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PTS. 1 Wer AOC Se eA Sere we whines aks wa dads s adaaiepumkee i iuda sccenouaabeeoanstvorsedl & Chua, L.S.L and L.G. Saw A new record of Dryobalanops beccarii (Dipterocarpaceae ) from Peninsular UMMA re Net it de dao a Sete ee ee Re a piositecn us pean das vn noxteh@eunden od sar onee cag iars ntekogs sogeepaapau Chua, L.S.L and L.G. Saw Conservation notes on Vatica yeechongii (Dipterocarpacae) from ae nae AOA erie Dh ou inh kus ook avon nine ules Oba08 Sevinoncpeplgnwws Dasennnasnetemnanwakasvodmeseand Chung, R.C.K., E. Soepadmo and A.L. Lim. The significance of pollen morphology in the taxonomy of Grewia and Microcos (Tiliaceae) in Peninsular Malaysia and Borneo ....................0.0sss000secceeee ee 02-239 Gusman, G. Arisaema fimbriatum (Araceae) and its intraspecific Variation ................eseeseeeeerereee ee e201 Hassan I. Amote On ordia premnifolia Rid). (Borapinaceat) «isi. ac<cccooeetessciencdpcvadsgense so vapess ser LOD Hay, A. and C. Herscovitch A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ...........ccceceeeeeeeees 27 Herscovitch, C. and A. Hay An unusual new species of Homalomena (Araceae) from New Guinea ...............::ceee 31 Hilliard, O.M., B.L. Burtt and M.H. Bokhari Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from Borneo...................0068 35 Kiew, R. Begonia peninsulae (Begoniaceae), a confused species from Malesia ..................::00000 61 Kiew, R. and C. Geri Begonias from the Bau limestone, Borneo, including a new SpeCi€s ..............:.::eeeeeeeeeees 113 Kiew, R. and I.M. Turner te ity Plants CnieiIC 10 wai APOE 1 55.0 yd ned a escies dn tod dctence c dbdvcenereetenereescdaveen as Bansel 1S Kloet, S.P. Vander Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ...........0:.:ccccesseeeeeees 163 Lee, S., A. Samsuri, P. Leong, Ali Ibrahim and A.-T. Gwee A botanical survey of Chek Jawa, Pulau Ubin, Singapore ..................sscscssessscccsestee se ees 271 Mabberley, D.J. New species of, and other notes on, Chisocheton and Walsura (Meliaceae)................... 189 Middleton, D.J. A new species and a new combination in Bornean Kopsia (Apocynaceae: APOCYNOIGEAREC)..........veseaceesianniesnantbuasivniiosaicncinesieh niniiov ch Geoedeeitns tual tse Middleton, D.J. A revision of Dyera (Apocynaceae: Rauvolfioideae) ...............csscccecccceeeeeesssreeee ee ee Pearce, K.G. New species and varieties of Symplocos (Symplocaceae) from Borneo...............:::0++ Pearce, K.G. Five new Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia ..................000 Skornickova, J., M. Sabu and M.G. Prasanthkumar A new species of Curcuma L. (Zingiberaceae) from Mizoram, India ...............eeee Skornickova, J., M. Sabu and M.G. Prasanthkumar Curcuma codonantha (Zingiberaceae) — a new species from the Andaman Islands, Tinta ....ceccacaccovecscvsdeccsaeescccencsxceensnvazses<wawebuuystadaes gone ee ace ne Ese ee ee eee aaa ea Vermeulen, J.J. and P. O’Byrne New species and new records of Southeast Asian Bulbophyllum (Orchidaceae)......... Vermeulen, J.J. and P. O’Byrne Six new species of Bulbophyllum (Orchidaceae) from Sulawesi ............:::cceeeeeeeeeree es Wong, C., G. Argent, R. Kiew, Ohn Set and Y.Y. Gan The genetic relations of Musa species from Mount Jaya, New Guinea, and a reappraisal of the sections of Musa (Musaceacy Tittt.J2 is. .nceresscsensteoastenes en ate te Zahid, M.S. Three new species of Porterandia (Rubiaceae) from Mount Kinabalu, Borneo.......... Obituary Mohamad Shah bin Hj Mohamad Noor - R.Kaiew............ 00. ...ccscccccececccceeeeee ee eeceee ones Dr Chang Kiaw Lan - K.M WOmtng.......020co: oe0ss0% «050 s00nse oysodnvsxsntranaaeen ian ...209 sone 219 Raa) oT | ae eee aC ear Ale EM hone Me ly SE NOAA T EO ANG Ha AER RW 0 g te AB Pye a tie A eA 0 oe MON WAR an sot as LOA LEON YALL GNA PO tes haben ae . . tte oe Aver Sneath © . ms ‘ a ° ‘ . a . ENE CO A AMENIO TT stants ote reed eg ta . ha is ae thous ax ee Sel raed “ “i a eee ly nner alg if lo es pale ge sees «5 hie at ae ee nee eta! oe Cae yar ah piso fuente 8 IRAE Sat iY PEDO A 08 AM CORAM ME AA A Bf RS PLN CHE NAO WA ah OO Woo OF hee ip ep wt ies ener? yp aig EA eat PO tal gat oOo 2 tag yp a me ee ee ers ere ee ne eee eee eae es eee GC ee eC i. et We ie ae " mnths Nene Mame Nr: NO ee Teene). eee 434 ee ees ee eee - ; ete A uA scold, sone A Qe et 5 fame eve ent 0374 OS eee A gin ee AE. at NO ieee OO a) synthe Oe OE Ne OE ge ‘ Ar ye oom tte he hace he de Se a Se eee rd Se OM Ar emt Sone (As ae en IS ek Gee Ie te July 2003 art 1). L. 55 (P% VO 4 = 5 res ? i ; NATIO Gardens Cluny va i R ny I e Z ani re Bot Singapo THE GARDENS’ BULLETIN The Garden’s Bulletin Singapore publishes papers on plant taxonomy (including revisions), horticulture, phytogeography, floristics, morphology, anatomy and related fields with emphasis on plants in the West Malesian region. EDITORIAL BOARD Dr Ruth Kiew (Editor) Singapore Botanic Gardens Dr P.Y. Tan (Assist. Editor) Singapore Botanic Gardens Dr S. C Chin Singapore Botanic Gardens Dr M.J.E. Coode Royal Botanic Gardens Kew, U.K. Dr R.T. Corlett University of Hong Kong Hong Kong Dr M.C. Roos National Herbarium of Netherlands Leiden University Branch, Netherlands Dr E. Soepadmo Forest Research Institute Malaysia Kepong, Malaysia Dr W.K. Tan Singapore Botanic Gardens The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. Neither the National Parks Board not the Editorial Board is responsible for the opinions or conclusions expressed by the contributing authors. The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including postage. Overseas subscribers are required to make payment in the form of bank drafts or international money orders in Singapore currency payable to National Parks Board, Singapore. Instructions for contribiting authors are found on the inside backcover. ‘ * 0 3 2003 ‘ et acl The Gardens’ Bulletin Singapore VOL. 55 (Part 1) July 2003 ISSN 0374-7859 CONTENTS Chua, L.S.L. and L.G. Saw A new record of Dryobalanops beccarii (Dipterocarpaceae) from Fane VSB Bs Sete fas ees tetrad! oc aes:! use 1, oily. dgee iy als CMa aweeeseceoe d Chua, L.S.L. and L.G. Saw Conservation notes on Vatica yeechongii (Dipterocarpaceae) from Be eRe Nh Nts os os dace pee damnns weg dees give ndeeeoebiod bani es ‘4 Chew, W.-L. SLMS th WIRMeSIal PigeraCege Beis tihs anys vein ches iencs'ssl bite qepehgemedovenxdeeoveheers.y0004 3 Hay, A. and C. Herscovitch A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ............... 27 Herscovitch, C. and A. Hay An unusual new species of Homalomena (Araceae) from New aN a BS a cn ae cctas as atid aanicd apuumpt nc cksen ipiscsie davcpibaiine 31 Hilliard, O.M., B.L. Burtt and M.H. Bokhari Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from EE re TURE Node tees et Ae ee A EP = Kiew, R. Begonia peninsulae (Begoniaceae), a confused species from Malesia..................04. 61 Middleton, D.J. A new species and a new combination in Bornean Kopsia AR ON NEN IN Fetes Niet nso a ia lsvannepecigeraneenasesseusvencasdncwnnese 65 Pearce, K.G. New species and varieties of Symplocos (Symplocaceae) from Borneo................ 69 Pearce, K.G. Five new Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia... sisccccck cited escarole Ste a 73 Skornickova, J., M. Sabu and M.G. Prasanthkumar A new species of Curcuma L. (Zingiberaceae) from Mizoram, India.................... 89 Wong, C., G. Argent, R. Kiew, Ohn Set, and Y.Y. Gan The genetic relations of Musa species from Mount Jaya, New Guinea, and a reappraisal of the sections of Musa (Musaceae) .............cscssecccsesseeeee eee iD] Kiew, R. and C. Geri Begonias from the Bau Limestone, Borneo, including a new species ................. 113 Vermeulen, J.J. and P. O’Byrne New species and new records of Southeast Asian Bulbophyllum (Orchidaceae) -.2.2i0c ssc csceceswsieattderstarresliecevave fess pest Le ee 125 Kloet, S.P. Vander Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ................... 163 Obituary Mohamad Shah bin Hj Mohamad Noor ................. 00. seessseeeeeeeeeeeee RY Kiiew-165 Date of Publication: 17 September 2003 Published by National Parks Board Singapore Botanic Gardens 1 Cluny Road Singapore 259569 Printed by Print Dynamics (S) Pte Ltd Garden’s Bulletin Singapore 55 (2003) 1-6 A New Record of Dryobalanops beccarii (Dipterocarpaceae) from Peninsular Malaysia L.S.L. CHUA AND L.G. SAW Forest Research Institute Malaysia, Kepong 52109, Kuala Lumpur, Malaysia Abstract Dryobalanops beccarii Dyer, Dipterocarpaceae, first described and recorded from Borneo, has now been documented from Johore, Peninsular Malaysia, where it grows on ridges in several compartments in the Panti Forest Reserve, Kluang Forest Reserve, and the Labis Forest Reserve. It is a large tree frequently reaching 40 m height and more than 1 m diameter. Introduction Two species of Dryobalanops, D. aromatica and D. oblongifolia, were recorded for Peninsular Malaysia (Ashton, 1982). The genus Dryobalanops is clearly distinguished from other dipterocarp genera by the presence of five wing-shaped, subequal fruit calyx lobes and closely parallel venation on its leaf blade. In Peninsular Malaysia, trees of both species are characterized in the field by scaly bark, peeling in large, irregular, thin flakes. In D. oblongifolia, leaves are oblong or oblong-lanceolate and the fruit calyx lobes are shorter than the nut, while in D. aromatica, leaves are oval or sub-ovate and the calyx lobes are longer. During a plant diversity enumeration exercise in the Virgin Jungle Reserves (VJR) network in Peninsular Malaysia, Ang Khoon Cheng and Kamarudin Saleh discovered in Kluang Forest Reserve (FR) and the VJR in Labis FR, Johore, trees that had leaves that closely resembled D. aromatica but the bark was distinctly different in that the outer bark was scaly, very shallowly fissured and had a laminate inner bark. It was clear that these populations belonged neither to D. aromatica nor D. oblongifolia but their identity could not be confirmed in the absence of flowers and fruits. A dipterocarp mast-fruiting episode occurred in Peninsular Malaysia during the second and third quarters of 2002. Taking advantage of this cyclic event, teams from the Botany and Seed Technology Units, Forest Research Institute Malaysia (FRIM), launched an extensive fruit collection exercise, targeting species that are rare and/or threatened. Fruiting trees in several of these Dryobalanops populations i) Gard. Bull. Singapore 55 (2003) enabled materials for herbarium specimens and fruits for ex situ planting to be collected. The identity of the species could be confirmed and proved to be D. beccarii Dyer. A full description of the species based on characters from these populations is provided below. Dryobalanops beccarii Dyer J. Bot. 12 (1874) 100; Ashton, Fl. Malesiana I (9) (1982) 375. Type: Malaysia, Sarawak, Matang, Beccari PB2944 (K, lecto). Large tree to 30 m tall, 63.7 cm diam., with small buttress to 1.5 m tall. Bark shallowly fissured to scaly, flaky and peeling off in pieces on the lowest part of the trunk immediately above the buttress, greyish-white; inner bark laminate, reddish-brown; sapwood yellowish brown, exuding resin when cut. Twigs 3 mm diam., glabrous, drying almost black. Leaves thickly coriaceous, glabrous, drying brown above, dark brown below; petiole 10-17 mm long, 1 mm thick, glabrous, drying almost black; blade broadly ovate, 3.5—6.3 x 2.4—3.7 cm, apex tapering to a 7-mm long acumen, base broadly cuneate to rounded, margin thickened, midrib prominently raised below, sunken above, secondary veins numerous, closely parallel, faint on both surfaces, smelling of camphor smell when crushed. Flowers not seen. Fruit calyx glabrous, shallow cup tapering gradually and cuneate at the pedicel, lobes 5, equal, oblong- spathulate, base to 2 mm, yellowish green when fresh, 4.0-6.5 x 0.6—1.0 cm, veins conspicuously raised on both surfaces, nut ovoid, glabrous, yellowish green when fresh, to 1.4 x 0.9 cm, style remnant 1 mm long, free from calyx except the base. Vegetatively, the leaf morphology of Dryobalanops aromatica and D. beccarii is very similar and hence it is difficult to differentiate them when sterile. However, the bole character as described above can readily distinguish the two species. In addition, the fruits of D. beccarii are different from D. aromatica (Table 1). Table 1: The bark and fruit characters of Dryobalanops aromatica and D. beccarii D. aromatica D. beccarii Outer bark long, thin, recurved, irregular shallowly fissured to scaly scales Inner bark fibrous laminate Fruit calyx size (cm) 4.0-6.0 x 0.8-2 4.0-6.5 x 0.6—1.0 Max. nut size (cm) 3% 1 14x09 Specimens examined: Johore: Compartment 6B in Panti FR, Damahuri S. FRI 43567, 7 August 2002 (KEP), Ang K.C. FRI 43651, 13 August 2002 (KEP); Kluang FR, Ang K.C. FRI 43653, 14 August 2002 (KEP). Dryobalanops beccarii in Peninsular Malaysia 3 Geographical distribution in Peninsular Malaysia In Peninsular Malaysia, Dryobalanops beccarii is known only from Johore from three localities, namely in compartments 6A, 6B, 7, 8A and 8B, Panti FR; in compartment 132, Kluang FR; and in compartment 841 in VJR, Labis FR. Prior to this discovery, D. beccarii was only known to occur in Borneo on leached sandy soils on coastal hills and inland ridges below 700 m (Ashton, 1982). Like D. aromatica and D. oblongifolia, it shows the same pattern of geographic distribution in southern Johore and west Borneo. Corner (1958) has drawn attention to the similarity in the floras of these two regions, which he termed the Riouw pocket. Ecology In Johore, this species is found in the lowland dipterocarp forest at about 75 m altitude, confined to ridges. They are large trees in the emergent and main canopies. On some ridges, it is co-dominant with Shorea curtisii Dyer ex King. In Kluang FR, it grows sympatrically with D. aromatica on steep hill slopes. In all the three sites, D. beccarii grows gregariously. In Borneo, the species is locally abundant on leached sandy soils on coastal hills and inland ridges below 700 m. Germination Three seed batches of mature fruits were collected from five trees in Panti FR and six trees in Kluang FR in August 2002 and germinated with the seed wings removed in the FRIM nursery (Table 2). Germination is deemed to begin when the radicle protrudes through the seed coat. Germination is epigeal. The hypocotyl elongates to c. 1.8 cm. The emergent cotyledons are bilobed, fleshy and unequal. The first two leaves are opposite. In some seedlings, branching may begin at the first node. Mean percentage germination was 73 + 12 s.d., the percentage varying from 62 to 86 (Table 1). All three seed batches showed a typical germination sigmoid curve (Fig. 1). The germination period for Dryobalanops beccarii seed batches was longer than that of D. aromatica (100% within 20 days, Ng 1991, Siti Asha er al. 1995, Tamari 1976) and D. oblongifolia (100% within 10 days, Siti Asha et al. 1995 and 33 days, Ng 1991). In addition to the longer germination period, the three seed batches of D. beccarii achieved a maximum germination percentage of only 86. Conservation notes In Panti FR, ground checks in August 2002 indicated that the boundary of compartments 6B, 8A and 8B, which forms part of the forest reserve boundary, is adjacent to land being developed for small-scale agriculture activities. Because agriculture has a propensity for expansion, it is recommended that the forest office regularly monitors this section of the boundary to ensure its integrity and prevent future conflict. Compartments 6B, 8A and 8B had been logged in 1982/83. These 4 Gard. Bull. Singapore 55 (2003) compartments are neither VJR nor are they above 1000 m elevation and hence have no legal protected status and may be licensed out for timber harvesting. Populations in Labis VJR and in Compartment 132, Kluang FR have a much brighter prospect as these forests are virgin and VJR and water catchment areas have protected status. Studies are currently being undertaken to determine the species’ population size and genetic structure. In view of the above scenario, Dryobalanops beccarii in Peninsular Malaysia is given the 2001 IUCN category of Endangered EN A3dB1. D. beccarii falls within the A3 category (there will be a projected or suspected population size reduction of 250% within the next 10 years or three generations, whichever is the longer);’d’ (the reduction can be based on actual or potential levels of exploitation); and B1 (its geographic range, in the form of the extent of occurrence, is estimated to be less than 5000 km’). Table 2. Percentage germination and survival in three seed batches of Dryobalanops beccarii Batch No. 2002-0502 (FRI 43567) 2002-0545 (FRI 43653) FRI 43651 Locality Panti FR, Compartment Kluang FR Panti FR, 6B Compartment 6B Collector Damahuri S Ang KC Ang KC Date planted 12 August 2002 16 August 2002 16 August 2002 No. sown 4] 617 78 Max. germination (%) 71 86 62 Germination period 4—32 4-3] 2-31 (days) No. days to achieve 6 9 23 50 % germination % seedling survival 22.8 72.8 100 after 1 month Dryobalanops beccarii in Peninsular Malaysia 5 100 Germination (%) Nn S Days —e— 2()02-0502 —m— 2002-0545 —&— FRI 43651 Figure 1: Germination curve for three different seed batches of Dryobalanops beccarii Acknowledgements We thank Hj. Ros and the foresters and rangers of the South Johor District Forest Office and Hj. Ahmad Fekri and staff of the Kluang District Forest Office for the permission and field assistance provided; Peter S. Ashton for confirming species identity; Kamarudin Saleh and Ang Khoon Cheng (FRIM) who first noted the presence of Dryobalanops beccarii and together with Damahuri Sabari, Mohd. Aidil Nordin, Mustapa Data and Ayau Kanik collected sufficient data for confirmation. Financial support from the Flora Malaysiana Centre Fund, Ministry of Primary Industries, Malaysia under Project 6: Conservation Monitoring System for Threatened Plants is gratefully acknowledged. 6 Gard. Bull. Singapore 55 (2003) References Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana. Ser. 1, 9: 237-552. Corner, E.J.H. 1958. An introduction to the distribution of Ficus. Reinwardtia 4: 15-45. IUCN, 2001. JUCN Red List Categories. Version 3.1. 1UCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. Ng, F.S.P. 1991. Manual of Forest Fruits, Seeds and Seedlings. Malayan Forest Records No. 34, Vol. 1. Forest Research Institute Malaysia. Kuala Lumpur. 400 PP. Siti Asha A.B., K.C. Ang, and S. Zaiton. 1995. Records of germination trials of dipterocarp seeds at FRIM, Kepong, 1979-1994. Research Data No. 4. Forest Research Institute Malaysia. Kuala Lumpur. 161 pp. Tamari, C. 1976. Phenology and seed storage trials of Dipterocarps. FRI Research Pamphlet No. 69. Forest Research Institute Malaysia. Kuala Lumpur. 73 pp. Garden’s Bulletin Singapore 55 (2003) 7-12 Conservation Notes on Vatica yeechongii (Dipterocarpaceae) from Peninsular Malaysia L.S.L. CHUA AND L.G. SAW Forest Research Institute Malaysia Kepong 52109, Kuala Lumpur, Malaysia Abstract The conservation status of Vatica yeechongii L.G. Saw, recently discovered from Peninsular Malaysia, is described. Introduction Vatica yeechongii was discovered along Sungai Tekala in Sungai Tekala Recreational Forest, which is located in Sungai Lalang Forest Reserve, Selangor, Peninsular Malaysia, 3° 03.485’ N, 101° 52.373’ E, alt. 79 m asl (Saw, 2002). Several weeks after its discovery in the type locality, a population was found in Compartment 4, Setul Forest Reserve, along the upper stream of Setul River, Negeri Sembilan (2°46.937’ N, 101°55.069’ E). The population was located in the forest margin of a logged-over forest adjacent to the trunk road from Mantin to Seremban town at an altitude of c. 192 m. Vatica yeechongii is very distinct in leaf characters and nut shape (Saw, 2002). The leaf blade is oblanceolate, 44-84 x 10—6.5 cm, thickly coriaceous, bullate above with 28—30 pairs of veins. The nut is ovoid. Ecology A medium-sized tree 8—15 m tall and 9-13 cm diameter, V. yeechongii occupies the dense understorey of lowland dipterocarp forest. In the type locality, nine trees were found on gentle earth banks of a free flowing stream near the public campsite. There were no trees further upstream. Details of its growth pattern are described in Saw (2002). Large dipterocarps such as Shorea leprosula Miq., S. parvifolia Dyer, S. macroptera Dyer, S. acuminata Dyer, S. pauciflora King, S. ovalis (Korth.) Blume, S. bracteolata Dyer, S. dasyphylla Foxw., Dipterocarpus crinitus Dyer and D. cornutus Dyer dominated the forest. Non-dipterocarps included /ntsia palembanica 4 Gard. Bull. Singapore 55 (2003) Mig. (Leguminosae), Saraca cauliflora Baker (Leguminosae), Terminalia subspathulata King (Combretaceae), Pometia pinnata J.R. Forst. & G. Forst. (Sapindaceae), Canarium patentinervium Mig. (Burseraceae), Elateriospermum tapos Blume (Euphorbiaceae), Palaquium gutta (Hook.f.) Baill. (Sapotaceae), Pimelodendron griffithianum (Miull.Arg.) Benth. (Euphorbiaceae), Endospermum diadenum (Miq.) Airy Shaw (Euphorbiaceae), Dyera costulata (Miq.) Hook.f. (Apocynaceae), Artocarpus lanceifolius Roxb. (Moraceae), A. scortechinii King, Gironniera subaequalis Planch. (Ulmaceae), G. nervosa Planch., Payena lucida A. DC. (Sapotaceae), Streblus elongatus (Miq.) Corner (Moraceae), Litsea costalis (Nees) Kosterm. (Lauraceae), Gynotroches axillaris Blume (Rhizophoraceae), Cratoxylum formosum (Jack) Dyer (Guttiferae), Scaphium macropodum (Miq.) Beumée ex Heyne (Sterculiaceae) and Pternandra echinata Jack (Melastomataceae). At Setul, the forest margin vegetation gradually gave way to a forest having a main canopy to c. 20 m tall, dominated by Shorea leprosula, S. macroptera, S. multiflora (Burck) Symington, S. acuminata, §. parvifolia, Parkia speciosa Hassk. (Leguminosae), Pometia pinnata, Campnosperma auriculatum (Blume) Hook.f. (Anacardiaceae), Artocarpus scortechinii, Pterocymbium tinctorium (Blanco) Merr. (Sterculiaceae) and Alstonia angustiloba Miq. (Apocynaceae). The understorey lacked palms and climbing rattans and the forest floor was covered with abundant dipterocarp seedlings. Herbs such as Tacca integrifolia Ker Gawl. (Taccaceae) and Costus speciosus (J. KOnig) Sm. (Costaceae) were present in scattered numbers. At least 70 V. yeechongii trees were found on steep slopes of riverbanks and on slopes away from the riverbanks. Trees here averaged 9.8 m tall (range 2—21 m), had a diameter at breast height c. 8.2 cm (range 1.6—18.8 cm) with a spreading crown. Its bark was whitish and smooth with horizontal rings. Those growing along the riverbanks had poor bole form and multiple branching was common while those further away had a better bole form. Germination Mature fruits were available from at least six trees in Sungai Tekala and one tree in Setul Forest Reserve from June to August 2002. Three seed batches, collected from two localities at different dates, were germinated with the seed wings removed and observed in the Forest Research Institute Malaysia, FRIM (Table 1). Germination is deemed to begin when the radicle protrudes through the seed coat. Germination of Vatica yeechongii is epigeal. The emergent cotyledons are bilobed, fleshy and equal. The hypocotyl elongates to c. 4.8 cm tall and the first two leaves are opposite. Mean percentage germination was 63.011 +31.207 s.d., the percentage varying from 42.1 to 98.9. All three seed batches showed a typical sigmoidal germination curve. Germination percentage on the first day of germination, i.e. day 5, was 38% for seed batch 2002-0510 and 18.7% for seed batch 2002-0503. Despite this promising Vatica yeechongii 9 Table 1. Percentage germination and survival of three seed batches of Vatica yeechongii Batch No. 2002-0503 (FRI 46613) 2002-0510 2002-0542 (FRI 46668) Locality Sungai Tekala, Sungai Sungai Tekala, Setul Forest Reserve, Lalang Forest Reserve, Sungai Lalang Negeri Sembilan Selangor Forest Reserve Collector Chung R Saw LG Chan YC & Ayau K Date planted 14 July 2002 28 July 2002 20 August 2002 No. sown 107 fie, 89 Max. germination (%) 42.1 48.1 98.9 Germination period 5-37 5-25 2-25 (days) No. days to achieve 45 32 2 40% germination % seedling survival 5.9 97.3 40 after 1 month beginning, however, these two seed batches took a longer time to achieve the 40% germination mark compared to seed batch 2002-0542 (Fig. 1). Preliminary germination results suggest that seed viability is higher in the population at Setul Forest Reserve but this could be due to a variety of factors e.g., fruit maturity, seed vigour, timing of fruit collection, tree health, pest infestation on fruits, etc. In addition, population sizes could also affect gene flow; a larger population is probably more genetically diverse. In situ Conservation Vatica yeechongii is so far known only from these two localities. Preliminary observations on population size suggest that this species is a rare endemic. The population at Sungai Tekala lies in recreational forest. Under the National Forestry 10 Gard. Bull. Singapore 55 (2003) 100 80 S60 = — = 40 = 5 Oo 2 0 12.3.4. 5 6.7, 8,9 1011 12.13)14 15.16.17, 18,19-98 2128 Days —o— 20()2-0542 —a— 2002-0503 —a— 2002-0510 Figure 1: Cumulative germination percentage for different seed batches of Vatica yeechongii Act, no harvesting of resources is permitted in such forests as they are strictly meant for recreation. Population loss will only take place if the Forest Reserve or parts of the reserve, through de-gazettement, are converted to non-forestry land use. For Sungai Tekala, such action is, however, highly unlikely as the area is in the vicinity of the water catchment for the Semenyih Dam, a principal dam providing water supply to Kuala Lumpur. In terms of man-made threats to the reduction in population size and area of occupancy, there are no immediate foreseeable pressures either from habitat loss or degradation or timber harvesting. We note that the practice of clearing undergrowth in areas designated for recreational use may affect regeneration and survival of populations. Careful observa tions on the individuals here suggest that the species has no potential commercial value, as trees do not attain significant harvestable size at maturity and has a poor bole form. The Forest Department of Central Selangor District has been alerted to the presence of the new species and steps are being taken to conserve the population. The population at Setul faces a different scenario. Although it lies in the forest reserve, it is adjacent to the main road. It has some form of protection by virtue of it being in Forest Reserve, but the fact that it lies very close to the road could pose Vatica yeechongii ‘il future conservation conflicts when road expansion is required. It is therefore recommended that more stringent protection measures be provided for this population. In view of the above scenario, Vatica yeechongii should be given the 2001 IUCN category of Critically Endangered CR A2cB1. It falls within the A2 category (inferred or suspected population size reduction of >80% over the last 10 years or three generations, whichever is longer); ‘c’ (a decline in area of occupancy, extent of occurrence and/or quality of habitat) and B1 (its geographic range, in the form of the extent of occurrence and area of occupancy, is estimated to be less than 100 km?). An attempt to estimate the population size of the species in each locality is currently being undertaken. From the viewpoint of conservation, the discovery of a new dipterocarp species in Selangor and Negeri Sembilan is remarkable because in comparison to other states, these two states encounter higher conflict between socio-economic development and conservation. Both populations face an acute need for non-forestry related land development and hence there exist tremendous pressures leading to habitat loss and degradation. The fact that a new species was discovered in this current era of diminishing forest areas merely shows that we do not know enough about our plant diversity and that long-term field observations are necessary. This can only take place with concerted botanical research programmes having full national, financial and personnel support. Ex situ Conservation One month after potting, 215 seedlings from the three batches survived, each bearing at least two pairs of leaves. Distribution to other gardens and arboreta is envisaged in the near future. Acknowledgements We thank Azid Adam, Mohd. Jantan, Borhan Mat Saad and Abu Kasim Omar, Selangor District Forest Office, Cheras, Kuala Lumpur and Hj. Mohamad Hj. Ismail of the Negeri Sembilan Barat District Forest Office for the permission and field assistance provided; Chan Yee Chong who discovered the species; Damahuri Sabari, Mohd. Aidil Nordin, Markandan Moorthy and the support staff of Botany Unit, FRIM, for the field collection and ex situ maintenance of plants. Financial support from the Flora Malaysiana Centre Fund, Ministry of Primary Industries, Malaysia under Project 6: Conservation Monitoring System for Threatened Plants is gratefully acknowledged. 12 Gard. Bull. Singapore 55 (2003) References IUCN, 2001. JUCN Red List Categories. Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK. Saw, L.G. 2002. A new species of Vatica (Dipterocarpaceae) from Peninsular Malaysia. Gardens’ Bulletin Singapore. 54: 247-251. Garden's Bulletin Singapore 55 (2003) 13-25 Studies in Malesian Piperaceae 3! WEE-LEK CHEW 12 Robinson Street Chatswood NSW Australia Abstract Thirteen species, mostly from New Guinea, are dealt with in this paper, of which 12 other species previously considered distinct are reduced to synonymy. No new taxon is proposed. Introduction This is part of a series of papers, albeit variously titled, resulting from continuing work initiated some 35 years ago. As mentioned previously (vide Chew, Journal Arnold Arboretum 53 (1972) 1-25), the work began by way of an undertaking to identify the collection of Piper made by the Royal Society Expedition to the Solomon Islands in 1965. This necessitated reference to the New Guinea species of Piper which then extended to an assessment of the taxonomic status of the more than 150 species described by Casimir de Candolle from that region and surrounding islands. I was encouraged to this much larger task by the fact that the type materials from the former German New Guinea held in the Berlin Herbarium are still extant, having been removed to safety prior to the destruction of the Herbarium, and assisted further by the availability of a vast collection of high quality specimens made since about 1945. Thus, I was enabled to determine (at the least to assess, in a few cases) the taxonomic status of some 108 of de Candolle’s species and to identify over 1,000 collections from New Guinea and the surrounding region. This resulted in the reduction of 75 Candollean names to synonyms. The assignment to synonymy of this large number of de Candolle’s species should not detract from the otherwise excellent work accomplished by this great taxonomist: he did not have the advantage of the availability of the vast range of contemporary collections that I have. Apart from the floristic account of the alpine species by van Royen in The Alpine Flora of New Guinea 3 (1982) 1267-1287, I am not aware of any major study having been undertaken on the New Guinea species since commencement of my work. ‘Following: Blumea 20 (1972) 145-149 and Blumea 37 (1992) 159-164. 14 Gard. Bull. Singapore 55 (2003) There still remains quite a number of Candollean species requiring determination of their taxonomic status. Additionally, large quantities of unnamed materials in many herbaria await identification, some of which may represent undescribed species. As this is a large undertaking, it may need the assistance of future botanists to continue the work. Unless otherwise stated, all collections cited in this paper have been seen and studied by me personally. 1. Piper abbreviatum Opiz in Presl., Rel. Haenk. 1 (1828) 157; Quisumbing, Philip. J. Sci. 43.1 (1930) 58, fig. 24 & 25; Chew, J. Arn. Arb. 53 (1972) 1. Type: Philippines, Luzon: Haenke s.n. (holotype PR). Synonyms: Piper minus K. Schum. & Lauterb., Fl. Schutzgeb., (1900) 258, syn. nov. Type: New Guinea, Ramufluss, Lauterbach 2611 (holotype B). Piper subnudilimbum C.DC., Bot. Jahrb. 55 (1918) 205, syn. nov. Type: New Guinea, Djamu, Schlechter 17347 (holotype B, iso K). Piper quintuplinervium C.DC., Candollea, 1 (1923) 203, 204; Candollea, 2 (1925) 224 syn. nov. Type: New Guinea, sine loc., Lauterbach 605 (isotype WRSL). Dioecious glabrous climber. Lamina shortly petiolate, elliptic, ovate to broadly ovate, c. 10 x 5 cm, length:breadth ratio c. 2:1, glabrous on both surfaces, apex acute to acuminate, base cuneate to rounded, symmetrical; lateral veins 4 pairs, the lowermost pair very short, arising from the base, the 2nd and 3rd usually long, arising from the midrib a little above the base, the distal 4th pair very short, often absent, arising from the midrib near the apex. Stipules to 1.5 cm long, often 1 cm long, usually as long as petioles. /nflorescences shorter than leaves, peduncle c. 1.5 cm long, usually as long as petioles; males thin, to 6 cm long, bracts orbicular, peltate, subsessile; females shorter and thicker, c. 2 x 0.8 cm, bracts orbicular, peltate, sessile. Male flowers 2-staminate; pedicel very short stout, hirsute; stamens c. 1 mm long, anthers reniform to subglobose, 2-valved, slightly shorter than filaments. Female flowers sessile; stigmas 3- or 4-fid, sessile. Fruits sessile, entirely concrescent at maturity. Distribution: Philippines, Java, Sulawesi, Maluku, New Guinea, Bismark Archipelago and Solomon Islands. Notes: Piper minus is a smaller-leafed version of P. abbreviatum and there are no essential differences that can justify their being kept apart. Kajewski 2047 has leaves that vary greatly from the narrowly ovate c. 4 x 1.5 cm to broadly ovate c. 6 x 4 cm. In Piper subnudilimbum, the type materials contain immature inflorescences. The leaves are large, much like those of the Philippine populations of P. abbreviatum, but the leaf bases are like the typical New Guinea plants with three pairs of lateral Malesian Piperaceae 3 15 veins. This difference is, however, not significant and cannot be relied on to distinguish P. subnudilimbum from P. abbreviatum. As for Piper quintuplinervium, the isotype shows two leaf forms: the narrowly ovate, small ones borne on end of free branches and the large pentagonal leaves borne at the nodes of the climbing parts of the plant. Other than small differences between P. quintuplinervium and P. abbreviatum, there is nothing significant to keep the two species apart. Specimens examined: NEW GUINEA: Djamu Distr., Schlechter 17347; Ramufluss, Lauterbach 2611; sine loc., Lauterbach 605. SOLOMON ISLANDS: Kajewski 2047 (A). 2. Piper arfakianum C.DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 127. Type: New Guinea, [rian Jaya: Arfak Mts., Angi Lake, L.S. Gibbs 5525 (isotypes K, L). Synonym: Piper pilosulinodum C. DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 128, syn. nov. Type: New Guinea, Arfak Mts., Koebre Ridge. L.S. Gibbs 5624 (holotype BM; isotype K). Climber, usually glabrous, rarely hairy. Leaves with 1—2 cm-long petioles. Lamina symmetrically ovate, c. 9 x 4 cm; lateral veins 3 pairs, arising from the base; apex long acuminate; base rounded, not auriculate. /nflorescence longer than leaves; peduncle longer than petiole. Bracts peltate, circular. Flowers sessile, crowded. Stigma 3-fid, reflexed, subsessile. Distribution: New Guinea. Notes: Piper pilosulinodum, also from the Arfak Mountains, is conspecific with P. arfakianum. Having compared the type materials of these two species and with other recent collections, | am unable to keep the species apart. Of the collections cited above, Kanehira & Hatusima 13708 is the closest match to the type material of P. arfakianum and van Royen & Sleumer 7456 is a good match to the type material of P. pilosulinodum. Piper arfakianum is closely related to P. macropiper Pennant from which it can be distinguished by the above enumerated combination of characters, especially the petiole being much longer than in the latter species and the lamina base not auriculate. Specimens examined: IRIAN JAYA: Arfak Mt., alt. 2200 m, Kanehira & Hatusima 13708, April (A, BO); alt. 2400 m, L.S. Gibbs 5525 (BM, K); Koebre Ridge, L.S. Gibbs 5624 (BM, K); Mt. Nettoti, alt. 1920 m, van Royen & Sleumer 7456, Nov (A, 16 Gard. Bull. Singapore 55 (2003) BRI, CANB, L, LAE). PAPUA NEW GUINEA: Central Division, Mt. Tafa, alt. 2300 m, Brass 4047 (BRI), 5029 (BRI), 5/76 (BRI); Chimbu Distr., Mt. Wilhelm, alt. 2600 m, Borgmann 292, Aug. (LAE); Milne Bay Distr., Mt. Dayman, alt. 2200 m., Brass 22534, May (A, CANB, LAE); Morobe Distr., Edie Creek, Ridsdale NGF 30266, Nov (LAE); alt. 2000 m, Womersley & Thorne NGF 12837, Jun. (BRI, CANB, LAE); Mt. Kaindi, alt. 2200 m, Brass 29732, May (LAE); alt. 2438 m: Coode & Katik NGF 32863, Jun. (BRI, CANB, LAE); alt. 2300 m, Millar NGF 23627, Aug. (BRI, CANB, LAE); alt. 2100 m, Streimann NGF 30867, Aug (BRI, LAE, NSW); Southern Highland Distr., Anga, alt. 2000 m, Schodde 1556, Jul. (A, CANB, LAE); Spreader Divide, alt. 2255 m, Streimann & Kairo NGF 45446, Nov (CANB, NSW). 3. Piper austro-caledonicum C.DC. in DC. Prodr. 16.1 (1869) 346; Schlechter, Bot. Jahrb. 39 (1906) 92: Moore, J. Linn. Soc. Bot. 45 (1921) 381; Guillaumin, Flore analytique et synoptique de la Nouvelle-Caledonie, Phanerogames. (1948) 90. Type: sine loc., Forster s.n. (syntype: BM). sine loc., Vieillard 1227 (syntype: P). Synonyms: Piper paitense Schltr., Bot. Jahrb. 39 (1906) 92, syn. nov. Type: New Caledonia, Mt Paita. Schlechter 14964 (isotypes L, P, WRSL). Piper comptonii S.Moore, J. Linn. Soc. Bot. 45 (1921) 381, syn. nov. Type: New Caledonia, Mt Panie. Compton 1805 (isotype BM). Piper peekelii C.DC., Bot. Jahrb. 57 (1922) 354, syn nov. Type: Neu-Mecklenburg, Namatanai, P. Peekel 322 (holotype B). Glabrous climber. Leaves petiolate. Lamina of lower leaves broadly ovate to somewhat rounded to cordate, c. 9 x 7 cm, those of the upper leaves usually smaller; apex shortly blunt-acuminate; base rounded to peltate, slightly unequal, glabrous on both surfaces. /nflorescences slender, as long as the leaves, peduncle as long as the petiole: bracts pedicellate. Male flowers 3-staminate, stamens subsessile. Female flowers with 3-fid stigmas. Distribution: New Guinea, Bismarck Archipelago, Solomon Islands and New Caledonia. Notes: Forster f. misidentified his collection from New Caledonia as Piper siriboa Forst. f. (auct. non L.). I have seen both Forster’s and Vieillard’s collections annotated by C.DC. and am confident that the collections belong to P. austro-caledonicum. Based on examination of the type specimens, P. paitense, P. comptonii and P. peekelii are all conspecific with this species because I cannot differentiate their type specimens from specimens of P. austro-caledonicum. They are therefore reduced to synonymy. Piper austro-caledonicum is closely related to P. elbertii C.DC. of Indonesia and P. Malesian Piperaceae 3 17 retrofractum Vahl of the Philippines. Further study is needed to assess their relationship. Specimens examined: NEW GUINEA: Torricelli, Schlechter 14381] (BO). BISMARCK ARCHIPELAGO: New Ireland (Neu-Mecklenburg), Namatanai, P. Peekel 322 (B). SOLOMON ISLANDS: Guadalcanal, Wanderer Bay, BS/P 12247 (LAE); Kolombangara, Kokove area, BS/P 7579 (LAE); New Georgia, Kibukibu River, BSIP 4819 (LAE); Ranongga Island, BS/P 15572 (LAE); Santa Ysabel, Jejevo River, BSIP 4700 (LAE), Sigana, Brass 3451 (A, BRI), Vuranimala village, BSJP 1540] (A, LAE). NEW CALEDONIA: Aoui, Mei, Baumann-Bodenheim 10156 (P), 10176 (A, BRI, P); Bois du Mois de Mai, R. Blanche, Baumann-Bodenheim 14071 (A, BRI,P), 15179 (A, BRI, P); Col de Vulcan, Baumann-Bodenheim 8259 (A, BRI, P); Dambea, Franc 3 (P), Franc 3A (L); Ermitage stream, Compton 149 (P); Mt. Paita, alt. 300 m, Schlechter 14964, Oct 1902 (L, P, WRSL); Mt. Panie, Compton 1805, Aug 1914 (BM); Noudoure: MacKee 2670 (A, P); Yaouhe, Schlechter 14786 (P). SINE LOC: Forster s.n. (BM); Vieillard 1227 (P); Schlechter 14787 (L, WRSL). 4. Piper bosnicanum C.DC. in Gibbs, Phyt. Fl. New Guinea, (1917) 207; Candollea 1 (1923) 221, as ‘bosniakanum’. Type: New Guinea, Schouten Islands, Bosnic, L.S. Gibbs 6277 (holotype K; isotype L). Glabrous climber. Lamina ovate, to 15 x 10 cm, with 3 pairs of lateral veins arising directly from the base. Petiole to 2 cm long. Inflorescence to 5 cm long, shorter than leaves; peduncular stalks c. 2 cm long, bracts oblong with sinuous margin. Female flowers subsessile, spirally arranged; stigmas 3-4-fid, sessile, strongly reflexed. Fruits subsessile, obovoid to ovoid, c. 10 x 6 mm broad, crowded. Distribution: New Guinea and the Solomon Islands. Notes: Piper bosnicanumiis very close to P. buruanum Migq. of Maluku: further study is needed to assess their relationship. Specimens examined: IRIAN JAYA: Schouten Islands, Bisnik, L.S. Gibbs 6277, Jan (K; L); Vogelkop Peninsula, Aifact River, van Royen & Sleumer 7260, Nov (L). PAPUA NEW GUINEA: Sepik Distr., Aitepe Subdistr., along Pieni River, Darbyshire & Hoogland 8007, Jun (CANB). SOLOMON ISLANDS: Bougainville Island, north of Buin, Craven & Schodde 256, Aug (CANB, LAE); Guadalcanal, Wanderer Bay, BS/P 9104 (LAE); Kolombangara, Bambari area, BSJP 751] (LAE); Malaita, Takwa, BS/P 10785 (LAE); New Georgia Group, Roniane Lagoon, BS/P 2918 (LAE); Wagina Island, BSIP 5493 (LAE). 5. Piper lessertianum C.DC., J. Bot. 4 (1866) 164; Quisumbing, Philip. Jour. Sci. 18 Gard. Bull. Singapore 55 (2003) 43.1 (1930) 36, fig. 10, including var. oblongibaccum (C.DC.) Quisumbing. Type: Philippines, Luzon, sine loc. Cuming 1343 (‘1342’), 1841 (holotype BM). Synonyms: Chavica lessertiana Miq., Syst. Piperac., (1843) 270 non Piper lessertianum C.DC. (1866): Piper pseudo-chavica C.DC. in DC., Prodr. 16.1 (1869) 351, nom. superfl. Type: Philippines: Luzon, Cagayan Province, Cuming 1343 (holotype G, not seen; isotype K). Piper biroi K.Schum. & Lauterb., Nachtr. Fl. Schutzgeb., (1905) 238, syn. nov. Type: New Guinea, near Malanaku, Biro 32, Oct. (holotype B). Piper oblongibaccum C.DC. in Leafl. Philip. Bot. 3 (1910) 777, Type: Philippines: Oriental Negros Prov., E/mer 9456 (not seen). Piper lineatipilum C.DC., Nova Guinea Bot., 8.6 (1914) 1009, syn. nov. Type: New Guinea, Mt. Hellwig, Rémer 962 (holotype L). Piper viridibaccum Trel., J. Arn. Arb., 9 (1928) 150, syn. nov. Type: New Guinea, Mowabula, L./J. Brass 1370 (holotype A, isotype BRI). Climber, dioecious, glabrous to villose. Leaves shortly petiolate, petiole to 1 cm long. Lamina narrowly ovate, ovate to broadly cordate, 12—32 x 5—17 cm; base unequally rounded, one lobe acute to narrowly rounded, the other auriculate, lateral veins 34 pairs, the distal pair arising about 2 cm from the midrib, the others almost directly from the base; apex often very long acute to acuminate. /nflorescences 5—12 cm long, less than 5 mm thick; peduncle slender, 3—8 cm long, much longer than the petiole and the floriferous part of the inflorescences. Male flowers with 2 subsessile stamens, with pedicellate peltate bracts. Female flowers with 3-fid rounded sessile stigmas. Fruits oblong, to 2.5 x 1.5 mm, borne crowded but not concrescent at maturity. Distribution: Philippines, Sulawesi, New Guinea (Irian Jaya and Papua New Guinea). Notes: This species is attributable solely to C.DC. who published it in 1866 basing on Cuming 1342 (in herb. BM) without reference to Miquel’s earlier species Chavica lessertiana which is based on Cuming 1343 (in herb. G). I am grateful to have the advice of J.F. Veldkamp of Leiden (pers. comm., 2003) that Cuming 1342 in C.DC.’s protologue of 1866 should read Cuming 1343 because Cuming 1342 refers to a species of Gramineae. Nevertheless, the authority of Piper lessertianum (1866) remains attributable solely C.DC. without reference to Miquel. Later, in DC. Prodr. 16.1 (1869) 351, when C.DC. combined Miquel’s Chavica lessertiana (1843) with his own Piper lessertianum (1866), he created the name Piper pseudo-chavica for the combined species. By contemporary rules, the name Piper pseudo-chavica is superfluous because his own Piper lessertianum (1866) is perfectly legitimate and should have been used instead. Malesian Piperaceae 3 19 Our Malesian species is not to be confused with Piper lessertianum (Miq.) C.DC. in DC. Prodr. 16(1): 258 (1869) based on Ottonia lessertiana Migq., which refers to an American species and, which binomial is a later homonym of our species. Piper lessertianum C.DC. (1866) is readily recognised by the petioles being very short and the peduncles very long, often more than five times longer than the petiole and the floriferous part of the inflorescence. Yet, it is most variable in leaf form with lamina ranging from very narrowly ovate with a long attenuate apex through ovate ob-pentagonal to broadly cordate or heart-shaped. Besides, leaves borne on the climbing parts are usually small and broadly cordate to heart-shaped while those borne on the freely hanging branches tend to be ob-pentagonal to very narrowly ovate with long attenuate apices. All these leaf forms have now been observed to occur on the same plant as evidenced by Vink & Schram BW 8&7]. In the circumstances, I have to assign to synonymy Piper biroi, Piper lineatipilum and Piper viridibaccum as they are distinguished purely on these leaf forms. For the same reason and after examination of collections authenticated by Quisumbing, I am unable to maintain var. oblongibaccum (C.DC.) Quisumbing. Specimens examined: PHILIPPINES: Luzon, Cagayan Prov., Cuming 1343 (BM, K); Mindoro, Mt. Halcon, Ramos & Edano 40676, Mar (NSW); Panay, Capiz Prov., Mt. Bulilao, Martelino & Edano 35706, Jun (NSW); Jaminden, Ramos & Edano 30861, 31272, May-Apr (NSW); Libacao, Martelino & Edano 35395 (NSW), 35442, May- Jun (NSW); Mt. Macosolon, Ramos & Edano 30776, Apr-May (NSW). SULAWESI: Kabaena Island, Elbert 3393 (BO). IRIAN JAYA. Wati, Mapon Distr., /jiri & Niimura 674, Apr (L); Wissel Lakes, Doglia, Vink & Schram BW 8&71, May (CANB, LAE). PAPUA NEW GUINEA: Milne Bay, near Mayu Island, Streimann NGF 28829, Apr (L, NSW): Morobe Distr., Asubazo, Kairo NGF 24403, Jul (L); Northern Division, Sibium Range, R. Pullen 5924, Sept (CANB). 6. Piper longipilum C.DC., Bot. Jahrb. 