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PSS ietrde wavs te che wygnet et or vas goat : at re vee ean Bea Phelan my phe ya hed 6 eA N MEET wee UREN ie FF Ve Pa tat eee ate re RS fe Seo EME 2ST ment ae ise gest aeets “eS a amet ct Pe gah Oi PNET ED Sy ee VIS Sean ca gy had ee Kan cians tants ym pease eta sad yee eae eee Lae VS eae Day oe Dee TaN ea Fete ate eas ates set intent at atten Seo! ett rete pewter be eyteee Awl dhe® byl Pea bee ey ceee vy wale Swe athe Harvard mihi 3 LIBRARY OF THE ARNOLD ARBORETUM HARVARD UNIVERSITY Digitized by the Internet Archive in 2014 https://archive.org/details/gardensbulletins450unse é. 4 P~ apse =f x at oe THE GARDENS’ BULLETIN VOLUME 50 Singapore Botanic Gardens 19 © Singapore Botanic Gardens Singapore, 1998 Applications for reproduction should be made to the Singapore Botanic Gardens, 1 Cluny Road Singapore 259569 DATES OF PUBLICATION OF THE TWO PARTS WERE: Part 1 Pp. 1-124 31 July 1998 Part 2 Pp. 125-336 31 December 1998 Singapore Gardens Bulletin 50 (1998) iti. CONTENTS Choo-Toh, G.T., S.L. Chaw, C.E.Z. Chan, D.H.W. Goh and E.Y.H. Lee. A survey of termites in the Singapore Botanic Gardens Rain Forest....... 171 — 183 Chung, R.C.K. New species of Helicia Lour. and Heliciopsis Sleumer (Proteaceae) LI MERAE SUBARU eee ae Clay Ua afafe rn ocapsbt st cabas neleata tousuulintessdnadssvoweap¥sosaeudeses ccs 151 — 160 Hay, A. A new species of Aglaonema Schott (Araceae) from Seger LL M ALANS Pesta Ree GUao rye Leen ache carci slant LL Ue weacuadensbawn tt ve rabsinndr cous 1—4 Hay, A. The genus Alocasia (Araceae-Colocasieae) in West Malesia PURI Re SHR Wee ee tha SENT ste got cle mem daatso ih Sukie date lancer ty sabes cana pede samt 220 — 334 Julia, S. and E. Soepadmo. New species and new record of Lithocarpus Blume (Fagaceae) from Sabah and Sarawak, Malaysia ............cccseeseeeeeee 125 — 150 Kiew, R. Niche partitioning in limestone begonias in Sabah, Borneo, including MPIEEMOMRUCINC CHENG pete Nay LAs cso tay Melty susie Wtcds eyluib ect dvess cdbglices stoned 161 — 169 Kiew, R. The unique elements of the limestone flora of Batu Tengar Cave Cie CeO a SAID AIMA IVA IANS Hed AN ae Wal A Hcy Lo DAN aubdel heghcseeocibaitsccosstvecancies teas 185 — 196 Kiew, R. and P. van Welzen. Codiaeum variegatum var. cavernicola var. nov. (Euphorbiaceae), the second Codiaeum from, BOMME6 \|....00..4.4.....0/..060 31 — 34 Kochummen, K.M. New species and varieties of Moraceae from Malaysia......... 197 — 219 Lim C. K. Unravelling Pinanga patula (Palmae) sensu Scheffer, Beccari and ANPP IAONTD ey MUNOR ED. eevee e ety Ss Wasdale va koteatesh Meare sbatl uaclath openeaetanevacenst uate ceddemaees 83 — 98 Lim C. K. Four new Pinanga Blume (Palmae) species from SDSS ULE TES WU FUE RSTO AARSgRere Mie Aen ve Rg ee GEILE eas UE By Oeire oeeMegaL 99 — 114 Reza Azmi and R. Kiew. Begonia lazat (Begoniaceae), a new culinary begonia ain Es OTM aD pects cael iret sata VS seasdueisns ia deeath des uk getuesbueahas wv dea teakiSa fee dace 43 — 48 Tan, B.C. The types and original specimens of published names of mosses preserved in the Herbarium of Singapore Botanic Gardens (SING)............ 49 — 57 Tanaka, N. The angiosperm flora of Singapore. Part 8 Cannaceae .......... cesses: 35 — 37 Turner, I. M. Nomenclatural changes for four Malayan species in Phrynium (Marantaceae), Solanum (Solanaceae), Stachyphrynium (Marantaceae) SCE CE Sie Sir (7S PAD OLACCRE )i\y cu. ciat eueat ones tupnce esbcesbiecesieraulnsccdoadiadeocensedesd sins 39 — 41 Turner, ILM. and M.R. Cheek. ‘Some New Eastern Gingers’ — a paper by H.N. Ridley containing descriptions of four species overlooked since BER ea PETLUHOP EN SIR WOON ee ea a ad oes adusladdeuapasvanedeseses 115 — 119 Singapore Gardens Bulletin 50 (1998) iv. Turner, I.M., C.M. Boo, L.M.J. Chen, J.P.S. Choo, A. Latiff and A. Zainuddin Ismail. The botany of the islands of Mersing District, Johore, Peninsular Malaysia. 2. The floras of Pulau Aur and Pulau Pemanggil, with notes on the smaller islamds\...0.) 1.2.0. 59 — 81 Zhu Hua, Wang Hong and Li Baogui. The structure, species composition and diversity of the limestone vegetation in Xishuangbanna, SW China............... 5 — 30 BOOK REVIEWS Flora Malesiana Series II — Ferns and Fern Allies, Volume 3 .............0..0.cceeeeeeeeee 335 — 336 Kochummen, K.M. Tree Flora of Pasoh Forest )....7 233.122 ee eee 121 — 122 Sornsamran, R. and O. Thaitong. Bryophytes in Thailand ......... eee ececeseeneeee 123 — 124 The Gardens' Bulletin Singapore VOL. 50 (Part 1) July 1998 ISSN 0374-7859 : NATIONAL PARKS BOARD Renal Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 4741165 Telefax: 4754295 abner scan ARet THE GARDENS’ BULLETIN The Gardens’ 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 Dr R.T. Corlett (Editor) University of Hong Kong Singapore Botanic Gardens Hong Kong Dr T.W. Foong Dr M.C. Roos (Assist. Editor) Rijksherbarium Singapore Botanic Gardens Leiden, Netherlands Dr S.C. Chin ' Dr E. Soepadmo Singapore Botanic Gardens Forest Research Institute Malaysia Kepong, Malaysia Dr M.J.E. Coode Royal Botanic Gardens Dr W.K. Tan Kew, U.K. Singapore Botanic Gardens The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. Neither the National Parks Board nor 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 contributing authors are found on the inside backcover. PREVA 1 UII The Gardens' Bulletin Singapore VOL. 50 (Part 1) July 1998 ISSN 0374-7859 CONTENTS Hay, A. A new species of Aglaonema Schott (Araceae) from Terengganu, Malaysia. .................. 1 Zhu Hua, Wang Hong and Li Baogui The structure, species composition and diversity of the limestone vegetation in Pola erga IMMA eS WN MO NM orto catrtilaaoieneewdes.fa ceeenpbiverboeles SF wbeslialibenaiichinasWUsverpddedebernanensednusavavonies 5 Kiew, R. and P. van Welzen Codiaeum variegatum var. cavernicola var. nov. (Euphorbiaceae), the second BG Asa oerad RUG OAUAV ER UN SSIMISR eee aL Ney ratele casrseneaats ane < leu Saesueeat aouk sbaliaelatd dh scvlsdannevaressarsabebopsiyeessy yeas St Tanaka, N. he anciospeun ora of Singapore Part 8 Canmaceae ..j..2.0...ccccseronccsnsssepossensesansnentcusonannese 35 Turner, I.M. Nomenclatural changes for four Malayan species in Phrynium (Marantaceae), Solanum (Solanaceae), Stachyphrynium (Marantaceae) and Boesenbergia ia acre eer ea ent Ce a IE Lye Mes: Poel Nese ceca nicat Rasa: neadeslab ont cus idess lipevuedeaerernands 39 Reza Azmi and R. Kiew Begonia lazat (Begoniaceae), a new culinary begonia from Borneo.......... ce eeeeeeeeeeees 43 Tan, B.C. The types and original specimens of published names of mosses preserved in the Metoamumot Singapore Botanic; Gardens (SING) 220s ic. csllicl diese cesslecataends bees 49 Turner, I.M., C.M. Boo, L.M.J. Chen, J.P.S. Choo, A. Latiff and A. Zainudin Ismail The botany of the islands of Mersing District, Johore, Peninsular Malaysia. 2. The floras of Pulau Aur and Pulau Pemanggil, with notes on the smaller islands.................. a0 Lim Chong Keat Unravelling Pinanga patula (Palmae) sensu Scheffer, Beccari and Ridley non Blume ..... 83 Lim Chong Keat Four New Pinanga Blume (Palmae) Species from Peninsular Malaysia ...............0008 99 Turner, I.M. and M.R. Cheek ‘Some New Eastern Gingers’ — a Paper by H.N. Ridley Containing Descriptions of Four Species Ovenooked emce their Publication 1m L9OD) s...f..clicscceckcstesensscnssnccsscscctecsootesceanes Lu5 BOOK REVIEWS Rocwummnen mali dl ree Poneo7 Paso: FOPeSt 5.5. .s0..0..3c.coneiuessesarntesndantaotacenesscacines S.C. Chin - 121 Ne REMAN Ra IRN 282 8 sec e ch kcdnos ope pov chap savkon th bavshanp eradediacausnbyevedecantendes Benito C. Tan - 123 Date of Publication: 31 July 1998 Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 259569 Printed by Oxford Graphic Printers Pte Ltd ayn ae 4 | Ate vt oe ay Ms F» ‘ ey a .o ' # ré iff a ny, Gil Toe. F War stgtes Pere ee aaa i* ike. ok: AAS 7 oh tai rh . ie @ a Pp 7 nm fos al , my Ji Ras i, A ithe Oe. i oy + i 16 - ag . v4 i P| i ny t rs vy ty tts > y wv bh Ls st rr. cee yi Ay 1 vt ot ng A : a hy, H a fey * { an nh Ni Re cn ‘ TaN an an Liv f ; nh MW ig het eas AA G) Bike A My vil + esha ny hia oly haat ih by hy , ; AP a ’ re aL j y wae $y) ds iy me i” 7 ( t 4 4 ic a i : hi vn 4 4 may AVE) } w juin ‘ 3 he’ bh rar es WAAAY TARO Hh al Nay es sani ath Bh. iy uae hint af i bea ih Mae a ibih iy ; ted ae me a 4 > ay rj ie riche yi ait ie ¥ f a Le ae a MR hie: F iN ee) tt , ‘ Nae | rele ey n } ity Ae Lo Au. hes) poolishd ti i a i! A fd vain RY fh WAAR ash te COP (RRC cnlcaty, yt Says ay) ie reas » Mn i | ute . ae a) ee Oe a “I ie 4a até bts 6 aoeh dealt va: Card Mala ei andy Pini nde Mh eka 4, Teo » } ‘ - , ‘ 4/) d : ’ ) ! aby oF - Vanes eH ne nes ee PARLE a, tae ‘ ' ah bias ira Ae a * eee ay ue | Edie esthdicte On il iy i Satie ae ar >A 1 , ’ nrc Atee | >» } ; OAL . ; 7: Mf ‘ ery PK t aa aie ‘h ad NC TEA eal lt ly gh i Sahih 4s. (Aah aaa CMA tant Sete oben a | ai + wy ' - ; ’ » 1 F fe A pie sdetepns ban en Bik eae oak da Pat, Saag ata te FRR RA Airis ol?) CAR it Oe hy AW Ayan Bile | — We : 5 " * } , ; sch eC eUTY hide nit Sy ed hae rédighe eRe y if caw Ox } ‘ i F . eh "y ae nay it cw) ey" ue i Ty. oh ni) hl - ie cay Mh aT ae on ' MA ean aa he bas vee Lanter 4 bie ata ee ,, r V4 a ( ay 4 ' thar ¥ ’ ee 7 way } Ad : hey Ler om by. aA me } i ‘ MS PES i ’ d ray ‘ | Ap! i at i¢ rea? (yw Ad ‘| 7" yA ay vt Ms ; zt es 2 Ny ) { ‘+f j 1 | io. Pants dant " ES mY a Nay , 4 an We Aa es ie dl ; Z iy Treat. d ' iv ij ee Wt ‘y H “al ae | Mm ’ yn he baa * uv ‘ ; wa ds an Bs a u¥ AB iba ; q Lay hf » . SOME Tt ey LUN ee bilaabinls i ei uf Mio a) in te Wal ) iim | "A | ee ek ¥. i i ae ny ah eat neni ¥ a i al a } Ps ets ih il syraers it ant «ere ; oa) gs Par ty de APN iB ran) ene en ie Ee ue Vale ro yr et f F vy A w | Vy pe oo ; ti sae are ne Om 4 ee tye G7 eee errr ft ape whe ie eine an r ‘ ttt bi P oun y Ea | pe yn vs ON TN nssilnt tis) Gardens’ Bulletin Singapore 50 (1998) 1-4. A New Species of Aglaonema Schott (Araceae) from Terengganu, Malaysia. A. HAY Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney 2000, Australia. Abstract Aglaonema flemingianum is described as new from Terengganu State in Peninsular Malaysia. The new species is illustrated and fitted into a previously published key to the species of Aglaonema. Introduction Aglaonema Schott (Araceae) is a genus of about 22 species of rainforest herbs, several of which are much prized as ornamental foliage plants since they are often variegated in nature (Jervis, 1980). The genus occurs in Indochina and southwestern China (9 species), West Malesia (6 species) and Central Malesia (incl. Philippines; 6 species), with only one species, A. marantifolium Bl., extending into East Malesia. Aglaonema was revised by Nicolson (1969), who divided the genus into two sections based on vegetative architecture: Sect. Chamaecaulon, with creeping branched stems, very short leaf sheaths and cataphylls subtending each petiole, and Sect. Aglaonema, with erect stems, longer leaf sheaths and cataphylls rarely present among the petioles. The new species belongs to the latter group. Nicolson also discussed the great variability of several species, and their tendency to be rather poorly differentiated from one another at the extremes of their variation. This new plant, however, is vegetatively so distinct from others as to warrant recognition as a discrete species, though known only from a single collection. Aglaonema flemingianum A. Hay sp. nov. Fig. 1. Ab aliis speciebus A glaonematis petiolo brevissimo omnino vaginato, vagina apice ligulata ad marginem membranacea, laminae nervis primariis numerosissimis superne impressis, inferne prominentibus, pistillis paucis differt. TYPUS: Cult. Hort. Reg. Bot. Sydney (Acc. No. 940284) originally collected from Malaysia, Terengganu, Sekayu, Ayer Terjun, Hay et al. 9216 (NSW, holo). NM Gard. Bull. Singapore 50(1) (1998) Erect herb to c. 30 cm tall; stem 1-2 cm diam., internodes c. 1 cm long, dark green, smooth; leaves in a tight rosette, subtended by cataphylls only at the beginning of a sympodium module; cataphylls 2—?3 becoming progressively elongate, up to c. 7 cm long; petiole c. 4 cm long, c. 4 mm wide at the apex, sheathing throughout its length and clasping the stem more or less throughout; wing of sheath 6-8 mm wide, not or hardly tapering distally, membranous, the very margin eventually (in oldest leaves) becoming dry, brown and scarious, the apex of each wing extending free for c. 1 cm and overlapping the lower part of the leaf blade; leaf blade ovate to narrowly ovate, to c. 22 cm by c. 8 cm, widest at the middle, mid- green, not glossy; apex of blade acute to rounded and very shortly (c. 5-8 mm) acuminate and mucronate for c. 2 mm, the base tapering and narrowly rounded; midrib basally c. 5 mm broad and distally tapering, flat and slightly raised on the adaxial side, distally becoming flush and then impressed in the distal quarter, abaxially conspicuously raised and rounded in cross- section; primary lateral veins numerous, c. 12 on each side of the midrib, inserted c.5 mm apart in the basal part of the midrib, further up c. 1 cm apart and distally c. 2 cm apart, diverging at c. 30° and gradually curving towards the leaf apex before running into the margin, adaxially impressed, abaxially prominent; secondary (interprimary) venation parallel to primary and flush ab- and adaxially; higher order venation forming an inconspicuous tessellate reticulum between the primary and interprimary veins; inflorescences to 2 together; peduncle concealed among leaf bases at anthesis, later extending somewhat to be exposed for c. 2 cm, subtended by short blunt cataphylls; spathe pale green, c. 3 cm long, broadly ovate, held boat-shaped at anthesis, c. 1.5 cm wide, open almost to the base, convolute in the lower 4 mm, inconspicuously keeled along the abaxial midline, the apex obtuse, mucronate for c. 2 mm; spadix somewhat exceeding the spathe, 3-5 cm long, stipitate for c. 5 mm at anthesis (elongating slightly afterwards), the stipe mostly adnate to the spathe, free in the upper 2 mm; female zone free, a. single whorl of pistils; ovaries subcylindric, c. 2 mm tall, 1 mm diam.; stigma discoid, cap-like, sessile, c. 2 mm diam.; male zone 2.5 cm long, 8 mm wide at base, tapering in the upper half to a blunt tip, at the base with a whorl of incompletely fertile stamens; stamens not ostensibly arranged into male flowers, close-packed, irregularly 4-lobed, c. 2 mm diam.; fruit unknown. Distribution and habitat: Endemic to Peninsular Malaysia and known only from the type collection from Terengganu, on the floor of wet lowland rain forest on slopes. Notes: Aglaonema flemingianum can be incorporated into Nicolson’s (1969) key to Aglaonema species thus: New Species of Aglaonema 3 Lesle Eka, Figure 1. Aglaonema flemingianum. Hay et al. 9216. a, habit; b, inflorescence; c, inflorescence with part of spathe removed; d, pistils and stamens. Scale bar to a = 4.5 cm; b = 2 cm;c = 1.3 cm; d=5 mm. 4 Gard. Bull. Sing. 50(1) (1998) 12. Venation differentiated into primary and secondary veins. 12a. Petiole about 1/Sth the length of the blade, sheathing throughout; blade with 24 primary Veins .)..53.:.2 600, hes A. flemingianum 12b. Petiole generally about or more than half the length of the blade, sheathing for about 1/2 to 4/Sth its length; blade with 4- LO primary VEINS: 3. chided acapa ieee 13 etc. as Nicolson (1969). Some difficulty may be experienced with lead 6 in that key however, since A. flemingianum falls rather between the two alternatives there, though it nevertheless matches the second alternative (spadix cylindric; spathe elongate) better than it does the first (spadix clavate; spathe globose). Insufficient material exists for the making, designation and distribution of isotypes. However, as soon as the plant from which the holotype was prepared flowers again, further material will be preserved for distribution to KEP, SING and other relevant herbaria. The new species is named in grateful recognition of Conrad D. Fleming who has generously supported many expeditions by Araceae botanists working in tropical Asia. Unlike many other Aglaonema species which are easily cultivated and fast growing, A. flemingianum is very slow-growing, but with very long-lived leaves. The dense rosette of almost sessile leaf blades may represent a useful feature for breeders of ornamental Aglaonema cultivars. Acknowledgements I thank Conrad D. Fleming for generous financial support of my fieldwork in Malaysia in 1994, when this species was first collected. I also thank Dr Jambari Hj. Ali, Head of the Department of Biology, Universiti Pertanian Malaysia, for allowing me to be affiliated with that department during my stay in Malaysia. Prof. Ruth Kiew, Mr Anthonysamy and Mr Roy Bangka provided generous hospitality and invaluable assistance with field work. I am indebted to Lesley Elkan for the botanical drawing, to Clare Herscovitch for technical assistance, Ian McLellan and nursery staff of the Royal Botanic Gardens Sydney for cultivating the living plants, and to Dr D.H. Nicolson for expert comment on the manuscript. References Jervis, R.N. 1980. Chinese Evergreens: Aglaonema Grower’s Notebook. Clearwater, Florida. Nicolson, D.H. 1969. A revision of the genus Aglaonema. Smithsonian Contributions to Botany. 