewhat shorter than the style. Nut trigonous with convex faces, tricostate, pyriform, shortly stipitate, verrucose in the upper part, pale brown, 134-2 by 114-134 mm; epidermal cells isodiametric hexagonal to roundish. Distr. Tropical Australia (Northern Territory, Queensland, Hammond I. in Torres Strait); in Malesia: New Guinea (Papua, Western Div.: Tarara, Wassi Kussa River). Ecol. In the only known Malesian locality in savannah forests, common in grass on ridges, at low altitude. 26. Fimbristylis furva R.Br. Prod. (1810) 228; BENTH. FI. Austr. 7 (1878) 318; BAILEY, Queens]. Fl. 6 (1902) 1766; Doin, Bibl. Bot., Heft 85 (1915) 460; S. T. BLAKE, J. Arn. Arb. 35 (1954) 219; KERN, Blumea 8 (1955) 121. — Iriha furva O.K. Rev. Gen. PlEZ soma. Perennial (always ?). Stems erect, tufted, obtus- angular, slightly compressed, glabrous and smooth, 10—30(—50) cm by 4-1 mm, the base clothed with 2—3 bladeless or very short-bladed, obliquely truncate sheaths ferrugineous-membranous at the mouth. Leaves shorter than the stems, flaccid, flat, obtuse or abruptly acuminate, glabrous, scabrid on the margins in the upper part, (1/4—)2—4 mm wide; ligule absent. Inflorescence simple or compound, relatively small, dense to rather loose, with 4—25 spikelets, 1-5 cm long and wide. Involucral bracts small, glume-like, or the lower 1-2 produced into a short point. Primary rays 4—6, obliquely erect but outer ones widely spreading, compressed, smooth, up to 3 cm; secondary rays very short, widely spreading. Spikelets solitary, ovoid or oblong-ovoid, terete, acute, brown, 4-8 by c. 2mm; rachilla winged. G/umes spiral, membranous, appressed, ovate or broadly ovate, obtuse, mucronulate, slightly keeled, with prominent midnerve, glabrous, nerveless sides, and scarious, ciliate margins, 244-3 by c. 2mm. Stamens 2-3; anthers oblong-linear, 1-114 mm; connective dis- tinctly produced, setulose at the top. Style tri- quetrous, pyramidally thickened at the base, sparsely ciliate, hispidulous at the base, 1’4-124 mm; stigmas 3, about as long as or longer than the style. Nut obtusely trigonous, obovoid, shortly stipitate, um- bonulate, smooth or scaly-verruculose, purplish black, %-/K, by Z-)4 mm; epidermal cells longi- tudinally oblong to almost isodiametric. Distr. Australia (N. Territory, Queensland in- clusive of the islands in Torres Strait); in Malesia: New Guinea (W. New Guinea: near Merauke; Papua: W. Div., Wassi Kussa R., Mai Kussa R.), Aru Is. (P. Trangan). Ecol. In damp grassland, marshy grounds, Melaleuca swamps, at low altitudes; locally often abundant. Note. The New Guinea plants are perennial. Some of the Australian collections I have seen (e.g. S. T. BLAKE 17548) are apparently annuals, which differ moreover from the perennials by the smaller glumes (114 mm), anthers (74 mm), style (1 mm), and nuts Y mm). 27. Fimbristylis lanceolata CLARKE, Kew Bull. add. ser. 8 (1908) 25; Domin, Bibl. Bot., Heft 85 (1915) 460; KERN, Blumea 8 (1955) 121. Perennial with short rhizome. Stems densely tufted, strongly compressed (almost ancipitous), glabrous, or pilose especially towards the apex, smooth, (10-)20-40 cm by 34-1)4 mm, the base clothed with 1—2 tubular, bladeless, 5-8 cm long sheaths. Basal /Jeaves shorter than the stems, erect, flat, rather rigid, obtuse or abruptly acuminate, glabrous, scabrid on the margins in the upper part, 114-2 mm wide; ligule absent; lower sheaths bladeless, with membranous margins, ferrugineous or brown. Inflo- rescence simple or subcompound, rather dense, with 2-9 spikelets, 114-5 cm long, 2—7 cm wide. Involu- cral bracts 2—3, very short, glume-like, cuspidate, 1—5 mm. Primary rays |—5, finally patent to arcuately reflexed, compressed, smooth, 14-2 cm; secondary rays when present very short, c. 14cm. Spikelets solitary, linear, terete, subacute, loosely many- flowered, 10-25 by 2 mm; rachilla winged. G/umes spiral, thinly membranous, oblong-ovate, very obtuse and notched, mucronulate just below the apex, scarcely keeled, densely reddish gland-dotted in the apical half, 3—5-nerved (midnerve prominent, lateral nerves obscure), fulvous or fuscous, 4-474 by 2)4-234 mm, the margins long-ciliate in the upper Y—-34. Stamens 3; anthers linear, 2-214 mm; con- nective distinctly produced, the appendage oblong- ovate, crested by conspicuous bristles. Style triquetrous, pyramidally thickened at the base, glabrous except for the hispidulous base, 3-324 mm; stigmas 3, shorter than the style. Nut very obtusely trigonous, broadly obovoid, shortly stipitate, hardly or not umbonulate, densely verruculose, at first whitish, ultimately blackish, 2%) by 4 )—*%% mm; epidermal cells obseure, longitudinally elliptic or oblong. Distr. North coast of Australia (Raffles Bay); in Malesia: SE. Celebes (Rumbia Distr.), S. Moluccas (Aru Is.: P. Trangan). Ecol. In monsoon-forests and savannahs, at low altitude. 28. Fimbristylis macassarensis STEUD. Syn. 2 (1855) 109; Mia. FI. Ind. Bat. 3 (1855) 318; Kern, Blumea 8 (1955) 122; in Back. & Bakh. f. Fl. Java 3 (1968) 462.— F. tenera (non SCHULT.) CLARKE, Philip. J. Sc. 2 (1907) Bot. 95; Merr. En. Philip. 1 (1923) 126. — F. corniculata MeRR. Philip. J. Sc. 7 (1912) Bot. 231; Fl. Manila (1912) 116; En. Philip. 1 (1923) 122. — Fig. 41. Annual. Stems densely tufted, compressed, obtus- angular, glabrous and smooth, 10-20cm by /4- 1 mm, the base clothed with 1—2 tubular, bladeless, up to 3 cm long sheaths. Basal /eaves half as long as the stems, flat, rather firm, acute, glabrous, 34- 14 mm wide, the margins scabrid in the upper part; Cy PERACEAE (Kern) 563 Fig. 41. Fimbristylis macassarensis StEUD. a. Habit, x 7, b. spikelet, x 6, c. glume, d. anther, both x 13, e. tip of anther, x 50, f. style, g. nut, both x 13,h. outer cells of nut, strongly enlarged (a~h BUNNEMEVER 10837). ligule absent. Inflorescence simple, rarely subcom- pound, with 2-10 spikelets, 1-3 cm across. Involu- cral bracts 1-2, membranous, ovate, with retuse, cuspidate apex, 2-5 mm long. Primary rays 1-4, finally patent to arcuately reflexed, compressed, smooth, up to 15 mm; secondary rays when present very short, up to 5m. Spikelets solitary, ovoid or oblong-ovoid, terete, obtuse, several-flowered, dull fuscous, 5-10 by 2—3mm. Glumes spiral, thinly membranous, ovate, mucronulate, scarcely keeled, densely glandular-puncticulate in the upper half, with strong green midnerve, nerveless sides, and margins long-ciliate in the upper 14-24, rounded, retuse or notched at the apex, 2~3 by 1/4-2 mm. Stamens (2-)3; anthers linear, 4-114 mm; con- nective distinctly produced, bristly at the top. Style triquetrous, slightly dilated at the base, hispidulous at the base, otherwise glabrous or sparsely ciliate, 1—2 mm long; stigmas 3, somewhat shorter than to as long as the style. Nut obtusely trigonous with convex sides, obovoid, subsessile, not or hardly umbonulate, verruculose or smooth, black, 4-74 by %-/4 mm; epidermal cells longitudinally elliptic or oblong. Distr. Malesia: Philippines (Central Luzon); SW. Celebes (Macassar; Mt Galesang near Malino), Madura (near Pamekasan). Ecol. Open grasslands, roadsides, at low altitudes. Notes. Only known from a few rather poor collections. The nuts are usually verruculose, smooth only in the specimens from Mt Galesang. In the Luzon plants the anthers are very small (?4 mm). F. macassarensis has nothing to do with F. tenera SCHULT., to which species the oldest Philippine collection was referred by CLARKE. It is very near to F. lanceolata CLARKE, from which it differs rather by quantitative than qualitative characters. 6. Sect. Heleocharoides BENTH. Fl. Austr. 7 (1878) 301. — Fimbristylis ser. Heleocharoides KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 100. Type species: F. pauciflora R.BR. 29. Fimbristylis dictyocolea S. T. BLAKE, J. Arn. Arb. 35 (1954) 209, f. 1; RayM. Dansk Bot. Ark. 23 (1966) 328.— F. cardiocarpa (non F.v.M.) Kix. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 9. Glabrous perennial with short rhizome. Stems tufted, erect, setaceous, obtusely 5-angular, striate, smooth or scabrid just below the inflorescence, 20-40 cm by 4-1 mm, the base clothed with 2-3 obliquely truncate, bladeless sheaths in front dis- integrating into fine, herringbone-shaped fibres. Basal /eaves few, shorter than to about as long as the stems, filiform, triquetrous, laterally compressed, smooth, }4—1 mm wide; ligule absent. Inflorescence consisting of a single spikelet. Spike/et erect, oblong or ellipsoid, terete, acute, densely many-flowered, pale green, 7-9 by c. 3 mm; rachilla not winged. Glumes spiral, membranous, oblong, obtuse, muti- cous, scarcely keeled, 1-nerved, hyaline-margined, 4—6 by 1/4—2 mm; lower 1-3 glumes larger, empty. Stamens 3; anthers linear; 2—2!4 mm long. Style 564 FLORA MALESIANA [ser. I, vol. 7? slender, compressed-triquetrous, incrassate at the base, minutely ciliolate, 3-5 mm long; stigmas 3, much shorter than the style. Nut trigonous with convex sides, obovoid, subsessile, umbonulate, ver- ruculose, shining, stramineous, c. | by 34mm; epidermal cells minute, hexagonal. Distr. Thailand (Chantaburi), Cambodia; in Malesia: New Guinea (W. New Guinea: Kurik near Merauke; Papua, W. Division: Tarara, Wassi Kussa R.; Mabaduan). Ecol. In open wet grasslands, in shallow pools, in savannah-forests, at low altitudes. Note. Allied to F. pauciflora, from which it can easily be distinguished by the coarser habit, the leaf- sheaths disintegrating into fine reticulate fibres, the larger spikelet, and the more oblong, obtuse, single- nerved glumes. 30. Fimbristylis pauciflora R.Br. Prod. (1810) 225; BENTH. FI. Austr. 7 (1878) 303; CLARKE, FI. Br. Ind. 6 (1893) 633: RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 90; Koorp. Exk. FI. Java | (1911) 198; ibid. 4, Atlas (1922) f. 249; Camus, Fl. Gén. I.-C. 7 (1912) 98, f. 13, 10; Riv. Fl. Mal. Pen. 5 (1925) 154; KUx. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 9: S. T. BLAKE, J. Arn. Arb. 35 (1954) 210. — Trichelostylis filiformis Nees in Wight, Contr. (1834) 102. — F. filiformis KunrtTH, En. 2 (1837) 221; Steup. Syn. 2 (1855) 106; Mia. FI. Ind. Bat. 3 (1856) 314; Boeck. Linnaea 37 (1871) 23. — F. pumila BENTH. in Hook. Lond. J. Bot. 2 (1843) 239 (p.p. ?); SrEUD. Syn. 2 (1855) 106; Mig. Fl. Ind. Bat. 3 (1856) 315.— F. malaccana Boeck. Flora 41 (1858) 597. — Chaetocyperus seta- ceus (non NEES) KuRZ, Nat. Tijd. N. I. 27 (1864) 223. — Iriha pauciflora O.K. Rev. Gen. Pl. 2 (1891) iS2. Perennial (always ?), glabrous. Stems densely tufted, finally forming large clumps some dm in diam., erect or finally prostrate, setaceous, striate- sulcate, smooth, greyish green, (5—)10—30cm by Y%-4 mm. Leaves all reduced to membranous, obliquely truncate, ferrugineous sheaths not disinte- grating into reticulate fibres, or 1-2 with a short liform blade sometimes up to 6cm long; ligule absent. Inflorescence consisting of a single terminal spikelet. Spikelet erect, oblanceolate or narrowly oblong, terete, acute, 4—9-flowered, 3-6 by 1- 1'4 mm; rachilla winged. Glumes spiral, cartilagi- nous, erect, lanceolate, acute, muticous, scarcely keeled, pale, 2/4-3 by c. 144mm; keel green, indistinctly 3—5(—7)-nerved, sides nerveless; lower (1-)2 empty glumes larger, almost as long as the spikelet, several-nerved, 3-4 by 134 mm. Stamens 1—2(—3); anthers linear, 114-2 mm long. Sryle tri- quetrous, slightly thickened at the base, sparsely and minutely ciliolate, 11/4-2!4 mm; stigmas 3, or in some flowers 2, much shorter than the style. Nut obtusely trigonous with convex faces, usually dorsiventrally compressed, sometimes biconvex (in the digynous flowers), obovoid, shortly stipitate, not or hardly umbonulate, whitish verruculose, stramineous to brown, 34-% by 4—-*%% mm; epidermal cells irreg- ular, isodiametric or elliptic. Distr. Mergui, Thailand, Indo-China, Ryu Kyu Is., Carolines, N. Australia and Queensland; in Malesia: Sumatra (incl. Riouw Arch., Lingga Arch., Banka and Billiton), Malay Peninsula, Borneo, Anambas and Natuna Is., Moluccas (Ternate, Am- boina, Ceram), New Guinea. A collection said to be from Java (HORSFIELD 89, BM) was probably mis- labelled. Ecol. In damp sandy and clayish localities both shaded and sunny, swamp margins, savannahs, often completely covering jungle-paths, at low altitudes, up to 800 m. Uses. In some parts of the Malay Peninsula, as in Penang, it makes a fair proportion of the weeds in the rice-fallows which are ploughed in for green manure (BURKILL). Vern. Rumput djénggol kambing, r. sapi, r. jambak, r. lumut, r. sarang puyoh, r. girah, M, r. djani, Riouw, kaju ménkirei, Banka. Note. BENTHAM’s description of F. pumila, which does not well accord with F. pauciflora, may partly refer to F. acicularis. The type collection (Amboina, leg. BARCLAY) is a mixture of the two species. 7. Section Signatae KERN, Blumea 8 (1955) 160. Type species: F. signata S. T. BLAKE. 31. Fimbristylis signata S. T. BLAKE, J. Arn. Arb. 35 (1954) 214; KERN, Reinwardtia 4 (1956) 96. — F. debilis F._v.M. Fragm. | (1859) 198; BENTH. Fl. Austr. 7 (1878) 315, non SteupD. 1855. — Iriha debilis O.K. Rev. Gen. Pl. 2 (1891) 753. Annual. Stems very slender, densely tufted, com- pressed, smooth, (10—)25—50 cm by 14-1 mm. Leaves reduced to 2—3 sheaths clothing the base of the stem, rather loose, acuminate, scarious-margined, gla- brous, pale green or stramineous, sometimes pro- duced into short, erect blades; ligule absent. Inflorescence simple or subcompound, small, with (1—)3—5(—9) spikelets, 1-274 by 1-314 cm. Involu- cral bracts 3—4, much shorter than the inflorescence, dilated at the base, up to | cm long. Rays up to 5, obliquely erect to horizontally spreading, com- pressed, smooth or slightly scabrid, 14-114 cm long. Spikelets solitary, oblong to linear-lanceolate, angu- lar, acute, loosely several-flowered, 6-10 by c. 2 mm. Rachilla broadly winged. G/umes spiral, membra- nous, appressed, ovate-lanceolate, acute, mucronate, sharply keeled, with prominent, ciliolate midnerve, nerveless sides, and margins microscopically ciliolate at the top, stramineous or pale brown, 334-4 by 2)4-3 mm (mucro excluded), the midnerve ex- current in a scabrid, ultimately spreading, 1-1/4 mm long awn. Stamens 2; anthers oblong-linear, 1-114 mm long. Style triquetrous, pyramidally thickened at the base, sparsely short-ciliate, 2-214 mm; stigmas 3, much shorter than the style. Nut obtusely trig- onous, pyriform, abruptly narrowed towards the base, subglose in the upper part, shortly stipitate, umbonulate, coarsely tuberculate, dark brown, c. 11% by 1 mm; epidermal cells obscure, almost isodiametric. Distr. Northern Australia; in Malesia: Philippines (Mindanao), New Guinea (Papua, W. Div.: Daru I.). Ecol. Onriver-side (Mindanao); frequent on damp soil in savannah-forest (Daru I.). 1974] Cy PERACEAE (Kern) 565 8. Section Abildgaardia (VAHL) BentuH. FI. Austr. 7 (1878) 299. — Abildgaardia VAHL, En. 2 (1806) 296. — Cyperus sect. Abildgaardia ENDL. Gen. PI. (1836) 119. — Fimbristylis ser. Monostachyae OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 53. Type species: F. monostachyos (L.) Hassk. (Cyperus monostachyos L. = F. ovata (BuRM. f.) KERN). 32. Fimbristylis ovata (BURM. f.) KERN, Blumea 15 (1967) 126; in Back. & Bakh. f. Fl. Java 3 (1968) 467. — Carex ovata Burm. f. Fl. Ind. (1768) 194; KUkK. Pfl. R. Heft 38 (1909) 103. — Cyperus mono- stachyos LINNE, Mant. 2 (1771) 180; Rotts. Descr. et Ic. (1773) 18, t. 13 f. 3. — Cyperus caribaeus RICH. ex Pers. Syn. | (1805) 65, nom. illeg. — Cyperus indicus RicH. ex Pers. /.c. — Abildgaardia mono- stachva VAHL, En. 2 (1806) 296; KUNTH, En. 2 (1837) 247; STEUD. Syn. 2 (1855) 72; Mia. Fl. Ind. Bat. 3 (1856) 296, inc/. var. rigidior STEUD. ex MIQ.; BOECK. Linnaea 37 (1871) 53.— Abildgaardia compressa PRESL, Rel. Haenk. | (1828) 179; Nees, Nov. Act. Ac. Caes. Leop.-Car. 19, Suppl. | (1843) 74; StTeup. Syn. 2 (1855) 72; Mia. Fl. Ind. Bat. 3 (1856) 297. — Abildgaardia javana Nees, Linnaea 9 (1834) 289, fide Ind. Kew. — F. monostachyos Hassk. Pl. Jav. Rar. (1848) 61 (‘monostachya’); STEUD. Syn. 2 (1855) 107; BENTH. FI. Austr. 7 (1878) 308; CLARKE, FI. Br. Ind. 6 (1893) 649; Philip. J. Sc. 2 (1907) Bot. 97; Koorp. Exk. Fl. Java | (1911) 200; ibid. 4, Atlas (1920)\eh 261); Camus, Fl) Gen. I-27 (1912) 122; Mer. En. Philip. | (1923) 124; Ripv. Fl. Mal. Pen. 5 (1925) 159; Back. Onkr. Suiker. (1928) 164, non t. 174; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 18. — F. compressa F.-ViLL. Nov. App. (1882) 307, non R. & S. 1817, nec Boeck. 1874. — Iriha mono- stachya O.K. Rev. Gen. Pl. 2 (1891) 751. Glabrous perennial with short rhizome. Stems densely tufted, compressed-trigonous, smooth, or scabrid at the top, somewhat incrassate at the base, (5—)15—40(-60) cm by 4-1 mm. Leaves basal, shorter than the stems, very narrow, rather abruptly acuminate, smooth, or slightly scabrid at the top, 24-1 mm wide; ligule absent. Inflorescence usually consisting of a single terminal spikelet, more rarely 1(—2) peduncled lateral spikelets added. Spikelets ovate or ovate-lanceolate, strongly compressed, ultimately often twisted, acute, shining stramineous or yellowish, 8—15(—30) by 4-6 mm; rachilla broadly winged. G/umes distichous, but at length often appear- ing subspiral by torsion of the rachilla, subcoriaceous, broadly ovate, acute, mucronate, with prominent 3-nerved keel and nerveless sides, 44-5 by 4— 44 mm; lowest 1-2 empty glumes awned, the awn usually not exceeding the spikelet, or the lowest with a short, 14-174 cm long blade. Stamens 2—3; anthers linear, 14-3 mm long. Style triquetrous, pyramidally thickened at the base, hyaline-fimbriate almost to the base, 2-4 mm long; stigmas 3, much shorter than the style. Nut obtusely trigonous, pyriform, sessile, umbonate, coarsely tuberculate or almost muricate except for the smooth base, very rarely quite smooth (not in Malesia), stramineous to greyish brown, 2-3 by 1)4—-2 mm; epidermal cells minute, isodiametric. Distr. Pantropical; throughout Malesia, but apparently very rare in the Malay Peninsula (Perlis: Kanga; Kedah: P. Langkawi; Singapore). Ecol. Sunny or partly shaded grasslands, sandy heaths, roadsides, teak-forests, at low altitudes, 0-900 m, rarely (in W. Java on Mt Gedeh and in New Guinea) collected at 1200 m. Vern. Babawangan, kukutjaian, S, sékétan, J, te léntejan, Md. 9. Section Fuscae Ouwi, J. Jap. Bot. 14 (1938) 571. Type species: F. fusca (NEES) C. B. CLARKE (Abildgaardia fusca NEES). 33. Fimbristylis cinnamometorum (VAHL) KUNTH, En. 2u0loS7). 229: Sreup. Syn. 2 (1855) 113; Borck: Linnaea 37 (1871) 35; KOK. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 9; S. T. BLAkg, J. Arn. Arb. 35 (1954) 220; KERN, Blumea 8 (1955) 123; ibid. 15 (1967) 433. — Scirpus cinnamometorum VAHL, En. 2 (1806) 278. — F. cvperoides R.BR. Prod. (1810) 228; KUNTH, En. 2 (1837) 244; CLarke, Fl. Br. Ind. 6 (1893) 650, incl. var. cinnamometorum CLARKE; Ill. Cyp. (1909) t. 44 f. 1-4; Bent. FI. Austr. 7 (1878) 317. — F. biflora Boeck. Linnaea 38 (1874) 393. — F. kamp- hoeveneri Boeck. Bot. Jahrb. 5 (1884) 505. — Iriha cinnamometorum O.K. Rev. Gen. Pl. 2 (1891) 753. Perennial with creeping, woody, |—1)4 mm thick rhizome. Stems solitary, approximate or somewhat remote, very slender, compressed-angular, smooth, 25-50 cm by 4-1 mm. Leaves at least half as long as the stem, erect, setaceous, often complicate, rigid, acute, antrorsely scabrid at the top, 4—)4 mm wide; ligule absent; lower sheaths horny. Inflorescence com- pound or decompound, loose, with many to numerous spikelets, 3-6 cm long. Involucral bracts 1—2, erect, much shorter to slightly longer than the inflorescence, the lowest up to 4cm. Primary rays 5-8, filiform, erect or obliquely erect, smooth. Spikelets solitary, oblong or oblong-linear, strongly compressed, acute, few-flowered, 4—5 by | mm; rachilla winged. G/umes distichous, thinly membranous, erect, ovate-lanceo- late, acute, muticous or apiculate, sharply keeled, with 3-nerved keel and nerveless, densely reddish gland-dotted sides, 234-4 by 14-2 mm. Stamens 3; anthers linear, (1—)1’4—2 mm. Sty/e slender, glabrous 566 except for the pyramidal, shortly hairy base, 3— 314 mm; stigmas 3, much shorter than the style. Nut trigonous with somewhat convex sides, obovoid or oblong-obovoid, shortly stipitate, umbonulate, ver- ruculose, stramineous to brownish, finely trans- versely lineolate by the oblong-linear epidermal cells, —Ao by A-Ao mm. /agao® cee Asia (Ceylon, India, Thailand, Indo-China) to S. China and tropical Australia; in Malesia: N. Sumatra (E. Coast Res.), Philippines (Luzon: Burgos), New Guinea (W. New Guinea: Cycloop Mts; NE. New Guinea: Sepik Distr.; Papua: Wassi Kussa R., Fly R., Mt Lawes near Port Moresby). Ecol. In swamps, savannah forests, on wet flats, at low altitudes, locally often abundant. Notes. It was already pointed out by S. T. BLAKE, /.c., that the Australian F. cyperoides R.BR. and the Asian F. cinnamometorum are not specifically distinct. From the type collection (KAMPHOEVENER 2485 from Teressa I. in KIEL) and BOECKELER’s accurate description it is evident that CLARKE wrongly referred F. kamphoeveneri BOECK. to the synonymy of F. fusca (NEES) CLARKE. 34. Fimbristylis adenolepis KERN, Blumea 8 (1955) 123, f. 3; ibid. 15 (1967) 433; in Back. & Bakh. f. Fl. Java 3 (1968) 468. Delicate, glabrous annual with fibrous roots. Stems tufted, setaceous, angular, smooth, 5-12 cm by 4-4 mm. Leaves much shorter than the stems, filiform, weak, flat, rather obtuse, abruptly acuminate, smooth, 14 mm wide; ligule absent. Inflorescence simple or almost so, loose, with (1—)3—5 spikelets, up to 2cm long and wide. Involucral bracts 2-3, shorter than the inflorescence, erect, the lowest Y-lem. Rays 1-3, erecto-patent, compressed, smooth, up to 12 mm. Spikelets solitary, lanceolate, strongly compressed, acute, few-flowered, 3—5 by 1 mm; rachilla narrowly winged. G/umes distichous, thinly membranous, erect, elliptic-ovate, muticous, densely reddish gland-dotted, 14-2 by 1mm. Stamen 1; anther oblong, c. 4 mm. Style triquetrous, pyramidally thickened at the base, glabrous, 1- 1/4 mm; stigmas 3, about half as long as the style. Nut trigonous, oblong-obovoid, subsessile, minutely umbonulate, verruculose, stramineous, transversely lineolate by the oblong-linear epidermal cells, 34— Yoyo mm. | | Distr. SE. and Peninsular Thailand, Indo-China (Cochinchina, Tonkin, Annam);in Malesia: Kangean I. (near Ardjasa). Ecol. In Kangean I. in a moist grass-field, abun- dant, at 25 m altitude. 35. Fimbristylis fuscoides CLARKE in Ostenf. Bull. Herb. Boiss. II, 5 (1905) 719: Kew Bull. add. ser. 8 (1908) 25; Camus, Fl. Gén. I.-C. 7 (1912) 124 p.p. (specim. siam.); RipL. Fl. Mal. Pen. 5 (1925) 160; KERN, Blumea 8 (1955) 125; ibid. 15 (1967) 433. — F. angustifolia RIDL. J. Str. Br. R. As. Soc. n. 59 (1911) 223. — F. erythradenia Camus, Not. Syst. 1 (1910) 247; Fl. Gén. I.-C. 7 (1912) 113. Perennial. Stems densely tufted, setaceous, smooth, 10-20 cm by 4-)4 mm. Leaves 4-)4 as long as the stems, erect, rigid, very narrow, flat, acute, glabrous, scabrid at the top, greyish or FLORA MALESIANA [ser. I, vole #7 glaucous, 14-74 mm_ wide; ligule absent. Inflo- rescence simple or compound, loose, with 4-18 spikelets, 2-4 cm long. Involucral bracts 2—3, very short, erect, dilated at the base, up to 1 cm. Primary rays 2-4, filiform, compressed, smooth, 1-2 cm. Spikelets solitary, lanceolate, strongly compressed, acute, few-flowered, 4-6 by 1-144 mm. Glumes distichous, membranous, erect, lanceolate, acutish, keeled, fuscous, densely reddish gland-dotted, with 3-nerved keel and nerveless sides, 3-334 by c. 14 mm; rachilla winged. Stamens 3; anthers linear, the anther-cells 1-1/4 mm, the connective produced into a subulate, smooth, 14 mm long appendage. Style triquetrous, glabrous except for the hairy, pyramidally thickened base, 2/4-3/4 mm; stigmas 3, much shorter than the style. Nut trigonous with somewhat convex sides, obovoid, shortly stipitate, obtuse, not umbonate, smooth, whitish, 4-24 by 24-)4 mm; epidermal cells minute, isodiametric. Distr. Thailand, Cochinchina, Cambodia; in Malesia only in the Western part: Malay Peninsula (Setul; Perlis: Bukit Ketri heath), Billiton, N. Borneo (Distr. Papar, Kimanis F.R.; Labuan). Ecol. Sandy fields, sandy places in heaths, at low altitudes. Vern. Rumput djani, Billiton. 36. Fimbristylis vanoverberghii KUk. Pfl. R., Heft 101 (1936) 631; KERN, Blumea 8 (1955) 127, f. 4; ibid. 15 (1967) 434. — Cladium cyperoides MERR. Philip. J. Sc. 7 (1912) Bot. 74, non F. cyperoides R.BR. 1810. — Mariscus fallax FERNALD, Rhodora 25 (1923) 53, non CHERMEZ. 1919. — F. fusca (non CLARKE) MERR. En. Philip. 1 (1923) 123 p.p. — F. nigrobrunnea (non THWaAITES) KUK. in Fedde, Rep. 51 (1942) 192. — F. fusca var. hispidissima KUK. Mitt. Thur. Bot. Ver. N.F.50(1943) 11. — Machaerinacyperoides KOYAMA, Bot. Mag. Tokyo 69 (1956) 63. Perennial with woody, shortly creeping-ascending rhizome covered with ovate, striate, fuscous sheaths. Stems solitary or somewhat tufted, obtusely trigonous, slightly compressed, glabrous or pilose, smooth, leafy at the base, 20-30 cm by )4-34 mm. Leaves much shorter than the stems, erect or slightly curved, flat or with inrolled margins, long-acuminate, with bristle-like, readily caducous tip, more or less hispid especially beneath, or glabrescent, 14-1 mm wide; ligule absent; outer sheaths coriaceous, shining fuscous or purplish, glabrous or pilose. Inflorescence simple or subcompound, obtriangular, rather dense, with (1—)5—15 spikelets, up to 4 cm long. Involucral bracts 3—4, erect, dilated at the base, hispid at least at the base, the lowest up to 2/4 cm. Rays up to 4, obliquely patent, compressed, glabrous and smooth, 1-2cm. Spikelets solitary or partly 2—3 together, oblong-ovate, strongly compressed, acute, 6—9- flowered, fuscous, 5—8 by 2-274 mm; rachilla broadly winged. G/umes distichous, chartaceous, obliquely erect, triangular-ovate, acuminate, apiculate, sharply keeled, 1-nerved, scabrid with short white hairs but finally glabrescent, whitish hyaline-margined, dull fuscous, 3—4 by 2—3 mm. Stamens 3; anthers linear, 1'4-2 mm. Style triquetrous, slightly incrassate at the base, glabrous, 2-3 mm; stigmas 3, about 4 as long as the style. Nut obtusely trigonous, obovoid, shortly stipitate, minutely umbonulate, sparsely verruculose, whitish or stramineous, 1-1/4) by 4-% mm; epidermal cells minute, isodiametric. 1974] Cy PERACEAE (Kern) 567 Distr. Malesia: N. Sumatra (between Sibosar and Parsoburan; W. Batuhuda; between Mt Piso-Piso and Toba Lake); Borneo (Brunei: Temburong R.); Philippines (Luzon: Bontoc Subprov., Bauco); New Guinea (W. New Guinea: Kebar; NE. New Guinea: Morobe Distr., Sattelberg). Ecol. In moist grasslands, on hill-sides, river- banks, 540—1320 m. 37. Fimbristylis eragrostis (NEES) HANCE, J. Linn. Soc. Bot. 13 (1873) 132'; KUx. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 9; S. T. Blake, J. Arn. Arb. 35 (1954) 215; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 117; KERN, Blumea 15 (1967) 434. — Abildgaardia era- grostis Nees & Mey. ex NEES in Wight, Contr. (1834) 95; KuNTH, En. 2 (1837) 249; Sreup. Syn. 2 (1855) 72; Boeck. Linnaea 37 (1871) 55.—F. nigrobrunnea THWaAITES, En. Pl. Zeyl. (1864) 434; CLARKE, FI. Br. India 6 (1893) 648; J. Linn. Soc. Bot. 34 (1898) 70; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 97; Camus, Fl. Gén. I.-C. 7 (1912) 120; Ript. Fl. Mal. Pen. 5 (1925) 159. — F. subtetra- stachya Boeck. Linnaea 37 (1871) 50. — F. pycno- stachya HANCE, J. Bot. Lond. 15 (1877) 338. — Iriha eragrostis O.K. Rev. Gen. Pl. 2 (1891) 753. — Iriha subtetrastachya O.K. I.c. — F. lepidota Camus, Not. Syst. 1 (1910) 247; Fl. Gén. I.-C. 7 (1912) 121, f. 14, 3—6. — F. tortispica TURR. Kew Bull. 1911 (1911) 348. — F. schlechteri KUK. Bot. Jahrb. 59 (1924) 50. Glabrous perennial with short woody rhizome in old specimens clothed with the remains of decayed leaf-sheaths. Stems solitary or somewhat tufted, rigid, acutely 4—5-angled, sulcate, smooth, or scabrid at the top, 30-70cm by 1—2(—3)mm, the base clothed with 2—3 tubular, 4—S(—15) cm long sheaths with short erect blades. Basal /eaves numerous, much shorter than the stems, subfalcate, firm, flat, exactly linear, with obtuse, rounded, apiculate top, spinulose-scabrid on the margins in the upper part, (2—)3—5 mm wide; ligule absent. Inflorescence com- pound or decompound, with several to numerous spikelets, 3-6 cm long, sometimes subcapitate. In- volucral bracts 2—4, very short, erect, dilated at the base, the lowest up to 1/4 cm. Primary rays (0—)3-6, erecto-patent to recurved, compressed, smooth, up to 4cm. Spikelets solitary or partly clustered, lanceolate, compressed or subterete, often contorted, acute, 8—16-flowered, stramineous to dark brown, 6—10(—15) by 2—4mm. Rachilla winged. Glumes distichous or subspiral, chartaceous, shining, gla- brous, broadly ovate to oblong-ovate, rather obtuse, distinctly mucronulate, keeled, 3-5 by 2/4-3 mm. Stamens 3; anthers linear, 1/4-2mm. Spe tri- quetrous, pyramidally thickened at the base, gla- brous or sparsely ciliate at the top, 2-3 mm; stigmas 3, about as long as the style. Nut trigonous, obovoid or broadly obovoid, shortly stipitate, umbonulate, verruculose, whitish or stramineous, 44-1 by 4-1 mm; epidermal cells minute, isodiametric. Distr. S. Asia, from Ceylon and India eastwards to S. China, Hainan and Formosa, southwards to tropical Australia (Queensland); in Malesia: Malay Peninsula (Kedah, Trengganu), Lesser Sunda Is. ‘ HANCE wrongly ascribes the authority of the basionym Abildgaardia eragrostis to VAHL. (Port. Timor), Aru Is. (P. Trangan), New Guinea (NE. New Guinea, Papua). Ecol. In savannahs and savannah-forests, forest- clearings, at low altitudes. Notes. In the wide sense here accepted a most polymorphous species. F. eragrostis s. str., F. nigrobrunnea and F. pycnostachya (= F. lepidota) look very different, but I am unable to trace dividing lines, as they are connected by numerous inter- mediates. Typical F. nigrobrunnea (= F. subtetrastachya) is characterized by the often clustered spikelets and the dark, imperfectly spirally arranged glumes. In its typical form unknown from Malesia. F. pycnostachya still more deviates from typical F. eragrostis by its thick rhizomes, robust stems, castaneous leaf-sheaths, coriaceous leaves, sub- capitate inflorescences, and subspiral dark glumes. In Malesia it is only known from Kedah (Kedah Peak). F. tortispica from N. Thailand and F. schlechteri from NE. New Guinea are in my opinion typical F. eragrostis, with solitary, light coloured spikelets and distichous glumes. 38. Fimbristylis fusca (NEES) CLARKE, Fl. Br. Ind. 6 (1893) 649; Philip. J. Sc. 2 (1907) Bot. 98; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 97 p.p.; Koorb. Exk. Fl. Java 1 (1911) 200; ibid. 4, Atlas (1922) f. 262: Camus, Fl. Gén. I.-C. 7 (1912) 123; Merr. En. Philip. 1 (1923) 123 p.p. (excl. syn. Cladium cyperoides Me_rR.); RIDL. Fl. Mal. Pen. 5 (1925) 160 p.p.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 18; S. T. BLAKE, J. Arn. Arb. 35 (1954) 211; KERN, Blumea 8 (1955) 125; ibid. 15 (1967) 435; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 118; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 468. — Gussonea cyper- oides PRESL, Rel. Haenk. 1 (1828) 183, t. 33, non F. cyperoides R.BR. 1810. — Gussonea pauciflora BRONGN. in Duperr. Voy. Bot. 2 (1829) 171, t. 34B, non F. pauciflora R.Br. 1810. — Abildgaardia fusca NEEs in Wight, Contr. (1834) 95; KUNTH, En. 2 (1837) 249; StEuD. Syn. 2 (1855) 72; Boeck. Linnaea 37 (1871) 54, excl. var. longifolia Boeck. — Abildgaardia cyperoides NEES in Wight, Contr. (1834) 95; KUNTH, En. 2 (1837) 249; Sreup. Syn. 2 (1855) 72. — Abild- gaardia pauciflora KUNTH, En. 2 (1837) 249; STEuD. Syn. 2 (1855) 73; Mia. FI. Ind. Bat. 3 (1856) 298. — Rhynchospora ? anomala Steup. [in Zoll. Syst. Verz. 1 (1854) 61, nom. nud.] Syn. 2 (1855) 149: Mia. Fl. Ind. Bat. 3 (1856) 337. — Iriha fusca O.K. Rev. Gen. Pl. 2 (1891) 753. — F. subfusca Camus, Not. Syst.1 (1910) 248: Pl: Gen: E-C: 7 (1912) 1232——F. rigidifolia RiwL. J. Str. Br. R. As. Soc. n. 59 (1911) 223: Fl. Mal. Pen. 5 (1925) 159. — F. stenochlaena KOK. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 11; S. T. Bake, J. Arn. Arb. 35 (1954) 220. — Fig. 42. Perennial with short, woody rhizome clothed with the remains of decayed leaf-sheaths. Stems tufted, rigid, angular-sulcate, often scabrid-pilose just below the inflorescence, otherwise glabrous and smooth, 20-50 cm by 4-1 mm. Leaves much shorter than the stems (often scarcely 14 as long), basal except for one somewhat higher on the stem, rather stiff, flat, exactly linear, abruptly pointed, glabrous, or pubes- cent especially beneath, scabrid at the top, (1—)2— 4 mm wide; ligule absent. Inflorescence compound to 568 FLORA MALESIANA [ser. I, vol. 73 Fig. 42. Fimbristylis fusca (NEES) CLARKE. a. Habit, x 14, a’. leaf tip, x 7, b. spikelet, x 5, c. glume, d. anther, e. nut with style and stigmas, x 10 (a—e BEUMEE A597). 1974] Cy PERACEAE (Kern) 569 supradecompound, loose, with several to very numer- ous spikelets, up to 10cm long. Involucral bracts 3-5, very short, glabrous or pilose, dilated at the base, the lowest rarely up to 3cm. Primary rays suberect, rigid, glabrous or pilose, up to 7cm. Spikelets solitary, lanceolate, strongly compressed, acute, 3—10-flowered, 5—10 by 2-2/4 mm; rachilla broadly winged. G/umes distichous, subchartaceous, obliquely erect, lanceolate, gradually acuminate, prominently keeled, l|-nerved, scabrid by very short, stiffish hairs, fulvous, with broad, glabrous, whitish hyaline margins, 4-6/4 by 2-2!4 mm. Stamens 3; anthers linear, 1)4-2 mm. Style very slender, tri- quetrous, pyramidally thickened at the base, gla- brous, 4-6 mm; stigmas 3, much shorter than the style. Nut trigonous with convex sides, obovoid, cuneate at the base, shortly stipitate, umbonulate, densely verruculose, whitish to brownish, ?4,—1 by %-% mm; epidermal cells isodiametric. Distr. From Nepal and India through Thailand and Indo-China to S. China and Japan (Shikoku, Kyushu); in Malesia: Sumatra, Malay Peninsula (Setul, Kelantan, Singapore), W. Java, Borneo, Philippines (Palawan, Luzon) Celebes, Moluccas (Ambon), and New Guinea. Ecol. In open, rather dry to wet grasslands, in Celebes collected on dry gravel-hills; in Java only known from low altitudes (40-100 m), in N. Sumatra and the Philippines ascending to 1300 m, in New Guinea to 1600 m. Notes. F. stenochlaena KO«. was based on BRASS 7840 from Papua, Lake Daviumbu. The specimens of this collection are stout and in habit similar to F. eragrostis, from which species they can readily be distinguished by the hairy, muticous, acute glumes, the much longer style, etc. According to KUKENTHAL F. stenochlaena differs from F. fusca by the broader leaves (2 mm) with light-coloured sheaths, the twice as long spikelets, the regularly distichous, long- acuminate, narrower glumes, and the many-flowered spikelets with all the flowers bisexual. S. T. BLAKE l.c., p. 223 gives the following key characters: F. fusca: glumes more than half as wide as long, 3—4 mm long; style about 3 mm long; culms 5- ribbed; leaves 2 mm wide; spikelets 114-2 mm wide. F. stenochlaena: glumes about half as wide as long, 4—5 mm long; style 4 mm long; culms many-ribbed; leaves 1/4—4 mm wide; spikelets 2~3 mm wide. Like in most Fimbristy lis spp. the width of the leaves and the number of flowers in the spikelets varies considerably. In the type-collection of F. fusca (WALLICH 3530) the glumes are 414-5!4 mm long, the style c. 5 mm. The flowers of F. stenochlaena are triandrous (like in F. fusca), not diandrous as KUKENTHAL wrongly indicates. In my opinion F. stenochlaena cannot be treated as a distinct species. Also in F. subfusca Camus and F. rigidifolia RIDL. the floral characters are those of F. fusca, so I take them for broad-leaved forms of this species. 39. Fimbristylis fulvescens (THWAITES) THWAITES, En. Pl. Zeyl. (1864) 434; CLarkg, FI. Br. Ind. 6 (1893) 650; J. Linn. Soc. Bot. 34 (1898) 72; Hook. /. in Trimen, Handb. a Ceylon 5 (1900) 62; KERN, Blumea 8 (1955) 127, ibid. 15 (1967) 435. — Abild- gaardia fulvescens sickens, En. Pl. Zeyl. (1864) 347. — Abildgaardia fusca var. BOECK. Linnaea 37 (1871) 55. — Fig. 43. Perennial (always ?). Stems densely tufted, very slender, sharply 4~—S-angled, ribbed, smooth, (20—)50-60 cm by c. 14 mm, the base clothed with 2—3, up to 7 cm long sheaths usually with very short blades. Basal /eaves erect, flat, acute, glabrous, scabrid on the margins at the top, up to 35 cm long, 1-2 mm wide; ligule absent. Inflorescence simple (very rarely one of the rays with a short secondary ray), loose, with 3-7 spikelets, 114-3cm long, 2—4 cm wide. Involucral bracts 3—5, erect or some- what excurved, setacous, dilated at the base, the lowest 4-1 cm. Rays 2-5, slender, obliquely patent to horizontally spreading, compressed, smooth, 5-15 mm. Spikelets solitary, lanceolate or oblong- lanceolate, strongly compressed, acute, 6-—10- flowered, 5—10 by 2—4 mm; rachilla broadly winged. Glumes exactly distichous, subchartaceous, obliquely erect, ovate-lanceolate, acute, 1-nerved, sharply keeled, pubescent by soft hairs, not or ate hyaline- margined, ciliolate in the upper part, 4/4—5 by 2/4-3 mm. Stamens 3; anthers linear, 134 mm. Style very slender, triquetrous, pyramidally thickened at the base, glabrous, 4 mm; stigmas 3, much shorter than the style. Nut trigonous, oblong-obovoid, shortly stipitate, umbonulate, sparsely verruculose, light brown, 1/4-1)4 by )4, mm; epidermal cells isodiametric. Distr. Insufficiently known. Ceylon; in Malesia: Malay Peninsula (Government Hill on P. Penang). Ecol. At 750 m altitude. Note. Closely related to F. fusca, from which it is possibly not specifically distinct. In Malesia only collected by KUNSTLER in 1881 and Curtis in 1889. longifolia 40. Fimbristylis fimbristyloides (F.v.M.) DRucE, Rep. Bot. Exch. Cl. Br. Isl. 1916 (1917) 623; S. T. BLAKE, J. Arn. Arb. 35 (1954) 221; KERN, Blumea 8 (1955) 129; ibid. 15 (1967) 437; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 117; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 468. — Abildgaardia fimbristyloides F.v.M. Fragm. 8 (1874) 273.—F. dallachyi F.v.M. ex BENTH. FI. Austr. 7 (1878) 309; BAcK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 18. — Iriha fimbristy- loides O.K. Rev. Gen. Pl. 2 (1891) 753. — F. lacei Turr. Kew Bull. 1911 (1911) 348. — F. disticha var. dallachyi CLARKE ex Domin, Bibl. Bot., Heft 85 (1915) 454. — F. disticha (non Boeck.) KUK. Bot. Jahrb. 59 (1925) 50.—F. straminea (non TurRR.) Ouwt, J. Jap. Bot. 14 (1938) 574; Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 60.— F. nanofusca TANG & WANG, FI. Reip. Pop. Sinic. 11 (1961) 229. Dwarf annual with fibrous roots. Stems tufted, ribbed, antrorsely scabrid-hispid at least at the top, leafy at the base, 7-12cm by 4-4 mm. Leaves much shorter than the stems, strongly falcate in the same direction (to the right or to the left), flat, abruptly acuminate, apiculate, antrorsely spinulose- scabrid on the margins, (l-)2mm _ wide; ligule absent. Inflorescence simple or subcompound, loose, with 3-10 spikelets, up to 4cm long. Involucral bracts 2—3, obliquely erect, much shorter than the inflorescence, up to 15(—25) mm. Primary rays 3—S, obliquely erect, scabrid-pilose, up to 3 cm; secondary rays when present very short. Spikelets solitary, lanceolate, strongly compressed, acute, 6—12- flowered, pale to dark brown, 4—10 by(1—)1'4-2 mm; —— — ; = 1974] rachilla winged. G/umes distichous, membranous, scabrid by minute hairs, ovate-lanceolate, acute, often mucronulate, prominently keeled, with ciliolate midnerve, nerveless sides, and broad hyaline margins, (2-)274-3 by c. 134 mm. Stamens (1—)2(—3); anthers ine ’%-'4 mm. Style triquetrous, Pyramidally thickened at the base, glabrous, (1-)1/4-174 mm; stigmas 3, much shorter than the style. Nut trigonous with convex sides, pyriform, umbonulate, abruptly truncate at the base which is conspicuously broader than the short but distinct stipe, smooth or ver- ruculose, whitish, 74-2“ by 14-74 mm; epidermal cells isodiametric. Distr. Burma, E. Thailand, S. China, Ryu Kyu Is., S. Korea, tropical Australia (Queensland); in Malesia: N. Sumatra, W. and Central Java, Madura, Kangean Arch., Lesser Sunda Is. (Flores), N. Borneo, Celebes, New Guinea. Ecol. In open wet grassy places, Imperata-fields, on hill-sides and in valleys, 25—825 m. 41. Fimbristylis intonsa S. T. BLAKE, J. Arn. Arb. 35 (1954) 221, f. 2; KERN, Blumea 8 (1955) 130; ibid. 15 (1967) 438. — F. disticha var. kurzii CLARKE, FI. Br. Ind. 6 (1893) 651. Annual with fibrous roots. Stems tufted, setaceous, acutely 5-ribbed, scabrid at the top, 5—15(—25) cm by c. ‘4mm, the base clothed with 1-2 tubular, short-bladed sheaths. Leaves much shorter than the stems, rather firm, falcate, flat, obtuse, apiculate, antrorsely spinulose-scabrid on the margins in the upper part, 1/4-2 mm wide; ligule absent. Inflo- rescence simple or subcompound, loose, with (1—)3— 8(—15) spikelets, 1—5 cm long. Involucral bracts 2—3, erect, spinulose-scabrid, the eye, often subfolia- ceous, up to 12 mm. Primary rays 3-7, scabrid, up to 3cm, the longer ones sometimes with 74-1 cm long secondary rays. Spikelets solitary, ovate-oblong, strongly compressed, sometimes contorted, acute, densely many-flowered, brown, (3—)5—10 by 1/4- 134 mm; rachilla winged. Glumes distichous, very small, subchartaceous, obliquely erect, very broadly ovate, obtuse, apiculate or minutely mucronulate, l-nerved, with sharp keel and ciliolate margins, densely pubescent by very short hairs almost all over, 134-2 by 114-134 mm. Stamens 3; anthers oblong- linear, c. %mm. Style triquetrous, pyramidally thickened at the base, glabrous, 1-1/4 mm; stigmas 3, much shorter than the style. Nut obtusely tri- gonous, with convex sides, obovoid or oblong- obovoid, shortly stipitate, umbonulate, densely verruculose, shining white, 74-4% by uy mm; epidermal cells isodiametric. Distr. Bengal; in Malesia: Sumatra (E. Coast Res., Tapanuli), New Guinea (W. New Guinea: Kebar Valley; Papua: W. Div., Lake Daviumbu, Middle Fly R.). Ecol. On wet grassy plains, scattered among tall grasses, at low and medium altitudes; in Sumatra up to 1000 m. 42. Fimbristylis malayana OHW1, Blumea 8 (1955) 96, f. 1; KERN, Blumea 15 (1967) 438. — F. fuscoides (non CLARKE) HENDeRS. J. Mal. Br. R. As. Soc. 17 (1939) 86. Perennial, glabrous except for the glumes, light green. Stems densely tufted, rigid, obtusely 3—4- angled or subterete, slightly compressed, smooth, 20-40 cm by /4- 1 mm, the base clothed with 1-2 CyPERACEAE (Kern) 571 bladeless or short-bladed, 1—-2cm long sheaths. Leaves }A—'% as long as the stems, falcate, flat, obtuse, apiculate, scabrid on the margins in the upper part, 114-2 mm wide; ligule absent. Inflorescence simple or subcompound, narrow, rather dense, with 2-7 spikelets, 3—4 cm long. Involucral bracts very short, often glume-like, mucronate, scarious- margined at the dilated base, the lower 1-2 some- times with a short blade up to 1/4 cm. Rays 2-4, suberect, rigid, slightly compressed, smooth, 1— 2(—3) cm. Spikelets solitary, lanceolate or linear- lanceolate, strongly compressed, acute, often contorted, 8-10 by 2-2)4 mm; rachilla broadly winged. Glumes distichous, chartaceous, suberect, oblong-ovate, long-acuminate, sharply keeled, with nerveless sides and whitish hyaline margins, minutely puberulous at the top, dull brown, 334-4 by 2 mm. Stamens 3; anthers linear, 1/4-17)4 mm. Style very slender, triquetrous, slightly pyramidally thickened at the base, glabrous, 2/4-3 mm; stigmas 3, much shorter than the style. Nut trigonous with somewhat convex sides, tricostate, obovoid or broadly obovoid, shortly stipitate, not or hardl ly umbonulate, smooth, stramineous to brown, 74-% by 74- 1% mm: epi- dermal cells roundish or elliptic. Distr. Malesia: Malay Peninsula (P. Langkawi). Ecol. On limestone screes in open places, espe- cially where wet, c. 60 m. 43. Fimbristylis disticha Boeck. Linnaea 38 (1874) 393: CLARKE, FI. Br. Ind. 6 (1893) 651, excl. var. kurzii CLARKE; KERN, Blumea 8 (1955) 130: ibid. 15 (1967) 438. — Iriha disticha O.K. Rev. Gen. Pl. 2 (1891) 753. — F. fuscoides (non CLARKE) CAMus, Fl. Gén. I.-C. 7 (1912) 124 p.p. (specim. cochinch.). Glabrous annual with fibrous roots. Stems solitary or tufted, acutely 4—5-ribbed, smooth, 10—35 cm by Y4-'% mm, the base clothed with 1—2 short-bladed sheaths. Leaves much shorter than the stems, rather rigid, falcate, flat, abruptly acuminate, scabrid on the margins in the upper part, 1-2 mm wide; ligule absent. Inflorescence subcompound to decompound, very loose, with several to numerous spikelets, 4—6 cm long. Involucral bracts 3—4, setaceous, often excurved, the lowest 4-214 cm. Primary rays 3-6, erecto-patent, filiform, angular, smooth, up to 4 cm. Spikelets solitary, oblong-lanceolate, strongl com- pressed, acute, ferrugineous, 4-6 by c. 1/4 mm; rachilla broadly winged. Glumes exactly distichous, very small, thinly membranous, glabrous, ovate, rather obtuse, minutely apiculate just below the apex, acutely keeled, with curved keel, distinct midnerve and nerveless sides, 1)/4—2(- 214) by c. Amm. Stamens 2(—3); anthers oblong, }4(-1) mm. Style triquetrous, pyramidally thickened at the base, glabrous, 1—1)4(—2) mm; stigmas 3, much shorter than the style. Nut trigonous, narrowly obovoid or obovoid, cuneate at the base, shortly stipitate, minutely umbonulate, more or less verruculose, whitish or stramineous, %—]{, by )4—/4 mm; epi- dermal cells isodiametric to suboblong. Distr. Burma, Central and Peninsular Thailand, Cochinchina, Tenasserim and Andamans, Teressa, Mergui, S. China; in Malesia: N. Sumatra. Ecol. In Sumatra in open grassy plains, 1200-— 1400 m. Note. Rather variable. The Chinese plants differ from the type collection by taller stems, longer glumes S72 FLORA MALESIANA [ser. I, vol. 7° (2?4 mm), anthers (1 mm), and styles (2mm), and by the obovoid, densely verruculose nuts. Several other collections connect the two extremes. 44. Fimbristylis calcicola KERN, Blumea 8 (1955) 129, f. 5, ibid. 15 (1967) 439. Very slender annual with fibrous roots, glabrous except for the glumes, pale green. Stems very densely tufted, setaceous, sharply 4-ribbed, smooth, 10—20 cm by 4-)K mm, the base clothed with 1—2 tubular short-bladed sheaths. Basal leaves much shorter than the stems, rather rigid, falcate, rather abruptly acuminate, 14-1 mm_ wide; ligule absent. Inflo- rescence simple or subcompound, loose, with 3—9 spikelets, up to 4cm long. Involucral bracts 3-4, erect with more or less excurved tip, setaceous, dilated at the base, the lowest 4-1 cm. Rays 2-S, filiform, obliquely patent, often upcurved, com- pressed, smooth, 1-274 cm. Spikelets solitary, lan- ceolate to almost linear, strongly compressed, acute, densely many-flowered, 3—7 (ultimately up to 12) by 1/4 mm; rachilla winged. Glumes exactly distichous, membranous, obliquely erect, ovate, acute, muticous or minutely apiculate, 1-nerved, sharply keeled, with straight keel, densely pubescent, ferrugineous, paler towards the margins, 14-134 by 1-1) mm. Stamens 2; anthers oblong-linear, c. 4 mm. Style slender, triquetrous, pyramidally thickened at the base, glabrous, 1 mm; stigmas 3, much shorter than the style. Nut very obtusely trigonous, oblong- obovoid to ellipsoid, shortly stipitate, minutely umbonulate, not verruculose, finely longitudinally striate and transversely lineolate by the transversely linear epidermal cells in 3—4 vertical rows on each face, at first whitish, finally yellowish brown; c. 74 by ‘A mm. Distr. Malesia: Malay Peninsula (Kedah: P. Langkawi). Ecol. On limestone screes. 10. Section Dichelostylis BENTH. FI. Austr. 7 (1878) 299. — Fimbristylis sect. Ferrugineae OHWI, J. Jap. Bot. 14 (1938) 573, p.p. — Fimbristylis ser. Ferrugineae OHW1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55, p.p. Type species: F. ferruginea (L.) VAHL (Scirpus ferrugineus L.). 45. Fimbristylis ferruginea (L.) VAHL, En. 2 (1806) 291; KuNTH, En. 2 (1837) 236; STeUD. Syn. 2 (1855) 118; Boeck. Linnaea 37 (1871) 16; BENTH. FI. Austr. 7 (1878) 312; CLarRKE, FI. Br. Ind. 6 (1893) 638; K. Sco. & Laut. FI. Schutzgeb. (1900) 197; CLARKE, Philip. J. Sc. 2 (1907) Bot. 94; Rip. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 92; CLARKE, Ill. Cyp. (1909) t. 42 f. 9-10; Koorp. Exk. Fl. Java 1 (1911) 199; ibid. 4, Atlas (1922) f. 255; Camus, Fl. Gén. I.-C. 7 (1912) 108, f. 16,9; Mere. En. Philip. 1 (1923) 123; KOK. Bot. Jahrb. 59 (1924) 48; 69 (1938) 258; RIDL. Fl. Mal. Pen. 5 (1925) 576; Back. Onkr. Suiker. (1928) 161, t. 168; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 23; S. T. BLAKeE, J. Arn. Arb. 35 (1954) 211; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 465. — Scirpus ferrugineus LINNE, Sp. Pl. (1753) 50. — Scirpus arvensis Retz. Obs. 4 (1786) 11. — F. arvensis VAHL, En. 2 (1806) 291.— F. marginata LABILL. Sert. Austr. Caled. 2 (1825) t. 16 f. 1. — F. trispicata STEUD. Syn. 2 (1855) 107; Mig. Fl. Ind. Bat. 3 (1856) 317. — F. cyrtophylla Mia. l.c. 325. — Triha ferruginea O.K. Rev. Gen. Pl. 2 (1891) 752. — F. polytrichoides (non R.BR.) RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 90 p.p.; Fl. Mal. Pen. 5 (1925) 154 p.p. — F. longispica (non STEUD.) CAMus, FI. Gén. LEn7 (1912) M07: Perennial with shortly creeping, woody rhizome. Stems approximate, forming tufts, rigid, striate, compressed, smooth, greyish green, 20-80 cm by 1/4-3 mm. Cauline leaves much shorter than the stems, rigid, erect, conduplicate or triquetrous, scabrid at the top, 2-10 cm by 4-14 mm, those of the sterile shoots much louger; lower sheaths blade- less, coriaceous, shining brown to castaneous, upper ones ferrugineous-membranous in front, ciliolate at the mouth; ligule a dense fringe of short hairs. Inflorescence simple or subcompound, usually con- tracted, with (1—)5—10(—25) spikelets, 3-5 cm long. Involucral bracts 2—3, the lowest suberect, shorter to slightly longer than the inflorescence, scarious- margined at the dilated base. Primary rays com- pressed, smooth, up to 2/4 cm. Spikelets solitary, ovoid to oblong-ovoid, terete, acute, densely many- flowered, dull brown, 5-20 by 3-4 mm; rachilla narrowly winged. Glumes spiral, subchartaceous, ovate to oblong, obtuse, scarcely keeled, apiculate, with green, l-nerved keel and nerveless sides, cili- olate at the upper edge, appressed-puberulous in the apical part, rarely subglabrous, 3-44 by 244-3 mm. Stamens 3; anthers oblong-linear, 1-1/4 mm. Style flat, slightly dilated at the base, densely ciliate, 2—3 by 7/4 mm; stigmas 2, shorter than the style. Nut biconvex, strongly compressed, obovate or oblong- obovate, shortly stipitate, obsoletely umbonulate, smooth, stramineous to fuscous, 1-1/4 by 4-1 mm; epidermal cells isodiametric or almost so. Distr. Pantropical, probably common through- out Malesia (but still unknown from the Lesser Sunda Is.). Ecol. In sunny wet localities with clayish soil, especially those subject to the influence of brackish water, rarely more inland near saline pools and mud-wells; locally often abundant; 0—100 m. Vern. Ba’ileu, Atjeh, rumput ruchut, Mal. Pen., sukét dot, sukét godokan, kodokan, J, kodoan, Bawean, gérinting, Karimundjawa, purum, W. Borneo. 46. Fimbristylis sieberiana KUNTH, En. 2 (1837) 237; STEUD. Syn. 2 (1855) 118; Mia. Fl. Ind. Bat. 3 (1856) 326; KERN, Blumea 8 (1955) 131. — F. ferruginea (non VAHL) DecNnE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 352; Descr. Herb. Timor. (1835) 1974] Cy PERACEAE (Kern) 573 34. — F. paucispicata F.v.M. Fragm. 1 (1859) 197. — F. ferruginea var. sieberiana BorcK. Linnaea 37 (1871) 17; CHERMEZ. in Humbert, Fl. Madag. fam. 29 (1937) 181. — F. ferruginea var. foliata BENTH. Fl. Austr. 7 (1878) 312, excl. syn. F. arvensis VAHL et F. tri- stachya R.BR. Closely related to F. ferruginea, from which it is distinguishable by the following characters: Cauline leaves well developed, less stiff, flat, up to 35cm by 114-2 mm; lower sheaths herbaceous, laminiferous, stramineous or ferrugineous; sheaths of the cauline leaves pilose, especially towards the top, finally more or less glabrescent. Lowest in- volucral bract as long as or usually distinctly over- topping the inflorescence, up to 10cm. Spikelets obtusish. Glumes very broadly ovate, usually dark castaneous, densely tomentose in the apical part, 3-4'% mm long and wide. Style broader, c. 74 mm wide. Nut broadly obovate or orbicular, distinctly stipitate (gynophore \Y%-2% mm), umbonulate, larger (1%-114 by 15-11% mm). Distr. Africa (Erythraea, S. Africa, Madagascar, Mauritius, Réunion), Asia (from Syria through Arabia and Persia to India), tropical Australia (Queensland); in Malesia very rare, only a few times collected in the Lesser Sunda Is.: Timor (LESCHENAULT; also recently collected near Tjamplong and Dili); Philippines (Mindanao, Cotabato Prov.: Buayan). Ecol. Probably less halophilous than F. ferruginea, but in Port. Timor in brackish marshes, at low altitude. Note. CLARKE, Fl. Cap. 7 (1898) 201 referred F. sieberiana to the synonymy of F. ferruginea, from which it is, however, clearly distinct. 47. Fimbristylis tristachya R.Br. Prod. (1810) 226; Kern, Blumea 8 (1955) 131; in Back. & Bakh. f. Fl. Java 3 (1968) 465, non THwaites, 1864. — F. marianna GAUDICH. in Freyc. Voy. Bot. (1826) 413, excl. varr.; KUNTH, En. 2 (1837) 236; STEUD. Syn. 2 (1855) 109; Kuk. Bot. Jahrb. 59 (1924) 5;S. T. BLAKE, J. Arn. Arb. 35 (1954) 212.— F. bispicata var. monostachya NEES in Hook. J. Bot. Kew Misc. 6 (1854) 29.— F. polymorpha BOECK. var. c BOECK. Linnaea 37 (1871) 16.—F. nutans (non VAHL) Naves, Nov. App. (1882) 307; VIDAL, Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 284. — F. maxima K. ScH. in K. Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 24, non K. Scu. & Laut. Fl. Schutzgeb. (1900) 196. — F. subbispicata (non NEES) CLARKE, FI. Br. Ind. 6 (1893) 634, quoad specim. Ind.; Philip. J. Sc. 2 (1907) Bot. 92, incl. var. caesia quoad specim. cit.; MERR. En. Philip. 1 (1923) 126. — F. annamica Camus, Not. Syst. 1 (1910) 245; Fl. Gén. I.-C. (1912) 95. — F. alleizettei CAMus, Not. Syst. 1 (1910) 291; Fl. Gén. I.-C. 7 (1912) 112.— F. marianna var. foenea KUK. in Fedde, Rep. 16 (1920) 432; Merr. En. Philip. 1 (1923) 124.— F. podocarpa (non NEEs) RIL. J. Str. Br. R. As. Soc. n. 59 (1911) 222; Fl. Mal. Pen. 5 (1925) 155. — F. schoenoides (non VAHL) BACK. Onkr. Suiker. (1928) 159 p.p.; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 23 p.p. Glabrous perennial with short rhizome. Stems tufted, rigid, sulcate-angular, compressed, smooth, or scabrid at the top, somewhat incrassate at the base, up to 65(—100) cm by 1-114 mm. Leaves about 4 as long as the stems, canaliculate, rigid, rather abruptly acuminate, scabrid on the margins in the upper part, greyish green or glaucous, 4-1/4 mm wide; sheaths brownish; ligule a dense fringe of short hairs. Inflorescence simple or subcompound, loose, with (1—)3—7(—11) spikelets. Involucral bracts 1-2, much shorter than the inflorescence, erect or suberect, scarious-margined at the dilated base, up to 3 cm (usually much shorter). Primary rays 0-5, com- pressed, smooth, up to 4cm. Spikelets solitary, ovate or ovate-lanceolate, terete, acute, densely many-flowered, (6—-)10—-25 by 4-6 mm; rachilla narrowly winged. Glumes spiral, chartaceous, ap- pressed, broadly ovate, obtuse, mucronulate, scarcely keeled, many-(up to 20-)nerved, fulvous-ferrugi- neous, 4—6 by 3-414 mm. Stamens 3; anthers linear, (114-)2-244 mm. Style flat, broad, slightly dilated at the base, densely long-ciliate in the upper half, 3-334 mm; stigmas 2, shorter than the style. Nut biconvex, obovate, distinctly stipitate (gynophore obconical, 4—)4 mm), not or hardly umbonulate, smooth, stramineous, (1-)14-1)4 by %-1 mm, obscurely reticulate by the isodiametric epidermal cells. Distr. From India to Micronesia and tropical Australia; widely distributed in Malesia: Sumatra (E. Coast Res.), Malay Peninsula (Setul, Perlis, Kedah, Selangor, Singapore), Java, Madura, Lesser Sunda Is. (Sumba, Port. Timor), N. Borneo (Labuan), Philippines (Palawan, Busuanga, Luzon, Guimara Is., Bohol, Mindanao), Celebes, New Guinea. Ecol. Open grassy fields, damp heaths, sometimes in teak-forests and rice-fields, usually below 200 m, in Sumba at 500 m, in W. New Guinea up to 540 m. Vern. Tihe-tihe, Sum. E.C., komes bu-u, Md. Note. F. subbispicata NEES & Mey. ex NEES from Japan, Korea, Ryu Kyu Is., Formosa and China, and F. pacifica Ouwi from Japan and Ryu Kyus are closely related to F. tristachya. Both were reduced to varieties of F. tristuchyva by KoyAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 114. 48. Fimbristylis schoenoides (RETZ.) VAHL, En. 2 (1806) 286; KunTH, En. 2 (1837) 222; STEuD. Syn. 2 (1855) 107; Mia. Fl. Ind. Bat. 3 (1856) 315; Boeck. Linnaea 37 (1871) 5, excl var. B; CLARKE, Fl. Br. Ind. 6 (1893) 634; Philip. J. Sc. 2 (1907) Bot. 92; RIDL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 90; Koorp. Exk. Fl. Java 1 (1911) 199; Camus, Fl. Gen. I.-C. 7 (1912) 99, f. 16, 1-2; MERR. En. Philip. 1 (1923) 125: Ripe. Fl. Mal. Pen. 5 (1925) 154; Back. Onkr. Suiker. (1928) 159, non t. 165; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 23 p.p.; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 465. — Scirpus schoenoides RETZ. Obs. 5 (1789) 14. — Scirpus bispicatus Roxs. F1. Ind. 1 (1820) 223. — Abildgaardia nervosa PRESL, Rel. Haenk. 1 (1828) 180; SteuD. Syn. 2 (1855) 73. — F. bispicata Nees in Wight, Contr. (1834) 97; Mia. Fl. Ind. Bat. 3 (1856) 317; Borck. Linnaea 37 (1871) 6. — Iriha schoenoides O.K. Rev. Gen. Pl. 2 (1891) 752. — Iriha bispicata O.K. l.c. 753. — F. longifolia S. T. BLAKE, Un. Queensl. Papers, Dept Biol. I, 13 (1940) 9. Glabrous perennial with short rhizome. Stems densely tufted, compressed, sulcate, smooth, often somewhat incrassate at the base, 10—45cm by Y4—-1 mm. Leaves shorter than the stems, very narrow, with involute margins, abruptly acuminate, spinulose- scabrous on the margins in the upper part, greyish green or glaucescent, 74—1 mm wide; lower sheaths 574 FLORA MALESIANA [ser. I, vole stramineous or ferrugineous; ligule a dense fringe of short hairs. Inflorescence consisting of a single terminal spikelet, or 1—2 lateral peduncled spikelets added, the 1-2 rays when present 1-214 cm. In- volucral bracts usually glume-like, the lowest some- times leafy and up to 4cm. Spikelets solitary, globose-ovoid or oblong-ovoid, terete, subacute, densely many-flowered, whitish ferrugineous to ful- vescent, 5—10 (ultimately up to 15) by 3-4 mm; rachilla narrowly winged. Glumes very regularly spiral, subchartaceous, appressed, very broadly ovate, usually somewhat broader than long, obtuse, muticous or apiculate, scarcely keeled, many-nerved almost over the whole breadth, 2A; 3 by 3- 34mm. Stamens 3; anthers oblong- linear, % mm. Style flat, dilated at the base, ciliate in the upper half, Ze 1/4 mm; stigmas 2, much shorter than the style. Nut biconvex, obovate, distinctly stipitate (gyno- phore obconical, c. 1% mm), umbonulate, smooth, at first whitish, brownish at maturity, VA-1% by 1-1/4 mm; epidermal cells minute, isodiametric. Distr. From SE. Asia (India, Thailand, Indo- China, S. China, Formosa) to tropical Australia (Queensland); sparingly introduced in America, see SVENSON, N. Am. FI. 18 (1957) 553; in Malesia: Sumatra, Malay Peninsula, Java, Kangean Arch., Madura, Lesser Sunda Is. (Tanimbar), W. and N. Borneo, Philippines (Palawan, Luzon). Ecol. In open grasslands, fallow rice-fields, at low altitudes, up to 400 m. Uses. Where in the Malay Peninsula it forms a fair proportion of the weeds of somewhat dry rice- fallows, it gets turned in at ploughing time to make green manure (BURKILL). Vern. Philippines: gumi-gumi, Tag. 49. Fimbristylis caesia Mig. Fl. Ind. Bat. 3 (1856) 315; KERN, Blumea 8 (1955) 134; in Back. & Bakh. f. Fl. Java 3 (1968) 465.— F. subbispicata NEES var. caesia CLARKE, Philip. J. Sc. 2 (1907) Bot. 92, haud quoad specim. cit.—F. monostachya (non HAssk.) BACK. Onkr. Suiker. (1928) t. 174. — Fig. 44. Probably annual; glabrous. Stems tufted, obtusely compressed-trigonous, striate, smooth, 20—40 cm by 44-1 mm. Leaves much shorter than the stems, flat, obtuse, smooth except for the margins scabrid in the upper part, glaucescent, 1— 14mm wide; sheaths pale green to cinnamomeous, hardly keeled, ligule a dense fringe of short hairs. Inflorescence simple, with 1-3 spikelets, the lateral spikelet(s) shortly peduncled (peduncles up to 1 cm). Involucral bracts glume-like when the inflorescence consists of a single spikelet, always short, 3—10(—40)mm. Spikelets solitary, oblong-ovoid, terete, very acute, densely many- flowered, often contorted, pale stramineous, brownish variegated, 10-20 by 3—4 mm; rachilla narrowly winged. Glumes spiral, subtristichous, chartaceous, broadly ovate-deltoid, obtuse, mucro- nulate, hardly keeled, many-nerved, with broad hyaline margins, glabrous, 414-5 by 4 mm. Stamens 3; anthers linear, 1 mm. Style flat, slightly dilated at the base, ciliate in the upper half, c. 2 by { mn; stigmas 2, much shorter than the style. Nut biconvex, Saha elliptic, smooth, long- -stipitate (g nophore 4% mm), stramineous, ba 2 by 140-14 mm, the apex emarginate by the c. 44mm wide style- scar; epidermal cells minute, isodiametric. Fig. 44. Fimbristylis caesia MiQ. a. Habit, x 4, b. spikelet, x 5, c. glume d. stamen, e. deflorate flower, f. nut, g. ditto from top, all x 10 (a- —g JUNGHUHN 532). 1974] CYPERACEAE (Kern) 375 Distr. Malesia: W.