55: 216 (1918); Candollea 1: 206 (1923). Type: New Guinea, Sepik, alt. 1-200 m, Ledermann 7565 (holotype B, isotype B). Climber, dioecious, lightly pilose. Leaves shortly petiolate, petiole to 2.5 cm long. Lamina ovate, c. 28 x 18 cm; base unequally cordate, lateral veins 5—6 pairs, the basal pair arising c. 1 cm from the base; apex acuminate. Male inflorescences (immature) c. 8 cm long, c. 3 mm thick; peduncle slender, c. 1.7 cm long. Male flowers with pedicellate peltate glabrous bracts. Female unknown. Distribution: New Guinea. Notes: The type material of this species consists of two sheets: the one with field 20 Gard. Bull. Singapore 55 (2003) notes and annotation by C.DC in 1917 is the holotype, the other is the isotype. A cursory examination of the type material suggests an alliance with Piper decumanum L. of which it might represent a hairy variant. As the type seems to be the only material there is of this species, I am unable to determine its status until I see more collections, especially of female plants. 7. Piper melula Trel., J. Arn. Arb. 9 (1928) 148. Type: Papua New Guinea, Vaitata River, L.J. Brass 1130 (holotype A; isotype BRI). Dioecious glabrous climbers. Leaves moderately petiolate, petioles 2-4 cm long, much longer than peduncles. Lamina irregularly ovate, broadly ovate to pentagonal, c. 12 x 8 cm; apex bluntly shortly acuminate; base rounded to shallowly cordate; lateral veins 3 pairs, the distal pair arising alternately from the midrib about 0.5—1 cm from the base, the other 2 pairs directly from the base; only the distal pair reaching to the apex, the 2" pair reaching to 1/2 the length of the lamina, the basal pair only to 1/3 of the length of the lamina. /nflorescences shorter than the leaves, males slightly longer than females. Male floriferous part to 6 cm long on a peduncle to 7 mm long; female floriferous part to 4 cm long on a peduncle to | cm long. Bracts circular, margin sinuate, peltate, very shortly pedicellate. Male flowers with strongly reniform anthers, dehiscing by long apical slits. Female flowers sessile, crowded; stigmas 3- to 4-fid, sessile, stigmatous part reflexed. Fruits almost entirely concrescent, free only at the apices. Distribution: Papua New Guinea. Notes: Piper melula has the same leaf venation pattern as P. plagiophyllum K.Sch & Laut., and in some cases even the lamina shape of that species, but it differs in being a climber amongst other characteristics. The infructescences of Piper melula appear similar to those of P. pachyarthrum K.Sch., but these species differ in other respects. Piper melula appears also to be related to P. elbertii C.DC. of the Lesser Sunda Islands and P. austro-caledonicum of the Solomon Islands and New Caledonia. There is a collection, Kellman ANU 1597 (CANB) from Mindanao in the Philippines, which is remarkably similar to Piper melula. However, as this collection has a very young inflorescence, I hesitate to identify it as this species. Specimens examined: PAPUA: Gulf distr., Tauri River, Schodde & Craven 4700, Mar (CANB); Vailala River, L.J. Brass 1130, Mar (A, BRI); Milne Bay distr., Kwagira River, Peria Creek, L.J. Brass 23997 (A, CANB, LAE), 24006, Aug (A, CANB, LAE). Malesian Piperaceae 3 a | 8. Piper novo-guineense Warb., Bot. Jahrb., 13 (1891) 284, K.Schumann & Lauterbach, Fl. Schutzgeb., (1900) 258. Type: New Guinea, Sattelberg bei Finnschlafen, Warburg 20740 (holotype B; isotype A). Glabrous climber. Lamina narrowly elliptic, narrowly ovate to broadly ovate or broadly pentagonal, 17—19 x 5—9 cm; apex shortly blunt acuminate to long acute; base cuneate to rounded, very slightly asymmetrical, not auriculate; lateral veins 3 pairs, the distal pair arising slightly alternately from the midrib c. 1—2 cm from the base, the other 2 pairs arising directly from the base, sometimes another faint short pair arising from the base; the veins on the underside, especially at the basal portion, often covered with minute reddish brown scales, petiole to 1 cm long. Male inflorescences thin, as long as the females, peduncle as long as the males. Male flowers with bracts as in the females; stamens subsessile with broadly crescent-shaped anthers. Female inflorescences to as long as the leaves, sometimes longer, usually to 15 cm long; peduncular stalk c. 2 cm long. Female flowers sessile, stigma 3-4-fid. Fruits sessile, oblong, crowded; bracts rounded peltate. Distribution: New Guinea. Notes: Piper novo-guineense is reported to be common in primary forest and older secondary forests, between 500 and 2000 m altitude. At higher altitudes, the plants tend to have smaller leaves. This species is closely related to Piper macropiper Pennant and especially to P. truncatibaccum C.DC. Specimens examined: IRIAN JAYA: Lake Habbema, L.J. Brass 10299, Oct (A). PAPUA NEW GUINEA: Morobe Distr., Boana Mt. Clemens 8467 (A); Gurokar, L.J/. Brass 29499, May (CANB, LAE); Huon Peninsula, Mt. Rawlingson, Hoogland 9054, Jun (A, BRI, LAE); Sattelberg, Warburg 20740 (B, A); Wantoat area, Kikiepa, Womersley & Thorne NGF 11875, Jun (BRI, LAE); Selileo, Hellwig 580, Apr (B); Sepik, near Kilifas village, Forman & Kumul NGF 48261, Mar (LAE). 9. Piper pallidilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1009. Type: New Guinea, Mt. Hellwig, alt. 2600 m, von Romer 1316 (holotype L). Synonym: Piper peracutilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1005, syn.nov. Type: New Guinea, Mt. Hellwig alt. 2600 m, von Rémer 1252 (holotype L). Climber, glabrous. Leaves shortly petiolate. Lamina ovate, 6-8 x 24 cm; base rounded, apex acuminate. Lateral veins 2-3 pairs, distal pair arising from the midrib a little 22 Gard. Bull. Singapore 55 (2003) above the base. /nflorescence shorter than leaves. Peduncular stalk 7-10 mm long, about as long as petiole. Males up to 4 cm long, females c. 2 cm long. Flowers borne crowded, sessile. /nfructescence congested, to 12 mm diam. as in Piper abbreviatum Opiz. Distribution: New Guinea. Notes: The type material of Piper peracutilimbum is indistinguishable from specimens of P. pallidilimbum. Therefore I see no reason to keep the two species apart. Piper pallidilimbum is very close to P. abbreviatum Opiz from which it differs in the lamina being furnished with fewer lateral veins, one of which issues from the midrib, and also by the peduncular stalk being somewhat filamentous. Specimens examined: IRIAN JAYA: Mt. Hellwig, alt. 2600 m, von Romer 1316 (L), 1252 (L); sine loc., Pulle 700 (L), 737 (L), 834 (L). 10. Piper pseudamboinense C.DC., Bot. Jahrb. 55 (1918) 206. Type: New Guinea, Keneyia, alt. 150 m, Schlechter 15445 (holotype B). Climber. Leaves very shortly petiolate. Lamina pentagonal to oblong, commonly c. 24 x 11 cm; apex bluntly acuminate, acumen to 1.5 cm long, base asymmetrically cordate, one side often forming a large lobe, the lobes often covering the node; lateral veins usually 5 pairs, the distal pair alternate, arising from the midrib c. 2—3 cm from the base, the 2° pair often arising also from the midrib but very close to the base; the other 3 pairs, often very short, issuing directly from the base; intercostals very prominent; upperside of lamina glabrous, undersides with short brown pubescence on the veins. Female inflorescences to as long as leaves, 20—24 cm long; the floriferous portion as long or to only slightly longer than the peduncular stalk; peduncle glabrous. Bracts circular, peltate, long pedicellate. Female flowers sessile, congested. Stigma 2-lipped, spreading, subsessile. Fruits sessile, free. Distribution: New Guinea. Notes: A lowland climber, Piper pseudamboinense 1s a little known species. Of the two collections identified as this species, NGF 25678 appears to be the best match to the holotype, the other being a more hairy variant. Piper pseudamboinense is very closely similar to P. amboinense (Miq.) C.DC. and from which it differs in the more prominent lateral veins and its stigma being 2-lipped. It is also closely similar to P. lessertianum C.DC. from which it can be distinguished by the 2-lipped stigma in addition to lamina and inflorescence characters. Malesian Piperaceae 3 23 Specimens examined: PAPUA NEW GUINEA: Morobe District, Buso River, NGF 25678 Aug (CANB, LAE); Oomsis Creek, T.G. Hartley TGH 10564, July (A, CANB, LAE); Keneyia, alt 150 m, Schlechter 18445 (B). 11. Piper reinwardtianum (Miq.) C.DC., DC. Prodr. 16.1 (1869) 354. Merrill, Inter. Rumph. Herb. Amb., (1917) 182. Basionym. Macropiper reinwardtianum Miq.., Linnaea 21 (1848) 481. Type: Indonesia, Maluku, Nussa Laut, Reinwardt, July 1821 (holotype L?). Synonym: Chavica reinwardtiana (Migq.) Miq., Ann. Mus. Bot. Ludg. Bat.1 (1863) 136. Dioecious shrub, glabrous to lightly pilose. Leaves with petiole 2.5—5 cm long. Lamina broad ovate to rounded, c. 16 x 15 cm, slightly asymmetrical; apex shortly acute, base asymmetrically truncate-cordate, lateral veins usually 34 pairs, upperside of lamina glabrous, undersides lightly pilose on the veins. /nfructescences to 11 cm long, peduncle glabrous to 2.5 cm long. Female flowers sessile, congested. Stigma 3, rarely 4, short thickset. Fruits sessile, somewhat obovoid, concrescent on the lower half. Distribution: Indonesia, Maluku. Notes: I have not seen any type material of this species which is probably in the Leiden Herbarium to which Reinwardt donated his collections from the region. The collections of Robinson and Teysmann cited herewith are all that I have seen of this species; and I must of neccesity rely on C. de Candolle and Merrill in the interpretation of this apparently rare species. In general growth habit, Piper reinwardtianum recalls that of P. sundaicum Blume, also a shrub from the same region. However, it has infructescences similar to P. mestonii Bailey of New Guinea and Australia from which it can be distinguished by the stigma being 3- or 4-fid, not bifid, and the plant a shrub, not a climber. In these respects, P. reinwardtianum appears to be an interesting species, and further investigation may yield insight into the phytogeograpical relationship between the three species. Specimens examined: INDONESIA: Maluku, Ambon, Robinson 60 (L). Ceram, Teysmann s.n. (L). 12. Piper subvirosum C.DC., Bot. Jahrb. 55 (1918) 215. Type: New Guinea, Alexishafen, Wiesenthal 68 (holotype B). 24 Gard. Bull. Singapore 55 (2003) Climber, twigs minutely pubescent. Lamina broadly ovate, 15—20 x 10-12 cm, upper surface glabrous, lower surface sparsely covered with short stiff hairs especially on lateral veins on basal part of the lamina; apex shortly acuminate, acumen 1.5—2 cm long; base broadly cuneate, rounded to truncate; lateral veins 3 pairs, distal pair alternate, arising from midrib a little above the base, the others directly from the base; intercostals prominent. Petiole to 2 cm long. Female inflorescence to 27 cm long, peduncle to 3 cm long, floriferous part to 24 cm long; inflorescences usually longer than leaves, peduncle longer than petiole; mature inflorescence to 7 mm diam. Female flowers sessile, not concrescent, with 3- or 4-fid stigmas. Distribution: New Guinea. Notes: Piper subvirosum is related to P. macropiper Pennant from which it can be distinguished by the much larger leaves and inflorescences, at least one pair of the lateral veins arising from the midrib, the lamina base not auriculate, and the peduncle not excessively longer than the petiole. Not much is known about this species. The two recent collections cited here are the closest match to the holotype. Specimens examined: IRIAN JAYA: Star Mts., Sibil Valley, alt. 1200-1500 m, Kalkman & Nicholas 4160, May (L). PAPUA NEW GUINEA: Central Division, Mt. Tafa, L.J. Brass 4127 (A); Alexishafen, Wiesenthal 68 (B). 13. Piper sundaicum Blume, Verh. Bat. Genoots. 11 (1826) 187, fig. 14; C.DC., DC. Prodr. 16.1 (1869) 352; Koorders, Verh. Kon. Ak. Wet. Amsterdams. 2, 14.4 (1908) 42, reprint; Koorders, Exkursionsflora van Java. 2 (1912) 26; Koorders- Schumacher, Exkursionsflora Atlas. 4.2 (1924) 445, fig. 718G. Type: Maluku, Ambon. Blume s.n., Leiden Herb. 908.363-43 (lectotype L, here designated). Synonyms: Chavica sundaica (Blume) Mig. Syst. Piperac., (1843) 274; Fl.Ind. Bat 1(2) (1859) 446, exclusive of the synonym P. a/bum Vahl. Piper amplilimbum C.DC., Candollea 1 (1923) 213; Schroeder, Candollea 3 (1926) 135; Chew, Blumea. 20 (1972) 145, fig. 1, syn nov. Type: Maluku, Ambon, Teysmann s.n. 1905 (holotype BO). Dioecious scandent shrubs, entirely glabrous. Twigs 5—8 mm diam.; internodes smooth; stipular scars prominent. Lamina symmetrically ovate, (23—)25—27 x 14-19 cm, somewhat coriaceous; apex acuminate, acumen to 2 cm long; base rounded; margin entire; lateral veins 34 pairs, very prominent, the distal pair arising alternately from the midrib at 2-3 cm from the base, the others directly from the base; intercostals Malesian Piperaceae 3 25 numerous, not straight, quite prominent. Petioles c. 2 cm long, smooth. Stipules to 8 x c. 1 cm, apparently not adnate to petioles. /nflorescences: male long, thin; female 11-13 x 0.7 cm, the peduncular stalk c. 1.5 cm long, glabrous. Bracts somewhat irregularly circular, peltate. Female flowers sessile; ovary pyramidal; stigmas 3-fid, short, sessile, slightly reflexed. Fruits ovoid, sessile, c. 3 cm long: crowded, not concrescent. Distribution: Indonesia, Maluku. Notes: Blume recorded this species as also occurring in Java. Koorders (1908, 1912) doubted the Javanese record as he failed to find any collection from that island. I have also been unsuccessful in finding any material of the species from Java. This is probably a very rare species: the type materials are all that is known of this species. I have compared the type materials of Piper sundaicum with that of P. amplilimbum C.DC., and I am unable to distinguish the one from the other. Piper sundaicum appears to have some relationship with Piper reinwardtianum from the same region (see above). Piper sundaicum has been identified with P. album Vahl by previous botanists. The two species are however, quite distinct: P. album Vahl is a climber, our species is not. Specimens examined: MALUKU: Ambon, Blume s.n. (L): Teysmann s.n. 1905 (BO). Acknowledgements I thank the curators of the following herbaria (identified by their acronyms) for loan of their collections, including type materials, for my study at the Herbarium of the Royal Botanic Gardens, Sydney where I held tenure as Senior Research Scientist from 1970 to 1975: A, B, BO, BRI, CANB, K, L, LAE, P, PR, PRC and WRSL. | sand gictiongaca seolnahoanioe nomena ome Pas yer bi rare are > ° lero < cf S10 = a1) eer s aé ona echate —— ; onnen. & angie ‘2m loony. Fernale , CAC = Ley’) 'v ? base 1 as bao Saponeve ed aris! ‘ Peeks eS. i cAlisi ie Fy p tl bars: ib nana vont ie jicwviriveds j= » Zt om Lang: ia nae rence: _—— apeg, hana ect iaibae «21 a sanburl fev ’ = git & tis ras. aa | = - : ict babys nan it in on - — 4p e pee; i i. - ETL CLL she 1 P nwyr > e >i ; or TS eas Bike ay ie vc Sh a4) vi Salaqit Ribs, RCT ti gh anh. dens) ‘Se i) wow * io) ziEag Ga Sisk} =m” Loan Soe fink ic cana Pi: a “he ac? en ees we AY ANB) ‘CU UINBA : ten bivie oe nfs * > a mri? a*? 0 Fi. alr J ‘ rT] } a AM. lor ay fy ery ty robin; ' ARO1TI loo th : Bi i] a, Joes f, sonst 58 a) U Tt ly i aaa so a3 : ce > ~ ea ’ “ ley T. fe A Ay? Jotien 4 9 is) Wks, Asm, Tesgondi - _ _ 7 7 - : Be u a 4H ny siti em neem wc f iy . *, me, 2C ath ty Sa 7 roi ish ui, fhe a 2. Oe Mhar a a - _ 7 J “i's i - Garden’s Bulletin Singapore 55 (2003) 27-30 A New Species of Schismatoglottis (Araceae) from Sabah, Malaysia A. HAY AND C. HERSCOVITCH Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia Abstract Schismatoglottis tahubangensis A. Hay & Herscovitch is described, illustrated and inserted into the latest key to Bornean Schismatoglottis species. Schismatoglottis tahubangensis A. Hay & Herscovitch, sp. nov. A Schismatoglottis platystigma M. Hotta spadice tenuiore, staminodiis inter pistilla, connectivo longiore differt. - Typus: Malaysia, Sabah, just outside Kinabalu Park, 100 m to Tahubang River along route to Marai-Parai, 11,000 m alt. [sic], 12 Sep 1993, J. Nais, Dolois, Harry & Masius SP 05312 (KEP, holo). Terrestrial herb c. 30 cm high. Stem creeping or decumbent, rooting between the nodes, c. 1 cm diam., with internodes c. 5 mm long. Leaves 6 together; petiole 10-17 cm, sheathing in the lower 1—2 cm, wings of sheath extending into a papery narrowly triangular ligule to 2.5 cm; blade 11—15 cm long, 3.5—5 cm wide, narrowly obovate, somewhat asymmetric, base truncate to acute, somewhat asymmetric, inserted into adaxial face of petiole, apex acute; midrib pronounced and minutely puberulent abaxially, with c. 10 primary lateral veins on each side diverging at c. 45°, interprimary veins somewhat conspicuous abaxially, obscure adaxially, fine. /nflorescence solitary, subtended by an inconspicuous papery cataphyll c. 6 mm long; peduncle 3 cm long. Spathe pinkish, 7 cm long, slender, weakly constricted; lower spathe 1.5 cm long, 3mm diam. Spadix very slender, cylindric for most of its length, 5.3 cm long, 1.5 mm diam.; female zone 1.2 cm long, (not clear whether adnate to spathe at base); pistils closely packed, subglobose, 0.3 mm diameter; stigma discoid, almost as wide as the ovary; interpistillar staminodes conspicuous, round and flat-topped or concave, drying pale yellow, of the same height and diameter as pistils; sterile interstice c. 2 mm long comprised proximally of 2 irregular whorls of looser packed, 0.5 mm diam., flat-topped staminodes, differentiated from interpistillar staminodes by larger size, slightly convex top, and dark yellow colour on drying, and distally of 2—3 irregular whorls of sterile 28 Gard. Bull. Singapore 55 (2003) stamens drying dark brown; male zone 1.2 cm long; stamens subsessile, more or less oblong ovate from above, more or less irregularly paired and oriented along long axis of spadix, c. 0.8 mm wide; thecae round, opening by a single apical pore; connective robust, columnar, flat-topped and somewhat impressed, slightly exceeding the thecae in height and about 1.5 times their diameter, rounded triangular from above with ends indented above thecae; appendix 2.8 cm long, tapering to a fine naked tip, composed of flat-topped, more or less rounded or irregularly polygonal staminodes c. 0.3 mm diam. similar to those of sterile interstice. Fruit unknown. Distribution: Malesia — Borneo, Sabah, Kinabalu (known only from the type). Habitat: Ona steep slope in unspecified vegetation type at c. 1,100 m alt. [Note that the label on the type records the altitude as 11,000 m, which 1s, of course, impossible]. Notes: This species somewhat resembles Schismatoglottis platystigma (Brunei) in having discoid stigmas and relatively small thecae on the edge of a broad connective. However, it is readily distinguished from that species by the presence of interpistillar staminodes, the subsessile stamens with a much higher connective and the more slender spadix. Schismatoglottis tahubangensis may be inserted into the key to Bornean Schismatoglottis provided by Hay & Yuzammi (2000) as follows: 17a. Inflorescence solitary or two together; anthers with broadly dilated connective Fe ee EO RES re ree ree Teeter en ee 18 17b. Inflorescences c. 4-6 together; anthers dumb-bell-shaped, with the connective a i a a ee a ERE NITED 18a. Interpistillar staminodes absent; appendix 1—2 cm long, blunt-tipped; Temburong, BOE? 2A ANE. RU TER G.. RUA ASS ORAS Se 18b. Interpistillar staminodes numerous; appendix c. 3 cm long, tapering to a fine CIN, PA MNANAIEL, DOIN. os os nog om eee i rmadtaesapasenn esata as <email S. tahubangensis 19 etc = 18 etc in Hay & Yuzammi (2000). Schismatoglottis tahubangensis 29 Figure 1. Schismatoglottis tahubangensis Herscovitch & A. Hay A. habit; B. ligule at base of petiole; C. detail of leaf venation; D. midrib detail; E. inflorescence with part of spathe removed: F. pistils with interpistillar staminodes; G. sterile interstice with staminodes and sterile stamens; H. stamens; I. staminodes of appendix. J. Nais, Dolois, Harry & Masius SP 05312. 30 Gard. Bull. Singapore 55 (2003) Acknowledgements We thank Josef Bogner for drawing our attention to the specimen on which this species is based, Dan Nicolson for commenting on the ms and and Lesley Elkan for the illustration. Reference Hay, A. & Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I — Schismatoglottis. Telopea. 9: 1-177. Garden’s Bulletin Singapore 55 (2003) 31-34 An Unusual New Species of Homalomena (Araceae) from New Guinea C. HERSCOVITCH AND A. HAY Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia Abstract Homalomena impudica Herscovitch & A. Hay is described, illustrated and inserted into the latest key to Homalomena species of New Guinea. Homalomena Schott was revised for New Guinea by Hay (1999). An overlooked specimen on loan to NSW from LAE represents a very distinctive undescribed species. Homalomena impudica Herscovitch & A. Hay sp. nov. Homalomena tenuispadix Engl. simulans sed inflorescentia femina spatham aequanti, inflorescentia mascula clavata tota exserta differt. - Typus: Papua New Guinea, West Sepik Province, Vanimo, Ossima, 28 Jan 1969, H. Streimann & A. Kairo NGF 39239 (holo, LAE). Erect multi-stemmed herb to c. 30 cm high. Stem condensed, 1.5 cm diam. Leaves to 8 together; petiole longer than blade, 17—32 cm long, glabrous, slender, sheathing in lower 6—7 cm; blade dull green above, green below, glabrous, sagittate to ovate- cordate,14-19 cm long, anterior lobe 12—16.5 cm, c. 8 cm wide, posterior lobes triangular to rounded, 2.5—4 cm long; midrib more prominent abaxially, with c. 8 pairs of lateral veins diverging at c. 45 °. Inflorescences to 9 together, each subtended by a narrow triangular papery cataphyll to 4 cm long; peduncle erect, later declinate, 4.5—6 cm long, longer than spadix. Spathe slightly constricted in bud, then narrow cylindrical, 2.0—2.8 cm long, 5 mm diam., with a conspicuous rigid darker sometimes reflexed apiculum 2—3 mm long. Spadix 2.7—3.6 cm long, much exceeding the spathe at anthesis, stipitate for c. 5 mm; female zone equalling to exceeding the spathe, 2.0—2.4 cm long; pistils very widely scattered along axis of female zone, becoming loosely and irregularly whorled distally, globose, c. 1 mm diam., infrapistillar staminodes absent; stigma button-like, sessile, 0.3 mm across; male zone narrow oblong-ovoid, 0.7—1.1 cm 32 Gard. Bull. Singapore 55 (2003) long, whitish, fully exserted from spathe; male flowers (-)2-merous; stamens more- or-less oblong to irregularly polygonal from above, c. 1 x 0.3-0.5 mm, the thecae overtopped by connective. Fruit unknown. Distribution: Malesia— New Guinea, Papua New Guinea, West Sepik Province (known only from the type locality). Habitat: In a dry stream bed in lowland rain forest at c. 30 m alt. Notes: This species resembles Homalomena tenuispadix Engl., but differs in the greatly elongated and lax female zone which equals to exceeds the spathe, and in the completely exserted and somewhat clavate male zone. The inflorescence at first resembles that of Schismatoglottis after the male zone has been exposed by the fall of the spathe limb, until it is noted that the spathe, as typical for Homalomena, 1s fully persistent. While some New Guinea species of Homalomena, such as H. davidiana A. Hay and others (see Hay 1999), characteristically have the male zone partially exserted ~ from the spathe after anthesis, this species extends the entire male zone beyond the apex of the spathe prior to anthesis. This characteristic, as distinct from the state where the male zone is exposed by the spathe being wide open ms anthesis, appears not to be recorded for any other species in the genus. Field notes describe the flower (sic) as white and the fruit green. The specimen includes fragments of declinate fruiting peduncle, but the infructescence is not present. It is unclear whether or not the colour observations refer to the spathe at different stages. The specific epithet, meaning lewd or shameless, alludes to the fully exposed male zone of the spadix. Homalomena impudica can be inserted into the key provided by Hay (1999) as follows: 10a. Spathe tip + hood-forming; spadix contained within the spathe after anthesis; peduncle tice Sorter tian tite PetiOles .2-. tcc cce cee ree ececee se H. lauterbachii 10b. Spathe with the tip reflexed; spadix exserted from the spadix during and/or after anthesis; peduncle c, 1/3—1/2 the leneth of (he petioles |. na beriecekes Gannann een 11 lla. Spadix with the male zone clavate and completely exserted from the spathe at anthesis). A SS a ee eae See H. impudica 1 1b. Spadix with the male zone cylindric-tapering, not exceeding the spathe at anthesis, partially exserted from spathe after anthesis ...................ssseeeseeeH. Cenuispadix 12 etc. = 11 etc. in Hay (1999) Homalomena impudica 33 Figure 1. Homalomena impudica Herscovitch & A. Hay. A. habit; B. inflorescence with part of spathe removed; C. tip of spathe; D. inflorescence in bud; E. pistils; F. stamens. Streimann & Kairo NGF 39239. 34 Gard. Bull. Singapore 55 (2003) Acknowledgements We thank LAE for the loan of the specimen on which this species is based and Catherine Wardrop for the illustration. Reference Hay, A. 1999. Revision of Homalomena (Araceae-Homalomeneae) in New Guinea, the Bismarck Archipelago and Solomon Islands. Blumea. 44: 41-71. Garden’s Bulletin Singapore 55 (2003) 35-60 Pleuroschisma, a New Section of Cyrtandra (Gesneriaceae) from Borneo O.M. HILLIARD’, B.L. BURTT! AND M.H. BOKHARI? ‘Royal Botanic Garden Edinburgh, EH3 5LR, U.K. *Multan, Pakistan Abstract Twelve Bornean species of Cyrtandra constitute a new section, Section Pleuroschisma, the name derived from an important diagnostic character, namely the unique fruit with a median septicidal split on each side (not reaching either apex or base of the fruit). Nine of the species and two varieties are newly described, C. angustielliptica Hilliard & B.L.Burtt, C. coacta Hilliard & B.L.Burtt, C. insolita Hilliard & B.L.Burtt, C. linauana B.L.Burtt, C. seganica Hilliard & B.L.Burtt, C. tesselata Hilliard & B.L.Burtt, C. tunohica Hilliard & B.L.Burtt, C. pendulifera Kraenzl. var. grossipilosa Hilliard & B.L.Burtt and C. sarawakensis C.B.Clarke var. Jongipilosa Hilliard & B.L.Burtt. Two species recognized remain nameless because the material is inadequate to typify a name. Introduction There are already 40 sectional names in Cyrtandra and the limits and interrelationships of the groups so designated are far from being understood; the addition of yet another sectional name to the list may seem somewhat irresponsible. Nevertheless, the fruit characters that are used to diagnose this section - the smooth fruit wall (due to a thin layer of sclerenchyma beneath the epidermis, Fig. 3), and the septicidal splits with inrolled margins (Fig. 1) form a unique combination in this small group. It is the lateral splits in the fruit-wall that form the basis for the sectional name Pleuroschisma; these splits, it should be noted, are septicidal and there is some reason to suppose that septicidal dehiscence may have been the primitive condition in the family as it is found in Ramonda, Haberlea, in the type species of Corallodiscus, all on the northern fringe of the subfamily Cyrtandroideae, and in the tribe Coronantherae, a basal tribe of subfamily Gesnerioideae found in New Caledonia, E. Australia, New Zealand and southernmost S. America. Its presence elsewhere in the family (e.g. in Leptoboea) may be secondary. It must be noted, however, that the lateral splits in this new section of Cyrtandra do not reach from apex to base of the fruit (Fig. 1) and do not give rise to free fruit-valves, as in a fully septicidal fruit. These features set section smaller. 36 Gard. Bull. Singapore 55 (2003) Figure 1. Mature fruit of Cyrtandra sarawakensis The two halves remain held together at apex and base. Burtt B4¢71]2 x4. Pleuroschisma apart from all the other sections. As so often in herbarium work on Cyrtandra, we have been much hampered by lack of flowers on many specimens, although mentioned in the collector’s field notes. Nevertheless, it is not the flowers that provide the diagnostic features of the Section: it is the fruits. We have therefore felt justified in naming C. tesselata (no. 6), C. seganica (no. 7) and C. insolita (no. 11), which we know only in fruit. In the eight species where we have seen flowers, the major floral characters are uniform: calyx divided almost to the base into five deltoid lobes, corolla c. 35-40 mm long, white, cream or palest yellow with two orange/yellow bars in the throat, puberulous outside, disc cupular, occasionally deeply excavated on one side, ovary crowned with a coma of relatively long hairs, otherwise so minutely gland-dotted or pustulate as to appear glabrous on casual inspection, style puberulous, stigma distinctly bilobed, lobes relatively large, clavate in outline. Leaf-venation provides a useful character in distinguishing some of the species, both the number of lateral veins on each side of the midrib, and the pattern described by the tertiary venation, of which three main types have been distinguished; for these we use the terms subscalariform, reticulate and pinnate (Fig. 2, A,B,C,D). In some species, the tertiary venation is invisible (e.g. C. insolita). Cyrtandra sect. Pleuroschisma af Figure 2. Types of tertiary venation found in Cyrtandra sect. Pleuroschisma A. subscalariform (C. sarawakensis, Burtt B1947, about actual size). B. finely reticulate (C. tesselata, Hotta 1417, slightly enlarged). C. coarsely reticulate (C. sp. nov. no. 12, Burtt B2658, x2). D. pinnate (C. coacta, Beaman 10656, slightly enlarged). 38 Gard. Bull. Singapore 55 (2003) The distinctive leaf anatomy, with tracheoids in the hypodermis, has already been illustrated (Bokhari & Burtt, 1970, Plate 2, G,H): the tracheoids may have either parallel or reticulate bars of thickening (Fig. 4). In some species (e.g. C. hoseana), similar tracheoids are found in the mesocarp of the fruit; spheroidal sclereids are commonplace and abundant (Fig. 3), making fruit-sectioning very difficult. In several species, polymorphic sclereids occur in the mesophyll of the leaf (C. tesselata, C. seganica, C. hoseana, C. angustielliptica); in C. penduliflora, they may be present or absent. The other species in section Pleuroschisma lack sclereids. (C. insolita and sp. no. 12 have not been examined). Protection of the apical bud in Cyrtandra is achieved in several ways. In section Pleuroschisma the bud is hidden between the erect petioles of the two fully developed uppermost major leaves on the stem. The following observations were made on herbarium material, and observations are needed on living plants to determine time-intervals in leaf development: when the developing leaf in C. /inauana measured 38 mm, the young apical bud was only 2.5 mm long. We examined the stem apex of C. linauana, removing the tip of the stem and rehydrating it. The very young major leaf of the apical bud is hidden by the conduplicate base of the minor (stipule-like) leaf of the uppermost pair of fully developed leaves. It is plane, measured 38 x 8 mm including the petiole (scarcely developed at this stage), and both surfaces are thickly clad in long silky hairs. The minor leaf has similar indumentum and 1s well developed, being 20 mm long. In contrast, the fully developed leaves below the apex show no trace of the juvenile indumentum except over the midrib and lateral veins on the lower surface. There is the same abrupt transition from a very small juvenile leaf to a fully developed one in five other species where we had stem-apices suitable for inspection (C. penduliflora, C. sarawakensis, C. coacta, C. angustielliptica, C. insolita). These species in section Pleuroschisma are all strongly anisophyllous. A comparable condition in isophyllous species was noted by Burtt (2001, p.399). Field observations by one of us (B.L.B.) found that in C. hoseana and C. penduliflora, as the new apical leaf begins to grow out, the petiole is held erect but the young lamina is pendulous, the hairs in C. hoseana being silvery white, while in C. penduliflora they are dark red. As the leaf blades enlarge, the hairs become more spread out, and are eventually shed, resulting in the blade of the mature leaf being glabrous above. Whether or not these young leaves perform any attractive function is not known, but they certainly catch the eye of the plant-hunter. Cyrtandra section Pleuroschisma Hilliard & B.L.Burtt sect. nov. ab sectionibus omnibus ad huc descriptis fructibus duris laevibus (strato annulari subepidermali schlerenchymatis uni- vel bi-seriato), parietibus in medio longitudinis fissuris duabus septicidalibus marginibus involutis notatis, foliis cellulis tracheoidalibus hypodermalibus praeditis facile distinguenda. Cyrtandra sect. Pleuroschisma WN | | 39 Ay 4 4 ie! = ~ i a> Ss —— ——— ~~ — ~ Figure 3. Transverse sections of part of fruit wall and septum A. Cyrtandra sarawakensis; B. Cyrtandra penduliflora. b. scl. = brachysclereid; fib. 1. = fibrous layer; m. scl. = macrosclereid; scl. p. = sclerified parenchyma [sclerenchyma]; sp. scl. = spheroidal sclereid; tr. = tracheoid [with reticulate thickening]. M.H. Bokhari del. Figure 4. Sclereid and tracheoid types. A. polymorphic sclereids. B. tracheoid with reticulate bars. C. tracheoid with parallel bars. M. Mendum after pencil sketches by M.H. Bokhari. 40 Gard. Bull. Singapore 55 (2003) Species typica: C. sarawakensis C.B.Clarke Distribution: Endemic to Borneo. Simple-stemmed perennial herbs (C. insolita bushy), leaves usually strongly anisophyllous, tracheoids, as far as is known, in hypodermis, inflorescence a dichasial cyme, bracts free, bracteoles present or absent, calyx divided almost to base, corolla medium-sized, white to palest yellow ground colour, disc cupular, ovary crowned with a coma of relatively long hairs, style pubescent, stigmatic lobes large, clavate, fruit with two septicidal grooves (later splits). Key to Species la. Lateral veins in largest mature leaves up to 10 each side of midrib ............... 2 ib, ‘Lateral veins 14-20 on each side of madrid... a eee ~ 2a. Hairs (lower leaf surface) on midrib and lateral veins very inconspicuous, short (to | mm), strongly appressed, pedicels c. 20 mm long,calyx lobes 10-15 tats... eae ee a ee ee 8. C. linauana 2b. Hairs (lower leaf surface) on midrib and lateral veins long, silky, strongly appressed, sometimes matted together to produce a papery surface, pedicels c. 5—8 mm, calyx lobes to'8 mii) ree eee 3 3a. Largest leaves c. 60-165 mm broad, petioles 75-135 mm, bracts c. 35— 40 x 10—16 mm, conspicuous, stronely veined. ::... 2.25.2... ae 3. C. coacta 3b. Largest leaves c. 36-60 mm broad, petioles 40-45 mm, bracts c. 5.5—7 x 1.8—2 mm, inconspicuous, only midrib visible.............. 10. C. angustielliptica 4a. Hairs (lower leaf surface) on midrib spreading: .... 200, clk. a. ..s0ek-seeeeesct mean 5 4b. Hairs (lower leaf surface) on midrib strongly appress................::sssecceeeeeeeees 10 5a. Peduncle c. 45-240 mm long, inflorescence lax, trailing..............eeeeeseeeeeeeeeeeeee Pie: omy te 1. C. penduliflora var. grossipilosa 5b. Peduncle c. 2-30 mm long, inflorescence neither lax nor trailing.................. 6 6a. Bracts c. 20-50 x 7-25 mm, 3-5 veins, clearly visible, calyx lobes 2A MAREE TOM ees Sead St ae a a leek ae eee 7 6b. Bracts c.15—18 x 2—3 mm, only midrib visible, calyx lobes ©. 6-10 TM TOTS oie occ eccce deed, Din, babdnetdaetesncs eee ttt. Aa state ge en ee 9 7a. Inflorescence a several- to many-flowered cyme, both bracts and bractéolés present... ...050..0. 00. 82. ee eee 8 7b Flowers 1-3, clustered and, at least when young, hidden by two foliaceous bracts, bracteoles wanting. ...............cssssssseeeeeeeeeeeeeeees 11. C. insolita 8a. Petioles 80-150 mm long, peduncles 5—30 mim...............cccceecesceeeeeeeeeeeeeeeeeeeeeeeees sauaind adboonvivesniNootecd seed dadana easel pl ee ee 2. C. sarawakensis var. longipilosa Cyrtandra sect. Pleuroschisma 4] 8b. Petioles 30—70 mm long, peduncles 0—2 mm............... 5. Cyrtandra sp. nov. 9a. Largest leaves c. 26-34 mm broad, lower surface displaying finely reticulate tertiary venation, petiole c. 20—25 mm long ............ 6. C. tesselata 9b Largest leaves c. 85—110 mm broad, lower surface displaying coarsely reticulate venation, petiole c. 45—50 mm long........... 12. Cyrtandra sp. nov. 10a. Peduncles c. 50-175 mm long, inflorescence lax, trailing... MPT eisintex. 0k). neo) er, SL. BAO... ae ee 1. C. penduliflora 10b. Peduncles c. 1-30 mm long, inflorescences neither lax nor mbit os Be es, a EO) 3 be ade RL a ail WR 11 ibaai ise. 60-20 ann booed cece UA heck coe tO RR ES 12 RAG ) aeaeee ond 1S bread a3 41h. een ahs; ae id is en NII ‘3 12a. ‘Tertiary venation on lower leaf surface subscalariform, hairs on stem, leaves (including petioles), bracts, bracteoles, pedicels and calyx appressed, all: hairs. separatel ncveli icc egl. wisn Ab sdae des 2. C. sarawakensis 12b. Tertiary venation more or less pinnate, hairs on all parts (as above) matted together giving a papery look to the surface................... 3. C. coacta 13a. Stem silky-villous, tertiary venation (lower leaf surface) coarsely reticulate, almost invisible, largest mature leaves c. 85—95 mm broad, foisbengthiz breath wo 0e3G: beniys. hd enh Qed 9. C. hoseana 13b. Stem pubescent, tertiary venation more or less subscalariform, leaves c. 35-60 mm broad, ratio length: breadth 4—-5.7:1........ eee 14 14a. —_ Largest leaves c. 35-50 mm broad, lateral veins 13—15 on each side of midrib, bracts 20—30 x 4—7.5 mm, at least midrib clearly ie aa oe eet hires. Dada. heed Sao r9 tate. a RSI 4. C. tunohica 14b. — Largest leaves c. 55-60 mm broad, lateral veins 18 on each side of midrib, bracts c. 20 x 4 mm, midrib scarcely ae ry ee ie Behar tit osiencssisvedass oonssanccsievcassse 7.C. seganica 1. Cyrtandra penduliflora Kraenzl., Mitt. Inst. Allg. Bot. Hamburg 7:100 (1927); B.L. Burtt, Notes Roy. Bot. Gard. Edinb. 30:36 (1970). Lectotype (Burtt, 1970): Kalimantan, Bukit Obat [c.0°56'N 113°20'E], 150m, Winkler 1328 (lecto HBG, isolecto E). Unbranched herb, stem to c. 600 mm long, 9-16 mm in diam., base often decumbent, rooting, prop roots present, young parts strongly appressed-pubescent. Leaves opposite, strongly anisophyllous, few forming a fan at apex of stem, reduced leaves stipule-like, c. 20-46 x 5-14 mm, lanceolate, acuminate, strongly appressed- pubescent, largest developed leaves c. 210-300 x 150—200 mm, very broadly elliptic, apex abruptly acute, base cuneate, very narrowly decurrent, briefly or to c. 40 mm, margins entire to obscurely to more distinctly irregularly serrate, lateral veins 1 1—13 on each side of midrib, tertiary venation more or less pinnate, upper surface glabrous 4? Gard. Bull. Singapore 55 (2003) at maturity, coarsely pitted, lower surface with fine appressed hairs on veins and blade; petiole 130-210 mm long, hairy as midrib. /nflorescence an axillary very lax dichasial cyme, loosely branched, trailing, tending to spring from axils of fallen leaves, peduncle c. 50-175 mm long, pubescent. Bracts (lowermost pair) c. 15—57 x 4-18 mm, lanceolate, acuminate, finely pubescent. Pedicels 6-20 mm long, puberulous. Calyx 5-lobed almost to base, lobes subequal, c. 6-14 x 1.5 mm, deltoid, outside puberulous, inside minutely gland-dotted. Corolla c. 37 mm long (no complete flowers seen, only a big bud), white with an orange-yellow blotch or two bars in throat, puberulous all round mouth. Stamens inserted c. 15 mm above base of tube, filaments c. 5 mm long, glabrous, anthers 3.5 x 1.8 mm, cohering face to face by a small ligature, connective glabrous or a very few glandular hairs. Disc 1 x 1.8 mm, cupular. Ovary 6 x 1 mm, minutely pustulate, coma of hairs at apex. Sty/e 12 mm, glandular-puberulous. Stigmatic lobes c. 1.5 x 1 mm (will enlarge), clavate. Fruit c. 20-45 x 5~7 mm, pericarp smooth. Seeds c.0.3 x 0.25 mm, testa red-brown. Notes: Cyrtandra penduliflora is a plant of the forest floor, often on steep banks, the stem supported by prop roots. It is distinctive by virtue of its trailing inflorescences, the peduncles mostly very long, the cyme always very lax, far laxer than in any other species of this group. The leaves are broad in relation to length, those of the type specimen, collected in southern Kalimantan, measuring roughly 300 x 200 mm, and exactly matched by those of Burtt 2610 from Sarawak collected 1° of latitude north of the type collection and at very nearly the same degree of longitude (1°56’N 113°06’E versus 0°56’N 113°20’E), but nearly all specimens have the leaves very broad in relation to their length. Tracheoids with parallel bars of thickening occur in the hypodermis; polymorphic sclereids are often present in the spongy mesophyll but they may be absent. Other specimens examined: Sarawak. Miri distr., Niah Forest Reserve, [c. 3°52'N 113°44'E], Anderson $31663 (E). Bintulu distr., Ulu Sungai Sinonok, 20-60 m, Hotta 14247 (E, KYO). Tatau, Batang Anap, Ulu Sg. Kana [c. 2°54'N 112°50'E], Mokhtar & Jugah 41779 (E). Sungai Kakus [c. 2°48'N 112°42'E], 30-80 m, Hirano & Hotta 220 (E, KYO). Ulu Kakus [c. 2°48'N 112°42'E], Anap, Haron S29980 (K, E). Ulu Segan [c. 2°30'N 113°E], N Setungan, c. 50 m, Ashton $22019 (E). Pelagus Rapids on Rajang [‘Rejang’], c. 2°10'N 113°E, Burtt & Woods B2559 (E). Sungai Bena [c. 1°56'N 113°06'E], a tributary of Sungai Sut, Burtt B26/0 (E). Sungai Jelok, near Bukit Sengkajang [1°14'N 111°31'E], Lanjak-Entimau P.F, c. 2100 ft, Chai S34034 (E, K). Bukit Ubah Ribu, Ulu Sungai Kaup, Lubok Antu [1°03'N 111°50'E], c. 200 ft, Chai S33761 (E). Cyrtandra penduliflora var. grossipilosa Hilliard & B.L.Burtt var. nov. a planta Cyrtandra sect. Pleuroschisma 43 typica pilis longis grossis patentibus in caulibus petiolis et subtus in costis (nec pilis tenuibus valde appressis) filamentorum apicibus et antherarum connectivo dense glanduloso-puberulo (nec glabro vel fere glabro) differt. Type: Sarawak, ascent to Gunung Mulu [4°O1'N 114°52'E], 13 vi 1962, Burtt & Woods B2060 (holo. E). Unbranched herb, stem up to | m long, 15 mm diam., base often decumbent, rooting, young parts villous, hairs coarse, spreading. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, largest developed leaves 190-370 x 85-160 mm, elliptic, apex acute to acuminate, base cuneate, very narrowly decurrent, margins entire to irregularly toothed, lateral veins 12—18 on each side of midrib, upper surface glabrous at maturity, coarsely pitted, lower surface with coarse spreading hairs scattered among short appressed ones on the midrib, similar appressed hairs on the blade; petiole 100—150 mm long, hairy as midrib. /nflorescence as in the typical plant, peduncle c. 45—240 mm long. Bracts c. 24-45 x 4-13 mm, finely pubescent. Calyx lobes c. 6-9 mm long, outside puberulous, inside minutely gland-dotted. Corolla (only one flower seen) 41 mm long, white to pale yellow with 2 orange-yellow bars in throat, outside puberulous, mouth glandular-puberulous inside, tube 27mm long, cylindric in lower part, expanded above, upper lobes c. 8 x 8 mm, lower lip c. 14 x 22 mm, median lobe 9 x 8 mm, all lobes suborbicular. Stamens inserted 22 mm above base of tube, filaments c. 5 mm long, glandular-puberulous in upper part, strongly twisted post anthesis, anthers 3 x 2 mm, cohering face to face by a small apiculus, connective densely glandular-puberulous. Disc 1 x 1.5 mm, cupular. Ovary 6 x 1.2 mm, very minutely pustulate, coma of hairs at apex. Style 13 mm long, glandular- pubescent. Stigmatic lobes 3 x 2.5 mm, clavate. Fruit c. 25-35 x 5 m, pericarp smooth. Seeds c. 0.4 x 0.2 mm, testa red-brown. Notes: This plant differs from typical Cyrtandra penduliflora in the coarse spreading hairs on the stem and along the midrib on the undersurface of the leaf (versus fine strongly appressed ones). The tertiary venation also differs subtly : in the typical plant, the tertiary veins are not clearly visible on the lower surface of the leaf and they are more or less pinnate in arrangement; in the variety they are clearly visible, subscalariform, with a fine reticulum of lesser veins between them. The leaves closely resemble those of C. sarawakensis, distinguished at once by its compact inflorescence. The label on Sands 5390 (cited below) described the leaves as ‘paler beneath with reddish-brown veins’; Church 2210 ‘whitish-green beneath with purple veins distinctly raised’. There is also a difference in the shape of the leaves, those of the typical plant being broader in relation to length than in the variety, ratio of length to breadth being 1.2—1.6:1 versus 1.7—2.5:1; in absolute terms c.150—200 mm versus 85—160 mm (largest leaves). 44 Gard. Bull. Singapore 55 (2003) Anatomically there is some difference in the thickenings of the tracheoids in the hypodermis: parallel bars in the typical plant, reticulate thickenings in the variety often mixed with others with parallel bars. The anther-connective in the typical plant is glabrous or nearly so, that in the variety densely glandular-puberulous. From what is known of the distribution of the species, the variety tends to be more northern and eastern than the typical plant though the few collections from Kalimantan indicate close proximity. Other specimens examined: Brunei. Temburong river at Wong Nguan rapids, 4°31'N L15°15'E, 120m alt., Coode 6649 (E). Temburong distr., Sungai Belalong, 4°30'N 115°10°'E, c. 100 m, Argent et al. 9/1 (E). Slopes above Temburong headwaters NE of Gunung Retak, 4°22'N 115°17'E, Sands 5309 (E). Tutong, Ladan Hills Forest Reserve, upstream from Belabau on E bank of Tutong river, 4°26'N 114°46'E, 20 m, Coode 6409 (E). Sungai Tokat, a branch of Sungai Batu-Apoi, 50-100 m, Hotta 13759 (E). Sarawak. Limbang [4°45'N 115°E], Ulu Medamit [4°27'N 114°55'E], Sungai Ensungei, Tanjong Long Amok, George et al. $4¢2368 (E). Gunong Api, Ulu Melinau, Tutoh, 4°07'N 115°15'E, 400 ft, Anderson S31784 (E). Melinau Gorge pathway, c. 4°05'N 114°50'E, Burtt & Woods B2239 (E). Gunong Mulu National Park, Gua Rusa, c. 4°02'N 114°50'E, Argent et al. 809 (E). Ulu Belaga, Sungei Semawat, 3°N 113°54'E, c. 250 m, Hansen 623 (E). Dulit Range [c.3°16'N 114°14'E], Sungai Sirui, 200 m, Awa & Yii $46629 (E). Valley of Sungai Keyan Linau-Balui divide, Sungai Nawai, c. 2°26'N 114°10'E, c. 2600 ft, Burtt B1 1456 (E). South of Punan Lusong, c. 2°28'N 114°12'E, Burtt B] 1294 (E). Kalimantan. Sintang, Bukit Baka National Park, Sungai Ella and environs, 0°38'S 112°17'E, 320m, Church 280 (A, E, HBG). Serawai, Uut Labang, 750 m, 0°36'6"S 112°38'56'E, Church 2210 (E). 2. Cyrtandra sarawakensis C.B. Clarke, DC., Monog. phan. 5:209 (1883); B.L. Burtt, Notes Roy. Bot. Gard. Edinb. 