1: 1-69. Gardens’ Bulletin Singapore 50 (1998) 5-30. The Structure, Species Composition and Diversity of the Limestone Vegetation in Xishuangbanna, SW China ZHU HUA, WANG HONG anp LI BAOGUI Xishuangbanna Tropical Botanical Garden, The Chinese Academy of Science, Mengla, Yunnan 666303 P.R.China Abstract The limestone vegetation in Xishuangbanna, tropical southwest China, includes three main vegetation types, six formations, and nine communities, which are described in detail with enumerations of forest profiles and species composition. Species diversity is discussed based on Shannon-Wiener’s indexes for each forest formation. Comparison between the limestone seasonal rain forests and the ones on non-limestone reveals that the limestone seasonal rain forest has a lower species diversity index value per unit area but higher community diversity than the rain forest on non-limestone. Introduction Limestone vegetation is one of the principal vegetation types in tropical Yunnan of southwest China. Because of the great diversity of habitat and topography, limestone vegetation is extremely diverse in community types and very rich in endemic taxa. However, limestone vegetation is even less well known than that not on limestone, owing to the rugged topography. About 19% (3600 km) of Xishuangbanna, the southern-most part of Yunnan, is limestone. Most of the limestone area is still covered by forests and although these have been studied (Liu, 1987; Xu et Jian 1987), little has been published in English. This paper is based mainly on three years’ field work on plots and is a phytosociological study of the limestone vegetation. General Geography Location and topography Xishuangbanna lies between 2109' and 2236' N, 9958' and 10150' E. The region, which borders Burma and Laos, is a mountainous area at the northern margin of mainland southeast Asia and the southern end of the Hengdwan Mountains (part of the Himalayas). Basically, the area has a mountain-valley topography with the mountains running north-south with lower elevations towards the south. Altitude varies from 480 m at the bottom of the lowest valley in the south to 2429 m at the top of highest 6 Gard. Bull. Singapore 50(1) (1998) mountain in the north. The limestone occurs mainly in the south-eastern part of Xishuangbanna as a basically north-south-trending tract and ranges in altitude from 600 m to 1600 m (Fig. 1). There are two main types of limestone topography. One is typical karst hills, which have rocky tops without soil, and slopes partially covered by thin soil. The other is usually much bigger mountains, which also have rocky tops without soil, but have slopes, especially the lower ones, covered by thick soil with fewer limestone outcrops. Because of the diversity of topography and the great site to site variance of soil depth and cover of outcrops, there is a wide range of micro-habitats, i.e. there is a great spatial heterogeneity in the limestone. Climate The region of Xishuangbanna has a typical tropical monsoon climate. In the limestone area, climatic change with altitude is conspicuous. The annual mean temperature is 22°C (600 m alt.) to 18.4°C (1600 m), and the annual temperature accumulation (the sum of daily temperature means where they are > 10°C) is 8000°C (600 m) to 6600°C (1600 m); the monthly mean temperature is 15.9°C (600 m) to 12.3°C (1600 m) for the coldest month and 25.7°C (600 m) to 22°C (1600 m) for the hottest month. The annual precipitation varies from 1200 mm to 1556 mm of which more than 80 percent falls during the rainy season which starts in May and lasts till the end of October. The Hengdwuan Mountains to the north of the region act as a huge barrier keeping out the cold air from the north in winter. Dense fog always exists for the whole of the dry season on the lower hills and in the valleys (average 146 foggy days per year and | mm precipitation per foggy day recorded in Mengla County in the south of the region), which compensates for the insufficient precipitation, so that a tropical moist climate occurs locally in spite of the fact that the region is controlled by strong monsoon climate and at a relatively high latitude and elevation. Methods Limestone vegetation is extremely diverse, especially because there are many communities, which are in different stages of succession. After initial floristic investigation (Zhu et al., 1996), the main and representative primary forest types (which occupied relatively large areas and are climax communities judged by field observation) were selected for establishing plots. For each selected forest type, one to several plots were laid out. The number of plots (or the total sampling area) for a forest type was Limestone Vegetation in Xishuangbanna “euIyD MS ‘euURQsUeNYsSTX UI OUO}SOUT] JO UOTINGISIG *[ aaNsIy Ot O01 a || $d019|NO auojsewl) yym Bay = SdOJ9\NO jNOYWM aseg auo|sew YUM Baly | ( ie reysuay \ — Suoysuir OF l SA 2S —— eS x. te \ ‘ | ; f re vidoawys, 8 Gard. Bull. Singapore 50(1) (1998) determined mainly depending on floristic variance of the forest type. Sixteen plots were used for the analysis described in this paper. Different sized plots were used for different forest types in different topographical sites Owing to considerations of phytocoenological minimal area and in some situations for facilitating field work. Plots were basically 2000-2500 m for seasonal rain forest (7 plots), 500-2000 m for seasonal moist forest (7 plots) and 100 m for dwarf forest (2 plots) on tops of hills or mountains. (It is difficult to fix plot size even for the same forest type because of the very rugged topography). In each plot, all trees were identified and their dbh. (minimum 5 cm), height and crown coverage measured. Each plot was roughly divided into 5 strips so that frequency of tree species could be calculated (except plots 102-16 and 102-15 in Tables 1 and 7, which were investigated by another botanist’s group without subdivision of the plots). Furthermore, in 3-5 subplots (in each plot) of 5 x 5 m (for seasonal rain forest) or 3 x 3 m (for others), saplings and shrubs were counted, and the cover of seedlings and herbaceous plants were estimated by Braun-Blanquet’s degree of abundance (Braun-Blanquet, 1932). Epiphytes and lianas were identified and abundance estimated by eye. Importance value indexes (IVI) devised by Curtis & McIntosh (1951) were calculated and shown in tables from data of plots except for Tables 1 and 7 in which percentage of total dominant density (% Dens.) and percentage of total dominant breast area (%BA) were calculated from data of plots 102-16 and 102-15. Shannon- Wiener’s indexes (Shannon-Wiener, 1949) for species diversity and Evenness Indexes of Pielou (1966) were calculated from data of plots. For all species in plots, specimens were collected and identified. Species authorities follow Flora Reipublicae Popularis Sinicae. Specimens are kept in the herbaria at KUN and HITBC as well as at SYS. Structure and species composition The primary limestone vegetation can be classified into three vegetation types i.e. tropical seasonal rain forest, tropical seasonal moist forest and tropical montane dwarf forest based on physiognomic, structural and floristic characters as well as habitats (Wang et al., 1997). The tropical seasonal rain forest occurs mainly in wet valleys and on lower slopes below 850- 900 m altitude. The tropical seasonal moist forest occurs mainly on middle slopes and tops of lower hills. However, the distribution of vegetation types is greatly affected and modified by local micro-habitats. Topography seems to have the stronger effect on distributional patterns of vegetation than elevation. For example, the tropical seasonal rain forest occurs Limestone Vegetation in Xishuangbanna 9 occasionally on the upper valleys near 1000 m altitude in some particular sites because of the temperature inversion appearing in the mountain areas. The tropical montane dwarf forest occurs on the tops of hills or mountains. Each vegetation type is further subdivided into formations. 1. Tropical Seasonal Rain Forest Like equatorial lowland rain forest, tropical seasonal rain forest has 3-4 indistinct tree layers. The top layer is mainly emergent trees more than 30 m tall (tallest up to 45 m) and has c. 30% of crown coverage; the second layer, up to 30 m tall with almost continuous crowns (70-80% coverage) and a greatest density of stems, is the main canopy layer; the third layer, 5-18 m tall, and with crown cover of c. 40%, consists of small trees and juveniles of species from the upper layers. In some sites, the third tree layer can be further divided into two sublayers: upper sublayer (10-18 m tall) and lower sublayer (5-9 m tall). Buttresses and cauliflory are common, and both big woody climbers and vascular epiphytes are abundant. The forest is mainly evergreen in spite of the fact that there are some deciduous trees in the emergent layer. This forest type occurs in wet valleys and lower slopes of hills or mountains and below 1000 m altitude. Tropical seasonal rain forest contains two main formations: la. Ravine seasonal rain forest This occurs in the wettest valleys and lower slopes as well as shaded slopes (usually northeast facing). It has fewer than 10% deciduous trees, either in number of species or in individuals and all exist in upper layer. Floristically the formation is characterized by Pometia tomentosa (Fig. 2). The similar forest type, which occurs on non limestone habitats in the region, was called “wet seasonal rain forest” in early Chinese botanical references (Qu, 1960), but the term “ravine seasonal rain forest” was preferred by recent authors owing to its valley habitat (Jin and Ou, 1997; Zhu et al., 1998). There are 90 tree species, 16 shrub species, 32 herbaceous species, 26 liana and 5 epiphyte species in the plots (cumulative area of 7400 m ). Two communities have been recorded: (i) Pometia tomentosa-Alphonsea monogyna community. This community occurs in the wettest bottom of valleys or on lower slopes, with Pometia tomentosa as dominant species of the upper tree layer. Its canopy is usually 35-40 m tall. Alphonsea monogyna is the dominant species and Pseuduvaria indochinensis the sub-dominant species in second tree layer. Horsfieldia pandurifolia is the dominant in the upper sublayer of the third tree layer ( 9-20 m tall) and Cleidion spiciflorum is the dominant in lower sublayer of the third tree layer (5-10 m tall). The understorey, with a 10 Gard. Bull. Singapore 50(1) (1998) / SS OS oF ah iM F i? | ye! WA Mail | /o Figure 2. Forest profile of ravine seasonal rain forest 1. Terminalia myriocarpa; 2. Pometia tomentosa; 3. Alphonsea monogyna; 4. Knema furfuracea; 5. Baccaurea ramiflora; 6. Garcinia cowa; 7. Syzygium latilimbum; 8. Barringtonia macrostachya; 9. Lasioccoca comberi var. pseudoverticillata; 10. Pittosporopsis kerrii; 11. Pseuduvaria indochinensis; 12. Pterospermum lanceaefolium; 13. Drypetes cumingii; 14. Horsfieldia pandurifolia; 15. Musa acuminata; 16. Trigonostemon thyrsoideum; 17. Ventilago calyculata; 18. Neottopteris nidus; 19. Rhaphidophora hongkongensis; 20. Combretum latifolium; 21. Fissistigma sp.; 22. Cleidion spiciflorum. Limestone Vegetation in Xishuangbanna 11 cover of 30-40%, consists mainly of saplings and young woody lianas. Pseuderanthemum polyanthum and Leea compactiflora are the commonest shrub species. The herb layer is developed with a cover of 60%. The commonest species are the ferns Crenitopsis fusipes and Bolbites heteroclida, the herb Ophiopogon latifolius and Piper boemerifolium, and the lianas Derris cudatilimbum, and Strychnos nitida. Its physiognomy and profile are almost exactly the same as those of wet seasonal rain forest on non- limestone in the region (Zhu, 1992, 1997). Most species of the community are also found in non-limestone seasonal rain forest, but the latter has many species not present on the limestone (Table 1). Table 1. Pomentia tomentosa-Alphonsea monogyna community Plot no.:102-16 Location: Meng-yue, Mengla Altitude (m): 700-720 Area of plot (m): 80 x 30 Aspect: NE Slope (degree): 0-5 Height of canopy: 40 m Coverage of vegetation: >90% No. of species (2 5 cm d.b.h.): 45 No. of stems: 140 Name of species % Dens. % BA % Dens.+% BA Pometia tomentosa 10.71 20.41 lt Ws Amoora tetratepala 0.71 21.36 22.07 Alphonsea monogyna 15.00 2.76 17226 Horsfieldia pandurifolia 6.43 me 12.14 Ficus altissima 0.71 9.98 10.69 Garuga floribunda var. gamblei Ot 8.56 O24 Cleidion spiciflorum 7.86 0.77 8.63 Diospyros hassellii. - 6.43 HONS) 8.62 Pseuduvaria indochinensis 6.43 1253 7.96 Glycosmis ferruginea 2.14 4.55 6.69 Litsea pierrei var. szemaois 2.86 3:25 611 Debregeasia squamata So, 0.85 4.42 Celtis timorensis 1.43 2.32 395 Prunus zippenliana 2.86 0.72 3.58 Picrasma javanica 2.14 0.74 2.88 Garcinia cowa 2.14 0.70 2.84 Erythrina stricta 1.43 E33 Bae (S Cryptocarya acutifolia 0.71 1,92 2.63 Macropanax dispermus 2.14 0.36 250 Canarium album 0.71 hk 2.42 Litsea dilleniaefolia 1.43 O75 2AS Elaeocarpus austroyunnanensis 1.34 0.67 2.10 Lasiococca comberi var. pseudoverticillata 1.43 0.45 1.88 Cont: 12 Gard. Bull. Singapore 50(1) (1998) Name of species % Dens. % BA % Dens.+% BA Diospyros nigrocortex 1.43 0.45 1.88 Antidesma montana 1.43 0.30 1.73 Tapiscia yunnanensis 0-71 0.94 HIE cs Pterospermum lanceaefolium 1.43 0.20 1.63 Laportea sinuata 1.43 0.15 1.58 Tetrameles nudiflora 0.71 0.67 1.38 Semecarpus reticulatus 0.71 0.64 1.35 Macaranga indica 0.71 0.57 1.28 Toona ciliata 0.71 0.43 | Dysoxylum lukii 0.71 0.40 itt Phaeanthus saccopetaloides 0.71 0.38 1.09 Dysoxylum binectariferum 0.71 0.27 0.98 Drypetes perreticulata 0.71 0.17 0.88 Canarium pimela 0.71 0.14 0.80 Phoebe puwenensis O74 0.11 0.82 Chisocheton sinensis 0.71 0.11 0.82 Trigonostemon thyrsoideum 0.71 0.11 0.82 Antidesma bunius O71 0.07 0.78 Sarcospermum arboreum 0.71 0.03 0.74 Sumbaviopsis albicans 0.71 0.02 0.73 Drypetes cumingii 0.71 0.02 0.73 Dichapetalum gelonioides | 0.71 0.02 0.73 Total 100 100 200 (11) Pometia tomentosa-Celtis philippensis var. wightii community. This community occurs near the bottoms of valleys and on lower slopes in somewhat less wet habitats, with rock outcrops usually covering more than 30% of the ground. It usually has Celtis philippensis var. wightii and Lasioccoca comberi var. pseudoverticillata as co-dominant species in the second tree layer and Pometia tomentosa as a dominant species in the upper tree layer. Sumbaviopsis albicans is the dominant in the upper sublayer of the third layer and Cleidion spiciflorum in the lower sublayer (Table 2). The understorey with a cover of 50% consists almost entirely of saplings. Only a few shrub species are recorded and the common ones are Psychotria siamica, Sauropus macranthus and Miliusa tenuistipitata. The herb layer has a cover of 30%, and the commonest are Tectaria cordatum (a fern), Pilea balansae and Piper polysyphorum. Ventilago calyculata var. trichoclada and Loeseneriella lenticellata are the commonest lianas. Rhaphidophora hongkongensis and Pothos chinensis are frequent epiphytes. This community is transitional toward lower hill seasonal rain forest in physiognomy and floristic composition. Limestone Vegetation in Xishuangbanna 13 Table 2. Pometia tomentosa—Celtis philippensis var. wightii_ community Plot no.: HW9203 HW9202 Location: Menglun Menglun Altitude (m): 700 740 Area of plot (m): 50 x 50 50 x 50 Aspect: NE NE Slope (degree): es 10 Height of canopy (m): a3 30 Coverage of vegetation (%): 100 100 No. of tree species (=>5 cm DBH): 25 19 No. of stems: 118 164 Name of species IvI” IVI Average Celtis philippensis var. wighttii 41.3 56.1 48.7 Lasiococca comberi var. pseudoverticillata 45.1 39.8 42.6 Cleidion spiciflorum 18.7 40.2 29.4 Sumbaviopsis albicans 24.7 30:7 pe Pometia tomentosa 11.8 18.5 (se Ficus altissima Dee —) 13.6 Neonauclea tsiana 12.5 22 12.4 Caryota urens 14.3 11.4 12.8 Amoora tetrapetala a9 15.6 1 a, Drypetes perreticulata 2.7 8.2 10.4 Mitrephora maingayi 8.8 10.2 95 Tetrameles nudiflora - ie eg 8.8 Terminalia bellerica 13.7 - 6.9 Garcinia xanthochymus 3.0 9.0 6.0 Mitrephora wangii yh — 4.9 Duabanga grandiflora 8.7 - 4.3 Alphonsea monogyne 8.2 +9) 4.1 Chukrasia tabularis var. velutina 7.6 ~ 3.8 Dysoxylum hainanensis + cae 3.8 Randia wallichii 5.9 + pss | Pterospermum lanceifolium 5.4 ~ 27, Pseudostreblus indica + 5.6 2.8 Ficus benjamina + 3:3 2.6 Morus macroura 4.0 — 2.0 Ficus glaberrima ae - 1.6 Dysoxylum lenticellatum a2 - 1.6 Glycosmis ferruginea - 3.0 LS Ficus cyrtophylla ~ 3.0 15 Diospyros hassellii - 3.0 5 Horsfieldia tetratepala + jabs Ls Laportea sinuata 2.9 ~ 1.4 Total 300 300 300 YTVI=% Density + % Frequency + % Dominance *)_: not recorded in the plot +: only saplings (<5 cm d.b.h.) or seedlings were recorded in the plot 14 Gard. Bull. Singapore 50(1) (1998) 1b. Lower hill seasonal rain forest Lower hill seasonal rain forest occurs in even less wet habitats mainly on lower hills and sometime on lower sun-facing (usually southwest) slopes. It has the same altitudinal range as formation la, but is never found in valleys. Deciduous trees make up 10-30% of the number of species or individuals and exist in upper layer and as emergents. The similar forest type occurs on non-limestone habitats in the region, and was also called “dry seasonal rain forest” (Qu., 1960), but the term “lower hill seasonal rain forest” was preferred recently considering its habitat.(Jin and Ou, 1997; Zhu et al., 1998). It has a canopy about 30 m tall and relatively clear stratification. The upper layer with a crown cover of 40-50%, is 20-30 m tall. The second layer, which is the main canopy layer, has a crown cover of 70-80% and 10-20 m tall. The third layer with a cover of c. 50-60% is 3-10 m tall. There are some scattered emergents such as Chukrasia tabularis var. velutina, Tetrameles nudiflora and Garuga floribunda var. gamblei (Fig. 3). There are 67 tree species, 12 shrub species, 13 herbaceous species, 32 liana and 4 epiphyte species in the plots (cumulative area of 7400 m’). M Figure 3. Forest profile of lower hill seasonal rain forest 1. Tetrameles nudiflora; 2. Celtis philippensis var. wightii; 3. Lasioccoca comberi var. pseudoverticillata; 4. Sumbaviopsis albicans; 5. Cleidion spiciflorum; 6. Alphonsea mollis; 7. Amoora tetrapetala; 8. Tarenna sylvestis; 9. Garcinia bracteata; 10. Metadina trichotoma; 11. Alphonsea monogyna; 12. Beilschmiedia yunnanensis; 13. Combretum latifolium; 14. Ventilago calyculata; 15. Tetrastigma henryi. Limestone Vegetation in Xishuangbanna 15 There are several other communities that occur on non limestone habitats in the region but only one community was recorded in the limestone: Celtis philippensis var. wighti—Lasiococca comberi var. pseudoverticillata community. This is the commonest community on lower slopes of limestone. The upper tree layer is dominated by Celtis philippensis var. wightii, with some scattered deciduous emergents. Lasiococca comberi var. pseudoverticillata is the dominant in the second layer. Sumbaviopsis albicans and Cleidion spiciflorum are still the dominants in the third layer (Table 3). The understorey with a cover of 30-50%, consists of saplings. Fewer true shrub species were recorded. The herb layer is very undeveloped and consists of seedlings and a lot of creeping lianas. The commonest creeping lianas are Derris caudatilinba and Loeseneriella yunnanensis. Big woody lianas, such as Combretum spp., Tetrastigma spp. Ventilago spp. etc., are frequent. Epiphytes are less frequent than in the ravine seasonal rain forest. Table 3. Lasiococca comberi var. pseudoverticellata—Celtis philippensis var. wightii community Plot no.: 94-03-01 93-12-03 9203 102-13 Location: | Mengyen Yingchan Yingchan Mengyen Altitude (m): 800 1000 1060 825 Area of plot (m): 50:x30 , 20(10 x10); 5(10.x 10), 40x 60 Aspect: SW NW SW W Slope (degree): 40 5-15 10 10 Height of canopy (m): 30 30 30 i) Coverage'(%):) 0" >90 90 95 90 Novof species (25cem'd.0.h:)) | 27 25 12 11 No. of stems 102 Saf 44 142 Name of species IVI IVI IVI IVI Average Lasiococca comberi var. pseudoverticellata 67.12 1261 101 io ea! tales Celtis philippensis var. wightii 23.64 44.18 30.37 ee 48.85 Chukrasia tabularis var. velutina 15.57 11.42 30.24 — 14.26 Garuga floribunda var. gamblei 9.66 32.16 8.97 - 12.70 Tetrameles nudiflora 40.67 - ~ - 10.17 Sumbaviopsis albicans 11.81 O35 7.64 6.39 8.8 Cleidion spiciflorum 10.67 6.67 Pol 9.3 8.55 Cont: 16 Gard. Bull. Singapore 50(1) (1998) Name of species IVI IVI IVI IVI Average Alphosea mollis 10.81 ~ 18.42 - 72 Tarena sylvestis - 14.1 yee | ~ 5.42 Bombax insignis 17.00 - - — 4.25 Metadina trichotoma - pe i Be Fe - 4.03 Amoora tetrapetala - 4.99 8.34 — 3.33 Laportea sinuata 5.84 piss) - _ 3.55 Syzygium szemaoensis - ~ - 12.8 as Fortunella polyandra ~ 127 + — 3.14 Garcinia bracteata 9:91 214 + - 3.01 Polyalthia cheliensis i ie a - + - 2.93 Alphonsea monogyna 6.31 3.45 — — 2.44 Croton crassifolium 3.09 — - 6.23 pe) Symphyllia silhetiana ~ ~ ~ 8.9 225 Beilschmiedia yunnanensis 6.66 Pie Ws - - 2.20 Caryota urens - - 8.42 - 2A Walsura robusta — - Tot ~ 1.89 Ficus conccina 6.24 — - ~ 1.56 Celtis bodinieri 5.95 ~ ~ - 1.49 Lagerstroemia tomentosa 5.79 ~ - - 1.45 Dracaena cochinchinensis 4.02 1.68 1.43 Ficus racemosa 5.34 - - - 1.34 Murraya tetramera - 5.10 + - 1.28 Ficus glaberrima 2.96 ita 2 an 1.28 Ficus virens 4.59 ~ - - 115 Vitex quinata var. puberula 3.14 a ~ ~ 0.79 Syzygium melanophylla - - - 3.0 0.75 Dysoxylum lenticellata 2.94 — — - 0.74 Mitrephora thorelii 2.94 - - - 0.74 Wrightia tomentosa 2.90 - — - 0.73 Ehretia tsangii 2,89 - aes ~ 0.73 Mitrephora maingayi - 2.38 - ~ 0.6 Derris robusta — — — VPA | 0.55 Diospyros yunnanensis - i ps — - 0.44 Mitrephora wangii ~ saya - ~ 0.43 Amoora calcicola - 1.68 + - 0.42 Aglaia testicularis ~ 1.66 - - 0.42 Randia acuminatissima + — — 1:55 0.39 Xeromphis spinosa ~ — - 1.40 0.35 Total 300 300 300 300 300 Limestone Vegetation in Xishuangbanna ry 2. Tropical Seasonal Moist Forest Tropical seasonal moist forest was recognized as a vegetation type based on its profile and its occurrence mainly on the middle and upper slopes ranging from 650-1300 m altitude on limestone. It usually has two distinct tree layers with the canopy 20-25 m tall although some scattered big trees can reach more than 30 m tall in some sites. Woody lianas are abundant and vascular epiphytes with small, thick leaves are common. Buttresses and cauliflory are relatively rare. This forest type is somewhat diverse in physiognomy and floristic composition because of the great diversity of micro-habitats on the mid and upper slopes of the limestone. This vegetation type was called “monsoon forest” in Chinese botanical references (Liu, 1987). The term seasonal moist forest is preferred because the forest is not equivalent to Schimper’s monsoon forest in many ways in spite of the fact that it is affected by seasonal dryness and contains a variable percentage of deciduous trees. Two main formations can be recognized: 2a. Tropical seasonal evergreen moist forest This formation occurs on upper slopes, shady slopes or tops of lower hills with more than 90% of rock outcrops from 650 m to 1300 m alt. The forest is evergreen, with two distinct tree layers. The upper layer with a crown cover of 40-60% is 15-25 m tall and the second layer with a crown cover of 70-80% is 3-15 m tall. Woody climbers are very abundant. Vascular epiphytes with small, thick leaves are frequent. There are 50 tree species, 8 shrub species, 10 herbaceous species, 10 liana and 11 epiphyte species in the plots (cumulative area of 3500 m ). It is intermediate between lower hill seasonal rain forest and montane dwarf evergreen forest. Two main communities have been recorded: (i) Osmanthus polyneurus—Dracaena cochinchinensis community. This community usually occurs on the upper slopes of mountains or hills above 1000 m altitude (Fig. 4). Osmanthus polyneurus is the dominant species in the top layer and Dracaena cochinchinensis is usually the dominant in the second layer (Table 4). The understorey consists of saplings and creeping lianas. The commonest lianas are Loeseneriella yunnanensis and Hiptage benhalensis. Herbaceous species of the family Urticaceae, such as Procris crenata, Elatostema spp. and Pilea spp., are abundant. Lithophytes are also frequent. 