—-E. Java, Philippines (Luzon). Apparently very rare. Ecol. In grassy fields, margins of rice-fields, among grasses on dry road-sides, in brushwood, also on heavy brackish moist clay, at low altitudes, up to 600 m. Note. This well-characterized species was ne- glected or misunderstood since MIQUEL published it. 50. Fimbristylis subalata KERN, Blumea 8 (1955) 133, f. 2; Reinwardtia 6 (1961) 46.— F. alata CAMuSs, Not. Syst. 1 (1910) 244 p.p.; Fl. Gén. I.-C. 7 (1912) 94 p.p. Glabrous perennial with horizontal, shortly creep- ing, up to 4cm long, woody rhizome clothed with ovate, striate, fuscous sheaths. Stems approximate, uniseriate on the rhizome, very slender, obtusely trigonous, striate-sulcate, smooth, 50-65cm_ by 1mm. Leaves few (c. 2) to the stem, somewhat shorter than the stems, rigid, erect, filiform, with inrolled margins, acute, smooth, or scabrid at the very top, /4—/4 mm wide: ligule short, membranous. Inflorescence consisting of a single terminal spikelet. Spikelet erect, ellipsoid or oblong-ellipsoid, terete, acute, many-flowered, ferrugineous, 15—35 by 4— 5mm; rachilla narrowly winged. Glumes spiral, chartaceous, appressed, ovate, obtuse, muticous or inconspicuously apiculate, scarcely keeled, with strong midnerve and about 10 less prominent nerves on either side, hyaline-margined in the upper part, 6-7 by 4’4-5mm. Stamens 3; anthers linear, 2’4-3 mm. Style flat, slightly dilated at the base, fimbriate-ciliate in the upper half, 314-4 by 14 mm: stigmas 2, shorter than the style. Ovary in the upper half with thinly membranous, transversely striate wings. Nut biconvex, obovate, conspicuously stipi- tate (gynophore Yy- ¥ mm), emarginate at the top by the %~-%% mm broad style-scar, whitish, c. 2 by 14-4 mm; epidermal cells minute, isodiametric; wings of the ovary indurate in the mature nuts and only discernible by their transverse striation different from the reticulation of the nut proper. Distr. Peninsular Thailand, Cochinchina, Cam- bodia; in Malesia: New Guinea (Papua: Lake Daviumbu, Middle Fly R.). Ecol. In savannahs, in small wiry tufts on wet grassy plains, at low altitudes. Note. Closely allied to F. alata Camus from Laos, which is an annual with spikelets 3 mm wide, glumes 4mm long and wide, a glabrous style 2 mm long, anthers 114-124 mm long, and nuts broadly winged even when mature. 11. Section Fimbristylis Fimbristylis sect. Eufimbristylis Boeck. Linnaea 37 (1871) 3.— Fimbristylis sect. Dichotomae OHW1, J. Jap. Bot. 14(1938) 573. — Fimbristylis ser. Dichotomae Onwl, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55. Type species: F. dichotoma (L.) VAHL (Scirpus dichotomus L.). 51. Fimbristylis dichotoma (L.) VAHL, En. 2 (1806) 287, excl. var.; Hassk. Pl. Jav. Rar. (1848) 64: FISCHER, Kew Bull. (1935) 150; S. T. BLAKE, J. Arn. Arb. 35 (1954) 213; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 39; J. Fac. Sc. Un. Tokyo III, 8 (1961) 111; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 466. — Scirpus dichotomus LINNE, Sp. PI. (1753) 50. — Scirpus diphyllus Retz. Obs. 5 (1789) 15. — F. diphylla VAHL, En. 2 (1806) 289; STEUD. Syn. 2 (1855) 116; BENTH. FI. Austr. 7 (1878) 311: CLARKE, Fl. Br. Ind. 6 (1893) 636, excl. var. pluri- striata CLARKE; Philip. J. Sc. 2 (1907) Bot. 93; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 91; CLARKE, Ill. Cyp. (1909) t. 42 f. 1-2; Koorp. Exk. Fl. Java 1 (1911) 199; ibid. 4, Atlas (1922) f. 253; Camus, FI. Gen. I.-C. 7 (1912) 103; Brown, Min. Prod. Philip. For. 1 (1920) 348; Ripv. Fl. Mal. Pen. 5 (1925) 155. — F. annua (non R. & S.) MerRR. En. Philip. 1 (1923) 121; KUx. Bot. Jahrb. 59 (1924) 47; Back. Onkr. Suiker. (1928) 160, t. 166; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 21. — F. pilosa (non VAHL) PRESL, Rel. Haenk. 1 (1828) 191, cf. MerR. Philip. J. Sc. 35 (1928) 5.— F. affinis PRESL, /.c.; F.-VILL. Nov. App. (1882) 308. — F. communis KUNTH, En. 2 (1837) 234: Mig. Fl. Ind. Bat. 3 (1856) 323.— ? Scirpus falcatus LLANos, Fragm. PI. Filip. (1851) 20; Merr. Sp. Blanc. (1918) 83. — ? F. brizoides SM. var. NEES in Hook. J. Bot. Kew Misc. 6 (1854) 28. — F. philippica Steup. Syn. 2 (1855) 116; Mia. Fl. Ind. Bat. 3 (1856) 324; F-ViLv. Nov. App. (1882) 308.— F. calocarpa STEuD. l.c. 117; Mig. l.c. 325. — F. ambigua SteEup. l.c. 117; Mia. l.c. 323. — F. circinnata STEUD. I.c. 116; MiQ. l.c. 324. — F. squarrosa (non VAHL) Mia. l.c. 319. — F. polymorpha Boeck. Linnaea 37 (1871) 14; SCHEFF. Nat. Tijd. N. I. 34 (1874) 56. — F. novae-britanniae BoEcK. Bot. Jahrb. 5 (1884) 93. — F. communis var. gracillima RipL. in Forbes, Nat. Wand. (1885) 521.— Iriha polymorpha O.K. Rey. Gen. Pl. 2 (1891) 752.— F. longispica (non STEUD.) CLARKE, Fl. Br. Ind. 6 (1893) 639; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 93; Fl. Mal. Pen. 5 (1925) 156 p.p. — F. germainii CAMus, Not. Syst. 1 (1910) 246; Fl. Gén. I.-C. 7 (1912) 105.— F. annua var. diphylla KUx. Act. Hort. Got. 5 (1929) 109; Bot. Jahrb. 59 (1924) 47. — F. ramosii Ktx. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 10. Annual, or perennial with very short rhizome. Stems slender to rather stout, tufted, angular, com- pressed especially below the inflorescence, striate, glabrous or pilose, smooth, 10—75(—100)cm_ by 1-2 mm. Leaves basal, from much shorter than to about as long as the stems, weak or rigid, flat, abruptly acuminate, glabrous or more or less pubes- cent, scabrid on the margins in the upper part, green or glaucous, 1)4-5 mm wide; ligule a dense fringe of short hairs. Inflorescence simple to decompound, loose or dense, with few to numerous spikelets, 576 FLORA MALESIANA [ser. I, vol. 7° up to 20cm long. Involucral bracts 2—5, much shorter to somewhat longer than the inflorescence. Primary rays obliquely patent, smooth, glabrous or pilose, up to 10 cm. Spikelets solitary or more or less aggregated, ovoid or oblong-ovoid, terete, acute, densely many-flowered, rufous, fuscous, or casta- neous, 5—10(—20) by 24—3(—5) mm;rachilla narrowly winged. Glumes spiral, chartaceous, very broadly ovate to oblong-ovate, obtuse, often mucronulate, scarcely keeled, glabrous, sometimes minutely cilio- late at the upper edge, with 3-nerved, green keel and obscurely few-nerved sides, 2—3(—4!4) mm _ long. Stamens 1-3; anthers oblong or linear, 14—1(-134) mm. Style flat, broad, hyaline-margined, slightly dilated at the base, ciliate at least in the upper half, 2-2'4(-4) mm; stigmas 2, somewhat shorter than the style. Nut biconvex, obovate or broadly obovate, shortly stipitate, umbonulate, smooth, more rarely sparsely verruculose, conspicuously trabeculate by 5—10 (rarely some more) longitudinal ribs on either face and numerous cross-bars, glistening white to stramineous, rarely brown, 1-114 by 34-1 mm; epidermal cells transversely elliptic to oblong. Distr. All warmer parts of the whole world, one of the most widely distributed species; very common in S. and E. Asia and throughout Malesia. Ecol. Open waste places, grassy road-sides, Imperata-fields, tea-plantations, teak-forests, more rarely in swamps; when growing abundantly often a troublesome weed difficult to eradicate completely, 0—1500(—1750) m. Uses. The leaves furnish a rather large quantity of forage with sufficient food-value. In the Philippines the stems are used for matting, but they are inferior to those of F. globulosa. Vern. Tjikukok, Atjeh, tihe-tihe, tihe solar, si marburi tano, imbulu tano, Sum. E.C., gujun kambing, djanggut kambing, ikal-ikal, S. Sum., rumput injep, Banka, rumput purun batu, r. parah, r. képala lalat, Mal. Pen., djuduluk, djampang muta, bulu mata munding, djukut mata munding, dodombadn, S, sukét kodokan, komis, J, rheba komés, komis bungkol, Md, burun, Borneo, werot intalun, Minah., tentarilomé, Talaud, fafoa, Sulu Is., mentembu, emsumi, Jappen- Biak; New Guinea: balimbuli, NE. New Guinea, kikisanki, Wapi, tumbu, Enga; Philippines: baliotas, taldgig, Bag., bubdging, gilal, Sub., tabtabin, tayok- tayok, Sbl., and many others. Notes. An exceedingly polymorphous species, burdened with hundreds of synonyms. Now the stems are filiform, then again rather robust; stems, leaf-blades, sheaths, and rays of the inflorescence are normally glabrous, but frequently long-hairy; the inflorescence may be widely branched with some hundred spikelets, or reduced to a few spikelets or a single one; size of the glumes, number of stamens, shape of the fruit, etc. are variable. An attempt to disentangle this polymorphism was given by KOYAMA, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 40. F. ramosii KUK., based on a very young collection (Ramos BS 7816 from Manila), is merely a hairy form of F. dichotoma. There is no close affinity to F. ferruginea, with which KUKENTHAL compares it. RIpDLey’s ‘F. longispica STEUD.” is a remarkable, coarse plant from the sandy sea-shore. Similar plants occur in Cochinchina (e.g. CLEMENS 3046 from the sandy sea-shore near Tourane). The stems are stout, up to 80 cm by 2 mm, the rigid leaves 3—4 mm wide, the involucral bracts short, the relatively few (c. 20) spikelets 4-5 mm wide, the glumes up to 44 mm long, the style 3-4 mm, the anthers 1- 14mm, with the connective distinctly produced and often somewhat bristly at the top. It is quite distinct from E. Asian F. longispica STEUD., but may represent a well-distinct race of F. dichotoma. An eastern race of F. dichotoma is F. depauperata R.Br., see below: ssp. dichotoma. — Perennial. Inflorescence usually compound or decompound with many to numerous spikelets. Glumes ovate or oblong-ovate, not ciliolate; cells in the upper part of the glumes oblong-linear. Anthers linear, c. 1(—134) mm. Style 2-274(—4) mm long. Stems, leaves and bracts sometimes stiffly hairy. Synonymy, distribution and ecology as above. ssp. depauperata (R.BR.) KERN, stat. nov. — F. depauperata R.Br. Prod. (1810) 227; Kunru, En. 2 (1827) 227; SteupD. Syn. 2 (1855) 120; BENTH. FI. Austr. 7 (1878) 311; Domin, Bibl. Bot., Heft 85 (1915) 455:S.T. BLAKE, Contr. Queens]. Herb. 8 (1969) 8. — F. spirostachya F.v.M. ex BENTH. FI. Austr. 7 (1878) 311.— F. diphylla var. depauperata CLARKE, FI. Br. Ind. 6 (1893) 637. — F. diphylla var. spirostachya CLARKE, I.c. — F. annua (non R. & S.) 8S. T. BLAKE, J. Arn. Arb. 35 (1954) 212-214. Annual. Inflorescence simple or compound, with few to several spikelets; spikelets pale brown, turgid. Glumes very broadly ovate, broader than long, often minutely ciliolate at the upper edges; cells in the upper part of the glume nearly square. Anthers oblong, 4-34 mm. Style short and broad, 1)4-1/4 by c. 4% mm. Stems, leaves, bracts, and rays of the inflorescence often softly hairy. Distr. Australia (N. Australia, Queensland), Micronesia; in Malesia: only in the extreme SE. part of the Archipelago and here much rarer than ssp. dichotoma: Lesser Sunda Is. (Timor), S. Moluccas (Tanimbar Is.: P. Jamdena), New Guinea. Ecol. Wet grasslands, savannahs, Melaleuca- forests, at low altitudes. Notes. By habit, shape of style, pubescence, etc. reminding one of F. tomentosa VAHL, but in the latter the nut is seated on a conspicuous gynophore, not strikingly longitudinally ribbed, and with epi- dermal cells in numerous rows. S. T. BLAKE, l.c., treats F. depauperata R.BR. (as F. annua R. & S.) as specifically distinct from F. dichotoma. To my mind it is rather a geographical race not identical with the European F. annua (ALL.) R. & S. The latter differs by the ovate glumes with different structure of the composing cells, the slender style half as broad as that of F. dichotoma ssp. depauperata, and is always a dwarf, subglabrous plant. The North American F. baldwiniana (SCHULT.) Torr. is also a closely related annual. Both F. annua and F. baldwiniana may better be treated as races of F. dichotoma s.l. 52. Fimbristylis tomentosa VAHL, En. 2 (1806) 290; S. T. BLAKE, Contr. Queens]. Herb. 8 (1969) 13. — F. affinis PresL, Rel. Haenk. 1 (1828) 191. — F. podo- carpa Nees & Mey. ex NEES in Wight, Contr. (1834) 98, p.p. typ.; Nov. Act. Ac. Caes. Leop.-Car. 19 1974] CYPERACEAE (Kern) Sy Suppl. 1 (1843) 77 p.p.; SteuD. Syn. 2 (1855) 117; CLARKE, FI. Br. Ind. 6 (1893) 638; Philip. J. Sc. 2 (1907) Bot. 94; Ill. Cyp. (1909) t. 42 f. 5-6; Koorp. Exk. Fl. Java 1 (1911) 199; Camus, Fl. Gén. I.-C. 7 (1912) 107; Merr. En. Philip. 1 (1923) 125; KERN, Blumea 8 (1955) 139; in Back. & Bakh. f. Fl. Java 3 (1968) 466. — F. cincta Nees in Wight, Contr. (1834) 98.— F. squarrosa (non VAHL) ZOLL. Syst. Verz. 1 (1854) 61 p.p. (specim. glaucescentia). — F. subtristachya STEUD. Syn. 2 (1855) 109. — F. diphylla var. pluristriata CLARKE, FI. Br. Ind. 6 (1893) 637; Ill. Cyp. (1909) t. 42 f. 3-4; CHERM. in Humb. FI. Madag., fam. 29 (1937) 179.— F. diphylla (non VAHL) K.ScH. & Laut. FI. Schutzgeb. (1900) 196 (quoad LAUTERBACH 348). — F. annua var. podocarpa KUx. Bot. Jahrb. 59 (1924) 5, 48. — F. schoenoides (non VAHL) BACK. Onkr. Suiker. (1928) t. 165, non p. 159.— F. diphylla var. podocarpa KUxk. Bot. Jahrb. 69 (1938) 257. — F. pluristriata BERH. Bull. Soc. Bot. Fr. 101 (1955) 376. — F. dichotoma ssp. podocarpa KoyAMA, Micronesica 1 (1964) 87.— Fig. 45. Annual with fibrous roots. Stems tufted, angular, compressed just below the inflorescence, smooth, 40-60 cm by 4-1 mm. Leaves shorter than to about as long as the stems, flat, abruptly acuminate, softly hairy, rarely glabrous, scabrid on the margins in the upper part, 2-3 mm wide; ligule a fringe of short hairs. Inflorescence simple or compound, loose, with 5—25 spikelets, very rarely reduced to a single spikelet, up to 8 cm long. Involucral bracts up to 5, the longest overtopping the inflorescence, up to 10cm. Primary rays obliquely erect, compressed, glabrous and smooth, the longest 2—4 cm. Spikelets solitary, ovoid or oblong-ovoid, rarely oblong, terete, acute, densely many- -flowered, brown, 5- 10(—15) by (2i4- )3—4 mm; rachilla narrowly winged. Glumes spiral, chartaceous, appressed, broadly ovate or orbicular, mucronulate, scarcely keeled, with strong midnerve and nerveless or faintly nerved sides, hyaline-margined, glabrous or with some soft hairs on the midnerve, 3!4-3!4 mm long and wide. Stamens (1—)2; anthers oblong-linear, 24-1 mm. Style flat, short and broad, hyaline-margined, grad- ually but slightly dilated towards the base, ciliate in the upper 74-24, UA4-1)4 by 4-/K mm; stigmas 2, shorter than the style. Nut biconvex, broadly elliptic to orbicular, prominently stipitate (the gynophore obpyramidal, often somewhat saucer-shaped, )4- 44mm long and wide), umbonulate, sometimes tuberculate, especially towards the obtuse, thickened edges, at first white, finally stramineous, y-1% by %o-17%% mm, reticulate by the impressed epidermal cells superposed i in 16—24 vertical rows on either face. Distr. Tropical Africa, Madagascar, Mauritius; from India through Farther India to S. China, Ryu Kyu Is., Micronesia and Queensland (Cape York Peninsula); widely distributed in Malesia, but much rarer than F. dichotoma: Malay Peninsula (a single collection from Pahang), W. Sumatra, W. Java (Tjikoya, Cheribon), E. Java (Madiun), Kangean Arch., Lesser Sunda Is. (Alor), Philippines (Luzon, Leyte, Bohol, Panay), N. Celebes (Tondano), Moluc- cas (Halmaheira), New Guinea, New Britain. Ecol. In open places or in light shade: moist grasslands, grassy road-sides, in and along rice-fields, forest-clearings, at low altitudes (usually below 200 m, in W. Sumatra at 600 m); in Khasia up to 1200 m. Vern. Mdtequila, Alor (probably for several Cyperaceae; see F. microcarya). Notes. I fail to distinguish between the specimens rightly named by CLARKE F. podocarpa NEES and those he referred to his var. pluristriata of F. diphylla. I consider them conspecific. F. cincta Nees from Mauritius differs from F. tomentosa only in some minor points (see WIGHT, Contr.). F. pluristriata (C. B. CLARKE) BERH. and F. subtristachya STEUD. are superfluous names for the African plant, which to my mind cannot be separated specifically from F. tomentosa. 53. Fimbristylis lineatisquaama OHwI1, Blumea 8 (1955) 109, f. 10. Perennial with short rhizome. Stems tufted, com- pressed-trigonous, striate, glabrous or sparsely soft- hairy, few-leaved at the base, 30-40 cm by }4-1 mm. Leaves half as long as the stems, rather stiff, curved, flat, rather abruptly acuminate, densely hairy in the lower part, scabrid on the margins, 1)4—2 mm wide; sheaths pale, densely hairy in the upper part; ligule a dense fringe of hairs. Inflorescence simple or com- pound, loose, with 4—10 spikelets, 2-5 cm long. Involucral bracts up to 5, the lowest shorter than to as long as the inflorescence. Primary rays few, compressed, smooth and glabrous, 2—4 cm. Spikelets solitary, oblong-ovoid, terete, acute, many-flowered, 7-10 (ultimately up to 14) by 2/4-3 mm; rachilla narrowly winged. Glumes spiral, membranous, oblong-ovate, rather acute, apiculate or mucronulate, scarcely keeled, glabrous, distinctly many-nerved almost over the whole breadth, brown, 3— 4, by 14-3 mm. Stamens 3; anthers linear, ye 124 mm, with distinctly p~ aduced, papillose-setulose append- age of the connective. Style flat, rather broad, slightly dilated at the base, densely ciliate in the upper half, 2—2)4 mm; stigmas 2, about as long as the style. Nut turgidly biconvex, with obtuse edges, oblong-obovate, broadly stipitate, umbonulate, re- ticulate by the roundish or transversely elliptic, slightly impressed epidermal cells superposed in 12-16 vertical rows on either face, greyish brown, 4-14 by Ao-%o mm. Distr. Malesia: Philippines (Sulu Is.: Golo). With certainty only known from the type-collection (MERRILL 11546); a poor collection from Luzon (Benguet Subprov., Pauai: CLEMENS 9145) may belong here. Notes. The description of F. urakasiana KUK. Bot. Jahrb. 59 (1924) 5, based on a collection from the Mariannas, Urakas Island (GIBBON 1158, not seen), agrees very well with that of F. lineatisquama, so that I presume that the former name is the correct one for this species. According to Koyama, J. Fac. Sc. Un. Tokyo II, 8 (1961) 113, F. urakasiana is conspecific with F. boninensis HayaTa, Ic. Pl. Form. 6 (1916) 109, f. 27 by longispica STEUD. var. boninensis OHW1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 80.— F. dichotoma ssp. longispica KOYAMA var. boninensis Koyama, l.c.], but this is very unlikely, as F. bon- inensis is a plant with contracted, head-like inflo- rescence, glabrous leaves, and broad, obtriangular- obovate nuts. F. boninensis is recorded for the Philippines: Batan Island, on the littoral cliff on the eastern foot of Mt Iraya, frequent: HATUSIMA 578 FLORA MALESIANA fser. I, vol Fig. 45. Fimbristylis tomentosa VAHL. a. Habit, x }4, b. spikelet, x 5, c. glume, d. anther, e. nut with persistent filaments and style, all x 10, f. outer cells of nut strongly enlarged (a—f BACKER 27124). 1974] CYPERACEAE (Kern) 579 28713 (not seen). See Mem. Fac. Agric. Kagoshima Un. V, 3 (1966) 60. In shape and texture of the nuts and in the setulose top of the connective F. lineatisquama shows much affinity to F. trichophylla. It can be distinguished by the long-ciliate ligule and the longer, more dis- tinctly nerved glumes. The delimitation against F. trichophylla and F. tomentosa requires further investigation. 54. Fimbristylis bisumbellata (ForskK.) Bus. Dodec. (1850) 30; FiscHER, Kew Bull. (1935) 149; Fl. Madr. 11 (1936) 1898, in corrig.; OHW1, Mem. Coll. Sc Kyoto Imp. Un. B18 (1944) 85; KERN, Blumea 8 (1955) 135; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 113; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 466. — Scirpus bisumbellatus Forsk. Fl. Aeg.-Arab. 1 (1775) 15. — Scirpus dichotomus (non L.) ROTTB. Descr. & Ic. (1786) 57, t. 13 f. 1. — F. dichotoma var. villosa VAHL, En. 2 (1806) 287. — F. dichotoma (non VAHL) KuNTH, En. 2 (1837) 225; Boeck. Linnaea 37 (1871) 12 p.p.; Mig. Fl. Ind. Bat. 3 (1856) 319; BENTH. FI. Austr. 7 (1878) 310; CLARKE, FI. Br. Ind. 6 (1893) 635; Philip. J. Sc. 2 (1907) Bot. 93; Koorp. Exk. Fl. Java 1 (1911) 199; Camus, FI. Gén. I.-C. 7 (1912) 102 p.p. — Scirpus pallescens Roxs. FI. Ind. 1 (1820) 229. — F. pallescens NEES in Wight, Contr. (1834) 101; Streup. Syn. 2 (1855) 111.— Iriha bisumbellata O.K. Rev. Gen. Pl. 2 (1891) 752. Annual with fibrous roots. Stems slender, densely tufted, trigonous, smooth, 7-25cm by )4-1 mm. Leaves shorter to somewhat longer than the stems, flat, abruptly acuminate, glabrous or pubescent beneath, scabrid on the margins in the upper part, 1—2 mm wide; sheaths membranous, stramineous or ferrugineous; ligule a dense fringe of short hairs. Inflorescence compound or decompound, loose, with many spikelets, 2—6cm across. Involucral bracts 2—3, suberect, the lowest as long as or overtopping the inflorescence. Primary rays 5-10, slender, smooth, 1—3 cm. Spikelets solitary, oblong-ovoid to narrowly oblong, angular, acute, 3-8 by c. 114 mm; rachilla narrowly winged. Glumes spiral, membra- nous, glabrous, broadly ovate, acute, mucronulate, sharply keeled, with 3-nerved, green keel, brownish, lineolate- puncticulate sides, ‘and hyaline margins, c. 1% by 1/4 mm. Stamen 1; anther oblong, 4 mm. Style slender, flat, with dilated base, ciliate in the upper half, ye 1 mm; stigmas 2, somewhat shorter than the style. Nut biconvex, broadly obovate or obovate, shortly stipitate, umbonulate, conspicu- ously trabeculate, stramineous, %<— YK by %- “mm: epidermal cells impressed, transversely oblong, in 5—9 vertical rows on either face. Distr. From the Mediterranean region through the Old World Tropics to Australia (N.S. Wales); in Malesia with certainty only known from E. Java (Modjokerto, along Brantas R.) and the Philippines (Luzon). There are also specimens in the Florence Herbarium from ‘Java, LABILLARDIERE’’, and in the British Museum from ‘“‘Malaya; probably Borneo, Lops” without precise localities. Ecol. Along rivers, on sandy river bars; in India a common weed of the rice-fields. Note. MERRILL merged the Philippine collections into F. dichotoma (L.) VAHL (as F. annua R. & S.), but it is one of the best characterized species of the difficult group, readily distinguishable by its angular spikelets and membranous, acutely keeled, mucro- nulate glumes. 55. Fimbristylis merrillii KERN, Blumea 8 (1955) 135, f. 6; in Back. & Bakh. f. Fl. Java 3 (1968) 466. — F. squarrosa (non VAHL) Merr. En. Philip. 1 (1923) 126 p.p. (quoad BS 26093). — F. annua var. gracilis Back. Onkr. Suiker. (1928) 160 p.p.; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 21 p.p. Annual with fibrous roots. Stems tufted, com- pressed-trigonous, smooth, often slightly pubescent at the top, leafy at the base, (2—)15—30(—40) cm by Y-1 mm. Leaves 4-4 as long as the stems, often curved, flat, rather abruptly acuminate, glabrous, or pubescent beneath, often ciliate on the margins, scabrid in the upper part, 1-2 mm wide; ligule a dense fringe of short hairs. Inflorescence simple or compound, loose, diffuse, with several spikelets but sometimes reduced to a single spikelet, up to 5(—10) cm long and wide. Involucral bracts 1-3, shorter than the inflorescence, often ciliate- puberulous at the dilated base, 1—3(—6) cm long. Primary rays up to 8, obliquely patent, glabrous and smooth, 1—4(—8)cm. Spikelets solitary, ovoid or oblong-ovoid, terete, acute, densely few- to several- flowered, 3—5(—7) by 14-2 mm; rachilla narrowly winged. Glumes spiral, appressed, subchartaceous, broadly ovate, rather acute, muticous or apiculate, slightly keeled, glabrous, with 3-nerved keel, nerve- ae shining rufous sides, and hardly hyaline margins, (VA- WA 2 by c. 14mm. Stamens 1(—2); anthers AOee Y%-'4 mm. Style slender, flat, not or hardly hyaline- margined glabrous or with ie 4 short cilia at the very top, >4—1 mm; stigmas 2, shorter than the style. Nut biconvex, obovate or broadly obovate, shortly stipitate, minutely umbonulate, sometimes sparsely verruculose, conspicuously trabeculate by the impressed, transversely oblong epidermal cells in 5—6(—9) vertical rows on either face, 4—34(-2{,) by Avo ™M. : , Distr. SE. Thailand, S. China, Queensland (Cape York Peninsula); in Malesia: N. Sumatra (Atjeh: Ketol Valley), throughout Java, Kangean Arch., Madura, Tanimbar Is., Philippines (Luzon, Panay, Mindanao), Celebes, New Guinea (Papua). Ecol. In swamps, wet rice-fields, grass-fields, teak-forests, at low altitudes (0-300 m), in Atjeh at 800-1000 m. 56. Fimbristylis tenuinervia KERN, Blumea 8 (1955) 137, f. 7; ibid. 13 (1965) 121. — F. diphylla Vaunt f. malasica CLARKE in sched. Probably annual. Stems densely tufted, very slender, obtusely trigonous, somewhat compressed, striate-sulcate, glabrous and smooth, leafy at the base, 15—35(—75) cm by c. 4 mm. Leaves somewhat shorter than to about as long as the stems, almost filiform, often with inrolled margins, acute, glabrous, or slightly puberulous on the sheaths, scabrid towards the apex, 1-114 mm wide; ligule a fringe of short hairs. Inflorescence simple or compound, very loose, with (3—)10—35 spikelets, (2—)3—8 cm long. Involucral bracts 3—5, erecto-patent, scarious-margined at the dilated base, glabrous or ciliolate, the lowest over- topping the inflorescence, up to 10 cm. Primary rays 5—7, very slender, erecto-patent, compressed, gla- brous and smooth, the longest up to 4 cm. Spikelets 580 FLORA MALESIANA [ser. I, vol. 7° solitary, oblong-lanceolate, terete, very acute, densely many-flowered, 4—5 by 1)4-1)4 mm; rachilla nar- rowly winged. Glumes spiral, appressed, chartaceous, glabrous, broadly ovate, apiculate, scarcely keeled, with prominent midnerve, faintly 5—8-nerved sides, and hyaline margins, shining brown in the apical part, much paler below, 2)/4-2%4 by 2-2/4 mm. Stamens (2—)3; anthers linear, (1-)1/4 mm. Style slender, flat, not hyaline- margined, hardly dilated at the base, ciliate in the upper half, 114-114 mm; stigmas 2, shorter than the style. Nut biconvex, narrowly obovate, shortly stipitate, umbonulate, trabeculate by 4—6 longitudinal ribs and numerous cross-bars, smooth, stramineous, ){)-?%) by /4- % mm; epidermal cells impressed, transversely oblong-linear. Distr. Insufficiently known; Malesia: Philippines (Central Luzon), NE. New Guinea (Sepik Distr.). Ecol. Grassy road-sides, moist meadows, growing in dense mats. i Fimbristylis perlaxa OHW1, Blumea 8 (1955) 101, Me Annual with fibrous roots. Stems very slender, tufted, compressed, obtusely trigonous, smooth, 30-50 cm by 4-1 mm. Leaves in the lower )4—)% of the stem, 14-14 as long as the latter, weak, flat, shortly acuminate to very acute, glabrous above, more or less pubescent beneath and on the pale green sheaths, 4-1/4 mm wide; ligule a dense fringe of hairs. Inflorescence compound, very loose, with c. 10-20 spikelets, 8-15 cm long. Involucral bracts erect, pubescent beneath, the lowest somewhat shorter than to about as long as the inflorescence. Primary rays very slender, suberect, compressed, glabrous and smooth, 4—6cm. Spikelets solitary, long-peduncled, oblong-ovoid, slightly angular, very acute, densely many-flowered, light brown, 6—8 by 2 mm; rachilla very narrowly winged. Glumes spiral, membranous, appressed, ovate, mucronulate, slightly keeled, glabrous, with 3-nerved keel, obsoletely “i 2-nerved sides and scarcely hyaline margins, 24 by 14mm. Stamens 2; anthers oblong, ye 44 mm. Style slender, flat, not hyaline-margined, slightly dilated at the base, sparsely ciliate at the top, 1 mm; stigmas 2, shorter than the style. Nut biconvex with acutish edges, broadly elliptic or suborbicular, shortly stipitate, umbonulate, finely cancellate by the roundish or transversely elliptic, slightly impressed epidermal cells in 15—18 vertical rows on either face, shining, stramineous or brownish, YZ by A mm. Distr. Malesia: Moluccas (Ceram), New Guinea (W. New Guinea: Mamberamo Distr.; NE. New Guinea: Sepik Distr.). Ecol. In swampy places at low altitudes. Note. By its very slender habit much resembling F. gracilenta, from which it is readily distinguishable by the hairy ligule (absent in F. gracilenta), the glabrous rays of the inflorescence (see, however, F. gracilenta var. psilopoda), and the different shape and texture of the nut. 58. Fimbristylis alboviridis CLARKE, Fl. Br. Ind. 6 (1893) 638; KERN, Blumea 8 (1955) 140; in Back. & Bakh. f. Fl. Java 3 (1968) 466.— ? F. annua var. pluristriata BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 21, non F. diphylla var. pluristriata CLARKE. — Fig. 46. Annual with fibrous roots. Stems slender, tufted, compressed, obtusangular, glabrous and smooth, 20-50 cm by )4-1 mm. Leaves half as long as the stems, sometimes slightly falcate, rather firm, flat, obtuse to rather acute, glabrous, scabrid on the margins in the upper part, greyish green, 1-2 mm wide; ligule a fringe of short, white hairs (see Note). Inflorescence simple or subcompound, loose, with few to several spikelets, 2-7 cm long. Involucral bracts 1—2, suberect, the lowest somewhat shorter than to twice as long as the inflorescence, up to 12cm. Primary rays few, suberect, compressed, smooth, up to 5cm. Spikelets solitary, ovoid to oblong-ovoid, terete, acutish, densely many-flowered, whitish or greyish green, often brownish variegated, 5—7 (ultimately up to 10) by c. 2)4 mm; rachilla narrowly winged. Glumes spiral, subchartaceous, glabrous, appressed, broadly ovate, obtuse, muticous or scarcely apiculate, not keeled, with greenish, 3-nerved keel and nerveless sides brownish except for the whitish basal part and margins, c. 2 mm long and wide. Stamen 1; anther oblong, c. /4 mm. Style slender, flat, slightly dilated at the base. ciliate in the upper 14-34, c. 1 mm; stigmas 2, shorter than the style. Nut biconvex, obovate, shortly stipitate, umbonulate, scaly-verruculose, obsoletely reticulate by the transversely elliptic or oblong, not impressed epidermal cells in 10-16 vertical rows on either face, stramineous, 1-1/4 by 4-24) mm. Distr. SE. Asia, from India to Malesia: Malay Peninsula (Kedah, Selangor), ? W. and E. Java, Philippines (Luzon). Ecol. Dry grass-fields, road-sides, at low altitudes. Note. The collections from the Malay Peninsula and the Philippines perfectly agree with the type- collection from India. In the Java collections the ligule consists of a few hairs only, the glumes are more acute, and the nuts slightly different in shape. They may represent a local race or a closely related species. 12. Section Rigidulae KERN, Blumea 8 (1955) 161. Type species: F. rigidula NEES. 59. Fimbristylis trichophylla RipL_. Fl. Mal. Pen. 5 (1925) 155; HENDerS. J. Mal. Br. R. As. Soc. 17 (1939) 86; KERN, Blumea 8 (1955) 140: ibid. 10 (1960) 648; Reinwardtia 6 (1961) 47. var. trichophylla. Perennial with short rhizome. Stems very slender, setaceous, densely tufted, decumbent or hanging in tresses over rocks, subterete, finely striate, glabrous and smooth, 15-60 cm by i JA mm. Leaves often very long (up to 50 cm), weak, filiform, very acute, with incrassate margins scabrid at the top, glabrous, 1974] CYPERACEAE (Kern) 581 Fig. 46. Fimbristylis alboviridis CLARKE. a. Habit, x 14, b. spikelet, x 5, c. glume, d. anther, e. deflorate flower, . nut, all x 10, g. outer cells of nut strongly enlarged (a—g BACKER s.n., Weltevreden a. 1903). 582 or sparsely hairy also on the sheaths, deep green, 4YA-% mm wide; ligule absent. Inflorescence simple or subcompound, loose, with (1—)3—5(—10) spikelets. Involucral bracts 1—3, erect, the lowest usually much overtopping the inflorescence, up to 8(—15) cm. Rays spreading, often curved, compressed, smooth, 1-2cm. Spikelets solitary, ovoid, terete, acute, shining brown, 4-6 (ultimately up to 10) by 2-2'% mm; rachilla very narrowly winged. Glumes spiral, membranous, ovate or broadly ovate, obtuse, apiculate, not keeled, obscurely many-nerved with prominent midnerve, glabrous or sparsely short- hairy, minutely ciliolate, brown with paler margins, 244-3 by 2-24 mm. Stamens 3; anthers linear, 1-1/4mm, with distinctly produced connective bristly at the top. Style flat, scarcely dilated at the base, densely ciliate in the upper half, 174-274 mm: stigmas 2, about as long as the style. Nut biconvex, oblong-obovate, or suboblong, shortly stipitate, scarcely umbonulate, smooth or sparsely verru- culose, minutely cancellate by the slightly impressed, roundish or transversely elliptic epidermal cells in 10-12 vertical rows on either face, brown, 1-114 by YA-%A mm. Distr. Malesia: Malay Peninsula, in the typical form described above only known from Pulu Langkawi. Ecol. In crevices of limestone rocks in dry open places, 60—240 m. var. erecta HOLTT. ex KERN, Blumea 8 (1955) 41. — Fimbristylis sp. prox. F. fusca HENDERS. J. Mal. Br. R. As. Soc. 17 (1939) 86. Less slender. Stems erect, subterete or bluntly trigonous. Leaves stiffer and paler, 1-1/4 mm wide. Inflorescence usually ampler, compound or sub- decompound, rarely simple, with up to 20 spikelets. Primary rays up to 7 cm. Anthers 1}4-2 mm. Distr. Peninsular Thailand; in Malesia: Malay Peninsula (Kedah: Pulu Langkawi, Gunong Baling; Kelantan: Gua Teja, Pahang: Gunong Senyum; Selangor: Bukit Takun near Kanching). Ecol. As the typical variety. Note. The Pulu Langkawi specimens of this variety have densely pubescent stems and leaves, the others are glabrous. In the specimens from Pahang (summit of Gunung Senyum, 1600 ft) the spikelets are somewhat larger than in the others, the nuts slightly broader (74 mm), and the appendage of the connective is smooth. 60. Fimbristylis rigidula NEEs in Wight, Contr. (1834) 99; Hook. J. Bot. Kew Misc. 6 (1854) 29; Steup. Syn. 2 (1855) 116; CLARKE, Fl. Br. Ind. 6 (1893) 640; J. Linn. Soc. Bot. 36 (1903) 242; Philip. J. Sc. 2 (1907) Bot. 95; Ill. Cyp. (1909) t. 42 f. 7-8: MerR. En. Philip. 1 (1923) 125, p.p.; KERN, Blumea 8 (1955) 141; Reinwardtia 6 (1961) 47: TANG & WANG, FI. Reip. Pop. Sin. 11 (1961) 91, t. 29 f. 9-12. — F. hanceana BoEcK. Linnaea 38 (1874) 394. — F. fer- ruginea (non VAHL) VIDAL, Phan. Cuming. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 284, p.p. Perennial with thick, woody, shortly horizontally creeping rhizome clothed with the fibrous remains of old leaf-sheaths. Stems closely uniseriate on the rhizome, compressed, conspicuously (often bul- bously) thickened at the base, smooth, 10—40(—70) cm by 1-1/4(-3) mm. Leaves about half as long as the FLORA MALESIANA [ser. I, vol. 7° Fig. 47. Fimbristylis semarangensis OHW1. a. Habit, x 5, b. spikelet, x6, c. glume, d. stamen, e. ditto, apex, f. deflorate flower, g. nut, all x12, A. outer cells of nut, strongly enlarged (a—~h KOORDERS 42656). 1974] CYPERACEAE (Kern) 583 stems, rigidulous, flat, abruptly acuminate, scabrid on the margins in the upper part, glabrous or (with their sheaths) pubescent, greyish green, 2—3(—5) mm wide; ligule absent. Inflorescence simple or com- pound, loose, often diffuse, with (4—)6—12(—35) spikelets. Involucral bracts 3—4, usually shorter than the inflorescence, sometimes slightly longer, erect, up to 6 cm. Primary rays smooth, compressed, with 1-3 spikelets, or again branched, 1—6 cm. Spikelets solitary or partly paired, ellipsoid or sub- globose, terete, very obtuse, densely many-flowered, fuscous to castaneous, 5-10 by 3—4 mm; rachilla very narrowly winged. Glumes spiral, membranous, ovate or broadly ovate, obtuse, apiculate, scarcely keeled, with prominent midrib and obscurely many- nerved sides, glabrous, or (in some of the continental Asian specimens) sparsely hairy and ciliolate at the apex, 3—4 by 2-3 mm. Stamens 3; anthers linear, 144-3 mm. Style flat, dilated at the base, long- ciliate in the upper half, 2-3 mm; stigmas 2, about as long as the style. Nut biconvex, obovate, minutely stipitate, umbonulate, smooth or sparsely verru- culose, finely reticulate by the isodiametric eae) cells in 20—30 vertical rows on either face, LY4-1) by 140-14 mm. Distr. From Nepal and N. India to N. Thailand and §. China; in Malesia: Philippines (Luzon, Mindanao). Ecol. In Luzon in wet open grassland at low and medium altitudes: in Thailand also in deciduous forests, in India up to 1800 m. Note. The specimens of MCGREGOR BS 32233 and Ramos BS 32811, cited by MERRILL /.c. under F. rigidula belong to F. insignis. 61. Fimbristylis sumbaensis OHw1, Blumea 8 (1955) 106, f. 9 Perennial with short rhizome. Stems slender, densely tufted, compressed-trigonous, obtusangular, striate, slightly puberulous or scabrid at the top, 30-40 cm by 4-34 mm. Leaves half as long as the stems, erect, often with circinnate top, rigidulous, flat, abruptly acuminate, at first slightly pubescent, soon glabrescent, scabrid on the margins at least in the upper part, greyish green or glaucous with brownish sheaths, 1-1/4 mm wide; ligule absent. Inflorescence subcompound to subdecompound, loose, with several to many spikelets, 3-10 cm long. Involucral bracts 3—4, the lowest somewhat shorter than the inflorescence, up to 8cm. Primary rays ascendent-erect, compressed, scabrid-pilose, up to 7cm.~ Spikelets solitary, ovoid or oblong-ovoid, somewhat angular, acute, densely many-flowered, dull greyish brown, 4-6 by 2-2/4 mm; rachilla narrowly winged. Glumes spiral, membranous, gla- brous, ovate, acutish, apiculate or minutely mucro- nulate, with c. 5-nerved, green keel, prominent midnerve, fulvous nerveless sides, and very broad hyaline margins, 2a- 3 by c. 2mm. Stamens 3; anthers linear, 1— VA mm. Siyle slender, flat, scarcely margined, ciliate in the upper half, retrorsely his- pidulous at the abruptly dilated base, c. 2 mm; stigmas 2, somewhat shorter than the style. Nut turgidly biconvex, broadly obovate, with acutish edges, shortly stipitate, minutely umbonulate, smooth, delicately trabeculate by longitudinal ribs and transverse lines, at first white, ultimately shining brown, *4 by ?4 mm; epidermal cells slightly im- pressed, transversely oblong-linear, in 8—10 vertical rows on either face. Distr. Malesia: Lesser Sunda Is. (Sumba: near Waingapu; here repeatedly collected, as yet not known.elsewhere). Ecol. Open or shaded meadows, grassy plains, abundant. Note. OHw! supposed it to be a hybrid of F. sericea and F. dichotoma. It has certainly nothing to do with F. sericea (which is not known from the Lesser Sunda Islands!), and because of its freely fruiting I do not see any reason to suppose hybrid nature. burnt-over 62. Fimbristylis semarangensis OHW1, Blumea 8 (1955) 106, f. 8: KERN, I.c. 141; in Back. & Bakh. f. Fl. Java 3 (1968) 467. — F. annua var. gracilis BACK. Onkr. Suiker. (1928) 160, p.p.; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 21, p.p. — Fig. 47. Glabrous annual with fibrous roots. Stems slender, tufted, compressed, obtusangular, smooth, 20—40 cm by c. 1 mm. Leaves 4—'% as long as the stems, erect, rather firm, flat, acute, scabrid on the margins in the upper part, 1-2 mm wide; ligule absent. Inflorescence subcompound or compound, very loose, with 5—20 spikelets, 3-6 cm long and wide. Involucral bracts 1-2, the lowest shorter than the inflorescence, 1-2cm. Primary rays obliquely ascending, com- pressed, smooth, up to 4cm. Spikelets solitary, narrowly ovoid, terete, acute, densely many-flowered, 6-8 by 2mm; rachilla narrowly winged. Glumes spiral, membranous, ovate, obtusish with rounded apex, muticous, scarcely keeled, with a midnerve not reaching the apex, nerveless, light brown sides, and broad hyaline margins, c. 2/4 by 14mm. Stamens (2—)3; anthers linear, 1 mm. Style slender, flat, not margined, slightly dilated at the base, ciliate in the upper 4-74, 114 mm long; stigmas 2, shorter than the style, 1 mm long. Nut biconvex, obovate, with acutish edges, shortly stipitate, um- bonulate, verruculose, finely reticulate by the minute, roundish or transversely elliptic, not impressed epidermal cells, c. 24 by 24 mm. Distr. Insufficiently known; Indo-China (Cam Ranh Peninsula, Kiét 224 in P); in Malesia: Central Java (Res. Semarang: Bledug-Kuwu; Res. Rembang: Kesongo). Ecol. Along mud-wells, in saline pools in small tufts scattered among Xerochloa imberbis R.BR., at low altitudes. Note. In habit very similar to F. merrillii, but well distinct by the eligulate leaves, the broad hyaline margins of the glumes, the longer anthers, the ciliate style, and the reticulate (not trabeculate) nuts. 13. Section Pogonostylis (BERTOL.) PAx in E. & P. Nat. Pfl. Fam. 2, 2 (1887) 113. — Pogonostylis BERTOL. Fl. Ital. 1 (1833) 312. — Fimbristylis sect. Squarrosae Onw1, J. Jap. Bot. 14 584 FLORA MALESIANA [ser. I, vol. 7° (1938) 573. — Fimbristylis ser. Squarrosae OHWI, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55. Type species: F. squarrosa VAHL. 63. Fimbristylis aestivalis (RETZ.) VAHL, En. 2 (1806) 288; KunTH, En. 2 (1837) 226; StEUD. Syn. 2 (1855) 110; Boeck. Linnaea 37 (1871) 11, excl. var.; BENTH. Fl. Austr. 7 (1878) 310; F.-ViLL. Nov. App. (1882) 308; CLARKE, FI. Br. Ind. 6 (1893) 637, p.p.; Philip. J. Sc. 2 (1907) Bot. 94; Ill. Cyp. (1909) t. 41 f. 14-15; Koorp. Exk. Fl. Java 1 (1911) 199; ibid. 4, Atlas (1922) f. 254; Camus, Fl. Gén. I.-C. 7 (1912) 106, f. 16, 7—8; MeERR. En. Philip. 1 (1923) 121; Back. Onkr. Suiker. (1928) 160, t. 167; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 22; S. T. BLAKE, J. Arn. Arb. 35 (1954) 212: Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 116, excl. varr.; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 467. — Scirpus aestivalis RETz. Obs. 4 (1768) 12. — F. dichotoma (non VAHL) PRESL, Rel. Haenk. 1 (1828) 191. — F. squarrosa (non VAHL) ZOLL. Syst. Verz. 1 (1854) 61, p.p. (specim. parva); De VriesE, Pl. Ind. Bat. Or. (1856-57) 141; MerRR. En. Philip. 1 (1923) 126, excl. BS 26093). — F. griffithiana StEuD. Syn. 2 (1855) 110.— F. tri- cholepis Miq. Fl. Ind. Bat. 3 (1856) 319. — Iriha aestivalis O.K. Rev. Gen. Pl. 2 (1891) 751. var. aestivalis. — Synonymy as above. Annual with fibrous roots. Stems very slender, densely tufted, setaceous, angular, smooth, 3—20 cm by %-A mm. Leaves shorter than the stems, filiform, flat or canaliculate, acute, densely soft-hairy also on the stramineous or ferrugineous sheaths, AA mm wide; ligule absent. Inflorescence com- pound, more rarely subsimple, loose, with many spikelets, 1-7 cm long. Involucral bracts up to 6, similar to the leaves, the lower 1-2 somewhat shorter than the inflorescence to slightly overtopping it. Primary rays filiform, obliquely patent, glabrous and smooth (see var. trichopoda !), up to 5 cm. Spikelets solitary, ovoid or oblong-lanceolate, angular, acute, densely many-flowered, greenish brown, 3-7 by 1-1/4 mm; rachilla narrowly winged. Glumes spiral, thinly membranous, subpatulous at the top, ovate, obtusish, mucronulate, with sharp, green, obscurely 3-nerved keel, prominent midnerve, and pubescent or glabrescent nerveless sides, ferrugineous, darker lineolate, A> V4 by %- 1 mm. Stamen 1: anther oblong, ie 14 mm. Style flat, not hyaline- margined, dilated at the base, sparsely ciliate at the top, glabrous or with a few short cilia at the base, 4-34 mm; stigmas 2, shorter than the style. Nut biconvex, with acute edges, elliptic or obovate, shortly stipitate, umbonulate, obscurely reticulate by the hexagonal, not impressed epidermal cells, smooth, shining stramineous, 14-24 by 7Z- 1, mm. Distr. From Ceylon and India through Thailand and Indo-China to China, Formosa, Japan and Amurland, southward to tropical Australia; in Malesia: Sumatra, W. and Central Java, Philippines (Luzon, Mindanao), N. Celebes, New Guinea (Papua). Ecol. In open damp places, in swamps, and especially as a weed in wet rice-fields, at low and medium altitudes, up to 1800 m. Vern. Djukut mumundingan, S; Philippines: bo- book, lamlamsit, Bon., sirau-sirau, Ik. Note. The specimens of ‘F. aestivalis’ from the Malay Peninsula in the Singapore Herbarium belong all to F. griffithii (I have not seen RIDLEY 43 from Pahang). var. trichopoda KERN, Blumea 12 (1963) 27. Rays and raylets of the inflorescence densely pilose. Distr. India; in Malesia: NE. bank of Sepik River. var. macrostachya BENTH. FI. Austr. 7 (1878) 310 (with ?); CLARKE, Philip. J. Sc. 2 (1907) Bot. 94; Domin, Bibl. Bot., Heft 85 (1915) 458; Merr. En. Philip. 1 (1923) 121. Stronger in almost every part. Stems about 30 cm tall. Leaves 1-114 mm wide. Lower bract distinctly overtopping the inflorescence. Spikelets 2 mm wide. Glumes 2/4 by 114 mm. Stamens 2. Style UA mm long. Nut more distinctly reticulate, 4 by mm. Distr. Queensland; in Malesia: Philippines (Leyte), JAGoR 1008, according to CLARKE, I.c., not seen. Note. CLARKE, I.c., thinks that this variety tends to F. bisumbellata, but this is certainly not the case. From the characters given above I get the impression that it is a polyploid derived from F. aestivalis var. aestivalis. 64. Fimbristylis griffithii Boeck. Flora 43 (1860) 241; KERN, Blumea 8 (1955) 142; Reinwardtia 6 (1961) 48; in Back. & Bakh. f. Fl. Java 3 (1968) 467. — F. aestivalis B glaberrima Boeck. Linnaea 37 (1871) 11, p-p. (quoad specim. Indiae or.). — F. aestivalis (non VAHL) CLARKE, FI. Br. Ind. 6 (1893) 637, p.p.; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 92; Fl. Mal. Pen. 5 (1925) 155. — F. dichotoma (non VAHL) CAMUS, Fl. Gén. I.-C. 7 (1912) 102, p.p.—F. aestivalis f. glabra KUxK. Bot. Jahrb. 59 (1924) 49, ex descr. Glabrous annual with fibrous roots. Stems slender but rigid, densely tufted, compressed, 3—5S-angled, smooth, (3—)10—30 cm by 4- 1 mm, the base clothed with 1—2 tubular, bladeless or short- bladed, obliquely truncate, 2-3 cm long sheaths. Leaves shorter than to as long as the stems, often falcate, scabrid on the often involute margins, with pale, stramineous lower sheaths, 1-2 mm wide; ligule absent. Inflorescence compound or subdecompound, loose, with many spikelets, 5-8 cm long. Involucral bracts up to 6, filiform, more or less recurved, usually much shorter than the inflorescence, much dilated at the base, scabrid, 1-3 cm. Primary rays several, obliquely erect, compressed-angular, smooth, up to Scm. Spikelets solitary, ovoid or lanceolate, angular, acute, densely many- -flowered, stramineous or brownish, 3-7 by 1-1)4(-1'4) mm; rachilla narrowly winged. Glumes spiral, thinly membranous, triangular-ovate, acutish, sharply keeled, with prominent midnerve excurrent in a short, more or less excurved mucro, nerveless sides, and hyaline margins, 1 Je 1 “a by c. 1mm. Stamens 1(—2); anthers sions a JA mm. Style slender, flat, not hyaline-margined, glabrous, 344-1 mm; stigmas 2, shorter than the style. Nut biconvex, with acute edges, obovate or broadly New Guinea, on 1974] CYPERACEAE (Kern) 585 obovate, shortly stipitate, umbonulate, smooth, shining yellowish brown, obsoletely reticulate by the transversely uae not impressed epidermal cells, Ao by A-A mm. a Bengal, Burma, Thailand, Indo-China, Tenasserim and Andamans; in Malesia: Central Sumatra, Malay Peninsula (Perak, Kelantan, Treng- ganu, Pahang, Malacca, Singapore), W. Java, Borneo, Celebes (Central: Posso Lake; SE. Penin- sula: Kendari), Moluccas (W. Ceram), W. New Guinea. Ecol. Wet or swampy places, river-banks, edges of lakes, road-sides, at low altitudes, up to c. 700 m. Vern. Rumput sérai, r. sérai jantan, r. salah pemakai, r. képala lalat halus, r. janggut kuau, Mal. Pen., kanta jambun, S. and E. Borneo. Notes. BOECKELER’s type specimen (Bengal, GRIFFITH in B) got lost by war action; his description perfectly matches the species described above (neo- type: GRIFFITH 6331, K). A collection of this species in the Copenhagen Herbarium (KAMPHOEVENER 2785 from Sambelong) was annotated by BOECKELER “‘Fimbristylis (Eufimbr.) aestivalis (VAHL) B glaberrima = F. lirosa PoeEpp. ex KUNTH’. I have not seen the S. American F. limosa, which is apparently very close to, but probably not conspecific with F. griffithii, as KUNTH, En. 2 (1837) 225, ascribes obovate-pyriform, often ver- ruculose nuts, and purplish leaf-sheaths to it. Often confounded or united with F. aestivalis, which is, however, a dwarfer, weaker, hairy plant lacking the tubular leaf-sheaths surrounding the base of the stem. 65. Fimbristylis squarrosa VAHL, En. 2 (1806) 289: KUNTH, En. 2 (1837) 224; Sreup. Syn. 2 (1855) 110; Mia. FI. Ind. Bat. 3 (1856) 319, quoad descr.; BOECK. Linnaea 37 (1871) 10; CLarKE, FI. Br. Ind. 6 (1893) 635; Ill. Cyp. (1909) t. 41 f. 8-10; Camus, Fl. Gén. I.-C. 7 (1912) 101, f. 16, 3-6; Koyama, Contr. Inst. Bot. Un. Montréal n. 70 (1957) 44. — Pogonostylis squarrosa BERTOL. FI. Ital. (1833) 312. — F. aestivalis var. squarrosa KOYAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 116. var. squarrosa.— Synonymy as above. Not in Malesia; see Notes. var. esquarrosa MAKINO, Bot. Mag. Tokyo (1903) 47; KERN, Blumea 8 (1955) 143; Reinwardtia 6 (1961) 48; in Back. & Bakh. f. Fl. Java 3 (1968) 467. — F. velata R.BR. Prod. (1810) 227; Kuntu, En. 2 (1837) 243; BENTH. FI. Austr. 7 (1878) 309. — F. propinqua R.Br. Prod. (1810) 227; KuNTH, En. 2 (1837) 243. — Iriha velata O.K. Rev. Gen. Pl. 2 (1891) 753. — F. squarrosa var. velata CLARKE ex CHEESEM. Man. New Zeal. Fl. (1906) 770. — F. makinoana Ouw\, J. Jap. Bot. 14 (1938) 578; Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 83. — F. aestivalis var. esquarrosa Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 116. Annual with fibrous roots. Stems slender, densely tufted, setaceous, compressed-subtrigonous, smooth, 10-25cm by }4-1 mm. Leaves shorter than the stems, very narrow, flat or canaliculate, acute, more or less soft-hairy at least on the sheaths, 4-1 mm wide; ligule absent. Inflorescence compound or decompound, loose, with several to many spikelets, 2-5 cm long. Involucral bracts 3—7, dilated at the base, soft-hairy, the lowest shorter than to about as long as the inflorescence. Primary rays up to 6, slender, filiform, smooth, up to 2/4 cm. Spikelets solitary, lanceolate, angular, acute, rather densely many-flowered, stramineous or ferrugineous, 4—7 by 1-144 mm; rachilla narrowly winged. Glumes spiral, thinly membranous, suberect, ovate, or oblong-ovate, acutish, acutely keeled, 3-nerved, with nerveless sides, and a strong midnerve excurrent in a straight or slightly excurved, up to 4 mm long mucro, glabrous or pubescent, 14-2 by 1 mm. Stamen 1; anther oblong, 4—)4 mm. Style slender, flat, not hyaline-margined, dilated at the base, sparsely ciliate at the top, >4—1 mm, the lower margin fringed with a whorl of long, pendent hairs closely appressed to the nut and covering 14-%{ of it; stigmas 2, shorter than the style. Nut biconvex, with acute edges, obovate, shortly stipitate, minutely umbonulate, smooth, obscurely reticulate by the not impressed, hexagonal epidermal cells, shining stramineous, %—*% by 14 mm. Distr. F. squarrosa var. squarrosa is widely dis- tributed in the tropical and warm-temperate regions of Asia (also in Japan and Korea, but absent from Malesia), S. Europe, Africa, and S. America (not in N. America); var. esquarrosa is known from Thailand, Indo-China, NE. and E. China, S. Korea, Japan, Polynesia, Australia, and New Zealand; very rare in Malesia: Sumatra (Toba Lake), W. Java (Mt Patuha), Philippines (Luzon: Cagayan). Ecol. Shores of lakes, on wet sandy soil, on Mt Patuha at 1600 m, along Toba Lake at c. 900 m. Notes. KoyAMA (1957) thinks that the E. and S. China Seas draw a clear line of demarcation between the areas of the two varieties, but in E. Asia there is a considerable overlapping. The collections mentioned by MERRILL, En. Philip. 1 (1923) 126 under F. squarrosa belong partly to F. aestivalis, partly to F. merrillii. The only Philippine collection of F. squarrosa var. esquarrosa I have seen is CURRAN FB 16783, distributed as F. aestivalis. F. squarrosa is intimately related to F. aestivalis, with which F. voN MULLER, Fragm. 9 (1875) united it. KOYAMA, 1961, followed voN MULLER, possibly with good reason. For, although typical F. squarrosa is clearly distinct from F. aestivalis by the much longer, strongly recurved mucros of the glumes giving the spikelets a squarrose aspect, and by the long pendent hairs of the style-base (taken by NEES for fibres of the pericarp !), the mucros of var. esquarrosa do not differ from those of F. aestivalis. So the trichomes of the style-base (absent or very short in F. aestivalis), the slightly longer style and the often somewhat larger nut are the only characters to distinguish between F. squarrosa var. esquarrosa and F. aestivalis. Anyway, by assigning specific rank to var. esquarrosa (F. velata), its taxonomic value is over-estimated. According to TANAKA the chromo- some number in var. squarrosa is n = 10, in var. esquarrosa n = 12. See also KERN, 1955 and 1961. 66. Fimbristylis gracilenta HANcE, J. Bot. Lond. 6 (1868) 89; CLARKE, J. Linn. Soc. Bot. 36 (1903) 237; Camus, Fl. Gén. I.-C. 7 (1912) 103; Kern, Blumea 8 (1955) 144; ibid. 10 (1960) 647: Reinwardtia 6 586 FLORA MALESIANA [ser. I, vol. 7° (1961) 49. — F. thorelii Camus, Not. Syst. 1 (1910) 246; Fl. Gén. I.-C. 7 (1912) 105. Annual with fibrous roots. Stems very slender, setaceous, tufted, slightly compressed, obtusangular, glabrous, or hairy at the top, 15-30 cm by 4-34 mm. Leaves somewhat shorter to longer than the stems, filiform, involute, abruptly acuminate, soft-hairy also on the sheaths, 4-1/4 mm wide; ligule absent. Inflorescence compound, very loose, with several to many spikelets, narrow, 3—10 cm long. Involucral bracts 2—3, erect, the lowest up to twice as long as the inflorescence, 6—12 cm. Primary rays 5—7, very un- equal, suberect, filiform, slightly compressed, pilose (like the short, suberect secondary rays), rarely glabrous (see Notes), up to 7 cm. Spikelets solitary, (narrowly) lanceolate, angular, acute, loosely many- flowered, dull brown, 3-8 by 1-1/4 mm; rachilla distinctly winged. Glumes spiral (see Notes), mem- branous, oblong-ovate, acutish, mucronate, keeled, glabrous, with 3—S-nerved keel (midnerve prominent) and nerveless sides, 2-2/4 by Lye 1/4 mm. Stamens 1(—2); anthers oblong- linear, 4 mm. Style slender, flat, not hyaline-margined, slightly dilated at the base, glabrous or sparsely ciliate, 1-1/4 mm; stigmas 2, shorter than the style. Nut biconvex, with acute, obscurely costulate edges, obovate, rounded or sub- truncate at the apex, smooth or sparsely white-scaly, shortly stipitate, minutely umbonulate, obscurely reticulate by the hexagonal or transversely elliptic, not impressed epidermal cells, shining, whitish to pale brown, 4-%, by c. 14 mm. Distr. S. China, Thailand, Indo-China; in Malesia: N. Sumatra (between Aek Bila and Aek Marbau; Rantau Parapat, Bila). Ecol. Grassy places, forest-clearings, altitudes. Notes. The rays and raylets of the inflorescence are generally pilose, a rare feature in Fimbristylis; in var. psilopoda KERN, Reinwardtia 6 (1961) 49, only known from NE. Thailand, they are glabrous. In a collection from Central Thailand (SMITINAND 6084) the upper glumes of the spikelets are regularly 2-ranked. See also the note under 57. F. perlaxa. at low 67. Fimbristylis argentea (ROTTB.) VAHL, En. 2 (1806) 294; KUNTH, En. 2 (1837) 223; STEUD. Syn. 2 (1855) 108; Mig. Fl. Ind. Bat. 3 (1856) 317; Boeck. Linnaea 37 (1871) 8; CLARKE, FI. Br. Ind. 6 (1893) 640; RipL. Mat. Fi. Mal. Pen. (Monoc.) 3 (1907) 93; Camus, Fl. Gén. I.-C. 7 (1912) 110, f. 15; 1; Ripe. Fl; Mal: Pen. 5 (1925) 157; KERN, Reinwardtia 6 (1961) 49. — Scirpus argenteus Rotts. Progr. (1772) 27; Descr. et Ic. (1773) 51, t. 17 f. 6. — Scirpus monander Rotts. Descr. et Ic. (1773) 50, t. 14 f. 3. — Iriha monandra O.K. Rev. Gen. PI. 2 (1891) 753. Glabrous annual with fibrous roots. Stems densely tufted, setaceous, trigonous, slightly compressed, smooth, 3—10(— 20) )cm by 4-34 mm. Leaves usually shorter than the stems, flat or canaliculate, rather abruptly acuminate, smooth or slightly scabrid at the top, glaucous, with stramineous sheaths, )4— -% mm wide; ligule absent. Inflorescence capitate, semi- globose to globose, with (2—)4—10(—25) spikelets, 44-1 cm across. Involucral bracts 2-4, somewhat dilated at the base, finally patent to reflexed, the lowest much longer than the inflorescence. Spikelets sessile, oblong-ovoid or cylindrical, slightly angular, acutish, very densely many-flowered, 4-10 by 1/4- 14 mm; rachilla narrowly winged. Glumes spiral, membranous, broadly ovate-deltoid, obtusish, mutic- ous or apiculate, slightly keeled, 3-nerved, with strong midnerve, and nerveless, silvery grey, fer- rugineous or fuscescent sides, c. 1 by 1 mm. Stamen 1; anther oblong, )4 mm. Sty le flat, not hyaline- margined, dilated at the base, minutely ciliolate in the upper part, 34 mm long; stigmas 2, somewhat shorter than the style. Nut biconvex, with acute edges, broadly obovate or suborbicular, shortly stipitate, minutely umbonulate, smooth or sparsely verruculose, obscurely reticulate by the transversely elliptic or oblong, not impressed epidermal cells, V4 by 4-4 mm, Distr. S. and SE. Asia: Ceylon, India, Bengal, Peninsular Thailand; in Malesia only known from a few localities in the Malay Peninsula (Kelantan: Kamposa; Trengganu: Kuala Trengganu). Ecol. In wet or swampy places, in the Malay Peninsula in damp hollows on the sea-shore. 14. Section Neodichelostylis Camus, Fl. Gén. I.-C. 7 (1912) 89. — Fimbristylis sect. Ferrugineae OHWI, J. Jap. Bot. 14(1938) 573, p.p. — Fimbristylis ser. Ferrugineae OHW1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55, p.p Type species: F. polytrichoides (RETZ.) R.Br. (Scirpus polytrichoides RETZ.). 68. Fimbristylis polytrichoides (RETZ.) R.BR. Prod. (1810) 226; KuntH, En. 2 (1837) 221; STEuD. Syn. 2 (1855) 106; Mia. Fl. Ind. Bat. 3 (1856) 315; BENTH. Fl. Austr. 7 (1878) 304; CLARKE, FI. Br. Ind. 6 (1893) 632; Bot. Tidsskr. 24 (1901) 88: RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 90, p.p.; CLARKE, Philip. J. Sc. 2 (1907) Bot. 92; Ill. Cyp. (1909) t. 40 f. 8-9: Koorp. Exk. Fl. Java 1 (1911) 198, non ibid. 4, Atlas (1922) f. 250; Camus, Fl. Gén. I.-C. 7 (1912) 96, f. 13,9; MerR. En. Philip. 1 (1923) 125; Rip. Fl. Mal. Pen. 5 (1925) 154, p.p.; BAcK. Onkr. Suiker. (1928) 159, t. 164; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 20; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 465. — Scirpus polytrichoides Retz. Obs. 4 (1786) 11; VAHL, En. 2 (1806) 248 (‘polythrichi- oides’). — Abildgaardia javanica StEuD. [in ZOLL. Syst. Verz. 1 (1854) 63 (‘Abilgardia’), nom. nud.] Syn. 2 (1855) 72; Mia. Fl. Ind. Bat. 3 (1856) 297; non A. javana NEES, 1834. — F. albescens STEUD. Syn. 2 (1855) 107; Mig. Fl. Ind. Bat. 3 (1856) 316. — F. subbulbosa Boeck. Flora 41 (1858) 598; Cyp. nov. 2 (1890) 39. — F. juncea (non R. & S.) Boeck. Linnaea 37 (1871) 4. — Iriha polytrichodes O.K. Rev. Gen. PEE SO 7as- Perennial (always?), forming dense tufts, glabrous (except for the ligule), glaucescent. Stems erect or curved, setaceous, sulcate, smooth, incrassate at the base, 5-30 cm by up to 1 mm. Leaves half as long to 1974] CYPERACEAE (Kern) 587 as long as the stems, setaceous, wiry, obtusish, complicate-canaliculate, smooth, with obliquely trun- cate, ferrugineous sheaths, /4—1 mm wide; ligule a row of short hairs. Inflorescence often consisting of a single, terminal or pseudo-lateral spikelet, more rarely 1 or 2 lateral spikelets added. Spikelets erect, solitary, ovoid-ellipsoid or pblong, terete, acute, densely many-flowered, pale brown, 5—15 by 2-3 mm, the subtending bract sometimes glume-like, but often with an erect blade as long as or slightly overtopping the inflorescence; rachilla narrowly winged. Glumes spiral, densely imbricate, thinly membranous, oblong- ovate or oblong, more than twice as long as broad, obtuse, scarcely keeled, muticous or apiculate with green midnerve, otherwise nerveless, 2?4-3 by 1- 114 mm; lowest 2 sterile glumes acute, often mucro- nate. Stamens 1-2; anthers linear. “4-1 mm. Style slender, flat, not hyaline-margined, slightly dilated at the base, sparsely ciliate at the top, 4-1 mm; stigmas 2, about as long as or longer than the style. Nut biconvex, with acute angles, cuneate-obovate or oblong-obovate, shortly stipitate, rounded at the apex (not umbonulate), smooth or verruculose, delicately reticulate by the minute, transversely elliptic or oblong, not impressed epidermal cells in c. 30 vertical rows on either face, greyish or blackish brown, %-1){, by 4-K(- 4%) mm. Distr. Tropical Africa, Asia, and Australia; in Malesia: Malay Peninsula (Kedah: P. Langkawi; Perak: Lumut Dindings; Pahang; Selangor; P. Penang), Sumatra, Java (along N. coast), Madura, Philippines (Luzon, Panay), New Guinea (Merauke, Port Moresby). Ecol. In open wet places, especially on sandy or muddy sea-shores and in rock-crevices near the sea, sometimes more inland on saline soil. Vern. Rébha kaproléam, Md. 69. Fimbristylis tenuicula Boeck. Linnaea 38 (1874) 385; CLARKE, Fl. Br. Ind. 6 (1893) 632. — Iriha tenuicula O.K. Rev. Gen. PI. 2 (1891) 753. Probably annual, with fibrous roots, growing in dense tufts. Stems erect or curved, setaceous or capillary, sulcate, smooth, 10-25 cm by c. )4 mm. Leaves much shorter than the stems, flattish but very narrow, weak, obtusish, smooth, glabrous or sparsely hairy, with obliquely truncate, cinnamomeous, hairy sheaths, /4—/4 mm wide; ligule absent. Inflorescence always consisting of a single, terminal spikelet. Spikelet erect, ebracteate, lanceolate or oblong- lanceolate, terete, acute, densely many-flowered, pale brown, 5-10 by 14 2mm. Glumes spiral, densely imbricate, thinly membranous, oblong-ovate or oblong, twice or more than twice as long as broad, scarcely keeled, minutely but distinctly mucronulate, ferrugineous, with green midnerve, otherwise nerve- less, glabrous, 2-234 mm long. Stamen 1; anther oblong- linear, “4-1 mm. Style slender, flat, not hyaline- -margined, dilated at the base, sparsely ciliate, 1-1/4 mm; stigmas 2, much shorter than the style. Nut biconvex, with acute angles, obovate, shortly stipitate, rounded at the apex (not um- bonulate), sparsely verruculose, reticulate-trabeculate by the transversely oblong, somewhat impressed epidermal cells in c. 15 vertical rows on either face, brown, 74-1 by %-* mm. Distr. Only known from a few localities in India, Tenasserim, Indo-China (Haut Dormai: near Blao):; in Malesia: S. Sumatra (S of Palembang, along Lake Pedammaran). Ecol. In Sumatra along shore of lake, at low altitude; in Indo-China in a swamp at c. 850 m. Notes. The collection Reliquiae Helferianae 146, distributed by the National Museum in Prague as F. polytrichoides, belongs here, and is in all proba- bility not from Calcutta, but part of HELFER’s type- collection from Tenasserim. In the Leyden Herbarium there is also an excellent collection of this species, inadequately labelled “Arch. Ind. leg. WaITz.” 