30:38 (1970). Lectotype (Burtt 1970): Sarawak, Kuching, July 1865, Beccari 153 (lecto FI, isolecto K, P). Unbranched herb, stem c. 110—230 x 7-20 mm, erect or decumbent and rooting, young parts densely appressed-pubescent, hairs c. 2 mm long. Leaves opposite, strongly anisophyllous, c. 3-8 forming a fan at apex of stem, reduced leaves stipule-like, c. 30 x 3 mm, lanceolate, acuminate, closely appressed-pubescent; largest developed leaves c. 225-300 x 57-125 mm, elliptic, apex very acute, base cuneate, very narrowly decurrent, margins subentire to serrate, lateral veins 16—20 on each side of midrib, tertiary veins subscalariform, upper surface glabrous at maturity, coarsely pitted, lower surface appressed-pubescent on blade, long (to 3mm) delicate appressed hairs on midrib and lateral veins (tawny or purple in life); petiole c. 80-150 mm, hairy as Cyrtandra sect. Pleuroschisma 45 midrib. /nflorescence a several-flowered congested cyme, borne mainly in axils of fallen leaves, peduncie c. 5—30 mm long, stout, puberulous. Bracts persistent, c. 20-35 x 7-20 mm, lanceolate, acuminate, margins entire or toothed, strongly veined, both surfaces appressed-pubescent; bracteoles similar but smaller. Pedicels c.10—12 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes subequal, 4.5—8 x 1—1.5 mm, narrowly deltoid, patent-puberulous outside, inside minutely gland-dotted. Corolla to c. 44 mm long, palest yellow, 2 orange-yellow bars on palate, outside puberulous, hairs acute, tube c.27 mm long, narrowly cylindric below, abruptly expanded above, two upper lobes c. 9 x 9 mm, lower lip 3-lobed, c.14 x 24 mm, median lobe c. 8 x 8 mm, all lobes suborbicular, glandular-puberulous inside, with patch of longer hairs adjacent to sinus of upper lobes. Stamens inserted 20-25 mm above base of tube, c.6mm long, strongly twisted after anthesis, minute glandular hairs fringing connective; lateral staminodes c. 2 mm long, posticous staminode c. 1 mm. Disc c.2x 2mm, cupular. Ovary c.7 x 1.4mm, minutely gland-dotted, coma of hairs at apex. Stigmatic lobes c. 3.5 x 1.2 mm, spathulate, conspicuous, stigmatic papillae relatively long. Fruit c. 20-25 x 4-5 mm, pericarp smooth. Seeds c. 0.25 x 0.2 mm, testa bright red-brown, reticulate. Notes: Cyrtandra sarawakensis was originally collected by Beccari at Kuching. Beccari had a hut on Mt. Matang, not far from Kuching, and this is possibly the type locality; Burtt & Woods B1947 (cited below) precisely matches the isolectotype specimen in Paris. The other collections seen came from the Semengoh Forest Reserve a few miles south of Kuching. Many species of Cyrtandra seem to have a very limited distribution and this may be the case here. The distinguishing features of C. sarawakensis are the many lateral veins, tertiary veins subscalariform and clearly visible especially on lower surface, hairs on veins (lower surface) strongly appressed, peduncles c. 5—30 mm long, bracts broad, strongly and conspicuously veined. It is a plant mainly of the forest floor. Tracheoids with parallel bars occur in the 1—2-layered hypodermis of the leaf, and there are no sclereids in the spongy mesophyll. Stomatal turrets are well developed. Other specimens examined: Sarawak. Mt. Matang [1°36'N 110°11'E], Burtt & Woods B1947 (E); Matang, Ulu Sungai Rayu, Lee $54099 (E); Semengoh Forest Reserve, 12 miles S of Kuching [1°28'N 110°22'E], Burtt & Martin B4712 (E); Semengoh Forest Reserve, Burtt & Woods B2487 (E). Cyrtandra sarawakensis var. longipilosa Hilliard & B.L.Burtt var. nov. a planta typica primum pilis in caule et costa grossis et patentibus (nec tenuibus et valde appressis) et calycis lobis plerumque longioribus 4.5—8 mm (nec 2.54 mm) et apicibus foliorum longe acuminatis (nec acutis) differt. 46 Gard. Bull. Singapore 55 (2003) Type: Sarawak [c.2°N 114°10'E, Hose Mts.], Melinau Community Forest, near base camp at Nanga Tunoh, Burtt & Martin B4772 (holo. E). Notes: The variety has been collected north and east of Kuching. It resembles Cyrtandra sarawakensis in nearly all features, including habit, number of lateral veins in leaves, subscalariform tertiary venation, bracts and other floral characters, but it differs in its long-acuminate (not merely acute) leaf tips, hairs on stem, petiole and midrib on under surface of leaves being coarse and spreading (not fine and closely appressed) and possibly also in length of calyx lobes, 2.54 mm (not 4.5—8 mm). The coarse spreading hairs are brightly coloured in life, and in the dried state the colour often persists in the cross walls of the hairs; the hairs on C. sarawakensis may also be coloured, but, being so much finer, the colour is more difficult to see in dried specimens. The occurrence of a variety with long coarse spreading hairs as opposed to appressed ones is paralleled in C. penduliflora, a species with tertiary venation indistinguishable from that of C. sarawakensis; C. penduliflora differs markedly in its lax inflorescence, and many specimens have fewer lateral veins in the leaves. Other specimens examined: Sarawak. Tatau, path to Bukit Buan, 300 ft. [c.2°56'N 112°55'E], Purseglove 5469 (E). Tatau, Ulu Anap [c. 2°56'N 112°49'E], Mokhtar S448]5 (E). Bukit Kana [2°42'N 112°54'E], 600-850 m, Hirano & Hotta 1299 (KYO); Bukit Kana, 50-150 m, Hirano & Hotta 15480 (E). Ulu Anap, Bukit Mersing [2°30'N 113°06'E], c.1000 m, Ashton $17679 (E). Hose Mts., Bukit Temedu [2°24'N 113°41'E], c. 1100 m, Ashton $19007 (E). Pelagus Rapids on Rajang River [‘Rejang’], also on Bukit Raya nearby, c. 2°10'N 113°E, Burtt & Woods B2544 (E). Bukit Raya, Smith § 27719 (E). SE end Hose Mts., Bukit Nibong, c. 2°6'W 113°42'E, Burtt & Martin B4845 (E). Sungai Bena, a tributary of Sungai Sut, c. 1°55'N 113°5'E, Burtt B2609 (E). Lundu distr., Gunung Gading [1°44'N 109°50'E], Burtt & Woods B2685 (E). Bukit Lumut, Ulu Amau [1°27'N 112°04'E], c. 950 m, Ashton $21264 (E). Bukit Goram, Ulu Sungai Kapit [1°34'N 112°45'E], Chai S36172 (E). 3. Cyrtandra coacta Hilliard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke venis lateralibus foliorum 9-15 (nec 15-20), venis tertiariis plus minusve pinnatis (nec subscalariformibus) pilis in bracteis et in foliorum pagina inferiore papyraceo- coactis (nec pilis singulatis manifestis) distinguenda. Type: Sabah, Ranau distr., road from Lohan to Mamut Copper Mine near Tank 47,6°O1'N 116°41'E, 1100 m, 9 vii 1984, Beaman 10656 (holo E). Cyrtandra sect. Pleuroschisma 47 Unbranched herb to c. 800 mm tall, stem horizonal at base then curving upwards, clad in a papery indumentum of matted hairs, leafy and floriferous on erect part. Leaves opposite, strongly anisophyllous, reduced leaf stipule-like, c. 25 x 2 mm, lanceolate, enveloped in papery indumentum; developed leaf c.180—360 x 60—110(—-165) mm, elliptic to broadly elliptic, apex acute to almost acuminate, base cuneate, very shortly decurrent, margins entire to obscurely or distinctly remotely serrate, lateral veins 9-15 on each side of midrib, tertiary veins more or less pinnate, upper surface glabrous, finely and closely pitted, lower surface papery, any loose hairs very delicate, toc.3mm long; petiole 75—135 mm, enveloped in papery indumentum. /nflorescence a several- flowered, very congested, axillary cyme, almost sessile. Bracts persistent, c. 35-40 x 10-16 mm, lanceolate, strongly veined, enveloped in papery indumentum, bracteoles similar but smaller. Pedicels c.5 mm. Calyx 5-lobed almost to base, lobes subequal, c. 3.5 x 1 mm, narrowly deltoid, tube c. 0.5 mm, outer surface with papery indumentum, inside minutely gland-dotted. Corolla c. 40 mm long, white or cream with 2 orange-yellow bars in throat below the two sinuses of lower lip, outside minutely puberulous, hairs acute, tube c. 25.5 mm, cylindric below for c.12mm then abruptly expanded, two upper lobes c. 9 x 6.5 mm, lower lip 3-lobed, c.14.5 x 12 mm, median lobe c. 9 x 7.5 mm, all lobes more or less suborbicular, glandular-puberulous inside and down tube to insertion of filaments. Stamens inserted c.19mm above base of tube, filaments c. 5 mm, strongly coiled post anthesis, glandular-puberulous at apex, anthers c. 2 x 1.5 mm, cohering face to face by a prominent ligature, connective densely glandular-puberulous; lateral staminodes c. 2 mm, posticous staminode c.lmm. Disc 1.5 x 2 mm, cupular. Ovary c. 8 xX 2 mm, minutely pustulate, coma of hairs at apex. Style c. 15 mm, glandular-puberulous. Stigmatic lobes c. 3 x 1.5 mm, clavate, conspicuous, stigmatic papillae relatively long. Fruitc. 20 x 6mm (Mendum & Lamb 234A), pericarp smooth. Seeds c. 0.5 x 0.25 mm, testa red-brown. Notes: This species has the aspect of Cyrtandra sarawakensis but is at once distinguished by the indumentum on the vegetative parts: on stem, leaves (including petioles), bracts, bracteoles, pedicels and calyx, the hairs are matted together (coactum meaning felt), giving a papery look to the surface; in C. sarawakensis, hairs on the corresponding parts are appressed, the individual hairs all separate. The very striking difference is easily seen along the midrib on the lower surface where the matted hairs of C. coacta form a papery-looking skin in sharp contrast to the strongly appressed but distinctly separate hairs in typical C. sarawakensis and the long coarse spreading hairs in var. Jongipilosa. The indumentum of C. hoseana and C. angustielliptica is not unlike that of C. coacta but both species differ, inter alia, in their narrow bracts lacking conspicuous venation. In C. coacta, there are 9—15 lateral veins on each side of the midrib, in C. sarawakensis 15—20, while the tertiary veins are more or less pinnate in C. coacta, 48 Gard. Bull. Singapore 55 (2003) subscalariform in C. sarawakensis. Furthermore, the leaves differ in anatomical detail: the tracheoids in the hypodermis of C. coacta have reticulate bars and there are no stomatal turrets; in C. sarawakensis the tracheoids have parallel bars and there are prominent stomatal turrets. The stamens may differ too, but too few flowers have been seen to be certain: in C. coacta, the connective in the anther and the top of the filament are densely glandular-puberulous; in C. sarawakensis both are glabrous or very minutely glandular. Cyrtandra coacta is known mainly from Sabah and the northern part of Sarawak (Yii et al. S5/691 came from the upper reaches of the Baleh River, in east central Sarawak), while C. sarawakensis is found in the southern part. Other specimens examined: Sabah. Sinsuron road, 850—900 m, 5°40'N 116°22'E, Mendum & Lamb 23 (E), Sinsuron road, Mendum & Lamb 23A (E); Lahad Datu distr., Research Centre, Ulu Sungai Segama, 200m, Argent et al. SAN 108203 (E). Mt. Kinabalu, eastern shoulder, 6°05'N 116°36-40'E, 3400 ft, Chew et al. 963 (E, K), Chew et al. 647 (E, K). Sarawak. Seventh division, Ulu Belaga, Sungai Semawat, 3°N 113°54'E, c. 250 m, Hansen 715 (E). NE slope Bukit Kama [2°42'N 112°54'E], c. 450 m, Hirano & Hotta 1239 (E, KYO). Ulu Sungai Balan, Bukit Batu Tiban, Yii et al. SS1691 (E). 4. Cyrtandra tunohica Hilliard & B.L.Burtt sp.nov. a C. seganica Hilliard & B.L.Burtt foliis minoribus (maxime 170—200 mm, nec 300 mm longis) et pro rata angustioribus (4—-5.7:1, nec 2.5-4.5:1), bracteis plerumque majoribus (20-30 x 4-7.5 mm prope basin, nec 20 x 4 mm), costa et venis lateralibus praecipue in pagina inferiore visibilibus (nec costa vix, lateralibus haud visibilibus), calycis lobis 2-4 x 1 mm (nec 4.5—-6 x1.5 mm) distinguenda. Type: Sarawak, Kapit distr., Sungai Melinau [Hose Mts.], Nanga Tunoh, c.2°3'40"N 113°38'20"E, c.450ft, Burtt B12664 (holo E). Unbranched herb, stem c. 5—6 mm in diam. near apex, height unknown, base decumbent and rooting, erect part toc. 150 mm long, upper part finely pubescent, soon glabrescent. Leaves opposite, crowded near apex of stem, slightly to strongly anisophyllous even on one stem, highly reduced leaves, when present, to c. 65 x 2mm, almost filiform, major developed leaves 170-200 x 35-50 mm, narrowly elliptic, apex acuminate, base narrowly cuneate, very narrowly and shortly decurrent, margins obscurely serrate to crenulate, lateral veins 13-15 (as few as 10 in minor leaves), tertiary venation subscalariform, upper surface glabrous at maturity, pitted, lower surface thinly hairy on blade, more densely so on veins, all hairs to c. 1 mm long, appressed; petiole 50— 70 mm long, hairy as midrib. /nflorescence: few-flowered congested cymes, axillary, Cyrtandra sect. Pleuroschisma 49 extending from near apex of stem to well down below the leaves, almost sessile. Bracts persistent, 20-30 x 4—-7.5 mm, lanceolate, long-acuminate, both surfaces puberulous, midrib and lateral veins clearly visible especially on lower surface; bracteoles similar but much smaller. Pedicels c. 1-8 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes subequal, 2-4 x 1 mm, narrowly deltoid, outside and margins closely appressed-pubescent, inside minutely gland-dotted. Corolla c. 30 mm long, cream with yellow marks on palate, outside puberulous, hairs acute, tube c. 20 mm long, lower half narrowly cylindric, upper part expanded, two upper lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe c. 5 x 3.5 mm, all lobes subrotund, glandular-puberulous inside, hairs extending down into throat. Stamens inserted c.14 mm above base of tube, filaments c. 4 mm long, strongly twisted, anthers 2 x 1.2 mm, cohering face to face by a small ligature, both anthers and filaments glabrous; lateral staminodes c. 2 mm, posticous staminode c. 0.5 mm. Disc 1—1.2 x 1.2—1.8 mm, cupular. Ovary c. 3.5—7 x 0.8—2 mm, very minutely gland-dotted, crowned with coma of relatively long hairs. Style 5-6 mm long, puberulous. Stigmatic lobes 3—4 x 1-2 mm, clavate. Fruit 10-18 x 4-6 mm, pericarp smooth, deeply grooved longitudinally. Seeds 0.3-0.5 x 0.2—0.3 mm, testa red-brown. Notes: The outstanding feature of Cyrtandra tunohica as we know it is variation in the degree of anisophylly; in the rest of the species in this section of Cyrtandra, the minor leaf is always reduced to a stipule-like outgrowth (with the caveat that several species are ill-known). In C. tunohica, the size of the leaves in each pair ranges from subequal (when the stem is much more leafy than is usual in this section as a whole) through degrees of reduction culminating in an extraordinarily long, almost filiform, stipule-like leaf; this occurs on a single stem. As the species is known from only one locality, the taxonomic value of this character remains to be assessed. The plants (very few) were found on the steep shady earth bank of a stream gulley, and were seen only once, despite further fieldwork in the area on this and other occasions. In its relatively narrow leaves, C. tunohica resembles both C. seganica and C. angustielliptica; it is easily distinguished from both by its bracts, 4-7.5 mm broad near the base (not 1.8—4 mm) and with both midrib and lateral veins raised on the lower surface and thus clearly visible (in contrast to nearly or quite invisible). It is further distinguished from C. seganica by its leaves, the largest 170-200 mm long (not c. 300 mm) and relatively narrower (ratio of length to breadth 4—5.7:1 versus 2.5—4.5:1), short (to c.1mm long) appressed hairs along the midrib on the lowersurface (not long coarse spreading hairs), and c.13—15 (not 18) lateral veins on each side of the midrib. Not only do the bracts differ from those of C. angustielliptica, but so do the leaves in several details: lateral veins 13-15 in major leaves (not 7-10), hairs on undersurface of leaves up to c.1mm long, sparse on the blade (in contrast to silky-villous, hairs c. 2 mm long, the blade eventually glabrous, hairs persistent on midrib), tertiary veins Clearly visible (not invisible or very nearly so). 50 Gard. Bull. Singapore 55 (2003) The leaf of C. tunohica has tracheoids with reticulate thickening in the hypodermis and lacks sclereids in the mesophyll; C. seganica and C. angustielliptica have similar tracheoids but differ in having polymorphic sclereids in the mesophyll. Other specimen examined: Sarawak. Kapit distr., Sungai Melinau [Hose Mts.], Nanga Tunoh, c. 2°3'N 113°38'E, Burtt B12665 (E). 5. Cyrtandra sp. nov. Unbranched herb, stem very stout, c. 12 mm diam., c. 300-600 mm tall, young parts villous, lower parts glabrous. Leaves opposite, few crowded near apex of stem, strongly anisophyllous, reduced leaves stipule-like, c. 40 x 2 mm, narrowly linear-lanceolate, villous; largest developed leaves c. 195-260 x 80-90 mm, elliptic or oblanceolate, apex abruptly acuminate, base narrowly cuneate, very narrowly decurrent, margins entire, lateral veins 16—18, tertiary veins finely reticulate, upper surface glabrous, coarsely pitted (at maturity; densely pubescent when juvenile), lower surface densely pubescent, veins pilose; petiole c. 30-70 mm long, villous. /nflorescence a many-flowered, highly congested dichasial cyme, axillary, forming bracteate clusters borne on the leafless part of the stem; peduncle very stout, to 2 mm long. Bracts persistent, primary pair c.20x7mm, lanceolate, acute, midrib and lateral veins prominent, outside densely pubescent, inside hairs mainly near apex and base; bracteoles similar but narrower, a pair subtending each pedicel. Pedicels c.5 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes c. 4 x 1 mm, outside puberulous, hairs patent, inside minutely gland-dotted. Corolla c. 30 mm long, white, 2 orange-yellow bars on palate, outside puberulous, hairs acute, tube c. 20 mm long, lower part narrowly cylindric, upper part expanded, two upper lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe c.5 x 4mm, all lobes subrotund, glandular-puberulous on lower lip, hairs extending down floor of tube, a median tuft of eglandular hairs at base of each upper lobe, a few glandular hairs towards outer margins. Stamens inserted c.17 mm above base of tube, filaments c. 4-5 mm long, twisted once near base, anthers 2.2 x 1 mm, cohering face to face by a conspicuous ligature, connective minutely glandular, glands extending briefly down back of filaments; lateral staminodes c. 2 mm, apex strongly hooked, tip globular, pale yellow (as anthers), posticous staminode c.1 mm. Disc 1.8 x 2 mm, almost cupular (deeply notched on one side). Ovary 7 x 2 mm, glabrous except for conspicuous coma. Style 10 mm (at anthesis), pubescent, hairs acute. Stigmatic lobes c. 2 x 1.5 mm (post anthesis), spathulate, stigmatic papillae conspicuous. Fruit 23 x 6 mm (one seen), pericarp smooth, split longitudinally, only apex and base cohering. Seeds c. 0.25 x 0.2 mm, testa red-brown. Notes: This species is allied to Cyrtandra tunohica and C. sarawakensis var. Cyrtandra sect. Pleuroschisma 34 longipilosa. It is distinguished from C. tunohica by the relatively long patent indumentum on stems, petioles and backs of leaves (versus hairs short, appressed), leaves c. 80-90 mm broad (not 35-50 mm) and many- (not few-) flowered inflorescence. It differs from C. sarawakensis var. longipilosa by its shorter petioles (30-70 mm, not 80—150 mm) and almost sessile inflorescence. Also, the pubescence on the undersurface of the leaf blade is dense and patent (not appressed). This is obvious in Haron $29978 (K), but not so well marked on the duplicate (E); it is also obvious in Jugah Kudi S23712 (K) where the finely reticulate tertiary venation shows up well. The two collections seen are not good: Haron S29978 was obviously pressed when badly withered; the sheet at Kew is particularly bad, but this is the one with a nearly complete corolla and several buds, from which the floral details were extracted. Only an old fruit survives on the sheet in Edinburgh, and the base of a fruit on Jugah Kudi S23712 appears to have eight loculi! The need for good collections before the species can be named is obvious. The leaves have a 1—2-layered hypodermis, and tracheoids with parallel bars in the hypodermis. Specimens examined: Sarawak, Anap, Ulu Kakus [c.2°30'N 113°E], Othman Haron $29978 (E, K); Balingian, Ulu Sungai Arip, Bukit [ju [roughly 2°40'N 112°40'E], Jugah Kudi §23712 (K, L). 6. Cyrtandra tesselata Hilliard & B.L.Burtt sp. nov. a C. tunohica (etiam folia anguste lanceolata gerente) caulibus et pagina foliorum inferiore villosis (nec breviter pubescentibus), petiolo 20—25 mm longo (nec 50—70 mm) villoso (nec pubescente), bracteis c.15 x 1.8—2 mm venatione invisibili (nec 20-30 x 4-7.5 mm venatione elevata conspicua), calycis lobis c.6—8 mm longis (nec 2.4 mm) differt. Type: Sarawak, Bintulu distr., en route (survey route) from Sungai Mah to Sungai Shinonok, Ulu Sungai Minah, alt. 40-150 m, Hotta 1/4117 (holo KYO). Herb, probably unbranched, height unknown, stem collected 50 x 10 mm, villous, hairs 3-4 mm long, leafy throughout, internodes c. 10 mm long. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 30 x 3 mm, narrowly lanceolate,long-acuminate, major mature leaves 150-230 x 26-34 mm, narrowly elliptic, apex gradually very acute, base narrowly cuneate, margins very obscurely serrulate, lateral veins 12—14 on each side of midrib, upper surface glabrous at maturity, pitted, lower surface villous, longest hairs (2-3 mm) on midrib, shorter on lesser veins, tertiary venation finely reticulate, all veins raised, hairs brownish (dried state) making venation very conspicuous; petiole 20—25 mm long, villous. Inflorescence a few-flowered axillary cyme, peduncle c. 4mm, very stout. Bracts c. 15x 1.8—2 mm, narrowly lanceolate, densely pubescent, venation obscure; bracteoles similar but 52 Gard. Bull. Singapore 55 (2003) Pedicels c.10mm (in fruit), pubescent. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes 6-8 x 1 mm in fruit, narrowly deltoid, outside and margins puberulous, hairs patent, inside minutely gland-dotted. Corolla not seen, only very young buds present, puberulous outside, hairs acute, lobes minutely glandular inside (tube scarcely developed). Stamens: filaments scarcely developed, anthers 2 x 0.8 mm, minutely glandular on margins of connective. Disc cupular. Ovary and style very young, style pubescent. Fruit c. 35 x 5 mm, pericarp smooth, traces of coma still visible at apex. Seeds c. 0.25 x 0.2 mm, testa bright red-brown. Notes: This species is so distinctive that lack of flowers should not preclude its formal recognition. The leaves are closely set down the 150 mm length of the piece of stem on the herbarium sheet, in contrast to most of its allies where the leaves appear to be quickly deciduous leaving a few, forming a fan, at the apex of the stem. Unfortunately, the collector gave no notes either on habit or habitat of his plant. The major leaves are short-petioled, narrowly elliptic, and at maturity glabrous above but villous below, with midrib, lateral and tertiary veins raised (the latter forming a fine reticulum) and conspicuous, partly owing to the dark hairs clothing them. Similar tertiary venation has not been seen in any other species in this section. Cyrtandra tesselata is diagnosed above against C. tunohica, which has similarly narrowly lanceolate leaves, but the indumentum on stem, petioles and leaf undersurface in C. tesse/ata is villous (not merely shortly pubescent), the petioles are remarkably short (20-25 mm versus 50-70 mm), the bracts are c. 15 x 1-2 mm, venation invisible (not 20-30 x 4—7.5 mm, venation raised and conspicuous), calyx lobes c. 6-8 mm long (not 24 mm). In sharp contrast to that of C. tesselata, the tertiary venation in the leaves of C. tunohica is rather indistinct, and tends to be subscalariform. The leaf anatomy of C. tesselata is unlike that of C. tunohica: vertically long tracheoids with parallel bars in the hypodermis, polymorphic sclereids in the mesophyll; the tracheoids in C. tunohica have reticulate bars and there are no sclereids in the mesophyll. 7. Cyrtandra seganica Hillard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke bracteis tantum c. 4 mm latis (nec 7-20 mm) sine venatione conspicua (nec costa et venis lateraliibus prominentibus) et folia angustiora prorata longitudinis (5.5:1 nec 2.54.5:1) facile distinguenda. Type: Sarawak, Bintulu distr., Ulu Segan [c. 2°30'N 113°E], rocky stream bank, 750 ft., Jlias Paie §27215 (holo E). Cyrtandra sect. Pleuroschisma 53 Unbranched herb, height unknown, stem c. 8 mm diam. near apex, apex densely pubescent, rest of stem quickly glabrous. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, c. 25 x 3 mm, lanceolate, acuminate, densely pubescent, hairs spreading; developed leaves c. 300 x 55—60 mm, elliptic, acute, base narrowly cuneate, very narrowly decurrent for c. 35 mm, margins entire, lateral veins c. 18 on each side of midrib, tertiary venation subscalariform, upper surface glabrous, pitted, lower surface with very small fine strongly appressed hairs on blade, coarse much longer (c. 1 mm) hairs on midrib and lateral veins, probably appressed and probably purplish in life (specimens well glued); petiole c. 55-70 mm long, hairy as midrib. /nflorescence: few-flowered congested cymes borne mainly in axils of fallen leaves, peduncle c. 4-8 mm long, very stout, puberulous. Bracts persistent, c. 20 x 4mm, narrowly lanceolate, acuminate, margins entire, both surfaces puberulous, hairs spreading, midrib scarcely visible; bracteoles similar but smaller. Pedicels c. 8-10 mm. Calyx lobed nearly to base, lobes subequal, 4.5—6 x 1.5 mm, narrowly deltoid, tube c. 0.5 mm, outside and margins puberulous, inside minutely gland-dotted. Corolla not seen. Fruit c. 18—36 x 4—5.5 mm, pericarp smooth. Seeds immature. Notes: Cyrtandra seganica is currently known with certainty only from the type collection. In the venation of the leaves, including the subscalariform arrangement of the tertiary veins, it resembles C. sarawakensis, but the leaves are narrower in relation to their length (ratio of length to breadth 5.5:1 versus 2.5-4.5:1). There is a striking difference in the bracts, only c. 4 mm broad near the base and with the midrib but faintly visible in C. seganica, in contrast to bracts c. 7-20 mm broad with prominent venation in C. sarawakensis. The leaves are similar anatomically in having tracheoids with parallel bars in the hypodermis, but differ in the presence of many polymorphic sclereids in the mesophyll of C. seganica whereas the leaves of C. sarawakensis lack sclereids. Two collections from further south and west resemble C. seganica in leaf-venation and in narrow bracts. They appear to differ in the shape of the leaves (oblanceolate) and also in the ratio of length to breadth (3.25—-4.6:1), this being within the range of C. sarawakensis. The leaves also appear to differ anatomically, in that the tracheoids have reticulate bars, not parallel bars; polymorphic sclereids occur in the mesophyll. These two collections come from Sarawak from Datu Permanent Forest [Bukit Datu? at 1°29'N 110°50'E], 420 m alt., on the ridge between two streams, Lee S§41904 (E) and 17" mile Bau/Lundu road, Gunong Undan (1° 27'N 109° 59'B), stream bank, 50m, Yii S45942 (E). Much more needs to be known about all three plants described here. 54 Gard. Bull. Singapore 55 (2003) 8. Cyrtandra linauana B.L.Burtt, species nova ex affinitate C. sarawakensis C.B.Clarke et C. hoseanae B.L.Burtt, ab ambabus nervis lateralibus foliorum 7-8 (nec 12—20) et calycis lobis 10-15 mm longis (nec usque ad 8 mm) differt. Type: Sarawak, Belaga distr., hill just N of Long Linau [c.2°40'N 114°E], Burtt 11478 (holo E; iso BO, L, SAR). Herb, stem simple, c.100 mm tall, 8 mm diam., puberulous, hairs very closely appressed. Leaves opposite, strongly anisophyllous, 5 pairs crowded near apex of stem, lower ones fallen, reduced leaf stipule-like, c. 35 x 5 mm, lanceolate, long-acuminate; developed leaf c. 210-300 x 82-83 mm, elliptic, somewhat falcate and eccentric, apex rather abruptly acuminate, base cuneate, very shortly and narrowly decurrent, margins entire, lateral veins 7-8 on each side of midrib, tertiary venation coarsely reticulate, upper surface mottled silver (living material), glabrous, pitted, lower surface with blade glabrous, midrib and lateral veins strongly appressed-pubescent, hairs very inconspicuous; petiole 70-80 mm, thickly appressed-pubescent. J/nflorescence an axillary dichasial cyme, flowers few, branches of cyme very short (3-4 mm), bearing only 4 flowers (apical bud on each branch suppressed), peduncle c. 5-8 mm, very stout. Bracts persistent, c. 20 x 2 mm, narrowly lanceolate, acuminate, both surfaces strongly appressed-pubescent, bracteoles similar but smaller, a pair subtending each pedicel. Pedicels c.20 mm, appressed-pubescent. Calyx 5-lobed almost to base, tube c.0.5 mm, lobes 10—15 mm, narrowly linear-lanceolate, outside appressed-pubescent, inside minutely gland-dotted. Corolla 26-30 mm long, white, throat yellow shading to orange, outside minutely puberulous, hairs acute, red, tube 17—18 mm long, cylindric in lower third, abruptly expanded above, two upper lobes 7—8 x 5 mm, lower lip 3-lobed, c. 9-12 mm long, median lobe 7—8 x 5 mm, all lobes oblong-elliptic, glandular- puberulous inside. Stamens inserted 8.5 mm above base of tube, filaments 2.5 mm, glabrous, anthers 2.2x 1.2mm, cohering face to face by a prominent ligature, connective fringed with stout red hairs; lateral staminodes 3 mm, posticous staminode 1.2 mm. Disc 1.5 x 2 mm, cupular but deeply excavated on one side. Ovary 5 x | mm, minutely pustulate, coma of hairs at apex. Style 7mm, glandular-puberulous. Stigmatic lobes 4x 1.8 mm, clavate, conspicuous. Fruit 24 x 3.8 mm, pericarp smooth. Notes: Cyrtandra linauana resembles both C. sarawakensis and C. hoseana in the size, shape and stance of its leaves but differs from both in having only 7-8 lateral veins on each side of the midrib (not 12—20). The tertiary venation also differs: in C. linauana it is rather indistinct and very coarsely reticulate though the veinlets may be more subscalariform towards the margins; in C. sarawakensis the veins are prominent and closely subscalariform; in C. hoseana they are very nearly invisible. Cyrtandra sect. Pleuroschisma 55 The tracheoids in the hypodermis have reticulate bars as in C. hoseana but there are no sclereids in the spongy mesophyll. In C. sarawakensis the tracheoidal bars are parallel and, as in C. linauana, there are no sclereids in the spongy mesophyll; C. hoseana has polymorphic sclereids. The inflorescences of all three species differ: in C. /inauana, the bracts are only c. 2 mm broad and the venation is not or scarcely visible; pedicels c. 20 mm long, making the inflorescence rather lax. In C. sarawakensis, the bracts are very conspicuous, being broad, venation prominent, pedicels c. 9-13 mm long, inflorescence congested. In C. hoseana, the bracts are c. 34 mm broad, venation inconspicuous, pedicels c. 5-10 mm long, inflorescence congested. Stout red hairs fringe the connective in C. linauana; in C. sarawakensis and C. hoseana, the connective is either glabrous or glandular. Cyrtandra linauana is known with certainty only from the type collection made at c. 2°40'N 114°E. There is another collection, made at c. 2°48'N 112°59'E, that is clearly very close to C. linauana, namely Hirano & Hotta 88] (KYO, E) from Bukit Keyan, Ulu Sungai Kakus, alt. 100-300 m. It differs in several details: leaves not falcate and with 10 lateral veins, otherwise identical with those of C. linauana, flowers fascicled in the leaf axils, pedicels c. 15 mm long, calyx lobes 6.5—9 mm. The most significant difference is the fascicled flowers in contrast to the shortly pedunculate dichasial cyme of C. linauana, where the long pedicels produce a very lax inflorescence. The leaves of C. angustielliptica have a deceptive similarity to those of C. linauana. That species is, however, distinguished by the short (to c. 1 mm) appressed and inconspicuous hairs on stem, petiole and midrib of leaves on lower surface (not 2—3 mm long, silky, appressed hairs matted together and easily seen), bracts c. 20 mm long (not 5—8 mm), connective of anthers fringed with stout red hairs (versus glandular puberulous ones all over). The inflorescence of C. angustielliptica is a nearly sessile, very congested, cyme. 9. Cyrtandra hoseana B.L.Burtt, Notes Roy. Bot. Gard. Edinb. 30:33 (1970). Type: Sarawak, Lambir Hills, c. 4°7'N 113°55'E, sandstone banks, Burtt & Woods B2365 (holo E, iso SAR). Unbranched herb, stem 20-150 cm long, c. 8 mm diam. near apex, hanging from sandstone cliffs and banks, sometimes supported by prop roots, young parts silky villous, hairs strongly appressed. Leaves opposite, strongly anisophyllous, few arranged in a fan at stem apex, reduced leaves stipule-like, c. 20 x 10 mm, broadly lanceolate, appressed silky-villous; largest developed leaves c. 255-320 x 85-95 mm, elliptic, apex abruptly acute, base cuneate, shortly and very narrowly decurrent, margins entire to obscurely serrulate, lateral veins 12-15 on each side of midrib, tertiary venation coarsely reticulate, upper surface glabrous at maturity, coarsely pitted, lower 56 Gard. Bull. Singapore 55 (2003) surface with long, strongly appressed, silky hairs on midrib and lateral veins, blade glabrous or almost so at maturity; petiole c. 45-60 mm, hairy as midrib. /nflorescence an axillary, very congested, cyme, mainly in axils of fallen leaves, few-flowered, peduncle 2-6 mm long. Bracts c. 15-18 x 34 mm, persistent but inconspicuous, lanceolate, acuminate, closely appressed-puberulous, only midrib visible on lower surface. Pedicels c. 5—10 mm long, puberulous. Calyx 5-lobed almost to base, lobes subequal c. 2.2-3.5 x 1.25 mm, deltoid, outside appressed-pubescent, inside minutely gland-dotted. Corolla c. 32 mm long, white or creamy white outside, 2 orange-yellow bars in throat, outside puberulous, hairs acute, red, tube c. 20 mm long, lower half narrowly cylindric, abruptly expanded above, two upper lobes c. 5 x 7 mm, lower lip 3-lobed, c.12.5 x 20 mm, median lobe c. 6 x 7 mm, all lobes subrotund, glandular- puberulous inside and for a short way down tube. Stamens inserted c. 16 mm above base of tube, filaments c. 5 mm, curved and twisted once, minute globular glands at apex, anthers c. 3 x 1.6 mm, cohering face to face by a small ligature, connective with a few glandular hairs and globular glands; lateral staminodes c. 1.2 mm, posticous staminode c.lmm. Disc 1 x 1.8 mm, cupular. Ovary 8 x 1.6 mm, very minutely gland-dotted, coma of hairs at apex. Style 10mm, glandular-puberulous. Stigmatic lobes not fully developed, c. 2 mm long, clavate. Fruit c. 27-45 x 5 mm, pericarp smooth. Seeds c. 0.4 x 0.25 mm, testa red-brown. Notes: So far, C. hoseana is known with certainty only from the Lambir Hills. Now that more material is available, it is clear that Burtt & Woods B2212 (E, SAR) from the Melinau Gorge, which was cited in the original description, is a distinct species, described here as C. angustielliptica. It is worth noting again that ‘the youngest leaf is very densely covered with shining silky hairs and it hangs downwards at the top of the shoot, like a tiny silvery flag’ (Burtt, 1970). In this species, the tertiary veins are scarcely discernible on the lower surface of the leaf, but form a coarse reticulum that may just be visible on the upper surface (compare the tertiary veins of C. sarawakensis). Tracheoids with reticulate bars occur in the 1—2-layered hypodermis, and a few polymorphic sclereids in the mesophyll. Other specimen examined: Sarawak. Lambir National Park, Sungai Lapoh and nearby, Burtt 11546 (E). 10. Cyrtandra angustielliptica Hilliard & B.L.Burtt sp. nov. a C. hoseana B.L.Burtt foliis 36-60 mm latis (nec 85-90 mm) apice acuminato (nec abrupte acuto), venis lateralibus utrinque costae 7-10 (nec 12-15), calycis lobis 5-8 mm longis (nec 2.2—3.5 mm) distinguenda. Cyrtandra sect. Pleuroschisma S¥ Type: Sarawak, Sungei Melinau Gorge, c. 4°5'N 114°50'E, sandstone hillock, Burtt & Woods B2212 (holo E). Unbranched herb, stem of unknown height, c. 6-7 mm in diam. near apex, young parts silky-villous, hairs strongly appressed, matted together. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 18 x 6 mm, lanceolate, silky-villous as stem, developed leaves up to c.10, largest 170-275 x 36-60 mm, elliptic, apex acuminate, base cuneate, margins entire to obscurely serrulate but teeth made conspicuous by apical tuft of hairs, lateral veins 7—10 on each side of midrib, tertiary venation obscure, upper surface glabrous at maturity, coarsely pitted, lower surface with long, strongly appressed, silky hairs on midrib and lateral veins, these also on blade but this eventually glabrous; petiole c. 40-45 mm, hairy as midrib. /nflorescence an axillary very congested cyme, flowers few, peduncle c. 2 mm. Bracts c. 5.5—7 x 1.8—2 mm, persistent but inconspicuous, lanceolate, acuminate, both surfaces densely appressed-pubescent. Pedicels c. 8 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm long, lobes subequal, 5—8 x 1.2 mm (in flower and fruit), deltoid, outside densely appressed-pubescent, inside minutely gland-dotted. Corolla c. 30-35 mm long, white or creamy white outside, 2 orange-yellow bars in throat, outside puberulous, hairs acute, tube c. 22—28 mm, lower half cylindric, upper part expanded, two upper lobes c. 5 x 5 mm, lower lip 3-lobed, median lobe c. 6 x 7 mm, all lobes subrotund, glandular-puberulous inside and short way down tube. Stamens inserted c. 16 mm above base of tube, filaments c. 5 mm, curved and twisted once, glandular-puberulous at apex, anthers c. 2 x 1.5 mm, cohering face to face by a small ligature, connective glandular-puberulous; lateral staminodes c. 2 mm, posticous staminode not seen. Disc c.1.5x 1.8mm, cupular. Ovary c.6x 1.2 mm, very minutely papillose, coma of hairs at apex. Style c.6 mm elongating to c.10 mm, glandular-puberulous. Stigmatic lobes c.3x 2mm, clavate. Fruit 25-48 x 5 mm, pericarp smooth. Seeds c. 0.3 x 0.2 mm, testa red-brown. Notes: In his original description of Cyrtandra hoseana, Burtt cited Burtt B2212 (above) as that species, but in the herbarium he later segregated it and further collections as differing in their narrower leaves. Not only are the leaves narrower, they are also shorter (170—275 x 36-60 mm versus 255-320 x 85-95 mm) and there are significantly fewer lateral veins than in C. hoseana (7-10 versus 12-15); also, the leaf tips are acuminate (not abruptly acute). They may differ further in the length of the calyx lobes (5—8 mm versus 2.2—3.5 mm, but see comments below). As in C. hoseana the tertiary veins are scarcely visible on the lower surface of the leaf; anatomically the leaves are similar, having tracheoids with reticulate bars in the hypodermis and polymorphic sclereids in the mesophyll. Cyrtandra angustielliptica is known only from the Melinau Gorge (in Gunung Mulu National Park) at c.100 m above sea level. A plant with leaves exactly like those of C. angustielliptica in all morphological details has been collected several 58 Gard. Bull. Singapore 55 (2003) times on the extreme headwaters of the Balleh, at c.1000—1700 ft altitude on a ridge between Sungai Balang and Sungai Balleh [1°35'N 114°30'E at 1200 ft by Anderson & Paie S28329 (E); at c.1000 ft by Anderson & Paie 28739 (E); at c.1000 ft by Anderson $2874] (E) and at 1700 ft by Paie S28409 (E, L)]. However, these specimens differ anatomically in having no sclereids in the spongy mesophyll (a character possibly of little taxonomic significance), and the calyx lobes are 2.5—5 mm long (not 5—8 mm), corolla 20—25 mm (not 30—35 mm), filaments and anthers glabrous (not glandular), and the disc unilateral (not cupular). Other specimens examined: Sarawak. Gunung Mulu National Park, Melinau Gorge, Burtt B8274 (E); Melinau Gorge, c. 100 m, Nielsen 537 (E). 11. Cyrtandra insolita Hilliard & B.L.Burtt sp. nov. nullae arcte affinis; pedunculo longo axillari bracteis duabus flores breviter pedicellatos 1—3 involucrantibus facile recognoscenda. Type: Sarawak, Kapit, Upper Rajang [“Rejang’] River, 1929, Clemens 21236 (holo K). Shrub (fide collector), 3 stems seen, 170-250 mm long, woody, leafy only at apex, young parts densely pubescent. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 20 x 3 mm at base, linear-lanceolate, villous; developed leaves few, c.160—190 x 55 mm, elliptic, apex long-acuminate, base cuneate, decurrent down petiole, margins obscurely serrulate, lateral veins c. 20 on each side of midrib, tertiary veins invisible, upper surface silky villous at first, soon glabrescent, coarsely pitted, lower surface densely pubescent, hairs very delicate, c.4mm long on midrib, shorter on blade; petiole c.10mm long, villous. /nflorescence mainly in axils of fallen leaves, 1—3-flowered, flowers springing from apex of peduncle and enfolded by bracts, peduncle 15-45 mm long, villous. Bracts 2, persistent, foliaceous, c. 35-45 x 10—22 mm, lanceolate, long-acuminate, 5-nerved, both surfaces villous; bracteoles wanting. Pedicels 4-5 mm, pubescent. Calyx 5-lobed almost to base, lobes subequal, 2 x 1.5 mm, deltoid, outside pubescent, inside gland-dotted. Corolla not seen. Anthers (in very young bud) 1.5 x 1.1 mm, glabrous. Gynoecium very young, but stigma clearly bilobed, disc cupular. Fruit c. 30 x 5 mm, pericarp smooth (young fruits retain an apical coma). Seeds c. 0.3 x 0.25 mm. Notes: Cyrtandra insolita (insolitus = extraordinary) is highly distinctive among other species in this Section, from which the pair of foliaceous bracts enfolding one to three flowers at the apex of the long peduncle immediately mark it out. The other species all have few- to many-flowered cymes. Also, these species are unbranched herbs of the forest floor; Mrs Clemens described her plant as ‘shrub in thickets’. Cyrtandra sect. Pleuroschisma aby 12. Cyrtandra sp. nov. Unbranched herb, stem woody, height unknown, 10 mm diam., pilose, hairs coarse, spreading, red-purple, as on other vegetative parts. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, c. 75 x 6 mm at base, linear-lanceolate, long-acuminate, pilose; largest developed leaves 300-350 x 85-110 mm, elliptic, apex acuminate, base narrowly cuneate, decurrent, margins finely and remotely toothed (teeth being extensions of the veins beyond the margins), lateral veins 15—16 on each side of midrib, tertiary veins coarsely reticulate, upper surface glabrous, pitted, lower surface with short, fine appressed hairs on blade, long coarse spreading hairs on midrib; petiole c. 45—50 mm long, hairy as midrib. /nflorescence a congested several-flowered cyme, solitary in axils of fallen leaves, peduncle c. 6-12 mm long, pubescent. Bracts persistent, c. 18 x 3 mm, lanceolate, acuminate, midrib visible on lower surface, both surfaces densely pubescent; bracteoles similar but smaller, a pair subtending each pedicel. Pedice/l c. 20 mm long (in fruit), densely puberulous. Calyx 5-lobed almost to base, tube c. | mm long, lobes subequal, c. 10 x 1.5 mm (in fruit), narrowly deltoid, outside densely puberulous, inside minutely gland-dotted. Corolla not seen. Fruit c. 30 x 5.5 mm, pericarp smooth, remains of coma visible at apex. Seeds not fully ripe. Notes: This plant needs to be re-collected. Gunung Gaharu lies c. 96 km SE of Kuching on the road to Serian. The specimen seen is a unicate, in fruit but without flowers. It much resembles Cyrtandra sarawakensis, particularly var. longipilosa, but is at once distinguished by the reticulate, not subscalariform, tertiary venation of the leaves, and by its much narrower bracts. Specimen examined: Sarawak, Gunung Gaharu, c. 2000 ft., Burtt 2658 (E). Acknowledgements We thank the curators of the herbarium at Royal Botanic Gardens, Kew (K), Botany Dept., Kyoto University (KYO), the Nationaal Herbarium Nederland, Universiteit Leiden Branch (L) for the invaluable loan of specimens. We are also grateful to several of our colleagues: Frieda Christie, electron microscope unit; Mary Mendum (Bates) for art work; Adele Smith for stepping into the breach in a typing crisis; and Debbie White, photographic unit. We also thank the Regius Keeper, Steve Blackmore, working facilities at the Garden. 60 Gard. Bull. Singapore 55 (2003) References Bokhari, M.H. and B.L. Burtt. 1970. Studies in the Gesneriaceae of the Old World XXXII: Foliar sclereids in Cyrtandra. Notes Royal Botanic Garden Edinburgh. 30: 11-21. Burtt, B.L. 1970. Studies in the Gesneriaceae of the Old World XX XIII: Some species of Cyrtandra chiefly from Borneo. Notes Royal Botanic Garden Edinburgh. 30: 23-42. Burtt, B.L. 2001. A survey of the genus Cyrtandra (Gesneriaceae). Phytomorphology Golden Jubilee Issue 2001: Trends in Plant Sciences. pp. 393-404. Garden’s Bulletin Singapore 55 (2003) 61-63 Begonia peninsulae (Begoniaceae), a Confused Species from Malesia RUTH KIEW Singapore Botanic Gardens, Singapore 259569 Abstract Begonia peninsulae Irmsch. subsp. peninsulae from Trengganu, Peninsular Malaysia, is synonymous with B. rajah Ridl. and B. peninsulae subsp. tambelanense Irmsch. is raised to specific rank as B. tambelanense (Irmsch.) Kiew. Introduction Irmscher (1929) recognised two subspecies within his Begonia peninsulae: ssp. peninsulae from Trengganu and ssp. tambelanense from the Tambelan Islands. These were both based on a mixture of plants cultivated in the Singapore Botanic Gardens and the Penang Botanic Garden. The ones collected from the Singapore Botanic Gardens in Ridley’s time (one from Trengganu; the other from the Tambelan Islands) have a clear provenance. However, later collections are without provenance and, in the case of those from Penang, are from batches that include many exotic ornamental begonias. Indeed, there is no evidence to suggest that they are plants of wild-collected origin. In addition, these two subspecies are clearly not conspecific as one has a cordate leaf and the other a peltate one. Subspecies peninsulae Irmscher cited one specimen from Trengganu (which was spelt ‘Tringganu’ on the label), which is without a number or information on the date and collector. The botanical name is written as ‘Begonia raja’. ‘Ridley’ has been added as the collector in a different hand after Irmscher’s publication. This specimen matches those of Begonia rajah Ridl. in characters of the leaf, inflorescence and fruit. Despite the leaf being exceptionally large (11.5 x 16 cm as opposed to 5.5—7.5 x 5.5—7.5 cm in wild plants of B. rajah), the leaf base, like B. rajah, is deeply cordate and, even in the dried state, the variegation is conspicuous with the veins lighter than the rest of the lamina. This specimen might be one of the 62 Gard. Bull. Singapore 55 (2003) plants originally collected in Trengganu (which Ridley spelt as Tringganu) or its progeny that was cultivated in the Botanic Gardens Singapore from where it was introduced into the ornamental trade in England. B. peninsulae ssp. peninsulae is therefore reduced to synonomy with B. rajah. The other specimens of ssp. peninsulae cited by Irmscher are very different in that they are all peltate (B. rajah never has peltate leaves). These specimens, Burkill Begonia No. 16 July 1917 (SING), Burkill 6114 31 Aug 1920 (SING), were cultivated in the Penang Botanic Garden and have no provenance. In leaf size, shape, colour, surface (‘coppery and bullate’) and margin they are identical to Begonia goegoensis N.E. Brown from Sumatra, which is one of the few Asian begonias that is easy to grow under local conditions. Begonia rajah Rid. Gard. Chron. II, 16 (1894) 213 & Fig. 31. Type: Native Collector s.n. 1892 “Tringganu’ (Trengganu) (holo K ex SING’) Synonym nova: Begonia peninsulae Irnscher ssp. peninsulae, Mitt. Inst. Allg. Bot. Hamburg. 