18 Gard. Bull. Singapore 50(1) (1998) 30 10 20 30 M Figure 4. Forest profile of Osmanthus—Dracaena community 1. Dracaena cochinchinensis; 2. Amoora tetrapetala; 3. Tarenna sylvestris; 4. Garcinia bracteata; 5.Mallotus philippinensis; 6. Diospyros yunnanensis; 7. Syzygium balsameum; 8. Photinia angusta var. hookeri; 9. Alphonsea mollis; 10. Engelhardtia spicata; 11. Clausena excavata; 12. Osmanthus polyneurus Limestone Vegetation in Xishuangbanna 19 Table 4. Osmanthus polyneurus—Dracaena cochinchinensis community Plot no.: 9207 9208 Location: Long-pa, Mengla Long-pa, Mengla Altitude (m): 1320 1420 Area of plot (m): 22 X20 25 x 20 Aspect: E 30 NW Slope (degree): 40 25 Height of canopy (m): 20 20 Coverage of vegetetion (%): 85 90 No. of tree species (>5 cm d.b.h.): Bs: pA No. of stems: af 42 Name of species IVI IVI Average Osmanthus polyneurus 74.83 90.86 82.84 Dracaena cochinchinensis 106.49 + D320 Tarenna sylvistris 24.14 £1.53 AWE ps Syzygiun sp. -- 24.22 111 Wightia tomentosa vrei) 14.9 11.35 Sterculia villosa - 19:75 9.88 Murraya tetramera 6.99 LiGy 9.03 Schefflera glomerulata ~ 1637 8.16 Mitrephora calcarea 9.56 5.99 7.82 Engelhardtia spicata 6.96 7.46 123 Alphonsea mollis not 6.55 6.93 Myrsine semiserrata + 132 6.76 Mallotus philippinensis 7.08 D102 6.35 Garcinia bracteata + 12.3 6.15 Celtis timorensis + 11.74 5.87 Ficus curtipes _ 9.34 4.67 Garruga pinnata W271 — 4.64 Ficus orthoneura 9.26 ~ 4.64 Kopsis officinalis - 8.18 4.09 Photinia arguta var. hookeri 8.08 - 4.04 Eriolaena kwangsiensis T79 - 50 Micromelum integerrimum vat.mollisimum 7.34 _ 3.67 Diospyros yunnanensis - Tih 3159 Clausena excavata 6.96 + 35 Ulmus lanceifolia + 6.7 5.35 Fortunella polyandra ~ 3,89 EM a) Schoepfia fragrans - 5.85 2.93 Wrightia laevis - 5.62 2.81 Total 300 300 300 20 Gard. Bull. Singapore 50(1) (1998) (11) Lasiococca comberi var. pseudoverticillat—Cleistanthus sumatranus community. This occurs only on dry slopes and the tops of lower hills in Menglun between altitudes 650-800 m. There are two tree layers, of which the upper layer is 16 to 23 m tall and has a coverage of 50%; the lower layer is 5—16 m tall and has a coverage more than 70 %. Lasiococca comberi var. pseudoverticillata is the predominant species in the upper tree layer and Cleistanthus sumatranus in the lower tree layer (Fig. 5). It abuts lower hill seasonal rain forest, which is on the lower slopes and in valleys. Some deciduous emergent trees, such as Tetrameles nudiflora, Garuga pinnata, and Chukrasia tabularis, are sparsely dotted through the forest (Table 5). The understorey is similar to the former community. ’ Pay) ff Nt ¥2 SA 4 hee 4 | aL IS Figure 5. Forest profile of Lasiococca—Cleistanthus community 1. Tetrameles nudiflora; 2. Cleistanthus sumatranus; 3. Laportea urentissima; 4. Pothos repens; 5. Sumbaviopsis albicans; 6. Garruga floribunda var. gamblei; 7. Mallotus paniculata; 8. Lasioccoca comberi var. pseudoverticillata; 9. Cleidion spiciflorum; 10. Sterculia lanceolata; 11. Murraya tetramera; 12. Musa acuminata; 13. Alocasia macrorhizza; 14. Colona floribunda; 15. Unknown; 16. Celtis philippensis var. wightii; 17. Santaloides roxburghii; 18. Sumbaviopsis albicans; 19. Dracaena cochinchinensis; 20. Aglaia parviridis; 21. Saurauia tristyla;22. Mitrephora thorelii; 23. Salacia polysperma; 24. Leea crispa; 25. Mallotus philippinensis; 26. Caryota monostachya; 27. Dead tree. Limestone Vegetation in Xishuangbanna wl Table 5. Lasiococca comberi var. pseudoverticellata—Cleistanthus sumatranus community Plot no.: 950506 Altitude (m): 750 Aspect: NW Height of canopy: 22 m No. of species (25 cm DBH): 29 Name of species Cleistanthus sumatranus Croton crassifolius Lasiococca comberi var. pseudoverticellata Celtis philippensis var. wightii Garuga pinnata Tetrameles nudiflora Glycosmis ferruginea Mayodendron igneum Tarena sylvestris Alphonsea monogyna Cipadessa baccifera Beilschmeidia yunnanensis Trigonostemon lyi Ehretia tsangii Syzygium cuminii Amoora tetrapetala Ficus orthoneura Wrightia tomentosa Amoora Stellata Zanthoxylum planispinum Ficus concinna Laportea basirotunda © Amoora calcicola Mitrephora calcarea Murraya microphylla Lagestroemia tomentosa Harpullia cupanioides Stereospermum tetragonum Lepisanthes sp. Total Location: Menglun Area of plot (m): 50 x 50 Slope (degree): 30 Coverage of vegetation: >95% No. of stems: 445 % Dens. % Freq. % B.A. IVI 45.6 7.8 12.66 66.1 8.5 Tee 37.5 53.9 27.19 1d 4.9 39.9 4.7 6.3 2143 323 2.0 7.8 6.9 16.8 0.2 1.6 jb 135 1.8 6.3 0.5 8.6 1.34 6.26 0.95 7.96 135 4.69 0.27 6.31 0.67 4.69 0.79 6.15 112 4.69 0.19 6.0 0.67 S10 0.29 4.09 0.67 a3 0.04 3.84 0.45 3:13 0.09 3167 0.45 3.13 0.04 3.62 0.22 1.56 1.14 2.92 0:22 1.56 0.66 2.44 0:22 1.56 0.13 1.91 0.22 1.56 0.10 1.88 0.22 1.56 O11 1.88 0.22 1.56 0.07 1.85 022 1.56 0.07 1.85 022 1.56 0.17 1.95 0.22 1.56 0.03 1.81 0.22 1.56 0.02 1.8 O22 1.56 0.01 1.79 0.22 1.56 0.01 1.79 0.22 1.56 0.01 1.79 0.22 1.56 0.01 1.79 100 100 100 300 i) i) Gard. Bull. Singapore 50(1) (1998) 2b Tropical seasonal semi-evergreen moist forest This formation occurs on much drier lower and middle slopes and in wide valleys within the range of 600-1200 m altitude. The forest is semi-evergreen with deciduous trees making up 30-60 % of the number of species and 35-70 % of the sum of cumulative importance value index (from plot data in Tables 6 and 7). The upper layer trees are usually deciduous with umbrella crowns and rough and thicker bark. The dominant species in the upper layer is usually Bombax insignis, but in some sites Colona floribunda, Tetrameles nudiflora or Erythrina lithosperma are either dominant or co-dominant. The second tree layer is evergreen. Small woody climbers are abundant but vascular epiphytes are less frequent. There are 80 tree species, 12 shrub species, 21 herbaceous species, 25 liana and 10 epiphyte species in the plots (cumulative area of 6150 m ). Two communities have been recorded: (1) Bombax insignis-Colona floribunda community. This occurs on lower and middle dry slopes and covers a relatively large area. Bombax insignis is dominant. In some sites Colona floribunda, Erythrina lithosperma are co-dominant species in the top layer. Pistacia weinmanifolia is usually dominant in the second layer (Table 6). The understorey consists of saplings, lianas and shrubs. Common shrub species are Murraya koenigii, Colebrookea oppositifolia and Allophylus hirsutus. Common lianas are Amalocalyx yunnanensis, Porana spectabilis and Acacia pinnata. Epiphytes are rare. Table 6. Bombax insignis—Colona floribunda community Plot no.: HW9201 93-12-01 94-03-02 Location: Huiwa Yingchan Mengyen Altitude (m): 980 1200 1000 Area of plot (m): 30 x 30 25 x 30 40 x 50 Aspect: SW SE SW Slope (degree): 37 10 45 Height of canopy (m): 20 20 22 Coverage of vegetetion (%): 95 u5 fis, No. of spp.(25 cm D.B.H.) 19 2 16 No. of stems 70 38 57 Name of species IVI IVI IVI Average Bombax insignis 22.82 - 98.89 40.57 Colona floribunda 36.36 67.23 6.25 36.61 Erythrina lithosperma ame 84 40.14 ~ 21.78 Cont: Limestone Vegetation in Xishuangbanna Name of species Lagerstroemia venusta Pistacia weinmannifolia Dracaena cochinchinensis Hymenodictyon excelsum Kydia calycina Schima wallichii Celtis philippensis var. wightii Mallotus philippinensis Spondias pinnata Ehretia tsangit Phyllanthus embelica Millettia tetraptera Ficus orthoneura Grewia eriocarpa Premna fulva Sarcosperma kachinensis Derris robusta Lithocarpus microspermus Helicia cochinchinensis Litsea glutinosa Sterospermum tetragonum Tetrasmeles nudiflora Sterculia villosa Lepisanthes senegalensis Eriolaena kwangsiensis Melia toosenden Engelhartia roxburghiana Alphonsea mollis Ficus hispida Toona ciliata Wrightia tomentosa Acrocarpus fraxinifolius Ulmus lanceifolius Phoebe puwensis Cipadessa baccifera Croton crassifolius Cratoxylon cochinchinensis Mayodendron igneum Zanthoxylum planispium Dolichandrone stipulata Radermachera microcalyx Elaeocarpus varunum Beilschmiedia yunnanensis Ficus glaberrima Total IVI IVI 64.28 ue 43.48 “ 10.84 “ ~ 2013 - 9.08 10.95 12.9 _ 19.07 ~ 7.29 ~ 14.79 11.75 — Cy, — ~ 11.26 10.87 ~ _ 127 _ 9.69 9.32 + — 9.3 - 8.86 _ 8.86 _ 8.65 ~ Lge = TiS 7.07 _ _ 6:71 + 6.54 _ 6.63 6.18 _ 5.40 2 5.02 _ 4.94 = 4.76 + 4.74 a 4.62 Le + oa, 300 300 pi, Average 21.43 20.9% 13.76 11.42 9.04 9.02 8.66 7.25 6.36 5.43 4.93 24 Gard. Bull. Singapore 50(1) (1998) (ii) Bombax insignis-Garcinia bracteata community. This occurs on lower drier gentle slopes or in wide valleys. Bombax insignis as emergent trees reaches up to 35 m tall. Garcinia bracteata and Dracaena cochinchinensis are co-dominant species in the second layer (Table 7). The understorey is similar to the former community. Table 7. Bombax insignis—Garcinia bracteata’ Community Plot no.: 102-15 Location: Mengyen, Mengla Altitude (m): 800 Area of plot (m): 50 x 50 Aspect: Slope (degree): 8-12 Height of canopy: 40m Coverage of vegetation: >90% No. of species (25 cm d.b.h.): 27 No. of stems: 115 Name of species % Dens. % BA % Dens. + % BA Bombax insignis 7.83 23.08 30.91 Garcinia bracteata 14.78 Tz 22.70 Dracaena cochinchinensis 11.30 9.02 20.32 Laportea sinuata 10.43 6.87 17.30 Tetrameles nudiflora 2.61 12.43 15.04 Glycosmis ferruginea 8.69 5.69 14.38 Celtis bodinieri Bey. 5.01 11.03 Sumbaviopsis albicans 6.09 273 8.82 Dysoxylum lukii 4.35 3.45 7.80 Phaeanthus saccopetaloides 4.35 3.36 Tale Vitex quinata var. puberula 2.61 a5 5.96 Ficus racemosa 2.61 20g 5a Cleidion spiciflorum 3.48 1.42 4.90 Tarenna sylvestris 2.61 2.09 4.70 Wrightia tomentosa 2.61 2.00 4.61 Wrightia pubescens 0.87 1.36 223 Ficus virens 0.87 95 1.82 Diospyros yunnanensis 0.87 0.90 ty | Garuga floribunda var gamblei 0.87 0.82 1.69 Clausena excavata 0.87 0.82 1.69 Croton crassifolius 0.87 0.78 1,65 Hymenodictyon excelsum 0.87 0.68 1:55 Garuga pinnata 0.87 0.49 1.36 Alphonsea mollis 0.87 0.34 Pea | Unknown sp. 0.87 0.34 1.21 Cipadessa baccifera 0.87 0.27 1.14 Polyalthia cheliensis 0.87 0.27 1.14 Total 100 100 200 Limestone Vegetation in Xishuangbanna DD 3. Montane Dwarf Forest Tropical montane dwarf forest occurs on the tops of hills and summits of mountains at an altitude range between 900-1600 m. The forest has only one dwarf tree layer with canopy height of 7-15 m. Epiphyte ochids and non-vascular epiphytes are abundant. Small woody climbers are also abundant in some sites. The forest is usually characterized by Agapetes burmanica, which has swollen roots for water storage. Two formations were recognised. 3a. Montane evergreen dwarf forest This occurs on shady tops of hills and summits of relatively higher mountains above 1000 m altitude. Pistacia weinmannifolia and Myrsine semiserrata are usually dominant or co-dominant. Lithophytic orchids are very abundant on rocks; creeping climbers are also abundant. Only a representative community was plotted and analysed because of difficult field work in the very rugged topography. Photinia angusta—Pistacia weinmannifolia community. This occurs mainly on limestone summits above 1200 m altitude. Photinia angusta and Pistacia weinmannifolia are co-dominant species (Table 8). Table 8. Photinia angusta-Pistacia weinmannifolia community Plot no.: 93-12-02 Location: Ying-chan,Mengla Altitude (m): 1380 Area of plot (m): 10 x 10 Aspect: SW Slope (degree): 20 Topography: on top of a hill Height of canopy: 7 m Coverage of vegetation: 95% No. of species (25 cm d.b.h.): 4 No. of stems: 6 Name of species %Dens. %Freq. % BA IVI Photinia arguta var. hookeri 33.33 33:33 43.72 110.38 Pistacia weinmanifolia 35:95 30:33 43.72 110.38 Myrsine semiserrata 16.67 16.67 8.69 42.03 Pterospermum proteum 16.67 16.67 3.87 S21 Total 100 100 100 300 Understorey *Abund. Freq. Understorey Abund. Freq. Agapetes burmanica 32 80 Derris caudatilimbum + 40 Eria hainanensis 22 100 Peperomia heyniana + 20 Hedychium villosum + 80 Bauhinia carcinophylla + 20 Fagopyrum tataricum + 80 Kalanchoe laciniata + 20 Tetrastigma delavayi 2.1 40 Pyrrosia adnascena + 20 Pilea platanifolia + 40 Eria javanica + 20 Clematis kerrii + 40 Campylotropsis pinatorum + 20 * Braun-Blanquet’s degree of abundance 26 Gard. Bull. Singapore 50(1) (1998) 3b. Montane semi-evergreen dwarf forest The formation occurs only on some dry tops of hills. Deciduous trees make up 40 % of the number of species and 60 % of the sum of cumulative importance value index. Epiphytes are rare but woody climbers are still abundant. Also only a representative community was plotted and analysed. Ficus neriifolia—Dracaena cochichinensis community. This occurs on dry and gentle tops of hills with an altitude of 900-1200 m. Deciduous species Ficus neriifolia is dominant (Table 9). Table 9. Ficus neriifolia-Dracaena cochichinensis community Plot no.: 93-12-04 Location: Yingchan,Mengla Altitude (m): 930 Area of plot (m): 10 x 10 Aspect: S Slope (degree): 15 Height of canopy: 15m Coverage of vegetation: 85% No. of species (25 cm d.b.h.): 5 No. of stems: 14 Name of species % Dens... Freq... \FoBA A. / PI Ficus neriifolia var. trilepis 42.86 40.00 44.50 127.36 Sterculia villosa 14.29 20.00 21.14 Sas Dracaena cochinchinensis Mine aN 20.00 3.78 52.35 Celtis philippensis var. wightii 7.14 10.00 20.23 eet Pistacia weinmannifolia 7.14 10.00 10:32 ZFS Total 100 100 100 300 Understorey *Abund. Freq. Understorey Abund. Freq. Schefflera glomerulata = 2.2 80 Phymatodes cuspidata + 80 Combretum punctatum — 2.2 60 Bauhinia carcinophylla + 40 Boehmeria siamensis Ze 80 Peperomia dindygulensis + 40 Pilea platanifolia ~ 100 Hedychium villosum ~ 20 Hoya pottisii + 80 * Braun-Blanquet’s degree of abundance Species diversity From the data, tree species diversity indexes were calculated and results presented in Table 10. The highest value of diversity index is for the Pometia-Alphonsea community of ravine seasonal rain forest, which occurs mainly on bottoms of wet valleys, while the lowest value appears in the communities of the montane dwarf forest, which occurs on upper slopes Limestone Vegetation in Xishuangbanna 27 Table 10. Species diversity of limestone vegetation Forest type Plot Area |'Alt, |) Habitat)", Sl")|'NS| NIH? | E (m?) | (m) | (?) I. Tropical seasonal rain forest a. Ravine seasonal rain forest al Pometia-Alphonsea Com. | 102-16 | 2400 | 700 | Wet valley | 0-5 | 45 | 140 | 3.2627] 0.8571 terrace HW9203) 2500 | 700 | Wet slope | 25 | 23 | 118 | 2.4269) 0.774 HW9202 | 2500 | 740 | Shade 10 | 19 | 164 | 2.0464 | 0.693 lower slope a2 Pometia- Celtis Com. b. Lower hill seasonal rain forest bl Lasiococca-Celtis Com. 940301 | 2500 | 800 | Shade 40 | 27 | 102 | 2.5277 | 0.7669 slope . Tropical seasonal moist forest a. Evergreen moist forest al Lasiococca-Cleistanthus 950506 | 2500 | 750 | Sun-facing | 30 | 29 | 445 | 1.7393 | 0.5165 Com. Slope b. Semi-evergreen moist forest bl Bombax-Garcinia Com. 102-15 | 2500 | 800 | Light slope} 8-12 27 | 115 | 2.8613 | 0.8682 III. Montane dwarf forest a. Evergreen dwarf forest 931202 | 100 | 1380) Top of hills} 20 |4 |6 | 1.3297/0.9592 Upper hill b. Semi-evergreen dwarf 931204 | 100 | 930 | slope 155) Sh bh WA FOL 0.855 forest | Sl: Slope; NS : Number of species (25 cm dbh); NI: Number of individuals (=5 cm dbh); H’: Shannon-Wiener’s diversity indices (Shannon-Wiener, 1949); E: Evenness indices of Pielou (1966) and tops of hills. —The communities on sun-facing steep slopes, which are usually consociations or associations with co-dominant species, such as the Lasiococca comberi var. pseudoverticillat -Cleistanthus sumatranus community, have relatively lower diversity index values than the communities on sun-facing gentle slopes. Compared with seasonal rain forests on non-limestone, the limestone seasonal rain forests show lower index values (Table 11) and this agrees with Cao’s results (Cao and Zhang, 1997). The communities of seasonal 28 Gard. Bull. Singapore 50(1) (1998) Table 11. Comparison of species diversity between the limestone seasonal rain forest and the seasonal rain forest on non-limestone Forest type Habitat 1. Tropical seasonal rain forest on limestone a. Ravine seasonal rain forest | 102-16 Wet valley terrace HW9203) 2500 |700 | Wet slope HW9202) 2500 |740 | Shade lower slope b Lower hill seasonal rain =| 940301 =| 2500 |800 | Shade slope forest | it Tropical seasonal rain forest on non-limestone a. Ravine seasonal rain forest | 940102 |2500 |650 | Wet valley slope | 5-10) 49 |108 | 3.586 | 0.9263 940103 |2500 |675 | Wet valley slope | 30 |57 |194 | 3.573 | 0.8727 940101 {2500 |700 | Wet valley slope | 25 |48 |96 | 3.599 | 0.9297 b Lower hill seasonal rain =| 931206 = |2500 |650 | Lower hill slope | 10 {52 | 182 | 3.3765 | 0.8545 forest 9201 2500 |680 | Lower hill slope | 30 |46 |207 | 3.1594 | 0.825 SI: slope; NS : Number of (25 cm dbh); NI: Number of individuals (25 cm dbh); H’: Shannon-Wiener’s diversity indices (Shannon-Wiener, 1949); E: Evenness indices of Pielou (1966) rain forest on non-limestone have almost identical values for diversity index and evenness, while the communities of limestone seasonal rain forest show a clear disparity in the values. This implies that the limestone seasonal rain forest has higher community diversity than the rain forest on non-limestone substrates. Acknowledgements This project was funded by the Chinese Academy of Sciences (part of the project KZ951-A1-104-03) and the Yunnan Science & Technology Committee. I thank Prof Xu Zaifu, Director of Xishuangbanna Tropical Botanical Garden and Prof Wu Zheng-yi, Prof Zhang Hong-da, my academic advisors, and my colleagues Mr Cao Min and Mr Zhang Jianhou for their great help with my research. I also wish to acknowledge the Limestone Vegetation in Xishuangbanna 29 Department of Plant Sciences, University of Cambridge and especially the Ecology Group in the Department for help of various kinds. I particularly thank Dr E. Tanner and Dr P.Grubb for their great help in analyzing my data and writing this paper during my year’s stay at the University of Cambridge as a visiting scholar. Finally, I particularly thank Dr T.C.Whitmore who has greatly supported and helped me in my research . References Braun-Blanquet, J. 1932. Plant Sociology (transl. G.D.Fuller and H.S.Conard), McGraw-Hill. Comp. New York, pp. 52-58. Cao, M. and J. Zhang. 1997. Tree species diversity of tropical forest vegetation in Xishuangbanna, SW China. Biodiversity and Conservation. 6:995-1006. Curtis, J.T. and R. P. McIntosh. 1951 An upland forest continuum in the prairie-forest border region of Wisconsin. Ecology 32:467-496. Jin, Z. Z. and X. K. Ou. 1997. The diversity features of plant community types in the tropical rain forest vegetation of Xishuangbanna, Yunnan. Acta Bot. Yunn. Suppl. TX:1-30 (in Chinese with English abstract). Liu, L.H. 1987. Rain forest. In: C.Y.Wu (ed.) Vegetation of Yunnan. Science Press, Beijing, pp.97—143.