70. Fimbristylis celebica OHW1, Blumea 8 (1955) 103, f. 6. — Fig. 48. Glabrous annual forming dense tufts. Stems setaceous, striate, slightly compressed, smooth, 5—10 by 4mm. Leaves about as long as the stems or somewhat longer, filiform, curved, very acute, smooth except for the scabrid top, with membranous, stramineous or brownish sheaths, 14—-!4 mm wide. Inflorescence always consisting of a single terminal spikelet. Spikelet erect, oblanceolate or narrowly oblong, angular, acute, few- or several-flowered, pale, fuscous-variegated, 5-7 by 2mm, usually ebracteate but sometimes subtended by an up to 1 cm long bract. Glumes spiral, thinly membranous, erect, oblong-lanceolate, acutish, sharply keeled, shortly mucronulate, with 3—5-nerved keel, whitish stramineous, fuscous-variegated, nerveless sides and hyaline margins, 3 by 1mm. Stamens (1—)2-3: anthers linear, 1-1/4 mm. Style very slender, flat, not hyaline- -margined, scarcely dilated at the base, glabrous, 1-1/4 mm; stigmas 2, somewhat shorter than the style. Nut biconvex, with acute edges, narrowly obovate, shortly stipitate, minutely um- bonulate, smooth, obscurely reticulate by the minute, not impressed epidermal cells, whitish, 7{,—*% by 4mm. Distr. Malesia: Celebes (Lake Tuwuti, Posso, Lake Matano). Ecol. On shores of lakes and in Imperata-fields, 300-400 m. Lake 71. Fimbristylis wetarensis OHwi, Blumea 8 (1955) HOSS ied. Densely tufted perennial. Stems erect, very slender, sulcate-angular, compressed, pubescent or glabres- cent, 30—45 cm by 14-7 mm. Leaves much shorter than the stems, rather firm, flat, abruptly acuminate, pubescent especially beneath, scabrid on the margins, with scarcely keeled, pubescent sheaths, 1-1/4 mm wide; ligule absent. Inflorescence consisting of a single terminal spikelet. Spikelet erect or slightly oblique, ebracteate, lanceolate, terete, acute, densely many-flowered, 7-15 (ultimately up to 20) by 3—4 mm; rachilla narrowly winged. Glumes spiral, appressed, membranous, ovate or oblong-ovate, acutish, mucronulate, scarcely keeled, i-nerved, glabrous, shining brown in the upper half. much paler below, hyaline-margined, c. 5 by 3 mm. Stamens 3; anthers linear, 2 mm. Style flat, scarcely dilated at the shortly setulose base, densely ciliate in the upper half, 4-414 mm; stigmas 2, somewhat shorter than the style. Nut biconvex, with acute edges, obtriangular-obovate, obtuse or subtruncate, shortly stipitate, scarcely umbonulate, densely verruculose 588 FLORA MALESIANA [ser. I, voli 7? Fig. 48. Fimbristylis celebica OHW1. a. Habit, x %, b. spikelet, c. two glumes, both x 8, d. anther, e. deflorate flower, f. nut, all x 16 (a—f Eyma 4113). all over, obscurely reticulate by the small, roundish or hexagonal epidermal cells, whitish or stramineous, VK by % mm. Distr. Malesia: Lesser Sunda Is. (Wetar, along the Meta Lerai). Ecol. In Eucalyptus savannah above the rain- forest bordering the river, 700—900 m. Note. Remote from the other spp. of the section; only known from the type collection, but hardly endemic in Wetar. 15. Section Nutantes Ouwi, J. Jap. Bot. 14 (1938) 573. — Fimbristylis ser. Nutantes OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55. Type species: F. nutans (RETZ.) VAHL (Scirpus nutans RETZ.). 72. Fimbristylis acuminata VAHL, En. 2 (1806) 285; KUNTH, En. 2 (1837) 221; Nes in Hook. J. Bot. Kew Misc. 6 (1854) 29; Steup. Syn. 2 (1855) 106; Mia. FI. Ind. Bat. 3 (1856) 314, excl. var. B; Boeck. Linnaea 37 (1871) 3; CLARKE, FI. Br. Ind. 6 (1893) 631; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 89; CLARKE, Philip. J. Sc. 2 (1907) Bot. 91, excl. AHERN 3390; Ill. Cyp. (1909) t. 40 f. 5-7; Koorp. Exk. Fl. Java 1 (1911) 198; ibid. 4, Atlas (1922) f. 251; Camus, FI. Gén. I.-C. 7 (1912) 95, f. 13, 8; MERR. En. Philip. 1 (1923) 121; Ripv. Fl. Mal. Pen. 5 (1925) 153; BAcK. Onkr. Suiker. (1928) 158, t. 163; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 19; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 464. — F. rhyticarya F.v.M. Fragm. 1 (1859) 215; BenTtH. FI. Austr. 7 (1878) 302. — Iriha acuminata O.K. Rev. Gen. Pl. 2 (1891) 751. — F. setacea (non BENTH.) CLARKE, FI. Br. Ind. 6 (1893) 632, p.p.; J. Linn. Soc. Bot. 34 (1898) 54, p.p.: Rip. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 89; Fl. Mal. Pen. 5 (1925) 153. — F. nutans var. minor Camus, FI. Gén. I.-C. 7 (1912) 96. Glabrous perennial with very short rhizome. Stems slender, erect, densely tufted, obtusely 3—4-angled, sulcate, smooth, 5-30cm by /4-1 mm. Leaves reduced to obliquely truncate, bladeless (or in the uppermost short-bladed) sheaths. Inflorescence con- sisting of a single terminal spikelet. Spikelet erect or slightly oblique, ebracteate, ovoid or lanceolate, terete, acute, many-flowered, 6-12 by 2-314 mm; rachilla narrowly winged. Glumes spiral, chartaceous, ovate, obtuse or acutish, scarcely mucronulate, slightly keeled, stramineous to brownish, with green, darker lineolate, 3—5(—7)-nerved keel, 3/4-4'%4 by 24-4 mm. Stamens 2-3; anthers oblong-linear or linear, 4-1/4 mm. Style flat, scarcely dilated at the base, ciliate in the upper half, 2-3)4 by 4-74 mm; stigmas 2, much shorter than the style. Nut biconvex, with obtuse edges, broadly obovate or orbicular, 1974] CYPERACEAE (Kern) 589 sometimes somewhat broader than long, with very short, broad stipe, not umbonulate, with 5—8 strong transverse wavy ridges, obscurely reticulate by the minute, not impressed, hexagonal epidermal cells, dirty stramineous or brownish, rarely dark brown, 14-13% mm long and wide. Distr. SE. Asia (from Ceylon and India to S. China), tropical Australia; in Malesia: Sumatra and adjacent islands P. Simalur, Mentawei Is., Banka, P. Lingga), Malay Peninsula, Java, Madura, Tanimbar Is., Borneo (W. Borneo, N. Borneo, Sarawak), Philippines (Luzon), New Guinea (W. New Guinea: Kurik near Merauke; Papua: Morehead R.). Ecol. In open wet and muddy places: swamps, rice-fields, river-banks, at low altitudes, up to 350 m. Vern. Balihin bu udeung, Simalur, rumput janggot keli, Mal. Pen.; Philippines: puydéng-usa, Tag., suru- sibuyas, Bik. 73. Fimbristylis nutans (RETZ.) VAHL, En. 2 (1806) 285; KunrtH, En. 2 (1837) 221; StEuD. Syn. 2 (1855) 106; Boeck. Linnaea 37 (1871) 5; BENTH. FI. Austr. 7 (1878) 303; CLARKE, FI. Br. Ind. 6 (1893) 632; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 89: Koorp. Exk. Fl. Java 1 (1911) 198; Camus, Fl. Gén. I.-C. 7 (1912) 96, f. 15, 10-11; Ripe. Fl. Mal. Pen. 5 (1925) 154; S. T. Brake, J. Arn. Arb. 35 (1954) 208. — Scirpus nutans Retz. Obs. 4 (1786) 12.— Iriha nutans O.K. Rev. Gen. PI. 2 (1891) 753. Glabrous perennial with very short rhizome. Stems slender, densely tufted, obtusely 3—4-angled, sulcate, smooth, 15—40(—70) cm by 4-1 mm. Leaves reduced to tubular, bladeless or short-bladed, stramineous or ferrugineous sheaths. Inflorescence consisting of a single terminal spikelet. Spikelet more or less nodding, ebracteate, ovoid or broadly ovoid, terete, obtuse or acutish, densely many-flowered, stramineous and castaneous variegated, 5-15 by 3—5 mm; rachilla narrowly winged. Glumes spiral, chartaceous, broadly ovate, very obtuse, apiculate, scarcely keeled, 3— 5(-9)-nerved in the centre, with shining brown, nerveless sides, 314-414 by 3-3/4 mm. Stamens 3; anthers linear, 114-2 mm. Style flat, relatively broad, liguliform, not or scarcely dilated at the base, ciliate in the upper half, 24-314 by 4-74, mm; stigmas 2, much shorter than the style. Nut biconvex or almost flat on the inner side, with obtuse edges, obovate, shortly and broadly stipitate, not or scarcely umbonulate, tuberculate towards the margins or throughout, with 3—S(—6) transverse wavy ridges, obscurely reticulate by the minute, hexagonal epi- dermal cells, whitish, 14-174 by 1-1/4 mm. Distr. From Ceylon and India through Farther India to §. China, Formosa, and the Ryu Kyu Is., southwards to tropical Australia;in Malesia: Sumatra (E. Coast Res.; Palembang), Malay Peninsula (Kelantan, Trengganu, Malacca, P. Besar, Johore, Singapore), Borneo (Sarawak, W. Borneo, N. Borneo, Labuan), New Guinea. Ecol. Damp sandy spots, moist places in forests, wet grass-plains, savannahs, wet rice-fields, swamps, also on brackish soil, at low altitudes (0Q—70 m). Vern. Rumput buntil, Borneo. 74. Fimbristylis acicularis R.BR. Prod. (1810) 226; BENTH. Fl. Austr. 7 (1878) 301; KERN, Blumea 8 (1955) 148. — Gramen polytrichum RumpH. Herb. Amb. 6 (1750) 17, t. 7 f. 1. — F. setacea BENTH. in Hook. Lond. J. Bot. 2 (1843) 239: StEuD. Syn. 2 (1855) 106; CLARKE, Fl. Br. Ind. 6 (1893) 632, p.p., incl. var. setacea CLARKE; Philip. J. Sc. 2 (1907) Bot. 91; Koorp. Exk. Fl. Java 1 (1911) 198; VALCK. Sur. Nova Guinea 8 (1912) 702: Camus, Fl. Gén. I.-C. 7 (1912) 96; MerR. En. Philip. 1 (1923) 126:S.T. BLAKE, J. Arn. Arb. 35 (1954) 208.— F. acuminata var. pumila Nees in Hook. J. Bot. Kew Misc. 6 (1854) 29. — Abildgaardia brevifolia StEUD. Syn. 2 (1855) 72; Mig. Fl. Ind. Bat. 3 (1856) 297. — Isolepis cochleata STEUD. Syn. 2 (1855) 100. — F. acuminata var. setacea Mia. FI. Ind. Bat. 3 (1856) 314; BENTH. Fl. Austr. 7 (1878) 301; KUk. Bot. Jahrb. 59 (1924) 47; ibid. 69 (1938) 257. — F. acuminata var. minor NEES ex Boeck. Linnaea 37 (1871) 4; RoLre, J. Bot. 24 (1886) 59 in nota (‘F. acutifolia var. minor’, sphalm.). — F. australica Boeck. Linnaea 38 (1874) 384, fide BENTH. — F. bursifolia (sphalm.) VIDAL, Phan. Cum- ing. (1885) 156; Rev. Pl. Vasc. Filip. (1886) 284. — Iriha acicularis O.K. Rev. Gen. Pl. 2 (1891) 753. Closely related to F. acuminata. Stems very slender, setaceous, spreading to decumbent, 5—20cm by 44mm. Leaves often all reduced to sheaths, but frequently some with well developed, filiform, acute, 44 mm wide blades scabrid on the margins near the top. Spikelet narrowly lanceolate, pale green, 4—6 by 1-1/4(—2) mm. Glumes membranous, the lower ones oblong, the upper ones oblong-ovate. Stamens 1-2; anthers linear, c. 1/4 mm. Style very slender, dis- tinctly dilated at the base, c. 2/4 mm long, 4) mm wide. Nut obovate, white or pale stramineous, Y-Aol-1) by Z-ZA(-/A) mm. Distr. Tropical Australia, Bougainville; in Malesia: Philippines (Luzon), S. Celebes (Makassar): Moluccas (Halmaheira, Buru, Amboina), New Guinea, Aru Is. Ecol. In open wet localities: edges of swamps, wet grass-fields, damp places in savannah-forests, trails in secondary forests, also on the sea-shore, at low altitudes, rarely up to 800 m. Notes. According to BENTHAM F. acicularis and F. setacea differ in the style (glabrous in acicularis, ciliate in setacea) and the number of stamens (1 and 3 respectively). In the type collection of F. acicularis the style is ciliate (“‘stylo basi dilatata nuda,” R. BROwN), and in both F. acicularis and F. setacea there are | or 2 stamens. I have not seen SCHULTZ 79, type collection of F. australica BOECK., in which the style is said to be glabrous. In the upper flowers of the spikelet I frequently find the upper flowers female, without a trace of stamens; cf. F. androgyna R.BR. 16. Section Mischospora (BoEcK.) Camus, Fl. Gén. I.-C. 7 (1912) 89. — Mischospora Borck. Flora 43 (1860) 113. — Fimbristylis sect. Tetragonae OHWI, J. Jap. Bot. 14 (1938) 571. Type species: Mischospora efoliata BOECK. (F. tetragona R.BR.) 590 FLORA MALESIANA [ser. I, vole 75. Fimbristylis tetragona R.BR. Prod. (1810) 226; KuntH, En. 2 (1837) 242; SreuD. Syn. 2 (1855) 108; BENTH. FI Austr. 7 (1878) 305; CLARKE, FI. Br. Ind. 6 (1893) 631; Philip. J. Sc. 2 (1907) Bot. 91; RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 88; CLARKE, IIl. Cyp. (1909) t. 40 f. 1-4; Koorp. Exk. Fl. Java 1 (1911) 198: Camus, Fl.) Gen. T.-C. 7 (1912) 93, f- 13, 6-7; Merr. En. Philip. 1 (1923) 127; Rw. Fl. Mal. Pen. 5 (1925) 153; BACK. Onkr. Suiker. (1928) 158, t. 162; KUK. Mitt. Thiir. Bot. Ver. N.F. 50 (1943) 8; OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 55; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 19; S. T. BLAKE, J. Arn. Arb. 35 (1954) 208; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 462. — F. cylindrocarpa KUNTH, En. 2 (1837) 222; STEUD. Syn. 2 (1855) 107; Kurz, J. As. Soc. Beng. 39, ii (1870) 85; Boeck. Linnaea 37 (1871) 7. — F. abjiciens Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.] Syn. 2 (1855) 107; Mia. Fl. Ind. Bat. 3 (1856) 316. — Mischospora efoliata Boeck. Flora 43 (1860) 113. — F. oxyrhachis Miq. Sum. (1861) 601. — F. arnottii THWAITES, En. Pl. Zeyl. (1864) 348. — Iriha tetragona O.K. Rev. Gen. Pl. 2 (1891) 753. Glabrous, glaucous perennial with short rhizome, or annual in unfavourable circumstances. Stems densely tufted, erect, rigid, quadrangular, strongly ribbed, smooth, 10-60 cm by 74-1 mm. Leaves of the flowering stems reduced to 2—3 obliquely truncate, membranous, ferrugineous or fuscous, 3—10 cm long sheaths, bladeless or the uppermost with a short, lanceolate blade; those of the sterile shoots filiform, c. 4mm wide. Inflorescence consisting of a single terminal spikelet. Spikelet erect, ebracteate, globose, ovoid or oblong-ovoid, terete, obtuse, very densely many-flowered, dull brown, 6-15 by 4-6 mm; rachilla ragged by narrow wings. Glumes spiral, closely imbricate, membranous, oblong-ovate or oblong, very obtuse, with rounded apex, muticous, concave, faintly many-nerved (3 central nerves more prominent), 3—5 by 2-3 mm. Stamens 1-3; anthers linear, 114-114 mm. Style subpersistent on the nut, flat, slightly dilated at the base which is nearly as wide as the nut, sparsely ciliate, dark brown, 2—3 mm; stig- mas 2 or 3, much shorter than the style. Nut plano- convex or subterete, oblong-cylindrical, slightly curved, conspicuously stipitate, trabeculate by the hexagonal or transversely elliptic, slightly impressed epidermal cells in c. 9 vertical rows on either face, stramineous, 114-2 by 74 mm; gynophore (stipe) spongious, (4—)!4—1 mm long. Distr. From Ceylon and India through Thailand and Indo-China to S. China, Formosa, and tropical Australia; in Malesia: N. Sumatra, Banka, Malay Peninsula (very local: Kuala Pahang, Singapore), W. and Central Java, Madura, Kangean, Lesser Sunda Is. (Lombok, Sumba), N. Borneo (Jesselton, Kuching), Philippines (Luzon), S. Celebes, New Guinea (very local). Ecol. On heavy soil in open, wet places: swamps, swampy grasslands, wet rice-fields, at low altitudes (usually below 300 m, in Sumba at 500 m, in Sumatra up to 1000 m). Vern. Komis, Md. 17. Section Dipsaceae Ouwi, J. Jap. Bot. 14 (1938) 571. — Echinolytrum Desv. J. Bot. 1 (1808) 20. — Fimbristylis sect. Echinolytrum OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 53. — Fimbristylis ser. Echinolytrum Koyama J. Fac. Sc. Un. Tokyo III, 8 (1961) 101. Type species: F. dipsacea (ROTTB.) CLARKE (Scirpus dipsaceus ROTTB.). 76. Fimbristylis dipsacea (ROTTB.) CLARKE, FI. Br. Ind. 6 (1893) 635; Philip. J. Sc. 2 (1907) Bot. 93; Il. Cyp. (1909) t. 41 f. 4-7; Merr. En. Philip. 1 (1923) 123; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 19: S. T. BLAKE, J. Arn. Arb. 35 (1954) 214; KERN, Blumea 9 (1958) 234; Koyama, J. Fac. Sc. Un. Tokyo III, 8 (1961) 118, incl. var. verrucifera KoyAMA; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 462. — Scirpus dipsaceus Rotts. Descr. & Ic. (1773) 56, t. 12 f. 1; VAHL, En. 2 (1806) 276; Boeck. Linnaea 36 (1870) 736; F.v.M. Descr. Not. 2 (1886) 35. — Echinolytrum dipsaceum Desv. J. Bot. 1 (1808) 21, t. 1; Nees in Wight, Contr. (1834) 96; Camus, FI. Gén. I.-C. 7 (1912) 130, f. 17, 3-5. — Isolepis dipsacea R. & S. Syst. 2 (1811) 119; Kuntn, En. 2 (1837) 205; STEUD. Syn. 2 (1855) 99; Mia. FI. Ind. Bat. 3 (1856) 309. — Isolepis verrucifera MAxiM. Prim. Fl]. Amur. (1859) 300. — F. verrucifera MAKktNo, Bot. Mag. Tokyo 9 (1895) 259; Ouwi, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 56. Glabrous annual with fibrous roots. Stems very slender, densely tufted, setaceous, angular, smooth, (2-)5—15(-25) cm by 4-)4 mm. Leaves short, fili- form, smooth, or scabrid at the top, 4—)4 mm wide: ligule absent; sheaths obliquely truncate, the lower ones bladeless or short-bladed, brownish. Inflo- rescence simple or subcompound, umbelliform, with up to 15 spikelets, but not rarely much reduced. Involucral bracts 3—5, shorter to longer than the inflorescence, dilated at the base. Primary rays up to 10, smooth, up to 2 cm. Spikelets solitary, subglobose or oblong, terete, obtuse, squarrose, very densely many-flowered, pale green, 3-6 by 2—3 mm; ra- chilla densely ragged by minute wings. Glumes spiral, thinly membranous, spreading, oblong or lanceolate, keeled, the blade c. 1 by 74 mm, with hyaline, obscurely 1—2-nerved sides and a strong green midrib excurrent into a /4—1 mm long recurved awn slightly scabrid at the top. Stamen 1; anther oblong, 4 mm. Style glabrous, slightly dilated at the base, ferrugineous, 14 mm long; stigmas 2, about as long as the style. Nut scarcely compressed, subterete, oblong-linear, very rarely narrowly obovoid, slightly curved, obscurely transversely lineolate by the minute, hexagonal or elliptic epidermal cells, fer- rugineous or brown, 14-34 by ){ mm (or, when obovoid, 7% by 14 mm), on either margin ornamented with a row of clavate appendages (glands?), which often fall off with age. Distr. Tropical Africa, S. and E. Asia; in Malesia very rare: S. Sumatra (near Palembang), Madura, Bawean, E. Borneo (Kutei), Philippines (Luzon, 1974] Cy PERACEAE (Kern) 3on Mindanao), Central Celebes (Lake Posso), New Guinea (Papua, W. Division). Ecol. Muddy lake-shores, river-banks, wet rice- fields, at low altitudes. Along Kastoba Lake (Bawean I.) forming a dense mat in association with F. aestivalis. Notes. Specimens with reduced inflorescence may be mistaken for Lipocarpha microcephala, which shares the squarrose spikelets and oblong-linear nuts with it. In Lipocarpha, however, the nuts are minutely punctulate (not transversely lineolate), never beset with gland-like appendages, and enclosed in two thinly membranous scales. DesvAUX (1808) took the pericarp for a utricle surrounding the fruit (like in Carex); on this supposed feature the genus Echinolytrum was based. The specimens collected in Borneo deviate from the other Malesian materials by their narrowly obovoid nuts; similar fruits I have seen in Indian plants. F. verrucifera is said to differ from F. dipsacea by the shorter and less recurved awns of the glumes and hence less squarrose spikelets. Similar plants occur in Africa; probably they even do not deserve varietal rank. 18. Section Actinoschoenus (BENTH.) KERN, Blumea 8 (1955) 160. — Actinoschoenus BENTH. in B. & H. Gen. PI. 3 (1883) 1058. — Arthrostylis subg. Actinoschoenus KUK. in Fedde, Rep. 53 (1944) 197. Type species: F. thouarsii (KUNTH) MERR. (Arthrostylis thouarsii KUNTH). 77. Fimbristylis thouarsii (KUNTH) Mer. En. Philip. 1 (1923) 127; KERN, Reinwardtia 6 (1961) 44. — Arthrostylis thouarsii KUNTH, En. 2 (1837) 284; MQ. Fl. Ind. Bat. 3 (1856) 335; KUx. in Fedde, Rep. 53 (1944) 199.— Arthrostylis chinensis BENTH. FI. Hongk. (1861) 397; KUk. in Fedde, Rep. 53 (1944) 198, incl. var. filiformis KUKk.—Arthrostylis _fili- formis THwaITEs, En. Pl. Zeyl. (1864) 352; BoEck. Linnaea 37 (1873) 524. — Actinoschoenus filiformis BENTH. in Hook. Ic. Pl. 14 (1881) 33, t. 1346; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 84; Fl. Mal. Pen. 5 (1925) 165, incl. var. rupestris RIDL. — Actino- schoenus chinensis BENTH. in Hook. Ic. Pl. 14 (1881) 33, t. 1346. — Actinoschoenus thouarsii BENTH. l.c.; CHERMEZ. in Humbert, Fl. Madag., fam. 29 (1937) 224, t. 20, f. 8-10. — F. actinoschoenus CLARKE, FI. Br. Ind. 6 (1893) 650, incl. var. chinensis CLARKE, Philip. J. Sc. 2 (1907) Bot. 98. — F. actinoschoenus var. thouarsii CLARKE, Ill. Cyp. (1909) t. 44 f. 5—10. — Actinoschoenus filiformis RipL. J. Fed. Mal. St. Mus. 6 (1915) 192.— F. filiformis H. PFEIFF. in Fedde, Rep. 29 (1931) 183, non KuNTH, 1837. — F. chinensis TANG & WANG, FI. Reip. Pop. Sin. 11 (1961) 106. Glabrous perennial with short rhizome. Stems very densely tufted, setaceous, trigonous, sulcate, smooth, (15—)30-75cm by 34-1 mm. Leaves re- duced to 2-3 bladeless or short-bladed sheaths, obliquely truncate, glabrous, stramineous or fer- rugineous, up to 10 cm long. Inflorescence capitate, globose, with up to 20 spikelets, 5-12 mm across. Involucral bracts very short, usually hidden by the spikelets, setaceous, the longest up to 8 mm. Spikelets sessile, stellately spreading, the lower ones finally refiexed, falling off as a whole, strongly compressed, acute, 1(—3)-flowered, 5—6 by c. 1 mm. Glumes 4-6, usually only 1 flower-bearing, distichous, the lowest ovate, >4mm, the upper ones gradually longer, lanceolate, acute, mucronate, with acute 3-nerved keel and nerveless sides, up to 5 mm; mucro more or less recurved. Stamens 3; anthers linear, 2—3 mm. Style triquetrous, slightly thickened at the base, glabrous or sparsely setulose at the top, 4-4/4 mm; stigmas 3. somewhat shorter than the style. Nut obtusely trigonous, ellipsoid or obovoid, sessile, not or hardly umbonulate, smooth or sparsely verrucose, reticulate, whitish to stramineous, 1/4—-1/4 by c. 1mm; epidermal cells hexagonal or transversely elliptic. Distr. Madagascar, Mauritius, Ceylon, Thailand, Indo-China, S. China; in Malesia: Banka, P. Lingga, Malay Peninsula (Johore, Pahang, Malacca, Perak, Kedah, P. Penang), Anambas Is., Borneo (Sarawak), Philippines (Culion, Busuanga); not in the SE. part of the Archipelago. Ecol. In sandy fields, secondary forests, on rocky slopes, in the Philippines back of the mangrove swamps, usually below 500 m, in the Malay Peninsula at 300—2100 m, occasionally lower. Vern. Série béni, Banka. Notes. As appears from the synonymy the syste- matic place of this species has been much disputed. The flower-structure is that of Fimbristylis but there are several indications that the relationship may lay elsewhere (eucyperoid anatomical structure, turbinate embryo, spikelets falling off as a whole) and that for these reasqns one could argument its reinstatement to generic rank. This was recently done by RAYNAL, Adansonia 7 (1967) 89—95, who asserts that the rachilla is persistent and it is only the glumes that fall off as a whole. For the Malesian plants he accepted the binomial Actino- schoenus filiformis (THWAITES) BENTH., because he thinks that they are specifically distinct from the Hongkong and the greater part of the Madagascar specimens (A. thouarsii sensu stricto). To me the differences are too slight for specific separation. To judge from gross morphological characters F. yunnanensis CLARKE, J. Linn. Soc. Bot. 36 (1903) 247; TuRR. Kew Bull. (1912) 425; KERN, Blumea 10 (1960) 648; Reinwardtia 6 (1961) 44, known from N. Thailand, Yunnan and N. India, is a close ally of F. thouarsii, but anatomical and embryographic details of this species are lacking. 392 Species of doubtful place 78. Fimbristylis fenestrata K UK. Bot. Jahrb. 59 (1924) 49. Annual. Stems weak, densely tufted, compressed, sulcate, few-leaved at the base, 10—20cm tall. Leaves shorter than the stems, setaceous, with pale brown sheaths. Inflorescence usually consisting of a single pseudolateral spikelet, sometimes a second spikelet on a reflexed peduncle added. Spikelet oblong-lanceolate, 4-5 by 1/4 mm. Bract erect, as though continuing the stem, overtopping the spikelet(s). Glumes spiral, ovate, shortly mucronate, pale ferrugineous with green keel. Style long, with pyramidally thickened base; stigmas 3, short. Nut trigonous, broadly obovoid, truncate and quasi- tridentate at the apex, minutely reticulate, stramineous. Distr. Malesia: NE. New Guinea, Sepik Distr. (LEDERMANN 12523, not seen). Ecol. On steep seepage in scrubby mountain- forest with few large trees, 1400-1500 m. Note. According to KUKENTHAL intermediate between F. cardiocarpa F.v.M. and F. spiralis R.BR., both occurring in Australia. Excluded or doubtful Abildgaardia javanensis GANDOGER, Bull. Soc. Bot. Fr. 66 (1920) 296. — “Java, JUNGHUHN’’. Not seen. The description is quite inadequate. Probably F. ovata (BURM. f.) KERN. Fimbristylis longispica STEUD. Syn. 2 (1855) 118; Mig. Fl. Ind. Bat. 3 (1856) 325; Koorpb. Exk. Fl. FLORA MALESIANA [ser. I, vol. 7° Java 1 (1911) 199.— The type collection ‘‘Hrbr. GOERING nr. 10. Java” (P!) is certainly not from Java, but from Japan. The species is closely related to, but in my opinion well distinct from F. dichotoma (L.) VAHL; see, however, KoyaAMA, J. Fac. Sc. Un. Tokyo III, 8 (1961) 112, who reduced it to subspecific rank. It is known from Japan, Korea, Quelpaert I., and China, but not from Malesia. CLARKE’s record for Singapore in Fl. Br. Ind. 6 (1893) 639 refers to a coarse, coastal form of F. dichotoma, which in outward appearance is similar to F. longispica, but differs in several details. RIDLEY’s ‘F. longispica’ is partly that form, partly F. insignis THWAITES. See also the notes under F. dichotoma. Fimbristylis separanda STEUD. ex JARD. Mém. Soc. Imp. Sc. Nat. Cherb. 5 (1857) 299, 324. — I have not seen the type. According to DRAKE DEL CASTILLO, FI. Polyn. Frang. (1893) 245, it is synonymous with F. nukahiwensis STEUD. Syn. 2 (1855) 117. In the Paris Herbarium there are some specimens determined by CLARKE as F. separanda, from “‘Baie de Sanpoong- Sumatra, détroit de la Sonde. Voyage de l’Astrolabe et de la Zélée 1838-1840; M. Homsron, 1841”. They represent a species related to F. dichotoma (L.) VAHL, with very short, narrow leaves, glumes hairy in the upper part, and nuts with 16-18 longitudinal series of epidermal cells on either face. The specimens of the Astrolabe expedition are often mislabelled, and it is very unlikely that HOMBRON collected the plants in S. Sumatra. (Sanpoong should possibly be Lampong). See also S. T. BLAKE, Contr. Queensl. Herb. 8 (1969) 10-12. 14. CYPERUS LInngE, Gen. PI. ed. 5 (1754) 26; Sp. Pl. ed. 1, 1 (1753) 44; Kunru, En. 2 (1837) 2; STEUD. Syn. 2 (1855) 2; Mig. Fl. Ind. Bat. 3 (1856) 254; Boeck. Linnaea 35. (1868) 436; ibid. 36 (1870) 271; B. & H. Gen. Pl. 3 (1883) 1043; C. B. CLARKE, J. Linn. Soc. Bot. 21 (1884) 33; Pax in E. & P. Pfl. Fam. 2, 2 (1887) 107; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 41; KUk. Pfl. R. Heft 101 (1935-1936) 41; KoyAMA, Quart. J. Taiwan Mus. 14 (1961) 159. — Mariscus GAERTN. Fruct. & Sem. 1 (1788) 11. — Juncellus C. B. CLARKE, FI. Br. Ind. 6 (1893) 594. — Chloro- cyperus RIKLI in Pringsh. Jahrb. 27 (1895) 563. — Duvaljouvea PALLA in Koch, Syn. ed. 3, 2 (1905) 2555. — Fig. 49-70. For the other generic names referred to Cyperus in the present treatment, see under the synonymy of the subgenera and sections. Perennial or annual herbs, tufted or with creeping rhizome. Stems erect or obliquely erect, solid, triquetrous, trigonous, or sometimes subterete, usually leafy only at the base, rarely halfway up. Leaves tristichous, narrowly linear, grass-like, rarely lanceolate or elliptic, the lower ones often scale-like, covering the base of the stem and the rhizome, rarely all reduced to their sheaths. Inflores- cence terminal, often anthelate (terminal spike or cluster overtopped by the lateral ones), simple to decompound, umbel-like, not rarely capitate by sup- pression of the rays, in a few spp. truly umbellate. Rays subtended by a foliaceous bract similar to the leaves, the base enclosed in a tubular, two-keeled prophyll (cladoprophyll; see below, note), the bracts usually approximate as though forming an involucre. Spikelets more or less compressed, quadrangular, or | 1974] CYPERACEAE (Kern) 593 subterete, 1-many-flowered; axis (rachilla) often winged by the decurrent base of the glumes, persistent or caducous (in the latter case spikelets falling off as a whole). Glumes distichous, rarely nearly so, usually 2 basal ones empty (bracteole and prophyll), rarely 3 or 4 so. Flowers bisexual, the uppermost of the spikelet often male or barren by reduction; very rarely male and female flowers in different individuals (dioecious; not in Malesia). Hypogynous bristles or scales absent. Stamens 3,2, or 1, the median one anterior, the others transversal: connective often produced into an apical appendage. Style continuous with the ovary, not or hardly incrassate at the base, caducous, 3-fid or 2-fid, very rarely almost undivided. Nut sessile or shortly stipitate, trigonous or lenticular (planoconvex, concave-convex, or biconvex). Distr. About 600 spp., the vast majority of them in the tropics and subtropics. In Malesia 76 spp., most of them of pantropical or paleotropical distribution. A few spp. are apparently endemic in New Guinea (C. neoguineensis, C. subpapuanus, C. cinereobrunneus, C. meistostylus, C. pachycephalus). Several spp. are very rare in Malesia and show large disjunctions (C. ohwii, C. exaltatus, C. alopecuroides, C. bulbosus, C. zollingeri, C. babakan, C. multispicatus, C. teneriffae, C. diaphanus, C. substramineus). Recent introductions are C. aromaticus and C. sphacelatus. Ecol. Most spp. are hygrophilous; they grow in moist or wet localities at low and medium altitudes. Only a few ascend to above 1600 m. The spp. of sect. Diffusi prefer moist forests. C. stoloniferus, C. radians, C. dubius, and C. pedunculatus grow on sandy sea-shores and in sandy dunes, those with creeping stolons often acting as sandbinders. C. procerus and especially C. malaccensis prefer muddy places within the influence of salt or brackish water. C. iria, C. difformis, C. pulcherrimus, C. halpan, and C. tenuispica are characteristic of inundated rice-fields, C. rotundus is a troublesome weed in gardens and on road-sides. Uses. Economically the genus is unimportant. The dried stems of some spp. (e.g. C. elatus, C. malaccensis, C. procerus) are used for making mats or as strings. C. brevifolius, C. kyllingia, and C. rotundus make rather good pasturage when they are young and grow extensively. C. flabelliformis and C. papyrus are grown as ornamentals in ponds. For C. flabelliformis see p. 618. Cyperus papyrus L., indigenous in Central tropical Africa, is readily recognizable by its up to 5 m tall, pithy stems at the base covered with bladeless sheaths, the umbellate inflorescence with very numerous, subequal rays, and the lanceolate involucral bracts much shorter than the inflorescence. It was early used for paper-making and cultivated for this purpose in Egypt, Palestine, and S. Europe. See CHIOVENDA, Mem. Istit. Bot. Modena 1 (1931) 1—120, t. 1—4. The tubers of C. esculentus and C. bulbosus are edible, but their use is not recorded for Malesia. Notes. Flower-structure in Cyperus is as in those Scirpus spp. which lack hypogynous bristles; Cyperus differs in that the glumes are distichously arranged (spirally in Scirpus). See note under C. pygmaeus, p. 635. There is no unanimity on the delimitation of the genus (type species C. esculentus L.). C. B. CLARKE, the founder of modern Cyperology, originally took Cyperus ina wide sense, but in his later publications he proposed the following subdivision: 1. Style 2-branched. ee EECIONOUS ois Eh oleh vas cane SNE teers Bie eek te eueadcen oa REE 2. Rachilla persistent. Boteuidatceally compressed oi) 0) Pi ee A OR eres Be NEEBOLSAlly COMpressed: see RR) ee a a. tt Pell ers 1. Style 3-branched. MGR DCESISECHE: 2) 5) kM! 5, Ne cog fee ee Me Oia 4. Rachilla deciduous. 5. Rachilla deciduous in one piece. POteMmes NOL winged sf. fo Me Me 245) Sy se ls Me) se) SOorisrus BoeOmies WINS a lg h ge 9 6 Neoareare 5. Rachilla breaking up into as many jointsasnuts . . . . . . Torulinium This classification is artificial in many respects. More recently two other (monotypic) genera have been segregated, Sphaeromariscus Camus for C. compactus, and Queenslandiella Domin (= Mariscopsis CHERM.) for C. hyalinus. In VALCKENIER SURINGAR’Ss excellent revision of Cyperus in the Malay Archipelago (1898), the genus is taken in its wide sense, and so it is in KUKENTHAL’S monograph in the Pflanzenreich (1935—36). With the exception of Courtoisia the genera accepted by CLARKE were here reduced to subgeneric rank. As also Juncellus, Mariscus and Torulinium are ill-defined even as subgenera, in the present treatment only Cyperus, Pycreus, and Kyllinga are accepted as such. See also O’NEILL, Rhodora 44 (1942) 43-47. VAN DER VEKEN, Bull. Jard. Bot. Brux. 35 (1965) 285-354, studied the embryos of 132 species of Cyperus belonging in different groups. The uniformity of the embryos appears to support the wide conception of the genus. The function of the cladoprophyll in Cyperus was pointed out by TH.Hotm, Am. J. Sc. 18 (1904) 304. It is by means of the cladoprophylls that the primary rays of the anthela attain their more or less spreading position. 594 FLORA MALESIANA [ser. I, vol. 7° Fig. 49. Cyperus rotundus L. a. Habit, x )4, b. spikelet, x 3, c. broadly winged rachilla, d. glume, e. anther, f. nut, all x 7 (a—f IBoeT 157). 1974] CYPERACEAE (Kern) 595 Before the flowers are fully developed the rays are erect and congested. When mature the tubular prophylls show a prominent swelling at their base on the adaxial side. This swelling consists of a rapidly developing tissue without chlorophyll. By the rapid growth of this local tissue the rays are pushed away from the central axis. KEY TO THE SECTIONS represented in Malesia The circumscription of several sections is wider than in KUKENTHAL’s monograph. It has been tried to bring the sectional names into accordance with the Rules (in the said monograph they are often not). 1. Nut with a face towards the rachilla, trigonous or, when lenticular, dorsoventrally compressed. Spp. 1—60. I. SuBG. CYPERUS 2. Rachilla persistent after the glumes and nuts have fallen off. 3. Spikelets spicately arranged on a more or less elongated rachis. 4. Spikes cylindrical, much longer than wide, with very numerous spikelets. Tall perennials with long, broad leaves and bracts. Inflorescence anthelate, large, compound. Stolons absent. 5. Stigmas 3. Nut trigonous. Spp.1-5. . . . . 1. Sect. Exaltati 5. Stigmas 2, or only in some flowers 3. Nut planoconvex, or ‘only some of them with a raised dorsal angle. Sp. Gdongr: . . 2. Sect. Alopecuroidei 4. Spikes ovoid or obovoid, usually less than thrice as long as wide, with much less numerous spikelets. Stigmas 3. Nut trigonous. 6. Rachilla broadly winged. 7. Rhizome horizontally creeping or emitting long stolons, often tuberiferous. Glumes densely imbricate. 8. Leaves and involucral bracts long. Stems trigonous. Spp.7—10 . . 3. Sect. Cyperus 8. Leaves reduced to bladeless or very shortly laminate sheaths. Involucral bracts short. Stems subterete. Sp.11. . . 4. Sect. Corymbosi 7. Rhizome very short, without long, tuberiferous stolons, ‘rarely annuals with fibrous roots. Glumes remote, not or scarcely imbricate. m9) Spikelets subquadrangular (rhomboidal in cross-section), distinctly compressed. Slender, medium- sized perennials or annuals. Spp.12-14_ . . 5. Sect. Subimbricati 9. Spikelets subterete, but slightly compressed. Perennials, often robust. Spp. 15-16. 6. Sect. Distantes 6. Rachilla not or hardly winged. 10. Stout or rather stout perennial with rhizomes emitting long stolons. Glumes rounded on the back, muticous or inconspicuously mucronulate, broadly hyaline-margined in the upper part. Spp. 17-20. 7. Sect. Proceri 10. Annuals with fibrous roots. Glumes distinctly keeled and mucronate. 11. Nut about as long as the subtending glume. Glumes orbicular to broadly ovate, rounded or emarginate at the apex. Style hardly any, stigmas short. Sp.21. . . 8. Sect. Iriae 11. Nut about half as long as the subtending glume. Glumes ovate, acuminate. Style and stigmas long. Spi 2274 al. . 9. Sect. Compressi 35 Spikelets digitately or ‘stellately arranged on a much abbreviated rachis (often : some solitary in the same inflorescence), or inflorescence capitate. Rachilla wingless or but narrowly winged. 12. Inflorescence anthelate (umbel-like), with more or less developed rays. Stigmas 3. Nut trigonous. 13. Leaves of the flowering stems reduced to their sheaths. Involucral bracts numerous (up to 20), sub- equal in length, somewhat distant from one another. Perennials with large, compound inflorescences. Sp. 23 . . . 10. Sect. Alternifolii 13. Leaves with well- developed blades. Involucral bracts less numerous, unequal in length, close together. 14. Perennials with very short rhizome; stolons absent. Involucral bracts long. Spikelets turgid- compressed. Glumes usually mucronate. 15. Nut with corky angles. Style flattened, fimbriate. Leaves septate-nodulose. Sp. 24. 11. Sect. Pseudanosporum 15. Angles of nut not corky. Style not flattened, glabrous or ciliate. Leaves not or inconspicuously septate-nodulose. 16. Stems longer than the inflorescence. Spp. 25-32 . . . 12. Sect. Diffusi 16. Stems very short, much shorter than the primary rays of the inflorescence. Sp. 33. 13. Sect. Radiantes 14. Perennials with stoloniferous rhizome, or small-sized, tufted annuals. 17. Glumes muticous or inconspicuously mucronulate. 18. Glumes cellular-reticulate. Annuals. Sp.34 . . . . . . . . +. . 14. Sect. Tenelli 18. Glumes not cellular-reticulate. 19. Nut much shorter than the subtending glume. Annuals or perennials. Spp. 35-37. 15. Sect. Halpani 19. Nut about as long as the subtending glume. Spikelets crowded into very dense clusters. Annuals. SDS Shy no oe kee fe 16.) Sect: Rusel 17. Glumes (in Malesian spp. ) ‘with conspicuous, spreading or recurved mucro, Dwarf annuals with narrow leaves 20. Glumes truncate, with 3-nerved keel and nerveless sides. Spp. 39-40 . . 17. Sect. Amabiles 20. Glumes attenuate, strongly nerved over the whole breadth. Sp.41 . . . 18. Sect. Aristati 596 FLORA MALESIANA [ser. I, vol. 7° 12. Inflorescence capitate, without any trace of rays. 21. Stigmas in all the flowers 3 (in sect. Anosporum style almost undivided). Nut trigonous. Perennials. 22. Angles of the nut not corky. Style deeply trifid. 23. Glumes mucronate. Lower leaf-sheaths scarious, inflated. Involucral bracts short, not or but slightly overtopping the inflorescence. Sp. 42. . . . 19. Sect. Humiles 23. Glumes muticous. Lower leaf-sheaths not inflated. Involucral bracts much overtopping the inflorescence. Sp.43. x (2p elise, omee 20. Sect, Blatystaclags 22. Angles of nut corky. Style almost undivided. Sp. 44. . 21. Sect. Cephalotes 21. Stigmas in flowers of the same inflorescence partly 2, partly 3. Nut lenticular, dorsoventrally com- pressed, or (in the same inflorescence) trigonous. Angles of the nut acute, often narrowly winged. Annuals. Spp. 45-46. 1 « « « « « 22. Sect Dichostyis 2. Rachilla disarticulating, falling off with the glumes and nuts. 24. Rachilla disarticulating only at the base; mature spikelets falling off entire. Stigmas 3. Nut trigonous. Perennials. 25. Internodes of the rachilla not corky. Spikelets rarely strictly 1-flowered. 26. Stem-base not strikingly pseudo-bulbous. 27. Spikelets several- to many-flowered, maturing at least 3 nuts. Glumes not tightly clasping the nut. 28. Spikelets oblong or lanceolate. Nut oblong-ellipsoid or oblong-obovoid, rarely narrower. 29. Glumes appressed. Spikelets subterete, turgid. Glumes rounded on the back. Stout plants with large inflorescences and septate-nodulose leaves. Spp.47-48 . . . . . 23. Sect. Pennati 29. Glumes patulous, at least at the top. Spikelets not turgid. 30. Rachilla broadly winged. Leaves broad. Sp.49 . . . 24. Sect. Thunbergiani 30. Rachilla not or but narrowly winged. Leaves narrow. Spp. GOA54 wu ee tute 2S Pinnati 28. Spikelets linear. Nut oblong-linear or oblong. 31. Spikes hemispherical or subglobose. Spikelets subulate, subterete. Spp. 52—53. 26. Sect. Flabelliformes 31. Spikes broadly cylindrical. Spikelets linear, compressed. Sp.54 . . . . 27. Sect. Strigosi 27. Spikelets few-flowered, maturing 1—2(—3) nuts. Glumes tightly clasping the nuts. Inflorescence simple, rarely subcompound. Spp.55=S7. . . 28. Sect. Mariscus 26. Stem-base pseudo-bulbous by turgid, membranous sheaths. Inflorescence capitate. Sp. 58. 29. Sect. Kyllingioides 25. Nut-bearing internode of the rachilla corky. Spikelets 1-flowered. Rhizome long-creeping. Sp. 59. 30. Sect. Remirea 24. Rachilla disarticulating at each node. Spikelets breaking up into short segments each bearing 1 nut tightly embraced by the much thickened wings of the rachilla.Sp.60. . . . 31. Sect. Diclidium 1. Nut with one of the two edges towards the rachilla, laterally compressed. Stigmas 2. 32. Rachilla persistent after the glumes and nuts have fallen off. Spp.61-69 . . . II. SuBG. PycReus 33. Epidermal cells of the nut isodiametric. Nut smooth. 34. Glumes muticous. 35. Stems decumbent at the base, rooting at the nodes, leafy in the lower 4—)4. Sp. 61. 32. Sect. Vestiti 35. Stems erect, leafy only at the very base. 36. Spikelets lanceolate or oblong. Glumes at least 3 mm long. Sp.62 . . . 33. Sect. Chrysanthi 36. Spikelets linear. Glumes at most 3 mm long. Spp.63-65. . . . . . . 34. Sect. Pycreus 34. Glumes mucronate. Spp. 66-67. . . . $35. Sect. Pomili 33. Epidermal cells of the nut longitudinally stretched. Nut rugulose with transverse wavy lines. Spp. 68—69. 36. Sect. Flavescentes 32. Rachilla disarticulating at the base. Spikelets falling off entire. Spp. 70-76. . III. SuBG. KYLLINGA 37. Inflorescence anthelate (umbel-like). Spikelets several-flowered. Sp. 70. . 37. Sect. Queenslandiella 37. Inflorescence capitate. Spikelets 1—2-flowered. S8aGlumesiwingless/Spps7l=75.... 2 PR Se oe Be oe os ow Oe se S85 Seeley 38: Glumes with winged keel: Sp! 76) Pde! s. See ee.) OLR UR | ea eee KEY TO THE SPECIES Only specimens with subterranean parts and ripe or almost ripe fruits are identifiable 1. Nut with a face against the rachilla (axis) of the spikelet, trigonous, or, when 2-sided, dorsoventrally compressed. 2. Style entire or but slightly notched at the top. Inflorescence a dense head. Nut corky on the angles below. 44. C. cephalotes 2. Style distinctly cleft into 2 or 3 stigmas. Other characters not united. 3. Stigmas 2 or in part of the flowers 3. Nut 2-sided or partly trigonous (several flowers should be dissected !). 4. Inflorescence with well-developed rays and oblong to cylindrical spikes. Stout perennial up to more than 1 m tall, with broad leaves up to 16mmwide. . . 6. C. alopecuroides 4. Inflorescence capitate, consisting of some sessile glomerules. Dwarf or medium-sized annuals, rarely up to 75 cm tall, with narrow leaves rarely up to 5 mm wide. 5. Glumes 3(—5)-nerved, with more or less spinulose keel, acute or minutely mucronulate. Stamens 1-2. Dwar, farely up to.25:enitéall) “eee = ih & le Rs Dam, Fae RY SoG: Cans y priiteris 1974] CYPERACEAE (Kern) 597 5. Glumes 7—9-nerved, with smooth keel, distinctly mucronate. Stamens 3. Usually medium-sized, up to 75cmtall_ . ee ee be een rae A ee DaAChYCepnaluS 3. Stigmas always 3. All the nuts trigonous. 6. Ripe nut tightly clasped by a much thickened. corky internode of the rachilla (in C. pedunculatus this internode is easily mistaken for an anomalous glume!). 7. Rhizome long-creeping. Spikelets 1-flowered, disarticulating at the base, falling off entire. 59. C. pedunculatus 7. Rhizome short. not stoloniferous. Spikelets several- to many-flowered. when mature breaking up miolshort sesmentsieach’contaiming anit) -)) 0,04 3 ee, 4 «6 «160: CC. odoratus 6. Rachilla-internodes not becoming corky. 8. Spikelets spicately arranged, i.e. at some distance from one another upona more or less elongated rachis. 9. Primary rays of the inflorescence very numerous (30-100). Flowering stems 2—5S m tall, obtusely trigonous or subterete, pithy, the base surrounded by bladeless sheaths. Involucral bracts much shorter than the umbellate inflorescence. . Pen Cue eee) (SCC P2593. C papyrus 9. Primary rays of the inflorescence much less numerous. 10. Spikelets 3—5 mm wide (mucros included). Glumes with strong, up to | mm long mucro. Annuals with fibrous roots. 11. Spikelets not squarrose, the mucros of the glumes at most slightly excurved; body of the glumes 3—4 mm long. Nut broadly ovoid, c. 1/4 by 1mm.Stamens3 . . . . 22. C. compressus 11. Spikelets finally squarrose by the strongly excurved mucros of the glumes; body of the glumes 1—2 mm long. Nut oblong-obovoid to almost linear. 74-1 by 14-4 mm. Stamen 1. 41. C. squarrcsus 10. Spikelets narrower and/or glumes muticous. 12. Stems with concave sides, almost 3-winged above, spongious, robust. 13. Glumes finally crispidly incurved all round, smooth. Inflorescence compound. Nut narrowly oblong ai-y . . 20. C. malaccensis 13. Glumes not crispidly incurved, hispid- scabrous on the keel at least above. Inflorescence simple. Nut broadly ellipsoid or fibowpida en) mess cli ane 2omeret Of 19:\C. habakan 12. Stems with flat or convex sides. 14. Leaves strongly septate-nodulose, coriaceous, very scabrous on the margins and keel. 15. Stems densely papillose (hand-lens!). Nut ellipsoid or subobovoid, 114-114 mm long. 47. C. javanicus 15. Stems not papillose. Nut oblong-linear, 2?4-2% mmlong. . . . . . 49. C. lucidus 14. Leaves not strongly septate-nodulose. 16. Each spikelet maturing several to many nuts. 17. Rachilla (axis) of the spikelet distinctly winged. 18. Spikes cylindrical, with long rachis, much longer than wide, with very numerous (more than 30) many-flowered spikelets. Tall perennials with broad leaves and bracts, and large, compound or decompound inflorescences. Rhizome short, not stoloniferous. 19. Glumes in the upper half with a broad, white, scarious margin, soon incised at the tip and bicuspidate, mucronulate somewhat below the apex. Spikelets suberect. Appendage of connective smooth . . =. 2 15: Cnutans 19. Glumes not scarious- margined, ‘with entire tip, muticous or with a short apical mucro. 20. Connective of anthers produced into a setulose appendage )4—/4 as long as the anther-cells. 21. Spikes dense, 3—5 mm thick. Spikelets suberect, linear, 1 mm wide. Cladoprophylls (tubular bracts surrounding the base of the i of the inflorescence) scabrous on the keel at least above. Nut % mm long . . 2. C. elatus 21. Spikes very dense, 6—8 (ultimately up to 10) mm wide. Spikelets obliquely patent, linear- oblong, 1/4 mm wide. Cladoprophylls smooth. Nut 1/4,-1/4 mm long . 3. C. ohwii 20. Connective of anthers but shortly produced, smooth. 22. Glumes muticous or apiculate. aces but slightly compressed, subterete, when ripe at right angles to the rachis. . . . I. C. digitatus Glumes with a short but distinct mucro. Spikelets distinctly compressed. eH Secondary rays of the inflorescence (in decompound inflorescences the tertiary rays) very short, hence the spike sessile or almost so. Spikelets suberect, completely hiding the rachis of the Spikezia see . 5. C. imbricatus 23. Secondary rays of the inflorescence up to 5cm long, hence the spikes at least partly distinctly peduncled. Spikelets obliquely patent to almost at — angles to the rachis, the latter therefore visible. . : . . 4. C. exaltatus 18. Spikes ovoid or broadly ovoid. Other characters not united. 24. Rhizome horizontally long-creeping or emitting slender to thread-like stolons (often lacking in herbarium specimens), often tuberiferous. Glumes imbricate, i.e. tip of each pinme distinctly (at least 14) overlapping the next higher glume on the same side of the spikelet. 25. Stems subterete in the lower part, triangular only just below the inflorescence. Leaf-blades short to almost wanting, rarely half as long as the stem. Lowermost involucral bract usually erect, as though continuing the stem, pushing aside the small inflorescence. 11. C. scariosus 598 FLORA MALESIANA [ser. I, vol. 7° 25. Stems triquetrous. Leaf-blades well-developed. 26. Involucral bracts distinctly separated from one another or only the lowermost obvious. Stolons thread-like, producing bulb-like tubers enclosed in a hard, black, irregularly splitting coat. Inflorescence often reduced to a simple or compound spike. 8. C. bulbosus 26. Involucral bracts close together or only the lowermost a little distant. Tubers not bulb-like, when young covered by membranous scales soon disintegrating into fibres. Inflorescence umbel-like. 27. Nut strongly compressed dorsally, the ventral side usually somewhat concave, the dorsal side with a raised angle. Ripe spikelets turgid, subterete. . 10. C. stoloniferus 27. Nut equally trigonous (in C. rotundus rarely maturing; examine ovary!). Ripe spikelets not turgid, strongly compressed. 28. Glumes with 7—9 prominent nerves equally distributed over the whole breadth (outer- most nerves near the margin), golden yellow to pale brown. Stolons very slender, yellowish, producing globose or ovoid tubers covered with a grey tomentum. 9. C. esculentus 28. Glumes with 5—7 nerves, the lateral nerves becoming much less prominent away from the keel, usually deep brown, rarely yellowish. Stolons stoutish, dark brown, forming ellipsoid tubers without grey tomentum .. _ 7. C. rotundus 24. Rhizome neither long-creeping nor emitting slender stolons, at most with very short, ascending surculi, never tuberiferous, or annuals with fibrous roots. 29. Nut turbinate-obovoid, truncate or even slightly pee at the top. Annual with fibrous roots. . . oie ee Ae we 2 SIS iGs7olingent 29. Nut not turbinate, not truncate. 30. Annual with fibrous roots. Glumes as a rule with a purplish spot on one or both sides giving the impression of a purplish stripe along the centre of the spikelet. 14. C. sphacelatus 30. Perennials with short rhizome. Glumes not purplish spotted. 31. Rachilla persistent on the rachis after the glumes and nuts have fallen off acropetally. 32. Glumes 3’4—4 mm long, with broad green keel and golden yellow to fulvous sides. Spikelets subquadrangular, i.e. rhomboidal in cross-section . . 12. C. tenuiculmis 32. Glumes 2-2/4 mm long, the keel not strikingly broad. Spikelets circular or elliptic in cross-section. 33. Spikelets narrowly linear, almost subulate, c. 1 mm wide, usually spreading at right angles. Glumes very distant, those on one side of the spikelet not overlapping. 16. C. distans 33. Spikelets oblong-linear, compressed, c. 2 mm wide, suberect. Glumes )4 or more overlapping . . 15. C. nutans 31. Rachilla disarticulating at the base, the spikelets falling off entire. Glumes 3-4 mm long, strongly nerved over the whole breadth. Nut 2- 2% mm long. 34. Glumes slightly imbricate (those on one side of the spikelet somewhat overlapping). Spikelets 1-114 mm wide. Nut oblong-obovoid . . . . 54. C. stenophyllus 34. Glumes distant, those on one side of the spikelet not overlapping. Spikelets subulate, c./4mm wide. Nut oblong-linear . . . . . .. . 53. C. dietrichiae 17. Rachilla not or hardly winged. 35. Glumes orbicular or broadly ovate, keeled, mucronulate, 1-1/4 mm long. Rachis of the spikes glabrous and smooth. Nut as long as the subtending glume. Estoloniferous. 21. C. iria 35. Glumes ovate or elliptic, rounded on the back, muticous, 2—3 mm long. Rachis of the spikes hispidulous or scabrid (sometimes but sparsely so). Nut about 7% as long as the subtending glume. Perennials with creeping stolons. 36. wel of the glumes scabrid at least towards the top. Inflorescence simple, with very dense spikes . Jet pe bide hee one Yrote cities £2 19s Caianrmen 36. Keel of the glumes smooth. 37. Inflorescence simple or subcompound. Rachis of the spikes sparsely scabrid to almost smooth. Spikelets 2/4—-3/4 mm wide. Anthers (1—)1/4-2 mm long. Nut 1/4 mm long. 17. C. procerus 37. Inflorescence compound. Rachis of the spikes rather to very densely hispidulous. Spikelets 2-214(-3) mm wide. Anthers 14-34 mm long. Nut 1-144 mmlong ._ . 18. C. pilosus 16. Each spikelet maturing 1—2(—3) nuts, rarely more than )% cm long, falling off as a whole. Spikes dense, exactly cylindrical to slightly obovoid, up to 1 cm wide. 38. Rhizome emitting slender stolons. Spikelets always bearingi nut . . . 57. C. paniceus 38. Rhizome not stoloniferous. Spikelets usually bearing 2—3 nuts. 39. Spikes exactly cylindrical, usually distinctly peduncled. Spikelets finally at right angles to the rachis or reflexed. Nut linear,0.5 mm wide. . 56. C. cyperoides 39. Spikelets attenuate to the base, sessile or almost so. Spikelets obliquely erect (see, however, var. maximus). Nut ellipsoid or oblong-ellipsoid, 0.6—0.9 mm wide . . 55. C. cyperinus 8. Spikelets digitately or stellately arranged, i.e. at practically the same level upon a much shortened 1974] CYPERACEAE (Kern) 599 rachis, forming several to numerous clusters, sometimes part of them solitary, or inflorescence capitate. 40. Involucral bracts numerous (up to 20), almost equal in length, evidently distant from one another. 23. C. flabelliformis 40. Involucral bracts much less numerous, unequal, close together. 41. Angles of nut much thickened by a corky tissue. Stout perennial with thick, funiliform roots and broad, septate-nodulose leaves. Style flattened, fimbriate . . . . . . 24. C. platystylis 41. Angles of nut not corky-thickened. 42. Stems shorter than the remarkably long primary rays of the inflorescence (those rays are easily mistaken for stems when the true stem is onlyafewcmlong!) . . . . . 33. C. radians 42. Stems longer than the inflorescence. 43. Glumes very small, #4—*% mm long, orbicular or broadly ovate. a globose, very dense, with MUSOU SoS. 4 go =o oo 6 89 8 © 6 . . . 38. C. difformis 43. Glumes larger. 44. Leaves septate-nodulose (see 46. C. fulvus!). 45. Clusters of spikelets (1’4-)2—4 cm across. Spikelets subulate, 1-114 mm wide. Glumes 3-414 mm long. Leaves 5— i) mmMWwiders wee . . 52. C. compactus 45. Clusters of spikelets at most 1 cm across. Spikelets 2 mm wide. Glumes 2-2/4 mm long. Leaves 2—4 mm wide. 46. Inflorescence compound. Leaves strikingly septate-nodulose, especially their lower, spongy sheaths. Spikelets 2—3 mm long, few-flowered. Nut oblong, y mm wide. 48. C. holoschoenus 46. Inflorescence simple. Leaves not strikingly septate-nodulose. Lower leaf-sheaths not spongy; stem-base almost bulbous. Spikelets 6-many-flowered, 5-20 mm long. Nut oblong- epevoid74=1 min wide: waren ity iets iMiaeh nit) og teh”. SIRIC Rallvas 44. Leaves not septate-nodulose. 47. Glumes truncate or emarginate, with 3-nerved keel, nerveless sides, and strong, more or less recurved mucro. Small annuals with very narrow leaves A 1 mm wide. 48. Stamens 2 or 3. Nut obovoid, 4-7 mm wide... 2) 2 94,1) 3955Cxcuspidatus 48. Stamen 1. Nut oblong, )4- Vi mm wide . : . . . 40. C. castaneus 47. Glumes not truncate or emarginate. Other characters not united. 49. Inflorescence capitate. 50. Glumes mucronate, strongly many-nerved over the whole breadth. 51. Base of the plant much thickened by inflated, scarious, purplish striate sheaths. Body of the glumes 2—3 mm long, cinnamomeous to purplish. Stamens3 . . 42. C. teneriffae 51. Base of the plant not thickened. 52. Body of the glumes 1—2 mm long, with strongly recurved mucro. Stamen 1. Nut 74-1 mm long . 41. C. squarrosus 52) Body of the glumes 3-4 mm long, the mucro not or slightly recurved. Stamens 3. Nut c. 14 mm LONG APMC AOR NR! loc ants mis . . 22. C. compressus 50. Glumes muticous. 53. Inflorescence a single, whitish, globose head 5-10 mm across. Glumes 1-134 mm long. Stamen 1. Nut 0.5—0.9 by 0. 25mm... . 43. C. pulchellus 53. Inflorescence a lobed head consisting of u to 6 confluent glomerules, 10—20 mm across. Glumes 3—4 mm long. Stamens 3. Nut 1-214 mm long. . . . 58. C. dubius 49. Inflorescence umbel-like, sometimes more or less contracted, but not truly capitate. 54. Nut very small, at most 4 mm long. 55. Glumes cellular- reticulate, narrowly winged on the keel, with strong midnerve and a faint nerve in the‘centre of-either sidéy 510: © a! Spomet) ong) leet S40 aquatilis 55. Glumes neither cellular-reticulate nor winged. 56. Involucral bracts very long and broad, the lowermost up to 50 by 1 cm, far overtopping the inflorescence. Many spikelets solitary. Sides of the a slightly sulcate. Stamen 1. Nut % mm long . . 32. C. multispicatus 56. Involucral bracts much shorter. Nut at most y mm n long. 57. Glumes straight, obliquely erect, the ripe spikelets not perspicaceous. Stamens 1, 2, or 3. Connective of the anthers produced into a setulose appendage . 36. C. halpan 57. Glumes spreading, the ripe spikelets perspicaceous. Stamen 1(—2). Appendage of connective smooth. 58. Glumes ultimately incurved. Nut sharply trigonous, with flat sides, c. 4 mm long, minutely puncticulate . . . 35. C. pulcherrimus 58. Glumes ultimately excurved. Nut. very obtusely trigonous, with convex sides, sub- globose, /4—/4 mm long, densely tuberculate. . . . . . . 37. C. tenuispica 54. Nut more than 1 mm long. 59. Nut linear-oblong. Spikelets narrowly linear,1 mm wide. . . . 50. C. angustatus 59. Nut ellipsoid or obovoid. Spikelets wider. 60. Anthers with smooth appendage of the connective. (Spp. 27—31 from New Guinea inade- quately known). 61. Glumes with scabrid keel and excurved top, 9—11-nerved. Leaves 8-11 mm wide. Style very short, %-’4mmlong . . . . . . . . . 29. C. subpapuanus 600 FLORA MALESIANA [ser. I, vol. 7° Glumes with smooth keel, not excurved. "62. Leaves 10-18 mmwide . pst bee eh. 28.) Cineornineensis 62. Leaves much narrower, 3—7 mm wide. 63°" Spikelets: 3—Smimgwidelras fan a Pe sic een es Vener bee compressa 63. Spikelets 114-2 mm wide. 64. Style distinct, “4-1 mm ee fimbriate. Stems scaberulous at the top. Glumes 24%4-3mm long . 30. C. cinereobrunneus 64. Style hardly any, 0.1— 0.3 mm long. Stems smooth. Glumes 2- 2/4 mm long. 31. C. meistostylus 60. Anthers with setulose apical appendage of the connective. 65. Style distinct, fimbriate. Glumes 11—17-nerved, ferrugineous to blackish brown. 27. C. pedunculosus 65. Style hardly any, stigmas long. 66. Stems wingless. Spikelets oblong, up to 20-flowered . . . . . 25. C. diffusus 66. Stems 3-winged. Spikelets ovoid, up to 8-flowered_ . . 4. 261Cstralatas 1. Nut with an edge against the rachilla, bilaterally compressed, 2-sided. Stigmas always 2. 67. Each spikelet maturing several to many nuts. Inflorescence often umbel-like, more rarely contracted to subcapitate. 68. Epidermal cells of the nut isodiametric, roundish or hexagonal. Nut under the lens puncticulate or finely reticulate. 69. Midnerve of the glumes excurrent into a distinct, usually more or less recurved mucro. 70. Nut 114,-124 mm long, c. | mm wide. Rachilla broadly winged, sspmeccpat at the base, hence spikelets bye. offentire . . . 70. C. hyalinus 70. Nut 4- Me by 4-4 mm. Rachilla almost or quite wingless, persistent on the rachis, the glumes and nuts seriatim deciduous. 71. Glumes with 3-nerved keel and nerveless sides. Stamens 1(—2) . . . . . . 66. C. pumilus 71. Also the sides of the glumes distinctly nerved.Stamens2. . . . . . . 67. C. neryulosus 69. Glumes muticous or at most with a very short, indistinct subula. 72. Glumes c. 4 mm long. Spikelets 4-5 mm wide, somewhat turgid. Stamens3. . . C. unioloides 72. Glumes 1/4-2(-2/,) mm long. Spikelets 1/,—3(—3/4) mm wide, flat. Stamens 1, = or a 73. Stems decumbent at the base, rooting at the nodes, leafy in the lower 14-4. Glumes with arcuate keel, usually with a depression on either side. Nut obovate to almost orbicular. 61. C. sanguinolentus 73. Stems erect, leafy only at the very base. Glumes with straight keel, without depressions. Nut oblong or oblong-obovate. 74. Nut with a median, longitudinal, shallow depression on either side. Stamen1 . 65. C. sulcinux 74. Nut with flat or slightly convex sides. Stamens usually 2, rarely 1. 75. Leaves flat. Glumes acute. Rachilla narrowly winged. Spikelets gradually tapering to the acute apex. Nut narrowly oblong, subtruncate at the top, minutely apiculate . 64. C. polystachyos 75. Leaves canaliculate, often almost filiform. Glumes obtuse. Rachilla straight, wingless. Spikelets exactly parallel-sided. Nut oblong-obovate to oblong-elliptic, not truncate, evidently apiculate. 63. C. flavidus 68. Epidermal cells of the nut longitudinally stretched, oblong or linear. Nut under the lens appearing wrinkled by the prominent short sides of the epidermal cells running into wavy or broken transverse lines. 76. Spikelets suberect even in fruit, 2 mm wide. Glumes mucronulate, stramineous-yellowish, 2 mm long. 69. C. substramineus 76. eal finally spreading, (2’4—)3—4 mm wide. Glumes muticous, fulvous, castaneous, or purplish, /o-3 mm long . . . 68. C. diaphanus 67. pach spikelet ennai maturing 1 nut, but not rarely some of them with 2 nuts in the same inflorescence. Inflorescence a whitish or greenish head of 1—S sessile glomerules with densely crowded, sessile spikelets. Rachilla disarticulating at the base, hence spikelets falling off entire. 17.. Keel ofthe ghimesdistinetlyiwinged’ shai ty: aye ae deade “art teed Tepaeiowal 46-76 C ayn 77. Keel of the glumes wingless. 78. Rhizome densely cespitose; stolons wanting. 79. Central glomerule of the inflorescence cylindrical, 10-12 mm long, lateral ones when present much smaller, subglobose. Nut truncate at the apex, 1 mm wide, ultimately black . 74. C. sesquiflorus 79. Central glomerule of the inflorescence subglobose, 5-8 mm long, lateral ones but slightly smaller. Nut not truncate, c. %mm wide, yellowish brownto brown. . . . . . . 75. C. triceps 78. Rhizome horizontally creeping. ENwlavelicral bracts \(5—\7=8 "cS et Pen oe ol Ot enicls 2 7722 1C caronaiens 80. Involucral bracts 3—4. 