8 (1929) 98. Type: ‘Tringganu’ without collector, number or date (K ex SING’). Subspecies tambelanense Irmscher described this subspecies (a begonia with peltate leaves) from a collection by Ridley made in 1895 from a plant that was cultivated in the Botanic Garden Singapore. This, in turn, was originally collected by Pereira from the Tambelan Islands, NW of Pontianak, Indonesian Borneo. It is very different from the specimen of his subsp. peninsulae (=B. rajah) from Trengganu as not only are the leaves peltate with a long acumen but the petiole and lower surface of the veins are conspicuously hairy. (Those of B. rajah are deeply cordate with a short acumen and are more or less glabrous). Again, Irmscher included plants of unknown provenance collected from both Gardens — Nur Begonia No. 2 (SING), Nur Begonia No. 3 (K, SING) both dated March 1917 from the Botanic ‘When Irmscher monographed the begonias of Peninsular Malaysia he borrowed the entire collection of the Singapore Botanic Gardens (SING) and these he annotated and cited in his monograph. He did not consult the collection at Kew (K) and none of their specimens is cited by him. A considerable number of unicate types are no longer to be found at SING but are now at K. We have no record of how they came to be transferred but they are definitely the Singapore types as they bear Irmscher’s original annotations. These are cited as K ex SING in this account. Begonia peninsulae 63 Gardens Singapore; and Burkill Begonia No. 1 (SING) July 1917, Burkill 6115 (SING) 31 Aug 1920 from the Penang Botanic Garden. These are among a gathering of begonias by Burkill that included ornamental begonias made over 20 years after the original Pereira specimen was introduced into Singapore. It is unlikely that the Tambelan plants had survived so long. These garden plants differ from the original collection by Ridley in their more densely pilose petioles and distinctly toothed leaf margin. Irmscher had already raised the possibility that these were pot plants (Topfpflanzen) and that they differed from the Tambelan plant in leaf size (6.5—17 x 8.5—24 cm), margin and acumen length. The Tambelan plant is therefore regarded as a distinct species and is here raised to specific rank. To add confusion, Irmscher also recognised a hybrid between the two subspecies, Burkill 6113 31 Aug 1920 (SING), because he considered it was intermediate between the two subspecies. This specimen is less hairy and has a less conspicuously toothed margin than Burkill 6/15 but it does not resemble B. goegoensis (Burkill 6114) at all. Begonia tambelanense (Irmsch.) Kiew, stat. nov. Basinym: Begonia peninsulae Irmsch. ssp. tambelanense Irmsch. Mitt. Inst. Allg. Bot. Hamburg. 8 (1929) 100. Type: Ridley s.n. 31 Aug 1895 Botanic Gardens Singapore originally collected by Pereira from Tambelan Is. (holo K ex SING’). Reference Irmscher, E. 1929. Die Begoniaceen der Malaiischen Halbinsel. Mitteilung aus dem Institut fur allgemeine Botanik in Hamburg. 8: 86—160. ate Ol eet ai is iy hak attests rote so otal Paotet ts, Niet sSelwenif? hd Sot et wet ob this aile — ost hy vei ears (aS NAEY Barnhyi eve ey ? “| : x ed Pe wr Atl A e ar a , +: - - ss .T a4 » | re i isd tines s tecvaeel oreviogs bbe OF 5s tyaielal , fOr nek TE SALTS Wilke 2k rid Bi ‘2 7a ee Melos rer? ot paanont 1S a! ‘ 4 » wai ~ he | ; ike ° pines o Li : oe | Thee rowed FOALS tit 5 aT: BSN missed her sap. poninsnl 7 7 a ww itin tt = if cto, _ Wear ate weal t A mien) peal 2 . nf suvorniadonat , — eer 1 snl oh ! meg C281 gud [2 12 yA Ss vet O04 (2581) zi sfhTt eit A ed te ; ® ‘ « pe Pt i i as f ok r } « a< Be 4 thus i a e % 1 a 4 : s . | vw : ’ . « Py rus sna, | r are Cited : A Garden’s Bulletin Singapore 55 (2003) 65—68& A New Species and a New Combination in Bornean Kopsia (Apocynaceae: Apocynoideae) DAVID J. MIDDLETON Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, U.S.A. Abstract A new species, Kopsia rajangensis D.J.Middleton, is described and a new combination, Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, is made. Introduction As preparation for an account of the Apocynaceae for the Tree Flora of Sabah and Sarawak, it is necessary to describe a new species of Kopsia and reduce Kopsia mitrephora Sleesen to a variety of Kopsia pauciflora Hook.f. Kopsia rajangensis D.J.Middleton, sp. nov. Frutex 1.2-4.6 m altus, ramis glabris. Folia elliptico-oblonga, 13.4—32 x 3.8—9.6 cm, nervis 9—25 paribus. Corollae tubus 21.5—25 cm, lobis 11-14 x 2.7-4.7 mm. Typus: Borneo, Sarawak, Kapit, Upper Rejang River, Clemens & Clemens 21221 (holo MO; iso A, BM, BO, NY, SAR) Small tree or shrub, 1.2—-4.6 m tall. Branchlets glabrous, not or sparsely lenticellate, terete. Leaves: petiole 4-9 mm long, glabrous; blade papery to subcoriaceous, elliptic or oblong, apex caudate, base obtuse to cuneate, 13.4—32 x 3.8—9.6 cm, 2.64.3 times as long as wide, midrib shallowly sunken or raised and with a central groove above, secondary veins 9—25 with 6—25 mm spacing, 40—60° from midrib, prominent or flat above, prominent beneath, clearly discernable from tertiary venation above and beneath, straight or slightly ascending near margin, tertiary venation prominent above, prominent or flat beneath, reticulate or sub-perpendicular to midrib and oblique to secondary veins, intramarginal vein looped and inset from margin, glabrous above and beneath. Inflorescences dichasial and then with cincinnate branches, 4-15 cm long with axes 14-140 x 1.4—2.1 mm, glabrous to densely puberulent; peduncle 2—76 66 Gard. Bull. Singapore 55 (2003) x 1.7—3.7 mm; pedicel c. 4 mm long; bracts persistent in inflorescence. Sepals ovate, apex rounded, 1.5—1.7 x 1.1—1.4 mm, 1.2—1.4 times as long as wide, ciliate, otherwise glabrous outside and inside. Corolla white; tube 21.5—25 x c. 2.3 mm, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside, 1.8—2 times as long as lobes, 12.6—16.7 times as long as calyx; lobes elliptic or oblong, apex obtuse or acute, 11—14 x 2.7-4.7 mm wide, 34.5 times as long as wide, not ciliate, glabrous outside and inside. Stamens 17-21 mm from corolla base which is 0.79-0.84 of corolla tube length; anthers 1.7—2 x 0.6—0.8 mm, 2.5—2.8 times as long as wide, 1.1—2 mm from corolla throat; filaments c. 0.7 mm long. Ovary 0.9 mm high, glabrous to densely pubescent; style 20 mm long; pistil head 0.8 mm long. Fruits falcate with small blunt projection near the base, 15—16 x 4 x 6.5—7 mm; projection 34 mm long. Distribution: Borneo - SARAWAK: endemic to the Rajang River area. Habitat: In hill forest and disturbed primary forest on rich clay soil. Notes: Merrill (1921) compared a specimen of this species, Haviland 3042, to Kopsia macrophylla Hook.f. but offered no further identification. Timmerman- Van der Sleesen (1959) was the first to note the peculiarity of the material and compared it to Kopsia lapidelecta Sleesen and Kopsia tenuis Leenh. & van Steenis, again without making a recommendation as to what it might be or describing a new species. Anderson (1980) and Ashton (1988) called this species Kopsia larutensis King & Gamble, a species known only from Peninsular Malaysia. Kopsia rajangensis differs from K. larutensis in its larger inflorescences, much larger flowers and relatively wider corolla lobes. Markgraf (1972) cited the specimens under Kopsia arborea Blume. As can be seen, there has been remarkably little agreement on the identification of the species around Kapit on the Rajang River, which the Clemenses spelt ‘Rejang’. Part of the problem has been the lack of good material. This new species is probably related to Kopsia lapidelecta and Kopsia sleeseniana Markgr. It differs from the former in its secondary venation, intramarginal vein and larger flowers, and from the latter in its smaller flowers and also its venation. Specimens examined: Sarawak: Kapit, Upper Rajang River, Clemens & Clemens 21211 (SAR), Clemens & Clemens 21221 (A, BM, BO, MO, NY, SAR); Pelagus Rapids, Ashton $17797 (L); Rajang River, Haviland 3042 (BM, SAR, SING); Bukit Raya, Smith S27738 (SAR). Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, comb. nov. Basionym: Kopsia mitrephora Sleesen, Blumea 10 (1960) 136. Type: G.H.S. Wood SAN 16118. Borneo, Sabah, Lahad Datu, Path between Sungai Sabahan and Sungai Bornean Kopsia 67 Dok (holo L; iso BRI). Notes: Kopsia mitrephora was formally described by Timmerman-Van der Sleesen (1960) following on from her earlier, but not validly published, account of the species (1959). In the earlier work she distinguished K. mitrephora on the basis of its stamens inserted in the lower part of the corolla tube, a character unusual in the genus, and on its delicate leaves. However, with the many more specimens available now, it is clear that the vegetative characters in this species are extremely variable and closely parallel the vegetative variation found in K. pauciflora. These variations include the size of the wings or angles on the stems, the thickness of the leaves, the length of the petiole and the degree of prominence of the venation. Despite long attempts to put some sort of order to this variation, and indeed there are character states concentrated in some geographical areas, no further taxa could be distinguished in either species. In addition, it became increasingly clear that the only character to distinguish K. pauciflora and K. mitrephora was the site of insertion of the stamens and that otherwise both species were remarkably similar, including in fruit characters. Therefore, varietal status is appropriate for Kopsia pauciflora var. mitrephora. Acknowledgments The study was partly carried out under the sponsorship of The Tree Flora of Sabah and Sarawak Project and funded by the Ministry of Science, Technology and Environment of Malaysia through IRPA Funding No. 30—08—01—01-005. The curators of the herbaria at BM, BO, L MO, NY, SAR and SING are thanked for either a loan of material or for their hospitality whilst the material was consulted. References Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forest Dept. Sarawak, Malaysia. Ashton, P.S. 1988. Manual of the Non—dipterocarp Trees of Sarawak. Forest Dept. Sarawak, Malaysia. Markgraf, F. 1972. Florae Malesianae Praecursores LIII. Apocynaceae II. 6. Urnularia, 7. Willughbeia, 8. Kopsia. Blumea. 20: 407-425 Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. Fraser & Neave, Singapore. Timmerman-Van der Sleesen, E.H.L. 1959. Preliminary revision of the Malaysian species of the genus Kopsia (Apocynaceae). Flora Malesiana Miscellaneous Records. 1: 1-15 68 Gard. Bull. Singapore 55 (2003) Timmerman-Van der Sleesen, E.H.L. 1960. In C.G.G.J van Steenis, Miscellaneous Botanical Notes X. Blumea. 10: 136—139 Garden’s Bulletin Singapore 55 (2003) 69-72 New Species and Varieties of Symplocos (Symplocaceae) from Borneo K.G. PEARCE 32, Lorong Kumpang 4, Kuching, Sarawak, Malaysia Abstract A new species of Symplocos, S. buxifolioides K.G.Pearce and two varieties, S. iliaspaiensis Noot. var. pedunculata K.G.Pearce and S. laeteviridis Stapf var. alabensis K.G.Pearce are described from Borneo. Introduction In his account of the Symplocaceae for the Flora Malesiana region, Nooteboom (1975, 1977, 1984, 1986, 1989) recognised 30 species with 19 varieties of the genus Symplocos Jacq. in Borneo. In the process of revising and preparing the manuscript for the Tree Flora of Sabah and Sarawak, one new species, Symplocos buxifolioides, and two varieties, S. iliaspaiensis var. pedunculata and S. laeteviridis var. alabensis are recognised. Descriptions and brief notes on the geographical distribution and morphological characters distinguishing these new taxa from those of previously known are given. 1. Symplocos buxifolioides K.G.Pearce, sp. nov. Symplocos buxifoliae Stapf similis a qua differt ramulis distincte modo gallice et anglice ‘zigzag’ et margine foliae revoluto praeter interdum ad apicem et e basi foliorum non altae obscure denticulatae. Typus: Chew & Corner RSNB 5882, Borneo, Sabah, Mt. Kinabalu, Mesilau (holo SING; iso SAN). Tree to 10 m tall, 10 cm diam. Twigs weakly, irregularly curved, smaller ones distinctly zig—zag with nodes closely set, slender, up to 4 mm diam., dark brown to black; young parts longitudinally ridged and sparsely appressed long—pilose; older parts becoming horizontally cracked, glabrescent, without prominently raised leaf scars. Leaves spirally arranged, coriaceous, glabrous above, hairy to virtually glabrous below, drying olive—brown above, yellow—green below; blade broadly ovate—elliptic, 2.54.5 x 1-3 70 Gard. Bull. Singapore 55 (2003) cm, base shortly attenuate, margin revolute except sometimes at apex, shallowly, obscurely toothed from near base, apex acute, shortly apiculate; midrib narrowly channelled above; /ateral veins (6—)7-& pairs, joining to form an intramarginal vein; intercostal venation reticulate, raised above and prominent below; petiole 3.5—5 mm long, sharply flanged to base. Inflorescences racemous, borne in the axils of upper leaves and below on leafless twigs; axis 2—5(—9) mm long, shortly, moderately dense appressed—pilose; bracts early caducous. Flowers 1-3; pedicels to 0.5 mm long; calyx tube c. 0.3 mm long, lobes broadly ovate, chartaceous, ciliate almost to apex, moderately dense to sparsely appressed—hairy, c. 1 mm long; corolla white, glabrous, 3.5—4 mm long; ovary sparsely to densely appressed hairy, 1.5 mm high. Fruits solitary on a pedicel to 9 mm long, ovoid to curved ellipsoid, 1-1.5 x 0.5—0.7 cm, hardly to shallowly ridged, sparsely appressed—pilose to glabrescent, drying yellowish brown, with persistent and erect calyx lobes. Distribution: Endemic in Borneo. Known only from Mt. Kinabalu, Sabah. Ecology: Upper montane forest at 2400-3660 m altitude. Other specimens examined: SABAH. Mt. Kinabalu: Hotta 3840 (SAN), Chew & Corner RSNB 5911 (SAN), Mikil & Aban SAN 46593 (SAN) and Aban et al. SAN 54254 (SAN). Notes: Symplocos buxifolioides is similar to S. buxifolia Stapf but differs from the latter in its slender twigs, which are distinctly and shortly zig—zag between the closely set nodes; young parts of the twigs sparsely appressed pilose; broadly ovate—elliptic leaves (not obovate—elliptic as in S. buxifolia) with revolute and closely, obscurely denticulate margin from near the base, acute to shortly apiculate apex (not obtuse to rounded), and 6—8 pairs of lateral veins (not 4—6); inflorescences borne on leafless twigs as well as in the axils of upper leaves; calyx tube c. 0.3 mm long, ciliate almost to the apex (not sometimes ciliate at base), and moderately densely to sparsely appressed—hairy (not glabrous); and sparsely to densely appressed—hairy ovary (not glabrous). This species is named for its close morphological similarity to S. buxifolia. 2. Symplocos iliaspaiensis Noot. var. pedunculata K.G.Pearce, var. nov. A var. iliapaiense differt racemis base ramosis ad 6.5 cm longis floribus (fere) sessilibus ad 2 mm pedicellatis. Typus: Fedilis SAN 95642, Borneo, Sabah, Kalabakan district (holo SAN; iso K, KEP, L, SAR, SING). Tree to 13 m tall. Leaves 7.5—20 x 2.5—6 cm, base cuneate, apex with acumen to 1.4 cm long. Racemes basally branched, to 6.5 cm long. Flowers virtually sessile or Symplocos from Borneo al with short pedicel to 2 mm long. Distribution: Endemic in Borneo. Sabah (Kalabakan, Sandakan and Tawau districts) and Brunei Darussalam. Ecology: In primary and logged—over mixed dipterocarp forest at altitudes to 80 m, on flat or undulating lands and ridges. Other specimens examined: SABAH. Sandakan district, Sigin SAN 56798 (KEP, SAR, SING), Leopold & Kodoh SAN 81396 (SAN, SAR), Leopold & Kodoh SAN 81424 (SAN, SAR); Lahad Datu Sinanggul SAN 56991 (SAN, SAR); Kinabatangan Meijer SAN 136314 (SAN), Pensiangan district Goh SAN 141166 (KEP, SAN). BRUNEI: Belait Wong WKM 1079 (SAN). Notes: This new variety may reach 13 m tall (not 3.3 m tall as in the typical variety). It is vegetatively very similar to var. iliaspaiensis but can be distinguished by its basally branched raceme or unbranched spike or fascicle of racemes 1.5 to 6.5 cm long bearing virtually sessile to shortly pedicellate (to 2 mm long) flowers. In contrast, var. iliaspaiensis has fascicles of sessile flowers with the axis entirely absent. This variety is named for its pedunculate inflorescences. 3. Symplocos laeteviridis Stapf var. alabensis K.G.Pearce, var. nov. Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. similis praeter ramunculos graciliores cum paginis infernis foliorum omnino glabros, foliis olivaceis, parvioribus, lanceolatis, ad 6.3 cm longis et 2.2 cm latis. Typus: Madani & Majawat SAN 119252. Borneo, Sabah, Penampang district, Gunung Alab (holo SAN, iso KEP). Glabrous treelet to 2 m tall. Twigs slender, glabrous. Leaves drying olive green; blade glabrous below, lanceolate, to 6.3 x 2.2 cm, base cordate; lateral veins 10 pairs; petiole c. 1 mm long. Distribution: Endemic in Borneo. Sabah, known only from Gunung Alab (the type specimen and Kamaruddin KMS 1431 (SAN, UKMS). Notes: The new variety is similar to Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. except that the twigs and lower leaf surface are entirely glabrous and it has more slender twigs and smaller (to 6.3 x 2.2 cm) lanceolate leaves that dry olive green. It is named for the locality where it was collected. 7? Gard. Bull. Singapore 55 (2003) Acknowledgements The work was carried out under the Tree Flora of Sabah and Sarawak Project and funded by IRPA Research Grant 30—08—01—01—005. The curators of the herbaria at K, KEP, SAN, SAR and SING are thanked for permission to examine specimens in their care as is H.P. Nooteboom for translating the diagnoses into Latin. References Nooteboom, H.P. 1975. Revision of the Symplocaceae of the Old World, New Caledonia excepted. Leiden Botanical Series. 1: 33-335. Nooteboom, H.P. 1977. Symplocaceae. Flora Malesiana. I, 8: 205-274. Nooteboom, H.P. 1984. Symplocos (Symplocaceae) from Bukit Raya. Blumea. 30:73-74. Nooteboom, H.P. 1986. Additions to Bornean Symplocaceae. Blumea. 31: 277-279. Nooteboom, H.P. 1989. Addenda, corrigenda et emendanda. Flora Malesiana. 1,10: 719. Garden’s Bulletin Singapore 55 (2003 ) 73-88 Five New Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia KATHARINE G. PEARCE DANIDA/SWMPI Project, Sarawak Forest Department, Kuching, Malaysia Abstract Five new Begonia species, Begonia kachak K.G.Pearce, B. kasutensis K.G.Pearce, B. niahensis K.G.Pearce, B. stichochaete K.G. Pearce and B. subisensis K.G.Pearce, were collected from limestone habitats in the Niah National Park. A key, descriptions and illustrations are provided Introduction Sarawak’s varied geology includes a number of isolated limestone outcrops dating from a range of geological epochs and varying in size. In north Sarawak the large Melinau formation falls partly within the Gunung Mulu National Park, while in west Sarawak there are a number of limestone outcrops around Bau and Serian and south towards Tebedu. Other exposures are scattered 1n the interior of Sarawak, for example, at Bukit Sarang (Tatau), Ulu Kakus and in the Middle Baram. Between Bintulu and Miri, in the Niah area about 16 km from the coast, there is an isolated limestone massif known as the Subis massif. In 1974, this massif and surrounding forest was constituted as the Niah National Park. The Bau limestone outcrops, being relatively accessible from Kuching, have been a favourite destination for plant collectors since the mid 19" Century and the Mulu limestone was the focus of intensive botanical collection during the Sarawak Forest Department and Royal Geographic Society Expedition in 1978. In contrast, the Subis massif was, until recently, somewhat neglected by plant collectors. G.D. Haviland, Curator of the Sarawak Museum, Kuching, and C. Hose, District Officer in Baram and Sibu, together made the first plant collections there in 1894. In 1932, P.M. Synge, a member of the 1932—-33 Oxford University Expedition to Sarawak, visited Subis, while in 1954, W.M.A. Brooke and Ahmad independently made collections in the area under the Sarawak Museum label. In 1962, B.L. Burtt with P.J.B. Woods and Chew W.L. collected there as did H.P. Fuchs in 1963 and G. 74 Gard. Bull. Singapore 55 (2003) Alphonso and Samsuri Ahmad of the Singapore Botanic Gardens in 1965. The Botany Unit staff, Sarawak Forest Department, made collecting trips to Niah on a number of occasions from the early sixties onwards, including J.A.R. Anderson, who had a particular interest in the limestone flora of Sarawak (Anderson, 1965) and who collected there in 1966 and 1972. Their collections included specimens of the five begonia species described below but which until now remained unnamed. All five species are endemic to the Subis limestone. The highest point of the Subis limestone massif is G. Subis. Bukit Kasut and Gua Pangomah are located on the west of the massif. G. Brangin and G. Bekajang are two discrete hills to the north and northeast of the massif, respectively. The latter is the site of the Niah Great Cave and the Painted Cave, which are important archaeological sites. The botanical field investigations in Niah National Park were carried out as part of the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life Master Plan Implementation through the Improved Management of Totally Protected Areas in Sarawak, Malaysia’ and enabled the five Begonia species to be re-collected from the limestone habitats and for detailed observation to be made. Key to Begonias in the Niah National Park la, Leat- ovate, oblong-ovate of Obovate = oo... 22.6 seaeese cena teen eee ib.) Leafwzeniform:to:sub+orbicular, sejcnic. 4 nell \enlineg beta eee 2a... ~Leaf 1.5 times.or moreastlong:as broad auiiiwe elim lab nehenel cee 2b. Leaf less than 1.5 times as long as broad ...................5..2.....+. 2. B. niahensis 3a. Leaf with lines of stiff hairs between veins ...................... 3. B. stichochaete 3b. Leaf without lines of stiff hairs between veins ...................... 5. B. subisensis 4a. Leaf more than 7 cm long and wide .....d28. Jena! dole edi.od bee 4b. Leaf less than 7 cm long and wide ...............ssccsseeeeeeee 4. B. Kasutensis I. Begonia kachak K.G.Pearce, sp. nov. Section Petermannia A Begonia speluncae Ridl. foliis majoribus ob internodia longiora non in fasciculis dispositis nec peltatis nec glabris differt. Typus: Great Cave, Niah, Miri District /.A.R. Anderson, S. Tan & E.Wright S26074 (holo SAR). Begonias from Niah National Park WS Figure 1. Begonia kachak A Habit, B Detail of leaf margin, C Male bud, D Male flower, E Androecium, F Stamens, G Female flower, H Style and stigma. (from $26074). 76 Gard. Bull. Singapore 55 (2003) Figure 1 Creeping herb or low root climber, roots fibrous, produced at the nodes in contact with the substrate. Stem and petiole with minute appressed to oblique brown hairs; distal end of young petiole, main veins on abaxial surface, margins of young leaf and main vein of stipule with stiff pink-brown trichomes to | mm long. Leafy stem 6-28 cm long, 3 mm diam., little branched, internodes 1—5 cm long. Stipules broadly triangular with a distinct midrib, 4.5 x 7 mm, margin entire, prolonged into an acuminate apex, persistent. Leaves alternate; petiole 1.8 cm in upper leaves, to 17 cm in lower leaves; lamina minutely papillose above and below, upper surface somewhat lustrous, olive- green to yellowish green especially near margins, lower surface pale green or in some specimens brownish pink with pale green veins, oblique, reniform to sub-orbicular, 7.5—12.5 x 7.5—15 cm, base cordate with the basal lobes hardly overlapping, rounded, 0.24.7 cm long, margin irregularly and indistinctly serrate, teeth setose, apex acute to acuminate, venation pinnate-palmate, main veins 4 pairs, branching about half way to the margin, with 2 more veins in the broader and one in the narrower basal lobe, sunken above, prominent beneath. /nflorescences axillary, protogynous, obliquely erect cymose panicles, 8.5—29 cm long, of which peduncle is up to 14 cm long, with one or two female flowers proximally and many male flowers distally; bracts pink, similar to stipules, 6.5 x 3 mm, persistent, bracteoles pale pink-brown, broadly ovate, 4 x 3 mm, apex setose, persistent. Male flowers with pink pedicel to 7.5 mm long, tepals 2, white translucent margins pink before anthesis, minutely papillose, orbicular, 7x 5.5 mm, margin entire, apex rounded, stamens yellow, c. 29, in a spherical cluster, joined in a torus c. 3 mm diam., filaments to 1 mm long, anthers pale yellow, obovate, c. 0.7 mm long, emarginate. Female flowers on pedicels 5.5 mm long, minutely hairy; ovary pink, oblong-obovoid, 10.5 x 7 mm, wings pink, 3, subequal, c. 1.5 mm wide; tepals 5, white translucent or palest pink, with scattered minute, deeper pink hairs, unequal, oblong with rounded apex, to 9.5 x 5.5 mm, margin entire; styles 3, free, each 2 mm to bifurcation from where papillose stigmatic surface forms a continuous twisted band. Fruits with decurved pedicel, 1—1.9 cm long; capsule asymmetric, broadly oblong-elliptic, to 1.9 cm x 1.6 cm, minutely hairy, papillose, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base and apex, two narrower wings to 5 mm wide, broader wing to 7 mm wide, style caducous. Seeds cylindrical, c. 0.3 x 0.2 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis. Distribution: Borneo — SARAWAK: endemic in the Subis limestone (G. Brangin, Niah Great Cave and Gua Pangomah). Habitat: Locally abundant in the shelter of overhangs in somewhat dry areas or Begonias from Niah National Park Vl creeping up rocks at cave mouths, infrequently at level ground in deep shade adjacent to limestone outcrops. Notes: This is a conspicuous begonia with large shiny leaves, commonly bearing a cluster of delicate, pale pink male flowers. It is easily observed on limestone outcrops along the Plank Walk to the Niah Great Cave and around the cave mouth. It takes its name from kachak, the Malay word meaning handsome. Specimens examined: SARAWAK: Gunung Subis, Niah B.L. Burtt & P.J.B. Woods B2010 (SAR), Mohidin S2 1603 (SAR); Great Cave K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude $78536 (SAR), Jemree Sabli S89062 (SAR); G. Brangin, Ulu Sg Subis Yii Puan Ching S40168 (SAR), Gua Pangomah J.A.R. Anderson S31691 (SAR, SING), Near Sg Subis K.G. Pearce $89463 (SAR); Niah Caves Alphonso & Samsuri A217 (SING), A222 (SING), A248 (SING). 2. Begonia niahensis K.G.Pearce, sp. nov. Section Petermannia A Begonia congesta Ridl. habitu erecto vel effuso (non cauli longo), capsula oblongo- ovoidea (non longa oblongaque) differt. Typus: Niah Caves B.L. Burtt & P.J.B. Woods B 2009 (holo SAR). Figure 2 Herb with stems to 75 cm long, branching near base with some spreading horizontally, some vertical and some drooping and then curving upward. Young stems and petioles with minute, erect translucent hairs. Older stems brown and semi-woody, to 1 cm diam., young stems brownish pink, internodes 2—8.5 cm long. Stipules green flushed brownish pink at centre, plicate, broadly ovate, 14 x 7 mm, margin entire with translucent hairs at base, apex acute, persistent. Leaves spirally arranged; petiole from c. 5 cm long in the upper leaves to 22 cm in the lower leaves, deep brownish pink; lamina medium green above, paler below, with veins on both surfaces deep pink at base and at the point where they branch, pinkish between, minutely papillose with a velvety appearance above, translucent hairy beneath, oblique, broadly ovate, 15—20 x 12—16.6 cm, base cordate with basal lobes not or hardly overlapping, rounded, to 9 cm long, margin irregularly and indistinctly serrate, translucent hairy, teeth with translucent setae, apex acute, venation pinnate-palmate, main veins 4—5 pairs, branching near base, with 2 or 3 smaller veins in broader basal lobe and 1 or 2 in narrower basal lobe. /nflorescences axillary, protogynous, obliquely upright, cymose panicles 3—8 cm long, of which the peduncle is 2.5—-6 cm long, with | or 2 pairs of female flowers at the base with many male flowers tightly congested distally, bracts 78 Gard. Bull. Singapore 55 (2003) plicate, pale green, broadly ovate, to 2—2.3 x 1.2 cm, persistent. Male flowers opening one at a time, pedicel obscured by congested bracts and male flower buds, tepals 2, white translucent, margins entire, orbicular, c. 6 x 7 mm, apex rounded; stamens yellow > 30, in hemispherical cluster, joined in a torus c. 1 mm diam., filaments c. 0.5 mm long, anthers yellow, emarginate, 0.8 mm long, opening by apical slits. Female flowers with pedicel c. 9 mm long; ovary yellow-green to pink, obovoid, 19 x 7 mm, wings 3, pale green, subequal, 1.5 mm wide; tepals 5, white translucent, subequal, broadly elliptic-obovate, 13 x 7 mm, margins entire except at fringed, obtuse apex, outer surface and margins minutely translucent hairy; styles 3, c. 1 mm long; stigmas bifurcated, stigmatic surface a minutely papillose, orange yellow spiral band. Fruits with a stiff, decurved pedicel, c. 1.5 cm long, pale green flushed pink at base; capsule oblong, to 3.2 x 1.4 cm, sparsely translucent-hairy, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, subequal, thin, broadest at apex, c. 4 mm wide, style caducous. Seeds cylindrical to obovoid, c. 0.3 x 0.2 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis. Distribution: Borneo - SARAWAK: endemic in the Subis limestone (G. Brangin, Niah Great Cave and Painted Cave areas). Habitat: Below cave mouths, rooted in pockets of soil on ledges and in crevices on steeply sloping limestone rock or in light shade under canopy of small trees. Notes: This robust and handsome begonia has large, asymmetric, velvety hairy leaves that sometimes have a bluish cast. The leaves are positioned more or less vertically against the limestone rock face in somewhat sheltered sites. It can be observed on limestone outcrops along the Plank Walk to the Painted Cave. It has been named for the type location - the Niah Caves. Its habit of drooping and upcurving stems and handsome, velvety leaves held vertically, give this species potential as an ornamental plant. Specimens examined: SARAWAK: G. Brangin Yii Puan Ching $40166 (SAR); Great Cave J.A.R. Anderson, S. Tan & E. Wright S26075 (SAR, SING), K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude §78537 (SAR); Near Sg Subis K.G. Pearce S89460 (SAR). 3. Begonia stichochaete K.G.Pearce, sp. nov. Section Petermannia A Begonia congesta Ridl. foliis angustioribus seriebus setarum inter venas gerentibus, capsulis longioribus oblongis (non oblongo-ovoideis) differt. Typus: Subis, Niah Caves Ahmad 14 (holo SAR; iso SING). Begonias from Niah National Park Figure 2. Begonia niahensis A Habit, B Detail of leaf undersurface, C Detail of leaf margin and upper surface, D Male bud, E Male flower, F Androecium, G Stamens, H Female flower, I Style and stigma from above, J Style and stigma. (A—C from Burtt 2009, D-I from $89460). 79 80 Gard. Bull. Singapore 55 (2003) Figure 3 Erect herb rooting at the base of the stem and at nodes along the horizontal rhizome which creeps below the soil surface and gives rise to shoots at intervals. Indumentum of stiff trichomes to 2.2 mm long, each on a raised conical base, pale brown to pinkish brown to deep brownish red on stem, young petiole and veins on lower and upper lamina surfaces, dense on young stem and petiole, less so on stipule margin and sparse on outer surface of stipule midrib and on the main veins of the upper lamina surface, in a row or rows between each pair of veins and on and between the teeth of the dentate margin, moderately dense on the main veins of the lower leaf surface. Upright stem to c. 42 cm long, 5—9 mm diam., unbranched, internodes 2.5—5 cm long to 8 cm long at base of stem. Stipules oblong-ovate with a distinct midrib, 17 x 11 mm, margin entire, apex setose, caducous. Leaves spirally arranged; petiole 0.7 cm in upper leaves, to 2.5 cm in lower leaves; lamina at first yellowish green then medium green on upper surface (rarely dark blackish-green), lower surface paler yellowish green or olive-green or deep pink or red, asymmetric, irregularly obovate, 11-19 cm long, narrower side to 2.5 cm wide towards apex, curving inwards towards base and hardly or not lobed at base, broader side 6—7.5 cm wide with a cordate base and a rounded basal lobe to 17 mm long, margin irregularly serrate (appearing almost praemorse), each tooth setose, with several setae between each pair of teeth, apex toothed, venation pinnate, c. 4 pairs of veins, branching with another 1-3 veins in basal lobes, impressed above, prominent beneath. /nflorescences axillary, protogynous, erect cymose panicles to 3.4 cm long, of which the peduncle is 2.3 cm, with 1—2 female flowers at base with distally many male flowers crowded in a terminal cluster; bracts similar to stipules, deep red, 13 x 10 mm, persistent; bracteoles pale brown to deep pink, broadly ovate, to 12 x 12 mm, apex setose, persistent. Male flowers witha green to deep pink pedicel with deep pink bristles, 7 mm long; tepals 2 or 4, the outer pair translucent white flushed pink at margin and base, the inner pair translucent white, at first finely hairy on outer surface near base, broadly elliptic, c. 4 x 3 mm, margin entire, apex rounded; stamens pale yellow, c. 32, in an ovoid cluster, joined in a torus c. | mm diam., filaments to | mm long, anthers obovate, to 1.5 mm long, shortly apiculate, opening by terminal pores which develop into slits. Female flowers solitary or in pairs; pedicels 5 mm long, medium pink with minute appressed to slightly oblique hairs; ovary pale pink tinged green with deep pink bristles, oblong-obovoid, to 24 x 16 mm, wings 3, deep pink to translucent white with margins flushed pale pink, subequal, c. 7 mm wide; tepals 5, translucent white flushed pale pink at margins or palest pink, deeper pink at base and on marginal teeth with scattered deeper pink hairs, on outer surface sparsely hispid, oblong with rounded apex, inner two larger, c. 6 x 2-4 mm, margin toothed from near base to fringed apex; styles 3, free to base, 3 mm long, bifurcating; stigmas forming a continuous twisted band, yellowish-green to bright yellow. Fruits with stiff, decurved pedicels, 5—9 mm long; capsule oblong,to 29 x 18 mm, 81 Begonias from Niah National Park es ARES or fad ony RS EDR Sag z ee = a oO o E 5 = ra + < a — o ss (e) : ra Oe & Ege ro] ees oa an =i -e m9 at ay ot = o vO Sa =. = as} cae ss o- Eee Ee gas s ea Soe, a ee alos & Sy was ce = e298 — ‘eres eo & > Fs ara waif. ae eee Es 2 Bm & ma <a 82 Gard. Bull. Singapore 55 (2003) sparsely hispid, minutely papillose, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, subequal, thin, narrowed to base, truncate distally, two narrower wings to 5 mm wide at midpoint and 6 mm at apex, broader wing to 6 mm wide at midpoint to 7 mm at apex, style caducous. Seeds cylindrical-spherical, c. 0.3 x 0.25 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis. Distribution: Borneo —-SARAWAK: endemic in the Niah National Park (Kuala Subis, the Sekaloh area and near the trail to Bukit Kasut). Habitat: This species occurs both in limestone habitats and undisturbed Mixed Dipterocarp Forest on loamy soil or brown clay as well as in secondary forest on alluvial soil. On limestone, it grows on low outcrops with little soil and in soil at the base of limestone cliffs in deep shade and damp conditions, where it is locally frequent. Notes: This erect begonia is characterised by its bristly leaves bearing rows of stiff hairs between the veins. It grows in shady sites, both on or near limestone outcrops and in forest away from the limestone. The species shows striking variation with the upper leaf surface ranging from mid-green to dark blackish-green. It is named for its bristly leaves. Specimens examined: SARAWAK: Niah National Park Bernard Lee S40075 (SAR), Kuala Niah near confluence with Sg Subis K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude S78539 (SAR); Route to Bukit Kasut K.G. Pearce & Narawi b. Johari S75597 (SAR); Sekaloh Jemree Sabli S8S9014 (SAR), K.G. Pearce S589267 (SAR). 4. Begonia kasutensis K.G.Pearce, sp. nov. Section Petermannia A Begonia conipila Irmsch. ex Kiew trichomatibus brevioribus sine basibus conicalibus, foliis suborbicularibus (non valde asymmetricis), punctis maculisque praesentibus, inflorescentiis contra folia insertis differt. Typus: Great Cave, Gunong Subis, Niah, Miri District J.A.R. Anderson S31940 (holo SAR; iso SING). Figure 4 Creeping herb with fibrous roots produced at the nodes. Stem and petiole with white minutely appressed hairs on distal end of petiole; main veins, lamina undersurface and margin with scattered pale orange-brown hairs to 0.7 mm long, more or less regularly arranged. Leafy stem toc. 15 cm long, 1.5 mm diam., not or hardly branched, internodes to 4 cm long. Stipules oblong-ovate with a distinct midrib, to 9 x 4mm, Begonias from Niah National Park 83 Figure 4. Begonia kasutensis A Habit, B Detail of leaf margin, C Leaf showing variegation, D Male bud, E Male flower, F Stamens. (A & B from $31940, C from $27269, D-F from $3/940). 84 Gard. Bull. Singapore 55 (2003) margin entire, apex setose, caducous. Leaves spirally arranged; petiole 1.5 cm in upper leaves, to 3.8 cm in lower leaves; lamina minutely papillose above and beneath, variegated, upper surface slightly lustrous, olive to dark green with irregular spots and blotches of pale grey-green between veins, lower surface reddish, sometimes light green towards base, slightly asymmetric, sub-orbicular, 4.7—6.2 x 3.9-5.5 cm, base cordate with rounded basal lobes not overlapping, to 8 mm long, margin irregularly, indistinctly and distantly serrulate, apex cuspidate, venation palmate, main veins 2 pairs, the two ‘middle’ ones branching near mid-point or towards margin, with 1—2 veins in each basal lobe, sunken above, slightly prominent beneath. Inflorescences \eaf-opposed, protogynous, racemes of cymules, to 11.5 cm long, including peduncle to 5.7 cm, with a single female flower at base and many male flowers distally; bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale green flushed pink, broadly ovate, 3.5 x 2 mm, apex setose, caducous. Male flowers with pink pedicel to | cm long; tepals 2, palest pink translucent, suborbicular, c. 7 x 4 mm, margin entire, apex rounded; stamens yellow, c. 20, ina more or less hemispherical cluster, joined in a torus 0.3 mm diam., filaments to c. 1 mm long, anthers bright yellow, c. 0.5 mm long, opening by pores, obovate, emarginate. Female flower not seen. Fruits with stiff, decurved pedicel, 7 mm long; capsule suborbicular, 1.6 x 1.2 cm, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base, truncate distally, the two narrower wings c. 1.5 mm wide, broader wing c. 2.5 mm wide, style caducous. Seeds cylindrical, c. 0.3 x 0.25 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis. Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great Cave, Bukit Kasut and G. Subis). Habitat: Infrequent, on limestone rock faces in shaded and dry crevices or in soil between limestone rocks on slopes of limestone hills up to where the canopy cover starts to thin out. Notes: This decorative begonia is characterised by its diminutive, sub-orbicular variegated leaves that have a pink undersurface, and by its delicate inflorescences. Its small size and elegant features give it potential as an ornamental species. It has been named for Bukit Kasut, one of the locations where it occurs. Specimens examined: SARAWAK: Subis Ahmad No. 3 (SAR, SING); Niah Cave Ahmad No. 65 (SING); Gunung Subis Jemree Sabli S89049 (SAR); Southern slopes of G. Subis, near Sekaloh River S. Tan & E. Wright § 27269 (SAR, SING); Route to Bukit Kasut K.G. Pearce & Narawi b. Johari S78596 (SAR). Begonias from Niah National Park 85 5. Begonia subisensis K.G.Pearce, sp. nov. Section Petermannia A Begonia pendula O. E. Schulz habitu majore caulibus erectis rigidisque, foliis setis inter dentes setiferos carentibus, fructibus latioribus oblongis differt. Typus: G. Subis ‘in Sekaloh River’, Niah, Miri District /.A.R. Anderson, S. Tan & E. Wright S27574 (holo SAR; iso SING). Figure 5 Cane-like, semi-herbaceous begonia with woody stems ascending, then held at 45°, drooping at apex, arising at intervals from a woody, branched rhizome c. 7-9 mm diam. bearing roots at the nodes, some stem bases bearing adventitious roots. Stem and petiole with minutely appressed brown hairs. Stems to >76 cm long, from 6 mm diam. at base to 1.5 mm at apex, bearing leaves only near apex, unbranched or with few branches, internodes 1.2 to >5 cm long. Stipules lanceolate, without a distinct midrib, 5.5 x 1.5 mm, margin entire, apex setose, caducous. Leaves spirally arranged; petiole 1—2.5 cm; lamina minutely papillose above and below, upper surface with a reflective sheen, medium to olive-green (those of young vegetative shoots variegated, olive-green with irregular spots of pale grey-green between veins), lower surface dull green or pinkish between veins to deep pink near veins, asymmetric, oblong-ovate, to c. 11.3 x 3.64.1 cm, lower half of leaf somewhat inwardly curved, base cordate with a rounded basal lobe to 1.3 cm long on one side, cuneate or slightly lobed at base on the other side, margin irregularly serrate, apex acute, venation pinnate-palmate, veins 3-4 pairs, branching except for the outermost two or three, flat or very slightly raised above, deep pink and prominent beneath. /nflorescences axillary, protogynous, cymose panicles held obliquely above the subtending leaf, axis pale to deep brownish pink, to 7.5 cm long with | or 2 female flowers at the base and up to 10 male flowers distally or sometimes, in the lower leaf axils, by reduction, only the female flowers present; bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale green flushed pink, lanceolate, 1.5 x 6 mm, apex setose, caducous. Male flowers with pink pedicel to 9 mm long; tepals 2, translucent white to palest pink, some flushed deeper pink at base, elliptic, 6.5 x 4mm, margin entire, apex rounded; stamens yellow, to 26, in a more or less obovoid cluster, joined in a 1 mm-diam. torus, filaments c. 1 mm long, anthers yellow, obovate, emarginate. Female flowers on a peduncle to 3 mm long; pedicels deep pink to 2.5 cm long, glabrous; ovary oblong, truncate, 1.6 x 2 cm, wings 3, subequal, c. 6 mm wide; tepals 5, palest pink flushed deeper pink at base, ovate, the outer two smaller than the inner three, c. 9 x 5 mm, margin entire, apex obtuse, glabrous; styles 3, bifurcated, 2-3 mm long to bifurcation, stigmatic surface orange with fine colourless papillae forming a continuous, twisted band. Fruits with flexuous pedicels, to 3 cm long; capsule oblong, 14 x 23 mm, locules 3, equal, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base, truncate distally, two narrower wings 10 mm wide, broader wing 11.5 mm wide, style caducous. 86 Gard. Bull. Singapore 55 (2003) Seeds cylindrical, c. 0.4 x 0.25 mm, surface areolate, areoles more or less equal sided at base and distally, and elongate along seed axis above base. Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great Cave and Bukit Kasut). Habitat: Locally frequent on limestone rock faces and the steep slopes of limestone hills in deep litter layer or between limestone rocks where the canopy cover starts to thin out. Notes: The leaves of this attractive cane begonia have a reflective sheen on the upper surface and are deep pink below. The young plant has variegated leaves. The inflorescence 1s delicate and, in fruit, the capsule dangles on a long, flexuous pedicel. It has been named for the type locality, G. Subis. Specimens examined: SARAWAK: Great Cave J.A.R. Anderson $31939, Bukit Kasut K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude S78538 (SAR). Acknowledgements This paper is based on botanical field investigations at the Niah National Park carried out under the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life Master Plan Implementation through the Improved Management of Totally Protected Areas in Sarawak, Malaysia’ (SWMPI). The author is most grateful to Julian T. Inglis, Chief Technical Advisor of the SWMPI Project, who has taken a lively interest in the collection and description of these begonias; the paper would not have been written without the inspiration and support provided by Dr Ruth Kiew, whose knowledge of and interest in begonias prompted a closer look at the species occurring at Niah National Park and who also checked the types at K and SING; and to Dr M.J.E. Coode for translating the diagnoses into Latin, and to Dr A. Radcliffe-Smth for suggesting ‘stichochaete’ as the appropriate name for that species. The author also thanks the staff of the Botany Unit, Forest Research Centre, and the Niah National Park, National Parks and Wildlife Department, Sarawak Forest Department, for assistance during collecting trips; the curator of SAR herbarium for permission to examine specimens; and Joseph Pau for his excellent botanical illustrations. Begonias from Niah National Park Figure 5. Begonia subisensis A Habit, B Juvenile shoot with variegated leaves, C Fruit. (A & C from $27574, B from S78538). 