(in Chinese). Pielou, E.C. 1966. The measurement of diversity in different types of biological collections. Theor. Biol. 13: 131-144. Qu, Z.X. 1960. The vegetation of Yunnan Nature Reserves. Journ. Yunnan Univ. (Nat. Sci.) 1:1-4. (in Chinese). Schimper, A. F. W. 1903. Plant-geography upon a Physiological Basis. Oxford University Press, Oxford. Shannon, C.E. and W. Wiener. 1949. The Mathematical Theory of Communication. Urbana: Univ. Illinois Press. Wane Zhu, and B.'Li. 1997. Vegetation on limestone in Xishuangbanna, southwest China. Guihaia 17(2): 101-117. (in Chinese with English abstract). Wu, C. Y. 1980. Vegetation of China. Science Press., Beijing, pp. 363-397 (in Chinese). 30 Gard. Bull. Singapore 50(1) (1998) Wu, C. Y. 1991. The areal-types of Chinese genera of seed plants. Acta Bot. Yunn. Supp. IV. (in Chinese with English abstract). Xu, Y. C. and H. Q. Jian (eds.) 1987. Reports of Multidisciplinary Investigation into Nature Reserves of Xishuangbanna. Yunnan Sci. Press. Kunming (in Chinese). Zhu, H. 1992. Tropical rain forest vegetation in Xishuangbanna. Chinese Geographical Science 2: 64-73. Zhu, H. 1997. Ecological and biogeographical studies on the tropical rain forest of south Yunnan, SW China with a special reference to its relation with rain forests of tropical Asia J. Biogeogr. 24:647-662. Zhu, H., H. Wang, and B. Li. 1996. La Wa ‘ in 7 J . hy 1 iV) 1 vis " ¥ Ny J ite A ] f -, i I t : ’ ye P iy F) a yeu A.) are Pat OA li, a riahiy ’ Wet Opt yr ‘ey a pal i2 of ane ar e's ih ik ON vir iy af ’ ri re tat Raed ae ok Pita MRP A ebiaa saa Sete Meee a ny Oe nee | et Ape atin Dy ied a i bar at rj 7 uy ) ‘ he rae { i i i 1 li y 2 ‘5 al \ : 4 Wa ae ‘ oo Le er ih prison savte y : s ' ny j aS ry ya ; oc : rt 7 ip My - j Timur iat hy } ri ia Wea aa 4 bh ie fia Ee i4 ’ ' f te pi ' , i % Att rf Neh Nes ' } Fo i ré ey Ls oh nt 4 PAI 4h ts J io Pe Tehran ease UL ® J i Kil Ye f : iy 4 fs vs Ram eh ATR ey Pin) Aleta te toy van € Oy ean eee ah Gardens’ Bulletin Singapore 50 (1998) 39-41. Nomenclatural Changes for Four Malayan Species in Phrynium (Marantaceae), Solanum (Solanaceae), Stachyphrynium (Marantaceae) and Boesenbergia (Zingiberaceae) I.M. TURNER Center for Ecological Research Kyoto University Shimosakamoto Otsu 520-0105 Japan Abstract A new name is provided to substitute for a later homonym in Marantaceae. Phrynium venustum I.M. Turner, nom. nov., replaces Phrynium gracile Holttum. Solanum maingayi Kuntze (Solanaceae) is shown to be the correct name for what has generally been referred to as Solanum sarmentosum Nees. Stachyphrynium cylindricum K. Schum. (Marantaceae) and Boesenbergia flava Holttum (Zingiberaceae) have to be considered new names because they were published as new combinations based on later homonyms. These illegitimate names, Phrynium cylindricum Ridl. and Gastrochilus flavus Ridl., are lectotypified. A new name for a Malayan Phrynium In his monograph of the Malayan Marantaceae, Holttum (1951) described a new species from the freshwater swamp forests of Johore as Phrynium gracile. Unfortunately, this combination had already been employed by Schumann for a species from New Guinea. A new name is provided for the Malayan plant. Phrynium venustum 1.M. Turner, nom. nov. Phrynium gracile Holttum, Gard. Bull., Singapore 13 (1951) 282, nom. illegit., non K. Schum. (1905). Type: Peninsular Malaysia, S.E.JJOHORE, Mawai-Jemaluang Road, Sungai Kayu Ara, E.J.H. Corner S.F.N. 29981, 5 January 1935 (holotype, SING!; isotype, K!). Holttum /oc. cit. gives the collecting locality of the type (Corner S.F.N. 29981) of this species as an uncertain site in Southeastern Johore, and refers to another, unnumbered, Corner collection from Sungei Kayu Ara. However, S.F.N. 29981 specimens at SING and K are both clearly labelled as originating from Sungei Kayu Ara. This anomaly is not easily explained. 40 Gard. Bull. Singapore 50(1) (1998) The correct name for Solanum sarmentosum The prickly, prostrate Malayan nightshade generally known as Solanum sarmentosum Nees cannot correctly be referred to as such, as the combination was used earlier by Lamarck. Ridley, in his Flora of the Malay Peninsula synonymized Solanum maingayi Kuntze to S. sarmentosum Nees. Inspection of the type of Kuntze’s species at Kew confirmed the correct placement by Ridley, and hence means that the Malayan plants must be referred to as Solanum maingayl. Solanum maingayi Kuntze, Rev. Gen. Pl. 2 (1891) 454. Type: Peninsular Malaysia, MALACCA, A.C. Maingay 1158, 1867 (holotype, K!). Solanum sarmentosum Nees, Trans. Linn. Soc. London 17 (1837) 58, nom. illegit., non Lam. (1794). The correct citation for Stachyphrynium cylindricum Stachyphrynium cylindricum is a member of the Marantaceae found growing on limestone in the Malay Peninsula (Holttum 1951). The combination was published by Schumann based on Ridley’s Phrynium cylindricum. However, this is an illegitimate later homonym. The International Code for Botanical Nomenclature Article 58.1 allows a new combination based on a later homonym to be considered as an avowed substitute (nomen novum) or a new name. As Schumann clearly intended to make a new combination based on the type of Phrynium cylindricum Ridl., Stachyphrynium cylindricum should be considered a nomen novum for the former. Ridley did not designate a holotype from among the two collections of Phrynium cylindricum he cited. I therefore take this opportunity of lectotypifying the species. Stachyphrynium cylindricum K. Schum., nom. nov., Pflanzenr. 4 (1902) 49; Holttum, Gard. Bull., Singapore 13 (1951) 278. Phrynium cylindricum Ridl., J. Straits Branch Roy. Asiat. Soc. 32 (1899) 178, nom. illegit., non Roscoe (1828). Syntypes: Peninsular Malaysia, Perak, Ipoh, C. Curtis 3318, August 1898 (SING!, 2 sheets); Kwala Dipang, H.N. Ridley 9787, 1898 (lectotype, selected here, K!). The correct citation for Boesenbergia flava An identical nomenclatural circumstance to the previous example surrounds the Malayan endemic ginger Boesenbergia flava. Holttum transferred the Nomenclatural Changes for Malayan Species 41 species from its original position in Gastrochilus, where it was a later homonym. Therefore, the combination must be considered a nomen novum with the same epithet as the illegitimate basionym. Gastrochilus flavus Ridl. is lectotypified. Boesenbergia flava Holttum, nom. nov., Gard. Bull. Singapore 13 (1950) 113. Gastrochilus flavus Ridl., Fl. Malay. Penins. 4 (1924) 248, ‘flava’, nom. illegit., non (Hook.f.) Kuntze (1887). Syntypes: Peninsular Malaysia, PERAK, Batang Padang, C. Curtis s.n. (lectotype, selected here, K!); Bujong Malacca, H.N. Ridley s.n., September 1898 (SING!); Bukit Kepaiyang, H.N. Ridley s.n., February 1904 (SING!). Acknowledgements The Director of the Singapore Botanic Gardens and the Keeper of the Herbarium, Royal Botanic Gardens Kew, kindly allowed the author access to the collections in their respective herbaria. Martin Cheek, Jill Cowley and Alan Paton are thanked for their help at Kew. An anonymous reviewer and Dr K. Gandhi provided invaluable assistance with nomenclature. References Holttum, R.E. 1951. The Marantaceae of Malaya. Gardens’ Bulletin, Singapore 13: 254-296. eyo aa ’ int a ie i HY } h . 7 MNT R i. fhe | i be We CT! at : Oar) Din P a%,° ; sal A Pair a mh ; wy) a, \) a). Es jag ' vp ace yee, ie agiois 2 if . were ally ’ P 4 ¥i cs fr he = Ry ~ , a\s Rese : ah it a Gardens’ Bulletin Singapore 50 (1998) 43-48. Begonia lazat (Begoniaceae), a New Culinary Begonia from Borneo Reza Azmt! AND RutTH KIEWw? 'WWE-Malaysia, Locked Bag No. 911, Jalan Sultan P.O., Petaling Jaya, Malaysia “Singapore Botanic Gardens, 259569 Singapore Abstract A striking new large-fruited begonia is described from the floodplain forest in the lower reaches of the Kinabatangan River, Sabah. Although rare, this begonia is known by some older local residents as a culinary delicacy when eaten with prawns and chilli. Introduction At just over 560 km long, the Kinabatangan River is the largest river in Sabah and, where it begins its lower course, flows through a floodplain that remains one of the largest forested floodplains in Malaysia (Figure 1). This floodplain is one of the most important conservation areas in Sabah as well as being a major nature tourist destination where the proboscis monkey, a Bornean endemic, is reliably and highly visible along some of its tributaries. Besides supporting a variety of wildlife, the Kinabatangan floodplain also includes an array of lowland habitats - remnant dipterocarp forest, freshwater swamp forest and open swamps, riverine forest, ox-bow lakes and limestone outcrops (Reza Azmi, 1996). In addition, a truly bewildering richness of natural resources characterises this vast floodplain. Reza Azmi (1996) recorded over a hundred useful plants with 22 species used for structural purposes (house and boat building, fencing, etc.), 12 for firewood, 35 as food, 73 in traditional medicine, and a further 27 species for miscellaneous uses (making fish-traps, parang (Malay=machete) handles, for ceremonial or pagan practices, etc.). There is, however, increasing pressure for forest conversion from the expansion of agricultural estates (Payne, 1989). The vulnerability of the floodplain region has prompted the Sabah Government to recognise the Kinabatangan floodplain region as one of the highest priority habitats for conservation. These floodplain habitats were surveyed by WWF-Malaysia in 1994. With the help of local residents, an initiative was started to document the uses of local plants by village residents living near the remnant floodplain Gard. Bull. Singapore 50(1) (1998) aa S49AIa speo. uofew uolzejueld ue}3e4 yaosvs (@AsaSaN Je1d4aLWUILUOD) 9A49S0N 3S9404 (@A49S9Y UO0IZI93041¢ 40 aj6une ul6a1A) BA49S9¥ 3S9404 Aaenjoues IsIPIIM pasodoid INN A ‘seore pojoojo1d sofeu pue Ayrpeso] 9d4q oy} JO UOTed0] oY) SUIMOYs UOISeI ULSURJeQeUTYy TOMO] OU, *T aansry os TC ge 2 es x LY \ Begonia lazat 45 forests (Reza Azmi, 1996). The new begonia described here was discovered during this study with the help of two knowledgeable elders from Buang Sayang village, which lies near the banks of the lower Kinabatangan River (Figure 1). The wife of one of our local collectors recounted how leaves of this plant are used as a vegetable and that it is delicious when cooked with prawns and chilli. Despite this begonia being known to the informants, attempts to relocate the plant or other individuals of it a year later proved fruitless and no other specimens have been discovered since the first collection. It is possible that this species may have a narrow habitat preference as it appears to be restricted to early secondary growth of indundated forest, the habitat where it was first discovered. Begonia lazat Kiew & Reza Azmi sp. nov. Holotype: Reza Azmi RA206 near Kampung Buang Sayang, Kinabatangan District, Sabah, Borneo (SAN, unicate, fruits in spirit). Figure 2. Ab aliis speciebus Begoniae Borneensibus in sectione Petermannia fructibus magnis bene distincta, sed combinatione petiolis longitudine laminis aequante, laminis latioribus quam longioribus et textura in sicco tenuissimis, inflorescentia compacta et fructu basi attenuatis. Cane-like glabrous begonia branching from base. Stems erect, reddish, slightly fibrous, 75—100 cm tall, 0.4 mm thick in dried state. Leaves alternate. Petiole as long as lamina, 9.5—12 cm long in fully grown leaves, dark red. Lamina of young leaves ovate (longer than broad) and almost symmetrical becoming suborbicular and slightly unequal when fully expanded, 9.5-12 by 12—12.5 cm, basal lobes rounded, 44.5 cm long (not overlapping), margin minutely serrulate, apex shortly acuminate, main veins 5, radiating from base, bifurcating two to three times before reaching the margin with up to 2 minor veins in basal lobes, veins impressed above and prominent beneath, in life glossy dark green with large and small silvery blotches arranged more or less in a line between veins, pale green beneath, succulent in life, drying tissue-paper thin and transparent. Stipules pale green, elliptic, up to 20 by 8 mm, entire, apex apiculate, drying thinly papery, caducous on the lower nodes. Inflorescence axillary, bisexual, a compact panicle with total length c. 2.25 cm, peduncle stout 1.5-7 mm long with 2-3 female flowers at base, male rachis erect c. 12-18 mm long lateral branches up to 3 mm long with dense clusters of male flowers. Bracts pale green, broadly ovate, 14 by 10 mm, partially enclosing the inflorescence, bracteoles similar in shape 46 Gard. Bull. Singapore 50(1) (1998) and decreasing in size towards apex of inflorescence. Female flowers: pedicel in flower 3 mm long, straight with the flower held horizontally, in fruit 5—7 mm becoming thickened and recurved so the fruit is pendant; ovary pale green, cylindric, c. 10 mm long and 5 mm wide, 3-loculate, placentas bilamellate, wings 3 and isomorphic; tepals white, 5, broadly elliptic, the largest 8.5 by 5 mm, apex rounded, entire; styles 3, bifid, falling in fruit. Male flowers: numerous, small, pedicels slender up to 11 mm long; tepals 2, glabrous, isomorphic, rosy red outside, broadly oblong, 6.5—8 by 5—5.5 mm, Figure 2. Begonia lazat. A Habit (x 0.3), B Inflorescence with male flowers (x 0.6), C Male bud (x 2), D Open male flower (x 1.3), E Androecium (x 3.3), F Stamen (x 7), G T.S. Ovary (x 0.6), H Seed (x 13). Begonia lazat 47 entire, apex rounded; androecium c. 50 stamens, sub-spherical, staminal column c. 4 mm long, filaments c. 1 mm long, anthers narrowly oblanceolate, 1 by 0.5 mm, apex trunctate. Capsule pale green ripening brown and papery, obconic, 3.5 cm long and 2.75 cm wide, wings thin c. 9 mm wide, base narrowing into pedicel, apex trunctate by the rounded wing angle. Seeds broadly oblong, less than twice as long as broad, c. 0.3 mm long. Distribution: Known only from type locality at Kampung Buang Sayang i 30'N 117° 30° E). Habitat: Growing on a lightly shaded, low earth bank in disturbed seasonally innundated forest close to Kinabatangan River, not common. Apparently intolerant of competition as it is eliminated by subsequent growth of a shrubby layer. Notes: Begonia lazat is one of several cane-like Begonia species with large fruits 3.5 cm long, such as B. erythrogyna Sands and B. tawaensis Mertr., which belong to section Petermannia. Like B. erythrogyna, it is atypical of this section in the male flower possessing two instead of four tepals. However, it is unique among Bornean begonias in possessing the following combination of characters: large fruits, which narrow towards the base and have a short stalk; extremely thin glabrous leaves, which are broader than long; petioles as long as the lamina; and inflorescences where the terminal part with small male flowers is extremely short. ‘Lazat’ (Malay=delicious) indicates the use of its leaves as a sourish vegetable, which together with chilli are cooked with prawns. It is not unique in this respect as several other Malesian begonia species, which have glabrous, tender leaves, are also used, particularly for flavouring fish and prawn dishes. Acknowledgements The field work, which led to the discovery of this species, was conducted under WWF-Malaysia Project MYS304/94 entitled ‘Conservation of the Kinabatangan Floodplain Flora, Habitats and the Role of Local Communities’. RA wishes to thank the Forest Research Centre, Sepilok, and SAN, who freely provided facilities to WWF during the period of study. RK thanks the Curators of the herbaria at BM, E, K, SAN and SING for permission to examine specimens in their care and for funding from the Ministry of Science, Technology and the Environment under IRPA Programme grant 08-02-04-025. The authors also thank Mdm P.H. Yap for the accomplished drawing. 48 Gard. Bull. Singapore 50(1) (1998) References Payne, J. 1989. A Tourism Feasibility Study for the Proposed Kinabatangan Wildlife Sanctuary. WWF-Malaysia. Reza Azmi. 1996. Protected areas and rural communities in the Lower Kinabatangan region of Sabah. Sabah Society Journal. 13: 1-32. Gardens’ Bulletin Singapore 50 (1998) 49-57. The Types and Original Specimens of Published Names of Mosses Preserved in the Herbarium of Singapore Botanic Gardens (SING) BENITO C. TAN School of Biological Sciences, National University of Singapore Singapore 119260 And The Herbarium, Singapore Botanic Gardens Cluny Road, Singapore 259569 Abstract A total of 51 types and original specimens of published names of mosses, mostly collected from West Malesia, are preserved at the herbarium of Singapore Botanic Gardens (SING). Information about the locality, collector and collector’s number, date of collection, nomenclatural status, and the currently accepted name, for each of the types and original specimens are presented. Introduction The well known herbarium of Singapore Botanic Gardens (SING) has a small, less known and little used collections of bryophytes. The specimens were collected mainly from Peninsular Malaya, Singapore, Sabah formerly British North Borneo and Sarawak. The entire bryophyte collections consist of about 10,000 packets, of which 51 are types and original material of published names of moss taxa. In addition, there are 13 moss specimens with unpublished herbarium names and marked as new species by the eminent English muscologist, Mr. H. N. Dixon, who also identified many of the moss collections at SING. Furthermore, a nearly equal number of Specimens were indicated to be types of various hepatic taxa. The information on the hepatic types at SING will be dealt with in a separate report. In preparing the list of moss types kept at SING, I have endeavored to confirm the nomenclatural status of all the specimens marked as types. Because many of the new species were described by Dixon, the holotypes are, presumably, with The Natural History Museum in London (BM) where the herbarium of Dixon was relocated after his death in 1944. The duplicate specimens at SING, even though represented by a larger quantity in several instances, can only be the isotypes or isoparatypes. It is interesting to note 50 Gard. Bull. Singapore 50(1) (1998) that some type packets at SING include handwritten field data not published in the species’ protologue. I also include in the present listing original materials of a handful of nomina nuda inadvertently published by Dixon based on the moss collections at SING. Often, this original material is needed to elucidate the intent of the author or to resolve the taxonomic enigma of a “nomen nudum”. Some have subsequently been shown to represent distinct species. Lastly, to help users of the type collections at SING, I have indicated the currently accepted name as a note under the binomial of the moss taxon concerned. Catalogue of Species Acroporium macroturgidum Dix. [Isotype] Malaysia, Pahang, G. Tahan, 28 Aug 1928, RE Holttum 20920. Acroporium surculare Dix. | [Isolectotype] Malaysia, Perak, Bidor Road, Tapah, Nov 1908, HN Ridley £55: Note: Recombined as Isocladiella surcularis (Dix.) B.C. Tan and H. Mohamed, the name of this species was lectotypified by Tan and Mohamed (1990). Breutelia kinabaluensis Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, 9000 ft, 16 Nov 1931, RE Holttum 25337. Calymperes carrii Dix. [Isotype] Papua New Guinea, Kanosia, Feb 1935, CE Carr 11473. Note: This taxon was treated as a synonym of Calymperes crassinerve (Mitt.) Jaeg. in Reese et al. (1986). The isotype was distributed as a number of Musci Selecti et Critici, Ser. WI , 1936. Chionoloma latifolium Dix. [Isotype] Malaysia, Langkawi, Pulau Dayang Bunting, 23 Aug 1925, RE Holttum 15130. Cladopodanthus microcarpus Dix. [Isotype] Malaysia, Sarawak, Mt. Dulit, Ulu Koyan, 22-25 Sep 1932, PW Richards M2034. Mosses at SING 51 Clastobryella asperrima Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, Pakka, 15 Nov 1931, RE Holttum 25646. Dicranoloma angustifrondeum Dix. [Isotype] Malaysia, Sabah, Tenompok, 4000 ft, 17 Jun 1925, CM Enriquez LOTTI. Note: A synonym of Dicranoloma braunii (Dozy & Molk.) Par. (Tan 1989). Dicranoloma brevicapsularis Dix. [Isotype] Malaysia, Pahang, G. Tahan, 12 Jun 1922, Mohd. Haniff Nur 7915 a. Note: A synonym of Dicranoloma billarderi (Brid. Ex Anon.) Par. (Tan 1989). Dicranoloma euryloma Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, Tenompok, Lumu-Lumu, 12 Nov 1931, RE Holttum 25633. [Isoparatype] Malaysia, Sarawak, Mt. Dulit, Ulu Koyan , 15 Sep 1932, PW Richards M1853. Note: A synonym of Dicranoloma assimile (Hampe) Ren. (Tan 1989). Diphyscium rhynchophorum Dix. [Isotype] Malaysia, Sarawak, Mt. Dulit, Ulu Koyan, 15 Sep 1932, PW Richards M1868. Note: The specimen bears the annotation of M. Manuel in 1978 as an isotype. According to Hyvonen (1989a), this species is a synonym of Diphyscium loriae C. Muell. Distichophyllum sinuosulum Dix. [Isotype] Malaysia, Perak, Birch’s Hill, 3 Mar 1924, [TH Burkill 12606. Note: A synonym of Distichophyllum cirratum Ren. & Card. (Mohamed and Robinson 1991). Endotrichella formosa Dix., nom. nud. [Original material] Malaysia, Pahang, Kota Gelanggi, 4 Aug 1929, MR Henderson 22405. Note: A synonym of Garovaglia elegans (Dozy & Molk.) Bosch & Sande Lac. (Mohamed and Tan 1988). 