81. Rhizome slender, 1—2(—3) mm thick. Stems rather approximate to very distant, rarely more than 30 cm tall, 4-1/4 mm thick, triquetrous with flat sides. Leaf-blades usually well-developed. Stamens 1—2(-3) . . . . 73. C. brevifolius 81. Rhizome stout, 3-4 mm thick. Stems approximate, 30-175 cm tall, 2-4 mm _ thick, sharply triquetrous, with more or less concave sides. Leaves reduced to the sheaths or only the upper 1—2 shortly laminate, Stamens'3 0-02"... ewehdee ec oot iat yee, al. Co melanospermany 1974] Cy PERACEAE (Kern) 601 I. Subgenus Cyperus Type species: C. esculentus L. 1. Section Exaltati KUNTH, En. 2 (1837) 70. — Cyperus sect. Longispicati c. Digitati VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 133. — Cyperus sect. Fastigiati KUK. Pfl. R. Heit 101 (1935) 53. Type species: C. exaltatus RETZ. 1. Cyperus digitatus Roxs. [Hort. Beng. (1814) 81, nom. nud.] Fl. Ind. 1 (1820) 209; CLARKE, FI. Br. Ind. 6 (1893) 618; VALcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 133, t. 1 f. 10, t. 4 f. 35; CLARKE, Philip. J. Sc. 2 (1907) Bot. 86; Koorp. Exk. Fl. Java 1 (1911) 191; ibid. 4 (1922) f. 226; Camus, Fl. Gen. I.-C. 7 (1912) 74, f. 9, 5—6; Merr. En. Philip. 1 (1923) 105; Back. Onkr. Suiker. (1928) 139, t. 137; KUx. Pfl. R. Heft 101 (1935) 55; S. T. BLake, J. Arn. Arb. 28 (1947) 213; BAcK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 38; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 480.— C. venustus (non R.BR.) NEES in Wight, Contr. (1834) 86; KuNTH, En. 2 (1837) 68; Mia. FI. Ind. Bat. 3 (1856) 280: Borck. Linnaea 36 (1870) 316, p.p. — C. strigosus (non L.) LLANos, Fragm. PI. Filip. (1851) 16. — C. auricomus (non SiEB.) BENTH. Fl. Austr. 7 (1878) 286; Naves, Nov. App. (1882) 305: CLARKE, J. Linn. Soc. Bot. 21 (1884) 188, p.p., t. 4 f. 31; CERON, Cat. Pl. Herb. Manila (1892) 177; K.ScuH. & Laut. Fl. Schutzgeb. (1900) 191. —C. auricomus var. microstachyus BOECK. in K.Sch. & Hollr. Fl. Kais. Wilh. Land (1889) 23. — C. racemo- sus (non RETZ.) K.ScH. & HOLLR. I.c. 24; K.SCH. & Laut. Fl. Schutzgeb. (1900) 192. — Mariscus sie- berianus var. evolutior (non CLARKE) RIDL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 73, p.p.; Fl. Mal. Pen. 5 (1925) 149, p.p. — C. elatus (non L.) Camus, FI. Gén. I.-C. 7 (1912) 70, incl. var. laxus CAMUS. Perennial with very short rhizome. Stems tall, tufted, obtusely trigonous, often triquetrous above, smooth, 50—150(—200) cm by 3—4(—7) mm. Leaves flat or the larger ones slightly plicate with 3 prominent nerves, coriaceous, scabrous on the margins, glau- cous. 4—10(—15) mm wide; lower sheaths purplish. Inflorescence compound or decompound, up to 40cm across. Involucral bracts 3-8, spreading, much overtopping the inflorescence, up to 60 cm long. Primary rays 6-10, unequal, smooth, the longest 15-30 cm; secondary ones 2—3 cm. Spikes digitately arranged, cylindric, rather loose, 3-6 cm by 1-3 cm; rachis glabrous, narrowly winged. Spike- lets spicately arranged, finally at right angles to the rachis, linear, but slightly compressed, 8—24(—44)- flowered, 5-20 by 1-14 mm; rachilla straight, persistent; internodes c. 4 mm; wings caducous, yellow, 4-14 mm wide. Glumes membranous, ap- pressed, elliptic-oblong, acute, apiculate, 134-2!4 by c. | mm, halfway imbricate; keel green or reddish brown, 3—5-nerved; sides yellowish to rufous, nerve- less. Stamens 3; anthers linear, 14 mm long; con- nective shortly produced, smooth. Stigmas 3. Nut trigonous, ellipsoid to oblong-obovoid, apiculate, yellowish brown, c. 1 by 4-4 mm. Distr. Pantronical: tropical America, rare in tropical Africa; from India, Farther India, S. China, and Formosa to Queensland; throughout Malesia, but here everywhere scattered. Ecol. In swamps, wet rice-fields, swinging bogs, on river-banks and in other open, wet places at low altitudes, 0-800 m. Vern. Mal. Pen.: bunga sadaian rumput, rémpara, rumput musiang, M; N. Borneo: pajong krah, rumput kara, r. triwak, M; rumput dékéng, J; kakanin kara, Menado; New Guinea: simbuai, Sepik. 2. Cyperus elatus LINNE, Cent. Pl. 2 (1756) 301; Sp. Pl. ed. 2 (1762) 67; CLARKE, J. Linn. Soc. Bot. 21 (1884) 189, excl. var. macronux CLARKE; FI. Br. Ind. 6 (1893) 618; VaLcK. SuR. Gesl. Cyp. Mal. Arch. (1898) 135, t. 4 f. 36; CLARKE, Philip. J. Sc. 2 (1907) Bot. 86; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 71; Koorpb. Exk. Fl. Java 1 (1911) 191; ibid. 4 (1922) f. 225; Camus, Fl. Gén. I.-C. 7 (1912) 70 (non f. 8, 1); Me_rR. En. Philip. 1 (1923) 105; Ripe. Fl. Mal. Pen. 5 (1925) 148; Back. Onkr. Suiker. (1928) 140, t. 138: KUK. Pfl. R. Heft 101 (1935) 59; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 38; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 479. — C. race- mosus RETZ. Obs. 6 (1789) 20; NEEs in Wight, Contr. (1834) 85; KuNTH, En. 2 (1837) 100; StEUD. Syn. 2 (1855) 52, p.p.; Mig. Fl. Ind. Bat. 3 (1856) 278; Boeck. Linnaea 36 (1870) 310. — C. scoparius (non Poir.) DECNE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 359; Descr. Herb. Timor. (1835) 31; Mia. FI. Ind. Bat. 3 (1856) 279. — C. bispicatus StEuD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.] Syn. 2 (1855) 49; Mia. in De Vriese, Pl. Ind. Bat. Or. (1856) 140; Fl. Ind. Bat. 3 (1856) 285; Naves, Nov. App. (1882) 303.— C. auricomus (non Stes.) RipL. in Forbes, Wand. (1885) 521.— C. radiatus (non VAHL) CAMUS, Fl. Gen. I.-C. 7 (1912) 72, f. 9, 1-4. — C. digitatus (non Rox.) RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 71; Merr. En. Born. (1921) 105; WINKL. Bot. Jahrb. 44 (1923) 523; Ripv. Fl. Mal. Pen. 5 (1925) 147. Perennial with very short rhizome. Stems tall, tufted, trigonous but triquetrous at the top, smooth, 50—100(—200) cm by (3—)4—7(-10) mm. Leaves flat or often somewhat plicate with the midrib prominent beneath and 2 lateral nerves strongly marked above, scabrous on the margins, bright green above, pale green beneath, 4—10(—15) mm wide; lower sheaths purplish to almost black. Inflorescence compound or decompound, up to 30 cm across. Involucral bracts 4—8, much overtopping the inflorescence, up to 60(—75) cm long. Keel of the cladoprophylls scabrous, at least in the upper part. Primary rays up to 10, unequal, smooth, up to 20 cm long; secondary ones 2—5 cm. Spikes digitately arranged, narrowly cylin- dric, dense, 2-7 cm by 3-5 mm; rachis glabrous, 602 narrowly winged. Spikelets spicately arranged, sub- erect, linear, compressed, 6—16-flowered, 3—6 by c. 1 mm; rachilla slightly flexuous, persistent; inter- nodes 4-74 mm; wings caducous, yellow, c. 4 mm wide. Glumes membranous, appressed, ovate, acute, mucronulate, 114-1374 by 1mm, 4-7 imbricate; keel green or reddish brown, 3—5-nerved; sides golden to rufous. Stamens 3; anthers oblong-linear, c. 4 mm long; connective produced into an apical appendage c. /4 as long as the anther-cells and setulose at the top. Stigmas 3. Nut trigonous, ellip- soid, apiculate, yellowish to greyish brown, 0.8—0.9 by 0.3—0.4 mm. Distr. From India and Farther India to Malesia: rather common in Sumatra, the Malay Peninsula, and Java; but a few times collected in the Lesser Sunda Is. (Sumba, Timor), Borneo, and Celebes:; rare in the Philippines (Luzon, Biliran, Leyte, Ticao). Apparently not reaching the Moluccas and New Guinea. Ecol. In swamps, wet rice-fields and meadows, on river-banks, and in other open wet places at low altitudes, 0-700 m, very rarely up to 1000 m. Uses. In Besuki (E. Java) and Gorontalo (Celebes) the leaves are used for making hats and mats. Vern. Bélumbungan, korisan, ombok, pérumbungan, rombok, si marappang-appang, umbut, M, babeleni, lilisungan, rumput adem, walingi, S, luru, sukét dékéng, J, bura, wlingi, Mad., pea-pea, tintilohuangga, Celebes: Philip.: kobong-kobong. Bis. 3. Cyperus ohwii KUK. in Fedde, Rep. 29 (1931) 197: Pfl. R. Heft 101 (1935) 60, f. 8 A-E: OHw1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 127; KERN, Rein- wardtia 2 (1952) 97, f. 1; KoyAmMa, Nat. Canad. 82 (1955) 205; Contr. Inst. Bot. Un. Montréal n. 70 (1957) 23; Quart. J. Taiwan Mus. 14 (1961) 166; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 479. — C. elatus L. var. macronux CLARKE, J. Linn. Soc. Bot. 21 (1884) 190; FI. Br. Ind. 6 (1893) 618; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 136; KUx. Pfl. R. Heft a (1935) 60, p.p., non CAMUS, FI. Gén. I.-C. 7 (1912) Closely allied to C. elatus. Usually more robust. Leaves often broader, up to 18 mm wide. Clado- prophylls with smooth keel. Spikes very dense (usually c. 70 spikelets to the spike), 6-8 mm (ulti- mately up to 10mm) wide. Spikelets somewhat broader, c. 1/4 mm wide, obliquely patent. Wings of the rachilla lanceolate, /4 mm _ wide. Glumes ovate to elliptic, (1/4-)124-2 by 1-1)4 mm. Anthers with a large, setulose appendage of the connective about half as long as the anther-cells. Nut oblong-ellipsoid, c. UZ by 7% mm. Distr. Insufficiently known, probably rare, though widely distributed in SE. Asia: Bengal, Indo-China; in Japan (Kyushu) probably introduced; in Malesia: W. Java (Danau swamp and Rawah Tembaga near Bekasi). Ecol. In swamps, swinging bogs, at low altitudes. 4. Cyperus exaltatus ReETz. Obs. 5 (1789) 11; VAHL, En. 2 (1806) 366; Mia. Fl. Ind. Bat. 3 (1856) 276; Boeck. Linnaea 36 (1870) 319; BENTH. FI. Austr. 7 (1878) 285; CLarkE, J. Linn. Soc. Bot. 21 (1884) 186, incl. var. amoena CLARKE; FI. Br. Ind. 6 (1893) 617; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 139, t. 4 f. 40; ? CLaRKE, Philip. J. Sc. 2 (1907) Bot. 86 [cef. Merr. En. Philip. 1 (1923) 109]; Koorp. Exk. FI. FLORA MALESIANA [ser. I, vol. 7° Java 1 (1911) 191; ibid. 4 (1922) f. 227; Camus, FI. Gén. I.-C. 7 (1912) 73, excl. var. digynus WILLIAMS; KUxK. Pfl. R. Heft 101 (1935) 64, f. 9 A-F; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 37; KERN, Reinwardtia 2 (1952) 99, f. 2; in Back. & Bakh. f. Fl. Java 3 (1968) 480.— C. venustus R.BR. Prod. (1810) 217; Mig. Fl. Ind. Bat. 3 (1856) 280; RiDv. in Forbes, Wand. (1885) 520. — C. altus NEES in Wight, Contr. (1834) 84; Mia. Fl. Ind. Bat. 3 (1856) 276. — C. digitatus (non ROxB.) VALCK. SuR. Nova Guinea 8 (1912) 699, p.p.—C. elatus var. macronux (non CLARKE) CaMus, FI. Gén. I.-C. 7 (1912) 72. Perennial with very short rhizome. Stems tall, densely tufted, trigonous, smooth, 100—180 cm by up to 8mm. Leaves flat or plicate with the midrib prominent beneath and 2 lateral nerves strongly marked above, coriaceous, scabrous on the margins and scaberulous on the upper surface, bright green above, pale green beneath, 8-15 mm wide; lower sheaths somewhat spongy, reddish to blackish brown. Inflorescence compound or decompound, up to 25 cm long. Involucral bracts 4—6, erecto-patent, up to 80 cm long. Primary rays 6—9, slender, unequal, smooth, up to 18cm long, secondary ones up to 5cm. Spikes digitately arranged, usually some of them solitary on slender peduncles, cylindric, loose to rather dense (up to c. 40 spikelets to the spike), finally 2-5 by 1-114 cm; rachis visible among the spikelets, somewhat flexuous, glabrous, narrowly winged. Spikelets spicately arranged, subdistichous, obliquely patent or almost at right angles to the rachis, oblong to linear, compressed, 10—20-flowered, 5-10 by VA-2 mm; rachilla straight, persistent; internodes c. cA mm; wings caducous, narrowly oblong, whitish or yellowish. Glumes membranous- chartaceous, patent, ovate or broadly ovate, obtuse, mucronate, 14-2 by 1)4-1/4 mm, 4 imbricate; keel green, 3—5(—7)-nerved especially above, sides nerveless or obscurely striate, shining brown or greenish. Stamens 3; anthers oblong, 4-74 mm long; connective shortly produced, smooth. Stigmas 3. Nut trigonous, ellipsoid, shortly apiculate, yellow or yellowish brown, ?4—% by 4-14 mm. Distr. Scattered in tropical Africa; rather common from India to Indo-China, E. China, and Japan, southward extending to S. Australia; doubtful for tropical S. America; in Malesia very rare: a few times collected in the Malay Peninsula (Singapore, Penang), W. Java (near Djakarta), the Lesser Sunda Is. (Timor), and W. New Guinea (Merauke). Ecol. In wet places at low altitudes: embankments in tidal forests, grassy swamps behind the beach-wall, in Timor in grass-flats at 400 m. Notes. Closely allied to C. imbricatus, from which it is sometimes difficult to be distinguished, as both are very polymorphous. C. exaltatus is often more robust, its spikes are less dense and peduncled, and the glumes more closely imbricate. From C. digitatus, also with patent spikelets, it can be distinguished by the broader spikelets, the broader, mucronate glumes, and the smaller nuts. KOYAMA opposed the E. Asian-Australian plants to the typical Indian ones as var. iwasakii (MAKINO) Koyama, Act. Phytotax. Geobot. 16 (1955) 11 [C. venustus R.BR. l.c.; C. iwasakii MAKINO, Bot. Mag. Tokyo 17 (1903) 49]. I am unable to draw a line between the Indian forms and those from Korea, Japan, China. Malesia. and Australia. 1974] Cy PERACEAE (Kern) The records for the Malay Peninsula are old: Singapore, 1853, ANDERSSON; Penang, 1904-06, BircH. The specimen from Sumatra, WaItz, mentioned by VALCKENIER SURINGAR, 1898, l.c., is much too young for identification. 5. Cyperus imbricatus Retz. Obs. 5 (1789) 12; LLANos, Fragm. Pl. Filip. (1851) 17; F.-ViLL. & NAVES in Blanco, FI. Filip. ed. 3, 4 (1880) 11; FIscHErR, Kew Bull. (1931) 262; KUk. Pfl. R. Heft 101 (1935) 69, f. 9 G-J, incl. var. elongatus KUk.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 37; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 480. — C. radiatus VAHL, En. 2 (1806) 369; Mia. FI. Ind. Bat. 3 (1856) 277; BoEcK. Linnaea 36 (1870) 317, incl. var. elongatus BoEcK.; CLARKE, J. Linn. Soc. Bot. 21 (1884) 185; Fl. Br. Ind. 6 (1893) 617; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 137, t. 4 f. 38, 39; CLARKE, Philip. J. Sc. 2 (1907) Bot. 85; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 70; Koorp. Exk. Fl. Java 1 (1911) 191; ibid. 4 (1922) f. 228; Brown, Min. Prod. Philip. For. 1 (1920) 348; Merr. En. Philip. 1 (1923) 107: Rip. Fl. Mal. Pen. 5 (1925) 147; Back. Onkr. Suiker. (1928) 140, t. 139. — C. involucratus Por. in Lamk, Encycl. 7 (1806) 253; DecNE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 360; Descr. Herb. Timor. (1835) 32.— C. verticillatus Roxs. F1. Ind. (1820) 209; Mia. Fl. Ind. Bat. 3 (1856) 276.— C. spicatus PRESL, Rel. Haenk. 1 (1828) 173; STEUD. Syn. 2 (1855) 52; Mia. Fl. Ind. Bat. 3 (1856) 287; NAveEs, Nov. App. (1882) 306; CLARKE, Philip. J. Sc. 2 (1907) Bot. 87; Kuk. Pfi. R. Heft 101 (1935) 73. — C. race- mosus RETZ. var. spiculis infuscatis NEES in Hook. J. Bot. Kew Misc. 6 (1854) 27.— C. obscurus NEES, l.c., in synon.— C. macrosciadion STEUD. Syn. 2 (1855) 37; Mig. Fl. Ind. Bat. 3 (1856) 277; NAveEs, Nov. App. (1882) 305.— C. anabaptistus STEUD. Syn. 2 (1855) 37; Mia. Fl. Ind. Bat. 3 (1856) 277; Naves, Nov. App. (1882) 305.— Dichostylis radiata PALLA, Allg. Bot. Zeitschr. 17, Beil. (1912) 8. Perennial with very short rhizome. Stems slender to robust, tufted, trigonous, smooth, 50—100(—150) cm by up to 8 mm. Leaves canaliculate, coriaceous, scabrous on the margins, pale green beneath, 5—10(—15) mm wide; lower sheaths somewhat spongy, stramineous to purplish black. Inflorescence com- pound or subdecompound, up to 20cm long and wide. Involucral bracts 3—5, obliquely erect to patent, up to 65 cm long. Primary rays 6—8, unequal, 603 stiff, smooth, up to 15 cm; secondary rays usually very short, rarely up to 3cm. Spikes digitately arranged, cylindric, very dense (usually 60—70 spike- lets to the spike), sessile or subsessile (rarely 2—3 cm peduncled), 2-3 cm by 5—10 mm; rachis glabrous, narrowly winged, hidden by the spikelets or almost so. Spikelets spicately arranged, suberect, rarely obliquely erect, oblong to oblong-linear, imbricate, compressed, 6—20(—30)-flowered, 4—6 by finally up to 2mm; rachilla straight, persistent; internodes )<- /4™mm; wings narrow, oblong, whitish hyaline, long-persistent. Glumes membranous, at first ap- pressed, finally obliquely patent, ovate to broadly ovate, obtuse, mucronate, without mucro 1-1/4 by c. 1mm, 4-% imbricate; keel green, faintly 3- nerved, sides fulvous to spadiceous, nerveless. Sta- mens 3; anthers oblong, c. 4 mm; connective shortly produced, smooth. Stigmas 3. Nut trigonous, slightly compressed dorsoventrally, ovoid to ellipsoid, shortly apiculate, yellowish brown, 0.6—0.8 by 0.4 mm. Distr. Pantropical; widely distributed in Malesia, not yet known from the Moluccas. Only recently collected in New Guinea; “Guinea” in MIQUEL, l.c. 277, does not relate to New Guinea, as VALCKENIER SURINGAR and KUKENTHAL supposed. Ecol. In swamps, wet rice-fields, on river-banks, and in other open wet places, at low altitudes, rarely up to 900 m. Uses. According to BROWN (1920) /.c., in some parts of the Philippines the outer portions of the stems are stripped, dried in the shade, and used for weaving mats, mattings, and screens. Vern. Bendong, N. Borneo, tintilo, Gorontalo; Philip.: alinang, dagk6é, 6bod-dbod, Bis., balabal- anguton, Tag., balayang, I\k., upopi, Ibn. Notes. A highly polymorphous species. The common Malesian form is figured in VALCKENIER SURINGAR’s t. 4 f. 38b, whereas the plant of f. 38a (not distinguished nomenclaturally) is very rare in Malesia (e.g. BACKER 8131, 24423). The two forms are connected by many intermediates. The latter one is possibly identical with C. imbricatus var. dense- spicatus (HAYATA) OHWI, Mem. Coll. Sc. Kyoto Imp. Un. B18 (1944) 128.— C. densespicatus HAYATA, Ic. Pl. Form. 6 (1916) 105. SURINGAR’s f. 39 represents an anthelule of a specimen determined by CLARKE as C. exaltatus var. amoena CLARKE, but which belongs to C. imbricatus, as was already pointed out by SURINGAR. 2. Section Alopecuroidei CHERM. Fl. Madag. fam. 29 (1937) 138. Type species: C. alopecuroides ROTTB. 6. Cyperus alopecuroides Rotts. Descr. Pl. rar. Progr. (1772) 20; Descr. & Ic. (1773) 38, t. 8 f. 2; Kunth, En. 2 (1837) 19; Mig. Fl. Ind. Bat. 3 (1856) 261; Boeck. Linnaea 36 (1870) 321 (var. 6 digynus, excl. var. « et y); BENTH. Fl. Austr. 7 (1878) 264; CLARKE, J. Linn. Soc. Bot. 21 (1884) 30, 74, t. 2 f. 12; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 72, t. 2 f. 21; Koorp. Exk. Fl. Java 1 (1911) 186, non ibid. 4 (1922) f. 200; KUx. Pfil. R. Heft 101 (1935) 71; Bull. Jard. Bot. Btzg III, 16 (1940) 300; in Fedde, Rep. 53 (1944) 101; KERN, Reinwardtia 2 (1952) 101; in Back. & Bakh. f. Fl. Java 3 (1968) 472. — Juncellus alopecuroides CLARKE, Fl. Br. Ind. 6 (1893) 595; Camus, FI. Gén. I.-C. 7 (1912) 38; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 42. Perennial with very short rhizome. Stems stout, tufted, trigonous, smooth, leafy at the much thickened base, up to 125 by 1 cm. Leaves flat or somewhat plicate with the midrib prominent beneath and 2 lateral nerves strongly marked above, coriaceous, scabrous on margins and nerves, 6-16 mm wide; lower sheaths spongy, cinnamomeous to fuscous. Inflorescence compound or decompound, up to 30 cm long. Involucral bracts 4-7, much overtopping 604 FLORA MALESIANA [ser. I, vol? the inflorescence, erecto-patent, up to 65 by 1 cm. Primary rays 5—10, unequal, smooth, suberect, rigid, up to 20 cm; secondary ones usually very short, up to 6cm. Spikes digitately arranged, oblong- cylindrical to cylindrical, divergent, very dense, 2-4 cm by 8—15 mm; rachis hidden by the densely crowded spikelets. Spikelets spicately arranged, ulti- mately at right angles to the rachis, somewhat turgid, ovate to oblong-lanceolate, 12—30-flowered, 4—6(—8) by 2-274 mm; rachilla straight, strongly compressed, persistent; internodes 4-14, mm; wings very narrow, long-persistent, whitish. Glumes membranous, con- cave, not keeled, with finally inrolled margins, obliquely patent, finally wide-spreading, ovate or elliptic-ovate, obtuse, apiculate or minutely mucro- nulate, 5—7-nerved in the centre, yellowish or golden with purple stripes, V4-2% by 1-14 mm, very densely (24-34) imbricate. Stamens 2(-3); anthers linear-oblong, 74-1 mm _ long; connective shortly produced, smooth. Stigmas in most flowers 2, not rarely 3 in some flowers of the same inflorescence. Nut dorsoventrally compressed, planoconvex or (in the trigynous flowers) with a raised dorsal angle, elliptic to slightly obovate, shortly apiculate, golden yellow or stramineous, 0.8—0.9 by 0.5—0.6 mm. Distr. In Africa from the Mediterranean region to the tropics, Madagascar; from India and Farther India extending to Queensland; West Indies (Guade- loupe); in Malesia very rare: Central Java (Depok E. of Jogjakarta, leg. JUNGHUHN, not recently collected), Lesser Sunda Is. (Bali, Sumba, Timor); not known from the Malay Peninsula (KUKENTHAL’s record, 1935, l.c., is erroneous). Ecol. In swamps and along margins of lakes, from the lowlands up to 1000 m (Bali). 3. Section Cyperus Cyperus sect. Bulbosi CLARKE, FI. Br. Ind. 6 (1893) 611.— Cyperus sect. Rotundi CLARKE, l.c. 614. — Cyperus sect. Tunicati CLARKE in Thiselt.-Dyer, Fl. Trop. Afr. 8 (1902) 314. — Cyperus sect. Esculenti KUKk. Pfl. R. Heft 101 (1935) 116. Type species: C. esculentus L. 7. Cyperus rotundus LINNE, Sp. Pl. (1753) 45; HAssk. Pl. Jav. Rar. (1848) 80; Mig. Fl. Ind. Bat. 3 (1856) 274: Boeck. Linnaea 36 (1870) 283; BENTH. Fl. Austr. 7 (1878) 279, p.p.; CLARKE, J. Linn. Soc. Bot. 21 (1884) 167, t. 2 f. 16; Fl. Br. Ind. 6 (1893) 614; VALcK. Sur. Ges]. Cyp. Mal. Arch. (1898) 107, t. 4 f. 23; CLARKE, Philip. J. Sc. 2 (1907) Bot. 85; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 68; Koorpb. Exk. Fl. Java 1 (1911) 190 (‘rotundatus’); ibid. 4 (1922) f. 216; Camus, Fl. Gén. I.-C. 7 (1912) 65; Back. Trop. Natuur 9 (1920) 174, f. 2; Merr. En. Philip. 1 (1923) 107; RiDL. Fl. Mal. Pen. 5 (1925) 145; Back. Onkr. Suiker. (1928) 135, t. 132; KUx. Pfl. R. Heft 101 (1935) 107, f. 13; S. T. BLAKE, Univ. Queensl. Pap. 2, 2 (1942) 8, t.4; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 33; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 479.— C. rotundus I bulbosus sive legitimus RUMPH. Herb. Amb. 6 (1750) 2, t. 1 f. 2. — Schoenus tuberosus Buro. f. Fl. Ind. (1768) 19. — C. curvatus (non VAHL) LLANos, Fragm. Pl. Filip. (1851) 15; F.-VILL. & Naves in Blanco, Fl. Filip. ed. 3, 4 (1880) 9.— Chlorocyperus rotundus PALLA, Allg. Bot. Zeit. 6 (1900) 201.— C. longus (non L.) K.ScH. & Laut. Fl. Schutzgeb. (1901) 192; VALCcK. Sur. Nova Guinea 8 (1912) 698.— C. bulbosus (non VAHL) Camus, FI. Gen. I.-C. 7 (1912) 64, quoad specim. cochinch. — Fig. 49. ssp. rotundus. — Synonymy as above. Perennial. Rhizome emitting long, slender, wiry stolons ending in a subglobose or ellipsoid, not or obscurely zoned, finally blackish tuber; scales of rhizome soon disintegrating and usually not present on last year’s growth. Stems slender, triquetrous, smooth, tuberous at the ULase, 15—30(—75) cm by 1-2 mm. Leaves several, flat, rigidulous, scabrid on the margins in the upper part, gradually acuminate, dark green above, light green beneath, 2-6 mm wide, lower sheaths reddish brown, soon disintegrating. Inflorescence simple or compound, up to 15 by 10 cm, usually much smaller. Involucral bracts 2—4(-—6), patent, as long as or overtopping the inflorescence, up to 30cm. Primary rays 3-9, very unequal, obliquely erect to spreading, slender, up to 10 cm. Spikes ovoid, loose to rather dense, with glabrous rachis. Spikelets spicately arranged, obliquely patent, linear, acute, strongly compressed, 10—40-flowered, sometimes strongly accrescent and up to 100- flowered, 1-3/4 cm by c. 2mm; rachilla flexuous, broadly winged, persistent; internodes c. 4% mm; wings persistent, c. 14mm wide. Glumes membra- nous, obliquely erect, ovate, keeled, subobtuse, rubiginous to deep brown, with narrow hyaline margins, 5—7-nerved over ee Y, Of either side (the midnerve disappearing or minutely excurrent just below the apex), with green keel more or less curved and slightly recurved at the top, 3-3/4 by c. 2mm, 4 imbricate. Stamens 3; anthers linear, c. 1 mm long, with small, smooth, reddish appendage of the con- nective. Stigmas 3. Nut trigonous, oblong-obovoid, apiculate, brownish to black, 144 by 4-4 mm, rarely maturing (plants reproducing almost exclu- sively by stolons). Distr. Widely distributed in the warmer parts of the whole world; very common throughout Malesia. Ecol. In sunny or lightly shaded localities: in lawns, along roads, in waste places, at low and medium altitudes, up to c. 1000 m. Often a serious pest in cultivated land. Uses. The young tubers are sometimes eaten and used in native medicine. The leaves furnish a rather good pasturage. Vern. Nut grass, E, téki,M,S,J, djukut beuti, muta, S, tjébuka, J, mota, Md, manggata, Gorontalo, rukut téki, wuta, Alf. Minah., kareha wai, Sumba, rumput haliya hitam, Mal. Pen.: Philip.: barsangd, 1974] CYPERACEAE (Kern) 605 Ilk., boto-botones, tarugug, Bik., galonalpas, kusung, malaapilid, motad, omadiung, onoran, sur-sur, Pamp., mutha, Tag. Notes. Var. salsolus CLARKE, J. Linn. Soc. Bot. 21 (1884) 171, is a small, coastal form. The Malesian specimens cited are to me normal C. rotundus. Specimens in which the spikelets are exceedingly elongated, up to 5 cm long, and up to 100-flowered, have been described as var. centiflorus CLARKE, l.c. (? var. elongatus BoEcK. Linnaea 36, 1870, 285). The strong accrescence of the spikelets is probably due to the fact that the flowers do not set fruit. The variety has no-taxonomical value. For C. distans x C. rotundus see p. 660. ssp. retzii (NEES) KUk. Pfl. R. Heft 101 (1935) 114, incl. var. disruptus KUK.— C. retzii NEES in Wight, Contr. (1834) 82; S. T. BLAKE, Univ. Queensl. Pap. 2, 2 (1942) 8, t. 5, non Poir. 1806. — C. rotundus var. pallidus BENTH. Fl. Austr. 7 (1878) 280, p.p.— C. disruptus CLARKE, Kew Bull. add. ser. 8 (1908) 12. — C. bifax CLARKE, I.c. 13.— C. rotundus var. pseudesculentus KUK. Bot. Jahrb. 59 (1924) 44. — °” C. weinlandii KUxK. Pfl. R. Heft 101 (1935) 131. Differs from ssp. rotundus by its stouter habit (stems 50-75 cm tall); the scales of the stolons usually persisting through the second year; the some- what broader spikelets (c. 2/4 mm wide when ripe): the thinly membranous, elliptic-oblong, 314-4 mm long, paler (usually ferrugineous) glumes, which soon spread and inroll, and are striate on either side for 4-74 of the breadth; the straight keel of the glumes; and its freely fruiting. Distr. India; widely spread in Australia; accord- ing to KUKENTHAL, /.c., in tropical S. America and widely spread in Africa; in Malesia: Lesser Sunda Is. (Alor), E. New Guinea, New Britain; a specimen collected in the Bogor Botanic Gardens on a compost heap may be introduced. Ecol. In moist localities: creek flats, silty flood- banks, sometimes as a weed, but never a pest. Notes. Difficult to distinguish from stout forms of C. rotundus ssp. rotundus, and hardly if at all from C. rotundus ssp. tuberosus (ROTTB.) KUK. (C. tuberosus Rotts. Descr. Pl. rar. Progr. 1772, 18; Descr. & Ic. 1773, 18, t. 7 f. 1). See S. T. BLAKE, I.c. 13, who takes C. rotundus, C. retzii, and C. tuberosus for separate species. The bright colour of the spikelets is similar to that of C. esculentus, with which species C. rotundus ssp. retzii is often confused. 8. Cyperus bulbosus VAHL, En. 2 (1806) 342; Mia. FI. Ind. Bat. 3 (1856) 274; Borck. Linnaea 36 (1870) 300: CLARKE, Fl. Br. Ind. 6 (1893) 611: VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 116, t. 4 f. 27, t. Sf. 8: Hook. f. in Trimen, Handb. Fl. Ceyl. 5 (1900) 22, t. 96; KUK. Pfl. R. Heft 101 (1935) 125, t. 15 f. C-E; S. T. BLAKE, Univ. Queensl. Pap. 2, 2 (1942) 9, t. 8: KERN, Reinwardtia 2 (1952) 107; in Back. & Bakh. f. Fl. Java 3 (1968) 478.— C. jeminicus (non ROTTB.) CLARKE, J. Linn. Soc. Bot. 21 (1884) 175, t.2 f. 17-18. Perennial. Stolons capillary, disappearing after having formed ovoid to fusiform, at first whitish, ultimately shining black, bulb-like, 1-1/4 cm long tubers; coat of tubers coriaceous, striate, splitting into some caducous, blackish segments. Stems arising from a tuber, slender, triquetrous, smooth, (S—)15— 30 cm by 4-1 mm. Leaves several, often recurved, as long as or longer than the stem, gradually acumi- nate, scaberulous in the upper part, 1—2(—4) mm wide. Inflorescence simple, often imperfect (reduced to a spike) or with a few short rays, 2—3 by 1—3(—S) cm. Involucral bracts (1—)2—3, patent to reflexed, 1-2 mm spaced or only the lowermost obvious, usually overtopping the inflorescence, up to 10 cm. Spikelets spicately arranged, patent, linear, compressed, 8—28- flowered, 1—3cm by c. 2mm; rachilla flexuous, broadly winged, persistent; wings persistent; inter- nodes c. 1 mm. Glumes chartaceous, obliquely erect, ovate to oblong-lanceolate, subacute, muticous or minutely mucronulate, 9—11-nerved, with green keel and shining reddish to castaneous sides, (3—)4 by 2mm, 4 imbricate. Stamens 3; anthers linear, 1/4-2 mm, with short, smooth, red appendage of the connective. Stigmas 3. Nut trigonous, obovoid to ellipsoid, obtuse, apiculate, ultimately black, c. 14 by 4 mm. Distr. Tropical Africa, S. Asia, tropical Australia ; in Malesia very rare: W. Java (Island of Damar Besar in Bay of Djakarta), Madura, Lesser Sunda Is. (Timor). Ecol. On dry sandy soil, in Malesia only near the sea. Notes. In the absence of the characteristic tubers, which is often the case in herbarium specimens, C. bulbosus may easily be confused with C. rotundus, but can be distinguished by the imperfection of the inflorescence, the spaced lower bracts, and the more distinctly nerved glumes. In countries where the species is common the young tubers are eaten. 9. Cyperus esculentus LINNE, Sp. PI. (1753) 45; BoEcK. Linnaea 36 (1870) 287; CLARKE, J. Linn. Soc. Bot. 21 (1884) 178; Fl. Br. Ind. 6 (1893) 616; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 120; KUK. Pfi. R. Heft 101 (1935) 116; S. T. BLAKE, Univ. Queensl. Pap. 2, 2 (1942) 9. Perennial. Stolons several, very slender, yellowish, clothed with ovate-lanceolate, acute, pale scales ending in ovoid to globose, c. 1 cm thick tubers which are transversely zoned when young and covered with a grey tomentum when ripe; after the tubers have been formed the stolons often disappear. Stems slender, rigid, triquetrous, smooth, 10-50 cm by 1—2 mm. Leaves several, somewhat shorter or longer than the stems, rather rigid, gradually acuminate, 3-6 mm wide; lower sheaths stramineous to reddish brown. Inflorescence simple or compound, rather loose to dense, (3—)5—10(—18) by (2—)4—6(—10) cm. Involucral bracts 3—6, obliquely patent, usually the lower 1-2 much overtopping the inflorescence. Primary rays 3—8, unequal, smooth, 3—10(—15) cm long. Spikes ovoid, with few to numerous spikelets: rachis glabrous. Spikelets spicately arranged, divari- cate, oblong-linear, subcompressed, obtuse, 8—16- flowered, 5—18 by c. 2 mm; rachilla slightly flexuous, broadly winged; internodes >4—1 mm. Glumes mem- branous, obliquely erect, hardly keeled, ovate to elliptic, very obtuse, sometimes minutely mucron- ulate, golden yellow to pale brown with whitish hyaline margins especially above, distinctly 7-nerved over their whole breadth, 24-3 by 2-2/4 mm, 14-¥% imbricate. Stamens 3; anthers linear, 1/4- 2mm, with short, smooth, red appendage of the 606 FLORA MALESIANA [ser. I, vol. 7? connective. Stigmas 3. Nut trigonous, obovoid to oblong-obovoid, obtuse, hardly apiculate, c. 1/4 by 1 mm. Distr. Widely distributed: from the Mediterranean region to S. Africa and through India and Farther India to N. Queensland; abundant in America; also cultivated; in Malesia only known from a single, suspected collection: E. Java, Tosari 1915, RIDLEY; see under Carex divulsa and C. muricata (C. pairaei). Ecol. “In fields along road”’. Note. In many countries the tubers, which contain sugar and oil, are roasted and eaten. Cultivation is not reported from Malesia. 10. Cyperus stoloniferus RETz. Obs. 4 (1786) 10; PRESL, Rel. Haenk. 1 (1828) 169; Mig. FI. Ind. Bat. 3 (1856) 265; Boeck. Linnaea 35 (1868) 489, excl. var.; Naves, Nov. App. (1882) 302; CLARKE, J. Linn. Soc. Bot. 21 (1884) 172; Fl. Br. Ind. 6 (1893) 615; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 115, t. 4 f. 26, t. 5 f. 5; CLARKE, Philip. J. Sc. 2 (1907) Bot. 85 (‘stoloni- fer’); Ill. Cyp. (1909) t. 19 f. 1-3; Rip. Mat. Fl. Mal. Fig. 50. Cyperus stoloniferus Retz. on the sandy beach near Kuala Takadai (Endau, eastcoast of Johore), occurring from below mean highwater level to about highwater springtide level. The trees are tjemara (Casuarina equisetifolia) (photogr. H. M. BURKILL, 1960). 1974] Cy PERACEAE (Kern) 607 Pen. (Monoc.) 3 (1907) 69; Koorp. Exk. Fl. Java 1 (1911) 190; ibid. 4 (1922) f. 218; Back. Trop. Natuur 9 (1920) 173, f. 1; MeRR. En. Philip. 1 (1923) 108; Ripe. Fl. Mal. Pen. 5 (1925) 145; Kuk. Pfl. R. Heft 101 (1935) 106; S. T. BLAKE, Univ. Queensl. Pap. 2, 2 (1942) 9, t. 7; J. Arn. Arb. 28 (1947) 213; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 33; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 478.— C. litoralis R.BR. Prod. (1810) 216.—C. tuberosus (non ROTTB.) KUNTH, En. 2 (1837) 50; Mor. Syst. Verz. (1846) 96.— C. lamprocarpus NEES in Hook. J. Bot. Kew Misc. 6 (1854) 27; Boeck. Linnaea 35 (1868) 490.— C. bulboso-stoloniferus StTEup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 18; Mia. Fl. Ind. Bat. 3 (1856) 266. — C. rotundus (non L.) BENTH. Fl. Austr. 7 (1878) 279, p.p. — C. bulbosus (non VAHL) CAMUS, FI. Gén. I.-C. 7 (1912) 64, incl. var. elatus CAMus, excl. specim. cochinch. — C. mayeri KU«. in Fedde, Rep. 29 (1931) 194; Pfl. R. Heft 101 (1936) 162, ex descr.—C. carrii KUK. Bot. Jahrb. 69 (1938) 256. — Fig. 50. Perennial. Stolons long-creeping, hardening into a woody rhizome forming stout, irregular tubers. Stems slender, distant, rigid, triquetrous, smooth, distinctly incrassate at the base, 15-50 cm by 1—2(—3) mm. Leaves gradually acuminate, scabrid in the upper part, glaucous, in dry localities rigid and narrow (1/4 mm), in wet places more flaccid and up to 4 mm wide. Inflorescence simple or subcompound, dense to rather loose, often small, rarely up to 6 by 5 cm. Involucral bracts 2—3, suberect, the lower 1 or 2 (much) overtopping the inflorescence, sometimes up to 30 cm. Primary rays 2—S, obliquely erect, very unequal, smooth, 1—6 cm. Spikes ovoid or broadly ovoid, 114-214 cm long and wide; rachis very short, glabrous. Spikelets spicately arranged, 3-1 to the spike, patent, linear-lanceolate. often some- what curved, turgid, almost terete, 8—20-flowered, 5-15 by 2-2'4 mm; rachilla slightly flexuous, broadly winged. persistent; internodes /4~7%, mm; wings hyaline, long-persistent. Glumes membranous, obliquely erect, ovate or broadly ovate. obtuse. not keeled, indistinctly 5—7-nerved, 2/4—3 by 2-2/4 mm, 74 imbricate; midnerve green, sides ferrugineous to rubiginous, margins (often glistening) white or yellowish hyaline in the upper part. Stamens 3; anthers linear, up to 2mm, with short, smooth, reddish appendage of the connective. Stigmas 3. Nut strongly dorsoventrally compressed, broadly ovate to ovate, convex with a raised angle on the dorsal side, often somewhat concave on the ventral side, obtuse, shining, dark brown to blackish, 1-1/4 by c. 1mm. Distr. Mauritius, Madagascar; from SE. Asia to Melanesia and Queensland; throughout Malesia, in the Philippines rare and local (Luzon, Panay). Ecol. On coastal sands, often acting as a sand- binder on dunes and beaches, occasionally also in salt, muddy places. Fig. 50. 4. Section Corymbosi KUNTH, En. 2 (1837) 53. — Cyperus sect. Brevifoliati CLARKE, F1. Br. Ind. 6 (1893) 611. Type species: C. corymbosus ROTTB. 11. Cyperus scariosus R.BR. Prod. (1810) 216; CLARKE, J. Linn. Soc. Bot. 21 (1884) 159; Fl. Br. Ind. 6 (1893) 612; Ill. Cyp. (1909) t. 15 f. 2-3; S. T. BLAKE, Univ. Queens]. Pap. 2, 2 (1942) 9, t. 2: KERN, Rein- wardtia 2 (1952) 103, f. 3; non VALCK. SuR. Nova Guinea 8 (1912) 699.— C. rotundus L. var. pallidus BENTH. FI. Austr. 7 (1878) 280, p.p.— C. diphyllus (non Retz.) VALCK. Sur. Nova Guinea 8 (1912) 698.— C. corymbosus Rotts. var. scariosus KUK. Pfl. R. Heft 101 (1935) 83. Perennial with creeping rhizome; stolons slender, rather short, up to 5cm, covered with lanceolate, reddish brown, striate sheaths, forming small tubers. Stems slender, almost terete in the lower part, gradually becoming trigonous upwards, somewhat incrassate at the base, rigid, smooth, 50-80 cm by 14-2 mm. Leaves few, up to 3, very short, rarely half as long as the stems, weak, gradually acuminate, scaberulous at the triquetrous top, 14-12 cm by c. 2 mm; lower sheaths stramineous to purplish. Inflo- rescence small, simple, usually thrown to one side by the erect lowermost bract, with short rays or reduced to a single spike, 2-5 cm long and wide. Involucral bracts much shorter than the inflorescence when leaves short, exceeding the inflorescence when leaves longish, erect or suberect, /4—6 cm, embracing the stem with minute, dark brown, scarious auricles. Primary rays filiform, erect or suberect, with few spike- lets, 2-3 cm. Spikelets spicately arranged. obliquely erect to patent, linear, strongly compressed, up to 26-flowered, 1—2(-2)4)cm by 2-2/4 mm; rachilla rather flexuous, broadly winged, persistent: internodes c. °4 mm; wings whitish hyaline, long- persistent. Glumes membranous, erecto-patent, ovate, obtuse, muticous, 2°4 by 1/4(-2) mm, /% imbricate; keel green, 3-nerved; sides faintly 3—4-nerved, stra- mineous, tinged with red; upper margin narrowly banded with brown. Stamens 3; anthers linear, c. 1/4 mm; connective shortly produced, smooth. Stig- mas 3. Nut trigonous, slightly obovoid to ellipsoid, broadly stipitate, shortly apiculate, brown, 1)4-114 by c. 4% mm. Distr. From Bengal and Farther India to tropical Australia; in Malesia: South coast of W. New Guinea; Papua: Central Distr., Kairuku Subdistr. Ecol. Coastal species, almost restricted to swampy, brackish or near-brackish localities. 608 FLORA MALESIANA [ser. I, vol. 7° 5. Section Subimbricati CLARKE, FI. Br. Ind. 6 (1893) 612.— Cyperus sect. Zollingeriani CLARKE, Kew Bull. add. ser. 8 (1908) 100. — Cyperus sect. Subquadrangulares KUK. Pfl. R. Heft 101 (1936) 127. Type species: C. zollingeri STEUD. 12. Cyperus tenuiculmis Boeck. Linnaea 36 (1870) 286; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 50; NAvEs, Nov. App. (1882) 306; KERN, Reinwardtia 3 (1954) 30; in Back. & Bakh. f. Fl. Java 3 (1968) 481.— C. rotundus (non L.) PRESL, Rel. Haenk. 1 (1828) 175; Mig. Sum. (1861) 260, 600.— C. longus L. var. B Mia. Fl. Ind. Bat. 3 (1856) 275.— C. rotundus L var. carinalis BENTH. Fl. Austr. 7 (1878) 280.— C. lucidulus (non KLEIN) CLARKE, J. Linn. Soc. Bot. 21 (1884) 99. — C. zollingeri (non STEUD.) CLARKE, FI. Br. Ind. 6 (1893) 613; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 118, t. 4 f. 28, excl. ZOLLINGER 2689; RipL_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 68; CLARKE, Philip. J. Sc. 2 (1907) Bot. 85; Ill. Cyp. (1909) t. 8 f. 1-2; Koorp. Exk. Fl. Java 1 (1911) 190; ibid. 4 (1922) f. 219; Camus, Fl. Gén. I.-C. 7 (1912) 67, f. 7, 3-5; MeRR. En. Philip. 1 (1923) 108, excl. Ramos 7672; RIDL. Fl. Mal. Pen. 5 (1925) 144; KUk. Pfl. R. Heft 101 (1935) 133, incl. var. condensatus KUK., excl. ZOLLINGER 2689; S. T. BLAKE, J. Arn. Arb. 28 (1947) 214; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 37; KERN, Reinwardtia 2 (1952) 107, f. 6.—C. scariosus (non R.BR.) VALCK. Sur. Nova Guinea 8 (1912) 699, non al. Perennial with short-creeping, woody rhizome. Stems solitary or subcespitose, slender but rigid, triquetrous, smooth, 20—70(—100)cm by 1-2 mm, the incrassate base surrounded by some purplish- striate, bladeless sheaths. Leaves several, shorter than the stems, canaliculate, rigid, very gradually acuminate in a long, setaceous point, scabrid in the upper part, c. 3(—6) mm wide. Inflorescence simple or subcompound, open, sometimes reduced to a single cluster. Involucral bracts 3—7, erecto-patent, the longest as long as or overtopping the inflorescence, up to 20cm. Primary rays 4—10, obliquely erect, very unequal, slender, smooth, up to 20 cm; second- ary rays when present up to 3 cm. Spikes broadly ovoid, up to 5 by 6cm, with 3—10(—15) spikelets; rachis glabrous or slightly scabrid. Spikelets spicately arranged, at first suberect, finally patent to hori- zontally spreading, linear or linear-lanceolate, slightly compressed, subquadrangular (i.e. rhomboidal in cross-section), 8—20(-rarely more)-flowered, 2—3(—5) cm by c. 2 mm, the lowest of each spike often with a setaceous, up to | cm long bract; rachilla flexuous, broadly winged, persistent, blackish, internodes c. 1% mm; wings oblong, caducous, whitish hyaline, YA mm wide. Glumes chartaceous, appressed, keeled, ovate or elliptic, obtuse or acutish, muticous or minutely mucronulate, 7—9-nerved, (3—)3)4(—4) by 2-2/4 mm, remote (not or scarcely imbricate); keel broad, green, sides golden yellow to fulvous, margins narrow, hyaline. Stamens 3; anthers linear, 1-1/4 mm long, the connective hardly produced. Stigmas 3. Nut triquetrous, ellipsoid or slightly obovoid, broadly rr ae aciy : apiculate, castaneous to blackish, 4-4-2 by %-1 mm. Distr. Tropical Africa, S. and SE. Asia to Micro- nesia and Australia; throughout Malesia, but every- where scattered. Ecol. In open grassland, savannahs, fallow fields, along grassy road-sides, sometimes in wet places in forests, on rather dry to swampy soil, 0—1600 m. Vern. Siani, Sum. W. C., kékuda, teleles, Gajo- lands, diyangere, New Guinea (Maniki); Philip.: daat-laua, Tag., pakama, Bag. 13. Cyperus zollingeri Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 17; Mia. FI. Ind. Bat. 5 (1856) 264; Boeck. Linnaea 36 (1870) 352; KERN, Reinwardtia 3 (1954) 28; in Back. & Bakh. f. Fl. Java 3 (1968) 480. — C. ramosii KU«K. in Fedde, Rep. 21 (1925) 326; Pfl. R. Heft 101 (1935) 136; Bot. Jahrb. 69 (1938) 255; KERN, Reinwardtia 2 (1952) 109, f. 7.— C. rubroviridis CHERM. Bull. Soc. Bot. Fr. 66 (1919) 350; KUx. Pfl. R. Heft 101 (1935) 135, f. 16 G-J.—C. sphacelatus RotTTB. var. tenuior CLARKE, FI. Trop. Afr. 8 (1901) 347. Annual with fibrous roots. Stems slender, tufted, triquetrous, smooth, up to 30cm by /4-1 mm. Leaves few, shorter than the stems, flat, flaccid, very gradually acuminate, scabrid towards the top, grass-green or light green, 1-3 mm wide; lower sheaths bladeless, membranous, stramineous, pur- plish striate. Inflorescence simple, loose. Involucral bracts 3—6(—9), obliquely patent, the lowest as long as or slightly overtopping the inflorescence, up to 12 cm. Spikes consisting of 1 erect terminal spikelet and 1—3(—6) ultimately horizontally spreading lateral ones, often some reduced to a single spikelet; rachis glabrous. Spikelets spicately arranged, linear, acute, slightly compressed, subquadrangular (i.e. rhom- boidal in cross-section), 6—16-flowered, 1-274 cm by c. 14 mm; rachilla flexuous, broadly winged; per- sistent; internodes c. 1 44mm; wings rather firm, soon caducous, stramineous, often brownish lineo- late. Glumes appressed or somewhat recurved at the top, remote (hardly overlapping the next higher one), ovate or elliptic, acutish, 7—9-nerved, 3-3/4 by c. 2 mm, the slightly excurrent midnerve setulose at the top; keel broad, green, sides stramineous to brownish, purplish lineolate, margins hyaline. Stamens 3; anthers oblong, 4-34 mm. Stigmas 3. Nut trigonous, turbinate-obovoid, truncate or somewhat depressed at the apex, broadly stipitate, brown to blackish, VA-1Y% by %-1% mm. Distr. Tropical Africa, Madagascar, tropical Australia (N. Australia, Queensland), in Malesia very rare: E. Java (Puger, ZOLLINGER 2689, type coll. of C. zollingeri, not collected afterwards), Lesser Sunda Is. (NW. Bali: Perapatagung), Philippines (Luzon: Ilocos Norte Prov., RAMOs 7672, type coll. of C. ramosii; Coron I.), New Guinea (W. New Guinea: Kebar Valley; Papua: Hisiu, Port Moresby). Ecol. In Java collected on road-sides, in Bali in 1974] open forest on limestone rocks at 100 m, in New Guinea in open sandy places at sealevel and on the sea-shore. 14. Cyperus sphacelatus Rotts. Descr. Pl. rar. Progr. (1772) 21; Descr. & Ic. (1773) 26; BOECK. Linnaea 36 (1870) 292; CLARKE, J. Linn. Soc. Bot. 21 (1884) 183; KUKk. Pfl. R. Heft 101 (1935) 129: BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246. p. 39; KERN, Reinwardtia 2 (1952) 107, f. 5; Blumea 8 (1955) 162: in Back. & Bakh. f. Fl. Java 3 (1968) 481. Annual with fibrous roots. Stems slender to rather firm, tufted, triquetrous, smooth, 10—30(—60) cm by 1-2 mm. Leaves few, shorter than the stems. rather rigid, flat or slightly canaliculate, gradually acuminate, scabrid in the upper half on margins and underside of midrib, 2-4mm wide. Inflorescence simple or compound, open. Involucral bracts 3-5, obliquely spreading, the lower ones overtopping the inflorescence, up to 20 cm long. Primary rays 3—5(—8), unequal, erecto-patent, slender, smooth, up to 10(—15) cm; secondary rays when present very short, 44-1 cm. Spikes broadly ovoid, usually c. 2 by 3 cm, sometimes up to 5 by 8 cm, with glabrous rachis. Spikelets spicately arranged, 3-10 to the spike, ultimately widely spreading or the lower ones some- what reflexed, linear-lanceolate, acute, slightly com- pressed, subquadrangular (i.e. rhomboidal in cross- section), 10—20(—50)-flowered, 1—2(—4)cm by c. 2 mm; rachilla flexuous, broadly winged, persistent: internodes 4-1 mm; wings whitish hyaline, c. 44mm wide. Glumes membranous, obliquely erect, ovate to elliptic, subobtuse, sometimes minutely Cy PERACEAE (Kern) 609 mucronulate, 7—9-nerved, 2'4-3 by 1/4-2 mm, /4 imbricate; keel green; sides stramineous, usually with a purple spot; margins white-hyaline. Stamens 3; anthers oblong, 14 mm long. Stigmas 3. Nut triquetrous, concave on the ventral side, ellipsoid or slightly obovoid, broadly stipitate, minutely apiculate, brown, 1/4-1)4 by 4-1 mm. Distr. Widely distributed in tropical Africa and tropical America, elsewhere introduced (Coro- mandel, Ceylon, Queensland, Tahiti), in Malesia: N. Sumatra, Malay Peninsula (P. Penang, Johore, Singapore). Northern Borneo (Kuching. Jesselton, Brunei, Labuan I., Nunukan), W. New Guinea (McCluer Gulf. Merauke), New Britain (Gazelle Peninsula), obviously naturalising. Ecol. In grassy fields, often as a weed on air-strips, in Brunei collected in Shorea albida peat-swamp. Notes. The earliest Malesian collection may be that of VESTERDAL (‘“Malacca’’, without date). In Sumatra it was first collected in 1922, in the other islands more recently. It is said to be collected in E. Java (Lawang) as early as 1905 by MOLHUuysEN, but since MOLHUYSEN’s collections include several plants certainly not native to Java, this statement is very doubtful. The species is very similar to C. tenuiculmis; it can be distinguished by its annual habit, the smaller. distinctly imbricate glumes, the oblong anthers, the somewhat smaller nuts, and usually at once by the purple spots on either side of the glumes (sometimes on one side only, rarely lacking), together giving the impression of a purple stripe along the centre of the spikelet. 6. Section Distantes CLARKE, Kew Bull. add. ser. 8 (1908) 99.— Cyperus sect. Longispicati a. Distantes VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 123. Type species: C. distans L. f. 15. Cyperus nutans VAHL, En. 2 (1806) 363; Mia. Fl. Ind. Bat. 3 (1856) 286; Boeck. Linnaea 35 (1868) 597; CLARKE, J. Linn. Soc. Bot. 21 (1884) 143, excl. specim. madag.; F1. Br. Ind. 6 (1893) 607; VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 126, t. 4 f. 30a, b, excl. specim. discrep., non f. 30c; Koorp. Exk. Fl. Java 1 (1911) 190; ibid. 4 (1922) f. 221; KUx. Pfi. R. Heft 101 (1935) 144, f. 5 A-D; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 38; KERN, Reinwardtia 3 (1954) 31; Koyama, Quart. J. Taiwan Mus. 14 (1961) 168; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 481.— C. calopterus Mig. FI. Ind. Bat. 3 (1856) 282. — C. eleusinoides (non KUNTH) K.SCH. & LAuT. Fl. Schutzgeb. (1900) 191; Merr. En. Philip. 1 (1923) 105, p.p. var. nutans. — Synonymy as above. Perennial with short rhizome. Stems tall, tufted, trigonous, triquetrous just below the inflorescence, smooth, up to 100(—150) by 1 cm. Leaves coriaceous, flat, the larger ones somewhat plicate with the midrib prominent beneath and 2 lateral nerves marked above, scabrous on the margins in the upper part, dark green or glaucous above, whitish striate beneath, 6—15 mm wide. Inflorescence compound or decompound, usually large. Involucral bracts 4—6, the larger ones far overtopping the inflorescence, up to 75 cm. Primary rays 6—10, very unequal, obliquely patent, rigid, smooth, up to 30 cm, the larger ones bearing 5—7 up to 6 cm long secondary rays. Spikes narrow, more or less penicillate, 3-4 cm by 5—10 mm, loose, with glabrous rachis and 15-25 spikelets. Spikelets spicately arranged, suberect, oblong-linear, distinctly compressed, 8—14-flowered, 6-14 by c. 2 mm; rachilla slightly flexuous, winged, persistent; internodes c. %mm; wings lanceolate, whitish hyaline, persistent, c. 14 mm wide. Glumes membra- nous, obliquely erect to patulous, keeled, oblong, obtuse, broadly whitish hyaline-margined in the upper half, soon incised at the tip and mucronulate by the excurrent midrib, 7-nerved, YA imbricate, 2-2/4 by c. 1 mm; keel greyish green, sides pale fuscous. Stamens 3; anthers oblong-linear, 4-1 mm, with distinctly produced, smooth appendage of the connective. Stigmas 3. Nut triquetrous, oblong to oblong-ovoid, apiculate, brown, c. 1/4 by 4 mm. Distr. Tropical West Africa (according to CLARKE and KUKENTHAL, not in HUTCHINSON & DALZIEL, Fl. W. Trop. Afr. 2, 1, 1931); from Ceylon and India to S. China, Farther India, and Malesia: in a few localities in Sumatra, W. and E. Java, Lesser Sunda Is. (Sumbawa, Flores, Alor), the Philippines (Luzon, Mindanao), Celebes, and New Guinea. Ecol. Swamps, wet rice-fields, moist places in 610 forests, margins of pools, river-banks, at low altitudes (up to 800 m, in New Guinea, Western Highlands, collected at c. 1500 m). Vern. Pérémpungan, M, N. Sum., djukut leuleutan, S, mau-mau, Alor; Philip.: gauang, salinganga, Bon. Note. For the differences with C. distans var. pseudonutans see there. The species is closely allied to C. distans, but in general appearance and several characters it shows affinity to the species of sect. Exaltati, especially to C. elatus, from which it is distinguishable by the loose spikes, the broader spikelets, the twice as long internodes of the rachilla, the smooth appendage of the connective, and the larger nuts. var. eleusinoides (KUNTH) HAINES, Bot. Bihar Orissa 5 (1924) 898; Koyama, Quart. J. Taiwan Mus. 14 (1961) 168, in nota. — C. eleusinoides KUNTH, En. 2 (1837) 39; Mia. Fl. Ind. Bat. 3 (1856) 270; BoEcK. Linnaea 35 (1868) 596; BENTH. FI. Austr. 7 (1878) 277; CLARKE, J. Linn. Soc. Bot. 21 (1884) 142; FI. Br. Ind. 6 (1893) 608; VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 128, t. 4 f. 31; CLARKE, Philip. J. Sc. 2 (1907) Bot. 84; Camus, Fl. Gén. I.-C. 7 (1912) 60, f. 6, 7-11; Merr. En. Philip. 1 (1923) 105, p.p.; KUk. Pfl. R. Heft 101 (1935) 144; Kern, Reinwardtia 2 (1952) 111. — C. xanthopus STEUD. Flora 25 (1842) 595; Syn. 2 (1855) 36, excl. pl. jav. Differs from var. nutans by: Inflorescence less compound. Primary rays suberect, the secondary ones often beset with spikelets almost to the base. Spikes shorter, denser. Spikelets usually fewer- flowered. Glumes less remote because of the slightly shorter, c. 14 mm long internodes of the rachilla, more distinctly mucronate (mucro 14—)4 mm). Nut more obovoid, 1.1—1.25 by 0.6—0.7 mm. Distr. Tropical Africa, S. and SE. Asia, Formosa, Ryu Kyu Is., tropical Australia, in Malesia very rare: Lesser Sunda Is. (Flores, Timor), Philippines (Luzon), New Guinea (Morobe Distr.). Ecol. In swamps, sometimes abundant, at low altitudes (in Timor at 750 m). Note. The African specimens I have seen look rather different from var. nutans, but the Asian often approach it very much, and no sharp line between the two varieties can be drawn. The collection from West Java determined by CLARKE as C. eleusinoides, and the greater part of the collections cited under this name by MERRILL, /.c., belong to var. nutans. The Philippine collection cited by KUKENTHAL, I.c., under var. subprolixus KUx. | take for a depauperate var. nutans. 16. Cyperus distans LINNE f. Suppl. (1781) 103; Boeck. Linnaea 35 (1868) 612; BENTH. FI. Austr. 7 (1878) 277; CLARKE, J. Linn. Soc. Bot. 21 (1884) 144; Fl. Br. Ind. 6 (1893) 607; VALcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 123, t. 1 f. 9, t. 4 f. 29; CLARKE, Philip. J. Sc. 2 (1907) Bot. 84; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 69; Koorp. Exk. Fl. Java 1 (1911) 191; ibid. 4 (1922) f. 222; Camus, FI. Gén. I.-C. 7 (1912) 60; MerR. En. Philip. 1 (1923) 105; Rint. FI. Mal. Pen. 5 (1925) 145; Back. Onkr. Suiker. (1928) 137, t. 134; KUx. Pfl. R. Heft 101 (1935) 137, incl. f. pachyanthos KUx.; S. T. BLake, J. Arn. Arb. 28 (1947) 214; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 35; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 481. — C. elatus (non L.) Rotts. Descr. & Ic. (1773) 37, t. 10; ZoLL. Syst. Verz. 1 (1854) 63; FLORA MALESIANA [ser. I, vol. 7° STEUD. Syn. 2 (1855) 49; Mia. Fl. Ind. Bat. 3 (1856) 284, incl. var. graminicola Miq.; Sum. (1861) 260, 600; Boeck. Flora 62 (1879) 551.— C. graminicola STEuD. [in Zoll. Syst. Verz. 1 (1854) 63, nom. nud.] Syn. 2 (1855) 49.— C. kurrii StEuD. Syn. 2 (1855) 38; Mia. Fl. Ind. Bat. 3 (1856) 279; NAves, Nov. App. (1882) 305.—C. digitatus (non RoxsB.) KUK. Bot. Jahrb. 69 (1938) 255, non al. Perennial with shortly creeping rhizome. Stems subcespitose, slender to rather stout, triquetrous, smooth, 10—50(—100) cm by 2—3(—5) mm, leafy in the lower 14-14. Leaves shorter than or as long as the stems, rigid to flaccid, flat, gradually acuminate, scabrous in the upper part, 5—8(—10)mm wide; lower sheaths purplish. Inflorescence compound or decompound, often large, usually open. Involucral bracts 4—6(—8), spreading, the lower ones overtopping the inflorescence, 20—40(—60) cm. Primary rays 6—12, obliquely patent, very unequal, rigid, smooth, up to 20 cm; secondary ones up to 8 cm. Spikes broadly ovoid-pyramidal, with slightly flexuous, glabrous rachis and c. 15 (5—20) spikelets. Spikelets spicately arranged, ultimately spreading at right angles, the lower ones often somewhat reflexed (see var.), narrowly linear, almost terete, 6—10(—30)-flowered, 1—2(—3) cm by c. 1 mm; rachilla flexuous, distinctly winged, persistent; internodes 1(-1/4) mm, wings lanceolate, whitish hyaline, caducous, c. 4mm wide. Glumes membranous, appressed, oblong-ovate or elliptic, hardly keeled, obtuse, remote (not imbri- cate), (134-)2(-244) mm by 1-114 mm; keel green, 3—5-nerved; sides nerveless, reddish brown, rarely stramineous; margins broad, whitish hyaline. Stamens 3; anthers oblong, 4-34 mm, with hardly produced appendage of the connective. Stigmas 3. Nut trigo- nous, oblong-cylindrical, broadly stipitate, apiculate, blackish brown under the greyish, detergible outer layer of cells, 114-1% by 4-4 mm. Distr. Widely spread over the warmer regions of the whole world; throughout Malesia and presumably everywhere common. Ecol. In wet or rather wet places: swamps, river-banks, grassy road-sides, wet rice-fields and meadows, open places in secondary forests; 0-1800 m. Vern. Pajungan, rumput pérémpungan, téki rawah, M, djukut djampang, ilat, lilik sunge, manjératan, §S, lingi, méndongan, wéligi, J, dlingi, M, rumput rusa, tali juru, Brunei, bobarai, Halmaheira, sinasoan, Talaud; Philip.: amiusan bakabakahan, muthang- kalabdu, Tag., barisanga, U1k., burabongdai, S.L.Bis., gagauan, Bon., hanganga, If., langalang, Bik., pulok- galo, talubak, Buk. Notes. Very variable in all its parts. The forms and varieties described have no or little taxonomical value, probably except for: var. pseudonutans KUxK. Pfl. R. Heft 101 (1935) 140. — Spikelets suberect, even in fruit, the spikes therefore more or less penicillate. — Ceylon, India, Formosa, Ryu Kyu Is., according to KUKENTHAL also in Africa; in Malesia: Singapore (KUKENTHAL, l.c., not seen), Central Java (Djapara, forest complex Ngarengan). — Strongly resembling C. nutans and confused with it by CLARKE; distinguishable by the remote, not imbricate glumes with nerveless sides and the long internodes of the rachilla. For C. distans x C. rotundus see p. 660. 1974] Cy PERACEAE (Kern) 611 7. Section Proceri KUNTH, En. 2 (1837) 72. — Cyperus sect. Longispicati b. Pilosi VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 128. — Cyperus sect. Latifolii CLARKE, Kew Bull. add. ser. 8 (1908) 99. Type species: C. procerus ROTTB. 17. Cyperus procerus Rotts. Descr. & Ic. (1773) 29, t. 5 f. 3; Mia. Fl. Ind. Bat. 3 (1856) 278; CLARKE, J. Linn. Soc. Bot. 21 (1884) 152; Fl. Br. Ind. 6 (1893) 610; VALCK. Sur. Ges]. Cyp. Mal. Arch. (1898) 131, t. 4 f. 34; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 70; Koorp. Exk. FI. Java 1 (1911) 191; ibid. 4 (1922) f. 223; Camus, Fl. Gén. I.-C. 7 (1912) 62; Rip. FI. Mal. Pen. 5 (1925) 147; Back. Onkr. Suiker. (1928) 138, t. 136; KUx. Pfl. R. Heft 101 (1935) 91; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 36; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 478. — C. ornatus R.BR. Prod. (1810) 217; BENTH. Fl. Austr. 7 (1878) 276; Naves, Nov. App. (1882) 303.—C. heynii BoEcK. Linnaea 35 (1868) 600; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 48. — Pycreus puncticulatus (non NEES) RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 61: Fl. Mal. Pen. 5 (1925) 140. — Pycreus baccha (non Nees) Camus, FI. Gén. I.-C. 7 (1912) 36. — Duval- jouvea procera H. PreirF. Mitt. Inst. Bot. Hamb. 7 (1928) 167. — C. puncticulatus (non VAHL) KUK. Pfl. R. Heft 101 (1936) 362, quoad specim. malacc.— Fig. 51. Perennial with stoloniferous rhizome. Stems stout, triquetrous, smooth, 70—125(—175) cm by 4—7(-—10) mm, the base clothed with spongious, greyish brown to purplish sheaths. Leaves canaliculate, spongious or coriaceous, firm, gradually acuminate, scabrid at the top, up to 9(—15) mm wide. Inflorescence simple or subcompound, loose, relatively small, 10-15 cm long. Involucral bracts 3-4, obliquely erect to spreading, the larger one(s) far overtopping the inflorescence, up to 50(—70) cm long. Primary rays 3—7, very unequal, more or less spreading, smooth, the longest 6—12(—20)cm; secondary rays when present very short. Spikes broadly ovoid, loose, 2—4 cm long and wide; rachis angular, (in Malesian specimens) more or less scabrous-hispid. Spikelets up to 10(—18) to the terminal spike, only 3—4 to the lateral ones, spicately arranged, patent, the lower ones at right angles to the rachis, oblong to broadly linear, often slightly curved, compressed, up to 40-flowered, 10—35 by 2/4-3!4 mm; rachilla straight, not or hardly winged, persistent; internodes c. 4 mm. Glumes membranous, patulous, ovate or elliptic, obtuse, hardly keeled, c. 7-nerved, reddish brown with broad hyaline margins in the upper part, 4 imbricate, 2/4-3 by c. 2 mm. Stamens 3; anthers linear, (1—)1/4—2 mm long, with very short, smooth appendage of the connective. Stigmas 3. Nut tri- quetrous, obovoid or ellipsoid, minutely apiculate, blackish brown, c. 114 by % mm. Distr. From Ceylon and India to Cochinchina, E. China, and Formosa, southward to Queensland, in Malesia rather rare: Malay Peninsula, Java and adjacent islands, N. Borneo, Philippines (Luzon). According to KUKENTHAL varieties in tropical Africa and Madagascar; see, however, CHERMEZON in Fl. Madag. fam. 29 (1937) 115 & 117. Ecol. In open, moist or wet, often brackish localities: swamps, pools, wet rice-fields, often near the sea, 0-50 m, a few times collected near Bogor (W. Java) at c. 250 m. Use. The tough stems split in three are sometimes used as strings or binder twines. Notes. In typical C. procerus the rachis of the spikes is smooth, while in all Malesian specimens it is more or less scabrous-hispid. The Malesian plants may therefore be referred to: var lasiorrhachis CLARKE, FI. Br. Ind. 6 (1893) 610: KOK. Pfl. R. Heft 101 (1935) 91.—C. procerus f. lasiorrhachis VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 131. — Axes of the spikes scabrous-pilose. — In the type of this variety (from Chota Nagpore, CLARKE 33818, K) the rachis is more densely hispid than in the Malesian plants. Erroneously the variety was described as having “axis of spikelets scabrous pilose’, which error was copied by KUKENTHAL. This variety is sometimes difficult to distinguish from stout specimens of C. pilosus. The inflorescence is less compound and more open than in that species, the spikelets are larger and more distant, the anthers longer, and the nuts larger. KUKENTHAL, l.c., mentions from Singapore (MAYER 328 p.p.) var. griffithianus (BOECK.) KUk. — C. griffithianus BorEcK. Linnaea 35 (1868) 601, according to him with compound inflorescence and oblong-elliptic glumes. I have not seen the type of C. griffithianus (GRIFFITH s.n. in herb. Boeck.) from India, nor MAYER’s Singapore collection. CLARKE reduced BOECKELER’s species to C. pilosus, but from the original description I might rather refer it to C. procerus var. lasiorrhachis. KAMPHOVENER 786 (C), also cited by KUKENTHAL, I take for typical C. procerus on account of the simple inflorescence and the smooth rachis. 18. Cyperus pilosus VAHL, En. 2 (1806) 354; BoEcK. Linnaea 35 (1868) 598, incl. var. muticus BOECK.; BENTH. FI. Austr. 