87 88 Gard. Bull. Singapore 55 (2003) Reference Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Symposium on Ecological Research in the Humid Tropics. Kuching, Sarawak. Sarawak Government and UNESCO. pp. 49-57. Garden’s Bulletin Singapore 55 (2003) 89-95 A New Species of Curcuma L. (Zingiberaceae) from Mizoram, India J. SKORNICKOVA, M. SABU AND M.G. PRASANTHKUMAR Department of Botany, University of Calicut, 673 635 Kerala, India. Abstract Curcuma rubrobracteata Skornickova, M.Sabu & Prasanthkumar sp. nov. is described from Mizoram, India. Introduction Exploration for gingers in Mizoram, NE India, carried out by the authors for the research project ‘Revision of Indian Zingiberaceae’ has brought an interesting species of Curcuma to light. The genus Curcuma L. naturally occurs in tropical and sub-tropical Asia with a few species extending to Australia and the South Pacific, but economically important or ornamental species are introduced elsewhere in the tropics. According to the latest records, Curcuma has the largest number of species in India, Thailand, Indonesia and China. The total number of species for this genus is still quite uncertain but is estimated at around 100 species (Sirirugsa, 1996; Larsen et al., 1998). From India, 29 species have so far been reported and accepted (Karthikeyan et al., 1989; Mangaly and Sabu, 1993). Several new species of Curcuma L. have been described in last 20 years from India (Skornickova and Sabu, 2002). While working on Indian Zingiberaceae in August and September 2002, the authors visited Mizoram, which is considered as the least surveyed corner of India, where an undescribed species of the genus Curcuma L. subg. Curcuma (Eucurcuma K. Schum.) was collected. Curcuma rubrobracteata Skornickova, M. Sabu & Prasanthkumar, sp. nov. Curcumae roscoeanae similis in coma inconspicua, spica c. 10 cm supra terram per vaginam protrudenti, bracteis floriferibus rubris, floribus luteo-aurantiacis, calyce trilobato, rhizomate repenti aromatico, lamina elliptico-lanceolata basi cuneata, infra glabra, supra pubescenti in venis principalibus elevatis differt. Typus: India, Mizoram, Lawngtlai District, on the way to Ngengpui Wildlife Sanctuary — Khomoi, 22° 30'N, 90 Gard. Bull. Singapore 55 (2003) 92° 46'E. 10.1X.2002. Skornickova & Prasanthkumar 86241 (holo MH; iso CALI, K, SING). Figure 1, Plate 1. Rhizomatous herb, up to 1.5 m high. Rhizome creeping, slender, 10—30 cm long, c. 1 cm diam., tan outside, scales triangular, papery, light brown, underground quickly decaying, whitish yellow inside, aromatic, smell resembling Kaempferia galanga L., taste very bitter, sessile tubers absent, root tubers 2 x 1.5 cm, white inside, distanced c. 5 cm away from main rhizome on 2 mm thick roots. Leafy shoot up to 1.5 m long, leaves 4—6, pseudostem green, c. 40 cm long. Leaves petiolate, ligule 1 mm long, light green, translucent, petiole green, glabrous 10-45 cm long (lower leaves with shorter petioles). Lamina elliptic-lanceolate 35—60 x 10-16 cm, adaxially green, pubescent along the raised veins, abaxially pale green, glabrous, base attenuate, tip acuminate 2 cm long, midrib green, glabrous. /nflorescence terminal, but protruding through the base of the pseudostem through lateral slits c. 3-10 cm above ground. Peduncle 5—10 cm long, 7 mm diam., whitish, without vegetative bracts. Spike 10 x 7-9 cm. Coma absent. Bracts 20-26 per spike, all fertile, bright red or light red, yellowish or yellowish green towards the base, bract 3.5 x 3.5 cm, glabrous, margin 0.3 mm hairy, hairs 0.1 mm long, subtending a cincinnus of 5—6 flowers, usually only 1—3 flowers per spike open at the same time. Bracteoles one per flower, 2—3.5 x 1-3 cm, hyaline, glabrous, white, translucent with reddish dots on the apical part of the biggest bracteoles. Flowers 6 cm long, yellow-orange, exserted, 1.5—2 cm longer than the bracts. Calyx 1.2 cm, white, translucent, hyaline, glabrous, 3-toothed. Corolla tube 3.74 cm, light orange, glabrous. Corolla lobes light yellow-orange, dorsal lobe 14 x 8 mm fringed by red on upper mucronate portion; lateral lobes 12 x 7 mm. Labellum 15 x 17 mm, periphery yellow-orange, centre deep yellow-orange, obscurely 3-lobed, middle lobe split about 2 mm. Lateral staminodes | x 1.1 cm; yellow-orange, hooded over the anther. Stamen c. 9 mm long, anther versatile. Anther 6 x 2.5 mm, orange. Anther spurs 3 mm long, incurved, orange. Anther thecae white, 5 mm long. Filament yellow-orange, 5 mm long, constricted, 4 mm at base, 2 mm at upper part. Ovary trilocular, 2 x 2.5 mm, pubescent with 0.6 mm long hairs; ovules many. Stigma white, closely appressed within the anther lobes. Epigynous glands 2, pale orange, 4 mm long, 0.9 mm diam. Fruits not seen. Flowering: August — September. Distribution: Hitherto known only from the type locality. From personal communication with other specialists we suspect this species to be identical with unidentified plants growing along the Thai-Burmese border. 91 Curcuma rubrobracteata 1 oe ee mee es ee -— i Figure 1. Curcuma rubrobracteata A. Habit; B. Inflorescence; C. Detail of leaf venation (adaxial side); D. Flower and subtending bract; E. Bracteole; F. Part of flower showing lateral staminodes, dorsal corolla lobe and anther; G. Part of flower showing labellum and anterior corolla lobes; H. Flower (side view); I. Calyx; J. Lateral staminode; K. Anther (front); L. Anther (side); M. Stigma; N. Epigynous glands and ovary; O. Ovary (cross section). Based on the type material Skornickova & Prasanthkumar 86241. del. J. Skornickova. 92 Gard. Bull. Singapore 55 (2003) Habitat: Growing in undergrowth in teak plantations and along roadsides. Etymology: This gorgeous species takes its name from the striking red bracts of its inflorescence. Other specimens examined: India, Mizoram, Lawngtlai District, on the way to Ngengpui Sanctuary — Khomoi, 22° 30' N 92° 46' E, 10.[X.2002, Skornickova & Prasanthkumar 86239 (CALI). Notes: The most obvious characters of this species are the creeping rhizomes, bright red bracts with no distinct coma and, in particular, the position of its inflorescence. Inflorescences with no distinct coma are reported for a few species e.g. Curcuma roscoeana Wall., C. albiflora Thwaites, C. ceratotheca K.Schum. and the recently described C. rhomba J. Mood & K. Larsen, which also possesses a small rhizome with linear growth (Mood and Larsen, 2001). The unique character of C. rubrobracteata is the position of its inflorescence, which is so far unknown in the genus Curcuma. Many of the earlier taxonomists made attempts to classify the genus into sections based on the position of the inflorescences. Roxburgh (1820) recognised two sections based on the lateral or central inflorescence, while Horaninow (1862) distinguished three sections, namely: I. Exanha (always lateral), I. Mesantha (inflorescence invariably terminal), and III. Amphiantha (inflorescences both terminal and lateral). However, the new species produces an inflorescence about 3—10 cm above ground, which breaks through the pseudostem through lateral slits. This feature has also been observed in the Zingiberaceae in the genus Plagiostachys Ridl., where the inflorescence is borne on a peduncle and projects from the side of the leafy stalk (Ridley, 1899). Smith (1985) pointed out that the inflorescence of Plagiostachys species, although pushed out laterally, is actually terminal on the leaf shoot and so shows affinity to the genus Alpinia Roxb., in which a few species, particularly A. hansenit R.M.Smith and A. havilandii K.Schum., have similar apparently laterally produced inflorescences. Recently, Kress et al. (2002) carried out molecular studies on phylogeny and classification of the family Zingiberaceae. The results based on analysis of DNA sequences of the nuclear internal transcribed spacer (ITS) and plastid matK regions suggest that the genus Curcuma, as it is accepted nowadays, is paraphyletic with Plate 1. Curcuma rubrobracteata 1. Detail of flower and spike — seen from above; 2. Detail of flower and dorsal corolla lobe — front view; 3. Detail of flower and bracts — lateral view; 4. Habit; 5. Detail of spike protruding through the pseudostem; 6. Creeping rhizome and root tubers. All photographs are of the type material Skornickova & Prasanthkumar 86241. Photo J. Skornickova. Curcuma rubrobracteata mh 94 Gard. Bull. Singapore 55 (2003) genera Hitchenia, Stahlianthus and Smithiatris, which also share cone-like inflorescences of few flowered, congested bracts, and that genus Curcuma is itself paraphyletic with three groups of species. Since the analysis was done with a limited number of samples (six Curcuma species), the authors recommended that more species of such large genus should be included in future molecular studies to shed more light on the difficult question of generic boundaries and allied genera. Acknowledgements The authors thank the Department of Science and Technology, Govt. of India for financial support (Order No. SP/SO/A-20/99 dt. 09.11.2001). We are also indebted to staff of Forest Department of Mizoram for their hospitality and help to reach otherwise inaccessible areas. The senior author thanks the Indian Council for Cultural Relations New Delhi, India, and the Ministry of Education of the Czech Republic for awarding a research fellowship, and to Singapore Botanic Gardens for providing facilities during her repeated visits. We also thank Dr. J. F. Veldkamp, National Herbarium of Netherlands, Leiden University Branch, and Dr. M. Svrcek, National Museum, Prague, for the help with the Latin diagnosis. References Horaninow. 1862. Prodromus Monographiae Scitaminearum. Petropoli (St. Petersburg), Russia. Karthikeyan, S., S.K. Jain, M.P. Nayar and M. Sanjappa, 1989. Florae Indicae. Enumeratio Monocotyledoneae. Botanical Survey of India, Calcutta. Pp. 289-299. Kress, J.W., L.M. Prince and K.J.Williams. 2002. The phylogeny and a new classification of the gingers (Zingiberaceae): evidence from molecular data. American Journal Botany. 89: 1682-1696. Larsen, K., J.M.Lock, H. Maas and P.J.M. Maas. 1998. Zingiberaceae. In: K. Kubitzk1. (Ed.) The Families and Genera of Vascular Plants. 4: 474-495. Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian species of Curcuma L. (Zingiberaceae). Rheedea. 3: 139-171. Mood, J. and K. Larsen. 2001. New Curcumas from South East Asia. The New Plantsman. 8: 207—217. Curcuma rubrobracteata 95 Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal Straits Branch Royal Asiatic Society. 32: 85-184. Roxburgh, W. 1820. Monandria monogynia. Flora Indica. Serampore, India. Pp. 1-84. Sirirugsa, P. 1996. The genus Curcuma of Thailand. In: T. L. Wu, Q. G. Wu and Z. Y. Chen (Eds.) Proceedings 2" Symposium on Family Zingiberaceae. Zhongshan University Press, China. Pp. 39-46. Skornickova, J. and M. Sabu, 2002. The genus Curcuma L. in India: Resumé and Future Prospects. In: A.P. Das (EIndiad.) Perspectives of Biology. Bishen Singh, Mahendra Pal Singh, Dehradun, India Pp. 45-51. Smith, R.M. 1985. A review of Bornean Zingiberaceae: 1 ( Alpinieae p.p.). Notes Royal Botanic Garden Edinburgh. 42: 261-314. Bee tll poenelitee a = " - Seytth the ot kro me ik ik. S mal chat erat ¥ oe pes ’ ry ° P| *» ’ ef lren i ~~ a ¢ te . at, ¢ ’ rt eek ees et Dal bs dime ide Lis features ore? jemi le ss © Anne Hey lecvgnrey © | Ay eey i. fi ‘Ai WA SiG “Bie Lime be ¢ i ini } mk psoas RPP ies ‘ . pam prereset oor bala no » Bogert a Garden’s Bulletin Singapore 55 (2003) 97-111 The Genetic Relations of Musa Species from Mount Jaya, New Guinea, and a Reappraisal of the Sections of Musa (Musaceae) CAROL WONG!', GEORGE ARGENT?’, RUTH KIEW’, OHN SET? AND YIK YUEN GAN! 'National Sciences Academic Group, National Institute of Education, Nanyang Technological University, 1 Nanyang Walk, 637616 Singapore "Royal Botanic Garden, Edinburgh, EH3 5LR, U.K. 3Singapore Botanic Gardens, | Cluny Road, 259569 Singapore Abstract Molecular analysis using amplified fragment length polymorphism (AFLP) confirms the status of Musa banksii F. Muell. as a subspecies of M. acuminata Colla and shows M. johnsii Argent to be genetically more similar to the Australian M. jackeyi W. Hill than to M. lolodensis Cheesman from New Guinea. In addition, AFLP analysis supports only two sections as genetically distinct, namely Sect. Musa (including Sect. Rhodochlamys) and Sect. Callimusa (including Sect. Australimusa). No material for Sect. [gentimusa was available for study. However, for practical purposes of grouping banana species, four informal groups are recognised — the ‘acuminata’ and ‘ornata’ groups within Sect. Musa, and the ‘coccinea’ and ‘textilis’ groups within sect. Callimusa. A key to the sections and informal groups is provided. Introduction Wild bananas are found throughout Asia and Malesia extending into Australia and the Pacific. They have become an increasingly conspicuous element of the vegetation as they invade forest margins along logging roads and openings in forest. As roads penetrate deeper into forests, new banana species are coming to light. Much still remains to be discovered about the relationships between wild banana species, which have been grouped into five sections (Cheesman, 1947; Argent, 1976) based on chromosome number and morphology. However, some of the species recently described do not fit comfortably into these sections (Argent, 2000, 2001), which calls into question their taxonomic validity. Also, molecular studies indicate that not all the sections are genetically uniform and distinct from each other (Jarret & 98 Gard. Bull. Singapore 55 (2003) Gawel, 1995; Wong et al., 2002). Bananas present a challenge to the collector who wishes to turn them into herbarium specimens, which unless accompanied by spirit material, detailed field notes and colour pictures, are often almost worthless. Molecular techniques are therefore particularly useful in this genus and have been used to study genetic diversity at the subspecific (Wong et al., 2001a), specific (Wong et al., 2001b) and sectional levels (Jarret & Gawel, 1995, Wong et al., 2002). AFLP is a DNA fingerprinting technique, which was developed by Vos et al., (1995). It is based on selective PCR amplification of DNA restriction fragments under stringent conditions. It can be used for DNA of any origin and complexity and is reported to be both reproducible and reliable (Vos et al., 1995). AFLP combines the reliability of RFLP with the power of PRC. In this study, AFLP (amplified fragment length polymorphism) was used to analyse the genetic relations of two species collected from Mt Jaya, Papua (formerly Irian Jaya), New Guinea: (a) to study the genetic relationship of Musa johnsii Argent, which is unique in the genus in having a fruit with a sterile mucilaginous pith chamber that occupies the distal third of the fruit (Argent, 2001); and (b) to reassess the status of M. banksii F. Muell., which was reduced to a subspecies of M. acuminata Colla by Simmonds (1956) but is still considered to be a distinct species by Argent (1976). In addition, the taxonomic status of the five sections of Musa is reviewed in the light of molecular studies. Materials and Methods Plant Materials Twelve samples were used for the present study (Table 1). These included Musa species from sections Musa, Rhodochlamys, Callimusa and Australimusa and a species of the related genus Ensete as an outgroup taxon. Voucher specimens were deposited in the herbaria of the Singapore Botanic Gardens and the Royal Botanic Garden Edinburgh. DNA Extraction Leaf tissue was used for AFLP analysis and prepared using a procedure from Zhang et al. (1997). Plant DNA was extracted using the CTAB method according to Reichardt and Rogers (1993) as outlined in Wong et al. (2001b). AFLP analysis The AFLP analysis was carried out according to Vos et al. (1995) with minor modifications. Restriction digests of genomic DNA with EcoRI and MselI were carried out at 37 °C for | h. Following heat inactivation of the restriction endonucleases, the genomic DNA fragments were ligated to EcoRI and MselI adapters overnight at 16°C 99 Musa in New Guinea & Section of Musa yeqes ‘nesurjayy ny wn UOISs9NUO_D suIUT] Wodsa1g = QW vISARIY ‘IIH S,Joseiy Ha vIskeleyy ‘Spue[ysiy Uos1owes; HO eIskeyeyy ‘yeqeg “wouay ‘yeg [eInynousy = dV a1odesuls ‘suopiey s1uvjog o10desurs as ysinquipy ‘uopiey oruejog [ekoy roxy | —-- qe eee Oads 81706661 THM ayant dV LTV SON SI]NXAI OWA €1900 UBUUISIOY SISUaPO]O] “W DSNUIJDAJSNY “JOOS DSN HA OL8b Ma [PI Suaospjoia “py ds 8h776661 ‘D9q SISUIAULOG " DSNU1]]DZ) JOOS DSN Ogu US YD ‘{ YOoH vauinsups ‘pw SKUUDIYIOPOY Y “JIS DSN OWA ‘us VO TNA “ Usyupg “Ww WO 919bMy pdinr019nu ‘dss e][[oD vipuNUNID * jh Ody Us YH pauls ‘dss e][OD vIpUIWINID “pw HD QTLr aa DIDIUNA “ASS BI[OD DIDUIUNID "Pw DSNJP “9S DSN JIA 79S00 juosry lsuyol pw OdS VLv8-96-6 UBUISIOYD (‘qxOY) wngiadns ajasuq —_————————————— — — eee soINOS “ON UOISSIDDY UuOXxeL OE ee ee, eee 2 ee eee Se eee ee ee ee Ee ee poipnys soroads ajasuq pue vsnpy T IQR], 100 Gard. Bull. Singapore 55 (2003) to generate template DNA for amplification. PCR was performed in two consecutive reactions. The template DNA generated was first pre-amplified using AFLP primers each having one selective nucleotide. The PCR products of the pre-amplification reaction were then used as template after dilution 5-fold in sterile water, for selective amplification using two AFLP primers, each containing three selective nucleotides. A total of eight primer combinations were used in this study (Table 2). The final PCR products were run on a 6% denaturing polyacrylamide gel in 1X TBE buffer. The EcoRI primers used were not radioactively labelled as in the original protocol. Instead, a modified silver staining method was used (Loh et al., 1999). Table 2 The sequence of primers used in the AFLP analysis. Name / Abbreviation GYY 101/ EA+ GYY 102/ EA- GYY 103/ MA+ GYY 104/ MA- GYY 105/E-A GYY 107/ E-AAC GYY 108/ E-AAG GYY 109/ E-ACA GYY 110/ E-ACT GYY 111/ E-ACC GYY 112/ E-ACG GYY 113/ E-AGC GYY 114/ E-AGG GYY 106/ M-C GYY 115/ M-CAA GYY 116/ M-CAC GYY 117/ M-CAG GYY 118/ M-CAT GYY 119/ M-CTA GYY 120/ M-CTC GYY 121/M-CTG GYY 122/ M-CTT Enzyme EcoRI EcoRI Msel Msel EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI Msel Msel Msel Msel Msel Msel Msel Msel Msel Type Adapter + Adapter - Adapter + Adapter - Primer +1 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +1 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Sequence (5’-3’) CTCGTAGACTGCGTACC AATTGGTACGCAGTCTAC GACGATGAGTCCTGAG TACTCAGGACTCAT GACTGCGTACCAATTCA GACTGCGTACCAATTCAAC GACTGCGTACCAATTCAAG GACTGCGTACCAATTCACA GACTGCGTACCAATTCACT GACTGCGTACCAATTCACC GACTGCGTACCAATTCACG GACTGCGTACCAATTCAGC GACTGCGTACCAATTCAGG GATGAGTCCTGAGTAAC GATGAGTCCTGAGTAACAA GATGAGTCCTGAGTAACAC GATGAGTCCTGAGTAACAG GATGAGTCCTGAGTAACAT GATGAGTCCTGAGTAACTA GATGAGTCCTGAGTAACTC GATGAGTCCTGAGTAACTG GATGAGTCCTGAGTAACTT Musa in New Guinea & Section of Musa 101 Data analysis For the diversity analysis, bands were scored as present (1) or absent (0) to form a raw data matrix. A square symmetric matrix of similarity was then obtained using Jaccard’s Similarity Coefficient Coefficient [x/(y-z)], where x is the number of fragments in common between two taxa, y is the total number of fragments scored, z is the number of fragments absent in both taxa, from the raw data matrix. Genetic diversity estimates (GDEs) were then calculated as 1 minus Jaccard’s Similarity Coefficient and used for cluster analysis using the Unweighted Pair Group Method with Arithmetic mean (UPGMA) technique of the NEIGHBOR program in PHYLIP version 3.5c (Felsenstein, 1993). The dendrogram was drawn using TREEVIEW version 1.6.1 (Page, 1996). Results AFLP Profiles The AFLP profiles generated using eight primer combinations contained highly informative bands, which distinguished all taxa examined and provided valuable information on genetic relationships. Figure | illustrates an AFLP profile obtained. Only unambiguous bands of size 50-500 base pairs were scored. A total of 201 bands were analysed out of which (95%) were polymorphic across all samples examined. On average, 27 bands were scored per primer pair. Fifty-nine unique bands were observed for the taxa examined (Table 3). All species except for Musa textilis were characterised by unique markers ranging from 1 to 18. M. johnsii is characterised by six unique bands, confirming it is a distinct species. M. banksii is characterised by four unique bands. The genetic markers observed will be useful for the development of probes in Musa breeding programmes. The genetic diversity estimates are shown in Table 4. Discussion Position of Musa johnsii This species has strikingly distinct morphological characters, such as the compact, subglobose head of schizocarpic fruits that are unique in the distal third being filled with pale pinkish orange, mucilaginous pith (Argent, 2001). AFLP analysis confirms it is a distinct species (Fig. 2). Argent (2001) suggested it was closely related to the schizocarpic M. lolodensis as the seeds of the two species are remarkably similar but Table 4 and Fig. 2 show it is genetically more similar to the Australian M. jackeyi than 102 Gard. Bull. Singapore 55 (2003) it is to the New Guinea M. lolodensis. All three species belong in Sect. Callimusa in the ‘textilis’ group (see below). This is another example where conspicuous morphological differences do not reflect genetic similarity (Jarret & Gawel, 1995). Status of Musa banksii Simmonds (1956) reduced this species to a subspecies of Musa acuminata based on experiment and field observations. However, Argent (1976) maintained it as a distinct species as it did not hybridise with M. acuminata subsp. malaccensis when they were grown together for many years in the Lae Botanic Garden, Papua New Guinea. This taxon also differs from the other M. acuminata subspecies in its female flowers having some fertile stemens, in the non-imbricating bracts in the male bud, and in producing a very large number of seeds (up to 400) as compared with 40-50 in subspecies from the lowlands of Peninsular Malaysia and Thailand (Simmonds, 1995). Results of several studies using molecular techniques are now available. Gawel & Jarret (1991) used cpDNA RFLP, which generated a phenogram with M. banksii embedded among the other subspecies of M. acuminata studied, indicating that it is not genetically distinct at the species level. Jarret & Gawel (1995) using total-DNA RFLP again found that M. banksii clustered with the other M. acuminata subspecies in their phenogram. Our study using AFLP showed M. banksii clustering with the other subspecies of M. acuminata (Fig. 2), again indicating that genetically it is not a distinct species and that subspecific rank would be more appropriate. However, the genetic diversity estimates (Table 4) indicate it is the least similar among the four subspecies studied, perhaps a reflection of a combination of its unique characters listed above and its most south-easterly distribution of all M. acuminata subspecies. Sections and groupings of wild banana species Cheesman (1947) created four sections within the genus Musa - sections Eumusa (now Musa), Rhodochlamys, Callimusa and Australimusa - as a convenient way of grouping the species. He noted that the sections were not of equal rank and that in some characters sect. Australimusa was intermediate between sect. Callimusa and sect. Musa, and that the division between sect. Musa and sect. Rhodochlamys was ‘unessential’. He speculated that sect. Musa and sect. Australimusa were the earliest to diverge and that sect. Rhodochlamys was an offshoot from sect. Musa. He was in two minds as to whether sect. Callimusa had diverged directly from sect. Australimusa or, as he considered the barrel-shaped seeds with a large oil/air space a very significant character, might be an earlier divergence from sect. Musa. Argent (1976a) described a fifth section, /gentimusa, with the single species M. ingens N.W. Simmonds. Subsequent to Cheesman’s work, new species have been described that do not conform to his concepts of the sections so that the distinction between the sections is becoming blurred. Molecular techniques have opened a new avenue of enquiry. Studies carried out so far (Jarret & Gawel, 1995; Wong et al., 2002, and this study) Musa in New Guinea & Section of Musa 103 Miz3 45 67 8°910l1 12 at =e eo Figure 1. AFLP profile generated using primer pair 1 (E-AAC, M-CAA). Lane M: pUC19/Hpall molecular weight marker, lane 1: Musa johnsii, lane 2: Ensete superbum, lane 3: M. lolodensis, lane 4: M. textilis, lane 5: M. jackeyi, lane 6: M. borneensis, lane 7: M. violascens, lane 8: M. banksii, lane 9: M. acuminata ssp. siamea, lane 10: M. acuminata ssp. truncata, lane 11: M. acuminata ssp. microcarpa, lane 12: M. sanguinea. Gard. Bull. Singapore 55 (2003) 104 PoweoIpur oie pUd ,¢ 94} JB poppe sapNosponu saNdgjos oy) ‘sod poseq Ja\depe-jasy: aspx PoyOIpuU oie pus ,¢ dy) Je poppe sopNosponu sandoyos oy) ‘sod poseq Jaidepe-[Yyooy : [YooTs. 6S ¢ I rd Cl v C ¢ t 0 I 8I 9 [BIOL t ; ; . : I ¥ c : . C : onl NB DOV v9 ¢ 2 7 . r 7 7 . 4 : "i G I OL) JOV GS 14 I P : ‘ I = I 4 : . I : LO LOV OV (4 ; i . 2 I 3 ‘ 7 ‘ 7 3 I ViLO DOV Le a I ¥ F 5 , 3 ? Fi “ : : C LY JIV 87 I ; . a 7 : I - : ; DVO VOV 61 I : ‘ E i ; : - ; ; I s : JVI OVV Ol OV € I c cl I C ¢ I . 5 Cl C *aVVO *JVV I aed Gee fe es ee ee eK eS Se SS CS sound 319d @ = = = & = 8 sce tn EY Ss a ee So = a § g § aS = = Poe ie 2 at oo S = SIOYIVUL a Gere “ES. + of Gee ee 2 eS ee ce anbrun jo .aquinu ae tone = = = of g a> * = S i [PIO], S S Ss Sy = =. ae Sd. a a y = S S S = [SSI [qooH Id OWL “SUOTIRUTQUIOD JOUWULId JYsto Suisn sisAjeue qT4{V uodn pojsa}ap UoXe} Yovo JO} OIpIOods spuvg JoyIeUW FeyNOsTOU oNbiun jo 1quINN *¢ IIqeL, 105 Musa in New Guinea & Section of Musa - DAUINSUDS "PW OLE'O - pdaps04ou “dss ‘nov "pw 6Lv'0 YIt'0 - pypouna) “dss “nov * py 8Svp0 LOVO 9PV 0 : paupis ‘dss ‘nop ‘Wy 887p0 6f70 8890 6S 0 . IISYUD *W Boo 64L0°-Se90 PLI0 ZO0E0 SUIISDIOIA "GW 969'0 LS9O0 8190 Iv90 6890 LPSO0 ‘ SISUJOUAOG “I cy90 St90 £090 v6SO 8190 IPSO S6t0 > 1dayoo! 88S0 9670 S90 c8SO OLSO YXLSO L6VO IvS0 - S1]1XA] “WW 9ILO 8020 8490 6990 O080 £990 £890 eSSO £2790 , SISUAPO]O] "W V6LO c9LO IVLO SLLO 17280 S6L0 tvLO Y99LO Y99L0 S880 js wingsadns * 5 CILO celO Sv9DO 9L90 8SL0 6090 6L7V0 6970 9790 Sc90 £6L0 ; suyol “py x S < < < < < S * < S 8 4 5 J 8 a 3 ES 8 a = > NID "Pl O} PaywelAdiqgge SI DIDUIUNID DSN “WatdJJooD AyUepWUIS s_presoer (SNUTPA[) - [| Usados SqdD ‘suoneurquioos sound g Suisn exe} ZT] UddMIOq (SYN) sawuyss AyIsIOAIP ONOUDS osIMJIVd poseq-q TV IY) JO ULI] “p IIqUL 106 Gard. Bull. Singapore 55 (2003) Ensete superbum Musa lolodensis Musa violascens Musa textilis Musa johnsii Musa jackeyi Musa borneensis Musa acuminata ssp. banksii Musa acuminata ssp. truncata Musa acuminata ssp. siamea Musa acuminata ssp. microcarpa Musa sanguinea 0.1 Figure 2. UPGMA cluster analysis of AFLP data generated by eight primer combinations for 12 taxa of Musa depicting patterns of genetic diversity. Scale depicts genetic diversity estimates (GDEs). Musa in New Guinea & Section of Musa 107 strongly support two main groups within the genus Musa, which correlate with chromosome number, i.e. those with a somatic number of x=11 (sect. Musa and sect. Rhodochlamys) and x=10 (sect. Callimusa and sect. Australimusa, including M. beccarii with a somatic number of n=9). On the other hand, they also show that sect. Rhodochlamys is not genetically distinct from sect. Musa, and that sect. Australimusa is not distinct from sect. Callimusa. No material of Musa ingens (sect. [gentimusa, somatic no. x=7) has been available for molecular analysis. The conspicuous distinction between sect. Musa and sect. Rhodochlamys is the tall pseudostem and pendulous inflorescences with dull coloured bracts, which mostly become revolute on fading, of the former, compared with the short pseudostems and erect inflorescences and brightly coloured bracts that do not become revolute of the latter. Apart from pseudostem height, these characters are related to the pollination syndrome: the flowers on pendulous inflorescences being pollinated by nectiferous bats and the erect ones by nectiferous birds (sunbirds and spiderhunters). Molecular studies show that many species classed within sect. Rhodochlamys are in fact more closely related to species within sect. Musa than they are to other species in sect. Rhodochlamys, which indicates that bird-pollination in bananas has evolved independently several times. Simmonds (1962) considered bat pollination ancestral to bird pollination in the genus Musa. Both Simmonds (1962) and Shepherd (1999) suggested that these two sections be amalgamated. Molecular studies (Jarret & Gawel, 1995; Wong et al., 2002 and this study) strongly support this view. Cheesman (1947) was most impressed with the distinct barrel-shaped seeds with the large oil/air space of sect. Callimusa, so much so that he thought it would have warranted generic status had there not been species in sect. Australimusa that shared characters of both sect. Musa and sect. Callimusa. However, as more species have been discovered, seed morphology has been shown to be much more variable (Argent, 2000). In general, sect. Ca/limusa includes species with erect inflorescences with colourful bracts compared with most species in sect. Australimusa with pendent inflorescences (although there are exceptions in both sections), again indicating that the erect inflorescence with bird-pollinated flowers has evolved several times. Molecular studies (Wong et a/., 2002; and this study) also illustrate that there is no genetic justification for keeping them separate as species in sect. Callimusa cluster among species of sect. Australimusa (Fig. 2). To put banana taxonomy on a firmer basis where the sections reflect genetic similarity among the species, three sections are recognised here: Section Musa (Baker) Cheesman, Kew Bull. (1948) 108. Basinym: subgen. Eumusa Baker, Ann. Bot, 7 (1893) 205, 208. Type: M. paradisiaca L. Syn: Sect. Rhodochlamys Baker, Ann. Bot. 7 (1893) 205. Type: M. ornata Roxb. 108 Gard. Bull. Singapore 55 (2003) Section Callimusa Cheesman, Kew Bull (1947) 112. Type: M. coccinea Andrews Syn. Sect. Australimusa Cheesman, Kew Bull. (1947) 110. Type: M. textilis Nee Section Ingentimusa Argent, Notes Roy. Bot. Gard. Edinb. 35 (1976) 111. Type: M. ingens N.W. Simmonds. However, Cheesman’s point that his four sections are a convenient way of grouping bananas still holds as the bird-pollinated species with colourful bracts in sect. Rhodochlamys and sect. Callimusa are important horticulturally, while the edible bananas fall into the other two groups, sect. Musa with the M. acuminata x M. balbisana hybrids and sect. Aastralimusa, which includes the Fe’i cultivar group. In addition, as Cheesman remarked, some groups are easily identified in the field. For example, most species in the ‘coccinea’ group are readily recognised by the pronounced scarious auricle along the leaf shoulder (Argent, 1976). For these reasons,we suggest that an informal grouping be adopted to satisfy this need and, to avoid confusion with the sections, should be named for the ‘type’ species for each group. Thus, sect. Musa would include the ‘acuminata’ and ‘ornata’ groups and sect. Callimusa the “coccinea’ and ‘textilis’ groups. Jarret & Gawel (1995) considered M. coccinea Andrews sufficiently distinct in their molecular study to be placed in a group of its own, though it should be noted that this result could be due to the fact that no other species in the ‘coccinea’ group was included in their study. In fact it differs from the rest of the group morphologically in lacking the pronounced rugose auricles but when other species in the ‘coccinea’ group are included, M. coccinea clusters with them (Wong ef al., 2002). In any case, if this argument is followed, it would end up with each of the unusual or anomalous species, such as M. johnsii or M. suratii Argent, each being placed in a group of their own, which defeats the purpose of grouping species in the first place. To show the diagnostic characters of each of these groups, a key to the sections and informal groups is given below: Key to Banana Groups (based on Cheeseman, 1947, and Argent, 1976) la. Section Musa. Chromosome number x=11. Bracts usually more or less sulcate, often more or less glaucous, rarely or never polished, convolute or more or less imbricate in the bud, usually strongly revolute on fading. Seeds often dorsiventrally COMMTCSSER, SOIICLMIES SIO OOS Eo sce eosuiccie ease paces DCA as kee 2 Musa in New Guinea & Section of Musa 109 2a. Pseudostems commonly exceeding 3 m tall. Inflorescences pendent or semi-pendent, fruits reflexing towards the base of the rachis. Bracts commonly dull-coloured, green, brownish or dull purple. Flowers many to a bract, in OO Ss, Rk Pe TE A se acuminata group 2b. Pseudostems commonly less than 3 m tall. Inflorescences erect, fruits pointing towards the apex of rachis. Bracts brightly coloured, often pink or red. Flowers few to a bract, usually in a single series............... ornata group 1b. Bracts plane, firm in texture, rarely or never glaucous, strongly imbricate in bud, polished or not, usually not revolute on fading. Seeds barrel-shaped or SE smee ty EG LZ, DANS ass.) GD, Sa, a A... J 3a. Section Ingentimusa. Chromosome number x=7. Pseudostems c.15 m tall. Leaf shoulders reflexed, non-scarious. Inflorescences pendent. Bracts not polished. Seeds strongly angled, 8—9 mm diam. 3b. Section Callimusa Chromosome number x=10. Pseudostems short or tall. Leaf shoulders scarious or non-scarious, never reflexed. Inflorescences erect or pendent. Bracts polished on the outside. Seeds barrel-shaped or strongly angled, up to 5 mm diam.................... 4 4a. Pseudostems commonly less than 3 m tall and inflorescences erect or sometimes pseudostems more than 3 m tall and inflorescences pendent (e.g. M. borneensis). Leaf shoulders usually with well-developed scarious auricle, rarely absent (e.g. M. coccinea). Seed often barrel-shaped or cylindrical with a well-developed oil/air space, rarely Sek a ice nee ada sibs sdaw rotates coccinea group 4b. Pseudostems more than 3 m tall. Inflorescence commonly pendent, sometimes erect (e.g. M. maclayi). Leaf shoulders commonly without a scarious auricle, sometimes scarious (e.g. M. lolodensis). Seeds subglobose or dorsiventrally compressed, sharply angled, without a well-developed oi1/ Bina pane cotati ah OWL AASTL.WOee!.. textilis group Acknowledgements We are indebted to the Directors of the Royal Botanic Garden Edinburgh and the Singapore Botanic Gardens for permission to collect leaf samples from their living collections; to the Academic Research Fund, National Institute of Education, Nanyang Technological University, for research grant RP 12/98/GYY; and GA thanks R.J. Johns, T. Utteridge and the Royal Botanic Gardens Kew for making possible field work on Mt Jaya. 110 Gard. Bull. Singapore 55 (2003) References Argent, G. 1976. The wild bananas of Papua New Guinea. Notes Royal Botanic Garden Edinburgh. 35: 77-114. Argent, G. 2000. Two interesting wild Musa species (Musaceae) from Sabah, Malaysia. Gardens’ Bulletin Singapore. 52: 203-210. Argent, G. 2001. Contributions to the flora of Mount Jaya VI. A new banana, Musa Johnsii (Musaceae) from New Guinea. Gardens’ Bulletin Singapore. 53: 1-7. Cheesman, E.E. 1947. Classification of the bananas. III. The genus Musa L. Kew Bulletin. 106-117. Felsenstein, J. 1993. PHYLIP (Phylogeny Inference Package) version 3.5c. Distributed by the author. Department of Genetics, University of Washington, Seattle. Gawel, N.J. and R.L. Jarret. 1991. Choroplast DNA restriction fragment length polymorphism (RFLPs) in Musa species. Theoretical and Applied Genetics. 81: 783-786. Jarret, R.L. and N.J. Gawel. 1995. Molecular markers, genetic diversity and systematics in Musa. In: Gowen, S. (ed.). Bananas and Plantains. Chapman & Hall, U.K. Chapter 3. Loh, J.P, R. Kiew, A. Kee, L.H. Gan and Y.Y. Gan. 1999. Amplified fragment length polymorphism (AFLP) provides molecular markers for the identification of Caladium bicolor cultivars. Annals of Botany. 84: 155-161. Page, R.D.M. 1996. TREEVIEW: An application to display phylogenetic trees on personal computers. Computer Applications in the Biosciences. 12: 357-358. Reichardt, M.J. and S.J. Rogers. 1993. Plant DNA isolation using CTAB. In: FM. Ausubel, R. Brent, R.E. Kingston, D.D. Moore, J.G. Seidman, J.A. Smith, and K. Struhl (eds). Current Protocols in Molecular Biology. Supplement 22, John Wiley and Sons, USA. Shepherd, K. 1999. Cytogenetics of the Genus Musa. International Network for the Improvement of Banana and Plaintain, Montpellier, France. Simmonds, N.W. 1956. Botanical results of the Banana Collecting Expedition, 1954-5. Kew Bulletin. 463-489. Musa in New Guinea & Section of Musa Lil Simmonds, N.W. 1962. The Evolution of Bananas. Longmans, U.K. Vos, P., R. Hogers, M. Bleeker, M. Reijans, T. van de Lee, M. Hornes, A. Frijters, J. Pot, J. Peleman, M. Kupier and M. Zabeau. 1995. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research. 23: 4407-4414. Wong, C., R. Kiew, J.P. Loh, L.H. Gan, S.K. Lee, Ohn Set, S. Lum, Y.Y. Gan. 2001a. Genetic diversity of the wild bananas Musa acuminata Colla in Malaysia as evidenced by AFLP. Annals of Botany. 88: 1017-1025. Wong, C., R. Kiew, Ohn Set, A. Lamb, S.K. Lee, L.H. Gan, Y.Y. Gan. 2001b. Sectional placement of three Bornean species of Musa (Musaceae) based on AFLP. Gardens’ Bulletin Singapore. 53: 327-341. Wong, C., R. Kiew, G. Argent, Ohn Set, S.K. Lee, Y.Y. Gan. 2002. Assessment of the validity of the sections of Musa (Musaceae) using AFLP. Annals of Botany. 90: 231-238. Zhang, W.P, J.F. Wendel and L.G. Clark. 1997. Bamboozled again! Inadvertent isolation of fungal rDNA sequences from bamboos (Poaceae: Bambusoideae). Molecular Phylogenetics and Evolution. 8: 205-217. 7 - arahas [ 7 Actin posaomra Gers wr ana 101 unindairae me ALAR 2001 aaa A he | fy Heed waa dhind nt. G, 2008. Two intent eis Ate aged Nive ies ks 2 apeaoe Sa ih ae i. ievgiaWM ai Bion Biya Ss et oe mernes blive + rn Pehl 1. 28 snarls tabs | Lie | Piterm a $3: 4 2 di00f 2D VY oe MO | A Oe ae AJ _— wi S TOT 2. “v9 pei Wis a eet oe} deta sora MRE, rst? Zone HOR WE ohiancehs e) gh pane ae as ms Ap haat vis aT “ard \4 vena, orn pen fais ree BES rf , Parola onlasiadl fea rage SRR ‘Me aed a y Aad, cin : saws ape cn 2 SONDUpT AVOh begas t to noitale if ~~ Oe 3 ‘6 7) niimboy A he shania (#9, volun . ‘ OP ’ < - | ; 2 rhencty hans - markers, ret 3 versity ade sy Bananas and Piansestirs. Chagnon ‘ ae 7 ® 4 4 ‘ « ‘ nm 2 - * ry | ~ Agi P > - a sion is t : 2 uy plified fr prem . 1 —* , _ 7 ; ait ~~ >~ ‘+ L. ‘ 7 the sdentificnt 161) o ) eee ea = wad \ re ; “a Fg p a. iy pray loge <a? al Oe: .- : : a" »- ae . Ww Bros tence, ss RS, + ‘ ae ton 2 esing CTAB t nF } ; y ‘ 4 Py 4* , ae 4 ¢ ; a pLUvppye sc rer Las > | od 2 = ,—— n Sirmmorkis, N.W. 1956. Botamieal ns le “ye Bille rit . 4r, 4489. Garden’s Bulletin Singapore 55 (2003) 113-123 Begonias from the Bau Limestone, Borneo, including a New Species RUTH KIEW' AND CONNIE GERI? 'The Herbarium, Singapore Botanic Gardens, Singapore * The Sarawak Biodiversity Centre, Kuching, Sarawak, Malaysia Abstract Six Begonia species (Begoniaceae) are found on limestone hills in the Bau area, Kuching, Sarawak. One, a new species, Begonia lailana Kiew & Geri, is described. A key and habitat notes for all species are provided. The male and female flowers of B. calcarea Ridl. and the female flowers of B. congesta Ridl., not previously recorded, are here described for the first time. Introduction In 1906, Ridley published a paper on the begonias of Borneo that included twenty species, of which five were new species from the Bau limestone (1° 25'N 110° 9'E), 35 km from Kuching, Sarawak. Most were based on his own collections. Two of these were imperfectly known so that descriptions of the male and female flowers of Begonia calcarea Ridl. and the female flowers of B. congesta Ridl. were lacking. As part of the recent biodiversity survey of the Bau limestone carried out by the Sarawak Biodiversity Centre, we collected specimens of begonias from 20 hills. All the species described by Ridley were recollected with the addition of a new species, which is described in this paper. Flowering material of B. calcarea was also collected and the flowers are described below. In addition, the habitats of the species, which are quite different, were recorded. The diversity of begonias on a single limestone hill can be attributed to individual species occupying different and non-overlapping niches (Kiew, 1998). Most of the hills at Bau are home to at least three species, B. pendula Ridl., B. rubida Ridl. and B. speluncae Ridl., with four species collected from Gunung (G.) Doya and G. Jebong and five from G. Kawa. Begonia rubida grows near the summit of limestone hills above the tree canopy exposed to sunlight or in light shade, while B. pendula grows below the tree canopy on damp, mossy, vertical shaded rock-faces. B. speluncae, as 114 Gard. Bull. Singapore 55 (2003) its name suggests, is common around cave mouths but also grows on dry rock-faces near the base of the hills where it occurs. The less common species include B. congesta, which grows on limestone boulders or damp low rock-faces near the cliff base, and B. calcarea and the new species, B. lailana Kiew & Geri, which both grow close to limestone hills on soil associated with the limestone. This soil is frequently more friable and more free-draining than soil derived from other rock types and often supports a different suite of plant species. It is significant that these last three species are less common than the others as this habitat is more susceptible to disturbance, particularly from agriculture, which encroaches towards the base of the hills. Several of these species are decorative and have ornamental value. Begonia calcarea is a handsome species with broad upstanding leaves, which are covered in striking long magenta hairs, particularly on the petiole. In addition, it has large, bright orange or orangey-red flowers with a wavy margin. B. pendula has attractive foliage —its leaves are variegated with silver-green spots. Ridley (1906) described B. speluncae as ‘one of the most attractive limestone species’ with its tufts of small round leaves. For identifying species, Ridley’s key is not entirely satisfactory as both B. pendula and B. rubida key out under ‘stems short, erect and leafy’. B. pendula is a creeping and often pendulous species and B. rubida has tall stems. A key to the Bau limestone species is therefore provided. Key to the Bau Limestone Begonias la. Stems erect and cane-like 2 1b. Stems rhizomatous or creeping ~ 2a. Leaves with veins prominently forked and narrowly diverging, fruits oblong 2. B. congesta 2a. Leaves with veins widely diverging when they fork, fruit much wider distally 3 3a. Leaves ovate, petiole at an angle to the midrib 5. B. rubida 3b. Leaves oblanceolate, petiole straight with the midrib 3. B. lailana 4a. Stems thin and creeping, leaves widely spaced, longer than broad, spotted silver-green 4. B. pendula 4b. Stem thick and rhizomatous, leaves tufted, as wide as broad, dark green, not spotted 5 5a. Robust plant, lamina more than 15 cm wide, petiole more than 15 cm long with dense long magenta hairs, flowers orange 1. B. calcarea Begonias from Bau, Borneo 15 5b. Diminutive plant, lamina up to 4 cm wide, petiole up to 7 cm long, hairs not long nor magenta, flowers white 6. B. speluncae 1. Begonia calcarea Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 260. Holotype: Sarawak, Gunung Bra’ang Haviland s.n. (not seen). Sect. Diploclinium Ridley (1906) described this handsome begonia from a fruiting specimen. It is a very striking plant due not only to its broad, upstanding leaves but also to its long magenta hairs, which cover the whole plant and are particularly striking on the petioles. It is unusual among Bornean begonias in having orange flowers (most begonias have pale pink or white ones). The flowers are produced below the leaves from the prostrate rhizome. They are described here from SBC 3854 for the first time. Inflorescences from the leaf axils or from the prostrate rhizome. Female flowers, 1—2 at one node, bracts c. 10 x 4 mm. Male flowers in umbels 2.5—5 cm long, rachis rosy pink to pale pink, male flowers 3-6; bracts lanceolate 7-10 x 2-4 mm, apex setose; bracteoles 2—6 x 1-3 mm. Male flowers with pedicel 4-19 mm long, tepals 4, orangey- red to deeper red around the margin, margin undulate, glabrous inside and out; isomorphic, outer two broadly oval to almost rotund, apex rounded, 7—11 x 5-9 mm, inner ones slightly narrower, 7—11 x 4-6 mm; stamen cluster hemispherical, c. 4-5 x 8 mm, almost sessile, stamens 25-35, filament c. 1.5—2 mm long, anthers deep yellow, obovate, c. 1.5—2 mm long, apex emarginate. Female flowers with rosy pink peduncle and pedicel, bracteole oval, c. 5 x 2 mm, apex acuminate, ovary c. 19 x 20 mm, glabrous, locules 3, placentas bifid; tepals 5, isomorphic, orangey-red, margin undulate, c.6x 7mm, stigma and styles c. 4 mm long. Fruits with peduncle and pedicel 20-40 mm long, capsule 11—21 x 15-21 mm, wings 3, equal, rounded, 4-12 x 5—7 mm, thinly fibrous. Seeds barrel-shaped, light brown, c. 0.2 mm long, collar cells (0.25—)0.75 length of seed. Doorenbos et al. (1998) doubtfully included Begonia calcarea in sect. Diploclinium. Examination of the female flower shows its ovary and fruit to have three locules and bifid placentas, which confirms its position within this section. Haviland recorded its habitat as ‘top of Mt Bra’ang, old jungle, limestone’, hence the name ‘calcarea’ that Ridley gave. However until our survey, no further specimens had been identified as this species so that there was some doubt that it was a limestone plant. It is certainly a rare species as, although 20 hills were surveyed, it was only found at the base of two of them. It is the most endangered among the Bau limestone species because its habitat is vulnerable to encroaching agricultural practices 116 Gard. Bull. Singapore 55 (2003) from farms close to the base of the hills. A similar specimen (Native Collector 1255) was collected from Gunung Dulit casting doubt as to whether this species is restricted to limestone and endemic to limestone in the Kuching District. Distribution: Malaysia, Sarawak - Kuching limestone and possibly also from non- limestone habitats on G. Dulit. Specimens examined: Gunung (G.) Angob Anderson §27513 (SAR); G. Kawa Connie Geri et al. SBC 3577 (SBC); G. Lanyang (G. Meraja) Burtt B8/54 (SAR), G. Manok Burtt B8134 (SAR), B8154 (SAR); G. Tabai Connie Geri et al. SBC 3854 (SBC). 2. Begonia congesta Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 253. Holotype: Sarawak, Bau Ridley s.n. (not seen). Sect. Petermannia This is a distinct begonia because its leaves are corrugate due to the many, narrowly bifurcating veins. As its name suggests, the inflorescence appears congested because the lateral branches are clustered at the base but above which there is a long (up to 14 cm long) rachis bearing male flowers. This rachis dies and falls before the fruits are mature and was obviously not present in the plants Ridley described. The plant is protogynous with two female flowers produced at the base followed by many small male flowers, which are produced later on short lateral branches and on the erect rachis. Male flowers with pedicel 5—7 mm long, with pale brown hairs c. 0.25 mm long; tepals 2, cream and rosy pink at base, minutely hairy outside, glabrous inside, upper rotund, 4-5 x 4—5 mm, lower broadly oval, 4-5 x 3—4 mm; stamen cluster hemispherical, c. 3—3.5 x 3-4 mm, joined at base in a stalk c. 0.5 mm long; stamens c. 35, filament c. 0.75 mm long, anther obovate, c. | mm long, apex not emarginate. Female flowers with pedicel 4-11 mm long with dense glandular hairs, ovary oblong, to 20 x 17 mm, wing 3-4 mm wide, isomorphic, rounded distally, locules and wings minutely hairy, locules 3, placentas bifid; tepals 5, isomorphic, oval, c. 18 x 6 mm, toothed in distal half, each tooth terminating in a hair, minutely hairy outside, glabrous inside; styles 3, c. 3 mm long, bifid, stigma a spiral papillose band. Fruits pendent, broadly oblong, to 2.5 x 1.7 cm, wings 3, equal, thin and fibrous. Seeds barrel-shaped, c. 0.3 mm long, collar cells c. 0.75 seed length. Ridley (1906) recorded the tepals as shorter than the stamens (surely an error as the stamens are less than 2 mm long) and the stamen number as 12. We counted Begonias from Bau, Borneo cig about 35 stamens. The species is endemic in the Kuching limestone. Ridley cited a specimen from the Niah limestone under this species but this has proved to be a distinct species (Pearce, 2003). Distribution: Malaysia, Sarawak - endemic in the Kuching limestone. Specimens examined: Bau Abang Mohtar S52906 (SAR), Brooke 9875 (SING), Purseglove P4468 (SAR, SING); Bidi Mamit S42147 (SAR); Bukit Kapur Burtt & Woods B188&1 (SAR); Bukit Krian Anderson $25137 (SAR); Fairy Cave Jugah et al. $70076 (SAR); G. Angob Anderson $27493 (SING); G. Batu Martin S39276 (SAR), Meekiong et al. SBC 2125 (SBC); G. Doya Julia et al. SBC 2062 (SBC), Meekiong et al. SBC 2244 (SBC), Raymond et al. SBC 3459 (SBC); G. Jebong Meekiong et al. SBC 2076 (SBC), Paul & Ilias $25626 (SAR); G. Kawa George S38280 (SAR), Meekiong et al. SBC 3113 (SBC); G. Krian Meekiong et al. SBC 1614 (SBC); G. Lanyang Julia et al. SBC 1224 (SBC), SBC 2938 (SBC); G. Pambur Julia et al. SBC 1270 (SBC); G. Ropith Donny et al. SBC 2001 (SBC), Julia et al. SBC 1295 (SBC); Seburan Anderson 8984 (SAR), S/4580 (SAR), Woods B2890 (SAR); G. Tai Ton Denis et al. SBC 1410 (SBC), Julia et al. SBC 726 (SBC); G. Tongga Burtt B&179 (SAR). 