52 Gard. Bull. Singapore 50(1) (1998) Fissidens albolimbatus Dix. [Isotype] Malaysia, Sarawak, Long Kapa, Mt. Dulit, Ulu Tinjar, 31 Aug 1932, PW Richards M1587. Note: A synonym of Fissidens ceylonensis Dozy & Molk. fide Tan and Iwatsuki (1989). Fissidens pachyphyllus Dix. [Isotype] Malaysia, Sarawak, Mt. Dulit, Ulu Koyan, 22 Sep 1932, PW Richards M2039. Fissidens perpellucidus Dix. [Isoparatype] Malaysia, Sarawak, Mt. Dulit, Ulu Tinjar, 29 Aug 1932, PW Richards M1589. Note: A synonym of Fissidens pallidus Hook. f. & Wils. fide Tan and Iwatsuki (1989). Forsstroemia rigida Dix. [Isotype] Papua New Guinea, above Port Moresby, Boridi, Nov 1935, CE Carr 13559. Note: The specimen was distributed as a number of Musci Selecti et Critici, Ser. V, 1938. The species was synonymized with Neolindbergia vitiensis (Bartr.) Enroth by Akiyama et al. (1991) who also reported the Carr specimen at BM as the lectotype. However, only one specimen (Carr 13559) was mentioned in the protologue (Dixon 1943), so there is no need for lectotypification. Hypnodendron copelandii Broth. var. latifolium Dix. [Holotype] Malaysia, Sabah, Mt. Kinabalu, Lamu-Lamu, 5000 ft, 18 Jun 1925, CM Enriquez 18146. Note: In the protologue, Dixon (1935: 96) indicated the holotype of his new variety to be at SING. The specimen was annotated by A. Touw in December of 1968 as Hypnodendron diversifolium Broth. & Geh. (Touw 1971). Hypnodendron wrayi Broth. ex Dix., nom. nud. [Original material] Malaysia, Perak, G. Batu Puteh, 4500 ft, L Wray Jr. 301. Note: Specimen annotated by A. Touw in December of 1968 as Hypnodendron diversifolium Broth. & Geh. (Touw 1971). Leptodontium kinabaluense Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, 13,400 ft, 14 Nov 1931, RE Holttum 25685. Mosses at SING D5 [Isoparatype] Ibid, RE Holttum 25688. Note: A synonym of Leptodontium flexifolium (Dicks.) Hampe (Zander 1993). Leptodontiopsis orientalis Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, 1,500 ft, 14 Nov 1931, RE Holttum 25668. [Isoparatype] Ibid, RE Holttum 25689. Mastopoma papillatum Dix., nom. nud. [Original material] Malaysia, Kelantan, G. Sitong, 2600 ft, 6 Mar 1924, Mohd. Haniff Nur 12234. Note: The name was published invalidly by Dixon in 1926. Macromitrium ochraceoides Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, between Kamborangah and Pakka, 7,200-10,200 ft, 13 Nov 1931, RE Holttum 25481. [Isoparatype] Ibid, below Pakka, 10,200 ft, 15 Nov 1931, RE Holttum 25663. Piloecium acroporioides Dix. [Isotype] Malaysia, Sarawak, Marudi (Claudetown), 300 m, Nov 1932, PW Richards M2672. Pogonatum euryphyllum Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, near Kamborangah, 7,200 ft, 13 Nov 1931, RE Holttum 25644. [Isoparatype] Ibid, Tenompok, 4,700 ft, 11 Nov 1931, RE Holttum 25344. Note: A synonym of Pogonatum cirratum (Sw.) Brid. ssp. macrophyllum (Dozy & Molk.) Hyvonen (Hyvoénen 1989b). Pseudoracelopus borneensis Dix. [Isotype] Malaysia, Sabah, Bettolan, near Sandakan, 21 Aug 1927, CB Kloss 19171. Note: A synonym of Pogonatum iwatsukii Touw (Touw 1986). Rhaphidostegium complanatulum Dix., nom. nud. [Original material] Malaysia, Perak, Padang Rengas reservoir, 18 Jan 1925, Mohd. Haniff Nur 14981. Note: The name was published invalidly by Dixon in 1926. Rhaphidostichum aquaticum Dix. [Isotype] Malaysia, Sarawak, G. Matang, 500 ft, 22 Jan 1930, RE Holttum 23165. 54 Gard. Bull. Singapore 50(1) (1998) Note: The species is now known as Papillidiopsis aquaticum (Dix.) Buck and B.C. Tan. Sclerohypnum riparium Dix. [Isotype] Malaysia, Pahang, Tahan River, 24 Aug 1928, RE Holttum 20089. Note: The species is now known as Sclerohypnumi littorale (Hampe) B.C. Tan (Tan 1991). Sphagnum cuspidatulum C. Muell. var. trengganuense A. Johnson [Holotype] Malaysia, Trengganu, G. Padang, 4,000 ft, Jun 1937, Moysey and Kiah 31023. Note: Eddy (1977) listed this trinomial under the synonymy of Sphagnum cuspidatum Hoffm. subsp. subrecurvum (Warnst.) Eddy. Sphagnum flaccidifolium Dix. ex A. Johnson [Holotype] Malaysia, Selangor, Telok Forest Reserve, 7" Milestone Klang, Carrick 500. [Paratypes] Malaysia, Selangor, Telok Panglima Garang, May 1933, RE Holttum 28317; Indonesia, Sumatra, near Palan Borae, 24 Feb 1933, RE Holttum 28127. Note: Annotated by A. Eddy in 1968 as Sphagnum subrecurvum var. The taxon is subsequently treated by Eddy (1977) as Sphagnum cuspidatum Hoffm. ssp. subrecurvum (Warnst.) Eddy var. Mae Mt Cees (A. Johnson) Eddy. Sphagnum holttumii Dix. ex A. Johnson [Holotype] Malaysia, Pahang, G. Tahan, 5 000 ft, 30 Aug 1928, RE Holttum 20906. [Paratypes] Malaysia, Pahang, G. Tahan, ca 5,000 ft, 30 Aug 1928, RE Holttum 20908; ibid, ca 6,000 ft, 31 Aug 1928, RE Holttum 20909; ibid, 6,000 ft, 31 Aug 1928, RE Holttum 20910; ibid, 3500-4500 ft, 28 Aug 1928, RE Holttum 20916; Cameron Highlands. 4800 ft, 1 Apr 1930, RE Holttum 23300; Johore, G. Pantai, 1600 ft, 9 Jun 1930, EJH Corner 23207. Note: All the above mentioned specimens bear the unpublished herbarium name, Sphagnum kedahense Dix. The species is treated as a synonym of Sphagnum perichaetiale Hampe (Eddy 1977). Sphagnum roseotinctum A. Johnson [Holotype] Malaysia, Kelantan State, G. Sitong, 2,600 ft, 6 Mar 1924, Mohd. Haniff Nur 12244. Note: The specimen was first reported by Dixon in 1926 as Sphagnum kelantanense, a nomen nudum. Johnson (1958) described it as a new species Mosses at SING 55 and Eddy (1977) reduced it to one of the many synonyms of Sphagnum perichaetiale Hampe. In the protologue (Johnson 1958), two paratype specimens of S. roseotinctum collected by Spare (no. 1439 and 1430) were mentioned, but I cannot find these two specimens at SING at present. Stephanodictyon borneensis Dix. [Isotype] Malaysia, Sabah, Mt. Kinabalu, Lobang, 19 Nov 1931, RE Holttum 25637. Note: The genus was combined with Trichostomum by Zander (1993) and the species is now known as T. borneensis (Dix.) Zand. Symphyodon complanatus Dix. [Isotype] India, Assam, watershed of Egar to Serpo, 5500 ft, 23 Jan 1912, IH Burkill 36208 (ex Herb. Hort. Bot. Calcuttensis). Syrrhopodon perakensis Dix. [Isotype] Malaysia, Perak, Lumut, Dindings, 1896, HN Ridley 449. Note: Annotated by H. Mohamed in September of 1983 as Syrrhopdon fallax Lac., a synonym of Syrrhopodn aristifolius Mitt. As asynonym, this taxon was not mentioned by Mohamed and Reese (1985) in their taxonomic revision of the genus for Malaysia and adjacent regions. Taxithelium bilobatum Dix. [Isotype] Malaysia, Perak, Bujang, Malacca, 1891, HN Ridley 739. Note: The species is now known as Glossadelphus bilobatus (D1x.) Broth. _ The isotype packet at SING has the collection number written as Holttum 739, probably an error. In the protologue, the holotype was indicated to be at Mitten Herbarium (Dixon 1924) Tayloria borneensis Dix. [Isotype] Malaysia, Sabah, Tenompok, 11 Nov 1931, RE Holttum 25329. References Akiyama, H., T. Koponen and D. H. Norris. 1991. Bryophyte flora of Huon Peninsula, Papua New Guinea. XLV. Neolindbergia (Prionodontaceae, Musci). Acta Botanica Fennica. 143: 77-89. Dixon, H. N. 1924. New species of mosses from the Malay Peninsula. Bulletin of Torrey Botanical Club. 51: 225-259. Dixon, H. N. 1926. A list of the mosses of the Malay Peninsula. Gardens’ Bulletin, Straits Settlements. 4: 1-46. 56 Gard. Bull. Singapore 50(1) (1998) Dixon, H. N. 1935. A contribution to the moss flora of Borneo. Journal of Linnean Society of Botany. 50: 57-140. Dixon, H. N. 1943. Alpine mosses from New Guinea. Farlowia. 1: 25-40. Eddy, A. 1977. Sphagnales of tropical Asia. Bulletin of the British Musuem (Natural History), Botany. 5: 359-445. Hyvonen, J. 1989a. The bryophytes of Sabah (North Borneo) with special reference to the BRYOTROP transect on Mount Kinabalu. VI. Polytrichaceae and Buxbaumiaceae (Bryopsida). Willdenowia. 18: 569- 589. Hyvonen, J. 1989b. A synopsis of genus Pogonatum (Polytrichaceae, Musci). Acta Botanica Fennica. 138: 1-87. Johnson, A. 1958. The genus Sphagnum in Malaysia. The Gardens’ Bulletin Singapore. 17: 312-324. Mohamed, H. and W. D. Reese. 1985. Syrrhopodon (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist. 88: 223- 254. Mohamed, H. and H. Robinson. 1991. A taxonomic revision of the moss families Hookeriaceae and Hypopterygiaceae in Malaya. Smithsonian Contributions to Botany. 80: 1-44. Mohamed, A. and B. C. Tan. 1988. A checklist of mosses of Peninsular Malaya and Singapore. Bryologist. 91: 24-44. Reese, W. D., T. Koponen and D. H. Norris. 1986. Bryophyte flora of the Huon Peninsula, Papua New Guinea. XIX. Calymperes, Syrrhopodon and Mitthyridium (Calymperaceae, Musci). Acta Botanica Fennica. 133: 151-202. Tan, B. C. 1989. The bryophytes of Sabah (North Borneo) with special reference to the BRYOTROP transect of Mount Kinabalu. II. Dicranoloma and Brotherobryum (Dicranaceae, Bryopsida). Willdenowia. 18: 497-512. Tan, B. C. 1991. Miscellaneous notes on Asiatic mosses, especially Malesian Sematophyllaceae (Musci) and others. Journal of Hattori Botanical Laboratory. 70: 91-106. Tan, B. C. and Z. Iwatsuki. 1989. The bryophytes of Sabah (North Borneo) with special reference to the BRYOTOP transect of Mount Kinabalu. VII. Fissidentaceae (Bryopsida). Willdenowia. 18: 591-602. Mosses at SING 57 Tan, B. C. and H. Mohamed. 1990. Novelties for Peninsular Malayan moss flora. Cryptogamie, Bryologique et Lichénologique. 11: 353-362. Touw, A. 1986. A revision of Pogonatum sect. Racelopus, sect. Nov., including Racelopus Dozy & Molk., Pseudoracelopus Broth. and Racelopodopsis Ther. Journal of Hattori Botanical Laboratory. 60: 1- a3: Zander, R. H. 1993. Genera of the Pottiaceae: mosses of harsh environments. Bulletin of the Buffalo Society of Natural Sciences. 32: 1- 378: ’ \ alike 5 Mw a ey . ¢ 2 at a4 7 J Vr Thi muh MAT shee I ata Pn & , vit 4 4 ul ae oes Pa ' 4 ad vbr, pha te ssa a 7 ih ah % ‘ Pre a + i ae a na’ 7 P § 7 7 f i : , A — vat Wp if A ¥ 4 Y, ts { } " by He < Lt ‘ hie ee TS Oe eh a f\ 7 ‘ f BAY ; i pl ( : MP . ; o e ‘ p of . ‘ f*. f ti q , as » Pi are ‘) «= i” J iy +a ee ie wal j a ) : I hi at AL 4 ae Z errs Heil ¥ a ive i eee ee ne pepe) Sy OEAMEL 0) ae, AO re eens oi Gardens’ Bulletin Singapore 50 (1998) 59-81. The Botany of the Islands of Mersing District, Johore, Peninsular Malaysia. 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands I.M. TuRNER!, C.M. Boo, L.M.J. CHEN, J.P.S. CHoo?’, School of Biological Sciences National University of Singapore Singapore 119260 A. LATIFF & A. ZAINUDIN ISMAIL Department of Botany Universiti Kebangsaan Malaysia 43600 Bangi, Selangor, Malaysia Abstract All the records for vascular plants found growing on Pulau Aur, its small neighbour Pulau Dayang, and Pulau Pemanggil, islands in the Mersing District of Johore, Peninsular Malaysia are collated and listed. Notes on the botany of some of the smaller islands in the vicinity are also presented. A total of more than 180 species are listed for Pulau Aur. It is notable as the only Malaysian locality for the rubiaceous tree Zuccarinia macrophylla. Other rare species recorded from the island include Selaginella plana (Selaginellaceae), Operculina riedeliana (Convolvulaceae), Thrixspermum carinatifolium (Orchidaceae), Rauvolfia sumatrana (Apocynaceae), Canarium hirsutum (Burseraceae) and Hymenodictyon orixense (Rubiaceae). A list of the 172 vascular plant species recorded as growing on Pulau Pemanggil is presented. Notable collections include Lasianthus barbellatus (Rubiaceae), Didymocarpus tiumanicus (Gesneriaceae), Mallotus moritzianus (Euphorbiaceae) and Margaritaria indica (Euphorbiaceae). Pulau Sibu is relatively well-known botanically. Details of some recent collections from Pulau Besar are given. Pulau Tengah is notable for records of Argusia argentea (Boraginaceae) and Schizachyrium sanguineum (Gramineae). PULAU AUR Introduction Pulau Aur (104° 32' E 2° 27' N) is the most remote of the Johore islands (Fig. 1), lying some 62 km from the Johore coast, about 74 km due east of Mersing. Aur covers about 19 km’, and the nearby Pulau Dayang about ' Present address: Center for Ecological Research, Kyoto University, Shimosakamoto, Otsu 520-0105, Japan. * Present address: Rutgers University, Cook Campus Foran 135, Picataway, New Jersey 08903, USA. 60 Gard. Bull. Singapore 50(1) (1998) 1.8 km’ (Fig. 2). The first botanical collections were made on Pulau Aur more than a century ago when J. Feilding visited the island in 1892. In April-May 1927, M.R. Henderson visited Pulau Aur, Pulau Dayang and Pulau Tioman. Henderson (1930) reported that the human population numbered about 400, which was a considerable reduction compared to a mid-nineteenth century estimate of 1400. Much of the native vegetation was cleared for cultivation when Aur was heavily populated, but since then secondary forest has developed on many areas away from the coasts. Except for a short note on some common plants (Marchette 1964), there has been no publication on the flora of Pulau Aur since Henderson’s paper. In this paper we collate all vascular plant records for Aur and Dayang that we have been able to verify from material in various herbaria (SING, SINU and UKMB) in Malaysia and Singapore. Recent collecting trips were made by a group from Universiti Kebangsaan Malaysia in January 1988, and by Boo, Chen and Choo in August 1996. P. TIOMAN P. Sembilang ; P. Seri Buat P. Jahat Fen P. Tokong Bahara P. Pemanggil P. Harimau P Gual “P Rawa P. Hujung P. Tengah MERSING. P. Besar P. Aur P. Dayang : Malay P. Ibui Peninsula Mainland P. Tinggi" 446 P. Sibu P. Sibu Tengah Figure 1. Map of the islands off the east coast of Johore. P. = Pulau, spot heights in metres. 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 61 Literature Records There are a few records for plants from Aur for which specimens have not been located recently. Balanophora abbreviata Blume was reportedly collected by Feilding on Aur (Ridley 1924, as Balanophora insularis), but the specimen seems to have eluded all recent authors. All other collections of this parasitic herb from Peninsular Malaysia come from Perak (Kiew 1978). Feilding’s Johore grass collections were written up for publication by Rendle (1894), probably based on specimens deposited in the Kew herbarium. Feilding is reported to have collected five species from Pulau Aur, probably Centotheca lappacea (L.) Desv., Coix lacryma-jobi L., Dactyloctenium aegyptium (L.) Willd., Panicum sarmentosum Roxb. and Setaria sp., but without reference to the specimens it is not possible to be absolutely certain. Tg Jongka Baii Tk Berhala 3 Tg Rekai Pasir Teluran 4 420 Bt Blakang Parang Tg Runjat PULAU AUR Tk Besar a 621 Tk Galing Bt Makum P. Pinang Tk Dalam Tk Ayer Nenek Tg Jongkar = Citi Pasir Lambing Figure 2. Map of Pulau Aur and its neighbouring islands. P. = Pulau, Tg = Tanjung, Tk = Telok, Bt = Bukit, spot heights in metres. 62 Gard. Bull. Singapore 50(1) (1998) Collections of Interest We have amassed records of over 180 species from Pulau Aur. These are listed in Appendix 1. Nomenclature follows Turner (1995) where possible. Probably the most important is that of Zuccarinia macrophylla, which is the only collection of this species (and genus) from Malaysia (Wong 1989). It is a small rubiaceous tree from lowland forest, and has not been recollected recently from Aur. Other very rare plants found on Aur include Selaginella plana (Wong 1983), Thrixspermum carinatifolium (Seidenfaden & Wood 1992), Canarium hirsutum (Turner et al. 1993) and Operculina riedeliana (this study). The apocynaceous shrub Rauvolfia sumatrana was previously reported from Perak alone in Peninsular Malaysia (Markgraf 1984). Our recent record from Pulau Aur extends the species to Johore. Our discovery of Hymenodictyon orixense on Aur is also notable biogeographically being the first record of this species from the east coast of the Malay Peninsula. PULAU PEMANGGIL Introduction Pulau Pemanggil is an island of about 12 km’ (Fig. 3) lying between Pulau Tioman and Pulau Aur in the South China Sea, about 50 km from the Johore mainland (Fig. 1). Pulau Pemanggil is one of the botanically least well-known of the islands of Mersing district. It was the subject of a paper by Latiff (1982), but otherwise little has been published regarding the vascular plants found on the island. Md. Noor and Samsuri collected on the island in 1966. Latiff and his students visited in January 1980 and Boo, Chen and Choo collected there in August 1996. We have collated all the records for vascular plants collected from Pulau Pemanggil in herbaria available to us (KEP, SING, SINU and UKMB) and a list of 172 species is given in Appendix 1. Notable Collections Lasianthus barbellatus, Didymocarpus tiumanicus and Mallotus moritzianus are very rare species in Peninsular Malaysia that are recorded from Pulau Pemanggil. Henderson (1930) reported the first two species as endemic to Pulau Tioman, but both have subsequently been found on Pulau Pemanggil. Didymocarpus tiumanicus has also been recorded from Pulau Tinggi (Turner 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 63 Batu Sudil Tg Batu Selat SR SOS we ON PEMANGGIL Tg Atok Tk Kador Tk Buau Pasir Sempit Tg Telaga Besdarah (rece ener men med Figure 3. Map of Pulau Pemanggil. Tg = Tanjung, Tk = Telok. et al. 1997). Margaritaria indica is confined to the Tioman island group in its Peninsular Malaysian range, having also been recorded from Pulau Sibu Tengah and two small islets to the northeast of Pulau Tioman. Mallotus moritzianus has been recorded from Trengganu as well as Pulau Pemanggil (Whitmore 1973). Not quite so rare, but still notable are the occurrence of Selaginella plana (Wong 1982), Aganosma wallichii (Middleton 1996) and Marsdenia acuminata (Ridley 1923, as Gymnema acuminatum). Deeringia polysperma is a species generally confined to limestone in Peninsular Malaysia (Chin 1979). Pemanggil does not have any limestone outcrops, and the presence of the species on the island is probably a reflection of the tendency to dry spells of the climate of Pulau Pemanggil and its neighbours, and the abundance of well-drained rocky substrates. Limestone, often being freely draining, tends to support species with ranges extending into drier climates. 