7 (1878) 275; CLARKE, J. Linn. Soc. Bot. 21 (1884) 148, incl. var. obliquus CLARKE et var. polyanthus CLARKE; O.K. Rev. Gen. Pl. (1891) 750, incl. f. pallidus O.K. et f. badius O.K.; CLARKE, Fl. Br. Ind. 6 (1893) 609; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 128, t. 4 f. 32; CLARKE, Philip. J. Sc. 2 (1907) Bot. 84; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 67; Koorp. Exk. Fl. Java 1 (1911) 191; ibid. 4 (1922) f. 224; Camus, Fl. Gén. I.-C. 7 (1912) 62, f. 6, 12; Merr. En. Philip. 1 (1923) 107; Rupv. Fl. Mal. Pen. 5 (1925) 144; Back. Onkr. Suiker. (1928) 138, t. 135; KUx. Pfl. R. Heft 101 (1935) 92, incl. var. contractus KUK.; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 34; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 477.— C. obliquus NEEs in Wight, Contr. (1834) 86; Boeck. Linnaea 35 (1868) 611; Naves, Nov. App. (1882) 305.—C. marginellus NEES in Wight, /.c. 83; in Hook. J. Bot. Kew Misc. 6 (1854) 28. — C. venustus (non R.BR.) Mor. Syst. Verz. (1846) 96.— C. piptolepis Steup. [in Zoll. Syst. Verz. 1 (1854) 63, nom. nud.] Syn. 2 (1855) 40; 612 FLORA MALESIANA [ ser: I, volyg% Fig. 51. Cyperus procerus Rotts. a. Habit, x 14, b. rachis of spike, c. spikelet, both x 5, d. persistent rachilla hardly winged, e. glumes, f. deflorate flower, g. anther, h. nut, all x 10 (a HEYNE s.n., b—h DANSER 6470). 1974] Mig. FI. Ind. Bat. 3 (1856) 279, incl. var. f, var. contractus MiQ. et var. pyrophilus Mig.; Miq. in De Vriese, Pl. Ind. Bat. Or. (1856) 139. — C. pennatus (non LAMK) ZOLL. Syst. Verz. 1 (1854) 62.—C. pauciflorus STEUD. Syn. 2 (1855) 34; Mig. Fl. Ind. Bat. 3 (1856) 275: Naves, Nov. App. (1882) 304. — C. hebes StEUD. Syn. 2 (1855) 315.— C. pyrophilus REINW. ex Mig. in De Vriese, Pl. Ind. Bat. Or. (1856) 139. — C. nutans (non VAHL), specim. discrep. VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 127, t. 4 f. 30 C.— Duvaljouvea pilosa PALLA in Koch, Syn. ed. 3, 3 (1905) 2555; Allg. Bot. Zeitschr. 17 (1912) Beil. 8. — Fig. 52. Perennial, the rhizome emitting slender stolons clothed with lanceolate scales. Stems stoutish, solitary or subcespitose, triquetrous, smooth or scaberulous just below the inflorescence, 10—50(—110) cm by up to 6 mm. Leaves shorter than to as long as the stems, canaliculate or somewhat plicate with 3 prominent nerves, weak to rather rigid, gradually acuminate, scabrous on margins and nerves in the upper part, 4—10(—13) mm wide; lower sheaths brown-purplish. Inflorescence compound, very variable in size and density. Involucral bracts (3—)4—5, obliquely erect to spreading, the larger ones far overtopping the inflorescence, up to 50cm. Primary rays (2—)6— 7(-10), very unequal, spreading, smooth, the larger ones 5—15(—20) cm with 3—6 short secondary rays. Spikes ovoid, 2—3 cm long, with angular, rather to very densely hispidulous or pubescent rachis. Spike- lets 6—25 to the spike, at right angles to the rachis or often reflexed, very rarely remaining suberect, elliptic to linear-lanceolate, compressed, 7—20(—50)- flowered, 5—10(-20) by 2-2)4(-3) mm; rachilla straight, wingless or almost so, persistent; internodes %-Y%,mm. Glumes membranous, patulous, ovate, obtuse, sometimes minutely mucronulate, hardly keeled, 5—7-nerved, 14-74 imbricate, with green midrib, stramineous to reddish brown sides, and broad, whitish hyaline margins in the upper part, 2-2)% mm by ec. 1/4 mm. Stamens 3; anthers oblong, ’%4-% mm long, with short, smooth, reddish ap- pendage of the connective. Stigmas 3. Nut triquetrous, obovoid or ellipsoid, broadly stipitate, apiculate, blackish brown, 1-1/4 by 4-24 mm. Distr. From Central Asia and Japan through the whole of tropical Asia and tropical Australia, very rare in tropical W. Africa; throughout Malesia, common and often abundant. Ecol. In open wet places; swamps, rice-fields, grasslands, river-banks, etc.; 0—1500 m. Vern. Rumput djéking, M, hilut, ilat, ilat walini, S, lumbungan sapi, J, rija-rija, rumput randjang, siban- gélulu, Sum. E. C., bola-bola tapongan, Batak, rombok, siangit padang, Sum. W. C., para-para, Riouw, subah péra, Siberut, madarong darat, Mal. Pen., mauflés, Timor, bundung, tali juru, Borneo: Dusun; Philip.: paragi, Sub.; New Guinea: buatak, Andjai, akarebok, Kapauku. Notes. For the differences with the very closely related C. procerus, see under that species. C. pilosus is very variable in size, colour of the glumes, number of the flowers in the spikelets, etc., but the species, varieties, and forms based on these differences have no taxonomic value and were already merged into the species by VALCKENIER SURINGAR. CLARKE mentions specimens with gla- brous or almost glabrous rachis of the spikes; they Cy PERACEAE (Kern) 613 have not yet been collected in Malesia. Some speci- mens from W. Java and the Kangean Archipelago have persistently erect spikelets and therefore peni- cillate spikes. They may represent a distinct variety analogous to C. distans var. pseudonutans. 19. Cyperus babakan STEuD. Syn. 2 (1855) 6; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 34: KERN, Reinwardtia 2 (1952) 105, f. 4; ibid. 3 (1954) 66; ibid. 6 (1961) 54; in Back. & Bakh. f. Fl. Java 3 (1968) 477.— C. babakensis StEuD. [in Zoll. Syst. Verz. | (1854) 62, nom. nud.| ex Mia. FI. Ind. Bat. 3 (1856) 257; Boeck. Linnaea 35 (1868) 521; CLARKE, FI. Br. Ind. 6 (1893) 610; Ill. Cyp. (1909) t. 15 f. 1-3; Camus, FI. Gén. I.-C. 7 (1912) 61; RIDL. Fl. Mal. Pen. 5 (1925) 144; Ktx. Pfl. R. Heft 101 (1935) 94.— C. pilosus VAHL var. babakensis CLARKE, J. Linn. Soc. Bot. 21 (1884) 151.—C. benghalensis CLARKE, l.c.— C. pilosus VAHL f. babakensis VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 129, t. 4 f. 33. — Duval- jouvea babakensis H. Preirr. Mitt. Inst. Bot. Hamb. 7 (1928) 167. Perennial, the rhizome emitting slender stolons clothed with broadly ovate-lanceolate, brownish scales. Stems stoutish, solitary, triquetrous with somewhat concave sides, smooth, up to 90 cm by 5-7 mm. Leaves few, shorter than to as long as the stems, flat or canaliculate, gradually acuminate, scabrous on the margins in the upper part, 5—10 mm wide; basal sheaths often bladeless, somewhat spongy. Inflorescence simple, relatively small. Involucral bracts 3—4, patent to reflexed, the lower 2 very long, much overtopping the inflorescence. Rays 2-5, unequal, erect or suberect, firm, short, rarely up to 5(—9) cm. Spikes ovoid or oblong-ovoid, very dense, up to 5 by 4cm; rachis hispidulous. Spikelets spicately arranged, patent to reflexed, oblong to linear, compressed, 10—20(—40)-flowered, 6—20 by 24-3 mm; rachilla straight, wingless or nearly so, persistent; internodes 14-34 mm. Glumes membra- nous, obliquely erect, boat-shaped, ovate, subacute, minutely mucronulate, 7—9-nerved, stramineous to reddish brown, 74 imbricate, 24-3 by 2 mm, the keel antrorsely hispid-scabrous at least towards the top; margins hyaline. Stamens 3; anthers oblong- linear, ¥%-1 mm long, with short, smooth, reddish appendage of the connective. Stigmas 3. Nut trique- trous, broadly ellipsoid or obovoid, apiculate, broadly stipitate, black, 1/4-1)4 by 0.8-0.9 mm. Distr. From Hindustan and Assam to Cochin- china and Malesia: Malay Peninsula (Wellesley, Perak, Kelantan, Pahang, Selangor, Malacca, P. Penang), W. Java (Babakan, Rangkasbetung), S. Borneo, the Philippines (Panay, Iloilo Prov.), Celebes (Lasoa), and W. New Guinea (Kurik near Merauke). Apparently a rare species. Ecol. In swamps and wet rice-fields, in light forests, at low altitude, up to 100 m. Note. STEUDEL, followed by MIQUEL, wrongly placed this well characterised species in sect. resp. subg. Pycreus. 20. Cyperus malaccensis LAMK, Ill. 1 (1791) 146: Mia. Fl. Ind. Bat. 3 (1856) 297; Boeck. Linnaea 35 (1868) 603; CLARKE, J. Linn. Soc. Bot. 21 (1884) 147; Fl. Br. Ind. 6 (1893) 608; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 121, t. 4 f. 22; Rip. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 70; CLARKE, Philip. J. Sc. 614 FLORA MALESIANA { ser. I; volar LB CE Ki Shue THU Pd) Nt Fig. 52. Cyperus pilosus VAHL. a. Habit, x }4, b. part of spikelet, x 5, c. glumes, d. rachilla, e. pistil, f. anther, also enlarged, g. nuts, all x 10 (a—g BACKER 27746). 1974] Cy PERACEAE (Kern) 615 Fig. 53. Cyperus malaccensis LAMK. The long, serpentine string-brushes consist of intertwined stem parts in rope of this species. They float and are set out perpendicular to the beach and serve for attracting young fish (bibit or bandeng) which are then bred in artificial brackwater fish ponds (tambaks) behind the beach. Pemalang, northcoast of Central Java; beach wall grown with Pandanus odoratissimus (photogr. VAN STEENIS). 2 (1907) Bot. 84; Koorp. Exk. Fl. Java 1 (1911) 190; ibid. 4 (1922) f. 220; Camus, Fl. Gén. I.-C. 7 (1912) 63; BRowNn, Min. Prod. Philip. For. 1 (1920) 346, t. 13, 14; MerR. En. Philip. 1 (1923) 106; RipL. FI. Mal. Pen. 5 (1925) 147; Back. Onkr. Suiker. (1928) 18671..1355> KK. Pil. R. Heft 101 (1935) 86S. 0. BLAKE, J. Arn. Arb. 28 (1947) 214; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 35; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 478. — C. spanio- phyllus Steup. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 21; Mig. Fl. Ind. Bat. 3 (1856) 267. — C. corymbosus (non Rotts.) Mia. l.c. 272. — C. tegetiformis (non RoxB.) BENTH. FI. Austr. 7 (1878) 278.— C. difformis (non L.) BLANCO, FI. Filip. ed. 3, 4 (1886) 302.— C. tegetum (non Roxs.) RIDL. J. Str. Br. R. As. Soc. n. 46 (1906) 222: MeRR. En. Born. (1921) 56. — Chlorocyperus malaccensis PALLA, Allg. Bot. Zeitschr. 17 (1912) Beil. 6 — Fig. 53. Perennial with stout stolons clothed with ovate, scarious, dark brown scales hardening into a woody rhizome. Stems approximate, robust, spongious, triquetrous with concave sides, almost 3-winged above, smooth, 60-175 cm by up to 12(—15) mm. Lower leaves on the flowering stems reduced to spongious, bladeless greyish black to purplish sheaths, the upper ones with very long (up to 20 cm) sheaths, often laminate but the blades always very short, the uppermost reaching up to halfway the stem; blades of the sterile shoots abruptly acuminate, scabrid at the top, 5—10(—18) mm wide. Inflorescence 616 compound or subdecompound, usually broader than long, sometimes congested, up to 10 by 15cm. Involucral bracts 3—4, flat, abruptly acuminate, scabrid at the top, much overtopping the inflo- rescence, shining green above, greyish beneath, the lowest usually erect, up to 30 cm by 8—15 mm, the others patent to reflexed. Primary rays 3—6(—10), very unequal, spreading, slender, smooth, 3—10 cm, secondary ones setaceous, c. 2cm. Spikes broadly ovoid, with glabrous rachis. Spikelets spicately arranged, 6-12 to the spike, linear, often somewhat curved, subterete, 1-3cm by 1 ya 14cm, 16- 20(—40)-flowered; rachilla straight, very narrowly winged, persistent: win S persistent, whitish or yellowish; internodes *%—%,mm. Glumes charta- ceous, ovate to elliptic. obtuse. muticous, not keeled (rounded on the back), indistinctly 5—7-nerved, c. \% imbricate, pale fuscous with yellowish margins, when dry crispidly incurved all round, 2-2/4 by 1-114 mm. Stamens 3; anthers linear, >4-1 mm. Stigmas 3. Nut trigonous, narrowly oblong, slightly compressed dorsally, hardly apiculate, dark brown to black, 134-2 by }4 mm. Distr. Widely spread in the eastern hemisphere, from Mesopotamia through India to S. China, Northern Australia, and Polynesia; throughout Malesia, in Java not rare along the muddy north coast, on the south coast only in Besuki (E.). Ecol. In moist localities, usually within the in- FLORA MALESIANA [ser. 1, vol fluence of salt or brackish water, a coloniser along muddy river-mouths, on mud flats, sandy foreshores covered by springtides; often abundant. Uses. The stems are often used for tying purposes and for making mats, baskets, and hats; the manu- facture of slippers of this material is carried on to a considerable extent in some towns of Bulacan Prov., Luzon (see Brown, I.c.). In Pekalongan (Central Java) the stems are used by the fishermen; pieces of the stems are plaited in ropes which are brought into the sea. The fry, attracted by the stems, is caught and then planted in fish-ponds. See VAN STEENIS, Trop. Natuur 29 (1940) 20, fig. Vern. Bundjung, wlingi laut, M, daréngdéng, S, kédot, sukét dem, J, kumbu, Sum., sélimbu, Johore, pea-pea, rumput kuluwing, Celebes, geida, Papua; Philip.: bagd-as, P.Bis., balangot, balongat, Tag., Bik., Bis., Pamp., barangot, Bik., talaid, Bag., tikog, Mbo. Note. C. malaccensis var. brevifolius BOECK. Linnaea 35 (1868) 604; KUx. Pfl. R. Heft 101 (1935) 87; KoyAMA, Quart. J. Taiwan Mus. 14 (1961) 171.— C.monophyllus VAHL, En. 2 (1806) 352; OHW1, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 136. BOECKELER wrongly cited C. spaniophyllus STEUD. (type ZOLLINGER 1209 from Java) as a synonym of this distinct E. Asiatic taxon, and therefore mentioned it for that island. It does not extend to Malesia. 8. Section Iriae KUNTH, En. 2 (1837) 38. iype species: C. ria L. 21. Cyperus iria LINNE, Sp. Pl. (1753) 45; Hassk. Pl. Jav. Rar. (1848) 83; Mia. Fl. Ind. Bat. 3 (1856) 269, incl. var. parviflorus MiqQ. et var. diaphaniria Miq.; Boeck. Linnaea 35 (1868) 595; BENTH. FI. Austr. 7 (1878) 276: CLARKE, J. Linn. Soc. Bot. 21 (1884) 137; Fl. Br. Ind. 6 (1893) 606; VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 105, t. 4 f. 16; CLARKE, Philip. J. Sc. 2 (1907) Bot. 83; RipLt. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 66; CLARKE, Ill. Cyp. (1909) t. 4 f. 2, non f. 1; Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 215; Camus, Fl. Gén. I.-C. 7 (1912) 59, f. 6, 4—6; Merr. En. Philip. 1 (1923) 106; Rip. Fl. Mal. Pen. 5 (1925) 143: Back. Onkr. Suiker. (1928) 135, t. 131; KUxK. Pfl. R. Heft 101 (1935) 150; S. T. BLAKE, J. Arn. Arb. 28 (1947) 214; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 34; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 479. — C. parviflorus (non VAHL) NEES in Wight, Contr. (1834) 87.—C. seminudus ZOLL. & Mor. in Mor. Syst. Verz. (1846) 96, non Roxs. 1820.— C. diaphaniria SteuD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 23.— C. nuttalii (non Torr.) LLANOs, Fragm. PI. Filip. (1851) 14; F.-VitL. & NAves in Blanco, FI. Filip. ed. 3, 4 (1880) 9.— Chlorocyperus iria Rik, Jahrb. Wiss. Bot. 27 (1895) 564; PALLA, Allg. Bot. Zeitschr. 17 (1912) Beil. 6. Annual, or perennial in favourable circumstances; roots yellowish red. Stems tufted, slender to rather stout, triquetrous, smooth, (5—)15—50(—80) cm by (1-)2-3(-5) mm. Leaves basal, shorter than to somewhat longer than the stems, flat or channelled, rather flaccid, gradually acuminate, scabrous on the margins in the upper parts, 3—6(—8)mm _ wide. Inflorescence greatly varying in size, simple or compound, usually loose, up to 20cm long. In- volucral bracts 3—S(—7), obliquely erect to patent, the larger 1-3 overtopping the inflorescence, up to 40 cm long. Primary rays 3—5(—8), very unequal, obliquely patent, slender, smooth, up to 10(—18) cm, the larger ones usually umbellately or paniculately branched, the secondary rays very short. Spikes narrow, oblong-ovoid, often elongate, rather dense or loose, with 5-25 spikelets; rachis flexuous, glabrous. Spikelets spicately arranged, oblong-linear, strongly compressed, obtuse, (in the Malesian speci- mens) erecto-patent, 6—20(—24)-flowered, 3—10(—13) by 1’4-2 mm; rachilla straight, wingless, persistent; internodes c. A mm. Glumes membranous, spreading- ascendent, orbicular or broadly ovate, not rarely broader than long, sharply keeled, rounded to emarginate at apex, very shortly mucronulate with green, arched, 3—5-nerved keel, cellular-reticulate, nerveless, golden to fulvous sides, and broadly whitish hyaline margins towards the top, 4-% imbricate, 14-11% by 1-1/4 mm. Stamens 2-3; anthers very small. oblong, c. 14 mm, with short, smooth, reddish appendage of the connective. Style hardly any; stigmas 3, 4—/4 mm long. Nut about as long as the subtending glume, triquetrous with slightly concave sides, obovoid-ellipsoid, broadly stipitate, hardly apiculate, shining dark brown to black, 1-114 by 4-3 mm. Distr. Widely spread in tropical Asia, extending northwards to Iran, Afghanistan, China, and Japan, 1974] Cy PERACEAE (Kern) 617 southwards to Australia, and westwards to tropical E. Africa; introduced and naturalised in the SE. United States and the West Indies; common through- out Malesia. Ecol. In open wet places, on road-sides, river- banks, and especially a characteristic weed in wet rice-fields; at low altitudes up to 700 m, rarely to 1200 m. Vern. Babawangan, S, dékéng wangin, njur-njuran, rumput djékéng kungit, umbung, J, djung padjungan, Md, blumbungan, papungan, rumput silupak, Sum. E. C., bua bua tapongan, Batak, kumis, Kangean; Philip.: alinang, paiung-paiung, sud-sud, taga-taga, Bik., okokiang, Bon. Notes. Very variable in size and the number of flowers in the spikelets. The extremes are connected by numerous intermediates and do not deserve nomenclatural recognition. LOHER 7159 from the Philippines represents f. chrysomelinus (LINK) KUK. Pfil. R. Heft 101 (1935) 151 [C. chrysomelinus Link, Hort. Berol. 1 (1827) 305], with setaceous stems and leaves, and the inflorescence reduced to 1—2 spikelets. It is probably a depauperated form, also of little or no taxonomical value. C. microiria STEUD., wrongly referred to C. iria var. parviflorus MiqQ., is a well-characterised, E. Asian species not extending to Malesia. 9. Section Compressi KUNTH, En. 2 (1837) 23. Type species: C. compressus L. 22. Cyperus compressus LINNE, Sp. Pl. (1753) 46: Hassk. Pl. Jav. Rar. (1848) 77; Miq. Fl. I d. Bat. 3 (1856) 263, incl. var. meyenii MiQ.; BoEcK. Linnaea 35 (1868) 517. incl. var. brachiatus BOECK.; CLARKE, J. Linn. Soc. Bot. 21 (1884) 97, incl. var. pectiniformis CriaRKE: FI. Br. Ind. 6 (1893) 605; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 82, t. 4 f. 6; CLARKE, Philip. J. Sc. 2 (1907) Bot. 84; RipL_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 67; Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 208; Camus, FI. Gén. I.-C. 7 (1912) 57, f. 5, 3-5; Merr. En. Philip. 1 (1923) 103; RIDL. Fl. Mal. Pen. 5 (1925) 144; Back. Onkr. Suiker. (1928) 132, t. 126; KUx. Pfl. R. Heft 101 (1935) 156, f. 4 A-D; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 32; KERN in Back & Bakh. f. Fl. Java 3 (1968) 480. — C. brachiatus Por. Enc.7 (1806) 259. — C. pectiniformis R. & S. Mant. 2 (1824) 128; Nees in Wight, Contr. (1834) 77. — C. meyenii Nees, Nov. Act. Ac. Caes. Leop.-Car. 19, Suppl. 1 (1843) 57. — C. humilis (non KUNTH) LLANOos, Fragm. PI. Filip. (1851) 13; F.-Vit_. & Naves in Blanco, FI. Filip. ed. 3, 4 (1880) 7. — Chlorocyperus compressus PALLA, Denkschr. K. Ak. Wiss. M.-N. KI. Wien 84 (1909) 451. Annual. Stems tufted, triquetrous, smooth, very variable in height. up to 50 cm by 1-2 mm. Leaves shorter to somewhat longer than the stems, flat or channelled, rigid. gradually acuminate, scabrous at the top, (14-)2—-4 mm wide; lower sheaths reddish brown. Inflorescence simple, open, not rarely reduced to a single sessile cluster. Involucral bracts 3-5, patent, the lower ones overtopping the inflorescence, up to 30cm. Rays 0-8, obliquely patent to widely spreading. slender. smooth. up to 15cm. Spikes broadly ovoid, with 3—7(—10) spikelets. Spikelets subdigitately arranged on the very short, up to 55 mm long rachis, obliquely patent to horizontally spreading or slightly reflexed, oblong to linear, acute, rhom- boidal in cross-section, strongly compressed, up to 40-flowered, 1-3 cm by 3—5 mm; rachilla flexuous. persistent, initially winged; wings very thinly mem- branous, caducous; internodes c. 34 mm. Glumes chartaceous, rigid, ovate, acute, strongly keeled. conspicuously multinerved, more than /% imbricate. with green keel, pale green to yellowish brown sides, and silvery-hyaline margins, 3-4 by 2-234 mm: mucro strong, often slightly excurved, up to 1 mm. Stamens 3; anthers small, oblong-linear, 74-1 mm, with scarcely produced, smooth appendage of the connective. Stigmas 3. Nut trigonous with incrassate angles, broadly obovoid, broadly stipitate, obtuse, shortly apiculate, shining dark brown to blackish. VA-17% by c. 1 mm. Distr. Pantropical; throughout Malesia. Ecol. A common weed of open grasslands. waste places, fallow rice-fields, road-sides, premises, sea- shores; in the lowlands and lower hills, rarely up to 900 m. Vern. Rumput kotor, rumput kuning. Banka, nanen, Deli, t@ki gedé. S, dékéng. J. lumhungan. Md. takimai. Mol.; Philip.: anand, Ibn., gisai-kalabau, tuhog-daldg, Tag., kaptos, Iv., sangsaiga, Bon. Note. Extremely tall or low specimens, and those with many-flowered spikelets, were described as separate species, subsequently reduced to varietal rank (see synonymy). They hardly deserve nomen- clatural recognition. A collection from Jappen-Biak, with very slender stems, capillary leaves, inflores- cences consisting of 1—4 spikelets only. and 2 involu- cral bracts the lower of which seemingly continues the stem, answers the description of var. capillaceus CLARKE in Urb. Symb. Ant. 2 (1900) 32, recorded from the Antilles and the Himalaya. Presumably it is merely a depauperated form. In African specimens the glumes are sometimes strikingly purple-spotted (like in C. sphacelatus). 10. Section Alternifolii KUNTH, En. 2 (1837) 31. — Cyperus sect. Textiles CLARKE, Kew Bull. add. ser. 8 (1908) 98. Type species: C. alternifolius L. 618 23. Cyperus flabelliformis Rotts. Descr. Pl. rar. Progr. (1772) 22; Descr. & Ic. (1773) 42, t. 12 f. 2; KunrTH, En. 2 (1837) 32; CLARKE, FI. Trop. Afr. 7 (1902) 336; Merr. Fl. Manila (1912) 110; En. Philip. 1 (1923) 105; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 475.—C. flabelliformis var. obtusangulus Boeck. Linnaea 35 (1868) 566. — C. alternifolius L. ssp. flabelliformis KOK. Pfl. R. Heft 101 (1936) 193. — C. alternifolius (non L.) BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 30. — C. alternifolius L. var. obtusangulus KOYAMA, Quart. J. Taiwan Mus. 14 (1961) 177. Perennial with stout, horizontal rhizome. Stems stout, densely tufted, obtusely trigonous to subterete, sulcate, scaberulous at the top, 50—-175cm by 3-8 mm. Normal leaves only present on the first sterile shoots, leaves on the flowering stems reduced to long, wide, brownish sheaths. Inflorescence large, decompound, 10-30 cm across. Involucral bracts numerous, up to 20, distinctly spaced, flat, obliquely patent to reflexed, rather abruptly acuminate, nearly equal in length, much overtopping the inflorescence, 25-50 cm by 8-15 mm. Primary rays numerous, subequal in length, slender, smooth, obliquely erect, 5—10cm long. Spikelets digitately arranged, stellately spreading, in clusters of 3—7(—15), ovate to oblong-linear, compressed, 10—40-flowered, 5—10 by c. 2mm; rachilla straight, wingless, persistent; FLORA MALESIANA [ser. I, vol. 7° internodes c. 4 mm. Glumes membranous, obliquely patent, ovate, obtuse, acutely keeled, mucronulate, faintly 3—5-nerved, with green keel and shining - ferrugineous sides often tinged with red, broadly hyaline- margined, densely (c. 34) imbricate, c. 2 by 114-14 mm. Stamens 3; anthers linear, c. 1 mm, the apical appendage of the connective setulose at the top. Nut trigonous, broadly ellipsoid or Dam obovoid, apiculate, yellowish brown, %-% b YA mm. Distr. Native to Arabia, tropical and South Africa; elsewhere often cultivated for ornamental purposes and sometimes running wild, e.g. in Queensland, New Caledonia, Hawaii, Ryu Kyu Is.; also in Malesia escaped from cultivation: Sumatra West Coast, W. Java, Philippines (Manila, Lamao), often naturalised. Ecol. Waste places, river-banks, wet rice-fields, along ditches, etc.; at low and medium altitudes, up to 1200 m. Vern. Umbrella plant, E, parapluplant, D, papa- yungan, M, mansiang babunga, Sum. W. C. Note. The very closely related C. alternifolius L. differs by its smooth stems, narrower leaves, lanceo- late glumes, and narrowly oblong, blackish nuts measuring c. 1 by }4 mm. It is native to Madagascar, Mauritius, and the Mascarenes; not found growing wild in Malesia. 11. Section Pseudanosporum CLARKE, J. Linn. Soc. Bot. 21 (1884) 117. — Cyperus sect. Natantes CLARKE, Fl. Br. Ind. 6 (1893) 597. Type species: C. platystylis R.Br. 24. Cyperus platystylis R.Br. Prod. (1810) 214; KuntTH, En. 2 (1837) 111; BENTH. FI. Austr. 7 (1878) 264; CLaRKE, J. Linn. Soc. Bot. 21 (1884) 27, 117, t. 1 f. 7-9; Fl. Br. Ind. 6 (1893) 598; VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 103, t. 4 f. 21; RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 63; Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 214; Merr. En. Born. (1921) 55; RipL. Fl. Mal. Pen. 5 (1925) 141; BAck. Onkr. Suiker. (1928) 134, t. 130: KUK. Pfl. R. Heft 101 (1936) 185, f. 21; S. T. BLAKE, J. Arn. Arb. 28 (1947) 215; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 31; KERN, Reinwardtia 2 (1952) 112, f. 8; in Back. & Bakh. f. Fl. Java 3 (1968) 472. — C. pallidus Nees [Linnaea 9 (1835) 284, nom. nud.] in Wight, Contr. (1834) 79; KUNTH, En. 2 (1837) 40.— Anosporum pallidum BOECK. Linnaea 36 (1870) 412; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 52. Perennial with very short rhizome; stolons wanting: roots thick, funiliform. Stems solitary or sut cespitose, stout, rigid, triquetrous, smooth, or scabrous on the angles above. up to 80(—110) cm by (3—)S—8 mm. Lvaves basal, channelled or flat, as long as or longer than the stems, gradually acuminate, septate, nodulose, coriaceous, very scabrous (cutting) on margins and midrib, glaucous or greyish green, 8-—12(—20) mm wide, lower sheaths bladeless, strongly keeled, cinnamomeous to purplish. Inflorescence compound or decompound, often with very numerous spikelets, depressed-corymbose or semiglobose, very dense to rather loose, up to 30 cm across. Involucral bracts 5-8, slightly distant, patent to reflexed, the larger ones much overtopping the inflorescence, up to 80cm. Primary rays up to 12, rigid, smooth, widely spreading, often slightly upcurved, up to 10cm; secondary rays divaricate, up to 4cm, tertiary ones when present very short. Spikelets often very numerous, digitately arranged, in clusters of 3-8, widely spreading, ovate to linear-lanceolate, acute, compressed but somewhat turgid, c. 5—20 by VA- 3 mm, up to 60-flowered; rachilla straight, wingless, persistent; internodes Ye 4A mm. Glumes firmly membranous, appressed, ovate to broadly ovate, obtusish, mucronulate, with rounded, strongly 3-nerved, green back, nerveless, cellular-reticulate, yellowish or brownish sides and narrow, whitish hyaline margins, very densely (?4-%%) ) imbricate, 2-244 by 14- 2mm. Stamens 3; anthers linear, c. 1mm, the connective produced into a bristly appendage. Style flattened, ciliate; stigmas 3, short. Nut unequally trigonous, dorsally compressed, with corky, much thickened angles, ellipsoid, apiculate, shining greyish brown to blackish with straw- coloured or yellow angles, 134-2 by 1 mm; ventral side somewhat concave, dorsal side with a raised angle. Distr. From Ceylon and India to Formosa (here only known from a single locality), and through Malesia to N. Australia, Queensland, and New South Wales; in Malesia very rare in Sumatra, the Malay 1974] Cy PERACEAE (Kern) 619 Peninsula, W. and E. Java, the Lesser Sunda Is. (Sumba), Borneo, Celebes, the Moluccas (Ceram), and New Guinea; not known from the Philippines. Ecol. Very wet places in swamps, on swinging bogs, sometimes in wet rice-fields; at low altitudes, usually below 500 m, in Sumatra (Atjeh) up to 1200 m. Vern. Para, Borneo (Kutei), kéebuamba, Sumba. Note. The corky tissue on the nuts enables dispersal by water. See RIDLEY, Disp. (1930) 238. 12. Section Diffusi Kuntu, En. 2 (1837) 25. — Cyperus sect. Incurvi KUK. Pfl. R. Heft 101 (1936) 216. Type species: C. diffusus VAHL. Spp. 27-32 from New Guinea are insufficiently known. 25. Cyperus diffusus VAHL, En. 2 (1806) 321; Mia. Fl. Ind. Bat. 3 (1856) 264; Boeck. Linnaea 35 (1868) 534: K. Sco. & HO tr. FI. Kais. Wilh. Land (1889) 24: CLARKE, FI. Br. Ind. 6 (1893) 603; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 97, t. 4 f. 17, 18, incl. f. microstachyus VALCK. Sur., excl. f. turgidulus VALCK. SuR.; CLARKE, Philip. J. Sc. 2 (1907) Bot. 83: RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 65: Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 213; Camus, Fl. Gén. I.-C. 7 (1912) 54; MerR. En. Philip. 1 (1923) 104; Ripv. Fl. Mal. Pen. 5 (1925) 142; Kox. Pfl. R. Heft 101 (1936) 208, incl. var. celebicus KUKk., excl. ssp. bancanus KUk.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 215; BAck. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 32; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 477.—C. sorzogonensis PRESL, Rel. Haenk. 1 (1830) 351. — C. scirpoides (non VAHL) PRESL, ibid. 1 (1828) 178. — C. longifolius (non Por.) DECNE, Nouy. Ann. Mus. Hist. Nat. Paris 3 (1834) 359; Descr. Herb. Timor. (1835) 31; KuNTH, En. 2 (1837) 30, quoad specim. timor.; Mig. Fl. Ind. Bat. 3 (1856) 265; RIDL. in Forbes, Wand. (1885) 520.— C. moestus KUNTH, En. 2 (1837) 31; Mia. Fl. Ind. Bat. 3 (1856) 265. — C. racemosus (non Retz.) Mor. Syst. Verz. (1846) 96: ZOLL. Syst. Verz. 1 (1854) 63; StEUD. Syn. 2 (1855) 52, p.p.; Mia. FI. Ind. Bat. 3 (1856) 270. — C. lagorensis STEUD. Syn. 2 (1855) 36, p.p.— ? C. auriculatus (non NEES) STEUD. I.c. 44, p.p.; Mia. FI. Ind. Bat. 3 (1856) 283.—C. holophyllus Mia. l.c., incl. var. celebicus M1q. — C. elegans (non L.) Boeck. Linnaea 35 (1868) 532, p.p., incl. var. moestus BOECK.; CLARKE, J. Linn. Soc. Bot. 21 (1884) 125, p.p. — C. kurzii (non CLARKE) CAMUS, FI. Gén. I.-C. 7 (1912) 54. var. diffusus. — Synonymy as above. Perennial with very short rhizome. Stems tufted, rather stout, trigonous below, triquetrous above. smooth, 15—80 cm by 3—5 mm. Leaves as long as or longer than the stems, channelled at the base, otherwise flat, flaccid, with 3 prominent nerves, rather abruptly acuminate, light green beneath, grassgreen above, 5—10(—20)mm wide; margins scabrous nearly throughout, nerves scaberulous in the upper half; lower sheaths purplish. Inflorescence decompound or supradecompound, diffuse, usually lax, up to 30cm across. Involucral bracts 4—10, spreading or reflexed, unequal, the larger ones much overtopping the inflorescence, up to S50 cm by 16 mm. Primary rays up to 20, unequal, spreading, rigid, smooth, up to 20 cm, secondary ones up to 4cm, tertiary ones short, setaceous. Spikelets digitately arranged, 2-9 together but often some solitary, oblong, turgid-compressed, (4—)6—12(—20)-flowered, (3—)7—10 by 2—3 mm; rachilla partly visible, straight, persistent, narrowly winged; internodes c. )4 mm. Glumes membranous, patulous, boat-shaped, broadly ovate to suborbicular, very obtuse, densely imbricate, with strongly 3-nerved, arcuate, green keel, fuscous, faintly nerved sides and whitish hyaline margins, 1/4(-174) by 1)4(—2) mm; _mucro slightly recurved, setulose at the top, 4—)4 mm. Stamens 3; anthers linear-oblong, rostrate by the produced connective setulose at the top, 4-1 mm. Style hardly any; stigmas 3, exserted from the glume. Nut triquetrous, ellipsoid or subpyramidal, often somewhat surpassing the body of the glume, acute, shortly apiculate, dark brown to dusky black, (ayuU4-LY mm by c. 24 mm. Distr. India, Farther India, S. China, Formosa, Solomon Islands: almost throughout Malesia (not known from the Lesser Sunda Is.). Ecol. In thickets, moist forests (often in deep shade) on river-banks, shady road-sides, at low altitude, rarely up to 700 m. Use. In Mindanao the roots are used as a medicine for diseased lips. Vern. Pukul sédapan, Riouw, parah-parah, rumput bumbat, rumput chukor karbau, Mal. Pen., tjékeng, J, tataboh, tutobok, N. Borneo, ibu,mom, New Guinea: Philip.: barsangd-bakir, Ik., gumi-gumi, haras, tuhog- daldag, Tag.,nipinpin-di-gubat,Sbl.,singao, Mindanao. var. macrostachyus Boeck. Linnaea 35 (1868) 534; KOK. Pfl. R. Heft 101 (1936) 209. — C. pubisquama STEuD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud. ] Syn. 2 (1855) 20; Mig. Fl. Ind. Bat. 3 (1856) 266; CLARKE, FI. Br. Ind. 6 (1893) 604; Philip. J. Sc. 2 (1907) Bot. 83; Camus, Fl. Gén. I.-C. 7 (1912) 56, f. 57. — C. lagorensis StTEUD. Syn. 2 (1855) 36, p.p.; Mia. FI. Ind. Bat. 3 (1856) 275. — C. calacaryensis STEUD. Syn. 2 (1855) 34; Mia. Fl. Ind. Bat. 3 (1856) 275; Naves, Nov. App. (1882) 304. — C. bancanus (non Mia.) NaAvEs, l.c. 302; CLARKE, Philip. J. Sc. 2 (1907) Bot. 83. — C. diffusus (non VAHL) CLARKE, J. Linn. Soc. Bot. 21 (1884) 127.— C. diffusus f. macrostachyus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 99, t. 4 f. 20. — C. diffusus var. pubisquama Hook. f. in Trimen, Handb. Fl. Ceyl. 5 (1900) 28; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 65, p.p.: Merk. En. Philip. 1 (1923) 104; Ripv. Fl. Mal. Pen. 5 (1925) 143. — Fig. 54. Spikelets usually linear, densely up to 30-flowered, less compressed than in var. diffusus, 8-12 by 2 mm. 620 FLORA MALESIANA [ser. I, vol. 7 thf) of AAT AT (| Sz Fig. 54. Cyperus diffusus VAHL var. macrostachyus BOECK. a. Habit, x %, b. spikelet, c. glumes, d. rachis of spikelet, e. deflorate flower, f. nut, all x 10 (a—f BACKER 4268). 1974] Cy PERACEAE (Kern) 621 Rachilla hidden by the glumes, also in fruit. Glumes closely clasping each other even at maturity, ap- pressed, the body somewhat larger than in var. diffusus (1/4 mm), the mucro shorter (4-14 mm). Nut usually smaller, 1-114 by 4-34 mm. Distr. & Ecol. As in var. diffusus; apparently much rarer. Note. Typical var. diffusus and var. macrostachyus look like different species, but they are connected by numerous intermediates precluding specific separation. 26. Cyperus trialatus (BOECK.) KERN, Reinwardtia 3 (1954) 32. f. 1,2; in Back. & Bakh. f. Fl. Java 3 (1968) 477. — Scirpus trialatus BoECK. Flora 42 (1859) 445: Linnaea 36 (1870) 721.— C. bancanus Miq. Sum. (1861) 599; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 65; Camus, FI. Gén. I.-C. 7 (1912) 56, f. 5, 10: RipL. Fl. Mal. Pen. 5 (1925) 143. — C. turgidulus CLARKE, J. Linn. Soc. Bot. 21 (1884) 130: Fl. Br. Ind.6 (1893) 604; Ill. Cyp. (1909) t. 13, f. 4-5. — C. diffusus f. turgidulus VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 100, t. 4 f. 19. — C. helferi (non BoEcK.) CAMus, Fl. Gen. I.-C. 7 (1912) 55. — C. diffusus ssp. bancanus KUK. Pfi. R. Heft 101 (1936) 209, excl. specim. philip. Differs from the closely allied C. diffusus by: Stems 3-winged. Leaves rather rigid. Inflorescence simple or compound, dense. Primary rays short, up to 4cm. Spikelets 8-16 together in globose clusters of 5-7mm g, ovoid, turgid, subterete, 6—8-flowered, 3—4 by 2 mm. Glumes 114-2 mm long, distinctly many-nerved, very shortly mucronulate. Nut larger, 114-124 mm long. Distr. SE. Asia: Thailand, Farther India, S. China, in Malesia only in the western part: Sumatra (East Coast), Banka, Billiton, Malay Peninsula (Setul, Wellesley, Perak, Pahang, Malacca, P. Penang, Singapore), Java (Karimundjawa Is.), S. & SE. Borneo. Fig. 55. Ecol. In secondary forests, wet rice-fields, on road-sides, only in the lowlands, up to 60 m. Fig. 55. Range of Cyperus trialatus (BOECK.) KERN. 27. Cyperus pedunculosus F. v. M. Fragm. 8 (1874) 266; BENTH. FI. Austr. 7 (1878) 272; CLARKE, J. Linn. Soc. Bot. 21 (1884) 131; VALCK. Sur. Nova Guinea 8 (1912) 697; KUK. Bot. Jahrb. 59 (1924) 43, incl. var. floribundus KUGxK.; ibid. 69 (1938) 256, incl. var. atrocastaneus KUK.; Pfl. R. Heft 101 (1936) 222: Mitt. Thiir. Bot. Ver. N. F. 50 (1943) 3, incl. var. alatus Ktx.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 218.— C. montis-sellae K. Scu. Bot. Jahrb. 18 (1894) 186; K. Scu. & Laut. Fl. Schutzgeb. (1900) 191.— ? C. papuanus RIDL. Trans. Linn. Soc. II, Bot. 9 (1916) 241; KUx. Pfl. R. Heft 101 (1936) 222. Perennial with very short rhizome. Stems tufted, usually rather stout, triquetrous with concave sides to distinctly 3-winged, smooth, (5—)10—45 cm by 174-5 mm. Leaves linear, canaliculate at the base, otherwise flat, longer than the stems, gradually attenuate in the long, triquetrous top, scabrid on nerves and margins, with 3 more prominent nerves, 6—20 mm wide; lower sheaths rubiginous. Inflores- cence simple or subcompound, usually loose. In- volucral bracts S—6(—8), patent, linear or linear- lanceolate, the longer ones much overtopping the inflorescence, up to 30(—45) by 2 cm. Primary rays 6-12, obliquely spreading, slender, smooth, 4— 10(—20) cm. Spikelets digitately arranged, 2—4(—7) together or some solitary, oblong-linear, obtusish, compressed though somewhat turgid, 10—40(—50)- flowered, 5—25 (—40) by 214-3 mm; rachilla straight, hardly winged, persistent; internodes 74-1 mm. Glumes subcoriaceous, patulous at the top, broadly ovate to ovate-lanceolate. not rarely broader than long, incurved, c. 14 imbricate, obtuse. minutely mucronulate, 11—17-nerved, ferrugineous to blackish brown, 2)4-3)4 by 2-3 mm. Stamens 3; anthers linear, c. 1 mm, with bristly appendage of the con- nective. Style ciliate, 24-114 mm. Nut trigonous, obovoid or ellipsoid, obtuse or acutish, blackish brown, 1)4-2 by 1-114 mm. Distr. Australia (E. Queensland); in Malesia: scattered throughout New Guinea. Ecol. In primary forests, on shady creek-banks and river-banks, 0—2000 m (according to KUKENTHAL on Mt Saruwaged up to 2700 m). Vern. Gudua, Papua. Notes. Very variable as to size and density of inflorescence, size and colour of glumes, and shape of nut. The varieties described by KUKENTHAL (var. floribundus: stout, with broad leaves and many- flowered spikelets; var. atrocastaneus with blackish brown glumes; var. alatus with distinctly winged stems) do not deserve nomenclatural recognition. C. papuanus RIDL. is only known from the type col- lection consisting of very young specimens collected on Mt Carstensz in W. New Guinea at c. 1300 m. According to RIDLEY it should be allied to C. babakan STEUD., with which species it has nothing to do. It is impossible to distinguish it satisfactorily from the other species of this badly known group. See for instance KUKENTHAL, I.c. 216-217, where C. pe- dunculosus and C. papuanus are opposed by the colour of the glumes. 28. Cyperus neoguineensis KUK. Bot. Jahrb. 59 (1924) 43 (‘neoguinensis’); Pfl. R. Heft 101 (1936) 229. — C. platyphyllos RiDv. Trans. Linn. Soc. II, Bot. 9 (1916) 242, non R. & S. 1817. — ? C. rigidulus Rit. l.c. 241, non VAHL, 1806.—? C. subrigidulus Ktx. Pfl. R. Heft 101 (1936) 230. Perennial with very short rhizome. Stems tufted, rather stout, smooth, 15—35cm by 2-3 mm (see note). Leaves few, linear-lanceolate, as long as or longer than the stems, flat, distinctly narrowed towards the base, gradually long-acuminate, scabrid in the upper part, 10-18 mm wide; lower sheaths 622 rubiginous. Inflorescence simple, contracted into a hemispheric head 1—4 cm across. Involucral bracts 3-5, lanceolate, patent, finally reflexed, up to 25 by 2cm. Rays hardly developed, up to 1 cm long. Spikelets in c. 4 dense clusters, patent, ovate- lanceolate or oblong-lanceolate, compressed but somewhat turgid, c. 10-flowered, 5—6 by 2/4-3 mm; rachilla straight, hardly winged, persistent; internodes c. 4mm. Glumes coriaceous, keeled, ovate, acumi- nate, strongly incurved, 74 imbricate, strongly 11—13-nerved, with pale fuscous sides and whitish hyaline margins, 3 by 2-2/4 mm. Stamens 3; anthers linear, c. 1 mm, with short, smooth appendage of the connective. Style short, 4%4—)4 mm; stigmas 3. Nut trigonous, ellipsoid, obtuse, reddish brown, 1/4 by Ao mm. Distr. Malesia: New Guinea (W.: Carstensz Peak; NE.: Sepik Distr.; Papua). Ecol. In open mountain forests, 1000—1200 m. Note. C. subrigidulus KUx. (based on C. rigidulus RIDL.), only known from the very poor type collection, most probably represents a depauperate state of C. neoguineensis with stems only a few cm tall. The structure of spikelets, flowers, and nuts is the same as in the latter species. RIDLEY’s inadequate de- scription contains several incorrect data (‘‘glumes nearly 1 mm long; nut obovate, distinctly attenuate towards the base; allied perhaps to C. stoloniferus RETZ.’’). 29. Cyperus subpapuanus KUK. in Fedde, Rep. 29 (1931) 196; Pfil. R. Heft 101 (1936) 222. — C. papuanus (non RIDL.) KUK. Bot. Jahrb. 59 (1924) 45. Perennial with very short rhizome. Stems tufted, firm, triquetrous, smooth, 30-60 cm by 2-3 mm. Leaves several, rigid, linear, canaliculate at the base, otherwise flat, longer than the stems, gradually long- acuminate, with 3 more prominent nerves, 8—11 mm wide. Inflorescence subcompound, loose, up to 8 by 6cm. Involucral bracts 3—4, patent, much over- topping the inflorescence, the longest up to 70 cm. Primary rays 6—7, spreading, slender, up to 4 cm; secondary rays very short. Spikelets subdigitately arranged, 3—7 together, stellately spreading, linear, subcompressed, 14—22-flowered, up to 20 by c. 2 mm; rachilla straight, hardly winged, persistent; internodes c. 1 mm. Glumes subcoriaceous, distant, slightly (up to )4) imbricate, oblong-ovate, with truncate or slightly emarginate, excurved apex, keeled, minutely mucronulate, scabrid on the upper part of the keel, with fuscous sides and whitish hyaline margins, 2)4-234 by 124-2 mm. Stamens 3; anthers linear, c. 1 mm, with short, smooth appendage of the connective. Style very short, /4-)4 mm; stigmas 3. Nut trigonous, ellipsoid, brownish, c. 124 by 1 mm. Distr. Malesia, only known from the type collection: NE. New Guinea, Sepik Distr. Ecol. Along brooklet in dense primary forest with few large trees, many Freycinetia and climbing Araceae. Note. Differs from C. papuanus RIDL. by the less numerous rays of the inflorescence and the more distant, distinctly nerved oblong-ovate glumes with scabrid keel and fuscous sides. 30. Cyperus cinereobrunneus KUK. Mitt. Thiir. Bot. Ver. 50 (1943) 3; S. T. BLake, J. Arn. Arb. 28 (1947) 216, f. 1 A; KERN, Reinwardtia 3 (1954) 34, f. 3. FLORA MALESIANA [ser. I, vol. 7° Perennial with very short rhizome. Stems tufted, rigid, triquetrous, scabrid at the top, up to 50 cm by 2 mm. Leaves several, linear, canaliculate at the base, otherwise flat, up to twice as long as the stems, gradually attenuate into the long, triquetrous top, scabrous on nerves and margins, with 3 more prominent nerves, 3!4—6 mm wide; sheaths purplish brown. Inflorescence subcompound, loose. Involucral bracts 4—6, patent, the longer ones far overtopping the inflorescence, up to 50 cm. Primary rays 5—10, spreading, slender, almost filiform, smooth or scabrid at the top, the longest up to 4. cm; secondary rays very short. Spikelets digitately arranged, 3-10 to- gether or some solitary, stellately spreading, oblong- linear, acute, compressed, 12—20-flowered, greyish brown, 6-10 by c. 2 mm; rachilla straight, hardly winged, persistent; internodes c. 4-74 mm. Glumes subcoriaceous, involute and distinctly keeled in the upper part, oblong-ovate, incurved, 74 imbricate, obtuse, apiculate, 15—17-nerved, VA-3 mm long; lower 3—4 glumes shorter, empty. Stamens 3; anthers oblong-linear, °4-1 mm, with short, smooth ap- pendage of the connective. Style 4-1 mm long, ciliate except for the base; stigmas 3, pilose, 4-24 mm long. Nut trigonous, ellipsoid, acute, with concave brown sides and strongly incrassate greyish angles, about half as long as the glumes, c. 1/4 by 4%-% mm. . Malesia: only known from the type collection: New Guinea: Papua, Western Div., Fly River. Ecol. In tufts on forest-floor. 31. Cyperus meistostylus S. T. BLAKE, J. Arn. Arb. 28 (1947) 217, f. 1 B; KERN, Reinwardtia 3 (1954) 36, f. 4. Perennial with very short rhizome. Stems tufted, slender, triquetrous, smooth, 10-35 cm by 2(—2)4) mm. Leaves few, linear, long-attenuate, flat, much longer than the stems, with 3 more prominent nerves, scabrid on margins and nerves in the upper part, 5—8 mm wide; lower sheaths purplish. Inflorescence subcompound or compound, rather loose or con- tracted, up to 5 cm across. Involucral bracts very unequal, 3 or 4 much overtopping the inflorescence, the lowest up to 30cm. Primary rays 4—S, rigid, smooth, up to 5cm, secondary ones very short. Spikelets subdigitately arranged, stellately spreading, 3-8 together, linear, acute, compressed, 8-—16- flowered, (5—)7—8(—12) by 1/4 mm; rachilla straight, hardly winged, persistent; internodes 14-74 mm. Glumes subcoriaceous, oblong-ovate, somewhat in- curved, /% imbricate, rounded at the top, muticous, 13—15-nerved, minutely setulose at apex, brownish red with scarcely hyaline margins, 2—2!4 mm long. Stamens 3; anthers linear, c. 74 mm, with short, smooth appendage of the connective. Style hardly any (rarely up to )4 mm); stigmas 3, pilose, rarely glabrous, c. 14mm. Nut trigonous, with rounded angles and concave sides, ellipsoid, acute, about 74 as long as the glume, brown, 174-1% by % mm. Distr. Malesia: New Guinea, scattered (W.: Mamberamo Distr.; NE.: Mt Michael; Papua: Dieni, Koitak1). Ecol. In rain-forests, usually below 500 m, on Mt Michael at c. 2000 m. Note. Very close to C. cinereobrunneus KUk. and possibly not specifically distinct. it differs by the 1974] Cy PERACEAE (Kern) 623 smooth stem, the less scabrid leaves, the less distinctly digitate, narrower spikelets, the smaller glumes, and the shorter style. 32. Cyperus multispicatus Boeck. Linnaea 38 (1874) 362: CLARKE, J. Linn. Soc. Bot. 21 (1884) 129; FI. Br. Ind. 6 (1893) 604; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 102; CLarkeE, Ill. Cyp. (1909) t. 13 f. 1-3; KUk. Pfl. R. Heft 101 (1936) 213; KERN, Reinwardtia 2 (1952) 114, f. 9; in Back. & Bakh. f. Fl. Java 3 (1968) 476. Perennial with very short rhizome. Stems tri- quetrous, smooth, 30-80 cm by 3—4 mm. Leaves flat, herbaceous, rather abruptly acuminate, with scabrous margins and scabrid nerves, grassgreen on both sides, 5-8 mm wide; sheaths membranous in front, purplish puncticulate. Inflorescence compound to supradecompound, diffuse, lax, up to 40 cm across. Involucral bracts 5—6, patent, the lowest up to 50 by 1 cm. Primary rays 8—many, very unequal, spreading, smooth, up to c. 15 cm, secondary ones up to 5(—7) cm. Spikelets very numerous, partly solitary and long- peduncled, partly 2—3 digitately fascicled, oblong- linear, compressed, 12—18(—30)-flowered, 6—8(—10) by 1'4-2 mm; rachilla straight, hardly winged, persistent; internodes 4-74 mm. Glumes membra- nous, somewhat excurved at the top, ovate or broadly ovate, obtuse, muticous or minutely mucronulate, with green, 3-nerved keel and nerveless, slightly sulcate, stramineous or ferrugineous, purplish punc- ticulate-striate sides, 114-114 by 1-114 mm, 14-4 imbricate. Stamen 1; anther oblong-linear, c. 4 mm; connective produced, bristly at the top. Style short; stigmas 3. Nut trigonous, ovoid or ellipsoid, granu- late, c. % by 4 mm. Distr. Obviously a very rare species, only known from the type collection (Tenasserim or Andamans), Assam, and Malesia: W. Java (Banten: Mt Halimun near Nirmala). Ecol. In Java in swampy open locality in primary forest, at 1500 m. Note. In sect. Diffusi the species stands rather apart by the numerous solitary spikelets, the nerveless grooved sides of the glumes, the single stamen (CLARKE, 1884, wrongly indicates stamens 2), and the small nut. There are strong affinities with sect. Halpani. 13. Section Radiantes VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 84. Type species: C. radians KUNTH. 33. Cyperus radians Nees & MeYEN [in Nees, Linnaea 9 (1835) 285, nom. nud.] ex KUNTH, En. 2 (1837) 95 (‘radicans’); Boeck. Linnaea 35 (1868) 515; CLARKE, J. Linn. Soc. Bot. 21 (1884) 100; Fl. Br. Ind. 6 (1893) 605; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 86, t. 5 f. 2-4; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 66; Fl. Mal. Pen. 5 (1925) 143; KUK. Pfl. R. Heft 101 (1936) 214, incl. var. griffithii KOK. et var. compositus KUK. — C. griffithii STEUD. Syn. 2(1855) 316; CLARKE, J. Linn. Soc. Bot. 21 (1884) 101. — C. macropus Miq. Sum. (1861) 599; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 84, t. 4 f. 7; t. 5 f. 1. — C. sinensis DEBEAUX, Act. Soc. Linn. Bord. 31 (1877) 14, t. 12; Camus, Fl. Gén. I.-C. 7 (1912) 52, f. 6, 1-3. Perennial with very short rhizome. Stems tufted, firm, rigid, trigonous, smooth, much shorter than the inflorescence, 1—5(—10) cm by c. 2 mm (see Notes). Leaves surpassing the stems, canaliculate to con- duplicate, rigid, often recurved, gradually acuminate, more or less scaberulous at the top, 2—S(—8) mm wide; lower sheaths purplish. Inflorescence simple or compound, loose. Involucral bracts 5—7, patent, shorter than to equalling the primary rays, up to 30cm long. Primary rays 4—7, remarkably long, spreading, firm, smooth, up to 30cm. Spikelets 8—15(—30) together in globose or subglobose clusters 8—20 (rarely more) mm @, sometimes a few binate or solitary, stellately spreading, lanceolate to linear, turgid, acute, 4—6(—32)-flowered, 5S—10(—20) by 3-4 mm; rachilla slightly flexuous, persistent, narrowly winged; internodes *4—*% mm. Glumes chartaceous, appressed, broadly ovate, keeled only towards the top, strongly 11—13-nerved, c. 3 mm long and wide, YZ imbricate, with short, more or less excurved mucro, green, straight or curved keel, rufous to sanguineous sides and narrow whitish hyaline margins. Stamens 3;, anthers linear, with short, smooth, reddish appendage of the connective. Style c. 1 mm, ciliate; stigmas 3. Nut trigonous, ellipsoid, broadly stipitate, very shortly apiculate, shining brownish black, 174-174 by c. 1 mm. Distr. SE. Asia: Farther India, S. and E. China, and Formosa, in Malesia restricted to the western part: Malay Peninsula, Banka, Borneo. Ecol. Coastal plant, growing in the sand of the sea-shore, in open dunes, and on wet rocks in the Terminalia-zone. Notes. The long rays of the inflorescence are easily mistaken for stems. The real stem is often extremely shortened and hardly visible. Highly variable species. In 1884 CLARKE dis- tinguished between C. griffithii and C. radians, which he even would like to place in different sections, but which he united in 1893. Also VALCKENIER SURINGAR took C. radians (C. griffithii sensu CLARKE 1884) and C. macropus (C. radians sensu. CLARKE 1884) for different species. The extreme forms are connected by a long series of intermediates. KUKENTHAL’S var. compositus was based on a specimen from Trengganu, leg. RIDLEY, merely a luxuriant plant of otherwise normal C. radians. 624 FLORA MALESIANA [ser. I, vol. 7° 5 14. Section Tenelli CLARKE, Kew Bull. add. ser. 8 (1908) 96. — Cyperus sect. Graciles BENTH. ex Kok. Pfi. R. Heft 101 (1936) 292, non CLARKE, FI. Trop. Afr. Type species: C. tenellus L. f. 34. Cyperus aquatilis R.Br. Prod. (1810) 213; S. T. BLAKE, Proc. R. Soc. Queensl. 51 (1940) 40; J. Arn. Arb. 28 (1947) 219; KERN, Reinwardtia 3 (1954) 46, f. 6. — C. trinervis R.Br. var. aquatilis KUK. Pfl. R. Heft 101 (1936) 294. Annual with fibrous, yellowish roots. Stems very slender, tufted, weak to rather stiff, strongly com- pressed, triquetrous at the top, smooth, 10—35 cm by 1(-2)mm. Leaves few, flat, weak, gradually acuminate, smooth, slightly septate-nodulose, 2—3 mm wide; lower sheaths scarious, stramineous or cinnamomeous, bladeless or shortly laminate. [n- florescence simple or subcompound, very loose. Involucral bracts (1—)2, erect or suberect, usually much shorter than the inflorescence, rarely over- topping it, the longest 3—10(—17) cm. Primary rays 3—5, suberect, slender, often filiform, one of them usually strongly elongated, up to 25cm. Spikelets digitately arranged, (1—)2—6 together, spreading or 8 (1902) 311. reflexed, oblong to oblong-linear, subobtuse, strongly compressed, 10—30-flowered, 4-15 by c. 3mm; rachilla nearly straight, wingless, persistent; inter- nodes c. 4 mm. Glumes thinly membranous, finally patulous, ovate, acute or apiculate, cellular-reticu- late, with strong midrib and a faint nerve in the centre of either side, 114-124 by c. 114 mm, when young is overlapping: centre (c. a the width) green, sides pale stramineous, hyaline, keel slightly arcuate with somewhat excurved top, narrowly winged, not serrulate. Stamens 2; anthers elliptic, c. 4 mm. Stigmas 3. Nut equally trigonous, ellipsoid to slightly obovoid, ara 4 as long as the subtending glume, broadly stipitate, not or hardly apiculate, minutely tuberculate, pale brown, c. % by /4 mm. Distr. N.and E. Australia, in Malesia: a few times collected in SW. New Guinea and Papua. Ecol. In wet places at low altitude, chiefly in coastal districts. 15. Section Halpani KUNTH, En. 2 (1837) 34 (‘Haspani’). Type species: C. halpan L. 35. Cyperus pulcherrimus WILLD. ex KUNTH, En. 2 (1837) 35; Mig. Fl. Ind. Bat. 3 (1856) 267; BoEck. Linnaea 35 (1868) 573; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 48: CLARKE, J. Linn. Soc. Bot. 21 (1884) 132: Fl. Br. Ind. 6 (1893) 600; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 94, t. 4 f. 12-13, incl. f. rectiglumis VALCK. SuR.; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 64; CLARKE, Ill. Cyp. (1909) t. 10 f. 1-3; Koorp. Exk. Fl. Java 1 (1911) 189, non ibid. 4 (1922) f. 211; Camus, FI. Gén. I.-C. 7 (1912) 48; RIDL. FI. Mal. Pen. 5 (1925) 142; Back. Onkr. Suiker. (1928) 133; KUx. Pfl. R. Heft 101 (1936) 242, incl. var. rectiglumis KUK.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 30; KERN, Reinwardtia 3 (1954) 36; in Back. & Bakh. f. Fl. Java 3 (1968) 476. — C. eumorphos Steup. [in Zoll. Syst. Verz. 1 (1854) 62. nom nud.] Syn. 2 (1855) 22; Mig. Fl. Ind. Bat. 3 (1856) 268. — C. silletensis (non NEES) Camus, FI. Gen. I.-C. 7 (1912) 47. Perennial, often flowering the first year; rhizome short, stolons wanting. Stems firm, rigid, tufted, trigonous below, triquetrous above, smooth, 10—40 cm by 1—2(—4) mm. Leaves flat or slightly canalicu- late, rigidulous, gradually acuminate. scaberulous at the top, 2—4(—6) mm wide; lower sheaths stra- mineous to cinnamomeous. [Inflorescence compound or decompound. rather open to very dense, 3—10(—20) cm across. Involucral bracts 3—6, somewhat (1—2 mm) distant, obliquely patent to horizontally spreading, 5—15(—25) cm. Primary rays up to 20, spreading, slender, smooth, 3—5(—12) cm, sometimes very short, secondary ones divergent, ye V4 cm, tertiary ones up to lcm. Spikelets digitately arranged in dense glomerules of 5S—20, oblong to linear, acute, strongly compressed, finally perspicacious because of the wide- “spreading glumes, 10—30(—50)-flowered, 3-10 by 1-1/4 mm; rachilla wingless, straight, persistent; internodes c. 4 mm. Glumes membranous, keeled, ovate, obtuse, minutely mucronulate just below the apex, with arcuate, 3-nerved, green keel, nerveless sides tinged with red or purple, and broad, hyaline margins, c. 1 by % mm, 24 ~Y, imbricate but finally wide-spreading and discrete: tip incurved when dry. Stamen 1; anther oblong-linear, with short, smooth appendage of the connective, c. 4 mm. Stigmas a Nut trigonous, broadly ellipsoid, distinctly broad- stipitate, shortly apiculate, stramineous, c. %4 by ’Amm. Distr. From Ceylon and India through Thailand and Indo-China to Malesia: not rare in the Malay Peninsula, rather common throughout Java, but a few times collected in Sumatra, Borneo, Celebes, and New Guinea; still unknown from the Lesser Sunda Is. and the Moluccas; apparently very rare in the Philippines; Luzon, Palawan, Tawitawi (not mentioned in MEeERRILL’s Enumeration, although already collected in 1906). Ecol. In open, moist or wet localities: swamps, pools, rice-fields, 0-750 m. Note. Whether or not the glumes in herbarium specimens are incurved depends on the degree of maturity of the spikelets. Before the glumes have curled inwards the spikelets are c. 114 mm wide, later on 1 mm. VALCKENIER SURINGAR’S J rectiglumis (with straight glumes and spikelets 1 4mm wide) was based on specimens with relatively young spikelets. 36. Cyperus halpan LINNE, Sp. PI. (1753) 45 (‘haspan’); Mor. Syst. Verz. (1846) 96; ZoLL. Syst. Verz. 1 1974] CYPERACEAE (Kern) 625 (1854) 63; Mia. Fl. Ind. Bat. 3 (1856) 267; BoEck. Linnaea 35 (1868) 574; excl. var. indicus; BENTH. FI. Austr. 7 (1878) 270; CLARKE, J. Linn. Soc. Bot. 21 (1884) 119, t.3 f. 23, 24, t. 4 f. 34; Fl. Br. Ind. 6 (1893) 600; VALCcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 91, t. 1 f. 6, t. 4 f. 9, excl. f. flavidus; CLARKE, Philip. J. Sc. 2 (1907) Bot. 82; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 64; Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 209; Camus, Fl. Gén. I.-C. 7 (1912) 50; Merr. En. Philip. 1 (1923) 105, p.p.; Ripv. Fl. Mal. Pen. 5 (1925) 142; Back. Onkr. Suiker. (1928) 132, t. 127; KUx. Pfl. R. Heft 101 (1936) 247, f. 28 E-G; S. T. Bake, J. Arn. Arb. 28 (1947) 219: Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 30; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 476. — C. micranthus PREsL, Rel. Haenk. 1 (1828) 178, non Nees. — C. minutiflorus PRESL, ibid. 1 (1830) 351; KuNtuH, En. 2 (1837) 102 (‘multi- florus’); Mig. Fl. Ind. Bat. 3 (1856) 287; NAveEs, Nov. App. (1882) 306. — C. dehiscens (non NEES) Naves, /.c. 303. — C. flavidus var. africanus CAMUS, Fl. Gén. I.-C. 7 (1912) 51. — Fig. 56-57. ssp. halpan. — Synonymy as above. Perennial, but often flowering the first year, with short rhizome and reddish roots. Stems slender, weak, solitary or tufted, triquetrous to almost 3-winged, often with a longitudinal median rib on each side, smooth, 10—40 cm. Leaves flat, smooth orscaberulous at the top, 2-5 mm wide; lower sheaths scarious, inflated, cinnamomeous or reddish; sometimes all the leaves reduced to ovate or lanceolate appendages of the sheaths. Inflorescence compound or decompound, loose to rather dense, up to 15 cm across. Involucral bracts 2—3, obliquely patent to widely spreading, all shorter than the inflorescence or one rather longer, up to 10(—15) cm. Primary rays numerous, up to 20, obliquely patent, slender, smooth, up to 15(—20) cm, secondary ones up to wy cm, tertiary ones when present very short. Spikelets digitately arranged, in clusters of 3—6(—10), stellately spreading, linear or linear-lanceolate, acute, strongly com- pressed, 10—30(—40)-flowered, 5-10 by 1-2 mm; rachilla completely hidden by the glumes or almost so, straight, persistent; internodes 14—!4 mm. Glumes membranous, suberect, keeled, oblong- ovate, rather obtuse, muticous, 1-1/4 mm long, 4-7 imbricate; keel 3-nerved, green, sides fuscous to Sanguincous or paler, nerveless. Stamen 1; anther oblong, c ‘4mm, with bristly appendage of the connective. Stigmas 3. Nut trigonous, broadly obovoid, broadly stipitate, shortly apiculate, densely granulate to verruculose (less distinctly so than in C. tenuispica), wee later on yellowish, 0.4—0.45 mm long and wide. Distr. Tropical and subtropical regions of the whole world; throughout Malesia. Ecol. In open, very wet places; a common and characteristic weed ofinundated rice-fields,0—1900 m. ssp. juncoides (LAMK) KUK. in Fedde, Rep. 23 (1926) 184; Pfl. R. Heft 101 (1936) 249.— C. juncoides LaMK, Ill. 1 (1791) 147. — C. halpan var. americanus Boeck. Linnaea 35 (1868) 575.—C. halpan f. evoluta VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 92, t. 4 f. 11. —C. halpan var. flaccidissimus KUx. Pfl. R. Heft 101 (1936) 248, saltem p.p. Differs from ssp. halpan by: Rhizome creeping. Stems usually more robust, up to 80(—100) cm tall, at the base surrounded by long, bladeless, reddish sheaths, rarely leaf-blades well-developed. Spikelets often longer, 1/4 mm wide; internodes of the rachilla 4A-)A mm. Glumes less densely imbricate, not com- pletely hiding the rachilla, narrower, mucronulate, 174-124 mm long. Stamens (2-)3; anthers linear- oblong. Nut slightly larger, 0.55—0.75 mm long. Distr. Pantropical. Ecol. Prefers more natural habitats than ssp. halpan: swamps, margins of lakes, muddy places in marshy forests, swampy grasslands, floating islands; at low and medium altitudes; on Bali, Mt Agung, near fumaroles at 3150 m. Vern. (of C. halpan s.l.). Rémbang, Sum. E. C., para-para, Riouw, para ayér, rumput bilis gantan, r. sumbo, r. tadah ambong, umbot-umbot, Mal. Pen., hiring banju, rumput kudung, Borneo; Philip.: bala- balangutan, manik-manikan, misai-kalabau, Tag., bar- sanga, Ilk., bungot-bungot, P. Bis., zaingal, Sub.; and many others. Notes. C. halpan is extremely polymorphic. The two subspecies are connected by numerous inter- mediates. LINNAEUS misread the vernacular name halpan as haspan. According to Art. 73 of the Code (example of Gluta rhengas) this orthographic error must be corrected. 37. Cyperus tenuispica STEUD. Syn. 2 (1855) 11; KUk. Pfl. R. Heft 101 (1936) 245, f. 28 A-D; KERN, Rein- wardtia 2 (1952) 116; ibid. 3 (1954) 38; in Back. & Bakh. f. Fl. Java 3 (1968) 476. — C. halpan (non L.) Rotts. Descr. & Ic. (1773) 36, t. 6 f. 2; MERR. En. Philip. 1 (1923) 105, p.p. — Scirpus n. 78 Rotts. Descr. & Ic. (1773) 58, t. 17 f. 3, non Scirpus autumnalis L. — C. caespitosus (non Poir.) LLANOs, Fragm. PI. Filip. (1851) 14 (‘caespitorus’); F.-VILL. & NAVES in Blanco, Fl. Filip. ed. 3, 4 (1880) 8. — ? C. halpan var. dicranolepis Mig. Fl. Ind. Bat. 3 (1856) 268, p.p. —C. halpan var. indicus Boeck. Linnaea 35 (1868) 574 (etiam quoad pl. jav. et born. ?).—C. flavidus (non Retz.) CLARKE, J. Linn. Soc. Bot. 21 (1884) 122, t. 3 f. 25; Fl. Br. Ind. 6 (1893) 600; Philip. J. Sc. 2 (1907) Bot. 82; Camus, Fl. Gén. I.