3. Begonia lailana Kiew & Geri, sp. nov. Differt a Begonia oblongifolia folis latioribus (8—10 cm nec 5 cm latis), floribus nec albis et fructis alis latioribus (8—11 mm nec 5 mm latis). Typus: Malaysia, Sarawak, Bau, Gunung Kawa. Connie Geri et al. SBC 3753 (holo SAR, iso K, L, SAN, SBC, SING). Section: Petermannia Figure 1 Cane-like begonia up to | m tall with several erect stems from the base, some plants branching near the base to form a bushy crown. Stem succulent becoming woody, reddish or greenish brown, swollen at nodes, up to c. 7 mm diam., glossy and glabrous or hispid with uniseriate translucent hairs c. 2 mm long, dense on the upper internodes and petioles. Stipules lanceolate, pale green, 15-27 x 6-10 mm, midrib prominent, margin entire, apex narrowly pointed, caducous. Leaves alternate, 7(—11) cm apart; petiole reddish, densely hirsute, 3—5 mm in upper leaves, 8—10 mm in lower leaves, thickened at the base, slightly grooved above, in line with the midrib, /amina in juvenile leaves silver-spotted between the veins, in mature leaves matt, plain light green with a red patch at junction with petiole, beneath whitish or sometimes deep red or reddish purple, oblanceolate, asymmetric, 12.5—17 x 8-10 cm, broad side 5—6.5 cm wide, 118 Gard. Bull. Singapore 55 (2003) upper surface glabrous or with scattered dark red bristles between veins and on the veins and midrib, the lower surface with lamina and veins glabrous or veins and midrib minutely hispid, succulent in life, papery when dry, base cuneate and subequal or unequal with the basal lobe of the larger side rounded, 0.5—1 cm long, margin minutely toothed, apex acuminate; venation palmate-pinnate with 2 pairs at base and 3 lateral pairs along midrib, branching towards margin, impressed above, beneath prominent and concolorous with lamina or sometimes red, basal lobe with 2 veins. Plant protogynous. /nflorescences axillary, erect, cymose panicle, 10-16 cm long of which peduncle is 0Q—3.5 cm long, greenish to dark red, minutely pilose, lowest node with 1—2 female flowers, above up to 11 short branches 1—1.5 cm long with many male flowers. Bracts green, ovate, 3-4 mm long. Male flowers with pale red pedicel 4—9 mm long, tepals 2, glabrous, deep pink or pale cream and deeper pink towards the base, rotund, 6—7 x 6-7 mm or oval 5—8 x 3—7 mm, apex rounded; stamen cluster 3—5 x 2-3 mm with a stalk c. 1 mm long, stamens more than 35, filament c. 0.5 mm long; anther pale yellow, obovoid, c. 1—-1.5 mm long, apex emarginate, dehiscing by longitudinal slits in upper half. Female flowers with pedicel 7-9 mm long, reddish green; ovary pale green sometimes with red veins on the wings, 13-18 mm long, locules 3, each with 2 axile placentas, wings 3, equal, 3-5 mm wide; tepals 5, pale peach to pale pink or green, sometimes deep ruby red at the base, slightly obovate, 6-12 x 3-8 mm, innermost smaller 4-10 x 3—7 mm, the margin entire or with 4-5 minute teeth towards the apex, apex acute, the outer surface sometimes with scattered translucent hairs; styles 3, bifurcating, spirally twisted, yellowish-green to pale green, 2—6 mm long; stigma a glistening papillose spiral band. Fruits with stiff, decurved pedicel 10-13 mm long, capsule glabrous, 2—2.7 x 1.8—2.8 cm, wings slightly unequal, slightly pointed at tip, thinly fibrous, widest wing 8—11 mm wide, narrowest 6-8 mm wide, dehiscing between locule and wing, style not persistent. Seeds barrel-shaped, light brown, c. 0.4 mm long, collar cells c. 0.4 length of seed. Distribution: Malaysia, Sarawak - endemic in Kuching limestone. Locally common on G. Aup, G. Poing and G. Taba. Habitat: At the base of limestone hills, in light shade. Notes: Begonia lailana is named in honour of YABhg. Datuk Amar Puan Sri Dr Hajjah Laila Taib, wife of the Chief Minister of Sarawak. This species belongs to sect. Petermannia because the ovary has three locules each with two placentas. It resembles B. oblongifolia in being a cane-begonia with obovate leaves, paniculate male inflorescences and oblong fruits about 2 cm long. However, it is distinct from this species in its leaves which are narrower, less than twice as long as broad (they are c. 5 cm wide and more than twice as long as broad in B. oblongifolia), in having fewer veins (8 in B. oblongifolia), larger, light red male Begonias from Bau, Borneo 119 7mm Figure 1. Begonia lailana Kiew and Geri. A foliage; B partial inflorescence; C male bud; D male flower; E stamen cluster; F. stamen; G & H female flower; I style and stigma; J. TS ovary. 120 Gard. Bull. Singapore 55 (2003) flowers (3 mm long and green in B. oblongifolia) and wider wings (c. 5 mm wide in B. oblongifolia). Begonia lailana populations and individuals show some variation in leaf colour and indumentum, and in ovary colour. Plants in some populations have completely green leaves (e.g. at G. Kawa), in others most plants have green leaves with some plants with red veins on the lower surface (e.g. at G. Tabai), or have most or some plants with a deep red or purplish red undersides to the leaf (e.g. at G. Poing and G. Aup, respectively). The shape of the silver spots on the juvenile leaves also varies between populations: either discrete (e.g. at G. Kawa and G. Poing) or coalescent (e.g. atG. Aup). Some populations have completely green ovaries (e.g. at G. Aup and G. Poing), or the wings of the green ovary have fine red veins (e.g. at G. Kawa and G. Tabai). Begonias are notoriously polymorphic for colour and patterning of the leaves so these differences are not considered significant taxonomically. While the stiff red bristles on the upper lamina surface are conspicuous, they are not always present in all plants, e.g. the G. Aup population has glabrous leaves, the Poing population has mostly glabrous individuals with a few with bristly leaves, while the G. Kawa population has plants with bristly leaves. The female flower may also vary in that the tepal margin may be entire or slightly toothed. However, this difference does not correlate with any other character and is therefore not considered taxonomically significant. Other specimens examined: Gunung (G.) Aup Connie Geri et al. SBC 3756 (SBC), Malcom et al. SBC 3303 (SBC); G. Batu Meekiong et al. SBC 2136 (SBC); G. Doya Julia et al. SBC 2055 (SBC), Raymond et al. SBC 3451 (SBC), SBC 3473 (SBC); G. Kawa Connie Geri et al. SBC 3753 (SBC), SBC 3592 (SBC, SING), Meekiong et al. SBC 3114 (SBC); G. Lanyang Julia et al. SBC 2901 (SBC, SING); G. Podam Malcom et al. SBC 3364 (SBC); G. Poing Julia et al. SBC 353 (SBC); Connie Geri et al. SBC 3755 (SBC), Malcom et al. SBC 1555 (SBC); G. Tabai Connie Geri et al. SBC 3757 (SBC), Julia et al. SBC 3414 (SBC). 4. Begonia pendula Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 257. Holotype: Jambusan Ridley 11772 (K). Sect. Petermannia This species is endemic in the Kuching District where it is common on most limestone hills. It creeps over vertical mossy cliff faces below the tree canopy with its thin stems rooting at the nodes. Ridley (1906) named it ‘pendula’ describing its habit as having trailing stems hanging down rock-faces. It is a decorative species as its bright green leaves are variegated with silver-green elongated spots between the veins that in some Begonias from Bau, Borneo 121 plants are discrete and in others coalescent. Some plants have leaves with the lower surface maroon, others are green. The male flowers are deep pink and the female flowers are red. Distribution: Malaysia, Sarawak - endemic in Kuching limestone. Other specimens examined: Bau Abang Mohrar et al. S52914 (SAR), Anderson 7799 (SAR, SING), Brooke 9891 (SING), Ridley 11774 (K); Bidi Clemens 20680 (SAR), Yii et al. SSO381 (SAR); Bukit Boring Yii et al. S50360 (SAR); G. Apin Donny et al. SBC 1360 (SBC), Jugah et al. S79954 (SAR), Makom et al. SBC 1107 (SBC), Malcom etal. SBC 1107 (SBC); G. Aup Julia et al. SBC 568 (SBC), Malcom et al. SBC 3301 (SBC); G. Batu Meekiong et al. SBC 2126 (SBC), Meekiong et al. SBC 2126 (SBC), SBC 2127 (SBC); G. Berloban Yii & Othman S$46234 (SAR); G. Doya Burtt BS1&7 (SAR), Julia et al. SBC 2027 (SBC), SBC 2144 (SBC), SBC 2154 (SBC), SBC 2162 (SBC), Sheavy et al. SBC 3056 (SBC); G. Jebong Donny et al. SBC 2095 (SBC), Meekiong et al. SBC 2074 (SBC), Paul & Ilias $25623 (SAR); G. Juita Connie Geri et al. SBC 3425, 3426 (SBC), Julia et al. 1293 (SBC); G. Pambur Julia et al. SBC 1263 (SBC) G. Podam Meekiong et al. SBC 3206 (SBC); G. Poing Julia et al. SBC 364 (SBC); G. Ropth Julia et al. SBC 2101 (SBC), Meekiong et al. SBC 3186 (SBC), Shaevy et al. SBC 2426 (SBC); G. Selabor Sinclair SFN 38471 (SING); G. Setiak Martin $38660 (SAR); G. Stulang Malcom et al. SBC 640 (SBC), SBC 643 (SBC); G. Tabai Burtt BS158 (SAR), Julia et al. SBC 1293 (SBC), SBC 2513 (SBC), SBC 2563 (SBC); G. Tai Ton Julia et al. SBC 2677 (SBC), Stevens et al. 204A (SAR), Yii et al. S51208 (SAR), $51210 (SAR); G. Tongga Donny et al. SBC 1777 (SBC); G. Umbut Julia et al. SBC 2622 (SBC); Lobang Angin Yii et al. S51265 (SAR); Saburan Anderson S$19119 (SAR). 5. Begonia rubida Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 256. Lectotype: Jambusan Ridley 12393 (K, here designated). Sect. Petermannia As the name suggests, this species has ruby-red male flowers and its stems are also bright red. It grows above the tree canopy on the shoulders of the summit, often on peaty soil among sparse vegetation in light shade to full sunlight. Its leaves are thick and succulent and are held vertically, unlike the leaves of the other cane-like begonias that grow in shaded conditions, e.g. B. Jailana and B. congesta, the leaves of which are held horizontally. This difference is probably a response to light conditions: the horizontal leaf surface will intercept more light in shaded conditions, while leaves held vertically will absorb less heat in bright sunlit conditions. 122 Gard. Bull. Singapore 55 (2003) Distribution: Malaysia, Sarawak — endemic in Kuching limestone. Other specimens examined: Bau Brooke 9832 (SING), Bidi Clemens 20681 (SAR); Bukit Jebong Johnson S29403 (SAR), Paul & Ilias $25617 (SAR); Bukit Manok Anderson $27416 (SING), Erwin & Paul S27416 (SAR); Bukit Numpang Chai & Seng S22861 (SAR); Bukit Regu Paul et al. $37392 (SAR); G. Apin Donny et al. SBC 1361 (SBC), Jugah et al. S79953 (SAR), Malcom et al. SBC 1108 (SBC); G. Batu Meekiong SBC 2136, 2137 (SBC); G. Juita Connie Geri et al. SBC 3427 (SBC); G. Meraja Julia et al. SBC 315 (SBC); G. Pambur Julia et al. SBC 1240 (SBC), SBC 1264 (SBC); G. Ropih Julia et al. SBC 2100 (SBC), Connie Geri et al. SBC 3516 (SBC); G. Setiak Martin $38670 (SAR), G. Taiton Julia et al. SBC 710 (SBC); G. Tongga Denis et al. SBC 1722 (SBC), SBC 1732 (SBC); G. Umbut Julia et al. SBC 2584 (SBC). 6. Begonia speluncae Ridl., J. Str. Br. Roy. As. Soc. 46 (1906) 258. Lectotype: Jambusan Ridley 11773 (K, here designated). Sect. Reichenheimia This begonia is, as its name suggests, a species that grows around cave mouths. It also grows on dry rock cliff faces at the base of hills. It is endemic in the Kuching limestone and has been collected from most hills in this area. Doorenbos et al. (1998) placed this species doubtfully in sect. Reichenheimia as the placenta type was not known. We confirm that it belongs to this section because the ovary and fruit have three locules and the placenta is unbranched, 1.e. one per locule. Other specimens examined: Bidi Anderson 75 (K), Brooks s.n. (K) Clemens 20644 (K, SAR), Mjoberg 177 (K); Bukit Pait Erwin & Paul S27429 (SAR); Bukit Selabor Paie S28044 (SAR, SING); Fairy Cave Rantai et al. S 70059 (SAR); G. Aup Julia et al. SBC 707 (SBC); G. Bar Rantai et al. S66034 (SAR); G. Batu Meekiong et al. SBC 2135 (SBC); G. Doya Julia et al. SBC 2159 (SBC), Shaevy et al. SBC 3047 (SBC); G. Jebong Lehman $30143 (SAR), Meekiong et al. SBC 2072 (SBC); G. Kawa Meekiong et al. SBC 2797 (SBC); G. Lanyang Julia et al. SBC 1229 (SBC), SBC 2905 (SBC), Denis et al. SBC 1439 (SBC); G. Manok Burtt BS132 (SAR); G. Meraja Julia et al. SBC 301 (SBC); G. Podam Malcom et al. SBC 814 (SBC); G. Poing Julia et al. SBC 355 (SBC); Quop Hewitt s.n. (K); Sungai Serian Jacobs 5177 (SAR). Begonias from Bau, Borneo 123 Acknowledgements We are grateful to the ASEAN Regional Centre for Biodiversity Conservation for funding The Biodiversity Survey of the Bau Limestone Project, of which this study is a part; to the curators of the herbaria at Kew and the Forest Department, Kuching, for permission to examine specimens in their care; to Sarawak Biodiversity Centre colleagues, past and present, for their help in the field, to Raymond Sylvester Bohari for preparing the illustration; and to M.J.S. Sands, Kew, for helpful suggestions on improving the manuscript. References Doorenbos, J., M.S.M. Sosef and J.J.RE. de Wilde. 1998. The Sections of Begonia. Wageningen Agriculture University Papers. 98-2. Kiew, R. 1998. Niche partitioning in limestone begonias in Sabah, Borneo, including two new species. Gardens Bulletin Singapore. 50: 161-169. Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia. Gardens Bulletin Singapore. 55: 73-88. Ridley, H.N. 1906. Begonias of Borneo. Journal of Straits Branch of Royal Asiatic Society. 46: 246-261. > . = - Diserib aaa a: Site ale ro i on sein tile eae — “navn ret ha © ; ya Ww erent ad ie ae eter ohadabed ‘i se Mii roc ih ly id i rte Ss 1 AANA by al 8? Rae 6 SA SIAC? Mal ioe ie shel oR hid sol ERS abe vate ants BA Balint u rae 4e; viii 5 SG wey it C) Cann eGerie ah SBC i nia ef al SB ¢ FiO (SE « og lsh SBC ur G, L ntn shalt 7 LS . L ) TAY wa J ' 4 LU bin Yoz0d | M2 a - : mes wherein J gurtion a ’ | ‘ mOY, ) agevhiy sae PV nt A cided SRGRSAN Hi ont spitmen stoi ~ 4 WOOD 7, Wid, nasil: if iano et j _ , ' i inno say Lee i AAS at May a Pur, SE PRG E/E . 322 : J ; i? YS TOISeO\” ree Henn og Cea 2 eases Chaat vt ox ena 4 ; ‘ ' ; H : a 1s chee +4 “yar | AA +4 wa On. (Ky Clemen * > Se ; y 7 Py , uy y A test ‘ na = ii : ', ‘* ; G:Aap lia wit § ‘ fe ekg éf ’ oe é |. SHC HOT SBC}! iG. 2(3BC}G ae a 3 Séet 123 {Si oy , SOC 2 ; Koyo > a ae SAIS. tS ay G. iS. IB t iy j G doit : 2 ey bi a 7 77 : my Of =e a), * F it?s ee . ere As ‘0D J . Orla é srobs SIP 2 oo i aoe - , ; oJ Jae ’ . - _ rs ' Garden’s Bulletin Singapore 55 (2003) 125-162 New Species and New Records of Southeast Asian Bulbophyllum (Orchidaceae) J.J. VERMEULEN Singapore Botanic Gardens Singapore 259569 AND F, 2, BYRNE 14 Clementi Cresent Singapore 599517 Abstract Seventeen new species of Bulbophyllum are described: B. tectipes (sect. Aphanobulbon, from Sulawesi), B. pseudopelma and B. simplex (both sect. Desmosanthes, from Sulawesi), B. flavorubellum (sect. Desmosanthes, from Sabah), B. decurrentilobum and B. iterans (both sect. Hybochilus, from Sulawesi), B. furcillatum (sect. Hybochilus, from Sumatra), B. anisopterum (sect. Monilibulbus, from Sabah), B. anakbaruppui and B. illecebrum (both sect. Osyricera, from Sulawesi), B. uncinatum (sect. Polyblepharon, from Sulawesi), B. catillus, B. lyriforme and B. sinapis (sect. Sestochilus, all from Papua New Guinea), B. coweniorum (sect. Sestochilus, from Laos), B. incisilabrum (sect. Sestochilus, from Sulawesi), and B. turpis (sect. Sestochilus, from Peninsular Malaysia). Bulbophyllum plumatum Ames, and B. spissum J.J. Verm. are newly recorded for Peninsular Malaysia. Introduction The collection of orchid species in Singapore Botanic Gardens appears an inexhaustible source of interesting plants. In this paper new species of the genus Bulbophyllum that recently came to light are described. Several more new species of the same genus, based on material from different sources (L, K) are added. Two Bulbophyllum species that are newly recorded for Peninsular Malaysia are discussed. 126 Gard. Bull. Singapore 55 (2003) Bulbophyllum sect. Aphanobulbon Bulbophyllum tectipes J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 1. Bulbophyllum tectipes J.J. Verm. & P. O’Byrne, inter species rhizoma patenti ad pendenti inflorescentia multiflora sect. Aphanobulbonis foliis 9.5—15 cm longis, pedunculo omnino bracteis 3 vel 4 in duabus seriebus imbricatis acute carinatis tecto notatum. — TYPE: Indonesia, Sulawesi, central part, SBG-O 1953 (SING, holo.). Roots creeping along the rhizome towards its point of attachment, then spreading over the substrate. Rhizome patent to + pendulous with stiff, + patent branches, up to 80 cm long, 2—2.5 mm diam., sections between pseudobulbs 8-9 cm long, bracts hardly persistent, distinctly keeled. Pseudobulbs distant, minute, cylindrical, 0.2—0.25 x 0.2-0.25 cm. Petiole 0.5—0.9 cm long. Leaf blade elliptic to ovate, 9.5—15 x 2.3—3.2 cm, index (length/width) 4.1—4.9; acute. /nflorescence patent, a rather dense raceme, 4.4—10.4 cm long, 14—26-flowered. Peduncle 1.4—2.2 cm, bracts 3-4, distichous, imbricate, the longest 14-20 mm long; acuminate, sharply keeled. Rhachis 3-8.2 cm. Floral bracts triangular, c. 3.8 x 1.5 mm, acute. Flowers not resupinate, not fully opening, many simultaneously. Pedicel and ovary c. 2.2 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat recurved, elliptic, c.6 x 2 mm, index c. 3; acuminate, margins entire, base broadly attached; thin, 3-veined, glabrous. Lateral sepals recurved, free, oblique, triangular, c. 6 x 2.8 mm, index 2.1—2.2; acute, otherwise as the median sepal. Petals porrect, elliptic, c.2.6 x 0.8 mm, index 3.2—3.3; acute, margins entire, slightly papillose towards the tip, base rather narrowly attached; thin but somewhat thickened towards the tip, 1-veined, surface glabrous. Lip recurved + half-way, general outline + elliptic, c. 2.8 x 1.4 mm, index c. 2 (all without artificial spreading); acuminate, margins entire; rather thin, glabrous; adaxially concave and with 2 conspicuous, rounded knobs near the base, with a median furrow up to half-way the length of the lip, surface slightly convex towards the tip; abaxially with a distinct, truncate ridge up to half-way the length of the lip. Column c. 1 mm long, column foot with a low ridge starting at the base of the stigma, and continuing over 2/3 of the length of the column foot, column foot without a tooth just above the ligament. Ste/idia porrect, triangular, c. 0.5 mm long, acute, along the lower margin with a distinct, triangular, subacute tooth. Anther abaxially with a slight crest distally, surface + glabrous, front margin not drawn out. Pollinia 2, semi-ellipsoid; no appendages present. Figure 1.Bulbophyllum tectipes J.J. Verm. & P. O’Byrne a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side, below: adaxial side; g. Pollinia, left: single, right: a pair. — All from SBG-O 1953 (living plant, spirit sample). Southeast Asian Bulbophyllum 127 Figure 1. Bulbophyllum tectipes J.J. Verm. & P. O’Byrne 128 Gard. Bull. Singapore 55 (2003) Colours: Plant glossy medium green. Flowers white. Habitat: Epiphyte in montane forest, alt. 1 100—1300 m. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Most similar to Bulbophyllum agapethoides Schltr. and B. falciculicorne J.J. Sm., also from Sulawesi. The three species share a relatively inflexible rhizome. B. tectipes is much larger vegetatively; the leaves are consistently almost twice as long and more than twice as wide as in the species mentioned. Most characteristic, however, are the conspicuous distichous, sharply keeled peduncle bracts that cover most of the peduncle. B. rhizomatosum Ames & C. Schweinf. (Borneo, Peninsular Malaysia), B. acuminatifolium J.J. Sm., and B. perpendiculare Schltr. (both from Sulawesi) have limply pendulous rhizomes. Bulbophyllum sect. Desmosanthes Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 2. Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne, a Bulbophyllo flammulifero rhachidi 0.5—1 cm longa floribus 4—8, stelidiorum margine basali sine ala antrorsa differt. - TYPE: Malaysia, Sabah, Interior Province, Nabawan, Vermeulen & Lamb 433 (K, holo., L, iso.). Roots present along the entire rhizome. Rhizome creeping, 1—1.4 mm diam., sections between pseudobulbs 2—5 cm long, bracts hardly persistent. Pseudobulbs distant, narrowly ellipsoid to subcylindrical, 1.7—3 x 0.3—0.6 cm. Petiole 0.3—0.4 cm long. Leaf blade elliptic, 3.2-4.5 x 1.2—1.9 cm, index (length/width) 2.3—2.7; obtuse. Inflorescences (sub-)erect, short racemes, 6—11 cm long, 4—8-flowered. Peduncle 5.5—10 cm, bracts 5—7, the longest 5—9 mm long. Rhachis 0.5—1 cm. Floral bracts ovate, c. 4.8 x 2.2 mm, acuminate. F/owers not resupinate, not fully opening, several open simultaneously. Pedicel and ovary c. 4 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat recurved with an incurved top, subtriangular, c. 9.5 x 2 mm, index 4.7-4.8; acute, margins entire, base broadly attached; rather thin, 3-veined, glabrous. Lateral sepals recurved, free, oblique, triangular, c. 9.5 x 2 mm, index 4.7-4.8; otherwise as the median sepal. Petals porrect, ovate-triangular, c. 2.2 x 0.8 mm, index 2.7—2.8; obtuse, margins slightly erose distally, base broadly attached; thin, 1-veined, glabrous. Lip recurved distally, general outline + ovate, c. 1.8 x 1.2 mm, index c. 1.5, acute, margins entire, very finely papillose proximally; thick, surface glabrous; adaxially slightly concave near the base, Figure 2. Bulbophyllum flavorubellum J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, above: two pairs, below: single pair. — All from Vermeulen & Lamb 433 (living plant, spirit sample). Southeast Asian Bulbophyllum 129 Figure 2. Bulbophyllum flavorubellum J.J. Verm. & P. O’Byrne 130 Gard. Bull. Singapore 55 (2003) with 2 obtuse ridges over most of the length of the lip, towards the base inconspicuous and diverging, towards the tip distinct and fused to a V-shaped structure; abaxially with a distinct, retuse ridge over most of the length of the lip, surface glabrous. Column c. 2.2 mm long, stigma without a tooth at its base, column foot with a small, obtuse knob just above the ligament. Stelidia porrect, falcate, triangular, c. 1 mm long, acute, along the lower margin with an inconspicuous, deltoid, obtuse wing. Anther abaxially without a crest, surface + glabrous, front margin drawn out into a flat, triangular, acute tip. Pollinia 4, the inner slightly shorter than the outer, all obovoid, no appendages present. Colours: Sepals yellow proximally, bright orange red distally. Petals very pale yellow. Lip orange. Column yellow. Flowers slightly scented. Habitat: Terrestrial or as an epiphyte close to the forest floor, in lowland kerangas forest. Alt. c. 300 m. Distribution: MALAYSIA: Sabah (1 specimen seen). Notes: On first sight, this could be taken for a exceptionally small form of Bulbophyllum flammuliferum Ridl., from Peninsular Malaysia and Borneo. That species, however, has a 2.5—10 cm long rhachis with 12-25 flowers, and a distinct, antrorse, rounded wing along the lower margin of the stelidia. Bulbophyllum pseudopelma J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 3. Bulbophyllum pseudopelma J.J. Verm. & P. O’Byrne, inter series speciebus inflorescentiis racemosis Sect. Desmosanthes labello brevi lato, stelidiis brevissimis notatum. — TYPE: Indonesia, Sulawesi, central part, SBG-O 3216 (SING, holo). Roots present along the entire rhizome. Rhizome creeping or straggling away from the substrate, 34.5 mm diam., sections between pseudobulbs 4.5—26 cm long, bracts little persistent. Pseudobulbs distant, ovoid, 2.54 x 1.5—2 cm. Petiole 0.6—1.1 cm long. Leaf blade elliptic-ovate, 9.5—17.5 x 1.9—2.5 cm, index (length/width) 5—7; obtuse. Inflorescences many, often tufted, along the entire rhizome, (sub-)erect, lax racemes, 2-5.5 cm long, 2-4-flowered. Peduncle 1—1.8 cm, bracts c. 3, the longest 4-5.5 mm long. Rhachis 0.4—3.5 cm. Floral bracts tubular proximally, 4-5 mm long, acute. Flowers not resupinate, not fully opening, several open simultaneously. Pedicel and ovary c. 4.5 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat recurved with an incurved top, elliptic-ovate, c. 7 x 3 mm, index c. 2.3—2.4; acuminate, margins entire, base rather broadly attached; thin, 3-veined, glabrous. Lateral sepals recurved, free, oblique, triangular, c. 7 x 2.5 mm, index c. 2.8; acute, base broadly Figure 3. Bulbophyllum pseudopelma J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e. Lip, abaxial side; f. Column and lip, lateral view; g. Anther, above: abaxial side, below: adaxial side; h. Pollinia, above: two pairs, below: single pair. — All from SBG-O 32/6 (living plant, spirit sample). Southeast Asian Bulbophyllum 131 Figure 3. Bulbophyllum pseudopelma J.J. Verm. & P. O’ Byrne 2 Gard. Bull. Singapore 55 (2003) attached; otherwise as the median sepal. Petals porrect, ovate, c. 3.2 x 2.8 mm, index 1.1-1.2; rounded, margins entire, base rather narrowly attached; thin, 3-veined, glabrous. Lip recurved distally, general outline ovate, c. 1.4 x 1.2 mm, index 1.1—1.2; rounded, margins entire; thick and soft; adaxially slightly concave near the base, with 2 rather distinct, obtuse ridges proximally somewhat diverging and running parallel up to c. 2/3 of the length of the lip, surface slightly convex towards the tip, glabrous; abaxially with a distinct, retuse ridge over most of the length of the lip, surface finely papillose in centre. Column c. 0.7 mm long, stigma with a transverse ridge at its base, column foot with an obtuse knob just above the ligament. Ste/idia minute, porrect, triangular, acute, along the lower margin with a distinct, antrorse, rounded wing. Anther abaxially with a distinct crest, surface + glabrous, front margin not drawn out. Pollinia 4, the inner slightly shorter than the outer, ellipsoid, the outer obovoid; no appendages present. Colours: Floral bracts greyish white. Flowers very pale greenish, pale yellow towards the base. Habitat: Epiphyte in montane forest, on the trunk and in the crown of a small tree. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Among the racemose species of sect. Desmosanthes, Bulbophyllum pseudopelma is easily distinguished by the short and wide, fleshy lip and the very short stelidia. In its general aspect this plant is reminiscent of B. reptans Lindl., a species widespread in continental SE Asia. Apart from being sturdier, it is distinct in the those characters mentioned above. In the general aspect of its inflorescence, B. pseudopelma resembles some species of sect. Pelma. Bulbophyllum simplex J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 4. Bulbophyllum simplex J.J. Verm. & P. O’Byrne, a Bulbophyllo trifloro J.J. Sm. sepalis 9—10 mm longis sine apice incrassato differt. — TYPE: Indonesia, Sulawesi, central part, SBG-O 3210 (SING, holo.). Roots present along the entire rhizome. Rhizome creeping or straggling away from the substrate, 0.8—1.5 mm diam., sections between pseudobulbs 5—28 cm long, bracts moderately persistent. Pseudobulbs distant, ovoid, 1.5—2.4 x 0.6—0.9 cm. Petiole 0.1—0.3 cm long. Leaf blade elliptic-ovate, 3.5—6.5 x 1.1—1.5 cm, index (length/width) 3.1—5; obtuse. Inflorescences often along the entire rhizome, (sub-)erect, subumbellate racemes, 1.8—2.5 cm long, 3—5-flowered. Peduncle 1.2—1.6 cm, bracts c. 2, the longest Figure 4. Bulbophyllum simplex J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, adaxial side; e. Lip, abaxial side; f. Column and lip, lateral view; g. Anther, left: abaxial side, right: adaxial side; h. Pollinia, above: two pairs, below: single pair. — All from SBG-O 32/0 (living plant, spirit sample). Southeast Asian Bulbophyllum 133 Figure 4. Bulbophyllum simplex J.J. Verm. & P. O’Byrne 134 Gard. Bull. Singapore 55 (2003) 4—5 mm long. Rhachis 0.2-0.25 cm. Floral bracts triangular, 3.2—3.6 x 1.2 mm, acute. Flowers partly resupinate, not fully opening, several open simultaneously. Pedicel and ovary 2.6—4.2 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat recurved with an incurved top, ovate, 9-10.5 x 3—3.3 mm, index 3-3.2; acuminate, margins entire, base rather narrowly attached; thin, 5-veined, glabrous. Lateral sepals recurved, free, oblique, triangular, 8—11.3 x 2.3-2.4 mm, index 3.44.4; acute-acuminate, base broadly attached; 3-4veined, otherwise as the median sepal. Petals porrect, elliptic, 3.5—4 x 2—2.2 mm, index 1.7—1.9; acuminate, margins slightly erose, base narrowly attached; thin, 3-veined, glabrous. Lip recurved, general outline + oblong, 1.6—1.8 x 0.9—1.2 mm, index 1.5—1.8 (all without artificial spreading), obtuse, margins finely papillose distally; rather thick; adaxially + flat near the base, with 2 obtuse ridges over most of the length of the lip, towards the base inconspicuous and diverging, towards the tip distinct and fused to a v-shaped structure, adaxial surface finely papillose distally; abaxially with a distinct, retuse ridge over most of the length of the lip, surface glabrous. Column c. 2 mm long, stigma without a tooth at its base, column foot with a small, obtuse knob just above the ligament. Stelidia porrect, falcate, triangular, c. 1 mm long, acute, along the lower margin with an inconspicuous, deltoid, obtuse wing. Anther abaxially with a distinct crest, surface + glabrous, front margin drawn out into a concave, rounded beak. Pollinia 4, the inner + as long as the outer, all subtriangular, no appendages present. Colours: Flowers completely white. Habitat: Epiphyte in montane forest, in the crown of a small tree. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Vegetatively and in the general aspect of the flowers Bulbophyllum simplex is fairly similar to B. triflorum J.J. Sm., from Java. That species differs in having consistently smaller flowers, with shorter sepals (3—4 mm long) with a thickened tip. B. concinnum Hook.f., from Thailand and west Malesia, differs in the same characters, as well as in having the pseudobulbs closer together and in having a narrower, ovate lip. Bulbophyllum sect. Hybochilus Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 5. Bulbophyllum decurrentilobum J.J. Verm. & P.O’ Byrne, a Bulbophyllo masarangico Schltr. labello lobis lateralibus distaliter lobo mediali sine sinu transientibus Figure 5. Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right: adaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left: single pair, right: two pairs. — All from SBG-O 3206 (spirit sample). Southeast Asian Bulbophyllum 135 Figure 5. Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne 136 Gard. Bull. Singapore 55 (2003) differt. - TYPE: Indonesia, Sulawesi, central part, SBG-O 3206 (SING, holo.). Roots mainly below the pseudobulbs. Rhizome creeping, 0.7—0.9 mm diam., sections between pseudobulbs 0.4—1 cm long, bracts not persistent. Pseudobulbs distant, ovoid, 0.3—0.5 x 0.2-0.45 cm. Petiole up to 0.05 cm long. Leaf blade elliptic, 0.6—-1 x 0.2-0.25 cm, index (length/width) 3-4; acuminate. /nflorescence erect to patent, 1.5—1.7 cm long, 1-flowered. Peduncle 0.8—0.9 cm, bracts 1, c. 1.5 mm long. Floral bracts tubular proximally, c. 1.3 mm long, + acute. Flowers not fully opening. Pedicel and ovary 1.1—1.2 mm long, basal node coinciding with the floral bract. Median sepal recurved, ovate, c. 7 x 1.3 mm, index 5.3—5.4; long acuminate, margins entire, base rather broadly attached; thin, 3-veined, glabrous. Lateral sepals oblique, c.8x 1.3 mm, index c. 6.1—6.2; otherwise as the median sepal. Petals porrect, elliptic, c. 1.8 x 0.7 mm, index c. 2.5—2.6; acuminate, margins erose distally, base rather narrowly attached; thin, 1-veined, glabrous. Lip recurved in the distal half, general outline ovate, c. 1.9 x 0.9 mm, index 2.1—2.2 (all without artificial spreading), margins entire; glabrous, 3-lobed; midlobe ovate, rounded, rather thin; adaxially somewhat convex and with a distinct, conical, obtuse knob near the base; abaxially without a ridge; lateral lobes erect, attached along the proximal half, c. 0.5 mm wide atc. 1/3 of the length of the lip, decurrent in front, rounded, thin. Column c. 0.6 mm long, stigma without a ridge at its base, column foot without teeth. Ste/idia porrect, ovate, c. 0.3 mm long, + truncate. Anther abaxially with a distinct, finely papillose crest, surface otherwise + glabrous, front margin drawn out into an incurved, triangular, subacute beak with entire margins. Pollinia 4, the inner slightly more than half as long as the outer, all obovoid; no appendages present. Colours: Sepals and petals purple, whitish near the base. Lip purplish. Habitat: Epiphyte in montane forest, on the trunk and on thin branches in the crown of small trees. Alt. 1800—2500 m. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Most similar to Bulbophyllum masarangicum Schltr., from the northern part of Sulawesi (Minahassa). The main difference between the two is in the shape of the lip: in B. decurrentilobum the front margin of the lateral lobes of the lip gradually merges with the midlobe, in B. masarangicum it does so with a distinct sinus. B. masarangicum also has smaller flowers, the sepals being 3—5 mm long. Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 6. Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne, a Bulbophyllo camptochilo J.J. Figure 6. Bulbophyllum furcillatum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, above: two pairs, below: single pair. — All from SBG-O 0922 (spirit sample). Southeast Asian Bulbophyllum 137 \ N N penwvssse Rieter or ol? - . . Figure 6. Bulbophyllum furcillatum J.J. Ve rm. & P. O’Byrne 138 Gard. Bull. Singapore 55 (2003) Verm. sepalis lateralibus papillosis, labello longioribus angustioribusque differt. — TYPE: Indonesia, Sumatra, Barisan Range, SBG-O 922 (SING, holo.). Roots usually one below each pseudobulb. Rhizome creeping, 0.8—1 mm diam., sections between pseudobulbs 0.35—1 cm long, bracts not persistent. Pseudobulbs distant, ovoid, 0.15—0.32 x 0.15—0.25 cm. Petiole up to 0.05—0.13 cm long. Leaf blade elliptic to ovate, 0.6—1.2 x 0.4—0.7 cm, index (length/width) 1.5—2.3; mucronate. Inflorescence erect to patent, 9-14 mm long, 1-flowered. Peduncle 0.3—0.7 cm, bracts 2, the longest 1.3—2.2 mm long. Floral bracts tubular proximally, c. 1.6 mm long, acuminate. Flowers not fully opening. Pedicel and ovary 2.8-4 mm long, basal node coinciding with the attachment of the floral bract. Median sepal + porrect, subtriangular, c. 2.8 x 1.3 mm, index 2.1—2.2; acuminate, margins finely papillose distally, base rather broadly attached; rather thin, thickened distally, 3-veined, adaxially finely papillose distally. Lateral sepals slightly oblique, obovate, c. 2.9 x 1.8 mm, index c. 1.6—1.7; + acute, base broadly attached; adaxially finely papillose except the proximal part of the upper half; otherwise as the median sepal. Petals porrect, + elliptic, c. 1.3 x 1 mm, index c. 1.3; subacute, margins erose distally, base narrowly attached; thin, 1-veined, glabrous. Lip recurved + half-way, general outline triangular, c. 1.5 x 0.5 mm, index c. 3; + obtuse, margins ciliate; thick; adaxially slightly concave near the base and with a rounded, glabrous ridge that is bifurcate proximally and runs up to c. 1/3 of the length of the lip, with 2 more distinct, rounded, papillose ridges diverging proximally and running up to c. 2/3 of the length of the lip, leaving a thin furrow in between them; abaxially without a ridge, surface glabrous in the proximal third, with long hairs elsewhere but with short hairs close to the tip. Column c. 0.9 mm long, stigma with a distinct, conical, obtuse tooth at its base, column foot without teeth. Stelidia porrect, triangular, c. 0.4 mm long, subacute, with a distinct, antrorse, triangular, subacute tooth along the lower margin. Anther abaxially with a distinct crest, surface + glabrous, front margin drawn out into a triangular, acute beak with entire margins. Pollinia 4, the inner slightly more than half as long as the outer, narrowly elliptic, the outer obovoid; no appendages present. Colours: Sepals and petals yellowish green, but lateral sepals blackish purple in the distal half except for the yellowish margins. Lip blackish purple. Habitat: Understorey epiphyte on tree trunks and branches. Alt. 1400-1500 m. Distribution: INDONESIA: Sumatra, Barisan Range (1 specimen seen). Notes: Most similar to Bulbophyllum camptochilum J.J. Verm., from Brunei. This species, however, has glabrous sepals, much narrower petals (index 2.4—2.5), and a wider lip (index 1.3—1.4) that is subrectangular in general outline. Figure 7. Bulbophyllum iterans J.J. Verm. & P. O’Byrne — a. Habit; b. top inflorescence; c. Flower; d. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; e. Lip, left: adaxial side, right: abaxial side; f. Column and lip, lateral view; g. Anther, left: abaxial side, right: adaxial side; h. Pollinia, above: single pair, below: two pairs. — All from SBG-O 321] (spirit sample). Southeast Asian Bulbophyllum 139 ~ (yet « ys) . A Pah TA ae ~% . Se * ae Pryor P's... I ore Cos Sees Cp fA ety SARE “ 4¢. AP Vl WIGAN MY TY WS Yy \\ om Figure 7. Bulbophyllum iterans J.J. Verm. & P. O’Byrne 140 Gard. Bull. Singapore 55 (2003) Bulbophyllum iterans J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 7. Bulbophyllum iterans J.J. Verm. & P. O’ Byrne, a Bulbophyllo errato Ames floribus c. 11 mm longis, petalis adaxialiter distincte pubescentibus, labello longo gracili differt. - TYPE: Indonesia, Sulawesi, central part, SBG-O 32/1] (SING, holo.). Roots below the pseudobulbs. Rhizome creeping, 0.9—1.4 mm diam., sections between pseudobulbs 0.4—0.9 cm long, bracts not persistent. Pseudobulbs close or distant, ovoid, 0.4—0.9 x 0.3-0.8 cm. Petiole up to 0.06 cm long. Leaf blade elliptic to ovate, 0.8—1.5 x 0.5—0.8 cm, index (length/width) 1.6—2; obtuse to acute. Inflorescence a congested raceme, 3.54.3 cm long, 1—7-flowered. Peduncle erect to patent, 2.3-2.8 cm, bracts 2—3, the longest 1.8—2.5 mm long. Rhachis slightly nodding, spindle-shaped, up to 0.3 x 0.1 cm. Floral bracts triangular, 2.2—3 x 2-3 mm, acuminate. Flowers not fully opening, each one developing only after anthesis of the previous. Pedicel and ovary c. 2 mm long, basal node on a 1.5—2 mm long stump. Median sepal slightly recurved, elliptic, 10.5—11.2 x 2.1 mm, index 5—5.4; acute, margins ciliolate, base broadly attached; thin, 3—5-veined, glabrous. Lateral sepals subtriangular, 11-12.2 x 2.5—2.7 mm, index 4.44.6; otherwise as the median sepal. Petals porrect, ovate, 7.6—7.9 x c. 1.9mm, index 44.2; acute, margins ciliate, base narrowly attached; thin, 3-veined, adaxial surface with hairs except near the base. Lip slightly recurved near the base, general outline ovate, slightly constricted near the base where the margins approach each other over the adaxial surface, c. 5.8 x 1.2 mm, index 4.8-4.9; truncate, margins ciliate; rather thick; adaxially concave near the base and with a distinct, narrow, rounded, ridge running from the constricted part of the lip, near the base, almost to the tip, adaxial surface glabrous; abaxially without a ridge, surface with stiff hairs near the base, and with softer hairs up to + half way the length of the lip. Column c. 1.8 mm long, stigma without a tooth at its base, column foot without teeth. Stelidia porrect, triangular, c. 1 mm long, acute, with minute, antrorse, triangular, obtuse tooth along the upper margin, close to the tip. Anther abaxially with a distinct, papillose crest, surface otherwise finely papillose, front margin drawn out into an incurved, truncate beak with entire margins. Pollinia 4, the inner c. 3/4 as long as the outer, ovoid, the outer ellipsoid; no appendages present. Colours: Flowers entirely dark purple. Habitat: Epiphyte in montane forest, on the trunk and in the crown of small trees. Alt. 1800—2500 m. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: While this species fits very well in sect. Hybochilus on account of the floral characters it has a racemose inflorescence, a character that is very rare in that Figure 8. Bulbophyllum anakbaruppui J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: single pair, right: two pairs. — All from SBG-O 3287 (spirit sample). 14] Southeast Asian Bulbophyllum Figure 8. Bulbophyllum anakbaruppui J.J. Verm. & P. O’Byrne 142 Gard. Bull. Singapore 55 (2003) section and the assumedly related sect. Polyblepharon. Only two other species in these sections display this character: Bulbophyllum erratum Ames (sect. Hybochilus, from the Philippines) and B. vexillarium Rid. (sect. Polyblepharon, from Papua New Guinea). B. iterans differs from both in having petals that are distinctly hairy on the adaxial side, and the long, slender lip. The name refers to the flowers that appear in succession on the inflorescence. Bulbophyllum sect. Leptopus Bulbophyllum spissum J.J. Verm. Bulbophyllum spissum J.J. Verm., Blumea 41 (1996): 360. — TYPE: Indonesia, Lingga Archipelago, Pulau Riau, O’Byrne LING 010 (L, holo.) Habitat: Epiphyte in forest. Alt. 200-1000 m. Distribution: MALAYSIA: Pahang, Pulau Tioman. INDONESIA: Lingga Archipelago, Pulau Riau. Notes: The Tioman plants represent the first record of this species for Malaysia. Material seen: MALAYSIA. Pahang, Pulau Tioman, SBG-O 737 (SING), 739 (SING). Bulbophyllum sect. Monilibulbus Bulbophyllum anisopterum J.J. Verm. & P. O’Byrne, sp. nov. Bulbophyllum anisopterum J.J. Verm. & P. O’Byrne, a Bulbophyllo stormii sepalo medio sepalis lateralibus 1/4-plo longiore differt. - TYPE: Malaysia, Sabah, Lamb 563/86 (K, holo.). Bulbophyllum stormii sensu J.J. Vermeulen, Orch. Borneo 2 (1991) 227 (pro parte, plate included), non J.J. Smith. Roots below the pseudobulbs. Rhizome creeping, c. 1 mm diam., sections between pseudobulbs c. 0.5 cm long, bracts not persistent. Pseudobulbs ovoid, basal half prostrate on and fused to the rhizome so that the new pseudobulbs arise c. half-way up the old, c. 0.9 x 0.4—0.5 cm. Petiole 0.5—0.8 mm long. Leaf blade elliptic, 1.8—2.5 x c. 0.5 cm, index (length/width) 3.5—5; obtuse. Inflorescence c. 6 cm long, 1-flowered. Peduncle erect to patent, 3.5—-4 cm, bracts c. 3, the longest c. 2.5 mm long. Floral bracts tubular, c. 2.5 mm, acute. Flowers not fully opening. Pedicel and ovary c. 18 mm long, basal node on ac. 2 mm long stump. Median sepal porrect, ovate, c. 9.5 x 2.8 mm, index c. 3.4; acuminate, margins entire, base rather widely attached; thin, Southeast Asian Bulbophyllum 143 glabrous. Lateral sepals recurved, free, oblique, c. 32 x 6 mm, index 5.3—5.4; caudate; otherwise as the median sepal. Petals porrect, ovate, c. 2 x 0.8 mm, index c. 2.5; obtuse, margins entire, base widely attached; thin, glabrous. Lip slightly recurved c. half-way, general outline ovate with a drawn-out top, c. 4.5 x 1.5 mm, index c. 3; rounded, margins entire; rather thick, glabrous; adaxially concave towards the base, convex and with the margins strongly recurved elsewhere; abaxially with a short, retuse ridge near the base. Column c. 2 mm long, stigma with a slight callus at its base, column foot without teeth. Ste/idia subulate, c. 1.5 mm long, acute. Anther abaxially with a distinct, rounded, papillose crest, surface + glabrous, front margin not drawn out. Pollinia 4, the inner less than half as long as the outer, all about ovoid, without appendages. Colours: Lateral sepals yellow, orange near the base. Petals translucent orange. Lip purple red. Habitat: Understorey epiphyte in wet montane forest, at 1200-1900 m alt. Flowering observed in Jan, and Jun—Jul. Distribution: MALAYSIA. Sabah, Ulu Padas (2 specimens seen). Notes: In Vermeulen (1991) I thought this was an aberrant form of the widespread and variable Bulbophyllum stormii J.J. Sm. Since then, I have found it again several times, and it is now clear that it is a distinct species. It differs from B. stormii in having a much shorter median sepal (about 1/3 to 1/4 of the length of the lateral sepals. B. anisopterum is most similar to B. hodgsonii Henders., from Peninsular Malaysia, but differs in having a much shorter median sepal, and caudate lateral sepals. Bulbophyllum sect. Osyricera (= sect. Globiceps) Bulbophyllum anakbaruppui J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 8. Bulbophyllum anakbaruppui J.J. Verm. & P. O’ Byrne, a Bulbophyllo hastifero Schltr. pseudobulbis quadrangulatis, petalis latioribus, labello ovato differt.