64 Gard. Bull. Singapore 50(1) (1998) THE SMALLER ISLANDS Introduction We have given accounts of the floras of Pulau Tinggi (Turner et al., 1997) and Pulau Aur and Pulau Pemanggil. There are many other islands, though mostly much smaller than the three already covered, in the Mersing District. In this final section our knowledge of these islands, including some unpublished data, is summarized. Pulau Sibu Pulau Sibu is the best studied island in terms of its botany. A detailed account of the flora and vegetation was given by Turner et al. (1993). Pulau Sibu has little undisturbed inland vegetation remaining, but good examples of a number of coastal vegetation types. The peculiar shrubby heathland found on hills overlooking the sea on Pulau Sibu was the topic of a study by Turner et al. (1995). Pulau Besar There has been little published about this island. On a brief visit in June 1996, Dr C.L. Loh of the Malaysian Nature Society made some plant collections. A list of the species collected is given in Appendix 1. There are some seagrass meadows in the waters surrounding Pulau Besar. Japar (1994) reports the presence of Cymodocea serrulata, Halodule uninervis, Halophila ovalis, Halophila spinulosa and Syringodium isoetifolium. Other Islands E.J.H. Corner visited a number of the small islands in the Tioman group on various trips in the period 1932-1936. He reported his botanical findings nearly fifty years later (Corner 1985). Most of the islets he visited were in Pahang, but he did make collections on Pulau Setindan and Pulau Tengah. Notable collections from Pulau Tengah were the first record from the east coast of Peninsular Malaysia for the small coastal shrub Argusia argentea (Messerschmidia argentea in his paper) and the first record from the Peninsula for the grass Schizachyrium sanguineum. Japar (1994) lists the following seagrass species from Pulau Tengah: Cymodocea serrulata, Halodule pinifolia, Halodule uninervis and Halophila ovalis. 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 65 In the Herbarium of the Singapore Botanic Gardens there are a number of collections made by Mrs Betty Molesworth Allen in July 1955 from Pulau Yu, an island south of Pulau Tinggi. Her ten collections are listed in Appendix 1. Acknowledgements The Malaysian Nature Society are thanked for their support of studies of the plant diversity of the Johore Islands. We are particularly grateful to Dr C.L. Loh for making collections during his visit to Pulau Besar. Dr Ruth Kiew kindly contacted Mrs Molesworth Allen to confirm which Pulau Yu she visited more than forty years ago. References Chin, S.C. 1979. The limestone hill flora of Malaya Il. Gardens’ Bulletin Singapore 30: 165-219. Corner, E.J.H. 1985. The botany of some islets east of Pahang and Johore. Gardens’ Bulletin Singapore 38: 1-42. Henderson, M.R. 1930. Notes on the flora of Pulau Tioman and neighbouring islands. Gardens’ Bulletin Straits Settlement. 5: 80-93. Japar, S.B. 1994. Status of seagrass resources in Malaysia. In Proceedings, Third ASEAN-Australia Symposium on Living Coastal Resources, Vol. 1 CLR. Wilkinson, S. Sudara & L.M. Chou (eds). Australian Institute of Marine Science, Townsville. Pp. 283-289 Kiew, R. 1978. The genus Balanophora in Peninsular Malaysia. Malayan ~ Nature Journal. 30: 539-549. Latiff, A. 1982. Notes on the vegetation and flora of Pulau Pemanggil. Malayan Nature Journal. 35: 217-224. Marchette, N.J. 1964. Notes on some flowering plants of Pulau Aur. Malayan Nature Journal. 18: 50-59. Markgraf, F. 1984. Florae Malesianae Praecursores LXIV. Apocynaceae VI. Rauvolfia. Blumea 30: 157-167. Middleton, D.J. 1996. A revision of Aganosma (Blume) G. Don (Apocynaceae). Kew Bulletin 51: 455-482. 66 Gard. Bull. Singapore 50(1) (1998) Rendle, A.B. 1894. Grasses from Johore. Journal of Botany 32: 100-104. Ridley, H.N. 1923. Flora of the Malay Peninsula. Volume 2. L. Reeve & Co., Ashford. Ridley, H.N. 1924. Flora of the Malay Peninsula. Volume 3. L. Reeve & Co., Ashford. Seidenfaden, G., & Wood, J.J. 1992. The Orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Fredensborg. Turner, I.M. (1995) A catalogue of the vascular plants of Malaya. Gardens’ Bulletin, Singapore 47: 1-757. Turner, I.M., Ong, B.L., & Tan, H.T.W. 1995. Vegetation analysis, leaf structure and nutrient status of a Malaysian heath community. Biotropica 27: 2-12. Turner, I.M., Tan, H.T.W., Kumar, P.P., Chua, K.S., & Haji Samsuri bin Haji Ahmad 1993. The vegetation of Pulau Sibu, Johore. Malayan Nature Journal 46: 169-188. Turner, I.M., Yong, J.W.H., A. Zainudin Ismail, & Latiff, A. 1997. The botany of the islands of Mersing District, Johore. 1. The plants and vegetation of Pulau Tinggi. Gardens’ Bulletin Singapore 49: 119-141. Whitmore, T.C. 1973. Euphorbiaceae. Tree Flora of Malaya. 2: 34-136. Wong, K.M. 1983. Critical observations on Peninsular Malaysian Selaginella. Gardens’ Bulletin Singapore 35: 107-135. Wong, K.M. 1989. Rubiaceae. Tree Flora of Malaya. 4: 324-425. 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 67 Appendix 1. A List of the Plant Species Recorded from Islands in the Mersing District, Johore. The species are listed with one representative collection for each. Where possible, for Pulau Aur, an indication of records from Pulau Aur and/or Pulau Dayang is given in brackets after each species. A ‘c’ in the margin next to an entry indicates a species only found in cultivation. Pulau Aur PTERIDOPHYTA Aspleniaceae Polypodiaceae Asplenium nidus L. - Boo, Chen & Choo Pyrrosia angustata (Sw.) Ching - Boo, 399 [P. Aur] (SINU) Chen & Choo 402 [P. Aur] (SINU) Pyrrosia lanceolata (L.) Farwell - Boo, Cyatheaceae Chen & Choo 257 [P. Aur & P. Dayang] Cyathea squamulata (Blume) Copel. -Boo, (SINU) Chen & Choo 386 [P. Aur] (SINU) Pteridaceae Dennstaedtiaceae Pteris ensiformis Burm.f. - Boo, Chen & Lindsaea cultrata ( Willd.) Sw. - Boo, Chen Choo 186 [P. Aur] (SINU) & Choo 384 [P. Aur] (SINU) Schizaeaceae Dryopteridaceae Lygodium circinnatum (Burm.f.) Sw. - Tectaria singaporeana (Hook. & Grev.) Copel.- Boo, Chen & Choo 420 [P. Aur] (SINU) Hymenophyllaceae Cephalomanes javanicum (Blume) Bosch - Boo, Chen & Choo 286 [P. Aur] (SINU) Marattiaceae Angiopteris evecta (G. Forst.) Hoffm. - Boo, Chen & Choo 383 [P. Aur] (SINU) Oleandraceae Nephrolepis acutifolia (Desv.) H. Christ. - Boo, Chen & Choo 382 [P. Aur] (SINU) Nephrolepis auriculata (L.) Trimen - Boo, Chen & Choo 217 [P. Dayang] (SINU) M.R. Henderson, S.EN. 18247 (SING) Selaginellaceae Selaginella plana (Desv.) Hieron. - M.R. Henderson, S.EN. 18365 (BM) Thelypteridaceae Pronephrium repandum (Fée) Holttum - Boo, Chen & Choo 388 [P. Aur] (SINU) Vittariaceae Vittaria ensiformis Sw. - Boo, Chen & Choo 379 [P. Aur] (SINU) SPERMATOPHYTA Acanthaceae c Justicia gendarussa Burm.f. - Boo, Chen & Choo 195 [P. Aur] (SINU) 68 Alangiaceae Alangium kurzii Craib - Boo, Chen & Choo 280 [P. Aur & P. Dayang] (SINU) Amaranthaceae Cyathula prostrata (L.) Blume - Boo, Chen & Choo 294 [P. Aur] (SINU) Amaryllidaceae Crinum asiaticum L. - Boo, Chen & Choo 144 [P. Aur] (SINU) Proiphys amboinensis (L.) Herb. - Boo, Chen & Choo 326 [P. Dayang] (SINU) Anacardiaceae c Anacardium occidentale L. - Boo, Chen & Choo 327 [P. Aur] (SINU) Annonaceae c Cananga odorata (Lam.) Hook.f. & Thomson - A. Latiff & A. Zainudin 2708 [P. Aur] (UKMB) Meiogyne virgata (Blume) Mig. - MLR. | Henderson, S.F.N. 18360 (SING) Friesodielsia affinis (Hook.f. & Thomson) D. Das - A. Latiff & A. Zainudin 2699 [[P. Aur] (UKMB) Friesodielsia kingii (J. Sinclair) Steenis - M.R. Henderson, S.F.N. 18235 (SING) Polyalthia cinnamomea Hook.f. & Thomson- M.R. Henderson, S.F.N. 18216 (SING) Apocynaceae Catharanthus roseus (L.) G. Don - Boo, Chen & Choo 248 [P. Aur & P. Dayang] (SINU) Cerbera manghas L. - A. Latiff & A. Zainudin 2668 [P. Aur] (UKMB) Cerbera odollam Gaertn. - Boo, Chen & Choo 314 [P. Dayang] (SINU) Rauvolfia sumatrana Jack - Boo, Chen & Choo 159 [P. Aur] (SINU) Gard. Bull. Singapore 50(1) (1998) Araceae Aglaonema simplex Blume - M.R. Henderson S.F.N. 18221 (SING) Rhaphidophora korthalsii Schott - Boo, Chen & Choo 380 [P. Aur] (SINU) c Syngonium podophyllum Schott - Boo, Chen & Choo 284 [P. Aur] (SINU) Araliaceae Arthrophyllum diversifolium Blume - M.R. Henderson, S.F.N. 18362 (SING) Schefflera elliptica (Blume) Harms - A. Latiff & A. Zainudin 2701 [P. Aur] (UKMB) Schefflera heterophylla (Wall. ex G. Don) Harms - M.R. Henderson, S.EN. 18238 (SING) Aristolochiaceae Aristolochia tagala Cham. - M.R. Henderson, S.EN. 18226 (SING) Asclepiadaceae Dischidia bengalensis Colebr. - Feilding s.n. (SING) Hoya verticillata (Vahl) G. Don - Boo, Chen & Choo 289 [P. Aur] (SINU) Boraginaceae Ehretia timorensis Decne. - M.R. Henderson, S.EN. 18203 (SING) Bromeliaceae c Ananas comosus (L.) Merr. - Boo, Chen & Choo 222 [P. Dayang] (SINU) Burseraceae Canarium hirsutum Willd. - M.R. Henderson, S.F.N. 18242 (SING) Cactaceae Opuntia monacantha Haw.- Boo, Chen & Choo 238 [P. Dayang] (SINU) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 69 Cannaceae c Canna edulis Ker Gawl. - Boo, Chen & Choo 213 [P. Dayang] (SINU) Capparaceae Cleome viscosa L. - Boo, Chen & Choo 151 [P. Aur & P. Dayang] (SINU) Cecropiaceae Poikilospermum suaveolens (Blume) Merr. - A. Latiff & A. Zainudin 2705 [P. Aur] (UKMB) Combretaceae Quisqualis indica L. - M.R. Henderson, S.EN. 18227 (SING) Commelinaceae Commelina diffusa Burm.f. - Boo, Chen & Choo 269 [P. Aur] (SINU) Murdannia nudiflora (L.) Brenan - Boo, Chen & Choo 163 [P. Aur & P. Dayang] (SINU) Compositae Elephantopus scaber L. - Boo, Chen & Choo 150 [P. Aur] (SINU) Vernonia cinerea (L.) Less. - Boo, Chen & Choo 179 [P. Aur] (SINU) Vernonia patula (Dryand.) Merr. - Boo, Chen & Choo 257 [P. Aur] (SINU) Wollastonia biflora (L.) DC. - Boo, Chen & Choo 220 [P. Dayang] (SINU) Convolvulaceae Ipomoea pes-caprae (L.) R.Br. - Boo, Chen & Choo 233 [P. Dayang] (SINU) Ipomoea pes-tigridis L. - Boo, Chen & Choo 265 [P. Aur] (SINU) Operculina riedeliana (Oliv.) Ooststr. - Boo, Chen & Choo 409 [P. Aur] (SINU) Crassulaceae Kalanchoe pinnata (Lam.) Pers. - Boo, Chen & Choo 196 [P. Aur & P. Dayang] (SINU) Cycadaceae Cycas ?siamensis Miq. - Boo, Chen & Choo 305 [P. Dayang] (SINU) Cyperaceae Scleria lithosperma (L.) Sw. - Boo, Chen & Choo 373 [P. Aur] (SINU) Dilleniaceae Tetracera indica (Christm. & Panz.) Merr. - Boo, Chen & Choo 168 [P. Aur] (SINU) Dracaenaceae Dracaena elliptica Thunb. - Boo, Chen & Choo 419 [P. Aur] (SINU) Ebenaceae Diospyros cauliflora Blume - M.R. Henderson, S.EN. 18276 (SING) Escalloniaceae Polyosma integrifolia Blume - Boo, Chen & Choo 421 [P. Aur] (SINU) Euphorbiaceae Claoxylon indicum (Reinw. ex Blume) Hassk. - Boo, Chen & Choo 279 [P. Aur] (SINU) Cnesmone javanica Blume - Boo, Chen & Choo 393 [P. Aur] (SINU) c Jatropha curcas L. - Boo, Chen & Choo 272 [P. Aur] (SINU) Macaranga gigantea (Rchb.f. & Zoll.) Miill.Arg. - Boo, Chen & Choo 353 [P. Aur] (SINU) c¢ Manihot esculenta Crantz - Boo, Chen & Choo 245 [P. Aur] (SINU) Melanolepis multiglandulosa (Reinw. ex Blume) Rchb.f. & Zoll. - Boo, Chen & Choo 278 [P. Aur & P. Dayang] (SINU) c Pedilanthus tithymaloides Poit. - Boo, Chen & Choo 293 [P. Aur] (SINU) Phyllanthus pulcher Wall. ex Miill.Arg. - Boo, Chen & Choo 189 [P. Aur] (SINU) 70 Flacourtiaceae c Flacourtia jangomas (Lour.) Raeusch. - Boo, Chen & Choo 405 [P. Aur & P. Dayang] (SINU) Flacourtia rukam Zoll. & Moritzi - A. Latiff & A. Zainudin 2665 [P. Aur] (UKMB) Flagellariaceae Flagellaria indica L. - Boo, Chen & Choo 172 [P. Aur] (SINU) Gnetaceae c Gnetum gnemon L. - Boo, Chen & Choo 331 P. Dayang Gnetum latifolium Blume var. funiculare (Blume) Marker. - Boo, Chen & Choo 290 [P. Aur] (SINU) Gnetum macrostachyum Hook.f. - M.R. Henderson, S.EN. 18250 (SING) Goodeniaceae Scaevola taccada (Gaertn.) Roxb. - Boo, » Chen & Choo 231 [P. Dayang] (SINU) Gramineae Cyrtococcum oxyphyllum (Steud.) Stapf - M.R. Henderson, S.EN. 18225 (SING) Guttiferae Calophyllum inophyllum L. - Boo, Chen & Choo 219 [P. Dayang] (SINU) Garcinia parvifolia Miq. - Boo, Chen & Choo 365 [P. Aur] (SINU) Hypoxidaceae Molineria latifolia (Dryand.) Herb. - Boo, Chen & Choo 374 [P. Aur] (SINU) Labiatae Leucas zeylanica (L.) R.Br.- Boo, Chen & Choo 339 [P. Dayang] (SINU) Leeaceae Leea indica (Burm.f.) Merr. - Boo, Chen Gard. Bull. Singapore 50(1) (1998) & Choo 162 [P. Aur & P. Dayang] (SINU) Leguminosae Abrus precatorius L. - Boo, Chen & Choo 239 [P. Dayang] (SINU) Bauhinia integrifolia Roxb. - M.R. Henderson, S.F.N. 18233 (SING) Canavalia cathartica Thouars - Boo, Chen & Choo 226 [P. Dayang] (SINU) Crotalaria pallida Aiton - Boo, Chen & Choo 267 [P. Aur & P. Dayang] (SINU) Crotalaria retusa L. - Boo, Chen & Choo 270 [P. Aur] (SINU) Crudia lanceolata Ridl. - Boo, Chen & Choo 410 [P. Aur] (SINU) Derris scandens (Roxb.) Benth. - Boo, Chen & Choo 341 [P. Dayang] (SINU) Desmodium gangeticum (L.) DC. - Boo, Chen & Choo 360 [P. Aur] (SINU) Desmodium heterocarpon (L.) DC. - Boo, Chen & Choo 381 [P. Aur] (SINU) Desmodium velutinum (Willd.) DC. - Boo, Chen & Choo 281 [P. Aur] (SINU) Flemingia strobilifera (L.) Roxb. - MLR. Henderson, S.F.N. 18240 (SING) Mimosa pudica L. - Boo, Chen & Choo 164 [P. Aur] (SINU) Mucuna biplicata Teijsm. & Binn. ex Kurz - Boo, Chen & Choo 425 [P. Aur] (SINU) Pueraria phaseoloides (Roxb.) Benth. - Boo, Chen & Choo 274 [P. Aur] (SINU) Senna alata (L.) Roxb. - Boo, Chen & Choo 145 [P. Aur] (SINU) Senna occidentalis (L.) Link - Boo, Chen & Choo 173 [P. Aur] (SINU) Loganiaceae Fagraea auriculata Jack - Boo, Chen & Choo 325 [P. Aur] (SINU) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 71 Malvaceae Hibiscus tiliaceus L. - A. Latiff & A. Zainudin 2 [P. Aur] (UKMB) Sida cordifolia L. - Boo, Chen & Choo 362 [P. Aur] (SINU) Sida rhombifolia L. - Boo, Chen & Choo 263 [P. Aur] (SINU) Urena lobata L. - Boo, Chen & Choo 304 [P. Dayang] (SINU) Melastomataceae Clidemia hirta (L.) D. Don - Boo, Chen & Choo 404 [P. Aur] (SINU) Melastoma malabathricum L. - Boo, Chen & Choo 193 [P. Aur & P. Dayang] (SINU) Memecylon caeruleum Jack - Boo, Chen & Choo 143 [P. Aur & P. Dayang] (SINU) Meliaceae Chukrasia tabularis A. Juss. - Boo, Chen & Choo 275 [P. Aur] (SINU) Dysoxylum cauliflorum Hiern - M.R. Henderson, S.EN. 18354 (SING) Menispermaceae Fibraurea tinctoria Lour. - Boo, Chen & Choo 424 [P. Aur] (SINU) Pericampylus glaucus (Lam.) Merr. - Boo, Chen & Choo 198 [P. Aur & P. Dayang] (SINU) Tinospora crispa (L.) Hook.f. & Thomson - Boo, Chen & Choo 275 [P. Dayang] (SINU) Monimiaceae Kibara coriacea (Blume) Tul. - Boo, Chen & Choo 422 [P. Aur] (SINU) Matthaea sancta Blume - Boo, Chen & Choo 418 [P. Aur] (SINU) Moraceae Ficus aurantiacea Griff. - A. Latiff & A. Zainudin 2677 [P. Aur] (UKMB) Ficus recurva Blume - Boo, Chen & Choo 375 [P. Aur] (SINU) Ficus superba (Miq.) Mig. - Boo, Chen & Choo 272 [P. Aur] (SINU) Myristicaceae Knema curtisii (King) Warb. - M.R. Henderson, S.F.N. 18220 (SING) Knema globularia (Lam.) Warb. - M.R. Henderson, S.EN. 18244 (SING) Myrsinaceae Ardisia elliptica Thunb. - Boo, Chen & Choo 320 [P. Dayang] (SINU) Ardisia lanceolata Roxb. - Boo, Chen & Choo 174 [P. Aur] (SINU) Ardisia lurida Blume - Boo, Chen & Choo 200 [P. Aur] (SINU) Ardisia solanacea Roxb. - M.R. Henderson, S.E.N. 18232 (SING) Myrtaceae c Psidium guajava L. - Boo, Chen & Choo 170 [P. Aur & P. Dayang] (SINU) Syzygium grande (Wight) Walp. - Boo, Chen & Choo 300 [P. Dayang] (SINU) c Syzygium jambos (L.) Alston - Boo, Chen & Choo 201 [P. Aur] (SINU) Syzygium pseudoformosum (King) Merr. & L.M. Perry - Boo, Chen & Choo 397 [P. Aur] (SINU) Nyctaginaceae Boerhavia diffusa L.- Boo, Chen & Choo 214 [P. Dayang] (SINU) Orchidaceae Cirrhopetalum puguahaanense (Ames) Garay et al. - Boo, Chen & Choo 429 [P. Aur] (SINU) Thrixspermum caranatifolium (Ridl.) Schltr. - J. Fielding s.n., 1892 (SING) Palmae Arenga westerhouti Griff. - Boo, Chen & Choo 356 [P. Aur] (SINU) Daemonorops melanochaetes Blume - M.R. Henderson S.F.N. 18358 (SING) c Metroxylon sagu Rottb. - Boo, Chen & Choo 205 [P. Aur] (SINU) Pandanaceae Freycinetia imbricata Blume - Boo, Chen & Choo 408 [P. Aur] (SINU) Passifloraceae Passiflora foetida L. - Boo, Chen & Choo 251 [P. Aur & P. Dayang] (SINU) Piperaceae c Piper betel L. - Boo, Chen & Choo 376 [P. Aur] (SINU) Portulacaceae Portulaca oleracea L. - Boo, Chen & Choo 192 [P. Aur] (SINU) Portulaca quadrifida L. - Boo, Chen & Choo 309 [P. Dayang] (SINU) Rosaceae Eriobotrya bengalensis (Roxb.) Hook.f. - M.R. Henderson, S.F.N. 18211 (SING) Rubus moluccanus L. var. angulosus Kalkman - A. Latiff & A. Zainudin 2692 {P. Aur] (UKMB) Rubiaceae Borreria laevicaulis (Mig.) Ridl. - Boo, Chen & Choo 229 [P. Aur & P. Dayang] (SINU) Geophila repens (L.) I.M. Johnst. var. asiatica (Cham. & Schldtl.) Fosberg - Boo, Chen & Choo 359 [P. Aur] (SINU) Hedyotis capitellata Wall. ex G. Don - M.R. Henderson, S.F.N. 18356 (SING) Hedyotis dichotoma Koenig ex Roth - Boo, Chen & Choo 371 [P. Aur] (SINU) Gard. Bull. Singapore 50(1) (1998) Hedyotis tenelliflora Blume - Boo, Chen & Choo 361 [P. Aur] (SINU) Hedyotis vestita R. Br. ex G. Don - MLR. Henderson, S.F.N. 18364 (SING) Hymenodictyon orixense (Roxb.) Mabb. - A. Latiff & A. Zainudin 2706 [P. Aur] (UKMB) Ixora javanica (Blume) DC. - M.R. Henderson, S.F.N. 18218 (SING) Ixora nigricans Wight & Arn. - MLR. Henderson, S.F.N. 18212 (SING) Ixora pendula Jack - Boo, Chen & Choo 202 [P. Aur] (SINU) Ixora umbellata Koord. & Valeton - M.R. Henderson, S.F.N. 18219 (SING) Kailarsenia tentaculata (Hook.f.) Tirveng. - ?Feilding 4082 (SING) Morinda citrifolia L. - Boo, Chen & Choo 247 [P. Aur] (SINU) Morinda elliptica (Hook.f.) Ridl. - MLR. Henderson, S.F.N. 18363 (SING) Morinda umbellata L. - M.R. Henderson, S.F.N. 18210 (SING) Ophiorrhiza discolor R.Br. - Boo, Chen & Choo 407 [P. Aur] (SINU) Paederia foetida L: - Boo, Chen & Choo 372 [P. Aur] (SINU) Pavetta naucleiflora R.Br. ex G. Don - M.R. Henderson, S.F.N. 18245 (SING) Tarenna costata (Mig.) Merr. - M.R. Henderson, S.F.N. 18375 (SING) Timonius wallichianus (Korth.) Valeton - Boo, Chen & Choo 391 [P. Aur] (SINU) Zuccarinia macrophylla Blume - M.R. Henderson, S.EN. 18213 (SING) Rutaceae c Citrus maxima (L.) Merr. - Boo, Chen & Choo 149 [P. Aur] (SINU) Glycosmis chlorosperma Spreng. - Boo, Chen & Choo 157 [P. Aur] (SINU) Glycosmis mauritiana (Lam.) Tanaka - Boo, Chen & Choo 313 [P. Dayang] (SINU) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands Hie) Glycosmis pentaphylla (Retz.) DC. - A. Latiff & A. Zainudin 2707 [P. Aur] (UKMB) Murraya paniculata (L.) Jack - Boo, Chen & Choo 207 [P. Aur] (SINU) c Triphasia trifolia (Burm.f.) P. Wilson - Boo, Chen & Choo 167 [P. Aur] (SINU) Sapindaceae Allophylus cobbe (L.) Raeusch. - M.R. Henderson, S.F.N. 18369 (SING) Cardiospermum halicacabum L. - Boo, Chen & Choo 161 [P. Aur] (SINU) Lepisanthes rubiginosa (Roxb.) Leenh. - M.R. Henderson, S.F.N. 18205 (SING) Lepisanthes tetraphylla (Vahl) Radlk. - M.R. Henderson, S.EN. 18372 (SING) Scrophulariaceae Lindernia crustacea (L.) F. Muell. - Boo, Chen & Choo 175 [P. Aur & P. Dayang] (SINU) Simaroubaceae Brucea javanica (L.) Merr. - M.R. Henderson, S.F.N. 18207 (SING) Eurycoma longifolia Jack - Boo, Chen & Choo 364 [P. Aur] (SINU) Solanaceae c Datura metel L. - Boo, Chen & Choo 142 [P. Aur] (SINU) Solanum erianthum D. Don - M.R. Henderson, S.EN. 18201 (SING) Sterculiaceae Leptonychia caudata (Wall. ex G. Don) Burret - M.R. Henderson, S.EN. 18355 (SING) Taccaceae Tacca palmata Blume - Boo, Chen & Choo 306 [P. Dayang] (SINU) Tiliaceae Triumfetta rhomboidea Jacq. - Boo, Chen & Choo 268 [P. Aur] (SINU) Umbelliferae Centella asiatica (L.) Urb. - Boo, Chen & Choo 335 [P. Dayang] (SINU) Urticaceae Pouzolzia zeylanica (L.) Benn. - Boo, Chen & Choo 190 [P. Aur] (SINU) Verbenaceae Callicarpa candicans (Burm.f.) Hochr. - M.R. Henderson, S.F.N. 18202 (SING) Callicarpa longifolia Lam. - Boo, Chen & Choo 155 [P. Aur & P. Dayang] (SINU) Clerodendrum laevifolium Blume - Boo, Chen & Choo 208 [P. Aur] (SINU) Gmelina elliptica Sm. - M.R. Henderson, S.F.N. 18209 (SING) Lantana camara L. - Boo, Chen & Choo 188 [P. Aur] (SINU) Peronema canescens Jack - Boo, Chen & Choo 153 [P. Aur] (SINU) Premna serratifolia L. - Boo, Chen & Choo 244 [P. Aur] (SINU) Stachytarpheta indica (L.) Vahl - Boo, Chen & Choo 221 [P. Dayang] (SINU) Vitex trifolia L. - Boo, Chen & Choo 227 [P. Dayang] (SINU) Violaceae Rinorea horneri (Korth.) Kuntze - MLR. Henderson, S.EN. 18217 (SING) Vitaceae Cayratia japonica (Thunb.) Gagnep. - Boo, Chen & Choo 253 [P. Aur] (SINU) Cayratia mollissima (Wall.) Gagnep. - M.R. Henderson, S.F.N. 18206 (SING) Cayratia trifolia (L.) Domin - A. Latiff & A. Zainudin s.n. [P. Aur] (UKMB) 74 Cissus repens Lam. - Boo, Chen & Choo 181 [P. Aur & P. Dayang] (SINU) Tetrastigma leucostaphylum (Dennst.) Alston ex Mabb. - Boo, Chen & Choo 392 [P. Aur] (SINU) Tetrastigma pisicarpum (Miq.) Planch.