-C. 7 (1912) 51, f. 4, 12, excel. var. africanus Camus. — C. halpan f. flavidus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 92, t. 4 f. 10, quoad specim. — Fig. 58. Annual with fibrous, reddish roots, or longer-lived in favourable circumstances. Stems slender, weak, solitary or tufted, triquetrous, smooth, (2—)5—40 cm by 1-2 mm. Leaves flat, gradually acuminate, smooth or scaberulous at the top, flaccid, 2-4 mm wide; lower sheaths bladeless or shortly laminate, scarious, stramineous to cinnamomeous. Inflorescence compound or decompound, relatively large, loose, 3-10 cm across. Involucral bracts 2—3, obliquely patent, the longest somewhat to conspicuously over- topping the inflorescence, up to 15(—25) cm. Primary rays 5—10, suberect to obliquely patent, slender, smooth, up to 10cm, secondary ones 2—2!)4 mm, tertiary ones when present very short. Spikelets digitately arranged in clusters of 3-9, stellately spreading, oblong to linear, rather acute, compressed, 8—30(—50)-flowered, 3—8(—13) by 1-114 mm;; rachilla slightly flexuous, soon visible between the glumes, wingless, persistent; internodes c. 14 mm. Glumes membranous, ovate, slightly keeled towards the top, 626 FLORA MALESIANA [ser. I, vol. 7° Fig. 56. Cyperus halpan L. a. Habit, x 74, b. spikelet, x 13, c—d. glumes, x 20, e. rachis, x 13, f. stamen, x 45, g. nut with style and one stamen, x 20 (a—g LORZING 13609). 1974] . Cy PERACEAE (Kern) 627 Fig. 57. Cyperus halpan L. (fine, pale) in marshy place, with in front Limnophila aromatica, surrounded by the tall grass Sorghum nitidum (siil). Lowland grass thickets at Plosokerep (Indramaju, W of Cheribon) (photogr. VAN STEENIS, 1935). obtuse, soon excurved, 4-1 by 4-3{mm: keel Stamens 1(—2); anthers oblong, “%—-%~ mm, the green, 3-nerved; sides nerveless, hyaline. purplish short apical appendage of the connective smooth. lineolate or with a purplish spot on one side; midrib Stigmas 3. Nut very obtusely trigonous with convex excurrent as a minute mucro just below the apex. sides, subglobose or broadly obovoid or ellipsoid, 628 FLORA MALESIANA [ser. I, vol / Ye ‘ Fig. 58. Cyperus tenuispica SteuD. a. Habit, x /4, b. fruiting spikelet, x 10, c. glume, x 20, d. wingless per- sistent rachilla, enlarged, e. pistil with two stamens, f. two deflorate flowers, with one or two stamens, all x 10, g. nut, x 20 (a—g BACKER 27362). 1974] Cy PERACEAE (Kern) distinctly broadly stipitate, not or hardly apiculate. densely tuberculate, marbly white, 4—-/4(—74) by AW mm. Distr. Widely distributed in the warm regions of the eastern hemisphere: tropical Africa, Ceylon, Nepal, India, Farther India, extending northward to China and S. Japan, southward to tropical Australia; in Malesia: Malay Peninsula (according to KUKENTHAL, /.c.), Sumatra, Java and adjacent islands, Philippines (Luzon); a few times also collected in the Lesser Sunda Is. (Lombok, Timor) and Celebes. Ecol. In very wet places, especially in inundated rice-fields, 0-500 m. 629 Notes. Often overlooked by confusion with C. halpan, but well characterised by the small, obtuse, patulous glumes with excurved top, the rachilla soon visible from the outside, the smooth anthers, and the minute nuts. CLARKE and KUKENTHAL consider it to be always a very short-living plant soon becoming yellowish or blackish. Small specimens flowering and fruiting in a short time are indeed characteristic of the regularly cultivated rice-fields. In wet, grassy places, fallow fields, etc. the plants grow much larger and may live for a considerable time, just as is the case in C. halpan. 16. Section Fusci KUNTH, En. 2 (1837) 37. — Cyperus sect. Difformes CLARKE, FI. Br. Ind. 6 (1893) 599. Type species: C. fuscus L. 38. Cyperus difformis LINNE, Cent. Pl. 2 (1756) 6; Amoen. 4 (1760) 302; Sp. Pl. ed. 2 (1762) 67; Mia. Fl. Ind. Bat. 3 (1856) 269; Boeck. Linnaea 35 (1868) 586: BENTH. Fl. Austr. 7 (1878) 268; CLARKE, J. Linn. Soc. Bot. 21 (1884) 133; Fl. Br. Ind. 6 (1893) 599; VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 96, t. 4 f. 15; CLARKE, Philip. J. Sc. 2 (1907) Bot. 82: Koorp. Exk. Fl. Java 1 (1911) 189; ibid. 4 (1922) f. 212; Camus, Fl. Gén. I.-C. 7 (1912) 49, f. 5, 1-2: Me_rr. En. Philip. 1 (1923) 104; Rupe. Fl. Mal. Pen. 5 (1925) 142; Back. Onkr. Suiker. (1928) 133, t. 129; Kok. Pfl. R. Heft 101 (1936) 237, f. 27 F—H, incl. var. breviglobosus KUK.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 29: KERN in Back. & Bakh. f. Fl. Java 3 (1968) 475. — C. subrotundus LLANOs, Fragm. Pl. Filip. (1851) 14; F.-Vitt. & NAveEs in Blanco, Fl. Filip. ed. 3, 4 (1880) 8. — C. goeringii STEUD. Syn. 2 (1855) 24; Mia. FI. Ind. Bat. 3 (1856) 271; Naves, Nov. App. (1882) 303. Annual with fibrous, reddish roots. Stems rather weak, tufted, triquetrous, smooth, 10—40(—65) cm by (1—)2—3 mm. Leaves flaccid, slightly canaliculate, rather abruptly acuminate, smooth or somewhat scaberulous on the margins at the top, 2-5 mm wide; lower sheaths stramineous to brown. Inflorescence simple or compound, usually rather lax, 114—-5(—7)cm across. Involucral bracts 2—3, patent, but the lowest often suberect (and then the inflorescence seemingly lateral), the 2 largest far overtopping the inflorescence, up to 25cm. Primary rays usually 5—9, spreading, smooth, up to 3(—5)cm, secondary ones when present up to | cm. Spikes globose, very dense, with very numerous spikelets, 8-15 (rarely 3—4) mm across. Spikelets stellately spreading, linear or oblong- linear, compressed but slightly turgid, obtuse, 10—30- flowered, 2/4-5 by 1-1)4 mm: rachilla straight, wingless, persistent; internodes c. /4 mm. Glumes very small, membranous, patulous, slightly keeled, orbicular to broadly obovate, very obtuse or some- what emarginate, muticous, with arcuate, green, faintly 3-nerved keel, purplish or sometimes pale, nerveless sides and broad whitish hyaline margins, muticous, ?4—%% by #4-1 mm, /% imbricate. Stamens (1—)2; anthers elliptic or elliptic-oblong, c. Ao mm. Style very short, 1% .—!% mm; stigmas 3, somewhat longer than the style. Nut triquetrous, ellipsoid to slightly obovoid, broadly stipitate, conically apicu- late, about as long as the subtending glume, shining stramineous or pale brown, 14-74 by c. 4 mm. Distr. Tropical and warm temperate regions of the Old World; Australia; in Central America presumably introduced; from S. Europe to Japan, S. Africa, and through India and Farther India to the Pacific Islands; in Malesia: rare in the Malay Peninsula and Sumatra, common throughout Java, not known from Borneo and the Moluccas, only a few times collected in the Lesser Sunda Is. (Bali, Timor, Alor, Tanimbar Is.), Celebes, and New Guinea: throughout the Philippines. Ecol. In very wet places: open grasslands, river- banks, etc., but especially a characteristic weed of wet rice-fields, often associated with C. halpan, C. iria, and C. sanguinolentus, 0-1400m (according to MERRILL 2000 m); flowering throughout the year. Vern. Djukut péndul,S,djébungan, ramon bréndélan, J, kamis, Kangean; Philip.: baki-baki, bankoan, gilamhon, Bis., ballayang, I\k., pukuungan, Bon. 17. Section Amabiles CLARKE, FI. Br. Ind. 6 (1893) 598. Type species: C. amabilis VAHL. 39. Cyperus cuspidatus KUNTH in H. B. K. Nov. Gen. & Sp. 1 (1815) 204: Boeck. Linnaea 35 (1868) 496; BENTH. FI. Austr. 7 (1878) 267; CLARKE, J. Linn. Soc. Bot. 21 (1884) 88; Fl. Br. Ind. 6 (1893) 598: VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 79, t. 4 f. 2; Koorp. Exk. Fl. Java 1 (1911) 188; KGx. Pfl. R. Heft 101 (1936) 261, f. 29 E, incl. var. angustifolius KUkK.; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 630 246, p. 31; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 475. — C. angustifolius NEES in Wight, Contr. (1834) 79. —C. pusillus (non VAHL) NFES, J. Bot. Kew Misc. 6 (1854) 27. — C. solutus STEUD. Syn. 2 (1855) 14; Miq. FI. Ind. Bat. 3 (1856) 263; NAves, Nov. App. (1882) 302. — C. castaneus f. sundaicus MiqQ. Fl. Ind. Bat. 3 (1856) 261. — C. squarrosus (non L.) F.v.M. Fragm. 8 (1874) 262; Camus, Fl. Gén. I.-C. 7 (1912) 58.—C. uncinatus (non Porr.) CLARKE, Philip. J. Sc. 2 (1907) Bot. 82; RmpDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 64; Camus, Fl. Gén. I.-C. 7 (1912) 47, f. 4, 5~7; Merr. En. Philip. 1 (1923) 108; Rip_. Fl. Mal. Pen. 5 (1925) 141; Back. Onkr. Suiker. (1928) 131, t. 124. — Dichostylis cuspidata PALLA, Bot. Jahrb. 10 (1888) 296. — Fig. 59. Fig. 59. Cyperus cuspidatus KUNTH. a. Habit, nat. size, b. part of spikelet, x 10(a—b VAN STEENIS 12018). Small annual with fibrous, ee roots. Stems tufted, trigonous, smooth, 2-15cm by )4-1 mm. Leaves very narrow, a setaceous, gradually acuminate, rigid, scaberulous in the upper part, 44-1 mm wide; sheaths membranous, striate, reddish brown. Inflorescence simple, often reduced to a single head. Involucral bracts 3—4(—6), patent, the longer ones overtopping the inflorescence. Rays 0-4, filiform, obliquely erect to spreading, smooth, up to 3cm. Spikelets digitately arranged, stellately FLORA MALESIANA [ser I, volhWe spreading, in clusters of up to 20, linear, strongly compressed, up ee 40(—60)-flowered, 74-1 vy —2) cm by (2—)274-3 mm (mucros included); rachilla slightly flexuous, wingless, persistent; internodes c. 4 mm. Glumes subchartaceous, finally obliquely patent, oblong-spatulate, 14 imbricate, the body strongly 3-nerved, emarginate at apex, with green, often purplish lineolate keel and nerveless, ferrugineous to castaneous sides, 1-174 by Me I mm, the strong mucro spreading to recurved, 74- 1 mm. Stamens (2—)3; anthers elliptic, 4-/4 mm. Stigmas 3. Nut trigonous, obovoid to oblong-obovoid, about half as long as the subtending glume, shortly apiculate, brown to castaneous, (4-)?4-3% by 4-% mm. Distr. Pantropical: tropical Africa, throughout India, extending northwards to S. China and S. Formosa, southwards to Queensland; from southern N. America through Mexico and the West Indies to Brazil; almost throughout Malesia (still unknown from New Guinea), but nowhere common. Ecol. In open, moist to rather dry localities, on sandy arable fields with previous soil, sometimes a weed in rice-fields, 0—1100 m. Vern. Rumput para-para, M, watjé namut, Alor; Philip.: salasa, Tag. 40. Cyperus castaneus WILLD. Sp. Pl. 1 (1797) 278; Boeck. Linnaea 35 (1868) 496; BENTH. FI. Austr. 7 (1878) 267; CLARKE, J. Linn. Soc. Bot. 21 (1884) 87; Fl. Br. Ind. 6 (1893) 598; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 81, t. 4 f. 4; RmpL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 63; Camus, Fl. Gén. I.-C. 7 (1912) 46, p.p., f. 4, 8-11; RipL. Fl. Mal. Pen. 5 (1925) 141; KUxK. Pfl. R. Heft 101 (1936) 264; KERN, Reinwardtia 2 (1952) 117.— Dichostylis castanea PALLA, Bot. Jahrb. 10 (1888) 296. — C. amabilis (non VAHL) Camus, FI. Gén. I.-C. 7 (1912) 45. Closely allied to C. cuspidatus. Spikelets narrower, c. 144mm wide (mucros included c. 2 mm), often multiflorous, up to 50(—80)-flowered. Glumes less patent, chestnut-brown, rarely ferrugineous: mucros often less recurved, shorter, ’-% mm. Stamen 1. Nut usually oblong with exactly parallel sides, c. 0.9 by 0.25(—0.3) mm, sometimes smaller and sub- obovoid. Distr. From India, Tonkin, and Cochinchina to tropical Australia, everywhere scattered, in Malesia: Banka, Krakatau, Malay Peninsula (Perak, Kelantan), N. Borneo. Ecol. In open sandy localities, up to 300 m. Note. Also closely related to and possibly not specifically distinct from C. amabilis VAHL, En. 2 (1806) 318. The latter species, not known from Malesia, is mainly characterized by the very shortly mucronulate, golden or ferrugineous glumes. How- ever, in an Australian variety of C. castaneus: var. brevimucronatus KUx. Pfl. R. Heft 101 (1936) 264, the glumes are also but shortly mucronate, and this variety is therefore only distinguishable from C. amabilis by the slightly smaller, dark glumes and the somewhat smaller nut and anther. 18. Section Aristati KUNTH, En. 2 (1837) 20. Type species: C. aristatus ROTTB. 1974] Cy PERACEAE (Kern) 631 41. Cyperus squarrosus LINNE, Cent. Pl. 2 (1756) 6; BENTH. FI. Austr. 7 (1878) 268: KERN in Back. & Bakh. f. Fl. Java 3 (1968) 473. — C. aristatus ROTTB. Descr. Pl. rar. Progr. (1772) 22; Descr. & Ic. (1773) 23, t. 6 f. 1; Mig. Fl. Ind. Bat. 3 (1855) 262; BoECK. Linnaea 35 (1868) 500; CLarkE, J. Linn. Soc. Bot. 21 (1884) 91; FI. Br. Ind. 6 (1893) 606; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 81, t. 4 f. 5; Koorp. Exk. Fl. Java 1 (1911) 188, 192, non ibid. 4 (1922) f. 207; Back. Onkr. Suiker. (1928) 131, t. 125; KUK. Pfi. R. Heft 101 (1936) 502, f. 55 F—J. — C. inflexus MUEHLENB. Descr. (1817) 16. — Mariscus squarrosus CxiaRKE, FI. Br. Ind. 6 (1893) 623, quoad synon. — Dichostylis aristata PALLA, Bot. Jahrb. 10 (1888) 296. — Mariscus aristatus CHERM. Bull. Soc. Bot. France 85 (1938) 366; BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 45. Small annual with fibrous roots, when bruised or dry smelling of fenugreek. Stems tufted, triquetrous to almost 3-winged, smooth, 1-10 cm by /4-1)4 mm. Leaves few, weak, canaliculate, gradually acuminate, nearly smooth, 1-2 mm wide; lower sheaths mem- branous, purplish. Inflorescence simple, often reduced to a single hemispherical head, or with 1-3 rays more or less developed. Involucral bracts 2-4, patent, at least 1 overtopping the inflorescence, up to 7 cm. Rays 0-3, filiform, smooth, up to 3(—5) cm. Spikes very dense, oblong-ovoid to subglobose, appearing echinate, 14-2cm @; rachis 1—S mm. Spikelets densely spicate, spreading, oblong to oblong-linear, strongly compressed, 6—30-flowered, 5—10(-15) by 3-4mm (awns included); rachilla straight or slightly flexuous, wingless; internodes Y-'A mm. Glumes membranous, patulous, elliptic- ovate to elliptic-oblong, the blades 1-2 by 4-1 mm, YA imbricate, strongly (S—)7—9-nerved over the whole breadth, ferrugineous to fuscous, tapering into a strong, recurved, (/4—)/4-1/4 mm long awn. Stamen 1; anther elliptic to elliptic-oblong, 14-34 mm. Stigmas 3. Nut very variable, trigonous, oblong- obovoid, oblong, or almost linear, 4-74 as long as the blade of the subtending glume, broadly stipitate, obtuse, very shortly apiculate, dull greyish brown, 74-1 by “’-A mm. Distr. Tropical Africa, extending to S. Africa; tropical Asia, to S. China; Australia; in the western hemisphere from S. Canada through the United States and tropical America to Argentina and Chile; in Malesia: Madura, Lesser Sunda Is. (Sumba, Timor). KOORDER’s statement that it occurs in the Javan mountains has never been verified. Ecol. In Madura and Sumba in sandy grassy fields, 1-50 m; in Timor on limestone-rocks, up to 900 m. Vern. Pun ana, Timor. Notes. The sheet in the Linnean herbarium marked “‘squarrosus” in LINNE’s handwriting con- tains two species, one of them represented by a tuft of complete plants, the other by a single in- florescence only. The description in Cent. Pl. 2 was undoubtedly drawn up from the complete specimens, which belong to the species described above. For this species CLARKE, KUKENTHAL, and others wrongly accepted ROTTBOELL’s younger synonym C. aristatus. The correct name for the fragment (C. squarrosus sensu CLARKE, KUK.) is apparently C. maderaspatanus WILLD. (not in Malesia). See also CLARKE, J. Linn. Soc. Bot. 21 (1884) 92; ibid. 30 (1894) 305; BENTH. FI. Austr. 7 (1878) 268 in nota. On account of the spikelets allegedly falling off as a whole, KUKENTHAL referred this species to subg. Mariscus. However, the caducity of the rachilla varies considerably. In the Timor specimens the glumes are more readily caducous than the rachilla. See also O’NEILL, Rhodora 44 (1942) 47. 19. Section Humiles Kuntu, En. 2 (1837) 51. — Cyperus sect. Rupestres CLARKE, Kew Bull. add. ser. 8 (1908) 96. Type species: C. rupestris KUNTH. 42. Cyperus teneriffae Por. in Lamk, Encycl. 7 (1806) 245; CLarKE, FI. Br. Ind. 6 (1893) 601; KUK. Pfi. R. Heft 101 (1936) 306, f. 34 H—K: S. T. BLAKE, Proc. R. Soc. Queensl. 51 (1940) 40; KERN, Rein- wardtia 2 (1952) 122, f. 11. — C. rubicundus VAHL, En. 2 (1806) 308; Boeck. Linnaea 35 (1868) 507; CiaRKE, J. Linn. Soc. Bot. 21 (1884) 104.—C. calcicola Doin, Bibl. Bot. Heft 85 (1915) 422, t. 17 f. 10-13 (‘calcicolus’). — Fig. 60. Annual with fibrous roots. Stems tufted, slender, triquetrous, smooth or scabrid above, 5—15(—20) cm by /4(-1) mm, surrounded at the base by broad, inflated, scarious, purplish striate sheaths. Leaves few, weak, flat or conduplicate, gradually acuminate, smooth or scaberulous at the top, 1(—2) mm wide. Inflorescence a single, hemispherical or subglobose head, up to 2/4 cm across. Involucral bracts 2-3, spreading, short, the lowest usually slightly over- topping the inflorescence, rarely up to 5 cm long. Spikelets digitately arranged, stellately spreading, 2-6(—15) together, oblong, obtusish, strongly com- pressed, up to 30(—40)-flowered, 6—20 by 3—4 mm; rachilla straight, wingless, persistent; internodes c. 44mm. Glumes membranous, patulous at the top, acutely keeled, ovate, acuminate, 9—13-nerved over the whole breadth, cinnamomeous to purplish with whitish hyaline margins, about 74 imbricate, the body 2-3 by 114-2 mm, the midrib excurrent in a recurved, c. 14 mm long mucro. Stamens 3; anthers oblong, c. 1mm, with short, smooth appendage of the connective. Style long (c. 1/4 mm); stigmas 3, much shorter than the style. Nut triquetrous, with somewhat concave sides, obovoid, attenuate towards the base, hardly apiculate, brown under the whitish or greyish outermost layer of cells, “a 1 by 4-4 mm. Distr. Widely distributed in Africa (from Teneriffe through tropical Africa to S. Africa and Madagascar), S. Arabia, India (Deccan Peninsula, Poona, Nilghiris), N. Queensland (see below); in Malesia: Lesser Sunda Is. (Timor). 632 FLORA MALESIANA [ser. I, vol. 7° Wag: ; WAR | \ | iy A e | WW | Ecol. In Timor in dry pastures on limestone hills, associated with C. squarrosus and C. hyalinus, up to 900 m. Vern. Pun métan, Timor. Notes. As in N. Queensland this species was observed on one only of the numerous karst hills near Chillagoe, BLAKE I.c. supposed it to have been introduced there. In Timor it is abundant on calcare- ous soil and it is rather unlikely that it was introduced. Some of the Timor specimens agree very well with the type collection of f. petraeus (STEUD.) KUK. Pfil. R. Heft 101 (1936) 307. — C. petraeus Hocust. ex STEUD. Syn. 2 (1855) 29 (Stems very short, 2 cm; spikelets few, shorter).— Most probably a de- pauperate form not deserving nomenclatural dis- tinction. Fig. 60. Cyperus teneriffae Por. a. Habit, nat. size, b. spikelet, x6, c. glume, x5, d. old rachilla, with persistent filaments, x5, e. stamen, x10, f. nut, x 14 (a-f WALSH 167). 20. Section Platystachyi KuntTH, En. 2 (1837) 44. — Sorostachys STEuD. Flora 33 (1850) 229; Syn. 2 (1855) 71. — Cyperus sect. Leucocephali CHERM. ex KUxk. Pfl. R. Heft 101 (1936) 276. Type species: C. leucocephalus RETZ. 43. Cyperus pulchellus R.Br. Prod. (1810) 213; KunrtH, En. 2 (1837) 110; StEuD. Syn. 2 (1855) 53; BENTH. FI. Austr. 7 (1878) 265; NAves, Nov. App. (1882) 301; F. M. BarLey, Queensl. Fl. 6 (1902) 1735; KERN, Reinwardtia 3 (1954) 39, f. 5 dextr. — C. leucocephalus (non Retz.) Nees in Hook. J. Bot. Kew Misc. 6 (1854) 28; CLARKE, J. Linn. Soc. Bot. 21 (1884) 107, p.p.; VIDAL, Phan. Cuming. Philip. (1885) 155; CLARKE, Philip. J. Sc. 2 (1907) Bot. 82; Mer. En. Philip. 1 (1923) 106; KUx. Pfl. R. Heft 101 (1936) 278, p.p. — Sorostachys kyllingioides STEUD. [Flora 33 (1850) 229. nom. nud.| Syn. 2 (1855) 71: Mia. Fl. Ind. Bat. 3 (1856) 296. — C. sorostachys Boeck. Linnaea 35 (1868) 588; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 48. Perennial with short rhizome; stolons wanting. Stems very slender, tufted, trigonous below, tri- quetrous above, smooth, 10—20(—40) cm by 74-1 mm, at the base surrounded by stramineous to brownish cataphylls. Leaves 1 or 2 to the stem, very narrow to almost setaceous, rigid, smooth or scaberulous at the top, 1(-114) mm wide. Inflorescence a single, globose, whitish or pale cinnamon-coloured head 5-10 mm g. Involucral bracts (2—)3(—4), much over- topping the inflorescence, patent, finally reflexed, the largest up to 10 cm. Spikelets numerous, ovate, strongly compressed, 8—12-flowered, (2—)3(—5S) by 14-2 mm; rachilla straight, wingless, persistent; internodes 14-14 mm. Glumes membranous, almost hyaline, obliquely patent, elliptic to oblong, obtusish, slightly keeled, faintly 3-nerved, whitish, purplish lineolate, 1-134 by 24-1 mm. Stamen 1, sometimes in some flowers 2; anther oblong, 4—)4 mm. Style ie 14 mm; stigmas 3. Nut trigonous, obovoid to oblong, slightly attenuate towards the base, shortly apiculate, yellowish to fuscous, 0.5—0.9 by c. 4 mm. Distr. Tropical Africa, Asia, and Australia, everywhere scattered ; in Malesia: Philippines (Luzon, twice collected). Ecol. In open damp places. Notes. By many authors united with C. leuco- cephalus Retz. Obs. 5 (1789) 11 (Kyllinga pierreana Camus, Not. Syst. 1, 1910, 290; Fl. Gén. I.-C. 7, 1912, 27; see KERN, Blumea 10, 1960, 642), which in my opinion is a distinct species occurring from India to Indo-China, mainly characterised by its semi- globose, paucispiculate heads, larger, truncate glumes, and larger, linear-oblong, black nuts. Due to the wrong citation of the Cuming-number collected between 1836 and 1840 (1617 should be 1417), MERRILL /.c. indicated as locality: Mindanao (Misamis). The species was again collected in Luzon in 1918. 1974] Cy PERACEAE (Kern) 633 21. Section Cephalotes VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 76. — Anosporum NEES in Wight, Contr. (1834) 70. — Hydroschoenus ZOLL. & Mor. in Mor. Syst. Verz. (1846) 95. — Trentepohlia Bokck. Bot. Zeit. 16 (1858) 249. — Cyperus subg. Anosporum CiaRKE, FI. Br. Ind. 6 (1893) 597. — Cyperus sect. Anosporum CLARKE in Thiselt.-Dyer, Fl. Trop. Afr. 7 (1902) 310. Type species: C. cephalotes VAHL. Fig. 61. Cyperus cephalotes VAHL. Habit, small specimen from Rawah Bening, x /4. 44. Cyperus cephalotes VAHL, En. 2 (1806) 311; Mia. Fl. Ind. Bat. 3 (1856) 271; BENTH. Fl. Austr. 7 (1878) 263; CLARKE, J. Linn. Soc. Bot. 21 (1884) 25, 34, t. 1 f. 1-6; Fl. Br. Ind. 6 (1893) 597; VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 77, t. 4 f. 1; CLARKE, Ill. Cyp. (1909) t. 6; Koorp. Exk. Fl. Java 1 (1911) 188; ibid. 4 (1922) f. 206; Camus, Fl. Gén. I.-C. 7 (1912) 44, f. 4, 1-4; Corrt, Trop. Natuur 23 (1934) 12, f. 8; KUx. Pfl. R. Heft 101 (1936) 291: Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 28; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 469. — C. monocephalus Roxs. FI. Ind. 1 (1820) 193. — Ano- sporum monocephalum NeErs, Edinb. New Phil. J. 17, n, 34 (1834) 263; in Wight, Contr. (1834) 92; Boeck. Linnaea 36 (1870) 411. — Hydroschoenus kyllingoides ZoL_. & Mor. in Mor. Syst. Verz. (1846) 95; ZOLL. Syst. Verz. 1 (1854) 63; Steup. Syn. 2 (1855) 71; Mig. Fl. Ind. Bat. 3 (1856) 296. — Trentepohlia bifoliata Boeck. Bot. Zeit. 16 (1858) 249.—C. monogynus BOECK. Linnaea 35 (1868) 565; SCHEFFER, Nat. Tijd. N. 1. 34 (1874) 47. — Anosporum cephalotes Kurz, J. As. Soc. Beng. 39, ii (1870) 84. — Ficinia foliaceo-bracteata PreirF. Rev. Gatt. Ficinia (1921) 35. — Fig. 61-62. Perennial. Rhizome emitting slender stolons rooting at the nodes. Stems solitary, rigidulous, trigonous, smooth, 15—40(—55) cm by 1-174(-2) mm, few-leaved in the lower )4. Leaves rather rigid, canaliculate at the base, otherwise flat, gradually narrowed into the triquetrous, scabrid top, grassgreen or greyish green, 2—-4mm wide; lower sheaths cinnamomeous to reddish. Inflorescence an ovoid or conical, lobed head consisting of 1-4 glomerules, 4-14 cm across. Involucral bracts 3—5, one long remaining erect, finally all widely spreading to reflexed, up to 30cm, the dilated base purplish striate, scariously margined. Spikelets numerous, Fig. 62. Cyperus cephalotes VAHL in a drifting mass of Pistia stratiotes in the lowland. Rawah Bening (S. Kediri, East Java) (photogr. CoERT, 1927). 634 FLORA MALESIANA [ser. I, vol. 7 erect to patent, ovate or lanceolate, often curved, 10—16(—24)-flowered, 4—6(—10) by 3—4 mm;; rachilla straight, wingless, persistent; internodes /4—/4 mm. Glumes subcoriaceous, ovate, very densely imbricate, pale brown to castaneous, 3—4 by c. 2 mm; keel acutely prominent to narrowly winged, often scabrid, strongly 3-nerved, produced into a short, slightly excurved mucro; sides obscurely several-nerved. Stamens 3; anthers linear, up to 1/44 mm. Style undivided or obscurely 3-lobed, 3—4mm_ long, gradually widened into the conical, 3-winged, serru- late base. Nut trigonous, ovoid, dorsally compressed, long-stipitate, apiculate by the c. 1mm _ long, persistent style-base, shining brown, 1-1/4 by %-1 mm; stipe of the nut at first narrowly winged, the wings ultimately strongly incrassate, corky, forming 3 yellowish ridges projecting up the angles of the nut. Distr. From India, Farther India, and S. China to tropical Australia, in Malesia: in a few localities in W. and E. Java, E. Borneo (W. Kutei), and New Guinea (Papua: Agu River branch of Middle Fly River). Ecol. On floating islands in tanks, the roots entangled in the mass of the decaying surface vegetation (Pistia, Salvinia, etc.); in W. Kutei floating in rivers; up to 100 m. Vern. Djukut bawangan, 8, gélunggung, W. Kutei. Notes. On this species with its curious style and nut the genera Hydroschoenus, Anosporum, and Trentepohlia were based. The corky thickenings on the nut were mistaken for a gynophore by PFEIFFER, /.c., who described a collection from W. Java as a new species of the African genus Ficinia; see KERN, Blumea 9 (1958) 222. The corky tissue on the nuts enables dispersal by water, like in C. platystylis. See RIDLEY, Disp. (1930) 238. KUKENTHAL, /.c., mentions the species also from ‘‘Malakka (GRIFFITH!)”. The GRIFFITH collection (n. 6172) originates from Burma. 22. Section Dichostylis (LesTIB.) BAILLON, Hist. Pl. fam. CXIX (1893) 338. — Dichostylis Lest1B. Ess. Fam. Cyp. (1819) 39, nom. illeg. — Scirpus sect. Dichostylis FieK, Fl. Schles. (1881) 473. Type species: Scirpus michelianus L. 45. Cyperus pachycephalus KERN, Reinwardtia 2 (1952) 119,f. 10; Blumea 13(1965) 123. — C. pygmaeus (non Rotts.) S. T. BLAKE, J. Arn. Arb. 28 (1947) 219, p. Annual. Stems tufted, rigidulous, triquetrous, smooth, up to 75cm by 1-3mm. Leaves flat, gradually acuminate, scabrid on the margins and nerves, (2—)3—4(—5) mm wide; basal sheaths brownish. Inflorescence consisting of several very dense glo- merules, contracted into a subtriangular-ovoid to subglobose, lobed head, 174-3 cm across. Involucral bracts several, patent, much overtopping the in- florescence, with reddish brown dilated base, the longest up to 40 cm. Spikelets often incurved and contorted, lanceolate, strongly compressed, 6—12- flowered, 4-8 by 1)4-2(-3) mm; rachilla straight, wingless, persistent; internodes c. 14 mm. Glumes pellucid-membranous, oblong, with smooth keel, distinctly 7—9-nerved, Yn imbricate, with pale stramineous sides and green nerves, the body 2-2)4 by 1 mm, the mucros of the lower glumes c. 4% mm, those of the upper ones ?4—*% mm, slightly excurved, smooth or sparsely setulose at the top. Stamens 3; anthers oblong, the cells c. 14 mm, the appendage of the connective c. 14, mm. Style up to 1 by c.'{_ mm, epapillose; stigmas (2—)3. Nut oblong, nearly always biconvex, strongly dorsoventrally compressed, pale brown, c. 1-114 mm long, 4 mm wide, 14 mm thick. Distr. Malesia: W. New Guinea, NE. New Guinea (Sepik Distr.). Ecol. Mud-banks along rivers and channels, on logs floating in lagoons, 50—90 m. Vern. Sera, Maibrat language, simbuai, Sepik. Note. Differs from C. pygmaeus by the stouter stems, the larger heads, the broader leaves, the less densely imbricate, distinctly nerved glumes, with smooth keel, the 3 stamens, the crested anthers, the broader styles, the number of stigmas (prepon- derantly 3), and the larger nuts. 46. Cyperus pygmaeus Rotts. Descr. & Ic. (1773) 20, t. 14 f. 4, 5; KUNTH, En. 2 (1837) 18; Mia. FI. Ind. Bat. 3 (1856) 261; BoEck. Linnaea 35 (1868) 493, excl. var. michelianus; BENTH. FI. Austr. 7 (1878) 262; Naves, Nov. App. (1882) 301; Crarke, J. Linn. Soc. Bot. 21 (1884) 28, 81, t. 24%. 10; VAnCGKe Sur: Gesl. Cyp. Mal. Arch. (1898) 74, t. 2 f. 22; Koorp. Exk. Fl. Java 3 (1912) 357; Back. Onkr. Suiker. (1928) 130, t. 123; S. T. BLAKE, J. Arn. Arb. 28 (1947) 219, p.p.; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 472. — Dichostylis pygmaea Nees, Linnaea 9 (1834) 289; PALLA in Koch, Syn. ed. 3, 3 (1905) 2557. — Juncellus pygmaeus CLARKE, Fl. Br. Ind. 6 (1893) 596; Philip. J. Sc. 2 (1907) Bot. 81; Camus, Fl. Gén. I.-C. 7 (1912) 38; Merr. En. Philip. 1 (1923) 114; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 41.— C. michelianus (L.) DELILE ssp. pygmaeus ASCHERS. & GRAEBN. Syn. 2, 2 (1903) 273; KUK. Pfl. R. Heft 101 (1936) 312, f. 3 F-G.— Kyllinga squamulata (non VAHL) Camus, FI. Gén. I.-C. 7 (1912) 26. Annual. Stems tufted, often pulvinate, triquetrous, smooth, 1-25 cm by 4-1/4 mm. Leaves canalicu- late, gradually acuminate, scabrid on the margins in the upper part, 1-2 mm wide; basal sheaths brown or reddish. Inflorescence consisting of several very dense glomerules, contracted into a triangular-ovoid to subglobose, lobed head up to 1/4 cm long and wide. Involucral bracts 2—7, patent, much over- topping the inflorescence, with dilated base, the longest up to 15cm. Spikelets often incurved and 1974] contorted, ovate to lanceolate, strongly compressed, 10—20-flowered, 3-5 by 1)4-2 mm; rachilla slightly flexuous, wingless, persistent; internodes c. % mm. Glumes thinly membranous, pellucid, distichous but often seemingly placed irregularly because of the twisted rachilla, lanceolate, keeled, acute or with a short (up to 4 mm long) mucro, 3—5-nerved, very densely (c. /£) imbricate, at first whitish, finally pale stramineous, c. 2 by 14 mm; keel more or less spinulose. Stamens 1—2; anthers linear, 14—)4 mm; connective hardly produced. Style 14-34 by /4 mm, epapillose; stigmas 2—3. Nut trigonous or plano- convex, oblong, pale brown, c. 1 by 14—/4 mm. Distr. Mediterranean region, E. Africa, from Asia Minor through S. and E. Asia to Australia; in Malesia very rare: in a few localities in W. and E. Java, the Philippines (Luzon, Mindanao), Central Celebes, and New Guinea. Ecol. Dried-up pools and ditches, fallow rice- fields, muddy river-banks, 0—700 m. Cy PERACEAE (Kern) 635 Notes. As a rule C. pygmaeus is digynous, but there are often some to several trigynous flowers among the digynous ones. Whether the nut is plano- convex or trigonous does not entirely correlate with the number of stigmas: there are often 2 stigmas to the trigonous nuts. The relations between this species and Scirpus michelianus L. (the latter not known from Malesia) constitute an interesting systematic problem. The two sometimes grow together and resemble each other deceivingly. They differ mainly by the phyllotaxy of the spikelets (glumes distichous in C. pygmaeus, spiral in Scirpus michelianus). BOECKELER considered them conspecific; he was followed by several authors (e.g. KUKENTHAL). KUNTH and CLARKE treated them as belonging in different genera. If Scirpus michelianus is not included in Cyperus, the sectional name Dichostylis, of which it is the type species, cannot be used in this genus. 23. Section Pennati KuNTH, En. 2 (1837) 79. — Mariscus sect. Turgiduli CLARKE, Fl. Br. Ind. 6 (1893) 623. Type species: C. pennatus LAMK. 47. Cyperus javanicus Houtt. Nat. Hist. 2, 13 (1782) Aanw. PI. (1), t. 88 f. 1; MeRR. J. Arn. Arb. 19 (1938) 321; S. T. BLAKE, J. Arn. Arb. 28 (1947) 222; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 474. — C. stuppeus Forst. f. [Fl. Ins. Austr. Prod. (1786) 89, nom. nud.| ex KUxK. Bot. Jahrb. 59 (1924) 45. — C. pennatus LAMK, Ill. 1 (1791) 144; Mia. FI. Ind. Bat. 3 (1856) 281; BENTH. FI. Austr. 7 (1878) 284; CLARKE, J. Linn. Soc. Bot. 21 (1884) 194; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 148, t. 6 f. 4; Koorp. Exk. Fl. Java 1 (1911) 187; ibid. 4 (1922) f. 203; Back. Onkr. Suiker. (1928) 142, t. 142; KUk. Pfl. R. Heft 101 (1936) 476, f. 53 A-~G. — C. canescens VAHL, En. 2 (1806) 355; Boeck. Linnaea 36 (1870) 340; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 51. — C. parviflorus VAHL, En. 2 (1806) 352. — Mariscus albescens GAUDICH. in Freyc. Voy. Bot. (1826) 415; CrLarkeE, FI. Br. Ind. 6 (1893) 623; Philip. J. Sc. 2 (1907) Bot. 88; Rip_. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 74; Fl. Mal. Pen. 5 (1925) 149. — C. holciflorus PRESL, Rel. Haenk. 1 (1828) 173; Mia. FI. Ind. Bat. 3 (1856) 282; Naves, Nov. App. (1882) 305.— C. firmus PresL, Rel. Haenk. 1 (1828) 173. — C. imbricatus (non Retz.) LLANOs, Fragm. Pl. Filip. 14 (1851) 17. — ? C. ovatus LLANOs, I.c. 15; F.-VILL. & NAVES in Blanco, Fl. Filip. ed. 3, 3 (1880) 10. — C. stig- matosus STEUD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 38; Mig. Fl. Ind. Bat. 3 (1856) 278; Naves, Nov. App. (1882) 305.— C. anomalus STEUD. Syn. 2 (1855) 37; Mia. FI. Ind. Bat. 3 (1856) 277; Naves, Nov. App. (1882) 305.— Duvaljouvea pennata PALLA, Denkschr. K. Ak. Wiss. M.-N. KI. Wien 84 (1909) 453. — Mariscus stuppeus Merk. Philip. J. Sc. 3 (1908) Bot. 398; ibid. 9 (1914) Bot. 62; Fl. Manila (1912) 113; Sp. Blanc. (1918) 80; En. Born. (1921) 57. — Mariscus pennatus Domin, Bibl. Bot. Heft 85 (1915) 440; Merr. En. Philip. 1 (1923) 113. — Mariscus javanicus MERR. & METC. Lingn. Sc. J. 21 (1945) 4; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 47. — Fig. 63. Perennial with very short rhizome; stolons wanting. Stems stout, tufted, obtusely trigonous or subterete, smooth, densely papillose, 30—70(—110) cm by up to 4mm. Leaves canaliculate, rigid, coriaceous, more or less spongy, septate-nodulose, very gradually acuminate, with drooping top, very scabrous on keel and margins by antrorse spinules, glaucous or greyish green, S—10 mm wide; lower sheaths shining, cinnamomeous to castaneous. Inflorescence large, compound or decompound, up to 15cm across. Involucral bracts (3—)5—7, widely spreading with drooping top, the lower ones much overtopping the inflorescence, up to 75 cm. Primary rays up to 12, patent, rigid, papillose, up to 10 cm, secondary ones short, divaricate. Spikes broadly cylindrical, obtuse, with up to 50 spikelets, 14-2 by 1-114 cm; rachis glabrous, up to 2 cm. Spikelets spicately arranged, at right angles to the rachis or the lower ones reflexed, oblong to lanceolate, hardly compressed, somewhat turgid, falling off as a whole, 6—10(—13)-flowered, 5—9(-13) by 2-214 mm; rachilla flexuous, broadly winged; wings ovate, yellowish white; internodes 4-1 mm. Glumes subcoriaceous, subpatulous, ovate or broadly ovate, rather acute, hardly keeled. cellular-reticulate, 7—9-nerved, 14-74 imbricate, at first pale, afterwards shining brown or purplish- lineolate, with green keel and whitish hyaline margins, 214-34 by 2-214 mm. Stamens 3; anthers oblong-linear, %4-1!4 mm long. Style 1-114 mm; stigmas 3, about as long as the style. Nut trigonous, ellipsoid or subobovoid, apiculate, shining, dark brown to black, 114-114 by %-%_ mm. Distr. From tropical Africa (very rare) and Madagascar through S. Asia to S. China, the Ryu Kyu Is. and Formosa to the Pacific islands, and through Malesia to tropical Australia; common throughout Malesia. Ecol. Characteristic of open, moist to swampy, usually salt or brackish localities near the sea: sandy foreshores, coral reefs, muddy creek-banks, inner 636 FLORA MALESIANA [ser. 1, volar Fig. 63. Cyperus javanicus Hout. a. Habit, x )4, b. spikelet, c. broadly winged rachilla, enlarged, d. glume, e. deflorate flower, f. nuts, all x 10 (a ELMER 10295, details KERN 8404). 1974] border of the mangrove; rarely more inland ; 0—250 m. See RIDLEY, Disp. (1930) 328. Vern. Rumput lingsing, s. sarai, r. suléngséng, séndayan, M, ilat badak, S, dékeng, djékéng, J, adjén-adjén, wlingi, Md, sosa, Timor, pépontak, Talaud, kikisa, New Guinea: Wapi lang.; Philip.: kai-kai, Mbo, obod-ébod, Ibn., tuhog-dalag, Tag. Notes. Very near to and possibly not specifically distinct from C. ligularis L., which is widely dis- tributed in West Africa and tropical America. KUKENTHAL, Bot. Jahrb. 69 (1938) 257, referred CarR 11390 to C. pennatus var. armstrongii (BENTH.) KUx. Pfl. R. Heft 101 (1936) 479 [C. armstrongii BENTH. FI. Austr. 7 (1878) 89], To me it is typical C. javanicus. See also S. T. BLAKE, J. Arn. Arb. 28 (1947) 223. 48. Cyperus holoschoenus R.Br. Prod. (1810) 215; BENTH. Fl. Austr. 7 (1878) 273; KUx. Pfl. R. Heft 101 (1936) 481; Bot. Jahrb. 70 (1940) 463: Bull. Jard. Bot. Btzg III, 16 (1940) 301, incl. var. fuscisquamatus KUK.; in Fedde, Rep. 53 (1944) 101. — Mariscus holoschoenus CLARKE, Kew Bull. add. ser. 8 (1908) 19; Doin, Bibl. Bot. Heft 85 (1915) 441. Perennial with short rhizome (according to BENTHAM I.c. rhizome thick, horizontal or creeping). Stems rigid, obtusely trigonous, smooth, leafy in the lower half, 30—60(—80) cm by up to 3 mm. Leaves coriaceous, canaliculate or conduplicate, septate- Cy PERACEAE (Kern) 637 nodulose, gradually acuminate, scabrid at the top. 3—4 mm wide; lower sheaths spongy, strongly septate- nodulose, fuscous to brown. Inflorescence compound or subdecompound, loose, up to 13cm across. Involucral bracts 3—4, patent, the lower ones much overtopping the inflorescence, up to 70 cm. Primary rays 5—8, obliquely patent, slender, up to 8 cm, secondary ones up to 2/4 mm, tertiary when present Y’-VA cm. Spikes contracted into dense, globular or ovoid clusters, with many spikelets, 4-7 mm g. Spikelets squarrose, ovate, rather acute, turgid. falling off as a whole, stramineous to brown, 4—8- flowered, 2—3 by 2 mm; rachilla straight, wingless or nearly so; internodes c. 74 mm. Glumes char- taceous, with patulous top, elliptic, obtuse, hardly keeled, minutely mucronulate just below the apex, (7—)9—11-nerved, )4 imbricate, stramineous to brown. c. 2 by 1 mm. Stamens 3; anthers linear, c. /4 mm, the connective produced into an ovate, c. 4%, mm long appendage. Style 1-114 mm; stigmas 3, about as long as the style. Nut trigonous. oblong. about as long as the subtending glume, abruptly apiculate, brown, 124 by 4 mm. Distr. Tropical N. Australia and Queensland, in Malesia: New Guinea (SW. New Guinea: Kurik nr Merauke; Papua: W. Division, Dagwa, Oriomo River; Morehead R.). Ecol. In swampy places at low altitudes; in Papua on shallow margins of a lagoon on savannah, 40 m. 24. Section Thunbergiani CLARKE in Thiselt.-Dyer, Fl. Cap. 49. Cyperus lucidus R.Br. Prod. (1810) 218; BoEcK. Linnaea 36 (1870) 355, p.p., excl. pl. brasil.; BENTH. Fl. Austr. 7 (1878) 283; KUk. Pfl. R. Heft 101 (1936) 448. — Mariscus lucidus CLARKE, Kew Bull. add. ser. 8 (1908) 18; Domin, Bibl. Bot. Heft 85 (1915) 442. Rhizome very short, thick; stolons wanting. Stems tufted, stout, prominently trigonous, smooth, not papillose, 30—100(—120) cm by up to 8 mm. Leaves often longer than the stem, rigid, coriaceous, strongly septate-nodulose, flat or canaliculate, very scabrous on the margins in the upper part and on the midrib beneath, very gradually attenuate, 5-20 mm wide; lower sheaths brown, finally dark brown to blackish. Involucral bracts 5—7, often very broad, patent, the lower ones far overtopping the inflorescence, purplish brown at the very base. Inflorescence large, com- pound, up to 10-radiate. Primary rays very unequal in length, compressed, smooth, the longest up to 20 cm, their tubular prophylls purple, up to 4 cm; secondary rays short, sometimes up to 2 cm. Spikes rather dense to dense, the terminal one oblong- cylindric, 2-3 cm wide, the lateral ones smaller, divergent. Spikelets obliquely to widely spreading, spicately arranged, oblong-linear to linear-lanceolate, flattened, acute, falling off as a whole, 6—20 by 114-2 7 (1897) 186. Type species: C. thunbergii VAHL. mm, 3—8-flowered; rachilla distinctly winged; wings whitish hyaline. Glumes erect, finally somewhat patulous, oblong-ovate or elliptic, acute, of a rich brown, with green keel and 2—3-nerved sides, the longest 4—6 mm. Stamens 3; anthers linear, 114-2 mm long; connective shortly produced. Stigmas 3. Nut oblong- -linear, triquetrous, acuminate, greyish brown, 2A-2% by % mm. Distr. Australia (Queensland, New South Wales, Victoria, Tasmania), in Malesia twice collected in New Guinea: Western Highland Distr., Wabag area, Sirunke. Ecol. In reed swamps, forming almost pure communities over large areas; c. 2500 m altitude. Note. According to KUKENTHAL, I.c., by far the greater part of the Australian specimens belong to: var. sanguineo-fuscus (NEES) KUK. Pfl. R. Heft 101 (1936) 449. — C. sanguineo-fuscus NEES, Ann. Nat. Hist. 6 (1841) 46, based on a Brazilian collection and a Tasmanian one. — Differing from typical C. lucidus by the more distant glumes and the linear-oblong nut (obovate in var. lucidus). The New Guinea specimens may belong here, but I have not seen BRown’s type of C. lucidus. 25. Section Pinnati KUK. Pfl. R. Heft 101 (1936) 451. Type species: C. angustatus R.BR. 638 50. Cyperus angustatus R.Br. Prod. (1810) 214; KuntTH, En. 2 (1837) 111; Steup. Syn. 2 (1855) 53; Boeck. Linnaea 38 (1874) 366; BENTH. FI. Austr. 7 (1878) 282; KUK. Pfl. R. Heft 101 (1936) 452. — Mariscus angustatus CLARKE, Kew Bull. add. ser. 8 (1908) 19; Domin, Bibl. Bot. Heft 85 (1915) 444. Perennial with very short rhizome. Stems slender but rigid, densely tufted, obtusely trigonous, smooth, striate, 30 cm by 2-3 mm. Leaves few, mostly shorter than the stems, narrowly linear, rigid, involute, not or hardly septate-nodulose, greyish green, scabrid on the margins in the upper part, 2-4 mm wide. Inflorescence compound, loose. Involucral bracts 3-6, the lower 1—2 much overtopping the inflores- cence. Primary rays 6—12, obliquely spreading, very unequal, slender, smooth, the longest up to 12 cm, secondary ones capillary, divaricate, up to 3 cm. Spikelets stellately spreading, in clusters of 3-10 together, narrowly linear, acute, scarcely com- pressed, 8—14(—20)-flowered, 6—12 by 1 mm; rachilla narrowly hyaline-winged, persistent. Glumes mem- branous, ovate, appressed or finally slightly spreading, obtuse, keeled, shortly mucronulate just below the apex, rather remote (c. 4 imbricate), with green keel and yellowish to golden brown sides, 7-nerved, c. 2 by 1/4 mm. Stamens 3; anthers short, linear- oblong, c. 4 mm long, with short, smooth, hyaline appendage of the connective. Style short; stigmas 3, longer than the style. Nut about as long as the subtending glume, triquetrous, linear-oblong, obtuse, apiculate, brown, c. 2 by 4 mm. Distr. Tropical Australia (N. Australia, Queens- land), in Malesia: once (1962) collected in S. New Guinea, near Kurik. Ecol. In wet localities, at low altitude. Notes. The species belongs to a most difficult group of Australian forms which needs to be mono- graphed. According to S. T. BLAKE, Proc. R. Soc. Queensl. 51 (1940) 42-47, KUKENTHAL’s arrangement of the forms and his key to them in Pflanzenreich is very unsatisfactory. The New Guinea specimens are much larger than the type, a single culm in BM, of which there seem to be no duplicates anywhere. Also in Australia the species has been rarely collected. CLARKE and KUKENTHAL placed C. angustatus and allies in the genus resp. subgenus Mariscus. Domi and S. T. BLAKE, I.c., pointed to the fact that in some species of the group the rachilla is persistent and the glumes are deciduous, while in others the FLORA MALESIANA [ser. I, vol? characters of subg. Mariscus and subg. Cyperus are combined. 51. Cyperus fulvus R.Br. Prod. (1810) 215; KUNTH, En. 2 (1837) 111; SrEuD. Syn. 2 (1855) 53; BENTH. Fl. Austr. 7 (1878) 274; KUx. Pfl. R. Heft 101 (1936) 456: S. T. BLake, J. Arn. Arb. 35 (1954) 237. — C. sieberi KUNTH, En. 2 (1837) 96; STeuD. Syn. 2 (1855) 51; Boeck. Linnaea 35 (1868) 608. — Mariscus fulvus CLARKE, Kew Bull. add. ser. 8 (1908) 18; Ill. Cyp. (1909) t. 30 f. 7-8; Doin, Bibl. Bot. Heft 85 (1915) 443. Perennial with very short rhizome. Stems rigid, densely tufted, trigonous, or triquetrous above, smooth or scabrid below the inflorescence, striate, (10—)30—60 cm by 2—3 mm, the base almost bulbous by broad, brown, many-nerved sheaths. Leaves crowded, mostly shorter than the stems, linear, rigid, keeled-complicate, septate-nodulose, light or greyish green, scabrous on the margins, (2—)4 mm wide. Inflorescence simple or subcompound. In- volucral bracts 4—S, patent, finally reflexed, lower 2 overtopping the inflorescence. Primary rays 5—10, rather firm, obliquely spreading, unequal, up to 5cm, secondary ones when present very short, divaricate. Spikes globose or hemispherical, very dense, with 6—numerous spikelets, 5-10mm oa. Spikelets stellately spreading, linear-lanceolate, rather acute, slightly compressed, 6—many- -flowered, 5-20 by 2—3 mm;rachilla very narrowly winged, deciduous or persistent. Glumes firm, broadly ovate, finally patulous, obtuse, shortly mucronulate, somewhat remote, with green keel and fulvous or golden brown sides, shiny, 7—9-nerved, 2-24 mm long. Stamens 3; anthers linear, 74-1 mm, with short, smooth ap- pendage of the connective. Style very short: stigmas 3, longer than the style. Nut somewhat shorter than the subtending glume, triquetrous, oblong-obovate, coni- cally apiculate, yellowish brown, densely puncticu- late, 174-2 by 74-1 mm. Distr. Widely distributed in NE. Australia, in Malesia once collected: New Guinea, Papua, near Quaipo (MACGREGOR in 1889). Not seen. Note. Highly variable. S. T. BLAKE, l.c., referred the Papuan collection to var. confusus (CLARKE) KOK. Pfl. R. Heft 101 (1936) 456. — Mariscus fulvus var. confusus CLARKE ex DomIn, Bibl. Bot. Heft 85 (1915) 444, although in his opinion it is doubtful how much importance can be assigned to the varieties that have been described. See also Doin, I.c. 26. Section Flabelliformes (CLARKE) KERN, comb. nov. — Mariscus sect. Flabelliformes CLARKE, FI. Br. Ind. 6 (1893) 624. — Cyperus sect. Subulati CLARKE, Kew Bull. add. ser. 8 (1908) 100. Type species: C. subulatus R.BR. 52. Cyperus compactus Retz. Obs. 5 (1789) 10; KOK. Pfl. R. Heft 101 (1936) 423, incl. var. macro- stachys Ktk. et f. decolorans Ktx., excl. var. pauciflorus KUx.; Bot. Jahrb. 69 (1938) 256554 3 BLAKE, J. Arn. Arb. 28 (1947) 221; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 473. — C. dilutus VAHL, En. 2 (1806) 357; Mig. Fl. Ind. Bat. 3 (1856) 285; Boeck. Linnaea 36 (1870) 354, incl. var. macrostachys BoECK.; CLARKE, J. Linn. Soc. Bot. 21 (1884) 193; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 146, t. 5 f. 3; Koorp. Exk. Fl. Java 1 (1911) 187; ibid. 4 (1922) f.202; Kix. Bot. Jahrb. 59 (1924) 45, incl. f. decolorans KUxK.; Back. Onkr. Suixer. (1928) 141, t. 141. — Mariscus microcephalus PRESL, Rel. Haenk. 1 (1828) 1974] CYPERACEAE (Kern) 639 @ioss Fig. 64. Cyperus dietrichiae Boeck. a. Habit, x 7%, b. spikelet, x 4, c. rachilla, x 8, d. glume, x 8, e. nut, x8 (a—e PARKINSON s.n., from New Britain). 640 FLORA MALESIANA [ser. I, vol. 7° 182; Mia. FI. Ind. Bat. 3 (1856) 290; CLARKE, FI. Br. Ind. 6 (1893) 624: Philip. J. Sc. 2 (1907) Bot. 88; RIpL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 74; FI. Mal. Pen. 5 (1925) 149. — Mariscus dilutus NEES in Wight, Contr. (1834) 90; MerR. En. Philip. 1 (1923) 112. — C. haenkeanus KuNTH, En. 2 (1837) 93. — C. quadriflorus LLANOS, Fragm. Pl. Filip. (1851) 17 (‘cuadriflorus’); F.-VitL. & NAves in Blanco, FI. Filip. ed. 3, 3 (1880) 12. — C. septatus StEup. [in. Zoll. Syst. Verz. 1 (1854) 62, nom. nud.] Syn. 2 (1855) 46; Mia. FI. Ind. Bat. 3 (1856) 284. — C. micro- cephalus Naves, Nov. App. (1882) 304, non R.BR. 1810. — ? C. grabowskianus BorcK. Bot. Jahrb. 5 (1884) 502. — C. luzoniensis LLANOs in Blanco, FI. Filip. ed. 3, 3 (1880) 12. — Sphaeromariscus micro- cephalus Camus, Not. Syst. 1 (1910) 239; Fl. Gén. I.-C. 7 (1912) 79. — Mariscus compactus BOLDINGH, Zakfl. Landbouwstr. Java (1916) 77; DRucE, Rep. Bot. Exch. Cl. 1916 (1917) 634: BAcK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 49. — Duvaljouvea diluta PALLA, Allg. Bot. Zeitschr. 17 (1912) Beil. 8. Perennial with very short rhizome; stolons wanting. Stems subcespitose, usually robust, obtusely trigo- nous or subterete, smooth, (15—)50{—100) cm by up to 6mm. Leaves rigid, strongly canaliculate, strik- ingly septate-nodulose, scabrous on the margins and midrib in the upper part, glaucous, 5—10(—12) mm wide; lower sheaths spongy, strongly septate- nodulose, reddish brown. Inflorescence usually large, compound or decompound, up to 20(—30) cm across. Involucral bracts numerous, patent, finally reflexed, the lower ones much overtopping the inflorescence, up to 1 m. Primary rays (6—)8—12, spreading, up to 18 cm long, secondary ones up to 3 cm, sometimes branched again. Spikes globose or subglobose, very dense, with numerous (—70) stellately spreading spikelets, 1—2(—4) cm across; rachis short, up to 4 mm. Spikelets squarrose, linear-lanceolate, almost subulate, subterete or slightly compressed, falling off asa whole, 4—8(—14)-flowered, 5-15 by 1-114 mm; rachilla flexuous, distinctly winged: internodes 1)4- 1°4 mm. Glumes membranous, convolute, with rounded or slightly keeled back, oblong-lanceolate, obtusish, muticous, 3-nerved in the green centre, with faintly 2—3-nerved, reddish brown sides (rarely paler), remote (4- 1/ inbricate), ), 3- 44 by 1-14 mm. Stamens 3; anthers oblong- -linear, %-1 mm. Style halfway 3- fid. Nut trigonous, oblong-linear, slightly convex on one side, apiculate-rostrate, brown, 14-2 by c. 4% mm. Distr. From India to S. China and Formosa; introduced into Reunion and Mauritius; throughout Malesia, in most parts common. Ecol. Swamps, wet rice-fields, swampy grassfields, coastal marshes, forest-edges, along ditches and river-banks, 0-500 m, sometimes up to 1000 m; often gregarious and then striking by the large, reddish brown inflorescences. Vern. Prumpungan, M (Deli), peperah, Alas-lands, djékéng, S, sukét téké, J, pajong krah, uru humbut, E. Borneo, totoboh, N. Borneo, kikisanki, New Guinea: Wampi lang.; Philip.: baki-baking-puld, S. L. Bis., durugi, Sub., giron, Bag., kadang-kadang, Bik., tikai, Tag. Note. I have not seen the GRABOWSKY collection from Borneo on which BOECKELER founded his C. grabowskianus, “‘species insignis in viciniam C. scabridis et C. ehrenberghiani’. 1 doubt whether KUKENTHAL’s reduction to C. compactus is right as the young plant was described as having cylindrical- ovoid, small spikes and 2-flowered spikelets. 53. Cyperus dietrichiae Boeck. Flora 58 (1875) 87; KOK. Bot. Jahrb. 59 (1924) 46; Pfl. R. Heft 101 (1936) 424, f. 48A—C. — C. trichostachys BENTH. FI. Austr. 7 (1878) 287. — Mariscus dietrichiae CLARKE, Kew Bull. add. ser. 8 (1908) 20. — Fig. 64. Perennial with very short rhizome; stolons wanting. Stems tufted, rigid, triquetrous, smooth, 30—80 cm by i-2mm. Leaves herbaceous, flat, gradually acuminate, scabrid in the upper part, (1—)2—4(—7?) mm wide; lower sheaths membranous, pale. Inflores- cence simple or subcompound, loose. Involucral bracts 3—7, patent, the lower ones as long as or overtopping the inflorescence. Primary rays 5—10, spreading, very slender, smooth, the longer ones up to 20 cm, often with 1—2 setaceous, patent or reflexed, up to 4cm long secondary rays. Spikes broadly ovoid, with 10-25 spikelets, 2/4—3(—4) cm _ wide; rachis 1-114 cm long. Spikelets filiform, subterete, acute, 4—8- flowered, 10-15 by 7/4 mm, the upper ones erect to spreading, the lower ones at right angles to the rachis to strongly reflexed, falling off as a whole; rachilla slightly flexuous, broadly winged; wings whitish hyaline; internodes c. 2 mm. Glumes remote (the top not reaching the base of the next higher one at the same side of the rachilla), appressed or only the apex somewhat patulous, narrowly oblong, acutish, not keeled, muticous, with green midnerve and rufous sides, 9—11-nerved, 3-34 by 1 mm. Stamens 3; anthers small, oblong- linear, c. “mm. Stigmas 3. Nut trigonous, oblong- linear, acuminate, apiculate, brown, 2-274 by YA-% mm. Distr. Tropical Australia (Queensland), in Malesia: New Britain, twice collected. Ecol. On black volcanic soil, at low altitude. Note. In habit, and by the narrow spikelets, the distant glumes, and the narrow nut, very similar to and therefore confused with C. distans L. f., in which, however, the rachilla is persistent, the spikelets are c. 1 mm wide, the lowermost spikelets never strongly reflexed, the glumes shorter, more obtuse, with nerveless sides, and the rachilla-internodes and nuts shorter. 27. Section Strigosi KUK. Pfl. R. Heft 101 (1936) 404. Type species: C. strigosus L. 54. Cyperus stenophyllus VALCK. Sur. Nova Guinea 8 (1912) 701, t. 114; KUk. Bot. Jahrb. 59 (1924) 47, incl. var. ornans KUK.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 221; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 474. — Mariscus flabelliformis (non KUNTH) CriaRKE, Philip. J. Sc. 2 (1907) Bot. 89. — Mariscus 1974] Cy PERACEAE (Kern) 641 tenuifolius (non NEES) CLARKE, l.c. 87; MERR. En. Philip. 1 (1923) 114.—C. ornans VALCK. Sur. Nova Guinea 8 (1912) 700, t. 113; KUx. Pfl. R. Heft 101 (1936) 418, incl. var. stenophyllus KUK.; Bot. Jahrb. 69 (1938) 256. Perennial with short rhizome; stolons absent. Stems tufted, slender, triquetrous, smooth, thickened at the base, 10-40 cm by 4-1 mm. Leaves weak to rather stiff, gradually acuminate, scabrid in the upper part, (1—-)2—4 mm wide; lower sheaths dark brown. Inflorescence simple, loose or more or less con- tracted, sometimes reduced to a single ray. Involucral bracts 3-5, patent, much overtopping the inflores- cence, up to 30(—50) cm long. Rays (1—)3—6, divergent, slender or very slender, smooth, up to 4 cm. Spikes broadly ovoid, with c. 8—25 spikelets, 1—2 by 3(-4) em:;rachis short, 1-174 cm. Spikelets at first obliquely erect, finally horizontally spreading to somewhat reflexed, linear, acute, slightly compressed, falling off asa whole, up to 12-flowered, 10-20 by 1— 1/4 mm; rachilla flexuous, broadly winged: wings hyaline, tightly clasping the nut; internodes 1/4-2 mm. Glumes membranous, appressed, oblong, subacute, minutely mucronulate, hardly or slightly imbricate, strongly 7—9-nerved, with green keel, stramineous sides, and hyaline margins, 324-4 by 124-2 mm. Stamens 3: anthers linear, 1-2 mm. Style 1-1)4 mm; stigmas 3. Nut trigonous, oblong- -ellipsoid, concave on the ventral side, with straight dorsal angle and convex lateral ones, shining, light brown, 2-2/4 by 4-1 mm Distr. Solomon Islands, Melanesia, in Malesia: Java (mainly in the eastern part), Lesser Sunda Is. (Babar), Philippines (Palawan, Luzon, Panay, Mindanao, Sulu Is.), Moluccas (Ternate), New Guinea. Ecol. Damp to dry places, usually under light shade: road-sides, open forests, river-banks, thickets, at low altitude, up to 750 m. Vern. Dékéng, djémbrahan raman, J; bakis-bakisan, Tag. Note. Near to the Australian C. bowmannii F.v.M. ex BENTH., which differs by the slightly narrower spikelets, the lighter coloured basal sheaths, and the remote glumes. Philip. : 28. Section Mariscus (VAHL) BENTH. FI. Austr. 7 (1878) 288. — Mariscus VAHL, En. 2 (1806) 372. — Cyperus sect. Umbellati CLARKE, FI. Br. Ind. 6 (1893) 620. Type species: Mariscus capillaris VAHL. 55. Cyperus cyperinus (RETZ.) VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 154, t. 6 f. 10; Koorp. Exk. Fl. Java 1 (1911) 187: Back. Onkr. Suiker. (1928) 143, t. 143: KUk. Pfl. R. Heft 101 (1936) 518; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 474. — Kyllingia cyperina RETZ. Obs.6(1791)21. — Mariscus cyperinus VAHL, En. 2 (1806) 377; CLARKE, FI. Br. Ind. 6 (1893) 621; Philip. J. Sc. 2 (1907) Bot. 87; RmDL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 73; CLARKE, Ill. Cyp. (1909) t. 22, f. 3-4; MerR. En. Philip. 1 (1923) 112; RIDL. Fl. Mal. Pen. 5(1925) 149: BAck. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 47.— Mariscus umbellatus (non VAHL) Mor. Syst. Verz. (1846) 98; ZOLL. Syst. Verz. 1 (1854) 63. — Mariscus sundaicus Mia. Fl. Ind. Bat. 3 (1856) 289. — Mariscus umbellatus f. contractus Mig. Sum. (1861) 600. — C. umbellatus (non BENTH.) NAves, Nov. App. (1882) 305. — C. sundaicus Naves, l.c. 306.— C. steudelianus (non BoEck.) NAVES, I.c. 306. — C. umbellatus f. cyperinus CLARKE, J. Linn. Soc. Bot. 21 (1884) 200.—C. manilensis BoECK. Bot. Jahrb. 5 (1884) 501. — C. paniceus (non Boeck.) O. K. Rev. Gen. Pl. 2 (1891) 750. — Mariscus umbellatus var. cyperinus CAMUS, Fl. Gén. I.-C. 7 (1912) 77.—C. cyperoides ssp. cyperinus KUK. Bot. Jahrb. 59 (1924) 46; Candollea 6 (1936) 424. Perennial with very short rhizome; stolons absent. Stems tufted, rigid, triquetrous, smooth, somewhat incrassate at the base which is clothed with the fibrous remains of old leaf-sheaths, 15—45(—60) cm by 1-3 mm. Leaves shining greyish green, rigid, flat or canaliculate, long-attenuate, scabrid on margins and keel, 3-6 mm wide; lower sheaths reddish to dark brown. Inflorescence simple (very rarely sub- compound), usually contracted and head-like. In- volucral bracts 5—10, obliquely patent to patent. much overtopping the inflorescence, the longest up to 40 cm. Rays 5—10, spreading, usually less than 1 cm, rarely longer than 2 cm. Spikes broadly cylindrical to obovoid, usu_lly attenuate towards the base, dense, with up to 80 spikelets, 1-2 cm by 8-10 mm. Spikelets obliquely erect, rarely more spreading, oblong-linear, subterete also in fruit, (1—)2—3(—4)- flowered, falling off as a whole, 3—5(—7) by 1-1/4 mm: rachilla broadly winged ; wings hyaline, tightly clasping the nut, c. /4 mm wide; internodes 14-2 mm. Glumes subchartaceous, appressed, elliptic, rather obtuse, muticous or minutely mucronulate, JA imbricate, with green, 3-nerved keel and brownish green to fuscous, 3—4-nerved sides, 3—4 by 174-2 mm. Stamens 3; anthers linear, 1— vy mm. Stigmas 3, longer than the style. Nut trigonous, ellipsoid or oblong- ellipsoid, slightly curved, shortly apiculate, brown, (1%-)2-2)4 by %- x) mm. Distr. From SE. Asia (India, Farther India, S. and E. China, Formosa, Ryu Kyu Is.) to Melanesia, Polynesia and Australia (Queensland); common throughout Malesia. Ecol. In open or partly shaded, somewhat moist to rather dry localities: secondary forests, old clearings, road-sides, river-banks, etc., at low and medium altitudes (0—-1300 m); on Mt Tengger (E. Java) up to 2100 m, on Mt Rindjani (Lombok) up to 2000 m. Vern. Rumput katélan, r. payong, r. pinang, M, walinian, S, papara, Talaud; Philip.: alusang, alusang- parang, ubud-ubud, Tag., bubuyankat, Sub., busikad- dako, C. Bis., hand, Iv., saka-dn, Ig., silal, Bik. Note. Typical, fruiting specimens can be dis- tinguished from the closely related C. cyperoides by the stiff leaves, the contracted inflorescence, the shorter and denser spikes, the obliquely erect, darker 642 FLORA MALESIANA [ser. I, vol. 7° spikelets, the broader, more obtuse glumes, and the broader, ellipsoid nuts; all these characters are very variable. See also varieties. var. maximus (CLARKE) KUK. Pfl. R. Heft 101 (1936) 520. — C. andersonianus BoECK. Bot. Jahrb. 5 (1884) 502. — Mariscus cyperinus var. maximus CLARKE, J. Linn. Soc. Bot. 34 (1898) 43. — C. cyperinus f. maxima CLARKE ex VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 156, t. 6 f. 11.—C. cyperoides ssp. cyperinus var. andersonianus KUK. Bot. Jahrb. 59 (1924) 46. Stems stouter, up to 3/4 mm thick. Bracts up to 12, very long, up to 60 cm. Rays of the inflorescence up to 16, often well-developed, up to 8 cm. Spikes very dense, up to 14cm wide. Spikelets patent, up to 8 mm, 4-flowered. Distr. India; in Malesia: Sumatra, Java, Borneo, Sumbawa, Mindanao. Note. Shows several characters of C. cyperoides, but the nut is that of C. cyperinus. f. curvatus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 156. Stems slender, c. 1 mm thick. Leaves narrow, 1—2 mm wide. Spikes short, c. 1cm long and wide. Spikelets curved. Distr. Malesia: W. Java (Wijnkoopsbaai, JUNGHUHN; Krawang, BLUME). Not _ recently collected. Note. By KUKENTHAL (Pfl. R. Heft 101, 1936, 519) referred to C. cyperinus var. laxatus (CLARKE) KUk., though the specimens on which the forma was based are cited (I.c. 526) under C. paniceus (RottB.) BOECK. By its slender habit similar to this species, but differing by the absence of stolons, the larger, often 2-flowered spikelets, and the larger nuts. The Malesian specimens cited by KUKENTHAL, I.c., under C. cyperinus var. pictus (NEES) KUK. and var. bengalensis (CLARKE) KUK. are in my opinion typical C. cyperinus. 56. Cyperus cyperoides (L.) O. KUNTZE, Rev. Gen. Pl. 3, 2 (1898) 333; KUk. Pfl. R. Heft 101 (1936) 514; S. T. BLAKE, J. Arn. Arb. 28 (1947) 224: KERN in Back. & Bakh. f. Fl. Java 3 (1968) 474. — Scirpus cyperoides LINNE, Mant. 2 (1771) 181. — Kyllingia sumatrensis RETZ. Obs. 4 (1786) 13.— Kyllinga umbellata Rotts. Descr. & Ic. (1773) 15, non t. 4 f. 2. — K yllingia umbellata var. sumatrensis WILLD. Sp. Pl. 1 (1797) 258. — Mariscus umbellatus VAHL, En. 2 (1806) 376; Mia. FI. Ind. Bat. 3 (1856) 288; Camus, FI. Gén. I.-C. 7 (1912) 76, excl. var. paniceus CLARKE et var. cyperinus CAMUS. — Mariscus sieberianus Nees [Linnaea 9 (1835) 286, nom. nud.] ex CLARKE, Fl. Br. Ind. 6 (1893) 622; Philip. J. Sc. 2 (1907) Bot. 88; Ill. Cyp. (1909) t. 23 f. 5-6; Merr. En. Philip. 1 (1923) 114; RipL. Fl. Mal. Pen. 5 (1925) 148. — Mariscus philippensis StEuD. Syn. 2 (1855) 66; Mia. FI. Ind. Bat. 3 (1856) 290; CLARKE, Philip. J. Sc. 2 (1907) Bot. 88. — Mariscus concinnus SCHRADER ex NEES, Fl. Bras. 2, 1 (1843) 47; Mia. FI. Ind. Bat. 3 (1856) 289. — C. umbellatus BENTH. Fl. Hongkong (1861) 386; Fl. Austr. 7 (1878) 289; F.v.M. Descr. Not. 7 (1886) 34; K. Sco. & Hovrr. FI. Kais. Wilh. Land (1889) 24; ScHum. in Warb. Bot. Jahrb. 12 (1891) 264, non Burm. f. 1768, nec Rox. 1820. — C. cylindrostachys BoECcK. Linnaea 36 (1870) 383, p.p.; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 151, t. 6 f. 6; Koorp. Exk. Fl. Java 1 (1911) 187, non ibid. 4 (1922) f. 204. — C. umbellatus var. cylindrostachys CLARKE, J. Linn. Soc. Bot. 21 (1884) 201. — C. rheedii F.v.M. Descr. Not. 7 (1886) 34. — C. incompletus (non LINK) O. K. Rev. Gen. Pl. 2 (1891) 749. — C. sieberianus SCHUM. in Engler, Pflanzenw. O. Afr. C (1895) 122; K. Scu. & Laut. FI. Schutzgeb. (1900) 193, p.p. — Mariscus cyperoides UrB. Symb. Ant. 2, 1 (1900) 164; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 47. — ? Mariscus rechingeri PaLLA, Denkschr. K. Ak. Wiss. M.