— TYPE: Indonesia, Sulawesi, central part, SBG-O 3287 (SING, holo.). Roots creeping along the rhizome towards its point of attachment, then spreading over the substrate. Rhizome + pendulous with + patent branches, up to 80 cm long, 2.5—4 mm diam., sections between pseudobulbs 3.8—7.4 cm long, bracts moderately persistent. Pseudobulbs distant, ovate, 2—3.4 x 0.9-1.3 cm, distinctly and obtusely 4-angled. Petiole 0.2-0.3 cm long. Leaf blade ovate, 8.5—12 x 2.1—3.1 cm, index (length/width) 2.8—5.3; acute. Inflorescence porrect, a rather dense raceme, 3.8—6.2 cm long, 13—17-flowered. Peduncle 2—2.5 cm, bracts c. 4, the longest c. 6.5 mm long. Rhachis, spindle-shaped, 2.2—3.2 x 0.38—0.43 cm. Floral bracts cordate, c. 4 x 5 mm, acute. Flowers not resupinate, arranged in regular spirals, not fully opening, many open simultaneously. Pedicel and ovary c. 2.2 mm long, basal node + flush with the surface of the rhachis. Median sepal + porrect, obovate, c. 4.3 x 3.3 mm, index c. 1.3; 144 Gard. Bull. Singapore 55 (2003) top cucullate, rounded, margins entire, ciliolate, base rather narrowly attached; rather thick, 3-veined, surface very finely papillose adaxially towards the tip. Lateral sepals adnate along the lower margins, oblique, ovate-triangular, c. 5.8 x 3.2 mm, index 1.8-1.9, top flat, subacute, margins slightly papillose, base broadly attached; thick, otherwise as the median sepal. Petals porrect, ovate, c. 3.4 x 1.7 mm, index c. 2; acute, margins entire, ciliolate, base narrowly attached; rather thin, 1-veined, surface very finely papillose. Lip slightly recurved + half-way, general outline ovate, c. 3 x 2.2 mm, index 1.3—1.4; rounded, margins entire; rather thick, glabrous; adaxially slightly concave near the base, with 2 inconspicuous, rounded ridges proximally somewhat diverging and running from c. 1/3 of the length of the lip up to near the tip; abaxially + without a ridge near the base. Column c. 2.2 mm long, stigma without a tooth at its base, column foot with 2 truncate wings near its base, and two more, retrorse, triangular, obtuse ones just above the ligament. Stelidia minute, abruptly bent downwards but with an antrorse. + acute tip, along the upper margin with a very large (far more conspicuous than the stelidia themselves), antrorse, wedge-shaped wing with a broadly rounded, erose top margin. Anther abaxially with a crest near its base, surface + glabrous towards the base, coarsely papillose towards the tip, front margin drawn out into a distinct, concave, triangular, rounded, coarsely papillose beak. Pollinia 4, the inner + as long as the outer, all ellipsoid-ovoid; no appendages present. Colours: Plant medium green. Flowers yellow, median sepal with thin red veins, petals with thin red veins and margins, lip slightly suffused with red proximally. Habitat: Epiphyte in a large tree in coffee and cocoa gardens near a village. Alt. c. 1800 m. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Suberect, patent or pendulous rhizomes occur only in two species of sect. Osyricera: Bulbophyllum salaccense Rchb.f., from West Malesia, and B. hastiferum Schltr, from Sulawesi. The latter is most similar to B. anakbaruppui, but differs in having well rounded, not angular pseudobulbs, much narrower petals and a hastate lip. In flower structure B. anakbaruppui resembles B. trigonobulbum J.J. Sm., from Sulawesi. That species, however, has a creeping or straggling rhizome, with many spreading roots. It also has narrower petals. Named after the children of the Toraja village Baruppu, who collected the plant from a large tree in the village gardens. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 9. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne, a Bulbophyllo osyricera Schltr. et Figure 9. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side, below: adaxial side; g. Pollinia, left: two pairs, right: single pair. — All from SBG-O 3215 (living plant, spirit sample). Southeast Asian Bulbophyllum 145 Figure 9. Bulbophyllum illecebrum J.J. Verm. & P. O’Byrne 146 Gard. Bull. Singapore 55 (2003) B. osyriceroidi J.J. Sm. sepalo mediali distincte concavo, petalis multo longioribus angustioribusque, labello parte apicali spathulato differt. - TYPE: Indonesia, Sulawesi, Central part, SBG-O 3215 (SING, holo.). Roots mainly below the pseudobulbs. Rhizome creeping, 1.8—2 mm diam., sections between pseudobulbs 0.8—2 cm long, bracts barely persistent. Pseudobulbs close together, ovate, 1—-3.5 x 0.7—1.1 cm, not angular. Petiole 0.7—2.8 cm long. Leaf blade obovate-elliptic, 5.5-12 x 0.8-1.3 cm, index (length/width) 4.2—12.8; acute. Inflorescence a rather dense raceme, c. 30 cm long, c. 24-flowered. Peduncle erect to patent, c. 21 cm, bracts c. 5, the longest c. 9 mm long. Rhachis nodding at the base, limply pendulous, not thickened, c. 9 cm long. Floral bracts + triangular, c. 3 x 3 mm, acute. F/owers not resupinate, not fully opening, many open simultaneously. Pedicel and ovary c. 3.2 mm long, basal node + flush with the surface of the rhachis. Median sepal somewhat recurved, elliptic-obovate, c. 7 x 3.5 mm, index c. 2; top cucullate, rounded, margins entire, papillose-ciliolate distally, base narrowly attached; very thin, 3-veined, surface glabrous. Lateral sepals adnate along the lower margins, recurved, top part slightly incurved, oblique, ovate-triangular, c. 7.5 x 2.1 mm, index 3.5—3.6, top flat, (sub-)acute, base broadly attached; thin, otherwise as the median sepal. Petals recurved to spreading, ovate, c. 7 x 0.9 mm, index 12—13; acute, margins entire, slightly erose near the tip, base narrowly attached; very thin, 1-veined, glabrous. Lip almost straight, general outline subspathulate with a slightly widened base, c. 6 x 2 mm, index c. 3, rounded, margins entire; proximally rather thick, distally very thick, almost circular in section, glabrous; adaxially concave near the base, with a transverse wall connecting the sides just above the ligament, top part distinctly convex; abaxially + without a ridge near the base, top part distinctly convex. Column c. 2 mm long, stigma without a tooth at its base, column foot with a distinct, transverse ridge near its base. Stelidia minute, abruptly bent downwards but with an antrorse, + acute tip, along the upper margin with a very large (far more conspicuous than the stelidia themselves), antrorse + rectangular wing with a truncate, slightly erose top margin. Anther abaxially with a crest near its base, surface + glabrous towards the base, papillose towards the tip, front margin drawn out into a distinct, concave, triangular, obtuse, papillose beak. Pollinia 4, the inner slightly shorter than the outer, ovoid; the outer ellipsoid; no appendages present. Colours: Sepals and petals translucent with thin, red veins. Lip greenish yellow, margins red distally. Habitat: Epiphyte in fairly mossy montane forest, on the trunk and in the crown of small trees. Alt. 2300 m. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: Bulbophyllum osyricera Schltr., from Java and Sumatra, and B. osyriceroides J.J. Sm., from Sumatra, are most similar to B. illecebrum. Both differ, however, in having a more or less flat median sepal, much shorter (less than half as Southeast Asian Bulbophyllum 147 long as the median sepal) and wider petals, as well as a lip with an ovate, rather than a spathulate top part. The diaphanous, widely spreading, antenna-like petals and the highly mobile, very thick and brightly coloured lip suggest a baiting of pollinators by simulating either prey or a sexual partner. The name refers to this. Bulbophyllum sect. Polyblepharon Bulbophyllum uncinatum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 10. Bulbophyllum uncinatum J.J. Verm. & P. O’Byrne, a Bulbophyllo bisepalo et B. stenochilo sepalis deorsum unciformibus, labello proximaliter gradatim dilatato differt. — TYPE: Indonesia, Sulawesi, SBG-O 3835 (SING, holo.). Roots all close to the base of the rhizome. Rhizome patent to hanging down perpendicularly, up to 25 cm long, c. | mm diam., bracts not persistent. Pseudobulbs ovoid-cylindrical, prostrate on and fused to the rhizome, so that the new pseudobulbs arise from near the top of the old, 0.9-1.3 x 0.2—0.3 cm. Petiole up to 0.5 mm long. Leaf blade ovate, 2—2.7 x 0.8—1.1 cm, index (length/width) 2.42.5; acute. Inflorescence c. 0.6 cm long, 1-flowered. Peduncle porrect, c. 0.1 cm, bracts 1, c. 2 mm long. Floral bracts tubular, c. 1.7 mm, acute. Flowers not fully opening. Pedicel and ovary c. 1.3 mm long, basal node coinciding with the attachment of the floral bract. Median sepal incurved at the top, ovate, c. 3.8 x 1.3 mm, index 2.9-3; subacute, margins entire, ciliate towards the tip, base widely attached; thin, surface glabrous. Lateral sepals recurved, fused along the lower margin, oblique, c. 3.3 x 1.2 mm, index 2.7—2.8; obtuse; margins sparsely ciliate; otherwise as the median sepal. Petals porrect, ovate, c. 1 x 0.7 mm, index 1.4—1.5; acuminate, margins entire, base widely attached; thin, glabrous. Lip recurved at about 1/3 of its length, general outline ovate with a slightly narrowed top, c. 2.3 x 0.6 mm, index 3.8—3.9; acute, margins ciliate towards the base, ciliolate-papillose towards the tip; thin; adaxially slightly concave towards the base and with a median ridge starting near the base and running over about 1/4 of the length of the lip, surface convex and with the margins recurved towards the tip, entire surface finely papillose; abaxially without a ridge near the base, glabrous. Column c. 0.9 mm long, stigma without basal tooth, column foot distinctly thickened and with a retrorse, conical tooth near the base. Stelidia triangular, c. 0.7 mm long, acute, with a slightly antrorse, obliquely deltoid, obtuse tooth along the lower margin. Anther abaxially with a distinct, rounded, papillose crest, surface + glabrous, front margin drawn out into a truncate beak with glabrous margins. Pollinia: 2, ovoid. Colours: Sepals and petals yellowish near the base, purple towards the tip, abaxially with minute, blackish spots. Petals translucent white. Lip purple. Habitat: Found in coppiced roadside trees, at 500 m alt. Flowering observed in 148 Gard. Bull. Singapore 55 (2003) Jan—Mar and Dec. Distribution: INDONESIA. Sulawesi: central part (1 specimen seen). Notes: Similar to Bulbophyllum stenochilum Schltr., from Papua New Guinea. B. uncinatum differs in having only a low median ridge on the adaxial side of the lip, without a retrorse tooth near the base of the lip. Bulbophyllum sect. Sestochilus Bulbophyllum catillus J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 11. Bulbophyllum catillus J.J. Verm. & P. O’Byrne, a Bulbophyllo lyriforme et B. cheiri columella lateraliter aspectu aliquot sigmatoide curvato differt. - TYPE: Papua New Guinea, Jongejan, P. cult. 614 (L, holo). Roots along the entire rhizome. Rhizome creeping, 4-5 mm diam., sections between pseudobulbs 4—6 cm long, bracts with veins persistent as coarse fibres. Pseudobulbs distant, ovoid, suberect, 1.5—2.0 x 0.8—1.2 cm. Petiole 1.8—2.5 cm long. Leaf blade elliptic, 11-13.5 x 5.5—6.5 cm, index (length/width) 1.8—2.3; rounded. Inflorescence c. 7 cm long, 1-flowered. Peduncle (sub-)erect, 1.2 cm, bracts c. 3, the longest c. 12 mm long. Floral bracts tubular, c. 14 mm, cuspidate. Flowers not resupinate, not fully opening. Pedicel and ovary 53—78 mm long, basal node on a 8—10 mm long stump, juga and valvae well-rounded. Median sepal somewhat incurved, elliptic to obovate, 21—31 x 8—11.5 mm, index 2.2—2.9; acute to acuminate, margins entire, base rather widely attached; rather thick, c. 1 1-veined, glabrous. Lateral sepals with the lower margins running more or less parallel, free, oblique, subtriangular, 20-28 x 9-16 mm, index 1.7—2.2, base widely attached; otherwise as the median sepal. Petals recurved to spreading, elliptic to obovate, 20—25 x 5.5—8 mm, index 3.2—3.8; (sub-)acute, margins entire, base rather narrowly attached; rather thin, c. 7-veined, glabrous. Lip recurved near the base, general outline ovate with a drawn- out top, 10-11 x 6-7 mm, index c. 1.4—1.9; obtuse, margins entire; top part thick, elsewhere thin, surface glabrous; adaxially widely concave towards the base; abaxially with a distinct, emarginate ridge over most of the length of the lip. Column 7-9 mm long, slightly sigmoid in profile, stigma without basal tooth, column foot without a tooth just above the ligament. Stelidia porrect, triangular, 1.5—2 mm long, obtuse, without teeth along the lower margin. Anther abaxially with a very distinct, conical crest, surface + glabrous, front margin drawn out into a rounded beak with erose margins. Pollinia 4, the inner as long as the outer, all sub-triangular, without appendages. Figure 10. Bulbophyllum uncinatum J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left: a pair, right: single. — All from SBG-O 3835 (spirit sample). 149 Southeast Asian Bulbophyllum Figure 10. Bulbophyllum uncinatum J.J. Verm. & P. O’ Byrne 150 Gard. Bull. Singapore 55 (2003) Colours: Unknown. Habitat: Unknown. Distribution: PAPUA NEW GUINEA (4 specimens seen). Notes: The slightly sigmoid profile of the column is diagnostic within the group of sect. Sestochilus with single, non-resupinate flowers. The lip widens abruptly towards the base into two thin wings, as in Bulbophyllum lyriforme, and B. cheiri. The top part, however, much shorter than in these species. Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 12. Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne, a Bulbophyllo microglosso in labelli latere adaxiali proximaliter pubescenti differt. -TY PE: Laos, SBG-O 3622 (SING, holo.). Roots mainly below the pseudobulbs. Rhizome creeping, 5-6 mm diam., sections between pseudobulbs 8—9 cm long, bracts with veins persistent as fibres. Pseudobulbs distant, ovoid to ellipsoid, erect, 44.5 x 2.3—2.8 cm. Petiole 44.5 cm long. Leaf blade elliptic, 9.5—19 x 4—5.7 cm, index (length/width) 2.3—3.4; obtuse. /nflorescence 9.5—11.5 cm long, 1-flowered. Peduncle erect to patent, 1.1—1.4 cm, bracts c. 2, the longest c. 11 mm long. Floral bracts tubular, 15—18 mm, cuspidate. F/owers resupinate, not fully opening. Pedicel and ovary 80-95 mm long, basal node on a 9-11 mm long stump, juga and valvae well-rounded. Median sepal somewhat incurved, ovate, c. 23 x 9 mm, index 2.5—2.6; acute, margins entire, base widely attached; rather thin, 8—9-veined, glabrous. Lateral sepals recurved, lower margin without a fold, free, oblique, ovate, c. 22 x 13 mm, index c. 1.7; acute-apiculate; otherwise as the median sepal. Petals recurved to almost spreading, ovate, c. 18 x 6 mm, index c. 3; acute, margins entire, base widely attached; rather thin, c. 5-veined, glabrous. Lip recurved near the base, general outline + ovate, c. 8 x 8 mm, index c. 1; subacute, margins entire; rather thin; adaxially concave towards the base, surface finely papillose- pubescent towards the base, glabrous elsewhere; abaxially with a distinct, emarginate ridge from base to tip, surface glabrous. Column c. 4 mm long, stigma without basal tooth, column foot without a tooth just above the ligament. Stelidia minute, porrect, deltoid, subacute, without teeth along the lower margin. Anther abaxially with a very inconspicuous crest, surface + glabrous, front margin drawn out into a deltoid beak with glabrous margins. Pollinia 4, the inner slightly shorter than the outer, all ovoid; a large, fleshy lump of tissue attached to the inner pair. Figure 11. Bulbophyllum catillus J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: abaxial side, below: adaxial side; g. Pollinia, left: a single pair, right: two pairs. — a. From LAE 77443 (herbarium material), with the flower reconstructed from Jongejan cult. 1216; b. From Jongejan cult. 1216; c—g. From Jongejan cult. 614 (all spirit samples). Southeast Asian Bulbophyllum tl Figure 11. Bulbophyllum catillus J.J. Verm. & P. O’Byrne 152 Gard. Bull. Singapore 55 (2003) Colours: Sepals and petals greenish with purple veins. Lip white, suffused with pinkish purple. Habitat: Unknown. Grows and flowers well in cultivation in lowland conditions as well as at 1000 m alt. Flowering observed in Jan—Mar and Dec. Distribution: LAOS: Locality unknown (1 specimen seen). Notes: Most similar to Bulbophyllum microglossum Ridl. which, however, has a glabrous lip. B. smitinandii differs in having two distinct patches of much longer hairs on the adaxial side of the lip, as well as a distinct central callus. Named after Bob and Chittima Cowen, Bangkok, Thailand, who successfully flowered this species in their nursery. Bulbophyllum incisilabrum J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 13. Bulbophyllum incisilabrum J.J. Verm. & P. O’Byrne, a Bulbophyllo rugoso Ridl. ovario cristis non-sinuosis, sepalo mediali petalisque rotundato differt. — TYPE: Indonesia, Sulawesi, central part, SBG-O 1879 (SING, holo.). Roots mainly below the pseudobulbs. Rhizome creeping, 4.5—6 mm diam., sections between pseudobulbs 0.9-1.6 cm long, bracts with veins persistent as fine fibres. Pseudobulbs close, ovoid-cylindrical, erect, 3.5—5.5 x 1.2—1.6 cm. Petiole 3-5.5 cm long. Leaf blade elliptic to obovate, 10—16.5 x 2.7-4.2 cm, index (length/ width) 3.2-4; obtuse. /nflorescence 11—12 cm long, 1-flowered. Peduncle erect to patent, 2.6—3 cm, bracts c. 3, the longest 14-18 mm long. Floral bracts tubular, 15—18 mm, cuspidate. Flowers resupinate, not fully opening. Pedicel and ovary 75—80 mm long, basal node coinciding with the floral bract, juga and valvae sharply crested, crests slightly curved towards the base of the flower. Median sepal somewhat recurved, ovate, c. 20 x 12 mm, index 1.6—1.7; rounded, margins entire, base narrowly attached; rather thick, 8—9-veined, glabrous. Lateral sepals + spreading, lower margin with a distinct fold slightly below half-way the sepal, free, oblique, ovate-triangular, c. 23 x 12 mm, index c. 1.9—2; subacute, base broadly attached; otherwise as the median sepal. Petals + recurved, ovate, c. 19 x 6 mm, index 3.1—3.2; rounded, margins entire, base broadly attached; rather thick, 8—9-veined, glabrous. Lip sharply recurved near the base and near the tip, general outline + ovate, c. 10.5 x 7.5 mm, index c. 1.4; rounded, margins entire; thick, surface very coarsely warty near the tip; adaxially concave near the base, with a median slit from base to tip, with 2 rather conspicuous, rounded ridges diverging proximally and running up to c. 2/3 of the length of the lip, surface glabrous; abaxially with a distinct, emarginate ridge from base to tip, surface glabrous but with a finely papillose patch proximally along the median line. Co/umn Figure 12. Bulbophyllum coweniorum J.J. Verm. & P. O’ Byme — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: abaxial side, right: adaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: two pairs, right: a single pair. — All from SBG-O 3622 (spirit sample). Southeast Asian Bulbophyllum 153 ~ooen & techy ah Bt Figure 12. Bulbophyllum coweniorum J.J. Verm. & P. O’Byrne 154 Gard. Bull. Singapore 55 (2003) c. 4mm long, stigma without basal tooth, column foot without a tooth just above the ligament. Stelidia minute, porrect, triangular, acute, along the lower margin below the base with a distinct, subtriangular, 3—3.5 mm long, obtuse tooth. Anther abaxially with a rather distinct crest, surface + glabrous, front margin drawn out into a concave, rounded beak with erose and papillose margins. Pollinia 4, the inner + as long as the outer, all ovoid; no appendages present. Colours: Sepals and petals yellow with red spots. Lip white, adaxially pink towards the margins. Habitat: Epiphyte in forest. Alt. 1100-1200 m. Flowering observed in Feb. Distribution: INDONESIA: Sulawesi, central part (1 specimen seen). Notes: This pretty species is most similar to Bulbophyllum rugosum Ridl. It differs in having the crests on the ovary only slightly curved distally (not distinctly sinuous), in having a rounded median sepal and petals, and in having widely spreading lateral sepals. Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne, sp. nov. — Fig. 14. Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne, a Bulbophylo cheiri loborum lateralium margine dorsali (proximali) concava labelli basi differt.— TYPE: Papua New Guinea, Jongejan, P. cult. 282 (L, holo). Vegetative parts not seen. Inflorescence c. 5 cm long, 1-flowered. Floral bracts tubular, c. 6 mm, cuspidate. Flowers not resupinate, not fully opening. Pedicel and ovary c. 40 mm long, basal node on a c. 5 mm long stump, juga and valvae well- rounded. Median sepal somewhat incurved, subtriangular, 23—31 x 6-8 mm, index 3.84.5; acute, margins entire, base rather widely attached; rather thick, c. 9-veined, glabrous. Lateral sepals recurved, free, oblique, 22—30 x 8-10 mm, index 2.5—3.4, base widely attached; otherwise as the median sepal. Petals spreading to reflexed, subtriangular, 21—28 x 2.24 mm, index 7—10.5; acute, margins entire, base widely attached; rather thin, c. 5-veined, glabrous. Lip slightly recurved near the base, general outline hastate-lyrate with a long drawn-out top part, margins slightly concave near the base of the lip, 17-19 x 7-8 mm, index c. 2.2—2.4; obtuse, margins entire; drawn out top part thick, elsewhere thin, surface glabrous; adaxially widely concave towards the base; abaxially with a distinct, emarginate ridge over c. 1/5—1/4 of the length of the lip. Column 6—8.5 mm long, stigma without basal tooth, column foot with a slight callus just above the ligament. Stelidia porrect, triangular, 1—1.8 mm long, obtuse, without teeth along the lower margin. Anther abaxially with a very distinct, flat crest, Figure 13. Bulbophyllum incisilabrum J.J. Verm. & P.O’ Byrne —a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, left: adaxial side, right: abaxial side; e. Column and lip, lateral view; f. Anther, left: adaxial side, right: abaxial side; g. Pollinia, left: single pair, right: two pairs. — All from SBG-O /879 (living plant, spirit sample). Southeast Asian Bulbophyllum 155 Figure 13. Bulbophyllum incisilabrum J.J. Verm. & P. O’ Byrne 156 Gard. Bull. Singapore 55 (2003) surface + glabrous, front margin drawn out into a widely rounded beak with entire margins. Pollinia 4, the inner as long as the outer, all about drop-shaped, without appendages. Colours: Unknown. Habitat: Unknown. Distribution: INDONESIA: Irian Jaya (4 specimens distributed from BO by J.J. Smith, presumably from Irian Jaya). PAPUA NEW GUINEA (1 specimen seen). Notes: Bulbophyllum lyriforme is most similar to B. cheiri Lindl.; it differs in having the lateral lobes at the base of the lip slightly bent backwards, so that their proximal margin is slightly concave. In B. cheiri this is invariably convex. Bulbophyllum sinapis J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 15. Bulbophyllum sinapis J.J. Verm. & P. O’Byrne, a Bulbophyllo macrantho in sepalis lateralibus latioribus (indice 2—3.5 versus 1.5—1.7) differt. - TYPE: Papua New Guinea, Jongejan, P. cult. 171 (L, holo). Bulbophyllum aff. hahlianum Schitr.: O’ Byrne, Lowland Orchids Papua New Guinea (1994) 456. Roots along the entire rhizome. Rhizome creeping, 4-5 mm diam., sections between pseudobulbs 8—12 cm long, bracts with veins persistent as coarse fibres. Pseudobulbs distant, ovoid-cylindrical, suberect, 1.5—2.0 x 0.5—0.8 cm. Petiole 24.5 cm long. Leaf blade elliptic to obovate, 12—19.5 x 3.3—7.4 cm, index (length/width) 1.95.4; obtuse. [Inflorescence 5—12 cm long, 1-flowered. Peduncle (sub-)erect, 0.6—2 cm, bracts 3—5, the longest 9-19 mm long. Floral bracts tubular, 10—16 mm, cuspidate. Flowers not resupinate, not fully opening. Pedice/ and ovary 65—88 mm long, basal node on a 13—16 mm long stump, juga and valvae well-rounded. Median sepal somewhat incurved, elliptic to ovate, 26-28 x 8-12 mm, index 2.1—3.5; acuminate, margins entire, base narrowly attached; rather thick, c. 11-veined, glabrous. Lateral sepals with the lower margins running more or less parallel, often touching at the tip, free, oblique, ovate-triangular, 16-22 x 10-13 mm, index 1.5—1.7, base widely attached; otherwise as the median sepal. Petals recurved to spreading, elliptic to obovate, 18-30 x 4.8-8 mm, index 2.7—5; acute to acuminate, margins entire, base narrowly attached; rather thin, c. 7-veined, glabrous. Lip recurved near the base, general outline almost hastate, 5—6.5 x 5—6 mm, index c. 0.8—1.2; obtuse, margins entire; thick, surface glabrous; adaxially hardly concave towards the base; abaxially with a distinct, emarginate ridge from base to tip. Column 4.5—6 mm long, stigma without basal Figure 14. Bulbophyllum lyriforme J.J. Verm. & P. O’ Byrne — a. Flower; b. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; c. Lip, above: adaxial side, below: abaxial side; d. Column and lip, lateral view; e. Anther, above: adaxial side, below: abaxial side; f. Pollinia, above: a single pair, below: two pairs. — All from Jongejan cult. 282 (spirit sample). Southeast Asian Bulbophyllum ‘57 ~ shee Figure 14. Bulbophyllum lyriforme J.J. Verm. & P. O’Byrne 158 Gard. Bull. Singapore 55 (2003) tooth,column foot without a tooth just above the ligament. Stelidia porrect, triangular, 1—1.5 mm long, obtuse, without teeth along the lower margin. Anther abaxially with a flat crest, surface + glabrous, front margin drawn out into a rectangular beak with erose margins. Pollinia 4, the inner as long as the outer, all sub-triangular, without appendages. Colours: Flowers yellow. Habitat: Epiphyte in lowland forest, coastal woodland. Alt. 0-100 m. Flowering observed in Nov. Distribution: PAPUA NEW GUINEA: New Britain (2 specimen seen). Notes: Most similar to Bulbophyllum macranthum Lind1.; differs in having shorter and wider lateral sepals (index 1.5—1.7; in B. macranthum 2-3.5). Bulbophyllum sinapis has uniformly mustard-yellow flowers (hence the name), differing conspicuously from the cream-coloured, heavily spotted flowers of B. macranthum. Bulbophyllum turpis J.J. Verm. & P. O’ Byrne, sp. nov. — Fig. 16. Bulbophyllum turpis J.J. Verm. & P. O’Byrne, a Bulbophyllo wrayi Hook. f. sepalo mediali acuto, columna basi dente clitoriformi differt.— TYPE: Malaysia, Pahang, Gunung Brinchang, SBG-O 060 (SING, holo.). Roots mainly below the pseudobulbs. Rhizome creeping, 7-8 mm diam., sections between pseudobulbs 2.8—3.5 cm long, bracts with veins persistent as fine fibres. Pseudobulbs close or distant, ovoid, erect, 4-5 x 2.2—3 cm. Petiole 16—18 cm long. Leaf blade elliptic, 28-29 x 4.8-—5 cm, index (length/width) 5.6—5.9; acute. Inflorescence a rather dense raceme, 40-47 cm long, 40—50-flowered. Peduncle erect to patent, 23—28 cm, bracts c. 7, the longest c. 28 mm long. Rhachis arching, 17-19 cm. Floral bracts triangular, 4.5—15 x 2—2.5 mm, subacute. Flowers + resupinate, + secund, not fully opening, many open simultaneously. Pedicel and ovary 5—11 mm long, basal node + flush with the surface of the rhachis. Median sepal + porrect, ovate, c. 9.2 x 4.8 mm, index 1.9—2; acute, margins entire, base rather widely attached; rather thick, 3-veined, glabrous. Lateral sepals somewhat recurved, oblique, triangular, c. 9 x 5 mm, index c. 1.8, base rather narrowly attached; otherwise as the median sepal. Petals porrect, subtriangular, c. 4 x 2 mm, index c. 2; obtuse, margins entire, base broadly attached; thin, 1-veined, glabrous. Lip reflexed at c. 3/5 of its length, general outline ovate with an oblong, drawn-out top part, c. 3 x 2 mm, index c. 1.5 (all without artificial spreading), margins entire; glabrous, + 3-lobed; midlobe oblong, truncate-emarginate, rather thick; adaxially concave and with 2 knob-shaped ridges Figure 15. Bulbophyllum sinapis J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, above: adaxial side, below: abaxial side; g. Pollinia, left: two pairs, right: a single pairs. — All from Jongejan cult. 171 (spirit sample). Southeast Asian Bulbophyllum 159 Figure 15. Bulbophyllum sinapis J.J. Verm. & P. O’Byrne 160 Gard. Bull. Singapore 55 (2003) touching over a median furrow near the base, with 2 more rather inconspicuous, obtuse ridges proximally somewhat diverging and running parallel from about 1/3 to 2/3 of the length of the lip, the space in between channelled; abaxially with an inconspicuous, retuse ridge up to about 1/3 of the length of the lip; /ateral lobes erect, attached along the proximal third of the lip, semi-circular, c. 1 mm high, rounded, thin. Column c. 2.8 mm long, stigma with a transverse ridge at its base, a longitudinal median ridge issuing from this, continuing up to + half-way the column foot, then splitting into two spreading lobes ending in a retuse tooth just above the ligament. Stelidia porrect, triangular, c. 1.2 mm long, obtuse to subacute, with a deltoid, acute tooth along the upper margin and a similar, obtuse tooth along the lower. Anther abaxially with a finely papillose crest, surface otherwise + glabrous, front margin drawn out into a porrect, triangular, acuminate beak with papillose margins. Pollinia 4, the inner slightly shorter than the outer, narrowly drop-shaped, the outer ovoid; no appendages present. Colours: Rhizome scales densely spotted with dark red. Sepals yellow, stained with red except near the base. Petals yellow. Lip yellowish green, top part yellow. Flowers strongly scented, reminiscent of overripe apples. Habitat: An understorey epiphyte in thick moss cushions in permanently wet montane forest. Bulbophyllum tahanense Carr and B. evansii Henders., like similar conditions and grow in the same area. Alt. 1600-2000 m. Flowering observed in Mar. Distribution: MALAYSIA: Peninsula (1 specimen seen). Notes: This species differs from Bulbophyllum wrayi Hook.f. in having acute leaf tips, as well as a distinct bifurcate tooth on the column foot. Bulbophyllum Incertae Sedis Bulbophyllum plumatum Ames Bulbophyllum plumatum Ames, Orch. 5 (1915) 184.— TYPE: Philippines, Mindanao, Bukidnon Subprovince, BSC (L. Escritor) 21479 (not seen). Bulbophyllum jacobsonii J.J. Sm., Blumea 1 (1935) 320.—TYPE: Indonesia, Sumatra, West Coast, Soengai Dareh, Jacobson E.R. cult. 2202 (Theunissen) (not seen). Habitat: In Malaysia found as an epiphyte in lowland swamp forest; in Sumatra found at 1000 m altitude. Distribution: MALAYSIA: Peninsula, Johor. INDONESIA: Sumatra. PHILIPPINES: Mindanao. Figure 16. Bulbophyllum turpis J.J. Verm. & P. O’ Byrne — a. Habit; b. Flower; c. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; d. Lip, above: adaxial side, below: abaxial side; e. Column and lip, lateral view; f. Anther, left: abaxial side, right: adaxial side; g. Pollinia, left: two pairs, right: single pair. — All from SBG-O 0060 (living plant, spirit sample). Southeast Asian Bulbophyllum 161 Simm x ny 7 Ms ie y: *: . i is ee 8, s i R: . ii Sy: fs a Te ees ce a eM Y _—-=—. -~ me Figure 16. Bulbophyllum turpis J.J. Verm. & P. O’Byrne 162 Gard. Bull. Singapore 55 (2003) Notes: The Johor plants represent the first records of this species for Malaysia and are the first fully documented records of this species in about 90 years. Plants of Bulbophyllum plumatum probably originating from Indonesia appeared in trade in some quantity a few years ago. The Johor plants differ slightly from these in having the lateral sepals coarsely transversely wrinkled. Material seen: MALAYSIA. Johor, Endau basin, km 56 on road to Mersing, SBG-O 2533 (SING), Johor, Rompin area, Hon Quik Pin (WOC 17,2002) 01 (SING). Garden’s Bulletin Singapore 55 (2003) 163-164 Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum S. P. VANDER KLOET Department of Biology, Acadia University, Wolfville, Nova Scotia, Canada B4P 2R6 In Vander Kloet (2002), I overlooked the fact that Sleumer had in 1941 already made combinations for the taxa in question under Vaccinium. Therefore the names I proposed are later homonyms and are illegitimate. The correct names for the taxa in question should be: Vaccinium ridleyi Sleumer, Bot. Jahrb. 71 (1941) 424. Agapetes parviflora Ridl., J. As. Soc. Str. Br. 61 (1912) 26 non Dunn (1903), nec Vaccinium parviflorum Andr. (1800); Agapetes micrantha Ridl., Fl. Mal. Pen. 2 (1923) 205, non Vaccinium micranthum Dunal 1839; Vaccinium micranthum Vander Kloet 54 (2002) 181 non Dunal (1839). Type: Ridley 15768 (holo SING!). Synonym: Vaccinium pseudodialypetalum Ng, Gard. Bull. Sing. 28 (1976) 231. Type: KEP 56673 (holo KEP). Vaccinium urophyllum Merr., Pap. Mich. Ac. Sc. 19 (1934) 184. Type: Yates 2427 (holo MICH; iso NY, US). Synonyms: Vaccinium longilingua Sleumer, Bot. Jahrb. 71 (1941) 424; Agapetes perakensis Ridl., Fl. Mal. Pen. 2 (1923) 205, non Vaccinium perakense Ridl. (1918); Vaccinium perakensis Vander Kloet, Gard. Bull. Sing. 54 (2002) 182 non. Ridl. (1918). Type: Ridley 5532 (SING!). Agapetes griffithii auct. non C.B.Clarke (1882): King, J. As. Soc. Beng. 74 (1906) II ak Acknowledgements I acknowledge the help and advice of J.F. Veldkamp (L) and I.M. Turner (SING) in the preparation of this erratum. 164 Gard. Bull. Singapore 55 (2003) Reference Kloet, S.P. Vander. 2002. Re-examination of Vaccinium dialypetalum (Ericaceae). Gardens Bulletin Singapore. 54: 179-183. Garden’s Bulletin Singapore 55 (2003) 165-171 OBITUARY Mohamad Shah bin Hj Mohamad Noor Mohamad Shah bin Hj Mohamad Noor, born in the staff quarters of the Singapore Botanic Gardens, was the son of Hj Mohamed Nur bin Mohamed Ghous. Like his father, he spent his working life in the herbarium exceeding his father’s record of 47 years’ service by one year. Born on 22™ January 1935, he received his education at the Anglo-Chinese School in Penang and the Tanjong Rhu Boys School in Singapore. After leaving school in 1954, his father encouraged him to follow in his footsteps and so Mat Shah began his career in the Herbarium in 1955 as a Plant Collector. When his father died in 1958, he took over his post as Herbarium and Museum Assistant. He rose to become a Research Officer. No sooner did he retire in 1993 than he was re-employed until his untimely death on 18" June 2003. Mat Shah’s father was exceptionally diligent (Burkill, 1958) and never lost the opportunity to collect plants. We even have a specimen of a banana he collected from the market in Mecca while he was on the haj! Mohamed Nur undertook the massive task of re-organising the herbarium to follow H.N. Ridley’s five-volume Flora of the Malay Peninsula. Many of the covers in use today still bear his handwriting. Besides his vast knowledge of plant names, he also amassed a vast collection of plants from all over Peninsular Malaysia. Mat Shah’s service in the Herbarium overlapped with his father’s for four years, which gave him an invaluable grounding both in plant identification and herbarium management. As a Plant Collector, Mat Shah accompanied the Director, J. W. Purseglove, on three expeditions to Sarawak in 1955, 1956 and 1957. The longest stint he spent in the field was when he joined the two Royal Society Expeditions to Gunung Kinabalu in 1961 and 1964 (see Appendix). Except for these expeditions, Mat Shah kept his own collecting books and numbering series and had collected more than 5,000 specimens by the end of his career. Not only did he collect in Sabah and Sarawak, he also collected extensively in the Peninsular Malaysia where he climbed many of the major mountains (Gunung Belumut, G. Bujang Melaka, G. Lawit, G. Ledang, G. Rabong, G. Tahan and G. Ulu Kali). He also collected continuously in Singapore, from time to time being involved in extensive surveys, such as that of plants in the Singapore Botanic Gardens in 1974 (96 numbers), the Seletar Reservoir area in 1971 (51 nos.), MacRitchie Reservoir area in 1976 (77 nos.) and the Pierce and Seletar Reservoir areas in 1981 (56 nos.). He has recounted some of his experiences in the field (Waterson, 2003): >My father taught me always to bring a mosquito net, because it can protect you against centipedes and scorpions, even snakes. One night on Gunung Tahan about nine or ten o’clock, I was lying under my net and I saw a very big cobra going up and over the 166 Gard. Bull. Singapore 55 (2003) top of the net. One of the porters got a parang but I told him not to kill it. Because the second thing my father taught me about the jungle was never to kill any animals. On Gunung Rabung, there were tigers. We met a very big one, but the kampung people assured us they were not man-eaters and never hurt anybody.” Mat Shah worked through the most turbulent period in the herbarium’s history as Singapore emerged from colonial status to independence and prosperity. There were more changes in Director in his 48 years than in the previous 80 years. The nadir was reached when rumours circulated worldwide that the Herbarium was no longer valued and was up for sale! He also experienced two major upheavals when the entire herbarium moved (Kiew, 2003). The first in 1963 was forced because the herbarium building was threatened with imminent collapse. The 400,000 specimens were moved inside the cupboards, the Gardens’ labourers providing the muscle to move the heavy, solid teak cupboards. The second was last year (2002) when the 650,000 specimens were moved out so that the new state-of-the-art Herbarium could be built on the same site. Sadly, Mat Shah will not be able to see the Herbarium settled into the new building in 2005. For the second move, Mat Shah worked steadily for more than six months with our team of ladies, Juriah Sabudin, Rohana Mohd Shari, Sariah Zelani and Sugaranthara Davi, to bring the family and generic concepts up-to-date and number the families according to D.J. Mabberley’s Plant Book and the genera and species alphabetically. At the end of six months each box was numbered with its family and sequence within the family. All the overflow of specimens (over 50,000) that had accumulated over the years that could not be fitted into the old herbarium cupboards was incorporated into the sequence. This systematic work paid off as it enabled the 10,000 boxes to be put straight onto the shelf in the correct order, all within 10 days. This re-ordering the Herbarium is a lasting legacy to Mat Shah’s dedicated work. Quiet, dedicated work characterised Mat Shah and his supervision of staff over the years was firm yet harmonious. He will be remembered by the many visitors to the Herbarium from all over the world for his courteous helpfulness. But it was most particularly for his extensive knowledge of plant names (Kiew, 2001) that made his service invaluable and led to his re-employment for eight years beyond retirement age. In 1993, he underwent a heart by-pass operation, which, although it curtailed his plant collecting activities, did nothing to dampen his dedication to herbarium work. He passed away peacefully in the early morning of 18" June 2003. He was due to finally retire in August 2003. His ability to identify plants and his knowledge of where plants grow in Singapore will be sadly missed. In the words of Dr S.C. Chin, Director: “With his passing, the tiny pool of specialist plant collectors and field botanists who can put a name to most things green, shrinks further.” He leaves his wife, Madam Jamilah binte Osman, whom he married in 1963, three sons and a daughter, and four grandsons and three granddaughters. 167 Obituary-Mohd Shah b.Hj Mohd Noor th 9. ) H.M. Burkill, November 19 Mohamed Shah in the old Herbarium. 168 Gard. Bull. Singapore 55 (2003) References Burkill, H.M. 1958. Obituaries. Haji Mohamed Nur bin Mohamed Ghous, B.E.M. Gardens Bulletin Singapore. 17: 331-336. Kiew, R. 2001. What is this plant? The Herbarium’s role in plant identification. Gardenwise. 17: 3-4. Kiew, R. 2003. The Herbarium moves ... again! Gardenwise. 20: 13-15. Waterston, R. 2003. Across the generations: Plant collectors and their careers at the Botanic Gardens. Gardenwise. 21: 12, 14. Ruth Kiew Singapore Botanic Gardens Singapore Appendix: Mohamad Shah’s Collecting Expeditions Outside Singapore (Bk Bukit, FR Forest Reserve, G Gunung, NP National Park, Sg Sungai) 1955 12-27 Sept. Sarawak — Kuching, Bk Kitang, Bau; Setapok, Sebanding; G Gading, Lundu, Sematan, Kampung Pueh (with J.W. Purseglove, specimens numbered under Purseglove’s collecting series) 1956 16 May-20 June. Sarawak — Bako NP, Telok Asam, Telok Tajor, Lokosan, Sg Serai, Telok Wakang, Telok Delima, Telok Lakei, Telok Lomi, Bk Buan, Tau Range, Bk Mayeng, Bk Merseng, Tatau (with Purseglove & H.B. Gilliland) 1957 2-18 Feb. Sarawak — Bako NP (with Purseglove) 6-9 May. Malacca — Fish Culture Research Station, Batu Berendam 18-22 Sept. Negeri Sembilan — Jendaram Estate, Nilai 1958 26 Jan—3 Feb. Pahang — Sabai Estate, Bentong 5-14 March. Perak — Bota Kiri FR, Ipoh (with Kadim bin Tassim) 2-9 April. Johore — Bk Paloh Estate (with Kadim) Obituary-Mohd Shah b.Hj Mohd Noor 169 1959 1 Feb. Johore — Kota Tinggi Waterfall (with H.M. Burkill) 28 Feb-8 March. Kelantan —- Kampung Gobek, Tamangan (with Kadim) 24 Aug. Selangor/Pahang — Fraser’s Hill (with Burkill and Mohd Noor bin Jamaat) 21-28 Sept. Johore — Kuala Sedili, Teluk Jereh, Tanjung Lembu, Sg Sedili Besar (with Mohd Noor and on 26" with Chang Kiaw Lan) 1960 15-19 Aug. Pahang — Fraser’s Hill (with Mohd Noor) 1961 18-26 May. Sabah — Pulau Gaya, Tanjung Aru, Bk Kepayang (with Kadim) 29May-16 Aug. Sabah — G Kinabalu (1* Royal Society Expedition, with Chew W.- L, specimens numbered under the RSNB series) 1964 10 Jan-20 May. Sabah — G Kinabalu (2" Royal Society Expedition, with Chew) 1965 3-9 Dec. Perak — Maxwell Hill, Bk Berapit, Batu Hampar, Bk Kemunting (with Sidek bin Kiah) 1968 16 Feb-7 March. Pahang — Taman Negara, Sg Tahan, Sg Teku, G Tuan Skeat, Kuala Keniyan, Gua Luas, Sg Redap, Sg Belar (with herbarium of Forest Research Institute Malaysia, KEP) 5 April. Johore — Mersing 1 Sept. Johore — Sg Pencharang, Bekok 1970 31 May. Johore — Sg Bantang, Labis F.R., Bekok (with E.A. Heaslett & Samsuri) 9-22 July. Pahang — Taman Negara, Ulu Sg Sat, Kuala Kelepah, Ulu Sg Sepia, Kuala Aur, Ulu Sg Tembeling, Kuala Tahan (with KEP & Mohd Noor) 18-19 Aug. Johore — Bk Tangga Tujoh, Lak Heng Eststate, Kota Tinggi (with R. Hill, Mohd Noor & Ahmad) 20-25 Sept. Johore — Sg Kahang, G Belumut (with KEP & Sanusi) 1971 29 Jan-4 Feb. Johore — Bk Pengantin, Sg Juasseh, Labis (with Ahmad) 22 April. Johore — Kota Tinggi Waterfall 5 Sept. Johore — Ulu Sg Segamat, Labis (with Samsuri Ahmad) 19 Sept. Johore — Padang Mulut, SE coast (with Ahmad) 170 Gard. Bull. Singapore 55 (2003) 1972 11-17 March. Kelantan — G Rabong, Ulu Sg Ketil (with KEP) 30 July. Johore — Sg Endau, G Janing (with Ahmad) 12-18 Aug. Pahang — Taman Negara, Bk Tersek, Kuala Trengan, Jeram Katak, Sg. Keniyan, Genut Tabing (with Ahmad, on 18" with Miss Lee Wai Chin) 30 Aug-2 Sept. Pahang — Fraser’s Hill (with Miss Geh Siew Yin) 23 Nov. Johore — G. Lesong (with Ahmad) 1973 29 Jan-4 Feb. Kelantan — Gua Musang, Bk Brangkat, Bertam, G Brong, Kampong Sta (with Mohd Ali) 11 Mar. Johore — G. Panti East (with Heaslett & Samsuri) 10-15 June. Selangor — Ulu Gombak, G. Ulu Kali, Anak Takun, G Bunga Buah (with Mohd Ali) 23-30 Nov. Johore — G Lesong, Sg Rompin (with Ahmad) 1974 22-25 Feb. Kelantan — Bk Baka, Machang (with Ahmad) 27 Feb-1 March. Kelantan — Bk Yong, Pasir Puteh (with Ahmad) 31 May-4 June. Trengganu —G Tebu, Jabi, Sg Chabang Dua, Sg Belantan (with Ahmad & Mahmud Awang) 1975 11-14 Feb. Perak — G Bujang Melaka, Sg Chenderiang (with Ahmad) 15 Feb. Pahang — Brinchang, Cameron Highlands (with Ahmad) 16-17 Feb. Perak — Sg Selung, Sg Hidup, Telok Anson (with Ahmad) 12-16 March. Trengganu — G Lawit, Kuala Sg Bok, Bk Yong (with British Museum Gunung Lawit Expedition, with Ahmad and Samsuri) 11-17 July. Johore — G Ledang; Ma’okil, Cha’ah; Sg Batu Badak (with Samsuri) 23-28 Oct. Pahang — Bk Berkelah, G Serudom, Kuantan (with Samsuri & Sidek) 1976 3-5 March. Kelantan — Bk Bongkok, G Ayam (with Samsuri) 8 March. Kelantan — Ulu Sg Sat. (with Samsuri) 1978 9-13 June. Perak — G Bubu, Keledang Saiong FR (with Mahmud Awang) 5 Oct. Trengganu — Rantau Abang (with Samsuri) 22-26 Nov. Trengganu — Abang area, Jambu Bongkok, Gong Balai, Tanjung Juara (with Sidek) 1981 15 Sept. Johore — Jason Bay, K. Sedili, Jemaluang Road (with Ali Rahim) Obituary-Mohd Shah b.Hj Mohd Noor 171 1984 18-21 July. Trengganu — Bukit Tangga (with Mahmud) 1989 10-15 Feb. Sabah - G Kinabalu (to summit) and Poring (with E.P. Tay, specimens numbered under Tay’s series) 20 Feb. Sabah — Nabawan (with E.P. Tay, specimens numbered under Tay’s series) 1991 23-26 July. Johore — Sg Linggui (with E.P. Tay and Ali Ibrahim, specimens numbered under Tay’s series) 1993 2-5 May. Kedah — Sg Badak, Changlun, Bk Perangin; Bk Enggang, Sik. ee ae! oe ee a | : 7 ae A i . ; a a ee - : i972 ‘~~ _ ‘ 11.27 March Kelantad ~—O Rebel vA" ihe vhs <— 7 * ain el atg Pi : 7 i July. Jomwe~ Sg I ange i i with Adenad ® Pal ner Pik ng ‘ _ ui ease Bh Tera . tT Opes ss nas sit — , dy ted ct’, anoney NNT SE add iv bart aie MAD <aataz | y * . » Hil ™ renee Aabete barr rete aw erderusn ‘ (Pied EVA x: pirth di oonduiiieaie “shal oe. rir) aswel = tah ind « i s] WOD Ts at oh » re otc? Bassok “eT rsbaw ieee has J in. By mgs Bu ne ee; Bank 48 30 148 nurigaedO " bet 92 — ah vats 2 b i > bh Aby tat) hee — _ , _ - *< fi > - id —T 1 a _ will Pu ma -) ; P 4 rn oy * pare -) bs A | ) with ; age % Eg). irene | Se Cheheng Dua;: ae ' | ; By, i, Ab uct ) - « a! 7 ~~. \ al : 4 Ma 7) (wih Rotah oie : ~ | - a _ : iwith Sani. be ' stl viel ae as . s — se _ : ; F im oa » eo 7 ‘ 5 Ti j re ol iweath 2 \ tahonud J ‘ ' eg, i} itciegen - aa. Ata { meri) iv Z ire } spha Bongitel : iu > Sa +9 1 ’ r.! i i | i I ay. KG Fr i. ™~ 5 7 a a) , ia | pa rar: Sorin ir 4 Sigipe 440) rv’ vi et allies) curteoc C 1% i p mbenittes! for gnihiicacion ¢sewtesee od iF oh Wha re Saniied, ¢ is ‘eed - af te toe, Tyee it porter , Ad Romar iype sll the ion Hine foe aad 1m [ AY ae prone try the P a ere andi this : ry paer ls wocepied, a diskrtin w, a @athors; Tha tiie ; Bib aie sition’) | | plea natare rus: bad ‘ A f SS 7 : ' r.¢ « | ’ | } ~< ‘ = Py i] : : j “ss ay}, etc, Wie . wat Aye Ale dertn . manic oe ie author sen | Che bomerin? ed 7? -_ C7 a4, Naha pid jams lg ‘ : =p v pen pao on a #0 to besuiks anc ie ag caplint, Alan rin nay | : : i 1190, The Diger Pe wt Theruedacaes 2 a”. clutions ia axoucrls sia vig tye ie r- 27 ” ." a, ifolia Bin i va Bok. Lape. Bat. sig is nimi Ms earn , = = ab Aes =i" Par ‘i J. al ¢ aa — ; a We cep . ust be. exceed sani, INSTRUCTIONS TO AUTHORS Manuscripts: The Gardens’ Bulletin publishes original findings and reviews of progress in the field of plant taxonomy, horticulture, and allied subjects. Contributions must be original and the material must not have been submitted for publication elsewhere. Two copies of the manuscript should be submitted, typed or type printed, and if typed, then the top copy must be one of the two. Type or print on one side only, with double-line spacing and a margin of at least 4 cm. Do not type all the letters of any word in capitals. Underline only in pencil: with a straight line for italic type face and wavy line for bold type face. Authors should see the layout of other papers recently published in this journal to ensure that papers submitted conform as closely as possible to the accepted pattern. 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Bot ingapore 55 (Part 2) December 2003 anic S VOL NW WN i it Mi Hh } i | | HI MA HHT Mi a i i Mi i Hi ii m HAH i) Wi i i HHI H| i] Mi i "a i i) ] ii} A i i if i AN 5 ' ri | mh a NIT i i 1 . aa HU Wi i i} HUE ii : WAN i} iI i Ty Hi) al Hi | : il i Ml i | il | ih | HN Wh il | HH UNH TILPTtBt ET: ihetil} ‘i — Ee ems i a a a a i“ a a L al il i The Gardens’ Bulletin Singapore VOL. 55 (Part 2) December 2003 ISSN 0374-7859 CONTENTS Kiew, R. and I.M. Turner Pale Amy NMI SMe MMIC DO SUID ADIOLS Foo oo cic seen onete cenananiscnsemsumenecceel 1D Hassan, I. A note on Cordia premnifolia Ridl. (Boraginaceae) .................000000+-185 Mabberley, D.J. New species of, and other notes on, Chisocheton and Walsura (Meliaceae) cia oe ack ame RT shes Ds sv:anviees'esinpsasenesdenereeaetvabes 189 Gusman, G. Arisaema fimbriatum (Araceae) and its intraspecific variation .............201 Middleton, D.J. A revision of Dyera (Apocynaceae: Rauvolfioideae) ................2 606+. 