- A. Latiff & A. Zainudin 2698 [P. Aur] (UKMB) Pulau PTERIDOPHYTA Aspleniaceae Asplenium affine Sw. - A. Latiff & A. Zainudin s.n. (UKMB) Asplenium polyodon G. Forst. - A. Latiff & A. Zainudin s.n. (UKMB) Asplenium robustum Blume - A. Latiff & A. Zainudin s.n. (UKMB) Dryopteridaceae Pleocnemia irregularis (C. Presl) Holttum — - Boo, Chen & Choo 95 (SINU) Lycopodiaceae Huperzia phlegmaria (L.) Rothm. - Md. Noor & Samsuri 24 (SING) Marratiaceae Angiopteris evecta (G. Forst.) Hoffm. - Boo, Chen & Choo 84 (SINU) Oleandraceae Nephrolepis acutifolia (Desv.) H. Christ - Boo, Chen & Choo 127 (SINU) Nephrolepis cordifolia (L.) C. Presl - R. Jaman 480 (UKMB) Nephrolepis hirsutula (G. Forst.) C. Pres] - Boo, Chen & Choo 55 (SINU) Pteridaceae Acrostichum aureum L. - Boo, Chen & Choo 112 (SINU) Gard. Bull. Singapore 50(1) (1998) Zingiberaceae c Alpinia galanga (L.) Sw. - Boo, Chen & Choo 258 [P. Aur & P. Dayang] (SINU) Alpinia mutica Roxb. - M.R. Henderson, S.F.N. 18228 (SING) Amomum uliginosum J. Konig - Boo, Chen & Choo 369 [P. Aur] (SINU) Pemanggil Pteris ensiformis Burm.f. - Boo, Chen & Choo 126 (SINU) Pteris venulosa Blume - R. Jaman 474 (UKMB) Selaginellaceae Selaginella plana (Desv.) Hieron. - Boo, Chen & Choo 77 (SINU) Selaginella willdenowii (Desv.) Baker - R. Jaman 479 (UKMB) Thelypteridaceae © Amphineuron terminans (Hook.) Holttum - Boo, Chen & Choo 77 (SINU) Vittariaceae Vittaria angustifolia Blume - A. Latiff & A. Zainudin s.n. (UKMB) SPERMATOPHYTA Amaranthaceae Deeringia polysperma (Roxb.) Mog. - Md. Noor & Samsuri 49 (SING) Amaryllidaceae c Hippeastrum reticulatum (L’Hér.) Herb. - Shafiee Daud s.n. (UKMB) Ancistrocladaceae Ancistrocladus tectorius (Lour.) Merr. - Shafiee Daud s.n. (UKMB) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands bs) Annonaceae c Cananga odorata (Lam.) Hook.f. & Thomson - Boo, Chen & Choo 76 (SINU) Apocynaceae Aganosma marginata (Roxb.) G. Don - Boo, Chen & Choo 66 (SINU) Aganosma wallichii G. Don - Md. Noor & Samsuri 54 (SING) Catharanthus roseus (L.) G. Don - Saidah Mamat s.n. (UKMB) Araceae Aglaonema nitidum (Jack) Kunth - Ramli Khamis s.n. (UKMB) Aglaonema simplex Blume - Ramli Khamis s.n. (UKMB) Epipremnum giganteum (Roxb.) Schott - A. Latiff & A. Zainudin 156 (UKMB) Schismatoglottis calyptrata (Roxb.) Zoll. & Moritzi - Boo, Chen & Choo 94 (SINU) Scindapsus beccarii Engl. - Boo, Chen & Choo 38 (SINU) Araliaceae Schefflera elliptica (Blume) Harms - Boo, Chen & Choo 16 (SINU) Schefflera oxyphylla (Miq.) R. Vig. - Boo, Chen & Choo 57 (SINU) Asclepiadaceae Hoya diversifolia Blume - Md. Noor & Samsuri 32 (SING) Marsdenia acuminata (Roxb.) I.M. Turner - Boo, Chen & Choo 120 (SINU) Cecropiaceae Poikilospermum suaveolens (Blume) Merr. - Md. Noor & Samsuri 38 (SING) Celastraceae Loesneriella macrantha (Korth.) A.C. Sm. - Md. Noor & Samsuri 19 (SING) Microtropis valida Ridl. - Md. Noor & Samsuri 41 (SING) Combretaceae Combretum tetralophum C.B. Clarke - Boo, Chen & Choo 13 (SINU) Quisqualis indica L. - Boo, Chen & Choo 46 (SINU) Terminalia catappa L.- Boo, Chen & Choo 106 (SINU) Commelinaceae Belosynapsis ciliata (Blume) R.S. Rao - Boo, Chen & Choo 48 (SINU) Commelina diffusa Burm.f. - Boo, Chen & Choo 70 (SINU) Compositae Blumea balsamifera (L.) DC. - Md. Noor & Samsuri 59 (SING) Complaya trilobata (L.) Strother - Boo, Chen & Choo 122 (SINU) Conyza bonariensis (L.) Cronquist - Md. Noor & Samsuri 55 (SING) Vernonia cinerea (L.) Less. - Boo, Chen & Choo 40 (SINU) Wollastonia biflora (L.) DC. - Boo, Chen & Choo 20 (SINU) Convolvulaceae c Ipomoea batatas (L.) Lam. - Arishah Hashim s.n. (UKMB) Ipomoea mauritiana Jacq. - Boo, Chen & Choo 45 (SINU) Ipomoea pes-caprae (L.) R.Br. - Boo, Chen & Choo 123 (SINU) Crassulaceae ¢ Kalanchoe laciniata (L.) DC. - Boo, Chen & Choo 41 (SINU) Kalanchoe pinnata (Lam.) Pers. - Boo, Chen & Choo 102 (SINU) Cycadaceae Cycas rumphii Mig. - Boo, Chen & Choo 141 (SINU) 76 Dilleniaceae Tetracera indica (Christm. & Panz.) Merr. - Boo, Chen & Choo 98 (SINU) Dracaenaceae Dracaena maingayi Hook.f. - Boo, Chen & Choo 61 (SINU) Euphorbiaceae Actephila excelsa (Dalzell) Miill.Arg. var. javanica (Miq.) Pax & K. Hoffm. - Md. Noor & Samsuri MN14 (SING) Antidesma cuspidatum Mill.Arg. - Boo, Chen & Choo 85 (SINU) Antidesma japonicum Siebold. & Zucc. - Rahim Hamid s.n. (UKMB) Breynia reclinata (Roxb.) Hook.f. - Boo, Chen & Choo 26 (SINU) Breynia vitis-idaea (Burm.f.) C.E.C. Fisch. - Md. Noor & Samsuri 9 (SING) Mallotus moritzianus Miill.Arg. - Md. Noor & Samsuri 17 (SING) Mallotus paniculatus (Lam.) Miill.Arg. - - Saidah Mamat s.n. (UKMB) Mallotus philippensis (Lam.) Miil.Arg. - Shafiee Daud s.n. (UKMB) c Manihot esculenta Crantz - Boo, Chen & Choo 81 (SINU) Margaritaria indica (Dalzell) Airy Shaw - Boo, Chen & Choo 18 (SINU) Melanolepis multiglandulosa (Reinw. ex Blume) Rchb.f. & Zoll. - Boo, Chen & Choo 72 (SINU) Phyllanthus pulcher Wall. ex Mill.Arg. - Boo, Chen & Choo 113 (SINU) Suregada multiflora (Juss.) Baill.- A. Latiff & A. Zainudin 148 (UKMB) Flacourtiaceae c Flacourtia jangomas (Lour.) Raeusch. - Boo, Chen & Choo 111 (SINU) Flagellariaceae Flagellaria indica L. - Boo, Chen & Choo 116 (SINU) Gard. Bull. Singapore 50(1) (1998) Gesneriaceae Didymocarpus tiumanicus (Ridl.) B.L. Burtt - Md. Noor & Samsuri MN72 (SING) Gnetaceae c Gnetum gnemon L. - Boo, Chen & Choo 90 (SINU) Gnetum latifolium Blume var. funiculare (Blume) Markgr. - Md. Noor & Samsuri MN68 (SING) Goodeniaceae Scaevola taccada (Gaertn.) Roxb. - Boo, Chen & Choo 105 (SINU) Gramineae Centotheca lappacea (L.) Desv. - Boo, Chen & Choo 69 (SINU) Oplismenus compositus (L.) P. Beauv. - Md. Noor & Samsuri 50 (SING) Guttiferae Calophyllum inophyllum L. - Boo, Chen & Choo 107 (SINU) Garcinia hombroniana Pierre - Md. Noor & Samsuri 25 (SING) Joinvilleaceae Joinvillea ascendens Brongn. & Gris. ssp. borneensis (Becc.) Newell - Arishah Hashim 43 (UKMB) Labiatae Leucas zeylanica (L.) R.Br. - Boo, Chen & Choo 34 (SINU) Ocimum tenuiflorum L. - Boo, Chen & Choo 23 (SINU) c Solenostemon scutellarioides (L.) Codd - Shafiee Daud s.n. (UKMB) Lauraceae Cassytha filiformis L. - Boo, Chen & Choo 25 (SINU) Litsea glutinosa (Lour.) C.B. Rob. - Md. Noor & Samsuri 6 (SING) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands ral Lecythidaceae Barringtonia asiatica (L.) Kurz - Boo, Chen & Choo 346 Leeaceae Leea indica (Burm.f.) Merr.- Md. Noor & Samsuri 4 (SING) Leguminosae Abrus precatorius L. - Boo, Chen & Choo 117 (SINU) Archidendron jiringa (Jack) I.C. Nielsen - Arishah Hashim 60 (UKMB) Bauhinia integrifolia Roxb. - Boo, Chen & Choo 9 (SINU) Canavalia cathartica Thouars - Md. Noor & Samsuri 60 (SING) c Cassia fistula L. - Saidah Mamat 32 (UKMB) Crotalaria pallida Aiton - Boo, Chen & Choo 29 (SINU) Derris scandens (Roxb.) Benth. - Boo, Chen & Choo 118 (SINU) Desmodium velutinum (Willd.) DC. - Boo, Chen & Choo 6 (SINU) Flemingia strobilifera (L.) Roxb. - Boo, Chen & Choo 133 (SINU) Mucuna biplicata Teijsm. & Binn. ex Kurz - Arishah Hashim 73 (UKMB) c Psophocarpus tetragonolobus (L.) DC. - Rahim Hamid s.n. (UKMB) Pueraria phaseoloides (Roxb.) Benth. - Boo, Chen & Choo 8 (SINU) Vigna marina (Burm.) Merr. - Boo, Chen & Choo 10 (SINU) Lythraceae Pemphis acidula J.R. Forst & G. Forst. - Shafiee Daud s.n. (UKMB) Malvaceae c Abelmoschus moschatus Medik. - Zainal Mustaffa s.n. (UKMB) Abutilon indicum (L.) Sweet - Rahim Hamid s.n. (UKMB) Hibiscus tiliaceus L. - Boo, Chen & Choo 100 (SINU) Sida acuta Burm.f. - Boo, Chen & Choo 24 (SINU) Thespesia populnea (L.) Sol. ex Corréa - Md. Noor & Samsuri 53 (SING) Urena lobata L. - Boo, Chen & Choo 11 (SINU) Melastomataceae Medinilla crassifolia (Reinw. ex Blume) Blume - Zainal Mustaffa s.n. (UKMB) Memecylon caeruleum Jack - Boo, Chen & Choo 47 (SINU) Memecylon edule Roxb. - Boo, Chen & Choo 30 (SINU) Pogonanthera pulverulenta (Jack) Blume - Boo, Chen & Choo 54 (SINU) Menispermaceae Pericampylus glaucus (Lam.) Merr. - Boo, Chen & Choo 32 (SINU) Stephania capitata (Blume) Spreng. - Boo, Chen & Choo 80 (SINU) Moraceae Artocarpus lanceifolius Roxb. - A. Latiff & A. Zainudin 132 (UKMB) Ficus callosa Willd. - Rahim Ahmad 65 (UKMB) Ficus deltoidea Jack - Arishah Hashim 66 (UKMB) Ficus drupacea Thunb. - Boo, Chen & Choo 63 (SINU) Ficus hispida L.f. - Boo, Chen & Choo 52 (SINU) Ficus schwarzit Koord. - A. Latiff & A. Zainudin 134 (UKMB) Ficus superba (Migq.) Miq. - Boo, Chen & Choo 43 (SINU) Ficus tinctoria G. Forst. ssp. gibbosa (Blume) Corner - Boo, Chen & Choo 7 (SINU) Streblus ilicifolius (Vidal) Corner - Boo, Chen & Choo 92 (SINU) 78 Myristicaceae Knema laurina (Blume) Warb. - Md. Noor & Samsuri 69 (SING) Myrsinaceae Ardisia elliptica Thunb. - Arishah Hashim 67 (UKMB) Ardisia oxyphylla Wall. ex DC. - A. Latiff & A. Zainudin 1394 (UKMB) Ardisia villosa Roxb. - Md. Noor & Samsuri 74 (SING) Myrtaceae c Psidium guajava L. - Boo, Chen & Choo 37 (SINU) Nytaginaceae Boerhavia diffusa L. - Boo, Chen & Choo 2 (SINU) Onagraceae Ludwigia hyssopifolia (G. Don) Exell - — Boo, Chen & Choo 87 (SINU) Opiliaceae Lepionurus sylvestris Blume - Md. Noor & Samsuri 67 (SING) Orchidaceae Cymbidium finlaysonianum Lindl. - Md. Noor & Samsuri 28 (SING) Dendrobium crumenatum Sw. - Boo, Chen & Choo 62 (SINU) Eulophia spectabilis (Dennst.) Suresh - Boo, Chen & Choo 109 (SINU) Phaius callosus (Blume) Lindl. - Md. Noor & Samsuri 26 (SING) Palmae Daemonorops angustifolia (Griff.) Mart. - Arishah Hashim s.n. (UKMB) Daemonorops sabut Becc. - R. Jaman 484 (UKMB) Daemonorops sepal Becc. - Shafiee Daud 49 (UKMB) Gard. Bull. Singapore 50(1) (1998) Nypa fruticans Wurmb - Boo, Chen & Choo 114 (SINU) Pandanaceae Pandanus dubius Spreng. - Boo, Chen & Choo 138 (SINU) Passifloraceae Passiflora foetida L. - Boo, Chen & Choo 71 (SINU) Rhamnaceae Gouania leptostachya DC. - A. Latiff & A. Zainudin 133 (UKMB) Rubiaceae Canthium glabrum Blume - A. Latiff & A. Zainudin 135 (UKMB) Chasalia chartacea Craib - Ramli Khamis 41 (UKMB) Hedyotis biflora (L.) Lam. - Boo, Chen & Choo 139 (SINU) Hedyotis dichotoma Koenig ex Roth - Boo, Chen & Choo 130 (SINU) Lasianthus barbellatus Ridl. - Md Noor & Samsuri 48 (SENG) Morinda citrifolia L. - Boo, Chen & Choo 101 (SINU) Prismatomeris tetranda (Roxb.) K. Schum. ssp. malayana (Ridl.) J.T. Johanss. - Md. Noor & Samsuri 23 (SING) Tarenna mollis (Wall. ex Hook.f.) B.L. Rob. - Shafiee Daud 47 (UKMB) Rutaceae Atalantia monophylla (L.) DC. - Boo, Chen & Choo 136 (SINU) Glycosmis mauritiana (Lam.) Tanaka - Boo, Chen & Choo 119 (SINU) Murraya paniculata (L.) Jack - Boo, Chen & Choo 110 (SINU) Sapindaceae Allophylus cobbe (L.) Raeusch. - Boo, Chen & Choo 51 (SINU) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 79 Sapotaceae Pouteria linggensis (Burck) Baehmi - Md. Noor & Samsuri 35 (SING) Simaroubaceae Brucea javanica (L.) Merr. - Md. Noor & Samsuri 12 (SING) Solanaceae Physalis ?pubescens L. - Boo, Chen & Choo 49 (SINU) Solanum nigrum L. - Boo, Chen & Choo 79 (SINU) Solanum torvum Sw. - Md. Noor & Samsuri 15 (SING) Stemonaceae Stemona curtisii Hook.f. - Boo, Chen & Choo 104 (SINU) Sterculiaceae Leptonychia caudata (Wall. ex G. Don) Burret - Md. Noor & Samsuri 42 (SING) Turneraceae Turnera ulmifolia L. - Boo, Chen & Choo 5 (SINU) Ulmaceae Trema tomentosa (Roxb.) Hara - Md. Noor & Samsuri 5 (SING) Verbenaceae Callicarpa longifolia Lam. - Boo, Chen & Choo 15 (SINU) c Clerodendrum calamitosum L. - Boo, Chen & Choo 4 (SINU) Clerodendrum inerme (L.) Gaertn. - Boo, Chen & Choo 27 (SINU) c Clerodendrum serratum (L.) Moon - Arishah Hashim 90 (UKMB) c Clerodendrum thomsonae Balf.f. - Md. Noor & Samsuri 11 (SING) Lantana camara L. - Boo, Chen & Choo 22 (SINU) Vitex trifolia L. - Boo, Chen & Choo 108 (SINU) Violaceae Rinorea bengalensis (Wall.) Kuntze - Md. Noor & Samsuri 43 (SING) Vitaceae Ampelocissus cinnamomea (Wall.) Planch. - Saidah Mamat s.n. (UKMB) Cissus nodosa Blume - A. Latiff & A. Zainudin 138 (UKMB) Cissus repens Lam. - A. Latiff & A. Zainudin s.n. (UKMB) Tetrastigma leucostaphylum (Dennst.) Alston ex Mabb. - A. Latiff & A. Zainudin 122 (UKMB) Tetrastigma pedunculare (Wall. ex Lawson) Planch. - A. Latiff & A. Zainudin s.n. (UKMB) Pulau Besar PTERIDOPHYTA Adiantaceae Adiantum stenochlamys Baker - C.L. Loh PB-20 (SINU) Blechnaceae Blechnum finlaysonianum Wall. ex Hook. & Grev. - C.L. Loh PB-30 (SINU) Stenochlaena palustris (Burm.) Bedd. - C.L. Loh PB-22 (SINU) Oleandraceae Nephrolepis auriculata (L.) Trimen - C.L. Loh PB-28 (SINU) 80 Schizaeaceae Lygodium circinnatum (Burm.f.) Sw.- CL. Loh PB-4 (SINU) SPERMATOPHYTA Anacardiaceae c Anacardium occidentale L. - C.L. Loh PB-37 (SINU) Apocynaceae c Thevetia peruviana (Pers.) K. Schum. - C.L. Loh PB-16 (SINU) Asclepiadaceae Hoya verticillata (Vahl) G. Don - C.L. Loh PB-10 (SINU) Cycadaceae Cycas rumphii Mig. - C.L. Loh PB-41 (SINU) Erythroxylaceae Erythroxylum cuneatum (Miq.) Kurz - C.L. Loh PB-42 (SINU) Euphorbiaceae Antidesma cuspidatum Miill.Arg. - C.L. Loh PB-3 (SINU) Breynia reclinata (Roxb.) Hook.f. - C.L. Loh PB-17 (SINU) Macaranga heynei 1.M. Johnst. - C.L. Loh PB-40 (SINU) Phyllanthus oxyphyllus Mig. - C.L. Loh PB-23 (SINU) Suregada multiflora (Juss.) Baill. - C.L. Loh PB-33 (SINU) Guttiferae Calophyllum inophyllum L.- C.L. Loh PB- 36 (SINU) Lecythidaceae Barringtonia macrostachya (Jack) Kurz - C.L. Loh PB-26 (SINU) Gard. Bull. Singapore 50(1) (1998) Leguminosae Dendrolobium umbellatum (L.) Benth. - C.L. Loh PB-9 (SINU) c Tamarindus indica L. - C.L. Loh PB-19 (SINU) Loranthaceae Dendrophthoe pentandra (L.) Mig. - C.L. Loh PB-13 (SINU) Malvaceae Hibiscus tiliaceus L. - C.L. Loh PB-14 (SINU) Moraceae Ficus fistulosa Reinw. ex Blume - C.L. Loh PB-38 (SINU) Moringaceae c Moringa oleifera Lam. - C.L. Loh PB-12 (SINU) Myrsinaceae Ardisia elliptica Thunb. - C.L. Loh PB-1 (SINU) Myrtaceae Rhodomyrtus tomentosa (Aiton) Hassk. _- CL. Loh PB-45 (SINU) Syzygium grande (Wight) Walp. - C.L. Loh PB-11 (SINU) Oleaceae Olea brachiata (Lour.) Merr. - C.L. Loh PB-31 (SINU) Opiliaceae Cansjera rheedei J.F. Gmel.- C.L. Loh PB- 8 (SINU) Polygalaceae Polygala paniculata L. - C.L. Loh PB-46 (SINU) 2. The Floras of Pulau Aur and Pulau Pemanggil, with Notes on the Smaller Islands 81 Rubiaceae Guettarda speciosa L. - C.L. Loh PB-18 (SINU) Timonius compressicaulis (Miq.) Boerl. - C.L. Loh PB-2 (SINU) Santalaceae Dendrotrophe varians (Blume) Mig. - C.L. Loh PB-48 (SINU) Sapindaceae Guioa pleuropteris (Blume) Radlk. - C.L. Loh PB-7 (SINU) Simaroubaceae Brucea javanica (L.) Merr. - C.L. Loh PB- 25 (SINU) Theaceae Gordonia multinervis King - C.L. Loh PB- 5 (SINU) Umbelliferae Centella asiatica (L.) Urban - C.L. Loh PB- 34 (SINU) Verbenaceae Vitex pinnata L. - C.L. Loh PB-29 (SINU) Pulau Yu PTERIDOPHYTA Oleandraceae Nephrolepis acutifolia (Desv.) H. Christ. - B. Molesworth Allen 2575 (SING) SPERMATOPHYTA Celastraceae Salacia chinensis L.- B. Molesworth Allen s.n., 30 July 1955 (SING) Guttiferae Calophyllum soulattri Burm.f. - B. Molesworth Allen s.n., 30 July 1955 (SING) Myrsinaceae Ardisia elliptica Thunb. - B. Molesworth Allen s.n., 30 July 1955 (SING) Rubiaceae Gynochthodes sublanceolata Mia. - B. Molesworth Allen s.n., 30 July 1955 (SING) Ixora javanica (Blume) DC. - B. Molesworth Allen s.n., 30 July 1955 (SING) Timonius compressicaulis (Miq.) Boerl. - B. Molesworth Allen s.n., 30 July 1955 (SING) Sapindaceae Guioa pleuropteris (Blume) Radlk. - B. Molesworth Allen s.n., 30 July 1955 (SING) Mischocarpus sundaicus Blume - B. Molesworth Allen s.n., 30 July 1955 (SING) Verbenaceae Premna serratifolia L. - B. Molesworth Allen s.n., 30 July 1955 (SING) Ck ’ ‘ ae : an : ae : ere tle. s 1 oe Mal ay ' 1? Sesh hes i ui e: be Pa ‘ Pyle f ' j 7 i te wy ‘4 a ' . _— Pyial ‘ 7 . " : P ; ne - ( 7 4 fd +, | ‘| | . - ; ha : ory aA j Pe Td - : i. ae : 7? . > . pepe eee b < mar ‘ are ’ Pale 36 6h ay ( : i eat | ' .* : Gardens’ Bulletin Singapore 50 (1998) 83-98. Unravelling Pinanga patula (Palmae) sensu Scheffer, Beccari and Ridley non Blume. LIM CHONG KEAT Palm Search Malaysia Project 215 Macalister Road, 10450 Penang, Malaysia Abstract Preparatory to a revision of the genus Pinanga Blume as found in Peninsular Malaysia, three taxa hitherto related to P. patula Blume by Scheffer, Beccari, and Ridley are discussed in the light of uncertainties pertaining to Blume’s species. The paper presents fresh nomenclatural notes on P. riparia Ridley, and describes P. auriculata Becc. var. merguensis Becc. as a new combination, and a new variety, P. auriculata Becc. var leucocarpa.. Introduction In 1838, Blume (Bull. Neerl. 1:65) introduced the genus Pinanga and described a number of new species, including P. patula Blume, based on specimens from the interior mountains of Sumatra collected by his friend Praetorius. The type location of P. patula has yet to be identified. From the text and illustration in his subsequent publication (Rumphia ii, 86, 87, t.115), the taxon was clearly characterised by having four to seven pairs of leaflets, and inflorescences having two rachillae. The herbarium specimens at Leiden (lectotype: Herb. Lugd. Bat. 900-182-241, L; syntype: 900-182- 241, L) display pinnate leaves with up to five pairs of leaflets, “spreading, falcate-lanceolate, acuminate” on laminas small enough to lie comfortably within the dimensions of a herbarium sheet (Plate 1), and are thus smaller than those of the taxa subsequently considered to be conspecific or varieties. Contemporaneously, Martius held a different view of Blume’s genus, and placed the new taxa under Seaforthia in his publication Historia Naturalis Palmarum (1837-1850). Later, in 1855, Miquel also disregarded Pinanga and relisted the lot under Ptychosperma, redescribing Ptycosperma patula (Flora van Nederlandsch Indie.3:26) presumably using the same original specimens, but in greater detail, indicating that the stem was three to four feet tall. As with Blume, he was silent on whether the taxon was solitary or clustering. Scheffer, then Director at Hortus Botanicus Buitenzorg (now Kebun Raya Bogor), subsequently revised Miquel’s account, reinstating Pinanga, including P. patula and the other Blume species with further descriptions (Natuurkundig Tjidschrift voor Ned. Indie. 32, 1871), freshly indicating 84 Gard. Bull. Singapore 50(1) (1998) that P. patula was stoloniferous (as distinct from being caespitose). He, however, appears to have added further cloudiness by reference to other taxa he considered to be conspecific or related, including P. inaequalis Blume, P. minor Blume, P. furfuracea Blume, and P. junghuhnii Blume. In 1876, he elaborated further on his Pinanga listing, and published photographs of the palms growing at Buitenzorg, including the clump labelled “P. patula”, which might well be the same still to be seen at present-day Bogor. I believe, however, that it is not the same as the Blume taxon, as will be explained below, and it can be suggested that Scheffer had not been familiar with the original species from the type location, and has misled Beccari and others in this identification. Beccari made three visits to Bogor, first meeting Scheffer on his way to New Guinea in 1871, and in 1874 when he became acquainted with the Javan flora. During these visits, he had so accepted P. patula sensu Scheff. as a distinct and stoloniferous species that during his third trip in 1878, when he travelled to Padang Pajang and Gunung Singalang he did not appear to have tried to find the Blume species. In 1885, after Scheffer’s death, he wrote up Reliquiae Schefferianae, obviously accepting P. patula sensu Scheff., and in Malesia 3, reconfirmed his concept of the taxon and its variety, P. patula var. junghuhnii Scheff., describing the latter as a “mountain form” of the species (citing his own specimen from Lubu-Raja, at 3000-4000 ft altitude). It would appear that after Praetorius, there had been no subsequent collections over the next 30 years or so; specimens by Korthals at Leiden are undated and without location notes. Collections from locations near Palembang were later made by Grashof (c. 1915), and by Teysmann probably earlier. In 1971-73, Dransfield collected from Gunung Tujuh and G. Kerinci at 1400-1900 m, but labelled his specimens (e.g. JD2689, K) tentatively “aff. P. patula”; whereas his specimen from Jambi (JD2555, BO, K) from a peat swamp were called “P. patula”, but they resemble more closely P. patula sensu Scheff. (as will be discussed later). In Sarawak, Beccari had begun to find innumerable new species, including the solitary and distinctive P. auriculata Becc. (Malesia. 