-N. K1. Wien 89 (1914) 499, ex descr. — Mariscus microcephalus var. pauciflorus PreEIFF. Mitt. Inst. Allg. Bot. Hamb. 7 (1928) 167, f. 1.— C. compactus var. pauciflorus KUK. Pfl. R. Heft 101 (1936) 424; cf. KERN, Rein- wardtia 2 (1952) 126. Perennial with very short rhizome; stolons absent. Stems tufted, rigid, triquetrous, smooth, incrassate at the base which is clothed with the remains of old leaf-sheaths, 20-75 cm by 1-3 mm. Leaves rather weak, flat, gradually long-attenuate, scabrid in the upper part, 3-6 mm wide; lower sheaths reddish brown. Inflorescence simple, rarely subcompound. Involucral bracts 5—10, patent, the lower ones much overtopping the inflorescence, up to 30cm. Rays 5-17, slender, obliquely patent, straight, smooth, up to 6(—10) cm. Spikes exactly cylindrical when in fruit, obtuse, very dense, with numerous spikelets, 2-4cm by 6-10mm. Spikelets at first obliquely erect, finally at right angles to the rachis and the lower ones often reflexed, linear, acute, subterete, maturing (1—)2(—3) nuts (upper flower often 3), falling off as a whole, 3/4—4 by 4-1 mm; rachilla straight, broadly winged; wings hyaline, tightly clasping the nut, c. ’%mm wide; internodes 1’4-2mm. Glumes membranous, appressed, oblong-ovate, rather acute, not keeled, muticous, 14 imbricate, with 3-nerved. green keel and stramineous sides, often ferrugineous lineolate, many-nerved, 3-3/4 by c. 1 mm. Stamens 3: anthers oblong-linear, °4—-1 mm. Style 4-24 mm; stigmas 3, longer than the style. Nut trigonous, linear, slightly curved, apiculate, rufous to casta- neous, 174-244 by 4 mm. Distr. Widely spread in the tropics and subtropics of Africa, Asia, and Australia; in the New World only in the West Indies, probably introduced; throughout Malesia. Ecol. Open or lightly shaded grassland, old clearings, secondary forests, road-sides, thickets, often a common weed in gardens, 0—1800 m. Vern. Téki idjem, M, sukét lumbungan, wlingin, J, djukut bébaléan, lilisungan, S, kumis, mota, muta, Md, /élassun, Atjeh, padang sila, Asahan, rumput plintang dapur, Banka, tétémung, S. Sumatra, mén- darong ékor tupai, Johore, bandang, kujuhu, kuriup sangan, Borneo, wawitiken, Ceram; New Guinea: etso, Mendi, simboru, Orokaiva, tidor, Enga; Philip.: kupiupi, mangilang, Sub., okokiang, Bon. Notes. It seems superfluous to distinguish nomen- claturally such states as var. subcompositus (CLARKE) Kx. Pfl. R. Heft 101 (1936) 516 [ Mariscus sieberianus var. subcompositus CLARKE, FI. Br. Ind. 6 (1893) 622; C. cylindrostachys f. subcompositus VALCK. SUR. Gesl. Cyp. Mal. Arch. (1898) 153 (t. f. 72)], with some of the spikes bearing a small secondary spike at the base; var. evolutior (CLARKE) KUx. Sinensia 1974] Cy PERACEAE (Kern) 643 3 (1932) 80 (Mariscus siberianus var. evolutior CLARKE. l.c.), a more robust form, and var. microstachys Ktx. Pfl. R. Heft 101 (1936) 518, with narrower spikes and smaller spikelets. KUKENTHAL reduced Mariscus rechingeri PALLA to a variety of C. stenophyllus VALCK. Sur. (it is also cited in the synonymy of C. ferax var. novae- hannoverae). The original material (New Britain, RECHINGER 4235) got lost during World War II: from PALLA’s description I might infer that it be- longed to C. cyperoides. 57. Cyperus paniceus (ROTTB.) BOECK. Linnaea 36 (1870) 381, p.p.; KUk. Pfl. R. Heft 101 (1936) 526, excl. specim. males. — Schaenoides paniceus ROTTB. Deser- Pl rar. Progr. (1772) 15, nom: provis: — Kyllinga panicea Rotts. Descr. & Ic. (1773) 15, t. 4 f. 1. — Mariscus paniceus VAHL, En. 2 (1806) 373; CLaRKE, FI. Br. Ind. 6 (1893) 620; Ill. Cyp. (1909) t. 22 f. 1-2. — Mariscus pullu StEuD. Syn. 2 (1855) 66. — C. umbellatus var. paniceus CLARKE, J. Linn. Soc. Bot. 21 (1884) 201. — Mariscus umbellatus var. paniceus Camus, Fl. Gén. I.-C. 7 (1912) 77. Typical C. paniceus differs from C. cyperoides and C. cyperinus by the following characters: Rhizome emitting very slender, c. 1 mm thick stolons covered with lanceolate, reddish brown striate sheaths. Stems very slender, (3-)5-30cm by c. 4mm. Leaves narrow, 1-2 mm wide. Inflorescence simple. Spikes sessile or subsessile. shortly cylindrical, 8-10 by 4-6 mm. Spikelets finally horizontally spreading. lanceolate, slightly curved, 2/4-3 by c. 4mm, always bearing 1 nut. Glumes 4, lanceolate-ovate. acute, obscurely nerved, reddish lineolate, the third nut-bearing, c. 24 mm, the fourth much reduced, sterile. Style very short, stigmas long, reflexed. Nut small, oblong, c. 1/4 by 14 mm. Distr. From Lower Bengal and Ceylon to Cochinchina. Not known from Malesia; the records in KUKENTHAL’s monograph for Java refer to C. cyperinus f. curvatus (see there), that for Luzon to C. cyperoides. In Malesia only: var. roxburghianus (CLARKE) KUx. Pfl. R. Heft 101 (1936) 526; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 474. — Mariscus paniceus var.roxburghianus CLARKE, Fl. Br. Ind. 6 (1893) 621. More robust, with thicker, shorter stolons, and stouter stems up to 60cm by 1-2 mm. Leaves 2-3 mm wide. Involucral bracts up to 25 cm long. Rays of the inflorescence up to 4cm. Spikes 8-15 by 6-8 mm. Spikelets 3- 44 by “mm. Nut-bearing glume 3—4 mm. Nut 2-2/4 by %-% mm. Distr. India; in Malesia only in a few localities in W., Central and E. Java, and in Madura. Ecol. In teak-forests, 0-150 m. Note. Possibly more related to C. cyperinus than to C. paniceus. 29. Section Kyllingioides KUNTH, En. 2 (1837) 94. — Mariscus subg. Bulbocaulis CLARKE, FI]. Br. Ind. 6 (1893) 619. — Cyperus sect. Bulbocaules KUxK. Pfl. R. Heft 101 (1936) 538. Type species: C. kyllingaeoides VAHL. 58. Cyperus dubius Rotts. Descr. & Ic. (1773) 20, t. 4 f. 5; Boeck. Linnaea 36 (1870) 336: CLARKE, J. Linn. Soc. Bot. 21 (1884) 197; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 150, t. 6 f. 5; Koorp. Exk. Fl. Java 1 (1911) 187; ibid. 4 (1922) f. 205; Kuk. Pfl. R. Heft 101 (1936) 563; KERN, Reinwardtia 6 (1961) 65; in Back. & Bakh. f. Fl. Java 3 (1968) 473. — C. kyllingaeoides VAHL, En. 2 (1806) 312; Mia. Fl. Ind. Bat. 3 (1856) 286, excl. pl. molucc. — Mariscus dregeanus KUNTH, En. 2 (1837) 120; CLARKE, FI. Br. Ind. 6 (1893) 620; Ill. Cyp. (1909) t. 21 f. 1-6; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 72; Camus, Fl. Gén. I.-C. 7 (1912) 75, f. 10, 1—4; Merr. En. Born. (1921) 57; RiDv. FI. Mal. Pen. 5 (1925) 148. — Mariscus irroratus Nees in Hook. J. Bot. Kew Misc. 6 (1854) 28. — Kyllingia multi- nervia STEUD. Syn. 2 (1855) 317. — Mariscus mariti- mus Miq. Sum. (1861) 600; cf. KERN, Reinwardtia 3 (1954) 59. — C. cruentus(non Rotts.) Boeck. Linnaea 36 (1870) 338, p.p.— C. maritimus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 160, t. 6 f. 13; Koorp. Exk. Fl. Java 1 (1911) 187, non Porr. 1806. — Maris- cus merrillii CLARKE, Philip. J. Sc. 2 (1907) Bot. 87: Merr. En. Philip. 1 (1923) 113. — Mariscus niveus Merk. Philip. J. Sc. 14 (1919) 369; En. Philip. 1 (1923) 113, non Schoenus niveus Murr. 1774. — Mariscus dubius KUK. ex FISCHER in Gamble, FI. Madras 9 (1931) 1644; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 45. — C. merrillii KUk. Pfl. R. Heft 101 (1936) 562. Perennial with short rhizome; stolons wanting. Stems densely tufted, rather slender, triquetrous, smooth, (S5—)15—45cm by 1-2(—3)mm, the base apparently bulbous by the striate, turgid, scarious, pale to brown lower sheaths, finally surrounded by their fibrous remains. Leaves weak, flat, gradually acuminate, slightly scaberulous at the top, (1/4-)2-5 mm wide. Inflorescence contracted into a very dense, lobed head consisting of 1-6 confluent spikes, broadly ovoid, subglobose or suboblong, 1—2 cm long and wide. Involucral bracts 3—5, weak, patent to reflexed, up to 30cm. Spikelets numerous, patent, ovate to oblong-lanceolate, acute, subterete, some- what turgid, oblique at the base, falling off as a whole, (2—)3—6-flowered, 4—7 by (1-)1’4-2!4 mm;; rachilla nearly straight, conspicuously winged: wings lan- ceolate, persistent, whitish hyaline. Glumes sub- coriaceous, broadly ovate to lanceolate, obtusish, muticous, 15—19-nerved over the whole breadth, slightly keeled, with at first whitish, finally stra- mineous to brown sides and hyaline margins, c. 74 imbricate, 3—4 by 1/4-3 mm. Stamens 3; anthers oblong-linear, 1-1/4 mm. Stigmas 3. Nut trigonous, ellipsoid or slightly oblong- ellipsoid, apiculate, castaneous, 174-214 by 4% mm. Distr. Widely distributed in the steppes and 644 FLORA MALESIANA [ ser. I, volar? coastal regions of Africa; from Ceylon and India through Further India to S. China: in Malesia: common on both coasts of the Malay Peninsula and on the sandy beaches of the East Coast of Sumatra; along the shores of the South China Sea (Victory I. near Borneo, Anambas and Natuna Is., Riouw Archipelago, Banka, Billiton, Merak I. in the Sunda Straits); also on the SW. coast of Java (Palabuan Ratu); Philippines (Luzon). See map in Reinwardtia 3 (1954) 61. Ecol. In W. Malesia only near the sea, usually on sandy beaches in fully exposed places, acting as a sand binder; the plant is probably sea-dispersed; see RIDLEY, Disp. (1930) 327. In Luzon on wet banks in ravines and on dry, open, rocky slopes at low altitudes. Vern. Umbut-umbut, Billiton. Notes. Owing to KUNTH’s misinterpretation of RuMPHIUS’s Cyperus longus, Herb. Amb. t. 2 f. 1, the species is also credited to the Moluccas by MIQUEL, VALCKENIER SURINGAR and KUKENTHAL. It does not reach eastern Malesia. MERRILL rightly reduced RUMPHIUS’s figure to Cyperus pedunculatus (= Remirea maritima). BOECKELER’s ‘Philippine’ record (as C. cruentus) was based on CUMING 2372, which was from Malacca; see Fl. Mal. I, 1 (1950) 121. 30. Section Remirea (AuBL.) KoyAMA, Quart. J. Taiwan Mus. 14 (1961) 162.— Remirea AUBL. Hist. Pl. Guian. Frang. 1 (1775) 44. Type species: Remirea maritima AUBL. 59. Cyperus pedunculatus (R.BR.) KERN, Act. Bot. Neerl. 7 (1958) 798, f. 4; KoYAMA, Quat. J. Taiwan Mus. 14 (1961) 190; KERN in Back. & Bakh. f. FI. Java 3 (1968) 473. — C. longus RumpH. Herb. Amb. 6 (1750) 5, t. 2 f. 1.— Remirea maritima AUBL. Hist. Pl. Guian. Frang. 1 (1775) 45, t. 16; KUNTH, En. 2 (1837) 139; Boeck. Linnaea 35 (1868) 435: Naves, Nov. App. (1882) 309; CLARKE, Fl. Br. Ind. 6 (1894) 677; Philip. J. Sc. 2 (1907) Bot. 103; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 99; CLARKE, Ill. Cyp. (1909) t. 102 f. 7-10; Koorpb. Exk. Fl. Java 1 (1911) 201; ibid. 4 (1922) f. 263; Camus, Fl. Gén. I.-C. 7 (1912) 155; BACK. Trop. Natuur 8 (1919) 8, f. 15; Me_rr. En. Born. (1921) 63; En. Philip. 1 (1923) 130; RIDL. Fl. Mal. Pen. 5 (1925) 169; HEYNE, Nutt. Pl. 1 (1927) 312: Ounwi, Bot. Mag. Tokyo 56 (1942) 209; KUx. in Fedde, Rep. 53 (1944) 206; S. T. BLAKE, J. Arn. Arb. 29 (1948) 100; Urrr. in Back. Bekn. F1. Java (em. ed.) 10 (1949) fam. 246, p. 48; non C. maritimus Por. 1806. — Remirea pedunculata R.BR. Prod. (1810) 236. — Remirea wightiana NEES in Wight, Contr. (1834) 92; in Hook. J. Bot. Kew Misc. 6 (1854) 29. — Mariscus capitatus STEuD. [in Zoll. Syst. Verz. 1 (1854) 63, nom. nud.] Syn. 2 (1855) 60: Mia. FI. Ind. Bat. 3 (1856) 288. — Mariscus pungens STEUD. Syn. 2 (1855) 60; Mia. FI. Ind. Bat. 3 (1856) 288. — Remirea rigidissima StTEuD. Syn. 2 (1855) 317. — Lipocarpha foliosa Mia. F1. Ind. Bat. 3 (1856) 332; Sum. (1861) 262.— Remirea distichophylla Boeck. Flora 41 (1858) 410. — Remirea maritima var. pedunculata BENTH. Fl. Austr. 7 (1878) 347; K. Sco. & Laut. Fl. Schutzgeb. (1900) 198; VALCK. Sur. Nova Guinea 8 (1912) 708; KUxK. Bot. Jahrb. 59 (1924) 53; Preirr. in Fedde, Rep. 29 (1931) 184; KUK. in Fedde, Rep. 53 (1944) 207. — C. capitatus (non Retz.) NaAves, Nov. App. (1882) 306. — Duvaljouvea maritima PALLA in Rech. Denkschr. K. Ak. Wiss. M.-N. KI. Wien 89 (1913) 500. — Fig. 65. Rhizome horizontally long-creeping, branched, rooting at the nodes, clothed with membranous, acute, brownish sheaths, 1—3 mm thick, the inter- nodes 3-—6cm long. Stems distant or somewhat tufted from the branched head of the rhizome, rigid, trigonous, smooth, 3-12cm by 1-2 mm, in the Malesian specimens usually well exserted from the leaves. Leaves crowded, rigid, canaliculate, often recurved, scabrid on the margins in the upper part, 4-5 mm wide at the base, very gradually narrowed into the triquetrous pungent top. Inflorescence head-like, consisting of some digitately arranged, sessile, short spikes. Involucral bracts 3—S(-8), patent to somewhat reflexed, the longest up to 8 cm, much overtopping the inflorescence. Spikes ovoid or ellipsoid, 8—15 by 7-10 mm. Spikelets sessile, densely crowded, ovoid, acute, slightly compressed, 1- flowered, falling off as a whole, 4—5 by 1)4-2 mm. Rachilla disarticulating above the basal 1—2 glumes; uppermost internode strongly flattened, similar to the glumes but nerveless, finally much thickened, corky, c. 3 mm long. Glumes broadly ovate, hardly keeled, many-nerved, acute or minutely apiculate, 3 lower ones empty, 2—3 mm long, the 4th flower- bearing, 3!4-414 mm long, the uppermost vestigial. Stamens 3; anthers linear, yellow, with short, conical appendage of the connective, c. 2mm long. Nut trigonous, oblong, slightly compressed, shining, castaneous to blackish, tightly enclosed in the upper internode of the rachilla, c. 24 by 34 mm. Distr. Pantropical; throughout Malesia. Map by MikGE & BoDARD, Bull. Inst. Franc. Afr. Noire A, 23 (1961) 704. Ecol. On sandy sea-shores and in sandy dunes near the coast, locally often abundant. Fig. 65. Taxon. The homology of the organ tightly clasping the nut has often been discussed: ROBERT BROWN (1810) took it for the incrassate flower- bearing glume. He was followed by nearly all later authors. KUNTH (1837) disagreed in assuming it to represent the thickened upper internode of the rachilla, but his undoubtedly correct interpretation was only sustained as late as 1922 (CHERMEZON, Bull. Soc. Bot. France 69, 1922, 812), whilst so recently as 1944 KUKENTHAL stuck to the interpretation as a but slightly transformed glume hardly different from the other glumes. From the facts mentioned below it can only be concluded that the nut is enclosed in the upper (or rather penultimate) rachilla-internode, and that the species does not belong in Rhynchosporeae, but in Cyperus as circumscribed in the present treatment. Cy PERACEAE (Kern) 645 Fig. 65. Cyperus pedunculatus (R.BR.) KERN, a characteristic, common, creeping and rooting sedge of the sandy beach, taken from above. Madura I. near East Java (photogr. JEswiet). a) The stamens are placed between the 4th glume and the ovary, not between the corky organ and the ovary; as a rule stamens in Cyperaceae are placed between the flower-bearing glume and the ovary. b) The glumes are all many-nerved, the corky organ is nerveless. c) The 4th glume is distinctly longer than the corky organ; in Cyperaceae the flower-bearing glumes are always the largest. The corky organ bears a cucullate appendage with in its axil a short but distinct continuation of the axis; this appendage must be the Sth, vestigial glume. e) The nut is flattened against the corky organ; trigonous nuts in Cyperaceae are always flattened against the rachilla, having an edge next the subtending glume. Anatomical and embryological characters corrob- d — orate exclusion from Rhynchosporeae; see VAN DER VEKEN, Bull. Jard. Bot. Brux. 35 (1965) 296. Notes. The fruits, completely hidden by the corky rachilla-internode, which is moreover wrapped in the upper glumes, are buoyant and easily dispersed by the sea. See CHERMEZON, Bull. Soc. Bot. France 71 (1924) 854, and RIDLEY, Disp. (1930) 328. The rhizome is fragrant; its use as an aromatic was mentioned by RUMPHIUS. In American and African specimens the stems are usually leafy almost to the top, in Asiatic and Australian ones they are mostly well exserted from the leaves. On this ground R. BRowN distinguished the Australian plants as Remirea pedunculata. KUKENTHAL, following BENTHAM, reduced BROWN’s species to varietal rank. As there is every gradation from included to exserted stems and no differences in spikelets, fruits, etc. exist, nomenclatural recogni- tion seems superfluous. 31. Section Diclidium (NegES) BENTH. FI. Austr. 7 (1878) 286. — Torulinium Desv. in Hamilt. Prod. Pl. Ind. Occ. (1825) 15. — Diclidium SCHRADER ex NEES, FI. Bras. II, 1 (1842) 51. — Cyperus sect. Feraces Kx. Pfl. R. Heft 101 (1936) 614. Types pecres? Cy ferax ib CARien. 60. Cyperus odoratus LINNE, Sp. Pl. 1 (1753) 46; VAHL, En. 2 (1806) 356: Boeck. Linnaea 36 (1870) 407: Danby in Exell, Cat. Vasc. Pl. S. Thomé (1944) 360; KERN in Back. & Bakh. f. Fl. Java 3 646 FLORA MALESIANA [ser. I, vol. 7° (1968) 473. — C. ferax L. C. Ricu. Act. Soc. Hist. Nat. Paris 1 (1792) 106; Boeck. Linnaea 36 (1870) 399: BENTH. FI. Austr. 7 (1878) 286; CLARKE, J. Linn. Soc. Bot. 21 (1884) 191; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 142, t. 6 f. 1; Koorp. Exk. Fl. Java 1 (1911) 186; ibid. 4 (1922) f. 201; Back. Onkr. Suiker. (1928) 141, t. 140; KUx. Pfl. R. Heft 101 (1936) 615, f. 6, K-P: S. T. BLAKE, J. Arn. Arb. 28 (1947) 226. — Torulinium ferax HAMILT. Prod. Pl. Ind. Occ. (1825) 15 (‘ferox’); MERR. En. Philip. 1 (1923) 114; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 48. — Torulinium confertum DESv. ex HamMILtT. Prod. Pl. Ind. Occ. (1825) 15; USsTERI, Beitr. Kenntn. Philip. Veg. (1905) 131; CLARKE, Philip. J. Sc. 2 (1907) Bot. 89; Ill. Cyp. (1909) t. 31 f. 1-4. — C. haenkei Presi, Rel. Haenk. 1 (1828) 172. — Diclidium elatum Nees in Hook. J. Bot. Kew Misc. 6 (1854) 27, non C. elatus L.—C. bracteolatus STEUD. Syn. 2 (1855) 49; Mia. FI. Ind. Bat. 3 (1856) 286. — C. nitidulus Boeck. Linnaea 36 (1870) 363; Scuerr. Nat. Tijd. N. I. 34 (1874) 51.— C. novae-hannoverae Boeck. Bot. Jahrb. 5 (1884) 91; Vatck. Sur. Nova Guinea 8 (1912) 701.— Mariscus ferax CLARKE, Fl. Br. Ind. 6 (1893) 624; RipL. J. Str. Br. R. As. Soc. 46 (1906) 223. — C. michauxianus (non ToRR.) VALCK. SUR. Gesl. Cyp. Mal. Arch. (1898) 143, t. 6 f. 2; Koorp. Exk. Fl. Java 1 (1911) 186. — C. ferax var. novae- hannoverae KU«K. Bot. Jahrb. 59 (1924) 46; Pfl. R. Heft 101 (1936) 618.— Mariscus sieberianus var. evolutior (non CLARKE) RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 73; Fl. Mal. Pen. 5 (1925) 149. Annual with fibrous roots (perennial under favour- able circumstances?). Stems usually stout, solitary or 2—3 together, trigonous, triquetrous in the upper part, smooth, 20-100 cm by up to 6mm, leafy up to 30cm above the incrassate base. Leaves sub- coriaceous, flat, shortly acuminate, scabrid on the margins, up to 12 mm wide. Inflorescence compound or decompound, rather loose to dense, 5—25 cm across. Involucral bracts 6—8, spreading, the larger ones far overtopping the inflorescence, up to 50 cm. Primary rays 7-12, obliquely patent, smooth, up to 20 cm; cladoprophylls usually with a short leafy blade. Spikes ovoid to oblong, 114-3cm wide; rachis narrowly winged, with 20—30(—60) spikelets. Spikelets spicately arranged, horizontally spreading to reflexed, subterete, linear, somewhat flexuous, 4-20-flowered, 5-25 by 1-1/4 mm, when mature breaking up into segments each containing 1 nut; rachilla flexuous, broadly winged; wings elliptic, at first hyaline, finally much thickened, corky, tightly clasping the nut; internodes 1-1/4 mm. Glumes rigid, chartaceous, at first appressed, finally with patulous top, broadly ovate or elliptic, obtuse, muticous or minutely apiculate, rounded on the back, rather distant, up to /4 imbricate (rarely the tip not reaching the base of the next glume above), with green, 5—9-nerved back, yellowish, reddish brown striate sides, and hyaline margins, 2-3/4 mm long. Stamens 3; anthers oblong-linear, 74-1 mm. Stigmas 3. Nut falling off with an internode of the rachilla and the next higher glume, held by the persistent wings, trigonous, oblong, or narrowly obovoid, slightly excurved, apiculate, greyish brown to blackish, 1/4-2 by 4-24 mm. Distr. Widely spread, in the warm regions of the whole world; throughout Malesia, but apparently nowhere common. Ecol. In marshes, wet rice-fields, along river- banks, usually at low altitudes, 0-200 m; a few records from 1900-2100 m: Arfak Mts; hot spring Kokoh Puti on Lombok I. Use. In Celebes used for making little mats. Vern. Rorisan, Sum., pédu pappa, Sulu Is., ratrieé, Tanimbar; New Guinea: omureh, Matapaili lang., borrongor, Biak; Philip.: biliran, S. L. Bis., pulakgalau, Sub. Note. Polymorphic. C. novae-hannoverae BOECK.., with few-flowered spikelets and narrow, dense spikes, cannot be separated satisfactorily from typical C. odoratus. In MAyR 230 from New Guinea, Arfak Mts, 1900-2100 m, the tips of the glumes do not reach the bases of those above, thus exposing a part of the rachilla, a character also found in the N. American C. engelmannii STEUD. II. Subgenus Pycreus (BEAUV.) Mig. FI. Ind. Bat. 3 (1856) 254. — Pycreus BEAUv. Pl. Oware et Benin 2 (1807) 48. Type species: C. polystachyos ROTTB. 32. Section Vestiti (CLARKE) KERN, comb. nov. — Pycreus sect. Vestiti CLARKE, Kew Bull. add. ser. 8 (1908) 94. — Cyperus sect. Sulcati KUK. Pfl. R. Heft 101 (1936) 379. Type species: C. sanguinolentus VAHL. 61. Cyperus sanguinolentus VAHL, En. 2 (1806) 351; KUk. Pfl. R. Heft 101 (1936) 385; KERN, Reinwardtia 3 (1954) 51; in Back. & Bakh. f. Fl. Java 3 (1968) 471. — C. eragrostis (non LAMK) VAHL, En. 2 (1806) 322; Mor. Syst. Verz. (1846) 96; Mia. Fl. Ind. Bat. 3 (1856) 256, incl. var. sanguinolentus MiQ.; BOECK. Linnaea 35 (1868) 443; Naves, Nov. App. (1882) 300; CLARKE, J. Linn. Soc. Bot. 21 (1884) 57; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 64, incl. f. sanguinolentus VALCK. Sur. et f. comosus VALCK. Sur., t. 2 f. 15, 18; Koorp. Exk. Fl. Java 1 (1911) 186, f. 15B; ibid. 4 (1922) f. 199; BAck. Onkr. Suiker. (1928) 130, t. 122. — Pycreus sanguinolentus NEES, Linnaea 9 (1834) 283; CLARKE, FI. Br. Ind. 6 (1893) 1974] CYPERACEAE (Kern) 647 590; Philip. J. Sc. 2 (1907) Bot. 79; RipL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 60; Camus, Fl. Gén. I.-C. 7 (1912) 30; Rm. Fl. Mal. Pen. 5 (1925) 139; Back. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 40. — C. concolor STEUD. Syn. 2 (1855) 6. — C. atratus StEuD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 12; Mig. Fl. Ind. Bat. 3 (1856) 259; Boeck. Linnaea 35 (1868) 446. — Pycreus eragrostis PALLA, Ann. Naturh. Hofm. 23 (1909) 204; Merr. En. Philip. 1 (1923) 110.— Pycreus latespicatus var. fagineicola CAMus, Not. Syst. 1 (1910) 240; Fl. Gén. I.-C. 7 (1912) 30. Annual, or perennial with short rhizome without stolons. Stems slender, tufted, decumbent at the base, trigonous, smooth, 1—4-noded at the base, rooting at the nodes, up to 4 clothed with stramineous to reddish brown sheaths, 5-60 cm by )4-2 mm. Leaves flat or canaliculate, gradually acuminate, scabrid on the margins in the upper part, 1—3(—4) mm wide. Inflorescence simple, either reduced to a cluster of few spikelets, or contracted into a dense head, or with 1—5 up to 5 cm long rays. Involucral bracts 2-5, obliquely erect to patent, the longer ones overtopping the inflorescence, the lowest sometimes erect, up to 15cm. Spikes ovoid to broadly ovoid, with 3-18 spikelets: rachis very short. Spikelets divergent, ovate to oblong-lanceolate, compressed, 8—26-flowered, 14-1 cm by 2—3(—3)4) mm; rachilla straight, wingless, persistent; internodes 74-3 mm. Glumes membranous, obliquely erect or finally patulous, ovate, keeled, obtuse, muticous, )4 imbri- cate, 114-274 by 1/4-2 mm; keel arcuate, green, 3—5-nerved: sides nerveless, whitish, ferrugineous, brownish, or almost black, usually with a longi- tudinal depression in the centre. Stamens 2 or 3 (the number constant in the same specimen); anthers oblong-linear, 14-1 mm. Style rather long; stigmas 2. Nut biconvex, laterally compressed, broadly obovate to orbicular, apiculate, brown to blackish, YLY by %-1){ mm; epidermal cells isodiametric. Distr. Widely distributed in the warmer parts of the eastern hemisphere; tropical Africa; from the Black Sea through C. and E. Asia to Japan and Australia; throughout Malesia, often common. Ecol. In wet open places: swamps, grassy fields: rice-fields, etc., 0—3125 m. KEY TO THE MALESIAN SUBSPECIES 1. Stamens 3. Leaves flat, weak, 2-4mm wide. Inflorescence capitate or nearly so, rarely the rays up to 3 cm. Involucral bracts patent. Spikelets (24-)24-3-3) mm wide. Glumes (1°4—)2- 2)4 mm long, distinctly depressed on either side, bordered by a well-marked sanguineous band. Nut medium-sized, (1—-)114-1)4 by %-1 mm. a. ssp. sanguinolentus 1. Stamens 2. 2. Nut relatively large, 14-14 by %o-1)4 mm. Stems very slender, often short. Leaves narrow, 1-1/4(-2/4) mm wide. Involucral bracts 2-3, the lowest usually erect or obliquely erect. Inflorescence capitate, with 3—9(—15) spikelets. Spikelets 234-314 mm wide. Glumes 2—2)4 mm long, without depressions, dark castaneous to almost black except fcr the pale keel and the narrow but distinct whitish hyaline margins, more rarely brown to pale brown. b. ssp. melanocephalus 2. Nut smaller, 44-1 by %4-% mm. Leaves flat, 2-4 mm wide. Involucral bracts spreading. In- florescence more or less open, or, when capitate, with many spikelets. Glumes 1/42 mm long, ferrugineous to whitish, often tinged with red. 3. Inflorescence more or less open, its rays up to 5cm. Glumes with a distinct depression on either side, ferrugineous, more or less tinged with red. Spikelets 2/4-2)4 mm wide. c. ssp. cyrtostachys 3. Spikelets densely crowded into a multispiculate head; rarely one of the rays somewhat elongated, up to 1 cm. Glumes without or with indistinct depressions, yellowish white, rarely somewhat ferrugineous or slightly tinged with red, appressed, hence spikelets only c. 2 mm wide. d. ssp. teysmannii a. ssp. sanguinolentus. — Synonymy see above. Distr. India, Farther India, Indo-China, China, Japan; in Malesia: Sumatra, Malay Peninsula, Java, Lesser Sunda Is.(Lombok, Flores), Celebes, Moluccas (Ambon), Philippines (Luzon). Ecol. Grassy places, swamps, margins of pools, etc., often in incultivated localities, more rarely in wet rice-fields, 800—2000 m, sometimes lower. b. ssp. melanocephalus (MiQ.) KERN, Reinwardtia 3 (1954) 55, f. 9. — C. melanocephalus Mia. FI. Ind. Bat. 3 (1856) 259. — C. eragrostis var. humilis Mia. l.c. 257. — C. eragrostis f. melanocephalus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 66. — C. eragrostis RUM TISSVIAV. CK SURE Gg09 iit mi ete sanguinolentus f. melanocephalus KUK. Pfl. R. Heft 101 (1936) 387. — C. sanguinolentus f. humilis KUK. l.c. 386.— ? C. globosus f. atrosanguineus KUK. Candollea 6 (1936) 422, ex descr. — C. globosus var. latisquamatus KUK. Mitt. Thiir. Bot. Ver. 50 (1943) 7. Distr. India (Khasia Mts, Sikkim, E. Himalaya), in Malesia: on several mountains in Java, the Philippines, and New Guinea. Ecol. In swampy grassy fields, etc., at high altitudes, from 1000 m upwards. Vern. New Guinea: Maitjop, Hattam lang., boh, Kapauku lang. c. ssp. cyrtostachys (MIQ.) KERN, Reinwardtia 3 (1954) 57. — C. eragrostis var. cyrtostachys Mia. F1. Ind. Bat. 3 (1856) 257; Boeck. Linnaea 35 (1868) 445 (‘cyrtolepis’); CLARKE, J. Linn. Soc. Bot. 21 (1884) 59, p.p. — C. eragrostis f. cyrtostachys VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 66, t. 2 f. 16. — C. sanguinolentus f. cyrtostachys KUx. Pfl. R. Heft 101 (1936) 387. Distr. Throughout Malesia. d. ssp. teysmannii (BOECK.) KERN, Reinwardtia 3 (1954) 57, f. 10. — C. teysmannii Boeck. Flora 58 (1875) 259. — C. eragrostis f. teysmannii VALCK. SUR. Gesl. Cyp: Mal. Arch. (1898) 67; t. 2 f. 17.— C. sanguinolentus var. teysmannii KUK. Pfl. R. Heft 101 (1936) 387. Distr. Malesia: Sumatra (Lampongs), Java and Madura, Lesser Sunda Is. (Bali), Philippines (Luzon). Ecol. Watersides, swampy places, wet rice-fields, 0-750 m. 648 FLORA MALESIANA (ser. 1, your 33. Section Chrysanthi CLARKE, Kew Bull. add. ser. 8 (1908) 95. — Cyperus sect. Lancei KUx. Pfl. R. Heft 101 (1936) 330. Type species: C. chrysanthus BOECK. 62. Cyperus unioloides R.Br. Prod. (1810) 216; BENTH. FI. Austr. 7 (1878) 260; CLARKE, J. Linn. Soc. Bot. 21 (1884) 60; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 71; KUx. Pfl. R. Heft 101 (1936) 338, f. 2B, 4 E-G; Kern, Reinwardtia 2 (1952) 124, f. 12; in Back. & Bakh. f. Fl. Java 3 (1968) 471.— C. angulatus NEES in Wight, Contr. (1834) 73; BoECK. Linnaea 35 (1868) 465. — Pycreus angulatus NEES, Linnaea 9 (1834) 283; CLARKE, FI. Br. Ind. 6 (1893) 593; Ill. Cyp. (1909) t. 4; Camus, Fl. Gen. I.-C. 7 (1912) 35. — Pycreus unioloides URB. Symb. Ant. 2 (1900) 164; Merr. En. Philip. 1 (1923) 111. Perennial (always?) with short rhizome; stolons wanting. Stems slender, somewhat tufted, rigid, triquetrous, smooth or scaberulous at the top, up to 90cm by 1-2 mm. Leaves flat to conduplicate, rigid, very gradually acuminate, with cartilaginous margins scabrid in the upper part, 2-4 mm wide; lower sheaths reddish to blackish brown. Inflorescence simple, loose to contracted, 3-7 by 6—9cm. In- volucral bracts 2-4, patent, the longest much overtopping the inflorescence, up to 20cm. Rays 3-6, suberect, smooth, up to 6cm. Spikes broadly ovoid, up to 3 cm long and wide, with 4—12 spikelets; rachis glabrous, up to 1 cm. Spikelets patent, oblong or lanceolate, compressed but slightly turgid, acute, 10—20(—30)-flowered, 8—15(—25) by 4-5 mm; rachilla flexuous, sche persistent; internodes °4— 4% mm. Glumes chartaceous, divergent, ovate to oblong, keeled, acute, muticous, 34 imbricate, 334-4 by 2/4-3 mm; keel acute, 3-nerved; sides nerveless, shining yellowish green to brown; margins not or hardly hyaline, undulate when dry. Stamens 3; anthers linear, up to 2 mm. Style long (1/4-3)4 mm); stigmas 2, up to 3 mm. Nut relatively small, biconvex, laterally compressed, short-elliptic to slightly obovate, shortly apiculate, shining black, 1-114 by 44-1 mm; epidermal cells isodiametric. Distr. Pantropic; in Malesia very local: N. Sumatra (Atjeh, East Coast Res., Tapanuli), W. Java, Philippines (Luzon, Mindanao), New Guinea. Ecol. In open localities: swamps, marshes, borders of lakes, river-banks, usually at medium altitudes (up to 1900 m), in New Guinea up to 2800 m. 34. Section Pycreus Pycreus sect. Polystachyi CLARKE, Kew Bull. add. ser. 8 (1908) 94. — Pycreus sect. Globosi CLARKE, I.c. 95 (‘Globosus’). Type species: C. polystachyos ROTTB. 63. Cyperus flavidus RETzZ. Obs. 5 (1789) 13; DECNE, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 359; Herb. Timor. Descr. (1835) 31. — C. globosus ALL. Fl. Pedem. Auct. (1789) 49, non Forsk. 1775; Boeck. Linnaea 35 (1868) 458; BENTH. Fl. Austr. 7 (1878) 260; CLARKE, J. Linn. Soc. Bot. 21 (1884) 47, incl. var. nilagiricus CLARKE; VALCK. SuR. Ges]. Cyp. Mal. Arch. (1898) 58, t. 2 f. 12, incl. f. strictus VALCK. SuR.; Koorp. Exk. Fl. Java 1 (1911) 186, 192: ibid. 4 (1922) f. 198; Back. Onkr. Suiker. (1928) 128,12 1205 Kuk Phe RR, Heft 101 (1936),352°+S, aL. BLAKE, J. Arn. Arb. 28 (1947) 220; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 471.—C. capillaris KOENIG ex Roxs. FI. Ind. 1 (1820) 198. — C. strictus Roxs. I.c. 203; Mig. Fl. Ind. Bat. 3 (1856) 258. — C. lanceolatus (non Porr.) PRESL, Rel. Haenk. 1 (1828) 167. — Pycreus globosus REICHENB. Fl. Germ. Exc. 2 (1830) 140; CLARKE, Philip. J. Sc. 2 (1907) Bot. 80, incl. var. nilagiricus CLARKE; CAMUS, F1. Gén. I.-C. 7 (1912) 31, f. 3, 10; Merr. En. Philip. 1 (1923) 110; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 41. — Pycreus capillaris Nees, Linnaea 9 (1834) 283; CLARKE, FI. Br. Ind. 6 (1893) 591. — C. vulgaris KUNTH, En. 2 (1837) 4; ? HAssk. Pl. Jav. Rar. (1848) 71; Mia. Fl. Ind. Bat. 3 (1856) 256, incl. var. polylepis MiQ.— C. mucronatus (non Rotts.) Mor. Syst. Verz. (1846) 95. — C. nilagiricus Hocust. ex STEUD. Syn. 2 (1855) 2; Boeck. Linnaea 35 (1868) 457. — C. jungendus StEup [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud.| Syn. 2 (1855) 3. — C. junghuhnii Mia. FI. Ind. Bat. 3 (1856) 260.— C. flavescens (non L.) BENTH. FI. Austr. 7 (1878) 259. — Chlorocyperus nilagiricus RIKL1, Jahrb. Wiss. Bot. 27 (1895) 564. — Chlorocyperus globosus PALLA, Allg. Bot. Zeitschr. 6 (1900) 61.—C. globosus var. oblonginux KUx. Mitt. Thiir. Bot. Ver. 50 (1943) 7. Annual with fibrous, yellowish roots, in favourable circumstances perennial with short rhizome; stolons wanting. Stems rigid, very slender, tufted, trigonous, smooth, 5-60cm by 1-1/4 mm. Leaves rigid, narrow, canaliculate, often almost setaceous, very gradually acuminate. smooth or slightly scabrid at the top, 1—2(—3) mm wide; lower sheaths ferrugi- neous to dark reddish brown. Inflorescence simple or subcompound, rather open to contracted into a single cluster. Involucral bracts 2—4, obliquely erect to patent, the lower 1-2 much overtopping the inflorescence, often seemingly continuing the stem, up to 25cm. Primary rays 3—6, erect or obliquely erect, very slender, smooth, up to 5 cm, secondary ones when present 1—3 mm. Spikes ovoid to broadly ovoid, with 5—20 spikelets; rhachis short, 3-10 mm. Spikelets spreading, the lower ones often somewhat reflexed, linear, exactly parallel-sided, strongly com- pressed, 20—40(—60)-flowered, 1—2(—3)cm by 2- 2/A(-3) mm; rachilla straight, wingless, persistent; 1974] Cy PERACEAE (Kern) 649 internodes 74—)4 mm. Glumes chartaceous, obliquely erect, sharply keeled, oblong-ovate, obtuse, muti- cous, 24 imbricate, 14-24 by 1-1/4 mm; keel green, 3-nerved; sides shining, stramineous, brown, or blackish, nerveless; margins whitish hyaline especially towards the top. Stamens 2; anthers oblong, c. 14mm. Style rather short; stigmas 2, longer than the style. Nut biconvex, laterally com- pressed, oblong-obovate to oblong-elliptic, distinctly apiculate, fuscous to dark castaneous, %-1/% by 24-74 mm; epidermal cells isodiametric. Distr. Widely distributed in the warmer parts of the eastern hemisphere: tropical Africa, from the Mediterranean region through Central and S. Asia to Australia; rather common in Malesia, but still unknown from the Malay Peninsula and Borneo. Ecol. In open, wet places: swamps, grassy fields, watersides, rice-fields, etc., from the lowland (here apparently rare) up to 2100 m, in New Guinea up to 2800 m. Vern. Rumput toyan, J; New Guinea: komaruem, Mendi, tampi, Enga; Philip.: tantanud, Bon. Notes. Often confused with C. sanguinolentus and C. polystachyos. From the former dis*‘nguished by the enodulose stem-base and the shape of the nut, from the latter by the more obtuse glumes, the straight, wingless rachilla, the linear spikelets, and the shape of the nut. Rather polymorphous. C. nilagiricus was mainly founded on the dark colour of the glumes, but CLARKE included all the forms with narrow spikelets whatever the colour. In Malesia the width of the spikelets varies between 2 and 3 mm, but the colour varies independently of the width. KUKENTHAL referred some apparently depauperate specimens from Java (not seen) to f. pauperior (BOECK.) KUK. Pfl. R. Heft 101 (1936) 355 [C. nilagiricus var. pauperior Hocust. ex Boeck. Linnaea 35 (1868) 458]. As to KUKENTHAL’s var. oblonginux I agree with BLAKE /.c. that there exists an intergrading series from oblong-elliptic nuts to the much more usual obovate ones. 64. Cyperus polystachyos Rotts. Descr. PI. rar. Prove. (1772), 21; Descr. & Ic. (1773) 39, t.. 11 Ff. 1; Hassk. Pl. Jav. Rar. (1848) 74; Mia. Fl. Ind. Bat. 3 (1856) 258: Boeck. Linnaea 35 (1868) 477; BENTH. Fl. Austr. 7 (1878) 261; CLARKE, J. Linn. Soc. Bot. 21 (1884) 51, t. 3 f. 27; VaLck. Sur. Gesl. Cyp. Mal. Arch. (1898) 61, t. 2 f. 13; Koorp. Exk. Fl. Java 1 (1911) 186, non ibid. 4 (1922) f. 196; KUx. Pfl. R. Heft 101 (1936) 367; S. T. BLAKE, J. Arn. Arb. 28 (1947) 220: KERN, Reinwardtia 3 (1954) 48; in Back. & Bakh. f. Fl. Java 3 (1968) 472. — C. rotundus floridus II mas RumMpH. Herb. Amb. 6 (1750) 2, t. 1 f. 2. — Pycreus polystachyos BEAUV. Pl. Oware & Benin 2 (1807) 48, t. 86 f. 2; CLARKE, FI. Br. Ind. 6 (1893) 592; Philip. J. Sc. 2 (1907) Bot. 80; RipL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 60; Camus, FI. Gén. I.-C. 7 (1912) 34; Ripv. Fl. Mal. Pen. 5 (1925) 139; BAcK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 40.— C. strigosus (non L.) LLANos, Fragm. PI. Filip. (1851) 16; F.-ViL_. & NAves in Blanco, FI. Filip. ed. 3, 4 (1880) 10. — ? C. teretifructus STEUD. [in Zoll. Syst. Verz. 1 (1854) 62, nom. nud. (‘teretifractus’)| Syn. 2 (1855) 3. — ? C. vulgaris var. teretifructus Mig. Fl. Ind. Bat. 3 (1856) 256. — Chlorocyperus polystachyus RIKI, Jahrb. Wiss. Bot. 27 (1895) 563. — Pycreus odoratus UrB. Symb. Ant. 2 (1900) 164; Merr. En. Philip. 1 (1923) 110, non C. odoratus L.— Pycreus holosericeus MERR. Philip. J. Sc. 12 (1917) 231, quoad specim., non C. holosericeus Link. — C. odoratus (non L.) BACK. Onkr. Suiker. (1928) 129, t. 121. Annual, or perennial with short rhizome; stolons wanting. Stems slender but rather firm, tufted, trigonous, triquetrous just below the inflorescence, smooth, 5—60(—90) cm by 1—2(—3) mm. Leaves weak to rather rigid, flat or canaliculate, gradually acuminate, scabrid on the margins in the upper part, (1—-)2—4 mm wide; lower sheaths membranous, light brown to purplish. Inflorescence simple or sub- compound, strongly contracted (sometimes into a single head) to open, 2-15 cm across. Involucral bracts 3—6, obliquely to widely spreading, the lower one(s) overtopping the inflorescence, the longest up to 20 cm. Primary rays 3—8, spreading, very short to slender, up to 7 cm, secondary ones when present very short. Spikelets 2—15 to the spike, fasciculate to divergent, linear-lanceolate. gradually tapering to an acute apex, strongly compressed, 8—50-flowered. )4- 24% cm by 1/4(—2) mm; rachilla flexuous, narrowly winged, persistent; internodes c. }4 mm. Glumes thinly chartaceous, obliquely erect, sharply keeled. elliptic-ovate, subobtuse, muticous or minutely mucronulate, 14 imbricate, 174-2)4 by c. 1 mm; keel green, 3-nerved ; sides nerveless, pale ferrugineous to fuscous, rarely castaneous; margins whitish hya- line. Stamens 2, very rarely in some or most flowers 1; anthers oblong or linear-oblong, /4—-1 mm. Style long: stigmas 2, about as long as the style. Nut 2-sided, laterally compressed, with flat to slightly convex sides, oblong with nearly parallel margins, somewhat asymmetric, abruptly rounded to truncate at the shortly apiculate apex, castaneous to black, 1-1% by %-)¥ mm; epidermal cells isodiametric. Distr. Widely spread in the warmer parts of the whole world, in the eastern hemisphere extending northward to the Mediterranean (see MEROLA, Delpinoa 10, 1957, 21-92), S. China, and Japan, southward to Australia; throughout Malesia, pre- sumably common everywhere, though only a few times collected in the eastern part. Ecol. In open, usually damp places: grassy fields, fallow rice-fields, road-sides, river-banks, also in the salt mud of the sea-shore, 0—1800 m. Vern. Rumput parah betina, Negri Sembilan, djukut bulu mata kibo, S, kankamut, N. Borneo, aduru, Talaud; Philip.: aluisang-pasigan, Tag., hand, Iv. Notes. Very variable, especially as to the size and density of the inflorescence. The varieties based on this variability have little or no taxonomic value, as there exists every gradation from the single dense cluster of suberect spikelets to the open inflorescence with rectangularly spreading spikelets. Plants with open inflorescences are rather common in Malesia, especially near the sea. They have been described as: var. laxiflorus BENTH. FI. Austr. 7 (1878) 261; CLARKE, J. Linn. Soc. Bot. 21 (1884) 53; VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 62; KUx. Pfi. R. Heft 101 (1936) 370. — C. paniculatus Rotts. Descr. PI. far.,'Progr. :(1772)223))Descr. i& Ices (1773) 402— Pycreus polystachyos var. laxiflorus CLARKE, FI. Br. 650 FLORA MALESIANA [ser. I, vol. 7° Ind. 6 (1893) 592; Philip. J. Sc. 2 (1907) Bot. 80; Camus, Fl. Gen. I.-C. 7 (1912) 35.— Pycreus polystachyos var. paniculatus MeRR. Fl. Manila (1912) 109; En. Philip. 1 (1923) 111. The glumes are usually straw-coloured, but often tinged with red. Specimens with strikingly dark- coloured spikelets were collected in N. Sumatra (Toba Lake) at 900—1350 m. The inadequately labelled specimen upon which f. longispiculatus VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 62, t. 2 f. 13a, was based, belongs to the N. American C. filicinus VAHL. See KERN, Rein- wardtia 3 (1954) 48. The Philippine specimens referred to the American var. leptostachyus BOECK. by KUKENTHAL, Pfl. R. Heft 101 (1936) 371 (as Pycreus holosericeus by MERRILL, I.c.), were misnamed. See KERN, !.c. I have not seen ZOLLINGER 456, type collection of C. teretifructus STEUD. According to ZOLLINGER it was collected in Java, according to STEUDEL in Japan. CLARKE (1884) and VALCKENIER SURINGAR referred it to C. globosus, CLARKE (1893), KUKENTHAL, and the Japanese authors to C. polystachyos. 65. Cyperus sulcinux CLARKE, J. Linn. Soc. Bot. 21 (1884) 56; VaALcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 63, t.2 f. 14, t. 3; KOK. Pfl. R. Heft 101 (1936) 364, f. 43; S. T. BLake, J. Arn. Arb. 28 (1947) 220; KERN, Reinwardtia 3 (1954) 46, f. 7; in Back. & Bakh. f. Fl. Java 3 (1968) 471. — Pycreus sulcinux CLARKE, Fl. Br. Ind. 6 (1893) 593; Philip. J. Sc. 2 (1907) Bot. 80; Camus, Fl. Gén. I.-C. 7 (1912) 34; Merr. En. Born. (1921) 58; En. Philip. 1 (1923) 111; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 40. — Pycreus substellatus CAMuS, Not. Syst. 1 (1910) 240; Fl. Gén. I.-C. 7 (1912) 29. — Pycreus odoratus var. holosericeus MERR. J. Str. Br. R. As. Soc. n. 76 (1917) 79; En. Born. (1921) 58, quoad specim., non C. holosericeus LINK. 7 Annual with fibrous, yellowish roots. Stems slender, tufted, trigonous below, triquetrous at the top, smooth, 5—30(—40) cm by up to 1 mm. Leaves rigid, flat or canaliculate, gradually acuminate, scabrid at the top, narrow, ’4—2 mm wide; lower sheaths purplish. Inflorescence relatively large, simple, loose, rarely subcapitate. Involucral bracts 3-4, obliquely patent, the lower one(s) overtopping the inflorescence, the longest up to 15cm. Rays 3-6, slender, often capillary, spreading, up to 7 cm, sometimes very short. Spikes broadly ovoid, with (3—)5—10(—14) spikelets; rachis glabrous, up to 5mm. Spikelets widely spreading, the lower ones more or less reflexed, lanceolate to linear, usually slightly curved, acute, strongly compressed, 10- 50(—68)-flowered, 1-3(— -444) cm by c. 14% mm; rachilla slightly flexuous, narrowly winged, per- sistent; internodes >%— 3% mm. Glumes membranous, obliquely erect, sharply keeled, obovate to oblong- obovate, obtuse, muticous, )Z—)4 imbricate, 14- 1°4(-2) by 1-1)4(-1/4) mm; keel fuscous, 3-nerved; sides ferrugineous, nerveless; margins hyaline. Stamen 1; anther linear-oblong, 4-74 mm. Style c. 4% mm; stigmas 2, short. Nut 2-sided, laterally compressed, on either side with a median longitudinal depression, oblong, somewhat asymmetric, truncate at the shortly spiculate apex, dark brown, 1/4(-1/4) by 4 mm; epidermal cells isodiametric. Distr. From India through Farther India to tropical Australia (Queensland), in Malesia very local: in a few localities in Java, Kangean Archipelago, Madura, Borneo (Sarawak, North Borneo), the Philippines (Palawan, Luzon, Mindanao), the Moluccas (Talaud Is., Buru, Ternate), Lesser Sunda Is. (W. Sumba), and New Guinea. Ecol. In open places on dry soil: fields, road-sides, etc., 0-1200 m. Note. According to CLARKE’s original description the flowers should be diandrous. I always found only 1 stamen in the numerous flowers dissected. 35. Section Pumili KUK. Pfl. R. Heft 101 (1936) 375. Type species: C. pumilus L 66. Cyperus pumilus LINNE, Cent. Pl. 2 (1756) 6; Amoen. 4 (1759) 302; Sp. Pl. ed. 2 (1762) 69; Mia. Fl. Ind. Bat. 3 (1856) 255; NAaves, Nov. App. (1882) 300; CLARKE, J. Linn. Soc. Bot. 21 (1884) 43, incl. var. punctatus CLARKE quoad pl. asiat. et f. borneensis CLARKE; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 55, t. 2 f. 10; Koorp. Exk. Fl. Java 1 (1911) 186, 192: ibid. 4 (1922) f. 197; Back. Onkr. Suiker. (1928) 128, t. 119; KUx. Pfl. R. Heft 101 (1936) 375, f.44 A-E: Kern, Reinwardtia 3 (1954) 50: in Back. & Bakh. f. Fl. Java 3 (1968) 470. — C. nitens RETz. Obs. 6 (1789) 13: NEEs in Hook. J. Bot. Kew Misc. 6 (1854) 28; Mig. Fl. Ind. Bat. 3 (1856) 255; BoEck. Linnaea 35 (1868) 483; VipAL, Phan. Cum. Philip. (1885) 155; Rev. Pl. Vasc. Filip. (1886) 283. — C. punctatus RoxB. Fl. Ind. 1 (1820) 197. — C. pulvinatus Nees & Mey. ex NEES in Wight, Contr. (1834) 74. — Pycreus pumilus Nees [Linnaea 9 (1835) 283, nomen] ex CLARKE, FI. Br. Ind. 6 (1893) 591, quoad basion., excl. descr.; Domin, Bibl. Bot. Heft 85 (1915) 417; Turritt, Kew Bull. (1922) 124.— Pycreus pulvinatus Nees, Linnaea 9 (1835) 283; CLARKE, Philip. J. Sc. 2 (1907) Bot. 79. — Pycreus nitens NEES, Nov. Act. Nat. Cur. 19, Suppl. 1 (1843) 53; CLARKE, FI. Br. Ind. 6 (1893) 591; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 61; Camus, FI. Gén. I.-C. 7 (1912) 33; Merr. En. Philip. 1 (1923) 110; Rip. Fl. Mal. Pen. 5 (1925) 140; BAck. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 39. —C. gymnoleptus STEUD. Syn. 2 (1855) 3; Mia. Fl. Ind. Bat. 3 (1856) 255. — Dichostylis nitens PALLA, Bot. Jahrb. 10 (1888) 296. — Pycreus hyalinus (non DoMtn) Merr. En. Born. (1921) 57.—C. silletensis (non NEES) ELMER, Leafl. Philip. Bot. 10 (1938) 3530. — Fig. 66. Annual with fibrous, yellowish roots. Stems slender, tufted, often almost filiform, triquetrous, smooth, 1-20cm by }4-1 mm. Leaves weak or somewhat rigid, canaliculate, gradually acuminate, scabrid in the upper part, 1-2 mm wide; lower 1974] Cy PERACEAE (Kern) 651 Fig. 66. Cyperus pumilus L. a. Habit, x 74, b. spikelet, x 6, c. rachilla, d. glumes, e. nut with stamen and style, all x 14 (SCHIFFNER Iter jav. 1596). sheaths stramineous to reddish brown. Inflorescence simple or subcompound. Involucral bracts 3-5, patulous, overtopping the inflorescence, the longest 2)4-15(—20)cm. Primary rays 3—6, divergent, slender, smooth, up to 3 cm, but often very short and then inflorescence head-like. Spikes ovoid or subglobose. dense, with 5—30 spikelets 1—2 cm 9; rachis short, up to 3 mm. Spikelets divergent, oblong to linear, subacute, strongly compressed, 8—30(—40)-flowered, 5-10(-13) by 1)4-2)4 mm (mucros_ included): rachilla straight, almost wingless, persistent; inter- nodes 14-'4 mm. Glumes membranous, obliquely patent, sharply keeled, ovate, 14 imbricate, the body (1-)1)4-13% by 1-114 mm; apex truncate or retuse: keel 3—S-nerved, green or reddish brown; sides nerveless, silvery, stramineous, or spadiceous; mucro erect or slightly recurved, /Z—)4 mm. Stamen usually 1, but not rarely in some or all flowers stamens 2: anthers oblong, c. 4 mm. Stigmas 2. Nut biconvex, laterally compressed, oblong-obovate, obtuse to truncate at the apiculate apex. shining brown to castaneous. 14-24 by 4-4 mm; epidermal cells isodiametric. Distr. India, Farther India, Formosa, S. China. Queensland (here known from 2 localities only): in Malesia: Sumatra (with certainty only known from Medan); in the Malay Peninsula and Java very local. but often abundant; in the Philippines from N. Luzon to Mindanao, in most or all islands; but a few times collected in the Lesser Sunda Is. (Flores), Borneo, Celebes, and the Moluccas (Ambon, Buru). Varieties in tropical America and Africa. Ecol. In open, damp to rather dry localities: fields, moist meadows, humid flats, etc.. often on sandy soil, 0-600 m. Note. According to Koorpers, /.c., in Java common in rice-fields and grassy localities in the plains and the mountains (quoted in KUKENTHAL’s monograph). However, it is neither common in Java nor does it occur in the mountains. 67. Cyperus nervulosus (KUK.) S. T. BLAKE, Proc. R. Soc. Queensl. 51 (1940) 41: J. Arn. Arb. 28 (1947) 221; KERN, Reinwardtia 3 (1954) 51. — C. breviculmis (non R.BR.) F.v.M. Fragm. 8 (1874) 267.—C. pumilus (non L.) BENTH. Fl. Austr. 7 (1878) 258; VALCK. SuR. Nova Guinea 8 (1912) 696. non al. — C. pumilus var. punctatus CLARKE, J. Linn. Soc. Bot. 21 (1884) 46, quoad pl. austr., non C. punctatus ROxB. — Pycreus pumilus var. punctatus Domin, Bibl. Bot. Heft 85 (1915) 417.—C. pumilus var. nervulosus KUkK. Pfl. R. Heft 101 (1936) 378. — Fig. 67. Closely allied to C. pumilus. Very slender. Spikes rather loose, with S—7(—10) spikelets. Spikelets linear, up to 44-flowered, (5—)10—18 by 2—3 mm. Glumes obovate-spatulate, widely patent. with dis- tinctly 2—3-nerved sides; mucro recurved, 4-34 mm. Stamens 2. Distr. N. Australia, NE. Queensland, in Malesia: New Guinea (south coast of W. New Guinea: Papua, Central Div.). Ecol. Damp savannah flats, dune hollows, 0—30 m. Note. As for the shape of the spikelets and glumes similar to C. squarrosus, which, however, is trigynous and has trigonous nuts. 36. Section Flavescentes KUK. Pfl. R. Heft 101 (1936) 395. — Cyperus sect. Latespicati KUK. l.c. 388. Type species: C. flavescens L. 68. Cyperus diaphanus SCHRADER ex R. & S. Mant. 2 (1824) 477; Bogck. Linnaea 35 (1868) 437; KERN, Blumea 10 (1960) 644. var. diaphanus. Only known from Nepal. Glumes very pale, sordidly white. Otherwise hardly different from the 652 FLORA MALESIANA [ser. I, yorker Fig. 67. Cyperus nervulosus (KUK.) S. T. BLAKE. a. Habit, nat. size, b. spikelet in fruit, x 10, c—d. glumes, x 20, e.rachilla, x 20, f. anther, x 50, g. nut, x 20 (a—g Brass 3731). 1974] CYPERACEAE (Kern) 653 common Malesian form, which may be referred to as: var. latespicatus (BOECK.) KERN, comb. nov. — C. latespicatus BoEcK. Flora 42 (1859) 441 (*433°); Linnaea 35 (1868) 467; CLARKE, J. Linn. Soc. Bot. 21 (1884) 40; VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 54; KUx. Pfl. R. Heft 101 (1936) 392; KERN, Reinwardtia 2 (1952) 124, f. 13; ibid. 3 (1954) 66. — Pycreus latespicatus CLARKE, Fl. Br. Ind. 6 (1893) 590; Ill. Cyp. (1909) t. 3; Camus, Fl. Gén. I.-C. 7 (1912) 30, excl. var. fagineicola Camus. — Chloro- cyperus latispicatus RIKLI, Jahrb. Wiss. Bot. 27 (1895) 564. Annual. Stems tufted, rigid, trigonous, smooth, 10-40 cm by 1-2 mm. Leaves flat, gradually narrowed upwards, 1-2/4 mm wide, the margins minutely scabrid in the upper part. Inflorescence simple, often contracted, sometimes with 2—3(—5) rays up to 7 cm long, or reduced to a single head. Involucral bracts 2—3(—4), patent, the lowest up to 15cm long. Spikes broadly ovate; rachis short, up to 5mm. Spikelets finally horizontally spreading, approximate, oblong, with nearly parallel margins, strongly compressed, subobtuse, !—2 cm by (2)4-)3— 4mm, 20—30(—40)-flowered; rachilla slightly flex- uous, wingless, persistent; internodes c. )4 mm. Glumes membranous, obliquely patent, keeled, ovate, obtuse, muticous, c. vA imbricate, (2/4-)3 by 134-2 mm; keel curved, green, 3(—5)-nerved; sides nerveless, shining fulvous to castaneous; margins somewhat undulate. Stamens 2; anthers oblong to oblong-linear, 44-1 mm. Style up to 114 mm; stigmas 2, about as long as the style. Nut biconvex, laterally compressed, obovate to broadly obovate or elliptic, broadly stipitate, apiculate, 1-14 by *%-1 mn, rugulose with transverse wavy lines, ultimately black; epidermal cells longitudinally oblong. Distr. From Central Asia to India and Farther India, in Malesia very rare: Sumatra (Res. Tapianuli and West Coast), Celebes (near Malino), Philippines (Luzon). Ecol. In grassy wildernesses and wet rice-fields, 300-1500 m. Notes. Strikingly resembling C. unioloides, but readily distinguished by the 2 stamens (3 in C. unioloides) and the rugulose nut. Taxonomically more important is probably: ssp. setiformis (KORSH.) KERN, comb. nov. — C. setiformis KorsH. Act. Hort. Petrop. 12 (1892) 405; KUK. Pfl. R. Heft 101 (1936) 393; OHwi, Mem. Coll. Sc. Kyoto Imp. Un. B 18 (1944) 158. — Pycreus setiformis B. ScHIsK. Fl. U.R.S.S. 3 (1935) 5. —C. latespicatus var. setiformis KOYAMA, Act. Phytotax. Geobot. 16 (1955) 11. Stems setaceous, 5—20cm tall. Leaves filiform. Inflorescence consisting of a single head, with 1— 3(—4) sessile spikelets. Bracts 1—2, the lowest erect. Spikelets 244-3 mm _ wide. Glumes rufescent to purplish. Distr. Amurlands, Japan, Korea, Manchuria, Bonin Islands, in Malesia: New Guinea (Western Highlands, Lake Inim, Lake Iviva). Apparently an eastern form of the species. Ecol. In swards of Juncus and Cyperus species, in wet organic mud, c. 2500 m. Vern. Tampi, Enga, New Guinea. 69. Cyperus substramineus KUxK. Pfi. R. Heft 101 (1936) 398. — C. stramineus NEES in Wight, Contr. (1834) 74, non Desr. 1820; CLARKE, J. Linn. Soc. Bot. 21 (1884) 39; VaLtckx. Sur. Gesl. Cyp. Mal. Arch. (1898) 54. — Pycreus stramineus CLARKE, FI. Br. Ind. 6 (1893) 589; Camus, Fl. Gén. I.-C. 7 (1912) As). Annual with fibrous, yellowish roots. Stems very slender, tufted, often slightly curved, obtusely trigonous, smooth, 10-35 cm by }4—1 mm. Leaves very narrow, canaliculate, gradually acuminate, slightly scaberulous at the top, 1 mm wide; lower sheaths purplish. Inflorescence simple, spike-like or with a few short rays, congested, consisting of 2—15 spikelets. Involucral bracts 2(—3), suberect, over- topping the inflorescence, up to 10cm. Spikelets suberect even in fruit, oblong to linear, acute, strongly compressed, up to 50(—70)-flowered, 2(—3)4) cm by 2 mm; rachilla straight, wingless, persistent; internodes c. }4 mm. Glumes membranous, obliquely patent, sharply keeled, ovate, rather acute, shortly mucronate, with green, 3-nerved keel, stramineous to yellow nerveless sides and whitish hyaline margins, c. 2 by 114 mm. Stamens 2; anthers linear, c. EA mm. Style c.*/4 mm; stigmas 2, about as long as the style. Nut biconvex, laterally compressed, elliptic to slightly obovate, asymmetric, shortly apiculate, transversely rugulose, brown to blackish, 44-1 by 34-34 mm; epidermal cells longitudinally oblong. Distr. From Khasia and Ceylon through India to Farther India and W. Malesia: Malay Peninsula (Kedah, P. Penang). Ecol. In moist localities: grassy fields, rice-fields, in India at 0—1000 m. Notes. Closely allied to the widely distributed C. flavescens L., which is unknown from Malesia. C. substramineus is usually slenderer, its leaves and bracts are narrower, the lowest bract is nearly erect so that the inflorescence seems to be lateral, the spikelets are not patent, the glumes more acute, and the number of stamens is always 2 (3 in C. flavescens). The roots are very aromatic, the stems and leaves extremely tough. III. Subgenus Kyllinga (RotTB.) VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 42. — Kyllinga Rotts. Wescr: 65ic..(1773) 12. Type species: Kyllinga brevifolia ROTTB. [ser. I, vol. 7° FLORA MALESIANA 654 37. Section Queenslandiella stat. nov. — Queenslandiella Doin, Bibl. Bot. Heft 85 (1915) Paris 25 (1919) 60. Type species: Queenslandiella mira DoMIN. > 415. — Mariscopsis CHERM. Bull. Mus. DoMIN) KERN ( (a KERN 7888, b—d BACKER 27180). Fig. 68. Cyperus hyalinus VAHL. a. Habit, x74, b. cluster of spikelets, x %, c. spikelet, x3, d. nuts x 13 1974] Cy PERACEAE (Kern) 655 70. Cyperus hyalinus VAHL, En. 2 (1806) 329; Mia. Fl. Ind. Bat. 3 (1856) 254; Boeck. Linnaea 35 (1868) 482: CLARKE, J. Linn. Soc. Bot. 21 (1884) 46; RIDL. in Forbes, Wand. (1885) 520; VALcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 57, t. 2 f. 11; Koorp. Exk. Fl. Java 1 (1911) 185; ibid. 4 (1922) f. 195; Kx. Pf. R. Heft 101 (1936) 498; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 470. — C. pumilus (non L.) NEEs in Wight, Contr. (1834) 74. — Pycreus pumilus CLARKE, Fl. Br. Ind. 6 (1893) 591, quoad descr.; CAMuS, FI. Gén. I.-C. 7 (1912) 32, excl. f. 3. — Pycreus hyalinus Doin, Bibl. Bot. Heft 85 (1915) 417 in obs., non Me_rr. En. Born. (1921) 57. — Queenslandiella mira Domin, Bibl. Bot. Heft 85 (1915) 416, t. 11 f. 7-13. — Mariscopsis suaveolens CHERM. Bull. Mus. Paris 25 (1919) 60. — Mariscopsis hyalinus BALLARD, Kew Bull. (1932) 457. — Queenslandiella hyalina BALLARD in Hook. Ic. Pl. 33 (1933) t. 3208; BACK. Bekn. FI. Java (em. ed.) 10 (1949) fam. 246, p. 43. — Fig. 68. Annual with fibrous roots. Stems tufted, tri- quetrous, smooth, 3—20cm by 4-1 mm. Leaves weak, flat, gradually acuminate, scabrid on the margins towards the top, 1-3/4mm wide. In- florescence simple, anthelate but often contracted and head-like. Involucral bracts 3-6, obliquely erect to spreading, the longer ones much overtopping the inflorescence, up to 20 cm. Rays 3—6, with 5—20 spikelets, often very short, up to 6 cm. Spikes loose to rather dense, 8—12 by 8—10 mm; rachis 4-winged. Spikelets spicately arranged, obliquely to widely patent, ovate to oblong, strongly compressed, 4— 7(—10)-flowered, 4-8 by 2-3 mm, falling entire: rachilla strongly flexuous. broadly winged, dis- articulating at the base; internodes c. 1 mm. Glumes membranous, broadly ovate or ovate, with more or less falcate, strongly 3-nerved, serrate-scabrous green keel, whitish or yellowish, hyaline, strongly 3- nerved sides reticulate by the wide cells, 2—3 by c. 2 mm (the more or less recurved, up to 4 mm long mucro excluded), c. 4 imbricate. Stamens 2; anthers elliptic to oblong, ’4—1 mm. Stigmas 2. Nut biconvex, laterally compressed, suborbicular to elliptic, usually asymmetric, truncate to emarginate at the apex, not or hardly apiculate, castaneous to black, 1%,-134 by c. 1 mm; epidermal cells isodiametric. Distr. Tropical E. Africa, Mascarene Is., India, Farther India, tropical Australia (Queensland); in Malesia: islands near Java (coral islands in the Bay of Djakarta, Madura, Kangean Archipelago), Lesser Sunda Is. (Timor, Tanimbar Is.), Moluccas (Ceram, Key Is.). Ecol. Sandy and calcareous localities near the sea, at low altitudes; in Timor also more inland, on calcareous hills, up to 900 m. Vern. Tualanga, krémé, Tanimbar Is. Notes. On account of its bifid style and laterally compressed, biconvex nut, several authors placed this species in subg. Pycreus. It is the type-species of the monotypic genus Queenslandiella DOMIN (= Mariscopsis CHERM.), which is characterized by the disarticulation of the rachilla, the bifid style, and the biconvex, laterally compressed nut. However, all these characters are also found in Cyperus subg. Kyllinga, and so Queenslandiella differs from most Kyllingae only in the umbelliform inflorescence (not from all species of this subgenus, see e.g. C. transitorius K Ux. with anthelate inflorescence). Shape and texture of the glumes also point to its close relationship with the species of subg. Kyllinga. KUKENTHAL referred C. hyalinus to subg. Mariscus in spite of the digynous flowers. It is certainly not closely related to any of the trigynous Mariscus species. Since C. hyalinus was collected in Timor already by R. BRown in 1803, it is unlikely that it was introduced there, as BALLARD supposes for Queens- land. C. squarrosus L. and C. teneriffae Porr., which accompany C. hyalinus both in Timor and Queens- land, show almost the same distribution. The strong odour of dried specimens resembles that of fenugreek (Trigonella foenum-graecum L.). 38. Section Kyllinga Cyperus sect. Eu-Kyllinga KUK. Pfl. R. Heft 101 (1936) 576. Type species: Kyllinga brevifolia ROTTB. 71. Cyperus melanospermus (NEES) VALCK. SUR. Gesl. Cyp. Mal. Arch. (1898) 50, t. 2 f. 8; Koorp. Exk. Fl. Java 1 (1911) 185; KUk. Pfl. R. Heft 101 (1936) 583; S. T. BLake, J. Arn. Arb. 28 (1947) 225: STEEN. Mt Fl. Java (1972) t. 14: 14.— Kyllinga melanosperma Nees in Wight, Contr. (1834) 91: Boeck. Linnaea 35 (1868) 419; CLARKE, FI. Br. Ind. 6 (1893) 588; Merr. En. Philip. 1 (1923) 115; RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 59; Fl. Mal. Pen. 5 (1925) 138; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 44; KERN in Back. & Bakh. /. Fl. Java 3 (1968) 470. — C. aphyllus Hassk. Cat. Hort. Bog. (1844) 24, quoad specim., nec Kyllinga vaginata LAMK nec C. aphyllus VAHL, 1798.— Scirpus aphyllus Hassk. I.c. 352. — Kyllinga vaginata (non LAMK) ZOLL. Syst. Verz. 1 (1854) 63, incl. var. major HASskK. nom. nud.; Miq. FI. Ind. Bat. 3 (1856) 290. — Kyllinga pungens (non LINK) CLARKE, Philip. J. Sc. 2 (1907) Bot. 78. Perennial with 2—4 mm thick, creeping rhizome covered by ovate-lanceolate. fuscous to blackish sheaths. Stems approximate, rigid, triquetrous, almost winged, smooth, 30—175 cm by 2—4 mm. Upper 1-2 leaves shortly to rather long-laminate, abruptly acuminate. the others reduced to membranous, purplish sheaths transversely corrugate on the anterior side. Inflorescence capitate, consisting of a single, ovoid to oblong-ovoid head, (6—)10—12(—16) by 6-88-10) mm. Involucral bracts usually 3, spreading to reflexed, up to 20 cm. Spikelets numerous, obliquely spreading, oblong-elliptic or ovate-elliptic, strongly compressed, 1(—2)-flowered, stramineous to fuscous, falling off as a whole, 3-44 by 1-1/4 mm. Glumes hyaline, ovate-lanceo- late or elliptic-ovate, mucronulate, with sharp, smooth or spinulose keel, Ist and 2nd small, 3rd 214-344 mm, with 3—4-nerved sides, 4th 3—4 mm, with 2-nerved sides. Stamens 3; anthers linear, 656 1-4 mm. Stigmas 2. Nut biconvex, laterally com- pressed, oblong to obovate, shortly apiculate, brown to black, 114-2 by %- 4mm. ssp. melanospermus. — Synonymy see above. Stems 30-175 cm by 2—4 mm. Leaves practically all reduced to membranous sheaths, only the upper 1-2 very shortly laminate. Inflorescence very dense. (6—)10—12(—16) by 6—8(—10) mm. Spikelets oblong- elliptic, 3— 4%, by 1 mm. Glumes ovate-lanceolate, the third 2)4-3)4 mm long, the fourth 34-4 mm, both with distinct ferrugineous nerves and smooth or sparsely spinulose keel. Nut oblong or elliptic- oblong, black. 