209 Skornickova, J., M. Sabu and M.G. Prasanthkumar Curcuma codonantha (Zingiberaceae) — a new species from the Andaman DART NINA wine tira lias rir ties He i Call aig bes Bie. bh aii ane . Poaddanerrsa ses Shs 219 Zahid, M.S. Three new species of Porterandia (Rubiaceae) from Mount Kinabalu, eae Oh bed pes eked Die piesa od rretee sarc. tiwlsh dues. Vawenshenedssiove cheaters 229 Chung, R.C.K., E. Soepadmo and A.L. Lim. The significance of pollen morphology in the taxonomy of Grewia and Microcos (Tiliaceae) in Peninsular Malaysia and Borneo .....................239 Vermeulen, J.J. and P. O’Byrne Six new species of Bulbophyllum (Orchidaceae) from Sulawesi ............257 Lee, S., A. Samsuri, P. Leong, Ali Ibrahim and A.T. Gwee A botanical survey of Chek Jawa, Pulau Ubin, Singapore ......................271 Obituary De Chane Kiaw Lan ...........:: 2 2e. sees oo. K.M Wong......309 Date of Publication: 15 December 2003 Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 259569 Printed by Print-Dynamics (S) Pte Ltd Gardens’ Bulletin Singapore 55 (2003) 173-184 Are Any Plants Endemic to Singapore? RUTH KIEW AND I. M. TURNER Singapore Botanic Gardens, | Cluny Road, Singapore 259569 Abstract Many taxa are reported in the literature as endemic to Singapore. Of the 19 investigated, seven are shown to be endemic. Of these six, Bolbitis xsingaporensis (Lomariopsidaceae), Flickingeria laciniosa (Orchidaceae), Spatholobus ridleyi (Leguminosae), Strychnos ridleyi (Loganiaceae), Tectaria griffithii var. singaporeana (Dryopteridaceae) and Thunbergia dasychlamys (Acanthaceae) are extinct, while Cryptocoryne xtimahensis (Araceae) is still alive in the Bukit Timah Nature Reserve. These seven taxa represent a mere 0.3% of Singapore’s vascular flora of about 2300 species. Anew combination is made for Hypobathrum coniferum (Ridl.) Kiew (Rubiaceae). Introduction In Ridley’s 1900 enumeration of Singapore seed plants, the first listing for Singapore, he noted 33 species as endemic to Singapore although he pointed out that most would probably be discovered in neighbouring countries and this proved to be the case. In addition, he indicated that some names he listed from the literature as coming from Singapore were either invalid names or that Singapore had been wrongly given as the place of origin. (The latter was not an unusual practice in the case of professional plant collectors who did not want the true origin of a valuable plant to be known). The preparation of the Flora of the Malay Peninsula, initiated at the Calcutta Botanic Garden, but eventually driven by the indefatigable Ridley, both while he was Director of the Botanic Gardens Singapore and later in retirement at the Royal Botanic Gardens, Kew, resulted in the description of many new species, some of which were known only from specimens collected from Singapore. The number of species named in honour of Ridley and the number of rare species that he collected (see Appendix) illustrate his diligence as a collector and that forest in Singapore was much more extensive and less disturbed in his day Since that period (1889-1925), a few new species apparently endemic to Singapore have been described often based on very old collections. Significant among these are two hybrids because hybrids often arise and survive in disturbed conditions so they may be a reflection of disturbance in the remaining forested areas in Singapore. 174 Gard. Bull. Singapore 55 (2003) Candidates for Endemic Species Among the species listed by Ridley (1900), one has proved to be endemic to Singapore. He described Dendrobium laciniosum, Orchidaceae (now known as Flickingeria laciniosa (Ridl.) A.D.Hawkes) from two plants collected in 1891 from Pulau Seletar. At one time, this species was confused with another orchid from Perlis, Peninsular Malaysia, but Seidenfaden & Wood (1992) believe the Perlis plant to be different. Incidentally, they erroneously record Pulau Seletar as in Johore, Peninsular Malaysia. Later, Ridley described three more species and a variety as endemic to Singapore: Didymocarpus perdita Ridl. (Gesneriaceae), Dischidia singaporensis Ridl. (Asclepiadaceae), Petunga conifera Ridl. (Rubiaceae) and Spatholobus ferrugineus (Zoll. & Moritzi) Benth. var. sericophyllus Ridl. (Leguminosae). Didymocarpus perdita collected from Seletar in 1889, Ridley (1923) described as ‘very rare, only 2 specimens seen in a ravine, now probably extinct’. This species has proved to be the same as D. punticulata Rid|. (now known as Henckelia punticulata (Ridl.) A. Weber), which grows in southern and eastern Peninsular Malaysia (Kiew, 1987). Dischida singaporensis is similarly a very rare species known only from one collection from the Changi Police Station in 1908. Later, D. singaporensis was collected from Endau-Rompin, Johore (Kiew, 1987). Petunga conifera, now known as Hypobathrum coniferum (Ridl.) Kiew (see Appendix), in Ridley’s time was known from a single tree in the Botanic Gardens Rain Forest, which at that time was called the Gardens Jungle (Ridley, 1923). Later, a specimen of this species was collected from Johore, disproving its status as endemic to Singapore. However, considering its extreme rarity and that is has not been recollected for more than a hundred years, it can be concluded that the Singapore population of this species is extinct. Spatholobus ferrugineus var. sericophyllus was originally described from four collections from Singapore (Ridley, 1922). However, this variety has since proved to be widespread (Ridder-Numan & Wiriadinata, 1985). In Peninsular Malaysia, for example, it has been collected from Perak, Selangor, Malacca and Negri Sembilan. From publications from Calcutta, five new species were described that were then known only from specimens collected in Singapore: Litsea ridleyi Gamble (Lauraceae), Piper flavimarginatum C.DC. (Piperaceae), Polyosma ridleyi King, Spatholobus ridleyi Prain ex King (Leguminosae) and Strychnos ridleyi King & Gamble (Loganiaceae). Litsea ridleyi, first collected in the 1880s, has been sporadically collected since, the latest being in 1982 from the Bukit Timah Nature Reserve. However, several specimens from Borneo (for example, Ridley s.n. 1892 from Bongaya and Senada Brunei 54121 from Baram) are conspecific with this species, which is therefore not Endemic Plants in Singapore i be, endemic to Singapore. Ridley (1924) reported the wild pepper, Piper flavimarginatum as only from Singapore, where he described it as ‘very common but rarely flowering’. However, it is widespread though not common in Peninsular Malaysia and has been collected from as far afield as Langkawi, Perak, Pahang, Trengganu, Negri Sembilan and Johore. It can therefore no longer be considered endemic to Singapore. It has not been recollected in Singapore since 1933. Another rare endemic, Polyosma ridleyi King (Escalloniaceae), was known from a collection made by Ridley in 1892 from Seletar and another by Cantley in the 1880s without the locality specified. Much later it was shown by Saw (1989) to occur in Johore and Brunei as well as Singapore. Spatholobus ridleyi is a distinctive species. It has only been collected from Singapore on three occasions and from nowhere else. It was last collected in 1899 and is an example of a rare Singapore endemic that is now extinct. Strychnos ridleyi is known from just two specimens, both from Singapore, the last collected in 1894. Ridley (1923) reported that it was ‘very rare in forests’. The most recent revision for the region by Leenhouts (1962) confirmed it as a valid species. S. ridleyi is another example of a rare Singapore endemic species that is now extinct. In the post-war period, new species were sporadically described either because genera were being revised, such as the ferns Bolbitis singaporensis Holttum (Lomariopsidaceae) and Tectaria griffithii (Baker) C.Chr. var. singaporeana Holttum (Dryopteridaceae); Thunbergia dasychlamys Bremek. (Acanthaceae); Psychotria ‘sp. 9’ K.M.Wong (Rubiaceae) or, in one case, was recently discovered, Cryptocoryne xtimahensis Bastmeijer (Araceae). In 1947, Holttum described the fern Bolbitis singaporensis and noted that it looked like a hybrid between what are now called Bolbitis appendiculata (Willd.) K.Iwatsuki ssp. appendiculata and B. sinuata (C. Presl) Hennipman that all grew together in one valley on Bukit Timah. In 1978, Hennipman formally designated this fern as a hybrid, Bolbitis xsingaporensis Holttum. As it is only known from Singapore, it is an endemic. Wee (1984) reported that it has not been seen ‘for a very long time’. It was last collected in 1943. It is probably now extinct. The case of the fern, Tectaria griffithii var. singaporeana is different. This variety was only ever known from a single plant collected in 1908 from Bukit Timah. The fact that it was not recollected by Holttum, who made an intensive study of ferns in Singapore, nor by other botanists suggests that it was a single aberrant plant that does not warrant taxonomic status. _ Thunbergia dasychlamys, a slender climber, is extremely rare and known from only two specimens from Singapore, both collected by Ridley. It has not been recollected since 1906. It is therefore a Singapore endemic, which is now extinct. 176 Gard. Bull. Singapore 55 (2003) Psychotria malayana Jack is a common and widespread species with fruits that dry conspicuously ribbed. However, some specimens that have fruits that are almost smooth and leaves that are broader and the base not tapered into the petiole were segregated from P. malayana by E.D. Merrill as recorded on his determination slips dated 1950 on the specimens. These specimens were formally recognised as distinct by Wong (1989) under the name Psychotria ‘sp. 9’. The status of this taxon requires detailed investigation to decide whether Psychotria ‘sp.9’ is in fact a distinct taxon or whether it is a southern form of P. malayana, something that is beyond the scope of this study. It is not, however, endemic to Singapore as Merrill assigned a specimen from Malacca (Ridley s.n. June 1892 from Gunung Mereng) to this taxon. In Singapore, this taxon was collected from several localities giving the impression that it was not rare. However, there are no specimens collected after 1906, which strongly suggests that the Singapore population is now extinct. Recently, Bastmeijer & Kiew (2001) described a new hybrid from Singapore, Cryptocoryne xtimahensis Bastmeijer (Araceae) from the Bukit Timah Nature Reserve. Its present status is endemic to Singapore as it has not been collected from elsewhere, but considering how readily species of Cryptocoryne hybridise, it is likely that in future it will be encountered in southern Peninsular Malaysia, where both the putative parents grow. Keng (1990) in his flora of Singapore cited two species, Polyosma kingiana Schltr. (Escalloniaceae) and Sabia erratica P.M. v.d. Water (Sabiaceae) as endemic to Singapore and both were listed as ‘strictly endemic to Singapore’ in Davis et al. (1995). The inclusion of P. kingiana appears to be an error as this species, together with the name it was originally described under, P. flavo-virens Ridl., was not recorded from Singapore by Ridley (1922). The case of Sabia erratica 1s an interesting one as van de Water (1980) clearly stated in describing the species that the label on the type specimen at Kew, which gave its origin as Bukit Timah, was clearly the wrong one as it described the plant as a tree 30 m tall whereas Sabia is a genus of climbers. The data on the label recorded the collector as Ngadiman (not Ngadiwan as given by van de Water) with the Field Number 36149 (which is a Singapore Field Number, not a Kepong number as reported by van de Water), collected from Tree No. 454 in Bukit Timah on 22 May 1940 and the specimen was identified as Scaphium (Sterculiaceae). The collection belonging to this label is in the Singapore Herbarium (SING) and represents a specimen of Scaphium macropodum (Miq.) Beumee ex Heyne, which is indeed a tree. The type specimen of Sabia was distributed from Kepong to Kew and at some point the label from the Scaphium specimen was attached to the Sabia specimen. Van de Water commemorated this mix-up in the specific epithet ‘erratica’ meaning wandering. Since Sabia erratica was described, two other specimens have been identified at Kew as belonging to this species. They are Makoto Togashi 622240 collected from Endemic Plants in Singapore 177 Cameron Highlands, Pahang, at 6500 feet on | Feb 1962 and T & P 6/4 collected from the Palas Tea Estate, Gunung Brinchang, Cameron Highlands, on | April 1979. There is a duplicate of the latter specimen in SING. It is therefore now clear that Sabia erratica is a very rare endemic from Cameron Highlands (probably confined to Gunung Brinchang) and is not found in Singapore. The status of Hanguana ‘Singapur’ (Hanguanaceae) suggested as a possible new taxon (Tillich & Sill, 1999) is discussed in the Appendix. It is not endemic. Discussion Out of the potential candidates, only seven can be considered endemics (Table 1). As mention above, two are hybrids that may have resulted from disturbance to the natural habitats in the last hundred years. One variety, Tectaria griffithii var. singaporeana, is doubtfully valid taxonomically as it is based on a single plant. Singapore does not in fact possess a flora that is distinct from the surrounding region but is a part of the flora found in southern Johore as well as showing affinities with the northern flora of Borneo. The endemic status of the species in Table 1 may not stand the test of time as the flora of Johore is relatively poorly collected compared with that of Singapore and it is likely that these species also occur there. It is interesting that three endemics are climbers, the life form most poorly represented in herbaria, because of the inaccessibility of their foliage, flowers and fruits high up in the canopy. Their endemic status in Singapore may therefore be an artefact of more intensive collecting. (Singapore has always had more botanists per km/? than any other country in the region). If a search is carried out in neighbouring countries, these climbers can be expected to be found. Table 1. Endemic species in Singapore Species Life form No. localities & date Status last collected Bolbitis xsingaporensis fern 1 1943 extinct Flickingeria laciniosa epiphyte | 1891 extinct Spatholobus ridleyi climber Z 1899 extinct Strychnos ridleyi climber d: 1894 extinct Thunbergia dasychlamys _ climber 2 1906 extinct Tectaria griffithii var. fern 1 1908 extinct singaporeana Cryptocoryne xtimahensis aquatic l extant 178 Gard. Bull. Singapore 55 (2003) Most of these endemic species were always very rare and are known from only one or two localities. Their extreme rarity meant that habitat destruction or disturbance would tip them into extinction. Most of this extinction apparently occurred about a hundred years ago as Singapore was intensively botanised in the 1920s and 1930s without these species being re-collected. Now only one of the seven still survives. Seven endemics for a flora of about 2300 species of vascular plants represents a very low level of endemism (0.3%), though it is higher (1.6%) if just ferns are considered. One would not expect the Singapore flora to include a large number of endemics as Singapore lacks many of the most biodiverse habitats with high endemism of the region, such as mountains, limestone hills and ultramafic areas. Conclusion This survey shows that seven species are endemic to Singapore and that of these six, Bolbitis xsingaporensis, Flickingeria laciniosa, Spatholobus ridleyi, Strychnos ridleyi, Tectaria griffithii var. singaporeana and Thunbergia dasychlamys, are extinct; only Cryptocoryne xtimahensis is still alive in the Bukit Timah Nature Reserve. The other eight species thought at one time to be endemic have been shown, as regional floras have become better known, to have a wider distribution and grow outside Singapore. Seven of these were always very rare and have not been re-collected in Singapore for many years, which indicates that their Singapore population is extinct, although the species continues to survive elsewhere. These include Dischidia singaporensis, Henckelia puncticulata, Hypobathrum coniferum, Piper flavimarginatum, Polyosma ridleyi, Psychotria ‘sp. 9’ and Spatholobus ferrugineus var. sericophyllus. A population of the eighth, Litsea ridleyi, still survives in Singapore in the Bukit Timah Nature Reserve but is very rare (Maxwell, 1983). The presence of the one extant endemic taxa and one rare species in the Central Catchment Reserve reinforces its value as a protected conservation area for the biodiversity of Singapore. The hybrid endemic to Singapore, Cryptocoryne xtimahensis, grows in two adjacent small pools within the Bukit Timah Nature Reserve. Its long-term survival is therefore vulnerable to any hydrological changes to that one stream. Active monitoring and management, particularly in dry periods, is therefore important. Acknowledgements This survey was prompted by Saifuddin Suran’s enquiry as to the status of the ‘endemic’ species mentioned in the Centres of Plant Diversity book. We thank Josef Bogner for drawing our attention to the Hanguana ‘Singapur’; S. Chan for permission to work in the Bukit Timah Nature Reserve; Saw L. G. for information on Sabia specimens at Kepong; C. Chia and Saifuddin Suran, Biodiversity Centre, NParks, for searching for Endemic Plants in Singapore 179 the location of Pulau Terijei; J.F. Veldkamp for help with tracing Hypobathrum names; and the Keeper of Herbarium at Kew for permission for R. Kiew to study specimens in his care, and to Lena Chan for comments on the manuscript. References Bastmeijer, J.D. and R. Kiew. 2001. A new Cryptocoryne hybrid (Araceae) from Bukit Timah Nature Reserve, Singapore. Gardens’ Bulletin Singapore. 53: 9- iF. Davis, S.D., VLH. Heywood and A.C. Hamilton. 1995. Centres of Plant Diversity. Vol. 2.Asia, Australasia and the Pacific. WWE and IUCN, U.K. Keng, H. 1990. The Concise Flora of Singapore. Gymnosperms and Dicotyledons. Singapore University Press, Singapore. Kiew, R. 1987. The herbaceous flora of Ulu Endau, Johore-Pahang, Malaysia, including taxonomic notes and descriptions of new species. Malayan Nature Journal. 41: 201-234. Leenhouts, P.W. 1962. Loganiaceae. Flora Malesiana. 1, 6: 293-387. Maxwell, J.F. 1983. New and interesting plant records for Singapore. Gardens’ Bulletin Singapore. 35: 191-198. Ridder-Numan, J.W.A. and H. Wiriadinata. 1985. A revision of the genus Spatholobus (Leguminosae-Papilionoideae). Reinwardtia. 10: 139-205. Ridley, H.N. 1900. The flora of Singapore. Journal of the Straits Branch of the Royal Asiatic Society. 35: 27-196. Ridley, H.N. 1922. Flora of the Malay Peninsula. Vol. 1. Reeve, U.K. Ridley, H. N. 1923. Flora of the Malay Peninsula. Vol. 2. Reeve, U.K. Ridley, H.N. 1924. Flora of the Malay Peninsula. Vol. 3. Reeve, U.K. Rintz, R.E. 1980. The Peninsular Malaysian species of Dischidia (Asclepiadaceae). Blumea. 26: 81-126. 180 Gard. Bull. Singapore 55 (2003) Saw, L.G. 1989. Sabiaceae. Tree Flora of Malaya. 4: 426-431. Seidenfaden, G. and J.J. Wood. 1992. The Orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Denmark. Tillich, H.-J. and E. Sill. 1999. Systematische Studien zur Morphologie und Anatomie von Hanguana Blume (Hanguanaceae) und Flagellaria L. (Flagellariaceae), mit der Beschreibung einer neuer Art, Hanguana bogneri spec. nov. Sendtnera. 6: 215-238. Water, Th.P.M. van de. 1980. A revision of Sabia (Sabiaceae). Blumea. 26: 1-64. Wee, Y.C. 1984. Common Ferns and Fern-Allies of Singapore. Malayan Nature Society (Singapore Branch), Singapore. Wong, K.M. 1989. Psychotria. Tree Flora of Malaya. 4: 396-399. Appendix: Taxonomic notes with the inclusion of specimens in the Singapore Herbarium (SING). Bolbitis xsingaporensis Holttum Basionym: Bolbitis singaporensis Holttum. Gard. Bull. Str. Settl. 11 (1947) 271. Type: Bukit Timah — Hullett s.n. March 1882 (holo SING, iso SING). Other specimens: Bukit Timah — C.G. Matthew 12567 1906 (SING), Holttum s.n. 27 Feb 1923 (SING), Holttum SFN 19799 12 Nov 1928 (SING), Holttum s.n. 12 Dec 1928 (SING), Holttum s.n. Feb 1943 (SING). Cryptocoryne xtimahensis Bastmeijer. Gard. Bull. Singapore. 53 (2001) 11. Type: Bukit Timah - Kiew et al. RK 5127 (holo SING). Other specimen: Bukit Timah - P. Blanc 91-1 (SING). Dischidia singaporensis Ridl. J. Str. Br. Roy. As. Soc. 61 (1912) 31. Type: Changi Police Station — Ridley s.n. 1908 (lecto SING). Note: The other specimen Ridley listed as belonging to this species, Rintz (1980) identified as Hoya lacunosa Bl. Flickingeria laciniosa (Ridl.) A.D.Hawkes. Basionym: Dendrobium laciniosum Ridl. J. Linn. Soc. 32 (1896) 242. Type: Pulau Seletar — Ridley s.n. 1891 (holo SING). Endemic Plants in Singapore 181 Other specimen: Pulau Seletar — Goodenough s.n. 1891 (SING). Hanguana ‘Singapur’ Josef Bogner (letter dated 5th Nov 2002) suggested that the Bukit Timah population of Hanguana, based on his observations of a sterile plant in the Munich Botanic Garden grown from seed collected in Bukit Timah, may be a new species, because unlike H. malayana (Jack) Merr. (Hanguanaceae) with which it had previously been identified, it was much smaller and did not produce long runners with young plants at the tip. Tillich & Sill (1999) compared the morphology and anatomy of this sterile plant, which they called H. ‘Singapur’, with three other Hanguana species from Borneo, H. major Airy Shaw, H. bogneri Tillich & Sill and H. ‘Borneo’. However, they did not compare the Bukit Timah plant with H. malayana. Hanguana malayana can be a confusing species as its extreme forms look very different. Plants that grow on lake or pond margins are robust plants that produce long runners (the aquatic form), while plants that grow in forest in wet hollows or on slopes (the terrestrial form) are more slender plants and their runners are much shorter. However, since there are so many plants that are intermediate between these two forms, Backer (1951) and Backer & Bakhuizen van den Brink (1968) concluded that the aquatic and terrestrial forms belonged to a single variable species. Unfortunately, neither of these characters (length of runners and robustness of the plant) can be seen on herbarium specimens and data on labels is also usually lacking. In deciding whether the Bukit Timah population represents a new species, it was necessary to compare it with the complete range of variation in Hanguana malayana from the aquatic to the terrestrial form. To this end, a study was initiated to find, firstly, a suite of characters that can be examined from herbarium specimens, that can be used to assess whether the terrestrial form is different from the aquatic one in any characters other than robustness and runner length, and, secondly, to test whether the Bukit Timah population, which grows in forest, falls within the range of variation of either of these forms. This study was carried out by R. Kiew, A.T. Gwee, P.K.F. Leong, Mohd Shah Mohd Noor and Samsuri Ahmad. Based on the literature, eleven characters were selected that could possibly separate the two forms, namely, leaf shape (ensiform v. lanceolate), indumentum (hairy v. glabrous), petiole (present or absent), lamina length, width, and the ratio of width to length, lamina tip (acute v. acuminate), inflorescence length (excluding stalk length), length of lowest inflorescence branch, indumentum of inflorescence, fruit shape (globose or fusiform) and fruit length. These were tested on herbarium specimens of both forms from Peninsular Malaysia. As with previous studies (Backer, 1951; Backer & Bakhuizen van den Brink, 1968), no character could be identified that would reliably separate the aquatic from the terrestrial form. The characters either showed continuous variation, such as lamina 182 Gard. Bull. Singapore 55 (2003) length and width or inflorescence length, or, if disjunct, such as fruit shape, did not correlate with other characters. The Bukit Timah Hanguana population grows in forest along streams and on slopes, i.e. the habitat of the terrestrial form. Fertile specimens in the Bukit Timah population were examined in the field for nine characters (leaf shape, leaf tip, petiole and lamina length, lamina width, inflorescence length, length of the lowest inflorescence branch, fruit shape and length). Results showed conclusively that they fell within the range of the Peninsular Malaysian population although in lamina length, they were in the lower range (mean length 57 cm, range 41-118 cm) compared with mean length 76 cm, range 20.5—153 cm for H. malayana in Peninsular Malaysia. Their smaller size might be explained by the nutrient-poor habitat of most of the population, which grows on the coarse sand in an eroded streambed. Those on the slope above the stream where there is a litter layer are larger plants but nevertheless still fall within the range of variation seen in Peninsular Malaysia. There was no difference for the other characters between the Bukit Timah and Peninsular Malaysian populations. A few plants growing in water on the streambed had short runners up to 20 cm long. This study therefore shows that the suggestion by Bogner that the Singapore population of Hanguana represents a new species cannot be upheld as it falls within the range of variation seen in H. malayana. Backer, C.A. 1951. Hanguana. Flora Malesiana. Ser. 1. 4: 248-250. Backer, CA. and R.C. Bakhuizen van den Brink. 1968. Hanguana. Flora of Java. 3: 20224: Henckelia puncticulata (Ridl.) A. Weber. Synonym: Didymocarpus perdita Ridl. J. Str. Br. Roy. As. Soc. 44 (1905) 54. Type: Seletar — Ridley s.n. 3 Nov 1889. (holo SING). Hypobathrum coniferum (Ridl.) Kiew, comb. nov. Basionym: Petunga conifera Ridl. J. Str. Br. Roy. As. Soc. 79 (1918) 81; Ridl. FI. Mal. Pen. 2 (1923) 85. Synonym: Hypobathrum sp. 11 K.M.Wong. Tree Fl. Mal. 4 (1989) 355. Invalid name: Hypobathrum coniferum Bakhf. in Keng, Concise Flora of Singapore Gymnosperms and Dicotyledons (1990) 156. Type: Singapore — Gardens Jungle Ridley 10722 1899 (holo SING). Notes: Bakhuizen f. never published the combination cited by Keng, which is therefore invalid. Hypobathrum coniferum resembles H. venulosum (Hook.f.) K.M.Wong but differs in its very short inflorescences (up to 1.5 cm long) and its larger and broader Endemic Plants in Singapore 183 leaves with more veins (15—18 x 6—7 cm with 7—9 veins) whereas H. venulosum has long inflorescences up to 8 cm long and leaves 9-13 x 3.5—5 cm with 5-7 pairs of veins. H. coniferum is therefore a valid species. All three specimens of H. coniferum were collected in flower so its fruit remains unknown. Other specimens: Singapore — Gardens Jungle Ridley 8427 April 1897 (SING); Peninsular Malaysia — Johore, Sungai Kayu Kiah SFN 32366 10 March 1937 (SING). Litsea ridleyi Gamble. Kew Bull. (1910) 317. Type: Reservoir Wood - Ridley 5101 1893 (lecto SING (flowers), here designated). Other specimens: Singapore (no locality) - Cantley s.n. 1880s (SING); Botanic Gardens - Cantley 193 (SING); Gardens Jungle Hill HI038 11 Aug 1970 (SING); Chan Chu Kang - Ridley s.n. 1894 (SING); Changi - Goodenough 3803 1892 (SING); Bukit Mandai - Ridley 3836 1892 (SING), Ridley 4701 1892 (SING), Goodenough 5064 1893 (SING), Ridley10833 22 May 1900 (SING); Holland Road - Hullett s.n. 22 May 1893 (SING); Bukit Timah Nature Reserve - Hamzah Tambi H23 14 July 1971 (SING), Mohd Noor MN1878 22 April 1974 (SING), Shah & Samsuri 3946 1976 (SING), Maxwell 82-61 28 Feb 1982 (SING). Notes: Among the Peninsular Malaysian species, it most resembles Litsea accedens BI. (syn. L. singaporensis Gamble) in leaf and inflorescence characters but the latter species is distinct in its fine reticulations that in dried leaves are raised. The description in Ridley’s flora (1924) contains an error for petiole length, which he describes as 0.15 inches long, when Gamble (1910) originally described it as 1.5 cm long. Piper flavimarginatum C.DC. Rec. Bot. Survey India. 4 (1912) 26. Type: Chan-chu-kan Ridley 3772. Other specimens: Bukit Timah — Ridley s.n. 1899 (SING), Corner SFN 3499 19 June 1938 (SING); Jurong — Corner s.n. 17 April 1933 (SING). Polyosma ridleyi King. J. As. Soc. Beng. 66 (1897) 304. Type: Seletar Ridley 3972 1892 (iso SING). Note: Known from a single collection. Psychotria ‘sp. 9’ K.M.Wong. Tree Fl. Mal. 4 (1989) 398. Specimens: Singapore (without locality) - Cantley 2807 1880s (SING), Ridley 11272 1901 (SING); Chan Chu Kang - Hullett s.n. 1896 (SING); Bukit Mandai - Ridley 6556 1894 (SING); Bukit Panjang (?Puyan) Ridley 12530 1906 (SING); Bukit Timah - Ridley 11238 21 Oct 1900 (SING); Bukit Timah Road - Ridley 904 (SING); Jurong - Ridley 5422 March 1891 (SING). 184 Gard. Bull. Singapore 55 (2003) Spatholobus ferrugineus (Zoll. & Moritzi) Benth. var. sericophyllus Ridl. Fl. Mal. Peni4( 1922). 574. Type: Upper Bukit Timah - Ridley 5578 (lecto SING). Other specimens: Kranji - Ridley 6394 1894 (SING), Sinclair SFN 40687 27 July 1955 (SING); Bukit Mandai Goodenough s.n. 1890 (SING); Mandai Road - Ridley s.n. 1900 (SING); Bukit Timah — Corner & Furtado s.n. 10 Nov 1929 (SING). Spatholobus ridleyi Prain ex King. J. As. Soc. Beng. 66.II (1897) 80. Type: Singapore Botanic Gardens Ridley 640] 1894 (iso SING). Other specimens: Chan Chu Kang - Mat 6795 1894 (SING); Gardens Jungle - Ridley s.n. 1899 (SING). Strychnos ridleyi King & Gamble. J. As. Soc. Beng. 74.II (1908) 621. Type: Tuas [Toas] Ridley 8313 4 May 1894 (isolecto SING). Other specimen: Pulau ?Terijei - Goodenough s.n. 1891 (SING). Notes: The name of the island (?Terijei) is illegible and we were unable to find an island in the Singapore/Indonesian waters off Singapore that has a name that could possibly match. Tectaria griffithii (Baker) C.Chr. var. singaporeana Holttum. Blumea 35 (1990) 551. Type: Bukit Timah - C.G. Matthew s.n. Jan 1908 (K). Note: This variety is known from a single plant so there are no specimens at SING. Thunbergia dasychlamys Bremek. Verh. K. Nederl. Akad. Wetensch. Afd. Natuurk. Sect Tio 14 (1955) 53. Type: Reservoir — Ridley s.n. 1906 (holo SING). Other specimen: Tanglin - Ridley s.n. 1896 (SING). Note: Bremekamp recorded ‘locality illegible’ for Ridley’s 1896 specimen. The label in fact reads ‘hedges in Tanglin’. Gardens’ Bulletin Singapore 55 (2003) 185-188 A Note on Cordia premnifolia Ridl. (Boraginaceae) HASSAN IBRAHIM Singapore Botanic Gardens Singapore 259569 Abstract Cordia premnifolia Ridl. is confirmed as a synonym of C. dichotoma G. Forst., not of C. cochinchinensis Gagnep., which does not occur in Malesia. C. premnifolia is lectotypified. Introduction This paper arises out of the exercise to verify the status of type specimens in the Singapore Herbarium (SING), in this case, to verify the type of Cordia premnifolia Ridl. The family Boraginaceae was revised by Riedl (1997) for Flora Malesiana, in which he regarded C. premnifolia as a synonym of C. cochinchinensis Gagnep. and stated no types were known for C. premnifolia. By including C. premnifolia in the latter, he considered that C. cochinchinensis occurs in Malesia. This paper discusses whether C. premnifolia should be included in the more widely distributed C. dichotoma G. Forst. as suggested by Ng (1989) or in C. cochinchinensis as suggested by Ried] C9979. Cordia premnifolia The three specimens that Ridley (1915) cited when he described Cordia premnifolia are all in the Singapore Herbarium. These specimens were collected in the Pahang river region, Peninsular Malaysia. Ridley (1915) described the leaves of C. premnifolia as ovate, obtuse or subacute, 6.5 x 4 in (16.5 x 10 cm), with the base as rounded or cordate and margins slightly undulate to distantly serrulate. These characters distinguished it from C. griffithii C.B.Clarke (now a synonym of C. dichotoma), which Ridley (1923) described as having ovate-lanceolate acute to acuminate leaves with a slightly cuneate base, 1.5-3.5 x 1-1.75 in (3.8-9 x 2.5-4.4 cm). A comparison of specimens of Cordia premnifolia and C. dichotoma from 186 Gard. Bull. Singapore 55 (2003) Peninsular Malaysia and Singapore shows the three syntypes of the former to have large, broad leaves up to 17 x 11 cm with a rounded to cordate base and entire to slightly serrulate margins; while the majority of the latter has smaller, narrower leaves, 6.5—11 x 3.5—9 cm, with a cuneate base and wavy margin. However, some specimens of C. dichotoma have broader leaves with a rounded base and an entire margin. In addition, even on a single specimen of C. premnifolia there is a great range in leaf size, for example, from 7.5—9.5 x 7 cm to 12.5—15 x 10-11 cm, thereby overlapping with those of C. dichotoma. There are no differences in fruit shape and size and both have the characteristic conical calyx cup. Taking the Peninsular Malaysian and Singapore population of C. dichotoma as a whole, the C. premnifolia specimens, although representing individuals with the largest leaves, fall within the range of variation for that species. Ng (1989) was therefore correct in including C. premnifolia as a synonym of C. dichotoma. He did not, however, lectotypify C. premnifolia. Riedl (1997) made Cordia premnifolia a synonym of C. cochinchinensis without giving a reason for his decision as well as without seeing any of the three syntypes at SING and, because of this, he considered that C. cochinchinensis occurred in Malesia. Cordia cochinchinensis In 1951, Johnston’s account of Cordia dichotoma highlighted its wide distribution ranging from southern China and northern India, south through Indochina and Malesia (including Peninsular Malaysia) to northeastern Australia. Gagnepain & Courchet (1914) first described C. cochinchinensis from specimens collected in Cambodia, Vietnam and Thailand (the Taleng Mountains in the Petchabouri region). Ried! (1997) recorded C. cochinchinensis as occurring in Malesia presumably because he considered C. premnifolia as a synonym of C. cochinchinensis. There is no other documentation of this species being found elsewhere in Malesia. For example, it is not included in the Flora of Java (Backer & Bakhuizen, 1965), the Tree Flora of Malaya (Ng, 1989), the Tree Flora of Sabah and Sarawak (Dayang Awa, 1996) or Merrill’s 1923 Enumeration of Philippine Flowering Plants. There are no specimens of Cordia cochinchinensis at SING but from the descriptions by Gagnepain & Courcet (1914) and Johnson (1951), C. cochinchinensis and C. dichotoma are very similar species with overlapping geographical ranges. The main difference appears to be in the fruit: ovoid and up to 30 mm long and 22 mm wide with a campanulate fruit calyx in C. cochinchinensis compared with ovoid and about 22 mm long and 13 mm wide with a broadly cup-shaped to an inverted cone- shaped fruit calyx about 6 mm long and 12 mm wide with obscurely lobed edges in C. dichotoma (including C. premnifolia). Unfortunately, there is as yet no Cordia premnifolia 187 revision of the Boraginaceae for the Flora of Thailand, where both species occur. It is beyond the scope of this paper to conclude whether C. cochinchinensis and C. dichotoma are conspecific. However, C. cochinchinensis does not occur in Peninsular Malaysia and therefore not in Malesia either. Cordia dichotoma G. Forst. Cordia dichotoma G. Forst., Fl. Ins. Austr. (1786) 18, nr. 110; Johnston, J. Arnold Arbor. 32 (1951) 8; Backer & Bakh. f., Fl. Java 2 (1965) 459; Ng, Tree Fl. Malaya 4 (1989) 60, Dayang Awa, Tree Fl. Sabah & Sarawak 2 (1996) 98, fig. 2; Riedl, Fl. Malesiana I.13 (1997) 74. Type: New Caledonia, Forster s.n., (not seen) Synonym: Cordia premnifolia Ridl., J. Str. Br. Roy. As. Soc. 68 (1915) 12; Fl. Mal. Pen. 2 (1923) 438. Type: Peninsular Malaysia — Tanjong Antan, Pahang, Ridley 2158 (lecto SING, here designated). Other syntypes examined: Peninsular Malaysia - PAHANG: Pahang River Burn Murdoch 186 (SING), Kuala Lipis Machado 11567 (SING). Notes: Ridley 2158 is chosen as the lectotype among the specimens it is the most complete having mature fruits. Acknowledgements I thank the Asean Regional Centre for Biodiversity Conservation for providing the grant for the Singapore Type Project and Ruth Kiew, Singapore Botanic Gardens, for support and guidance in preparing this paper. References Backer, C.A. and R.C. Bakhuizen van den Brink 1965. Boraginaceae. Flora of Java. 2: 457-459. Dayang Awa, A.L. 1996. Boraginaceae. Tree Flora of Sabah and Sarawak. 2: 97-100. Gagnepain F. and L.D.J. Courchet. 1914. Boraginacées. Flore Generale |’ Indo-Chine. 4: 203-204. 188 Gard. Bull. Singapore 55 (2003) Johnston, I.M. 1951. Studies in the Boraginaceae, XX, Representatives of three subfamilies in Eastern Asia. Journal of the Arnold Arboretum. 32: 8-11. Merrill, E.D. 1923. Boraginaceae. An Enumeration of Philippine Flowering Plants. 3: 373-374. Ng, F.S.P. 1989. Boraginaceae. Tree Flora of Malaya. 4: 60-62. Riedl, H. 1997. Boraginaceae. Flora Malesiana. I, 13: 68-79. Ridley, H.N. 1915. New or Rare Malayan Plants. Journal Straits Branch of Royal Asiatic Society. 68: 12. Ridley, H.N. 1923. Boragineae. Flora of the Malay Peninsula. 2: 438-439. Gardens’ Bulletin Singapore 55 (2003) 189-200 New Species of, and Other Notes on, Chisocheton and Walsura (Meliaceae) D. J. MABBERLEY Nationaal Herbarium Nederland, University of Leiden, The Netherlands; 'National Herbarium of New South Wales, Royal Botanic Gardens Sydney, Mrs Macquaries Road, Sydney 2000, Australia' Abstract Chisocheton maxilla-pisticis Mabb., from Sabah, Malaysia, and Palawan, Philippines, and C. velutinus Mabb., from Sarawak, Malaysia, and Brunei, are described as new. A third new Chisocheton species, from Sarawak, is so far known only from a single fruiting specimen. Walsura pinnata Hassk. sensu T.Clark includes a number of distinctive Bornean entities, two of which are here afforded or re-afforded specific rank: Walsura decipiens Mabb., from north- ern Borneo and W. grandifolia Ridl. from the Sarawak limestone. Chisocheton nicobarianus Debnath & Sreek. is reduced to C. macrophyllus King, which is lectotypified here, while Walsura yunnanensis C.Y.Wu from China is reduced to W. pinnata sensu lato (W. cochinchinensis (Baill.) Harms). Introduction In preparing the text of Meliaceae for the Tree Flora of Sabah and Sarawak, it be- came clear that new Chisocheton species signaled in the author’s monograph (1979; see also 1995) could now be described using materials collected since its publication. Examination of Borneo materials assigned by Clark (1994) to Walsura pinnata Hassk., a perplexingly variable species, has shown that two distinct species, one of them new, are better segregated from it. In passing, notes on allied taxa from outside Borneo are included. Chisocheton Blume 1. Chisocheton maxilla-pisticis Mabb., sp. nov., Fig. | Chisocheton, C. macrophyllo King affinis, indumento dense-fulvo, foliolis costis paucioribus, pseudogemmula hiante vix circinnata, maxilla pisticis simulante, floribus pusillioribus, differt. Typus: Malaysia, Sabah, Telupid, Pinangah F.R., 10 July 1993, Sawan Tingki SAN 136830 (holo SAN!, iso KEP!, SAR!). ' Address for correspondence 190 Gard. Bull. Singapore 55 (2003) Rather pachycaul tree with sparsely branched crown to 45 m tall; fluted bole to 90 cm diameter with steep buttresses to 1(—3) m tall. Bark smooth to scaly, markedly lenticellate; inner bark reddish brown; sapwood pale yellowish white. Twigs brown, with longitudinal cracks, cicatrices conspicuous, latex white. Leafy twigs 1.2-1.5 cm diameter, fulvous-tomentellous. Leaves in dense terminal spirals, to 1.3 m long, pseudogemmulate; petioles 8.5—16.5 cm long, more or less angled or grooved, like rachis, indumentum sericeous to pilose; leaflets in up to at least 17 pairs, oblong- ovate, weakly falcate, to 21 x 8.5 cm, rather fleshy (Pennington), young ones brownish or reddish, densely pilose on midrib and costae, also abaxially scattered between veins, bases asymmetrical, rounded or, particularly in juveniles, cuneate, apices acute to shortly acuminate, costae c. 17 on each side, spreading, rather prominent adaxially in sicco, petiolules to c.9 mm long, densely pilose; pseudogemmula of rather unfurled leaflets, densely long-hairy. Thyrse to 50 cm long, narrowly paniculiform, tomentellous; peduncle c. 18 cm long, branches rather distant, to 10 cm long, squarrose; ultimate branchlets cymulose, many-flowered; pedicels tomentellous. Calyx c.2 mm tall, cupular, sericeous, margin truncate to weakly 4-lobed. Petals 4 or 5, c. 11 mm long, c. 1.8 mm wide, linear-spathulate, sericeous without, glabrous within, imbricate, apices concave (corolla clavate in males). Staminal tube swollen at mouth, weakly adherent to petals at base, hairy at least along interlobe sutures near apex without, hairy within, margin with 6—8 linear, more or less 2- or 3-toothed, lobes to 2.5 mm long; anthers 5—8(9), c. 2 mm long, oblong, locellate, basifixed, slightly exserted. Disc obscure. Ovary 4-locular, sericeous; style sericeous in proximal 5/6; stylehead subcylindrical with flattened apex. Infructescences to 30 cm long, axillary or on twigs behind leaves. Capsule spherical, to at least 9 cm diameter, reddish brown, latex white; pedicel to 9 mm across. Seeds unknown. Other material seen: SABAH: Kinabatangan, Tedong, 21 July 1962, Singh SAN 31087 (K!,L!,SAN!,SAR!); Lahad Datu, Kretam, 28 May 1954, Wood SANA 4769 (KEP!, L!, SING!), Lahad Datu, road to Taliwas, Danum Valley, 24 June 2000, Madani et al. SAN 143512 (SAN!); Ranau, Lohan F.R., 28 Oct 1963, Pennington 7934 (FHO!, K!, KEP!, L!, SAN!); Sandakan, Sepilok, boundary between Compts 4 & 5,22 Oct 1963, Pennington 7915 (FHO!, L!, SAN! = Meijer SAN 39141 (SAN!)), Sg. Daghat 1960 Coupe, 18 Oct 1961, Meijer SAN 27861 (SAN!), “Keratam lineated land’, 27 May 1983, Sigin & Lideh SAN 97058 (SAN!); Tawau, Kalabakan, 30 miles WNW of Tawau, 24 April 1954, Wood SAN A 3696 (KEP!, L!, SING!). KALIMANTAN: Loa Haur, Figure 1. Chisocheton maxilla-pisticis. A, flowering twig; B, young leafy shoot, C, older leaflet; D, adaxial view distal part of staminal tube; E, flower bud; F, longitudinal section of flower bud; G, abaxial view of distal part of staminal tube; H, fruit. (A from SAN 31087, B from SAN 136830, C from Pennington 7915, D—-G from SAN 31087,H from SAN A 4769.) 191 Chisocheton and Walsura 192 Gard. Bull. Singapore 55 (2003) west of Samarinda, 60 m, 12 May 1952, Kostermans 6834 (A!, K!, KEP!, L!); Nunukan (northern part), 2 Dec 1953, Kostermans 8897 (K!, L!, SING!); W Kutei, Kelindjau River, 12 June 1954, Kostermans 9571 (K!,L!,SING!), bb 16168 (SING!); SE Borneo, Tidoeng vicinity, 15 m, bb 17894 (L!); Berouw, 400 m, 4 July 1934, bb 19239, (L!). PHILIPPINES: Palawan, Pagdanan Range, Ibangley Brookside Hill, 21 Apr. 1984, Ridsdale SMH 496 (KEP!). Habitat: Lowland dipterocarp forest. Notes: Some older sterile and fruiting specimens were previously referred with a ‘?’ (Mabberley, 1979: 346) to the closely allied Chisocheton macrophyllus King, which does not occur in the Tree Flora area. The specific epithet, meaning shark-jaw, refers to the open pseudogemmula characteristic of the leaf of this tree. 2. Chisocheton velutinus Mabb., sp. nov., Fig. 2 Chisocheton, C. erythrocarpum affinis, sed inflorescentia longiore facile separanda est. In ser. Sandoricocarpis indumento velutino distincta est. Typus: Wong 1536, Brunei, Temburong Dist., Bt. Belalong, west ridge from summit, 650 m, 23 July 1989 (holo SAN!; iso KEP!, SAR!, SING!). Tree to 25 m tall; bole to 35 cm diameter with small buttresses. Bark smooth, medium brown to greyish, hoop-marked, inner bark pinkish to reddish brown; sapwood straw; buttresses to 1.3 m x 7.5 cm. Leafy twigs 1.2—2.5 cm diameter. Leaves to 1 m long, pseudogemmulate; petioles 10-15 cm, more or less angled, like rachis, fulvo- velutinous; leaflets in up to at least 12 pairs, oblong, to 24 x 9 cm, minutely pubescent on sunken midrib, densely velutinous on abaxial surface, bases asymmetrical, rounded, apices shortly acuminate, lateral veins 15—17 on each side, spreading, rather prominent adaxially in sicco, petiolules to 9 mm long. Thyrse to 40 cm long, paniculiform, velutinous; branches rather distant, to 13 cm long; ultimate branchlets cymulose, few-flowered; pedicels densely pubescent. Calyx c. 6 mm tall, cupular, pubescent, green, margin very obscurely 4-lobed to truncate. Petals 4(5), linear-spathulate, to 16 mm long, imbricate, puberulous to pubescent without, glabrous within, cream. Staminal tube cream, weakly adherent to petals at base, more or less hairy without, margin with 6—8 2-toothed lobes to 1 mm long; anthers 6—8, oblong, c. 3 mm long, Figure 2. Chisocheton velutinus. A, flowering twig; B, flower buds; C, longitudinal section of flower bud; D, abaxial view of staminal tube; E, adaxial view of staminal tube; F, young new shoot; G, older leaflet; H, fruit. (A—E from Wong 1536, F—H from Kirkup et al. 940.) — 193 Chisocheton and Walsura ss = a mene ELC =F ~ -~-+4e4-7 —-TF 194 Gard. Bull. Singapore 55 (2003) locellate, basifixed, apices within the tube. Disc obscure. Ovary 4-locular, sericeous; style more or less glabrous; stylehead shortly cylindrical. Infructescences to 30 cm long, axillary or on twigs behind leaves. Capsule globose, c. 5 cm diameter (immature), rostrate when young, velutinous, yellow turning red. Other material seen: Restricted to Borneo. SARAWAK: Ulu Luak, Sg. Setap road, Miri, 15 May 1964, S 21307 (K!, SAR!, SING!); Kapit, Ulu Merit, Ng. Sebatong, 6 Oct 1969, S 28793 (K!, SAR!, SING!). BRUNEI: Temburong, Bt. Belitun, 29 Jan 1994, Kirkup et al. 940 (K!, KEP!, SAN!, SAR!, SING!). KALIMANTAN: Balikpapan, Kostermans 10024 (L!); Peak of Balikpapan (G. Beratus), sandstone, 700 m,9 July 1952, Kostermans 7383 (K!, SING!); Sg. Wain region, N of Balikpapan, 30 m, Kostermans 4175 (SING!). Habitat: Rain forest at altitudes to 650 m. Notes: Some of the older fruiting specimens cited above (Kostermans 7383, Kostermans 10024 and S 28793) were tentatively placed under Chisocheton species B (Mabberley 1979: 372; 1995: 186). The remaining specimens from the Tree Flora region cited there (S$ 25844 from Kapit, Melinau, Ulu Sapurau, Bt. Salong, Sarawak (K!,SAR!), with flowers are in bud) is the flowering element used in that description and remains the only flowering material I have seen of that tree. Apparently conspecific with it is the fruiting specimen S 4551/5 (from the 7" Division, Belaga, Linau, Sg. Iban in Sarawak (FHO!, L!, SAR!), collected as C. medusae). 3. Chisocheton sp. nov. aff. diversifolius Mig. Collected as Aglaia affinis Merr. [= A. odoratissima Blume], this fruiting collection of a 4 m tall treelet (PC. Yii S 48450, Sarawak, 7" Division, Batang Balleh, Bukit Melatai, Camp 2; FHO!, SAR!) has delicate twigs with 7-jugate leaves, with very narrow multi-veined leaflets 5—6 times as long as wide, and a very delicate fruiting rachis c. 20 cm long with pubescent spherical fruits 3-4