1/4-2 by 4-3{ mm. Distr. Tropical and subtropical Africa, S. Asia, Fiji; widely spread in Malesia, but nowhere common: Sumatra, Malay Peninsula, Java, Philippines (Luzon, Mindanao) Lesser Sunda Is. (Flores), Celebes. New Guinea. Ecol. Swamps, wet grassy fields, seepage hollows, open wet places in forests, usually 500-2000 m, sometimes up to 2200 m, one record (S. Sumatra, Lampongs) from 200 m. Vern. Rumput buwong, Sum. W.C., djukut bulu mata, djukut péndul,S, peaad-peaa, tudju-tudju.Celebes, kokwakokwa, New Guinea: Kapauku. ssp. bifolius (M1Q.) KERN, Reinwardtia 3 (1954) 62. f. 12. — Kyllinga bifolia Miq. F1. Ind. Bat. 3 (1856) 293. — C. brevifolius f. vaginatus VALCK. Sur. Ges]. Cyp. Mal. Arch. (1898) 47, t. 2 f. 4, p.p. — Fig. 69. Stems up to 75 cm by c. 2mm. Lower 1-2 leaves 10—25 cm by 3—4 mm. Inflorescence 6—10 by 6—8 mm, less dense than in ssp. melanospermus. Spikelets ovate-elliptic, 3-3/4 by 174-114 mm. Glumes elliptic- ovate, the first 24-3 mm long, the second 3-3/4 mm, with evidently serrate- spinulose keels. Nut broadly elliptic to obovate, 114 by 1-14 mm. Distr. Malesia: Malay Peninsula (Pahang), Java, Lesser Sunda Is. (Bali), New Guinea (Papua). Ecol. As in ssp. melanospermus, 1100—3000 m. Note. In many respects intermediate between ssp. melanospermus and C. brevifolius. From the former it can be distinguished by the usually shorter stems, the fairly developed leaf-blades, and the different shape of spikelets and nuts: from the latter by its much stouter stems, relatively smaller leaf- blades, larger spikelets and nuts, and triandrous flowers. 72. Cyperus aromaticus (RIDL.) MattrF. & KUK. Pfi. R. Heft 101 (1936) 581: KERN, Reinwardtia 3 (1954) 62; Koyama, Bot. Mag. Tokyo 83 (1970) 186. — Kyllinga polyphylla WILLD. ex KUNTH, En. 2 (1837) 134, non C. polyphyllus VAHL. — Kyllinga aromatica RIDL. Trans. Linn. Soc. II, Bot. 2 (1884) 146. Very near to C. melanospermus. Involucral bracts more numerous, (5—)7(—8). Inflorescence subglobose, consisting of a terminal head and some smaller sessile ones in the axils of the involucral bracts, all confluent into a head c. 1 cm o. Glumes more distinctly mucronate. Distr. Native to E. Africa; in Malesia introduced and naturalized in Singapore I.; the earliest collection is from 1941; nowadays very abundant. Ecol. In grassy open places. Notes. Also introduced into Ceylon, Solomon FLORA MALESIANA [ser. I, vol. 7° Is., Samoa and in Fiji, where it is spreading rapidly and has become a noxious weed of pastures in the wet zone. See PARHAM, Plants of the Fiji Islands (1964) 296. The Malesian specimens belong to the stout var. elatus (STEUD.) KUk. Pfl. R. Heft 101 (1936) 582. — Kyllinga elata SteuD. Syn. 2 (1855) 70, non C. elatus L.— Up to 90cm. Stems rigid, strongly compressed, 2—3 mm _ thick. Involucral bracts up to 7 mm wide. Spikelets 4 mm long. 73. Cyperus brevifolius (RotTB.) HAssk. Cat. Hort. Bog. (1844) 24: VaLcK. Sur. Gesl. Cyp. Mal. Arch. (1898) 46, t. 2 f. 3, incl. f. subtrifoliatus, f. gracilis et f. pumilus VALCK. Sur., excl. f. vaginatus; Koorp. Exk. Fl. Java 1 (1911) 185, non ibid. 4 (1922) f. 193; KOK. Pfl. R. Heft 101 (1936) 600, incl. f. pumilus et f. firmiculmis KUxK.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 225: Kern in Back. & Bakh. f. FI. Java 3 (1968) 470. — Schoenus coloratus LINNE, Sp. Pl. ed. 2 (1762) 64, p.p., non C. coloratus VAHL, 1806. — Schaenoides brevifolius RottB. Descr. PI. rar. Progr. (1772) 15, nom. inval. (cf. Montreal Code art. 20, note 1). — Kyllinga brevifolia Rotts. Descr. & Ic. (1773) 13, t. 4 f. 3; Mig. Fl. Ind. Bat. 3 (1856) 291; Boeck. Linnaea 35 (1868) 424; Naves, Nov. App. (1882) 300: CLARKE, Fl. Br. Ind. 6 (1893) 588; Philip. J. Sc. 2 (1907) Bot. 78; RmpL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 58: CLARKE, Ill. Cyp. (1909) t. 1 f. 1-4; Camus, Fl. Gén. I.-C. 7 (1912) 24, f. 3, 3-6; MerR. En. Philip. 1 (1923) 114: Rupe. Fl. Mal. Pen. 5 (1925) 138; Back. Onkr. Suiker. (1928) 145, t. 145; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 45. — Kyllinga cruciformis SCHRADER ex R. & S. Mant. 2 (1824) 137: Boeck. Linnaea 35 (1868) 426. — Kyllinga sororia KUNTH, En. 2 (1837) 131; Mia. FI. Ind. Bat. 3 (1856) 293. — Kyllinga gracilis KUNTH, En. 2 (1837) 134: ZoLi. Syst. Verz. (1854) 63, p.p-; Mig. Fl. Ind. Bat. 3 (1856) 292; Naves, Nov. App. (1882) 300. — Kyllinga pumilio STEUD. [in Zoll. Syst. Verz. 1 (1854) 63, nom. nud.] Syn. 2 (1855) 67: Mig. Fl. Ind. Bat. 3 (1856) 292. — Kyllinga monocephala (non Rotts.) ZOLL. Syst. Verz. (1854) 63, p.p. — Kyllinga longiculmis Mia. FI. Ind. Bat. 3 (1856) 292; Naves, Nov. App. (1882) 300. — Kyllinga rigidula StTEUD. Syn. 2 (1855) 71, quoad specim. philip.; Miq. Fl. Ind. Bat. 3 (1856) 294; NAves, Nov. App. (1882) 300. — Kyllinga squarrosa STEUD. Syn. 2 (1855) 68; NAves, Nov. App. (1882) 300. — Kyllinga caespitosa NEES var. robusta Boeck. Linnaea 35 (1868) 413; SCHEFFER, Nat. Tijd. N. I. 34 (1874) 45; VmpaL, Phan. Cum. Philip. (1885) 155; Rev. Pl. Vasc. Filip. (1886) 283.— Pycreus pumilus var. substerilis CAMUS, Not. Syst. 1 (1910) 241; Fl. Gén. I.-C. 7 (1912) 33, p.p. — Kyllinga colorata DRUCE, Rep. Bot. Exch. Club Br. Isl. 1916 (1917) 630. — C. crypsoides KERN, Reinwardtia 2 (1952) 128, f. 14. — Fig. 70. Perennial; rhizome horizontally creeping, sub- terraneous or close upon the ground, covered by ovate-lanceolate, reddish-brown scales, 1-3 mm thick, the internodes very variable in length (de- pendent on soil-conditions). Stems approximate to distant, rigidulous, triquetrous, smooth, 3—40 cm by 4- i ’4 mm. Leaves usually well-developed, weak to somewhat rigid, canaliculate, scabrid on the margins in the upper part, grassgreen, 1-3 mm wide, very rarely only the upper one with a very short 1974] Cy PERACEAE (Kern) 657 SS =x SSS = SSS Se Fig. 69. Cyperus melanospermus (NEES) VALCK. Sur. ssp. bifolius (M1Q.) KERN. a. Habit, x |4, b. spikelet, c. glumes, d. deflorate flower, e. nut, f. prophyll, all x 10 (a—f VAN STEENIS 4360). 658 Fig. 70. Cyperus brevifolius (ROTTB.) HAssk. Habit of creeping rooting specimen in dry lawn at Djakarta, x JZ (BACKER 33612). FLORA MALESIANA [ser. I, vol. 7° blade. Inflorescence capitate, usually consisting of a terminal, globose to ovoide, 5-10 mm long and wide head, at first greenish, finally stramineous to fuscous, rarely with 1—2 smaller, sessile heads at the base; rachis narrowly conical. Involucral bracts 3-4, rarely 2 or 5—6, spreading to reflexed, but the lowest often remaining erect, 3—6(—20) cm long, very rarely all erect and very short (c. 2 cm). Spikelets closely packed, spreading or obliquely erect, elliptic- oblong or oblong-lanceolate, strongly compressed, 1(—2)-flowered, (2—)3(—4) by c. 1 mm, falling off as a whole. First glume elliptic-lanceolate, Y-1 mm, 2nd ovate, 1mm, 3rd and 4th boat-shaped, strongly compressed, mucronulate, acutely keeled, with 2—3- nerved sides and more or less spinulose, rarely smooth keel, resp. (1/4-)2)4(—3) mm and (2—)3(—4) mm long. Stamens 1—2(—3); anthers linear, c. 1 mm. Stigmas 2. Nut biconvex, laterally compressed, obovate or elliptic, apiculate, yellowish brown, 1-1/(-1/4) by 4-% mm. Distr. Widely spread in the tropical and warm- temperate regions of the whole world; very common throughout Malesia. Ecol. In sunny or partly shaded localities, along roads, in grassy fields, damp forest clearings, on river-banks, etc.; 0—-1500 m, rarely up to 1900 m (see var. stellulatus). Use. Like C. kyllingia sometimes used as a fodder for cattle and horses; food value satisfactory, but yield scanty. Vern. Djukut péndul, téki, S, rumput sadanan, téki rowo, J, komes, Md, rumput kapas, Sumatra, badjawa, Timor, wutu intalun, N. Celebes, kokolguli, Aru Is.; Philip.: bibi-inok, Bon., kadkadot, Ig., pugo-pugo, C. Bis.; New Guinea: punum, Wapi, ylampi, Enga. Notes. Extremely variable: the characters of the (certainly edaphic) forms distinguished by VALCKE- NIER SURINGAR and KUKENTHAL are not constant. C. crypsoides KERN from Celebes, with lower leaves reduced to bladeless sheaths, only the upper one with a short, rarely more than 2 cm long blade, and erect, linear to lanceolate, pungent involucral bracts, I now consider a form of C. brevifolius occurring in very wet localities. Systematic value may have: var. stellulatus (VALCK. Sur.) KUx. Pfl. R. Heft 101 (1936) 603. — C. brevifolius f. stellulatus VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 48, t. 2 f. 5. — Kyllinga fuscata Miq. Fl. Ind. Bat. 3 (1856) 294. — Kyllinga intermedia (non R.BR.) MerR. En. Philip. 1 (1923) 115. — Kyllinga brevifolia var. stellulata Onwt, Bot. Mag. Tokyo 56 (1942) 199. Dwarfish. Inflorescence small, with fewer (c. 20) spikelets, 4—5S(—7) mm across, finally fuscous. Glumes with smooth keel. Stamens 3 or 2. Nut 14-14% by 4-1 mm, brown to castaneous. Distr. Malesia: Central and East Java, Lesser Sunda Is. (Lombok), Philippines (Luzon, Mindoro, Negros, Mindanao), W. New Guinea (Arfak Mts). Ecol. Open marshy places, grassy fields, open Casuarina forests, 1400-2300 m. 74. Cyperus sesquiflorus (ToRR.) MATTF. & KUK. Pfi. R. Heft 101 (1936) 591, f. 6 E-J. — Kyllinga sesqui- flora Torr. Ann. Lyc. N. York 3 (1836) 287. 1974] Cy PERACEAE (Kern) 659 In Malesia only: var. subtriceps (NEES) KoyAMA, Quart. J. Taiwan Mus. 14 (1961) 191. — Kyllinga cylindrica NEES in Wight, Contr. (1834) 91, incl. var. subtriceps NEES; Boeck. Linnaea 35 (1868) 415, p.p.; CLARKE, Fl. Br. Ind. 6 (1893) 588, excl. specim. malacc.; Philip. J. Sc. 2 (1907) Bot. 79; Camus, Fl. Gén. I.-C. 7 (1912) 23; Merr. En. Philip. 1 (1923) 115; Back. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 44, p.p.; non C. cylindricus CHAPM. 1878. — C. viridulus VALCK. SUR. Gesl. Cyp. Mal. Arch. (1898) 51, t. 2 f. 7, 9, non Boeck. 1868 nec Kyllingia viridula HOCHST. ex A. Ricu. — Kyllinga odorata VAHL var. cylindrica KUK. in Merr. J. Str. Br. R. As. Soc. 76 (1917) 80; MERR. En. Born. (1921) 58; KUx. Candollea 6 (1936) 422. — C. sesquiflorus var. cylindricus KUK. Pf. R. Heft 101 (1936) 593; Bot. Jahrb. 70 (1940) 463; KERN, Reinwardtia 3 (1954) 65.— C. kernianus OHWI & Koyama, J. Jap. Bot. 30 (1955) 126; Koyama, Act. Phytotax. Geobot. 16 (1955) 35, t. 4 f..B.— C. sesquiflorus ssp. cylindricus (NEES ex WIGHT) KOYAMA, Bot. Mag. Tokyo 83 (1970) 187. Perennial with very short rhizome. Stems tufted, triquetrous, smooth, seemingly thickened at the base by reddish brown cataphylls, 10-30 cm by 1 mm. Leaves rigidulous, flat, gradually acuminate, scabrid on the margins in the upper part, 2-4 mm wide. Inflorescence capitate, whitish, finally straw- coloured, consisting of a terminal, cylindrical head 10-12 by 4-5mm, and 0-2 lateral, subglobose, sessile heads much smaller than the terminal one. Involucral bracts 3—S, finally reflexed, up to 10 cm. Spikelets numerous, patent, elliptic, strongly com- pressed though somewhat turgid, maturing 1 nut, 2-2)% by 1-1/4 mn, falling off as a whole; rachilla cylindric, disarticulating at the base. Glumes hyaline, Ist and 2nd small, 3rd and 4th nearly equal in length, broadly ovate, acute or apiculate, with smooth or hardly spinulose, sharp keel, resp. strongly 9—11- and 5-—7-nerved, 2—244 mm. Stamens 2; anthers oblong, 4-34 mm. Stigmas 2. Nut biconvex, laterally compressed, obovate or broadly obovate, obtuse, shortly apiculate, black, c. 1/4 by 1 mm. Distr. Tropical Africa; in SE. Asia from Ceylon and India to Yunnan, Formosa, and Malesia: scattered in Sumatra (Atjeh, Tapanuli, East Coast Res.), Java, Lesser Sunda Is. (Bali), N. Borneo, Philippines (Luzon, Mindanao), Aru Is. (Trangan I.), and New Guinea. Not known from the Malay Peninsula (see C. triceps). Ecol. In open grasslands, usually 1000—1600 m, rarely at low altitudes. Note. The typical var. sesquiflorus, occurring in tropical Africa, the warm parts of N. and S. America, and Australia, differs mainly by the larger, 3-3!4 mm long spikelets. 75. Cyperus triceps ENDL. Cat. Hort. Ac. Vindob. 1 (1842) 94; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 50, t.2 f.6; KUk. Pfl. R. Heft 101 (1936) 578; KERN, Reinwardtia 3 (1954) 61.— Schaenoides triceps Rotts. Descr. Pl. rar. Progr. (1772) 15, nom. inval. (cf. Montreal Code, art. 20, note 1).— Kyllinga triceps Rotts. Descr. & Ic. (1773) 14, t. 4 f. 6; BOECK. Linnaea 35 (1868) 413, p.p.; CLARKE, FI. Br. Ind. 6 (1893) 587.— Kyllinga pumila StEuD. Flora 35 (1842) 596, non Naves, Nov. App. (1882) 300. — Kyllinga cylindrica (non NEES) CLARKE, FI. Br. Ind. 6 (1893) 588, quoad specim. malacc.; RIDL. Mat. FI. Mal. Pen. (Monoc.) 3 (1907) 59; Fl. Mal. Pen. 5 (1925) 137. Perennial with short rhizome; stolons wanting. Stems densely tufted, slender, sometimes setaceous, obtusely trigonous, smooth, 5—20(—40) cm by up to 1 mm, the incrassate base covered by brownish, desintegrating sheaths. Leaves flat or slightly con- duplicate, weak, gradually acuminate, 1—2(—3) mm wide. Inflorescence capitate, consisting of (1—)3(—S) dense, sessile heads; central head subglobose, obtuse, 5—8 by Smm, lateral ones globose, somewhat smaller; rachis cylindrical. Involucral bracts 3-4, widely spreading to reflexed, up to 10 cm. Spikelets numerous, patent, oblong, strongly compressed, pale green to whitish, 1-flowered, c. 2 by %% mm, falling off as a whole; rachilla very short, disarticu- lating at the base. Glumes hyaline, ovate-oblong, acuminate or submucronulate, with sharp, smooth or hardly spinulose keel; Ist and 2nd glume small, 3rd 7-nerved, c. 134mm, 4th glume 5-nerved, c. 2 mm; nerves less prominent than in C. sesquiflorus. Stamens 2; anthers oblong-linear. Stigmas 2. Nut biconvex, laterally compressed, oblong, apiculate, yellowish brown to brown, 1-1/4 by 4 mm. Distr. Widely spread over tropical Africa, India, and Farther India to S. China and Australia (Queensland, New South Wales); in Malesia once collected (Singapore, 1885); probably an introduc- tion. Ecol. In Singapore as a weed on a sandy lawn. Notes. Confused with C. sesquiflorus, from which it is distinguishable by the shape of the inflorescence, the narrow spikelets, and the narrow, brown fruits. Although the basionym K yllinga triceps ROTTB. is illegitimate, the correct name of this species in Cyperus is C. triceps ENDL. (1842), the synonym Kyllinga bulbosa BEAUV. (1804) already in 1842 not being transferable to Cyperus (art. 66 note 2 and art. 72 Montreal Code). ROTTBOELL’s specific epithet was nomenclaturally superfluous because the name Scir- pus glomeratus L. was cited in synonymy (art. 63), but the type of Kyllinga triceps is not that of Scirpus glomeratus, as ROTTBOELL indicated a definite type (“Dominus K@nigius legit.’’). 39. Section Alati KUK. Pfl. R. Heft 101 (1936) 604. Type species: Kyllinga alata NEES. 76. Cyperus kyllingia ENDL. Cat. Hort. Ac. Vindob. 1 (1842) 94; KUk. Pfl. R. Heft 101 (1936) 606, f. 64 C-D, incl. f. humilis, f. tenuis et f. subtriceps KUK.; S. T. BLAKE, J. Arn. Arb. 28 (1947) 225; KERN in Back. & Bakh. f. Fl. Java 3 (1968) 469. — Gramen capitatum RuMPH. Herb. Amb. 6 (1750) 8, t. 3 f. 2. — Scirpus cephalotes JacQ. Hort. Vind. 1 (1770) 42, t. 97, non C. cephalotes VAHL, 1806. — Kyllinga 660 FLORA MALESIANA [ser. I, vo monocephala Rotts. Descr. & Ic. (1773) 13, t. 4 f. 4; ZOLL. Syst. Verz. 1 (1854) 63, p.p.; Mia. Fl. Ind. Bat. 3 (1856) 291; Boeck. Linnaea 35 (1868) 427, incl. var. subtriceps KUNTH et var. mindorensis BOECK.; CLARKE, Fl. Br. Ind. 6 (1893) 588; Philip. J. Sc. 2 (1907) Bot. 77; RipL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 58; oa Ill. Cyp. (1909) t. 2 f. 1-2; Camus, Fl. Gén. -C. 7 (1912) 25, f. 3, 1-2; Merr. En. Philip. 1 (1923) 1s: RIDL. Fl. Mal. Pen. 5 (1925) 138; Back. Onkr. Suiker. (1928) 144, t. 144; Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 44.— Kyllinga gracilis (non KUNTH) ZOLL. Syst. Verz. 1 (1854) 63, p.p. — Kyllinga triceps (non RoTTB.) BLANCO, FI. Filip. (1837) 34 (‘Killinga’); StEUD. Syn. 2 (1855) 72; Naves, Nov. App. (1882) 330; K.Scu. & Laut. Fl. Schutzgeb. (1900) 194. — Kyllinga brevifolia (non Rotts.) Mor. Syst. Verz. (1846) 95. — C. leucocephalus HaAssk. PI. Jav. Rar. (1848) 87, non Retz. 1789. — Kyllingia gracilis KUNTH var. capitulo globoso NEES in Hook. J. Bot. Kew Misc. 6 (1854) 28. — K yllinga mindorensis STEUD. Syn. 2 (1855) 67; Mig. Fl. Ind. Bat. 3 (1856) 292; Naves, Nov. App. (1882) 300.— C. mono- cephalus F.v.M. Fragm. 8 (1874) 271; VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 44, t.2 f. 1; Koorb. Exk. Fl. Java 1 (1911) 185; ibid. 4 (1922) f. 193, 194, non Roxs. 1832. — Kyllinga cephalotes DRUCE, Rep. Bot. Exch. Club Br. Isl. 1916 (1917) 630. Perennial; rhizome horizontally creeping, covered by ovate-lanceolate scales, 1-2 mm thick, the inter- nodes very variable in length (dependent on soil- conditions). Stems approximate to distant, rigidulous, triquetrous, smooth, 5—45 cm by 1-1/4 mm. Leaves well-developed, flaccid or rigidulous, canaliculate, scabrid on the margins in the upper part, grassgreen, 2—4(—5S) mm wide. Inflorescence capitate, consisting of a terminal, ovoid-globose or ellipsoid head 8—12 by 6-10mm, with usually 1-3 much smaller, sessile heads at the base, at first snowy white, turning fulvous after anthesis; rachis conical. In- volucral bracts 3-4, spreading to reflexed, the lowest up to 30cm. Spikelets very closely packed, spreading, obliquely ovate- -elliptic, strongly com- pressed, 3-3)% by (1-)1/4(-124) mm, falling off as a whole. First and second glume narrow, membranous, 1-1/4 mm long; other glumes boat-shaped, strongly compressed, apiculate or mucronulate, acutely keeled, broadly winged on the keel from somewhat above the base, green in the upper part of the keel, otherwise white, usually reddish brown puncticulate; wings ciliate-serrate, YA- Zi mm broad; 3rd glume en- closing a ¢ flower, 2)4-3 mm long, 4th glume with narrow, cuneate base (in fact the winged rachilla), enclosing a 3 or ¢ (sometimes a barren) flower, 3-314 mm, 5th glume when present small, sterile. Stamens 3; anthers linear, 4-1 mm. Stigmas 2. Nut biconvex, laterally compressed, oblong or oblong- obovate, apiculate, yellowish brown to castaneous, 14-14 by 4% mm. Distr. Common in the warmer parts of Asia, less so in tropical Africa and Australia, very rare in tropical America; common throughout Malesia. Ecol. In sunny or somewhat shaded waste places, road-sides, open grasslands, secondary growths, etc., at low and medium altitudes, rarely up to 1750 m. Use. In those regions where grass is scantily available in the dry monsoon, the plant is appreciated as a fodder for cattle and horses; the food value is satisfactory. Vern. Teki, teki badot, M, djukut péndul bodas, S, meélaran, sukét wudélan, udél-udélan-alit, J, ping ajuping, Md, rumput tai, Trengganu, rumput kutu radja, Lingga, andor si tiga bulung, Asahan, kémbili- kémbili, Karo-Batak, sélang sajong, Karo, idjar kapas, Enggano, rumput iput-iput, Banka, kakamut, kingkimut, tomatong, N. Borneo, teki sela, tinti lamonu, Celebes, nésar leteng, Wetar, dodopala, tittingapu, Moluccas; Philip.: anuang, mustra, mutha, Tag., barubotones, bolobotones, borobotones, boskad, bosoboténes, malabatones, sud-sud, Bis., botoncillo, Sp., borsa ngadadakkel, Uk., baki-baki, pungos, S. L. Bis., bosikad, C. Bis., busikad, P. Bis., katutu, Mag., kuru-kamoting-orig, Bik., sangsangitan, Bon., uli-uli, Bag.; New Guinea: lemisaharu, Kutubu. Hybrid Cyperus distans L. f. x C. rotundus L. A Philippine collection from Negros, Prov. of Negros Oriental, Dumaguete (Cuernos Mts), June 1908, ELMER 10369, is in all respects intermediate between C. distans and C. rotundus, which species were collected in the same locality at the same time. There can be little doubt that it is a hybrid of the said species. The completely sterile spikelets are strongly accrescent, c. 5 cm long, 114 mm wide, and c. 50-flowered. Glumes c. 2/4 mm long, less broadly hyaline-margined than in C. distans, more obtuse than in C. rotundus, and but slightly overlapping. Internodes of the rachilla c. 1/4 mm long. Anthers c. 1 mm long. Doubtful and Excluded A fairly great number of Cyperus species has been recorded for or described from Malesia which cannot be placed satisfactorily. Most of them must be referable to some of the species accepted above, but they are not determinable from the descriptions, and the specimens on which the records were based are unavailable. This is especially so with several names in NAvEs’s Novissimae Appendix (1882). Several of PRESL’s records in Reliquiae Haenkeanae (1828-30) “in insula Luzon” are certainly not Philippine, but American. See also MERRILL, En. Philip. 1 (1923) 108-109, 111, 116, and KUKENTHAL, Pfl. R. Heft 101 (1936) 626-627. Cyperus albus PRESL, Rel. Haenk. 1 (1828) 175; KUNTH, En. 2 (1837) 102; StEUD. Syn. 2 (1855) 52; Mia. FI. Ind. Bat. 3 (1856) 287; NAves, Nov. App. (1882) 306. — Luzon. Cyperus alternifolius L. — VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 103. Native to Madagascar. The old specimen in L from “Java”, without any details, was certainly mislabelled. Cyperus brunnescens BOECK. Cyp. Nov. 2 (1890) 3; CLaRKE, FI. Br. Ind. 6 (1893) 619. — “Herb. Mus. botan. Hauniensis.-Mis. Voigt. Singapore.” — In all probability VoiGT never visited Malesia, and the type locality should read “‘Serampore”’ (in Bengal), where most of VoIGT’s specimens were collected. The type is a poor fragment. See KUx. Pfl. R. Heft 101 (1935) 159. “Cyperus clavatus LAMK.” — USTERI, Beitr. 1974] Cy PERACEAE (Kern) 661 Kenntn. Philip. Veg. (1905) 131. LAMaRcK did not publish this name. The specific epithet was mis-spelt. Cyperus corymbosus Rotts. — Mia. Sum. (1861) 600; NaAves, Nov. App. (1882) 303. — Not known from Banka, nor from the Philippines. Cyperus flavescens L. — Boeck. Linnaea 35 (1868) 438, quoad pl. Luzon; Naves, Nov. App. (1882) 301. — A widely distributed species, but in Asia extending eastward to the western Himalaya, not reaching Malesia. The records for Indo-China are also wrong. See KERN, Blumea 10 (1960) 641. Cyperus flavicomus Micux. — NaAves, Nov. App. (1882) 301. — A N. American species not occurring in Malesia. Cyperus haematodes ENDL. — STEUD. Syn. 2 (1855) 44: Mia. Fl. Ind. Bat. 3 (1856) 282; NAves, Nov. App. (1882) 305. — The Philippine record is due to an error of STEUDEL. The type is from Norfolk Island. Cyperus hexastachyos B pendulus Nees in Wight, Contr. (1834) 82. — “Java, Manilla”. Referred by Kuk. Pfl. R. Heft 101 (1936) 125 to C. bulbosus VAHL, which is unknown from the Philippines. Cyperus kiikenthalii MERR. J. Arn. Arb. 19 (1938) 322. — C. javanicus KU«K. in Fedde, Rep. 29 (1931) 194; Pfl. R. Heft 101 (1936) 319, non Houtt. 1782. — Juncellus kiikenthalii BACK. Bekn. Fl. Java (em. ed.) 10 (1949) fam. 246, p. 42. — “Java, W. HILLEBRAND.” Erroneously localized (see Fl. Mal. I, 1, 1950, 232). The type was destroyed during World War II. Cyperus laevigatus L.— Mug. FI. Ind. Bat. 3 (1856) 260; Naves, Nov. App. (1882) 301. — Not known from Malesia. A specimen in S labelled “e Java, leg. MELLERBORG s.n.”” was apparently wrongly localized. Cyperus lanceolatus PotR. var. compositus PRESL, Rel. Haenk. 1 (1828) 167; KUxk. Pfl. R. Heft 101 (1936) 350. — Luzon, Sorsogon. — An American plant not occurring in the Philippines. Cyperus lanceus THUNB. — VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 70, t. 2 f. 20. — A native of Africa. The specimens cited l.c. (in L, without authentic label) do not originate from Malesia. See KERN, Reinwardtia 2 (1952) 122. Cyperus longus L. — Mig. FI. Ind. Bat. 3 (1856) 275; Naves, Nov. App. (1882) 304.— A Eurasian species not extending to SE. Asia. The records for Indo-China (Fl. Gén. I.-C. 7, 1912, 68) refer to specimens of C. corymbosus Rotts. See KERN, Blumea 10 (1960) 643. Cyperus luzonensis PRESL, Rel. Haenk. 1 (1828) 174; KunTH, En. 2 (1837) 101; Steup. Syn. 2 (1855) 52; Mig. Fl. Ind. Bat. 3 (1856) 287; NAves, Nov. App. (1882) 306. Cyperus luzulae (L.) Retz. — VALCK. Sur. Gesl. Cyp. Mal. Arch. (1898) 89, t. 4 f. 8. — Courtoisia cyperoides (non Nees) Mig. Fl. Ind. Bat. 3 (1856) 295.— “Java or Sumatra” (L). Supposed to be collected by JUNGHUHN, but no authentic label extant. Occurrence in Malesia at JUNGHUHN’s time is very unlikely. Cyperus melanospermus var. plurifoliatus KUxK. Pfl. R. Heft 101 (1936) 585.— “Java, HILLEBRAND.” Wrongly localized. See Fl. Mal. I, 1 (1950) 232. Cyperus minutiflorus PRESL, Rel. Haenk. 1 (1830) 351; KUNTH, En. 2 (1837) 102; StEupD. Syn. 2 (1855) 52; Mia. Fl. Ind. Bat. 3 (1856) 287; Naves, Nov. App. (1882) 306. — C. micranthus PReEsL, Rel. Haenk. 1 (1828) 178, non R. & S. 1824. — C. breviflorus A. Dietr. Sp. Pl. 2 (1833) 316. — Luzon, fide PRESL. Cyperus pangorei Rotts. — NAves, Nov. App. (1882) 304. — Not known from the Philippines. Cyperus philippensis PResL, Rel. Haenk. 1 (1828) 174; KUNTH, En. 2 (1837) 101; StEup. Syn. 2 (1855) 52; Mia. Fl. Ind. Bat. 3 (1856) 287; NAves, Nov. App. (1882) 306. — Luzon, fide PRESL. Cyperus sexflorus R.BR.— Naves, Nov. App. (1882) 304. — An Australian species not known from the Philippines. Cyperus stenostachyus (non BENTH.) VALCK. SUR. Gesl. Cyp. Mal. Arch. (1898) 112, t. 4 f. 25, t. 5 f. 7.— “Prob. Java.”’ The specimens on which this record was based belong to C. longus var. badius (DesF.) ASCH. & GR., a Mediterranean plant not occurring in Malesia. See KERN, Reinwardtia 3 (1954) 27. Cyperus tegetiformis RoxB. — Naves, Nov. App. (1882) 303. — Not occurring in the Philippines. Cyperus tenuiflorus (an RoTTB.?) VALCK. SuR. Gesl. Cyp. Mal. Arch. (1898) 111, t. 4 f. 24, t. 5 f. 6: Koorp. Exk. Fl. Java 1 (1911) 196; ibid. 4 (1922) f. 217.— A poor, very old, inadequately labelled collection from W. Java(““Gedogan, bamboo-grove’’), was determined by CLARKE as C. tenuiflorus ROTTB. and as such mentioned by VALCKENIER SURINGAR. Without having seen the specimens KUKENTHAL, Pfl. R. Heft 101 (1935) 103, referred SURINGAR’s record to C. mitis STEUD. (C. tenuiflorus ROxB., non ROTTB.) and mentioned two other Java collections of this species (JUNGHUHN, N. J. ANDERSSON), which I have not seen. I cannot identify the specimens in the Rijksherbarium, which belong in the extremely difficult sect. Cyperus, but are only in very young flower. Cyperus tuberosus RoTTB. — KUNTH, En. 2 (1837) 50; Mia. Fl. Ind. Bat. 3 (1856) 265. — ? Luzon, fide KUNTH. — The record must be erroneous as the species is not known from the Philippines. Kyllinga pumila (non StEuD.) NAves, Nov. App. (1882) 300. — STEUDEL’s name is a synonym of Cyperus triceps (ROTTB.) ENDL., which is unknown from the Philippines. See p. 659. 15. TETRARIA BEAUvV. Mém. Inst. Fr. (Acad. Sc. Paris) 1812, II (1816) 54; BAILLON, Hist. Pl. 12 (1894) 374; CLARKE, Fl. Cap. 7 (1898) 275; TuRRILL, Kew Bull. (1925) 72: 662 FLORA MALESIANA [ser. I, vol. 7° f revel ‘DF | Fig. 71. Tetraria borneensis KERN. a. Habit, x 7%, b. spikelet, x 8, c. glume, x 4, d. deflorate flower, e. sterile flower, f. nut, all x 8,g. anther, x 16 (a PURSEGLOVE 5072, b—e, g KOSTERMANS 10737, Jf PURSEGLOVE 5559). 1974] CYPERACEAE (Kern) 663 KUk. in Fedde, Rep. 48 (1940) 195; Levyns, J. S. Afr. Bot. 13 (1947) 73. — Elynanthus Nees, Linnaea 7 (1832) 520; ibid. 9 (1835) 298: Boeck. Linnaea 38 (1874) 253; B. & H. Gen. Pl. 3 (1883) 1063, vix BEAUV. ex LesTiB. Essai Fam. Cyp. (1819) 32. — Fig. 71-72. Perennial herbs with short woody rhizome. Stems tufted, erect, terete, trigonous or triquetrous. Leaves basal or also cauline, setaceous, subulate or flat, sheaths long, those of the cauline leaves often tubular, dark; ligule sometimes present. Inflorescence paniculate, narrow. Bracts leafy, not sheathing, or with tubular sheaths. Spikelets several to many, solitary or in clusters, oblong- lanceolate, more or less compressed, 2(—3)-flowered. Rachilla very short, straight, not produced beyond the upper flower. Glumes 4—12 (usually 5—7), distichous or subdistichous, the lower ones (usually 4—6) empty, small, the Fig. 72. Tetraria borneensis KERN gregarious in thickets on kerangas soil in Bako National Park. Sarawak, with young Ploiarium and Dacrydium in background (photogr. SLEUMER, 1963). 664 FLORA MALESIANA [ser. I, vol. 7° upper ones flower-bearing. Flowers usually 2, close together, the lower male or functionally male (gynaeceum abortive), the upper bisexual. Hypogynous bristles (OQ—)2—9(—14), delicate, sometimes shortly plumose. Stamens 3, 4, 6, or 8; anthers linear, with distinctly produced connective, often auricled at the base (ears without pollen). Style slender, continuous with the ovary, dilated at the base, with 3, 4, 6, or 8 stigmas; style-base persistent on the nut. Nut usually trigonous, ovoid-oblong or obovoid, with a (sometimes conspicuous) sterile apex (the persistent style-base). Distr. About 35 spp., most of them in extra-tropical S. Africa, a few in Australia, in Malesia 1 sp. in Borneo. 1. Tetraria borneensis KERN, Blumea 9 (1958) 222, f. 2. — Fig. 71-72. Stems rather stout, obtusely trigonous, smooth, c. 14-2 m by 2-4mm (5-6mm at the base), the incrassate base covered with castaneous sheaths and their fibrous remains. Basal leaves coriaceous, much shorter than the stems, conduplicate at the base, otherwise canaliculate or flat, gradually narrowed into along triquetrous point, scabrous on the margins. 4-10 mm wide; cauline leaves 3—4, shorter than the basal ones, with tubular castaneous 4—8 cm long sheaths; the emarginature at the orifice acute, v- shaped, with overlapping margins. Inflorescence loose, 40-90 cm long, consisting of 4-8 distant fascicles of branches; branches 2—4 together, erect, unequal, ancipitous, scaberulous. Lower bracts simi- lar to cauline leaves but shorter, 5—10 cm long, the upper gradually shorter. Spikelets numerous, solitary, shortly peduncled, narrowly lanceolate, acute, 2- flowered, 7-8 by 1-1/4 mm. Glumes 7, subdistichous, castaneous; lower 4 empty, ovate, shortly aristate; flower-bearing glumes oblong, c. 6 mm long, asper- ous towards the apex; uppermost glume thinly membranous, empty. Lower flower male, or func- tionally male (with abortive gynaeceum), upper bisexual. Bristles 2—3, capillary, white, minutely scaberulous at the top. Stamens in both flowers 6; anthers linear, rufescent, inconspicuously auricled at the base, 2mm long, with a c. 1/4 mm long scabrous appendage of the connective. Persistent style-base narrowly pyramidal, antrorsely scabrous, stramineous, c. 2mm. Stigmas 3. Nut trigonous, obovoid, attenuate at the base, smooth, castaneous, 24-3 by 1-144 mm. Distr. Malesia: Borneo (E. Borneo, Kelindjau River region; Sarawak: Lobok Pasir, Telok Asam; W. Borneo: bank of Sendabai lakes, Mt Kenepai). Ecol. Marshy ‘padang’, on sandy soil, at c. 100 m, locally very common and a dominant in this type of vegetation (E. Borneo); in Sarawak also in open heath forest (Bako National Park) and abundant but localized in the centre of peat swamp forest (ANDERSON, Gard. Bull. Sing. 20, 1963, 216). Note. In view of the distribution of the other species the occurrence of a Tetraria in the equatorial lowland of Borneo is very surprising. The group of genera to which Tetraria belongs is extremely difficult, and the genera have been circumscribed in very different ways. The subdistichous glumes, the short straight rachilla, the barren lower flower, the number of stamens, the long appendage of the connective, the persistent style-base, the delicate hypogynous bristles, etc., of the Bornean plant compel to inclusion in the genus Tetraria. There are, however, strong affinities to Costularia pilisepala. 16. COSTULARIA CLARKE [in Dur. & Schinz, Consp. Fl. Afr. 5 (1895) 658, nom. nud.| in Thiselt.- Dyer, Fl. Cap. 7 (1898) 274; Kew Bull. add. ser. 8 (1908) 125; KUxk. in Fedde, Rep. 46 (1939) 13-76. — Lophoschoenus StaprF, J. Linn. Soc. Bot. 42 (1914) 177; PFEIFF. in Fedde, Rep. 23 (1927) 343. — Fig. 73. Perennial herbs with short woody rhizome. Stems stout, erect, obtusangular, nody. Leaves basal and cauline, linear, broad, coriaceous; sheaths of the cauline leaves tubular. Inflorescence paniculate, loose, consisting of S—10 distant fascicles of partial panicles. Bracts similar to the cauline leaves, but shorter. Spikelets numerous, solitary or 2—3 together, oblong-lanceolate, acute, compressed, usually 2-flowered, both flowers or only the upper one fertile. Rachilla short, not or only shortly produced beyond the upper flower. Glumes distichous, lowest 3-16 empty, small. Hypogynous bristles usually 6, ciliate or plumose. Stamens 3; anthers linear, yellow or ferrugineous; connective + produced. Style slender, continuous with the ovary, the base incrassate, conical or pyramidal, persistent, hispidulous; stigmas 3. Nut trigonous, sessile, indistinctly tessellate, rugulose, or almost smooth. 1974] Cy PERACEAE (Kern) 665 Fig. 73. Costularia pilisepala (STEUD.) KERN. a—a’. Habit, x 4, a". lower branch, nat. size, b. spikelet, x 3, c. uppermost internode of rachilla with sterile glume, d. deflorate spikelet, left lower 3 flower with abortive pistil, right bisexual flower, x 5, e. deflorate upper flower, f. LS of nut with 3 filaments and 6 plumose bristles (after BRONGNIART). 666 FLORA MALESIANA [ser. I, vol. 7° Distr. In the circumscription of KUKENTHAL the genus comprises about 20 spp., 2 of them in S. Africa, 10 in Madagascar, the Mascarenes and the Seychelles, 9 in New Caledonia, and 1 in Malesia. Note. KUKENTHAL divided the genus into 3 subgenera, of which in Malesia only subg. Lophoschoenus (STAPF) KUK. in Fedde, Rep. 46 (1939) 26, characterized by the comose fibrous sheaths of the basal leaves, the densely plumose hypogynous bristles, the long-produced connective of the anthers, and the sessile, obsoletely tessellate nut. 1. Costularia pilisepala (STEUD.) KERN, FI. Mal. I, 5 (1957) 420; Blumea 9 (1958) 234.— Carpha arundinacea BRONGN. in Duperrey, Voy. Coq. Bot. 2 (1829) 169, t. 30, non Costularia arundinacea KUK. — Asterochaete arundinacea KUNTH, En. 2 (1837) 312, p.p. (excl. pl. nov.-caled.); STEUD. Syn. 2 (1855) 155, p-p.; Mig. Fl. Ind. Bat. 3 (1856) 338. — Restio pilisepalus STEUD. Syn. 2 (1855) 256.— Carpha urvilleana GAUDICH. [ex NEES, Linnaea 9 (1835) 300, nom. nud.| ex Boeck. Linnaea 38 (1874) 272. — Lophoschoenus urvilleanus STAPF, J. Linn. Soc. Bot. 42 (1914) 180; Merr. En. Born. (1921) 63; PFEIFF. in Fedde, Rep. 23 (1927) 346, in nota. — C. urvilleana KUk. in Fedde, Rep. 46 (1939) 28; S. T. BLAKE, J. Arn. Arb. 29 (1948) 95. — Fig. 73. Stems obtusely trigonous to subterete, striate, smooth, leafy, 120-150 cm by 4-6mm, the base densely clothed with the brown fibrous remains of decayed leaf-sheaths. Basal leaves long, gradually narrowed into a long triquetrous point, sharply keeled, up to 8 mm wide, with revolute margins; margins and keel serrulate-scabrous; cauline leaves 3, narrower; sheaths 4—5 cm long, brown or fusces- cent, the emarginature at the orifice semicircular, the margins not overlapping. Panicle 35-60 cm long, narrow, loose to rather dense, compound; branches in fascicles of 1-3, erect, slender, compressed- trigonous to ancipitous, smooth; branchlets filiform. Spikelets numerous, mostly solitary, peduncled, oblong-lanceolate, acute, 2-flowered, 6-7 by c. 1’4 mm. Uppermost internode of the rachilla slightly elongated, bearing a sterile glume. Glumes 9-10, the lower ones ovate-lanceolate, the upper lanceolate, c. 5 mm, reddish brown, with acute scaberulous keel, minutely pubescent in the upper part, the lower S—6 empty, shortly awned. Upper flower 9’, the lower one male with abortive gynaeceum, or male, sometimes both flowers “. Hypogynous bristles 6, twice as long as the nut, c. 5 mm, flexuous, attenuate towards the apex, densely plumose up to the top. Stamens 3; anthers linear, with a moderately long, smooth or slightly antrorsely scabrous appendage of the con- nective. Style-base long-conical, pale, hispidulous, persistent. Nut surrounded by the persistent bristles, obovate-elliptic, trigonous, ferrugineous, c. 2mm long (immature). Distr. Malesia: N. Borneo (Mt Kinabalu, common) and West New Guinea (Waigeo I.; P. Rawak; Hollandia and vicinity.; Mt Cycloop). Ecol. In brushwood, in swampy localities, on stony patches of dry open slopes; in New Guinea up to 350 m, on Mt Kinabalu 1650—2640 m. 17. CARPHA BANKS & SOL. ex R.Br. Prod. (1810) 230, p.p.; Boeck. Linnaea 38 (1874) 265-273, p.p.; KUK in Fedde, Rep. 47 (1939) 108. — Fig. 74. Perennial herbs with short woody rhizome. Stems tufted, erect, obtusely trigonous, leafy. Leaves basal and cauline, usually narrow. Inflorescence paniculate. Spikelets oblong-lanceolate, compressed, densely crowded. Rachilla short, straight, not produced beyond the flowers. Flowers in each spikelet 1—2(-3), 3, fertile. Glumes 3—6, distichous, the lower ones Small, empty. Perigone (in Mal.) consisting of 6 plumose bristles. Stamens 3, rarely 2; anthers linear, sulphureous; connective with short, conical appendage. Style slender, continuous with the ovary, the base incrassate, persistent; stigmas 3. Nut oblong, trigonous, tessellate or minutely puncticulate, cuspidate by the per- sistent style-base. Distr. In KUKENTHAL’s circumscription the genus comprises 11 spp., 9 of them endemic in Africa, 1 in Japan, and 1 circum-South Pacific, in Malesia: New Guinea. Levyns(J.S. Afr. Bot. 13, 1947, 81) refers the S. African spp. to the genera Asterochaete and Trianoptiles. 1. Carpha alpina R.Br. Prod. (1810) 230; KUNTH, En. 2 (1837) 322; Sreup. Syn. 2 (1855) 159; Bogck. Linnaea 38 (1874) 269: Hook. Ic. Pl. 13 (1877) 14, t. 1216; BENTH. Fl. Austr. 7 (1878) 381; F.v.M. Trans. R. Vict. n.s. 17 (1889) 35; CLARKE, Kew Bull. (1899) 114; VaLck. Sur. Nova Guinea 8 (1912) 706: PreiFF. in Fedde, Rep. 29 (1931) 178; KUx. in Fedde, Rep. 47 (1939) 112; S. T. Biake, J. Arn. Arb. 29 (1948) 93.— Rhynchospora alpina SPRENG. Syst. Veg. 1 (1825) 195.— Chaetospora alpina F.v.M. Fragm. 9 (1875) 39. — Fig. 74. Stems slender but rigid, often curved, 5—25 cm, obtusangular, sulcate-striate, smooth, 5—30 cm by c. 1 mm, leafy towards the base. Basal leaves rigid, often recurved, flat or slightly canaliculate, obtuse, with slightly scaberulous margins, 1-3 mm wide; 1974] Cy PERACEAE (Kern) 667 Fig. 74. Carpha alpina R.BR. a. Habit, x 74, b. spikelet, x 4, c. deflorate flower, x 6 (a—c SCHODDE 1827). sheaths shining brown; cauline leaves with tubular sheaths. Panicle contracted, 114—5 cm long; branches short, solitary or 2 together, smooth. Bracts 2-3, leaf-like, overtopping the inflorescence; inner ones scale-like. Spikelets few, 2—3 together, unequally peduncled, oblong-lanceolate, finally turbinate, ob- tuse, 1-flowered, 8-12 by 2—2)4 mm. Glumes 4-5, distichous, chartaceous, narrowly lanceolate, stra- mineous, glabrous, almost obtuse, the lower 2-3 empty; above the flower often a narrow empty glume. Hypogynous bristles 6, exceeding the nut, 8-9 mm long, rigid, ferrugineous, plumose almost the whole length, antrorsely scabrous at the top. Stamens 3. Style-base long, incrassate at the base, glabrous, persistent. Nut oblong, 3-4 by 34-1 mm (beak excluded), trigonous, shortly stipitate (stipe c. 4 mm), brown, finely puncticulate, beaked by the persistent style-base, surrounded by the persistent, finally spreading bristles. Distr. SE. Australia, Tasmania, New Zealand, Auckland Islands; KUKENTHAL treats the South American C. schoenoides BANKS & SOL. ex HooK. f. as a variety of C. alpina; it is evidently closely allied and probably a geographical race. In Malesia: W. New Guinea (Mt Wilhelmina) and E. New Guinea (Mt Sarawaket, Mt Amungwiwa, Mt Wilhelm, Kubor Range, Mt Hagen, Mt Sugarloaf, Mt Giluwe, Mt Piora, Mt Dickson, Mt Scratchley, Mt Albert Edward, Mt Knutsford). Ecol. Microtherm; on alpine grasslands, sandy banks of streams, in marshy hollows, and on alpine seepage slopes, often a pioneer in shallow tarns, 3000-4200 m. Vern. Era, New Guinea (Mendi lang.). 668 FLORA MALESIANA [ser. 1, voR@r Excluded Carpha paniculata PuiL. Linnaea 29 (1858) 80; Boeck. Linnaea 38 (1874) 268. = Schoenus rhyncho- sporoides (STEUD.) KUK. in Fedde, Rep. 44 (1938) 19. A Chilean species, not belonging in Carpha, but in Schoenus. RIKLI (in Jahrb. Wiss. Bot. 27, 1895, 567) mentioned it for the Philippines. Obviously he took the name of the author PHILIpP! for the location. 18. LEPIDOSPERMA LABILL. Nov. Holl. Pl. Sp. 1 (1804) 14; KUx. in Fedde, Rep. 50 (1941) 19—S0, 112-128. — Fig. 75-77. Perennial herbs with short, woody rhizome, often stoloniferous. Stems tufted, usually nodeless, terete, compressed, or 3—4-angular. Leaves basal, equitant, similar to the stems. Inflorescence paniculate. Bracts shorter than the inflorescence, the lowest with a short pungent blade, the upper ones gradually shorter. Spikelets oblong-lanceolate. Flowers usually 2—4, the upper one ¢@, the lower one(s) sterile, or 3 by reduction. Rachilla short, straight. Glumes 4—7, spirally arranged, imbricate, scaberulous at the top. Perianth consisting of 6 biseriate, white hypogynous scales, which are narrow, hyaline (therefore easily overlooked in young flowers), under the fruit broadened and incrassate, spongy, ovate to lanceolate. Stamens 3, connective distinctly produced. Style continuous with the ovary, 3-fid; style-base persistent, at first bulbous to dome- shaped, ultimately depressed-hemispherical. Nut oblong or oblong-elliptic, obtusely trigonous, crusty, often smooth and shining when ripe. Distr. About 40 spp., often difficult to delimit, mostly Australian, 1 endemic in New Zealand, 2 in New Caledonia. Fig. 76. In Malesia only: 1. Lepidosperma chinense NEES & MEYEN [Linnaea 9 (1834) 302, nom. nud.] in Kunth, En. 2 (Jan.—June 1837) 320; in Hook. & Arn. Beech. Voy. (July—Aug. 1837) 228; NeEs, Nov. Act. Caes. Leop.-Car. 19, Suppl. 1 (1843) 117; Steup. Syn. 2 (1855) 158; Boeck. Linnaea 38 (1874) 329, p.p.; CLARKE, Fl. Br. Ind. 6 (1894) 676; RiDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 98; StapF & TURRILL in Gibbs, J. Linn. Soc. Bot. 42 (1914) 179; Rip. J. Fed. Mal. St. Mus. 6 (1915) 193; Fl. Mal. Pen. 5 (1925) 167, excl. var.: KOK. Bull. Jard. Bot. Btzg III, 16 (1940) 306, excl. pl. nov.-guin.; in Fedde, Rep. 50 (1941) 123. — Cladium borneense CLARKE in Stapf, Trans. Linn. Soc. II, Bot. 4 (1894) 245. — Cladium arfakense RENDLE in Gibbs, Arfak (1917) 90, p.p. — Mariscus borneensis FERN. Rhodora 25 (1923) 52.— L. striatum (non R.Br.) OHwi, Bot. Mag. Tokyo 56 (1942) 207. — Machaerina borneensis KoyAMA, Bot. Mag. Tokyo 69 (1956) 62. — Fig. 75, 77. Stems slender to rather stout, rigid, (25—)40— 150(—250)cm by 1-3mm, subterete or slightly angular, smooth, few-leaved at the base. Leaves shorter than the stems, almost terete, with a shallow, very inconspicuous groove, rigid, glaucous, pith chambered by numerous approximate transverse partitions; sheaths castaneous, lower ones bladeless. Panicle narrowly oblong, dense, often somewhat interrupted at the base, 3-10 cm long, consisting of several oblong-lanceolate, suberect spikes. Involucral bracts spathe-like, shortly sheathing, blackish brown, rounded at the apex, the lowest ending in a short pungent blade. Spikelets crowded, oblong-lanceolate, 5—6 mm long, 2-flowered. Glumes 5—7, firm, lanceo- late, obtuse, black with ferrugineous margins, the empty ones mucronulate, the upper ones subacute. Hypogynous scales small, connate at the base, whitish, 14-34 mm long. Nut 214-3 by 1-1/4 mm, apiculate, shining ferrugineous, purplish dotted. Distr. Coastal regions of S. China, in Malesia: Malay Peninsula (Perak: G. Kerbau; Trengganu: G. Padang; Pahang: G. Tahan, G. Tapis, G. Tebu; Kelantan: G. Sagi; Malacca: Mt Ophir; Johore: G. Blumut), North to Central Sumatra (Mts Losir, Goh Lembuh, Sago, and Talang), Karimata Is., N. Borneo (Mt Kinabalu), West New Guinea (Arfak Mts). Fig. 76. Ecol. Mountains: in heaths, on open rocks and in marshy meadows, often forming pure stands, (900—)1500—3000 m. In S. China also in rice-fields. Notes. Closely allied to the W. Australian L. striatum R.BR., and hardly distinguishable from the E. Australian L. neesii KUNTH. OuHwl, l.c., referred KANEHIRA & HATUSIMA 13555 and 13678 from West New Guinea (Arfak Mts) to L. striatum. As in both collections the glumes are obtusish and mucronate, and the hypogynous scales very small, they belong in my opinion to L. chinense. Brass 4644 from Papua, Wharton Range, cited in Bull. Jard. Bot. Btzg III, 16 (1940) 306 and in Fedde, Rep. 50 (1941) 123 under Lepidosperma chinense, belongs to Machaerina gunnii (HOOK. f.) KERN. Doubtful Lepidosperma waigiuense STEUD. Syn. 2 (1855) 158; KUK. in Fedde, Rep. 50 (1941) 41. Distr. Malesia: Waigeo I., near West New Guinea, D’URVILLE. Note. I have not seen the type specimen. It is very doubtful whether STEUDEL’s incomplete descrip- tion refers to a species of Lepidosperma. Nothing is said about the presence or absence and the shape of 1974] Cy PERACEAE (Kern) 669 & Lb ! Z a laut Fig. 75. Lepidosperma chinense NEES & MEYEN. a. Habit, x 4, a’. LS of stem, enlarged, b. cluster of | Fig. 76. Range of the genus Lepidosperma LaBILL.. spikelets, c. spikelet, d. pistil, all x5, e. nut with the numbers indicating the number of species in each persistent perianth scales (3 visible) and three fila- | area. Of L. chinense Nees & MEYEN the localities are ments, x 7/4. indicated by dots. 670 FLORA MALESIANA [ser. I, voli Fig. 77. Lepidosperma chinense Nees & MEYEN in stands of stiff clay in mountain heaths (b/angs) of Mt Losir, at c. 2300 m alt. (photogr. VAN STEENIS). a perigone. The description of the nut (“‘achenio... points to Machaerina than to Lepidosperma. See stylo basi persistente tomentoso coronato’’) rather Machaerina glomerata in note on p.698. 19. TRICOSTULARIA Nees in Lehm. Pugillus 8 (1844) 50; in Lehm. PI. Preiss. 2 (1846-1847) 83; BENTH. FI. Austr. 7 (1878) 382; in B. & H. Gen. Pl. 3 (1883) 1064; Pax in E. & P. Pfi. Fam. 2, 2 (1887) 115; KGxK. in Fedde, Rep. 53 (1944) 212. — Fig. 78. Perennial herbs with short rhizome. Stems tufted, erect, terete or trigonous. Leaves basal, rarely 1 or 2 distant on the stem, often reduced to the sheaths; ligule absent. Inflorescence paniculate, spike-like to much branched. Spikelets solitary or clustered, sessile or peduncled, compressed, oblong-lanceolate, 1—2(—3)-flowered, the flower(s) fertile, or the lowest male or sterile. Rachilla persistent, straight, with very short internodes. Glumes 4—6, distichous, mem- branous, l-nerved, keeled, glabrous, pale brown, 2—4 lower ones empty. Perianth consisting of (3—)6 lanceolate to linear, short, flat, hyaline, whitish or finally ferrugineous scales not thickened under the fruit. Stamens 3; anthers linear, with distinctly produced connective. Style continuous with the ovary, caducous, 3-fid. Nut small, obovoid or pyriform, sessile, trigonous, brown with 3 pale ribs, hispid at the top; pericarp thin. Distr. Small genus (c. 5 spp.), all Australian, one species extending to S. Asia. Note. Allied to Schoenus and Lepidosperma; Schoenus is mainly characterized by the elongated, prominently zigzag upper internodes of the rachilla and the reduction of the upper flower in the spikelet; Lepidosperma can be recognized by the thick spongy perianth-scales under the fruit and the glabrous nuts. The circumscription of the genus here accepted is that of BENTHAM in the Flora Australiensis. CLARKE and KUKENTHAL restricted Tricostularia to 3 extra-Malesian spp., and removed T. undulata and T. paludosa to Cladium, Schoenus, or Costularia, which in my opinion is contrary to their natural relationships. 1974] Cy PERACEAE (Kern) 671 Fig. 78. Tricostularia undulata (THW.) KERN. a. Habit, x 2%, b. bract enclosing cluster of spikelets, c. spikelet, x 4,d. anther, x 6, e. deflorate flower, x 8. f. nuts, x 12, g. nut, with partially removed pericarp, showing ovule extending into hairy part of nuts, x 6 (a CaRRICK JC/130, c—e Monp1 163, b, f (left), g BUNNEMEUER 1483, / (right) SINCLAIR & K1iAH SF 40429). 672 FLORA MALESIANA [ser. I, vol. 7° 1. Tricostularia undulata (THW.) KERN, Act. Bot. Neerl. 8 (1959) 267. — Cladium undulatum THw. En. Pl. Zeyl. (1864) 353; CLaRKE, FI. Br. Ind. 6 (1894) 674; RIDL. Mat. Fl. Mal. Pen. (Monoc.) 3 (1907) 86; CLARKE, Ill. Cyp. (1909) t. 83, f. 6-11; Camus, FI. Gén. I.-C. 7 (1912) 152, f. 19, 2-8; MeRR. En. Born. (1921) 62; RipL. Fl. Mal. Pen. 5 (1925) 167; Ui1TIEN, Rec. Trav. Bot. Néerl. 32 (1935) 194; KUx. Bull. Jard. Bot. Btzg III, 16 (1940) 311; in Fedde, Rep. 51 (1942) 162, incl. var. pulchrum KUx.; S. T. BLAKE, J. Arn. Arb. 29 (1948) 98. — Carpha junciformis BOECK. Linnaea 38 (1874) 267. — Lepidosperma zeylanicum LINDL. ex BOECK. |.c. 332. — Chaetospora fimbri- styloides F.v.M. Fragm. 9 (1875) 34. — T. fimbri- styloides BENTH. F1. Austr. 7 (1878) 384. — Schoenus fimbristyloides F.v.M. First Cens. (1882) 128. — T. borneensis RIDL. J. Str. Br. R. As. Soc. n. 59 (1911) 52, nom. nud.— Cladium undulatum var. fimbri- styloides Doin, Bibl. Bot. Heft 85 (1915) 473. — Lepidosperma chinense var. alpinum RIDL. J. Fed. Mal. St. Mus. 6 (1915) 193.— Cladium pulchrum RIDL. |.c. 192; Fl. Mal. Pen. 5 (1925) 167. — Machae- rina pulchra KOYAMA, Bot. Mag. Tokyo 69 (1956) 65.— Machaerina undulata KoyaMa, l.c. 66.— Fig. 78. Forming dense tussocks; roots thick. Stems slender, rigid, obtusely trigonous, sulcate-striate, smooth, leafy only at the incrassate base, 30-90 cm by 1-2 mm. Leaves basal, shorter than to as long as the stems, rigid, strongly ribbed, obtusish, deeply chan- nelled because of the involute margins, scaberulous, 1-3 mm wide. Panicle much branched, rather diffuse to dense, consisting of 4—5 distant to approximate fascicles of branches. Lowest bract similar to the leaves, shorter to slightly longer than the inflo- rescence, upper ones decreasing in length; sheaths short, reddish brown. Branches 2-3 together, un- equal, more or less exserted from the sheath, rigid, flexuous, compressed, scaberulous. Spikelets almost sessile in clusters of 2—8, 1-flowered, 4-5 by 1- 14mm. Glumes 4, oblong-lanceolate, acute, the upper 4—5 mm long, the lower 2 much shorter, c. 2mm. Perianth-scales (3—)6, lanceolate to linear- lanceolate, densely covered with white antrorse short hairs, usually 14 mm long, rarely about as long as the nut. Anthers c. 2mm; appendage of the connective triangular to oblong. Nut obtusely trigo- nous, slightly reticulate-rugulose, castaneous to blackish, with 3 whitish ribs, hispidulous at the top, VA-V4 by 4%-% mm. Distr. From N. Australia to Ceylon, Thailand and Indo-China’, in Malesia: Sumatra (Palembang), Banka, Billiton, Malay Peninsula (Trengganu: Padang Kandis, G. Padang; Pahang: G. Tahan; Setul), N. Borneo & Sarawak, W. & S. Borneo, Labuan, Anambas Is., Karimata Is., New Guinea (NE. New Guinea: Sepik Distr., Green River; Papua: Wassi Kussa R.; West. Distr., Morehead Patrol Post). Ecol. In open sandy heaths, savannah-forests, slightly damp spots on sandstone rocks, 0—1600 m. Vern. Rumput anjai, Billiton. Note. As will be evident from the synonymy the systematic place of this species has often been dis- cussed. CLARKE and KUKENTHAL considered it a member of Cladium subg. Baumea (Machaerina in the present treatment), but there cannot lie its true affinity. The leaves are spirally arranged in a basal rosette, flat hypogynous scales are found nowhere in Machaerina, and also the small 3-ribbed fruit with thin marcescent exocarp puts it out of this genus. As a rule the upper flower in Machaerina is reduced; in Tricostularia undulata the spikelets are always 1- flowered, but I never find a small barren glume above the fertile one. BENTHAM (1878) remarked that there might be question of a distinct genus, but in my opinion shape and texture of glumes, nuts, and hypogynous scales unmistakably point to its con- generity with the other Tricostularia species. Excluded Tricostularia paludosa (R.BR.) BENTH. Fl. Austr. 7 (1878) 382. — Chaetospora paludosa R.BR. Prod. (1810) 233.— Costularia paludosa CLARKE, Kew Bull. (1899) 114; ibid. add. ser. 8 (1908) 47. CLARKE’s record for New Guinea of this Australian species was based on a collection by GIULIANETTI from Papua; this appeared to belong to Schoenus curvulus. In passing I remark that the systematic place of T. paludosa has also been much disputed. CLARKE placed it in Costularia, but it does not fit the modern circumscription of this genus. Its original assignment to Schoenus (resp. Chaetospora) was maintained by KUKENTHAL, who placed it in sect. Helothrix of this genus, but in doing so he made the section very heterogeneous, as T. paludosa widely differs by the reverse distribution of sexes in the spikelet (in Schoenus it is the upper flower which is reduced), and by the short straight rachilla neither elongated nor flexuous between the flowers. KUKENTHAL’s argu- ment against placing in Tricostularia (in which genus according to him the glumes should be spirally arranged) is ineffective, as both in Schoenus and Tricostularia the glumes are distichous. 20. SCHOENUS LINNE, Sp. Pl. 1 (1753) 42; Gen. Pl. ed. 5 (1754) 26; Boeck. Linnaea 38 (1874) 273; BENTH. Fl. Austr. 7 (1878) 352; in B. & H. Gen. Pl. 3 (1883) 1062; CLARKE, Kew Bull. add. ser. 8 (1908) 12; KUk. in Fedde, Rep. 44 (1938) 5; l.c. 161; ibid. 48 (1940) 246. — Chaetospora R.BR. Prod. (1810) 232; Boeck. Linnaea ' The record in Fedde, Rep. 51 (1942) 163: “Formosa: TAKOW” was based on HENRY’s List of Plants from Formosa (1896) 105; it refers, however, to a collection of Schoenus falcatus R.BR. (see OHW1, Mem. Coll. Sc. Kyoto Imp. Un. B18, 1944, 28). 1974] Cy PERACEAE (Kern) 673 38 (1874) 287. — Helothrix Nees, Ann. Mag. Nat. Hist. 1, 6 (1841) 45. — Cyclo- campe STEUD. Syn. 2 (1855) 156; Mia. FI. Ind. Bat. 3 (1856) 339 (sphalm. ‘Cyclo- carpa’); non BENTH. in B. & H. Gen. PI. 3 (1883) 1063. — Lophocarpus BOECK. Allg. Bot. Zeitschr. 2 (1896) 110, non Link, 1795. — Neolophocarpus CAMUs, Fl. Gén. I.-C. 7 (1912) 148, quoad syn. — Fig. 79. Perennials with woody, abbreviated or shortly creeping rhizome, more rarely annuals. Stems erect, or ascending and rooting at the nodes, terete or obtusely trigonous, striate or sulcate, usually smooth. Leaves either all basal, or basal and cauline, linear, often setaceous, canaliculate, sometimes reduced to mucronate sheaths; ligule absent; basal sheaths open, dark-coloured; sheaths of the cauline leaves closed, tubular, often bearded at the mouth. Inflorescence terminal, racemose or paniculate, consisting of some distant fascicles of branches subtended by a leafy bract, or contracted and head-like. Spikelets solitary or clustered, (in the Malesian spp.) compressed, lanceolate to oblong-lanceolate, sometimes falcate, usually few-flowered. Rachilla straight and with very short internodes in the lower part (between the empty glumes), the upper internodes (between the fertile glumes) elongated and prominently zigzag. Glumes distichous, acropetally caducous, I-nerved, keeled, frequently dark-coloured, the lower ones empty, the upper (flower-bearing) ones usually decurrent on the rachilla (‘rachilla winged’), the terminal one often strongly reduced and empty. Flowers in the hollows of the zigzag rachilla, usually bisexual, but the uppermost often more or less reduced (male or functionally male). Perigone consisting of up to 6 filiform or linear-lanceolate, ciliate to plumose or antrorsely scabrous bristles, or absent. Stamens (1—)3(—6), (in Mal. (2—)3); anthers linear, with more or less produced glabrous connective. Style slender, continuous with the ovary, not or hardly dilated at the base, brown, caducous (often the very base remaining on the nut as an apiculus not separated from the nut proper by a constriction). Stigmas 3 (very rarely 2). Nut sessile or shortly stipitate, ovoid, obovoid, or ellipsoid, trigonous, often 3-ribbed, very rarely biconvex, glabrous, or hispidulous in the upper part, smooth, rugulose, or scrobiculate; epidermal cells isodiametric to oblong. Distr. About 80 spp., mainly distributed over SE. Asia and Australia; some spp. extend eastward to New Zealand, New Caledonia, and Micronesia, a few others westward to Europe; a few spp. occur in extratropical South America; S. ferrugineus L. is almost cosmopolitan. The centre of development of the genus is to be found in continental Australia (more than 60 spp.). In Malesia 13 spp., 11 of which occur in New Guinea. Ecol. In Malesia the members of sect. Calostachyi, Paniculati, and Nudicaules are mainly inhabitants of savannahs and savannah forests in the periodically dry regions; the species of sect. Helothrix occur only at high altitudes. Notes. The 9 species LINNAEUS originally assigned to Schoenus have in the course of time all been transferred to other genera by VAHL, BROWN, LINK, NEES, and KUNTH. As more recent authors only two of them (S. nigricans L. and S. ferrugineus L.) recognized as true Schoenus spp., it seems advisable to accept S. nigricans L. as the type species of the genus. R. Brown restricted Schoenus to those species in which the perianth is absent, and created the genus Chaetospora for the species in which the perianth is represented by hypogynous bristles. BENTHAM pointed out that often in the same species the bristles may be present or absent, and consequently he referred all sor tall spp. to Schoenus. His circumscription of the latter genus was accepted by nearly all subsequent authors. Schoenus as circumscribed by BENTHAM and CLARKE is undoubtedly a rather homogeneous group, mainly characterized by the peculiar structure of the rachilla (lower internodes very short, upper ones elongated and zigzag). It is easily discernible from the other Cyperaceous genera in Malesia with exactly distichous arrangement of the glumes as follows: (a) Hypogynous bristles are always absent in Cyperus and Fimbristylis, but often present in Schoenus. (b) Long-plumose bristles are characteristic of Malesian Carpha and Costularia spp., but in Malesian Schoenus spp. only found in S. nitens. 674 FLORA MALESIANA [ser. I, vol. 7° Specific delimitation is extremely difficult throughout the genus. BENTHAM’s remark that further study from the living plants or from specimens gathered in all the different stages of development of the flowers is required, is in force up till the present day. KEY TO THE SECTIONS represented in Malesia 1. Spikelets large, (in the Malesian sp.) 20-25 mmlong.Sp.1 . . . . . . . IJ. Sect. Calostachyi 1. Spikelets much smaller. 2. Leaves reduced to the mucronate or very shortly laminate sheaths, which (in the Malesian spp.) are bearded at the mouth. Glumes with woolly-ciliate margins. Spp. 2-4. . . 2. Sect. Nudicaules 2. Leaves with well developed blades, their sheaths not bearded at the mouth. Glumes glabrous. 3. Inflorescence paniculate, compound or decompound. Spikelets numerous, often falcate. Glumes ferrugineous to brownish. Nut hispidulous at the top. Spp. 5—7 . . 3. Sect. Paniculati 3. Inflorescence racemose or capitate. Spikelets few, straight. Glumes usually purplish or blackish. Nut glabrous. 4. Hypogynous bristles scabrous. Inflorescence racemose, sometimes contracted and head- like. Rhizome cespitose. Spp. 8-11 ‘ . 4, Sect. Helothrix 4. Hypogynous bristles plumose. Inflorescence capitate. Rhizome. creeping. Sp. 12. 5. Sect. Repentes KEY TO THE SPECIES 1. Spikelets 20-25 mm long. Nut 3-3/4 mm long . . 1. S. calostachyus 1. Spikelets and nuts much smaller, the former up to 12 mm long, the latter up to 2 mm. 2. Leaves reduced to mucronate or shortly laminate sheaths (i.e. leaf-blades very short in proportion to the size of the plant, more rarely a few of them elongated’. Sheaths of leaves and bracts bearded at the mouth (more or less glabrescent with age). Glumes woolly-ciliate, at least when young. 3. Stems at least partly with 1—2 leaf-sheaths in or about the middle, very thin, /4—1(—1/4) mm thick. Glumes dark brown, densely puncticulate (gland-dotted). Nut papillose on the angles at the top, otherwise smooth, brown, blackish blotched . . . 2. S. sparteus 3. No leaf-sheaths between the basal sheaths and the bracts. Stems generally thicker, (1 ay )2—4 mm across. Glumes not puncticulate. Nut not papillose, concolorous. 4. Glumes blackish purple. Nut dull stramineous, more or less rugulose . . . 3. S. melanostachys 4. Glumes dark brown. Nut shining milky white, smooth . . 4, S. laevinux 2. Leaves with well-developed prominent blades. Sheaths not bearded at the mouth. Glumes glabrous. 5. Inflorescence paniculate, compound or decompound, at least part of the primary branches with several to numerous spikelets. Spikelets ferrugineous to light brown. Nut setulose on the angles at the top. 6. Nut scrobiculate by the wide, pitted epidermal cells, rufous, only slightly asymmetric. Bristles usually present. Te pt flat and 5—7 mm wide in the basal part, tapering into a very long subulate point. Spikelets —12 mm long, more or less falcate. Inflorescence stiffly erect . . . 5. S. falcatus Ds feta filiform almost from the base. Spikelets 5—6 mm long, not or hardly falcate. Inflorescence more or less drooping . . 6. S. punctatus 6. Nut quite smooth (epidermal cells obscure, not pitted), milky white, distinctly asymmetric. Bristles absent. Spikelets 5-6 mmlong . . . . 7. §. delicatulus 5. Inflorescence racemose or almost so, or capitate; primary ‘branches when present with 1-3 spikelets. Spikelets purplish, only in S. maschalinus often pale. Nut glabrous. 8. Hypogynous bristles absent. Spikelets9-11mmlong . . . . IIL. S. longibracteatus 8. Hypogynous bristles present. Spikelets up to 8 mm long, often much shorter. 9. Hypogynous bristles plumose in the lower part. Rhizome creeping; stems distant or wigre tufted. Inflorescence capitate. . . i . S. nitens 9. Hypogynous bristles scabrous, not plumose. Rhizome cespitose, stems densely tufted. nice racemose, in S. setiformis strongly contracted and head-like. 10. Inflorescence dense, almost acer (occasionally with a fascicle of 1-3 spikelets lower down). Nut smooth, brown . . . . . 10. S. setiformis 10. Inflorescence loose, racemose. "Nut slightly scrobiculate, whitish. 11. Spikelets 6-8 mm long, purple. Anthers 2—3 mm. Nut 4 2 mm long, Stems erect, rigid, 10-40 cm tally