3, 1886: 134-135), which he clearly considered to be distinct from the clustering P. patula sensu Scheff.. Viewing Beccari’s own collections in Florence, we can observe interesting but curiously variable determinations of specimens sent to him between 1866 and 1892, and later, with particular reference to the Malayan ones. Several, which he labelled as P. patula have to be regarded as incorrect or dubious. He also began to coin new varieties, which were not published, as far as I have been able to discover, e.g. “var. kalamantanica”, “var. lianggagangensis”, and “var. borneensis”. The last- named was presented as P. patula Blume forma borneensis by Winkler 85 Unravelling Pinanga patula (Palmae) (1913), together with P. patula Blume var. microcarpa Becc., also from Borneo. Following Beccari’s wishes, posthumously Martelli published P. patula Blume var. merguensis Becc. (in Martelli, 1934), and in 1935 relisted: P. patula var. celebica Scheff. (which he proposed to be synonymous with P. inaequalis, P. minor, and P. furfuracea - a three-way puzzle to be resolved elsewhere), P. patula var. gracilis Scheff. (synon. of P. gracilis Blume), P. patula var. junghuhnii Scheff. from Sumatra, and introduced P. patula Blume var. riparia Becc. in Martelli, the last-named being a reduction of P. riparia Ridley — which will be discussed later. Plate 1. Pinanga patula Blume (lectotype: Praetorius, Herb. Lugd. Bat. 900-182-241, L). By courtesy of Rijksherbarium, Leiden. 86 Gard. Bull. Singapore 50(1) (1998) § 2 eat Plate 2. Pinanga riparia Ridley, cultivated in Singapore Botanic Gardens, c.1934. By courtesy of Singapore Botanic Gardens. Plate 3. Pinanga paradoxa (Griff.) Scheff., Plate 4. Pinanga riparia Ridley, leaves. leaves (*H0942). Unravelling Pinanga patula (Palmae) 87 Significantly, especially in the context of this account, Beccari determined the specimen Ridley 3158 from Kuala Tenok, Pahang collected in 1891 as P. patula Blume, thus “importing” the nomenclature into Peninsular Malaysia. Another specimen collected in 1892 from Ulu Bubong, Perak (King’s Collector 10702, K, FI and CAL) was similarly cited by Beccari and J.D. Hooker (and propagated as such by a fine drawing in the Bentham Trust) further contributed to the spread of this error. I am positive, however, that this is a specimen of P. pectinata Becc. & J.D. Hooker, which is distinct, as will be discussed more fully in my revision (in prep.) of Pinanga in Malaya. Hooker had asked Beccari to collaborate on the palm section of Flora of British India but, although identified as precedent co-author, it is believed that Beccari had not responded to the invitation. From the correspondence between the two, it emerges that in 1886, Hooker had wanted Beccari (who was then preoccupied with the third volume of his | own Malesia) to go to Kew to work on the Scortechini material; in September 1891, he offered Beccari 15 pounds sterling to provide diagnoses and descriptions of the Indian species. In the above-mentioned Flora itself, J.D. Hooker published P. pectinata based on King’s Collector 4393 together with other Pinanga taxa, but called P. patula Blume “a doubtful species”! Ridley, in Materials for a Flora of the Malay Peninsula (1907) and Flora of the Malay Peninsula (1925) reinstated P. patula as a Malayan species, citing his own Kuala Tenok collection, and the Ulu Bubong one mentioned above, thus leaning on Beccari’s authority. Here, sensu Ridley, three different taxa are being confused. Although his Pahang specimen was of a solitary species, Ridley described the lot as “tufted”. He, however, chose to ignore Beccari’s efforts to sink his P. riparia (1905), but in this paper, P. patula var. riparia (1935) will revert to being a synonym. This also contradicts Whitmore’s taxonomic note (Principes. (1970) 14:125), where he incorrectly deemed P. riparia to be synonymous with P. pectinata, but suggested that P. patula sensu Ridley was distinct. It is obviously urgent and desirable to seek out the “real” P. patula and to collect fresh herbarium and live specimens (for propagation) from the probable type location in Sumatra, on the mountains. The prominent clumps labelled as this taxon in Kebun Raya Bogor and also those previously in the Singapore Botanic Gardens (Plate 2) - as shown in a photograph c.1934 by a Captain Johnstone, which correspond with herbarium specimens originally labelled “P. disticha”, but determined by Furtado in 1929 as “P. patula var.” are indistinguishable from P. riparia Ridley, which is a stoloniferous species found in low and wet places, and should now be recognised as such. It appears to adapt well to garden conditions, as evidenced at Bogor, and is indeed a handsome horticultural attraction. 88 Gard. Bull. Singapore 50(1) (1998) Dransfield (1974) believes that his specimen (JD 3590, BO, K, L, SING) from Bengkulu at 500 m altitude matches with the type; his, however, has larger leaves and inflorescences with three or more branches. Another specimen (JD 2679, BO, K) collected at 800 m from Kepahiang, Bengkulu (which occupies six sheets), displays various forms of leaf dissection, including one that does seem similar to the Praetorius specimen at Leiden. I would be inclined, however, to defer a definitive verification of P. patula Blume until field visits to the “interior mountains of Sumatra” yield more conclusive results. My suspicion is that the elusive palm may prove to be closer in appearance to P. paradoxa (Griff.) Scheff. (Plate 3) and P. salicifolia Blume, from Peninsular Malaysia and Borneo respectively. The Praetorius specimens have slender stems of similar dimensions to these. With regard to the species of relevance to Peninsular Malaysia, I propose to address the following three taxa: P. riparia (also found in Thailand, and probably once in Singapore), P. patula var. merguensis (now known to be widespread in South Thailand and in Perlis), and P. patula sensu Beccari and Ridley non Blume (in Thailand, Peninsular Malaysia and Singapore). I. Pinanga riparia Ridley. J. Roy. As. Soc. Str. Br. 44 (1905) 201. Type: Johor: Sg. Tebrau, 1903, Ridley 11518 (SING — lectotype here chosen, K iso). Synon : P. patula Blume var. riparia Becc. in Martelli. Nuov. Giorn. Bot. Ital. NS 42 (1935) 71. Type: Johor, Kukub, 1909, Ridley 14170 (FI, SING). Notes: Viewing the herbarium specimens in Leiden, I became convinced that Ridley’s taxon is not related to P. patula Blume. Although they may both be clustering species, P. riparia is distinctly stoloniferous, and has laminas which are usually larger, have more numerous leaflets, and broader apical ones (Plate 4). From Ridley’s accounts and field familiarity, I was also positive that the Bogor clump is P. riparia and not Blume’s taxon, which, as conjectured above, neither Scheffer nor Beccari had the opportunity of verifying from live specimens. The error becomes clear from Beccari’s account of P. patula (Malesia. 3: 139-140), from which we learn that the Bogor live specimen had been collected from Banka by Teysman (and was similar to Beccari’s own find at Sungai Bulu in Padang), both undoubtedly from riverine habitat. Beccari further commented on Teysman’s Bornean collections from Kapuas and Sg. Landak, which he felt were varieties or other forms of P. patula sensu Scheffer. Presented with Ridley’s specimen 14170 collected from Kukup in 1909, he obviously could only treat it as a variety of P. patula sensu Scheffer. Unravelling Pinanga patula (Palmae) 89 John Dransfield (pers. comm.) believes that in Borneo, there may be other swamp-dwelling, stoloniferous taxa which relate with P. riparia, and might even be conspecific; indeed, collections of P. patula var. borneensis, and other specimens from Kalimantan, Brunei and Sarawak have to be reexamined (also in relation to the puzzling P. furfuracea) - an interesting prospect for further research in that domain. P. riparia is easily identifiable after acquaintance in the field; in its natural habitat, it is practically rheophytic. The shiny leaves vary not only in size, but also in number of pinnae, and the petiole and rachis are sometimes glaucous. The stolons arise often at a distance from the main plant, and the nodal sections of the stems are green, light or darker, and “unwoody”, often to 4m in height. The deflexed inflorescence has usually two rachillae distinctively purple (coral red initially), with elliptical light green drupes (Plate 10), turning red to black. Specimens in herbaria have sometimes been mislabelled as P. singaporensis Ridley (with which P. riparia is often sympatric in Johor), and which in turn has often been misidentified as P. pectinata. : Distribution: Thailand: Narathiwat (viz. Phengklai & Niyomdham, 1991); Peninsular Malaysia: Terengganu (Saw Leng Guan pers. comm.), Pahang, Selangor, Negri Sembilan, Johor; Indonesia: S. Sumatra, Banka. Habitat: peat swamps, river banks; not rare, but endangered by habitat destruction. Specimens examined: Thailand: Narathiwat, 1974, Larsen 33092 (K); Peninsular Malaysia: Selangor, Tanjong Karang, 1937, Nur 34126 (SING); Johor: Sg. Tebrau, 1903, Ridley 11518 (Type, K, SING), 1906, Ridley 13235 (K, SING), Kukub, 1909, Ridley 14170 (Type of P. patula var. riparia Becc. in Martelli, FI, SING), Sg. Sedili, 1935, Corner 29239 (K). 2. Pinanga auriculata Becc. var. merguensis (Becc. in Martelli) C.K. Lim comb. nov. Synon.: P. patula Blume var merguensis Becc. in Martelli. Atti. Soc. Toscana Scienze Naturali Res. Pisa Memorie 44 (1934): 125,126; (“merguiensis”: Nuov. Giorn. Bot. Ital. NS 42 (1935):71). Type: Myanmar: Mergui, Tarapon, 1911, Meebold 14380 (two sheets), (WSRL). 90 Gard. Bull. Singapore 50(1) (1998) Notes: From wider field observations and collections, this taxon is confirmed as widespread from Mergui and along the west coast of Peninsular Thailand — where indeed it had been collected by Kerr, Whitmore and others — and within Perlis, where it was collected in 1995 by L.G. Saw and C.K. Lim (H1837, H1840 KEP), and noted as a new record for Peninsular Malaysia (Lim, Principes 42: 115). It is a solitary species clearly unrelated to the Sumatran taxon, P. patula Blume, as discussed earlier. Observing the striking similarity in habit and habitat of P. auriculata Becc. (1886), which Beccari collected at Kuching (holotype PB589, FI), I propose to transfer var. merguensis to varietal status under it. Although I had felt an earlier hesitance and reluctance to “cross the Sunda shelf” to relate Peninsular Malaysian and Bornean Pinanga species, recent field trips to Sarawak have provided new perceptions. Furthermore, the two taxa display many features in common, and it would seem that Pinanga taxa with affinities to P. auriculata may be quite widespread in the western Malesian region. The bifid eophyll and juvenile leaves (Plate 6) are quite indistinguishable within the group (and similar also with P. limosa Ridley); the prophylls are also similar, and dry into papery tatters. Ligules or auricles subtending from the leafsheath, where the petiole splits away, are often variable even in P. auriculata var. auriculata, and may not always be prominent. | The leaflets of var. merguensis (and of another new variety to be described below) differ from those of P. auriculata var. auriculata, which are more numerous and longer; in the variety, these are more sigmoidal (Plates 7), with pinnae that may be closely or more distantly spaced. They both have inflorescences usually with four to six rachillae; in var. merguensis the infructescence, often profuse and abundant, has drupes which are distinctively shiny and almost translucent, wine-red, (Plate 11) before ripening black. Meebold’s fine specimens (Plate 5) has been well curated at Wroclaw (earlier known as Braslav). Beccari, who designated the type in 1913, originally annotated it as “P. patula Bl. forma merguensis Becc.”. Distribution: Myanmar: Mergui; Thailand: Ranong, Trang, Surat Thani, Phuket, Satun; Peninsular Malaysia: Perlis. Habitat: hill forests or lowland, riverine; not rare. Specimens examined: Thailand: Ranong, 1918, Kerr 16386 (K), 1927, Kerr 11763 (K), Trang, Khao Chong Nat. Park, 1979, Dransfield JD 5451 (K). Peninsular Malaysia: Perlis, Mata Ayer F.R., 1995, C.K. Lim H1837, H1840 (KEP), 1996, C.K. Lim H1942* Unravelling Pinanga patula (Palmae) 9] FRO A PP Eg { — ‘f f Ore - Bt. %, rape foke be Pit DAT + aie x [te xO . f j ; ae patele 1) [Ly 1 4uidethubAs 1% Plate 5. Pinanga auriculata Becc. var. merguensis (Becc. in Martelli) C.K. Lim (holotype: 1911, Meebold 14380, WSRL). By courtesy of WSRL. 92 Gard. Bull. Singapore 50(1) (1998) Plate 7. Pinanga auriculata var merguensis, leaves and fruit (*H1837). Unravelling Pinanga patula (Palmae) 93 *Note: Within this account, as in my other taxonomic papers, certain specimens (prefix: H) currently kept in the Palm Search Malaysia collection are cited to supplement herbarium collections examined. Although it is intended eventually to deposit more specimens in the major reference herbaria, many items represent field records of the in situ conservation status, which the PSM project is in the process of monitoring. 3. Pinanga auriculata Becc. var leucocarpa C.K. Lim var. nov. A varietate typica fructibus albidis in statu immaturo distinguibilis. Typus: Pahang, Kuala Tenok, 1891, Ridley 3158 (holotypus, SING; isotypus, FI, K) Synon nov.: P. patula sensu Ridley non Blume, Ridley. Materials for a Flora of the Malay Peninsula. 2 (1907) 143; P. bowiana Hodel. The Palm Journal 134 (1997) 35. (Type: Thailand: Narathiwat, 1997, Hodel & Vatcharakorn 1608, BK) Stem, size and habit similar to P. auriculata var. merguensis; similarly, leaves divided with six to eleven pairs of leaflets, sigmoidal, with four nerves, sometimes bullate, glabrous, light or dark green, lighter on underside; prophyll drying papery; inflorescence infrafoliar, pendent, with two to six rachillae, usually light green; floral pits distichously arranged, flowers not examined; drupes globose, 6 x 8 mm, distinctively creamy white with green tips when immature, ripening red to black. Geographical range: only along east coast of peninsular Thailand and Peninsular Malaysia, and Singapore. Notes: As discussed in the Introduction, the specimen from Pahang collected by Ridley was incorrectly determined by Beccari, who may not have known that the Malayan taxon was distinctively solitary. Ridley had later described it as “tufted” or with “several” stems, and also confused it with P. pectinata. Whitmore (1973: 92) was obviously aware that Ridley’s descriptions of P. patula were faulty, and from his field observations pointed out that it was a solitary species (Plate 8). In habit and leaf form, which are pronouncedly sigmoidal (Plate 9), it could sometimes be confused with P. auriculata var. merguensis, which it closely resembles, but can be distinguished by the fruit, which are more globose and creamy white (with green tips) when immature, or nearly mature (Plate 12) — hence the varietal epithet. The fruit is also reminiscent of those of P. limosa, which is a diminutive species Gard. Bull. Singapore 50(1) (1998) 94 Plate 8. Pinanga auriculata Becc. var. leucocarpa C.K.Lim, at Merapoh, Pahang. Plate 9. Pinanga auriculata var. leucocarpa, leaves and inflorescence (*H0583). Unravelling Pinanga patula (Palmae) Plate 12. Pinanga auriculata var. leucocarpa, inflorescence and fruit (*H1259). 95 96 Gard. Bull. Singapore 50(1) (1998) with spicate inflorescences. It should be noted that when dried, drupes tend to look quite similar to those of P. /imosa but are more elliptical or fusiform, and no longer globose. Both varieties of P. auriculata are quite variable in robustness and size — seen fruiting at heights varying from 2 to 4 m. In the inflorescence of var. /eucocarpa, the rachillae are usually light green, but coral red variants have been observed, with immature drupes not the usual creamy colour, but red; this rare variation has also been noticed in P. limosa. In an earlier paper (Lim, 1998), I determined that P. bowiana Hodel was conspecific with “P. patula Blume” — more correctly, with P. patula sensu Ridley non Blume. In the light of further research on the basionym, both will now be reduced to synonymity under P. auriculata var. leucocarpa,. In his account, Hodel did not seem aware of the many herbarium collections of the Malayan taxon, or of those collected previously in the Narathiwat area. His description also lacked the essential mention of the fruit and diagnostic colour of the drupes, but from familiarity with his collection sites, I feel sure that his specimen is of this particular variety. The Ridley specimen from Pahang is for Malesian taxonomy important and historical, and has drawn with it numerous other collections designated similarly. For this reason, it continues to serve as the type for the new variety. The many herbarium specimens hitherto labelled P. patula by Ridley and others, however, may now have to be redesignated, and sorted out to differentiate var. leucocarpa from var. merguensis, the convenient initial guide being the collection site, and better, from clear evidence of the inflorescence and fruit. | As a result of more extensive field monitoring, territoriality becomes a useful indicator where it can be reasonably sure that certain taxa are localised. In geographical range, var. Jeucocarpa is widespread along the east coast of Peninsular Malaysia from Singapore and Johor to Kelantan, and in the Narathiwat area of Southern Thailand, but has so far not been found in the western side of the Peninsula where var. merguensis is common, from Perlis northwards. Distribution: Thailand: Narathiwat; Peninsular Malaysia: Kelantan, Terengganu, Pahang, Johor; Singapore. Habitat: hill forests or lowland, swamp; not rare in Peninsular Malaysia. Specimens examined: Thailand: Narathiwat, 1997, Hodel & Vatcharakorn 1608, BK (Holotype of P. bowiana Hodel); Peninsular Malaysia: Terengganu, Kemaman, 1935, Corner30165 (K, SING), Ulu Setiu, 1977, J. Dransfield JD5175 (K), G. Padang, 1969, 7.C. Whitmore FRI 12749 (KEP), Sg. Kerbat, 1971, 7-C. Whitmore FRI 20222 (KEP), Kelantan, Bukit Batu Unravelling Pinanga patula (Palmae) 97 Papan, 1935, Henderson 29525 (SING), Pahang, Temerloh, 1967, Palmer 18 (SING), K. Kenyam, 1985, J. Dransfield JD6224 (K); Singapore: Bukit Timah, 1902, Ridley s.n., (SING). Acknowledgements To the Directors of the Herbaria at BK, BO, CAL, FI, K, KEP, L, SING and WSRL special thanks, and for permission to reproduce illustrations of types. For invaluable taxonomic advice, consultation and discussion: Dr. Chin See Chung, Dr. Ed de Vogel, Dr. John Dransfield, Dr. Ruth Kiew, Dr. C. Nepi, Dr. Saw Leng Guan, Dr. Wanda Stojanowska and Dr. Tim Whitmore. To the members of the Palm Search Malaysia crew, for their field efforts and support. References Beccari, O.D. 1885. Reliquiae Schefferianae. Ann. Jard. Bot. Buitenz. 2. Beccari, O.D. 1886. Nuovi studi sulle palme asiatiche. Malesia. 3:134-135, 139-140. Blume, C.L. 1838. Bulletin des Sciences Physiques et Naturelles en Neerlande. EGS. Blume, C.L. 1838-39. Rumphia. 2 (18): 87, t. 115. Dransfield, J. 1974. Notes on the palm flora of Central Sumatra. Reinwardtia. 8: 519-531. Hodel, D. 1997. New species of palms from Thailand. The Palm Journal 134: 35. Hooker, J.D. 1892. Palmae. Flora of British India 6: 411-412. Lim, C.K. 1998. Notes on recent palm species and records from Peninsular Thailand. Principes. 42: 110-119. Martelli, U. 1934. Generi, specie e varieta nuove di palme gerontogee della triba Arecaceae lasuate. inedite dal Dr. O. D. Beccari ed. Ordinate a cura di U. Martelli. Toscana Science Nature 44: 125,126. Martelli, U. 1935. La sinonimia dele palme gerontogee della tribu delleAreceae. Nuovo Giornale Bot. Ital. 42 : 71. Miquel, F.A.W. 1855. Flora van Nederlandsch Indie. 3: 19, 26. 98 Gard. Bull. Singapore 50(1) (1998) Phengklai, C. and C. Niyomdham. 1991. Flora in Peat Swamp Areas of Narathiwat. p. 263. Phikul Thong Study Centre, Bangkok. Pichi Sermolli, R.E.G. and C.G.G.J. van Steenis 1983. Dedication. Flora Malesiana. 9: 7-44. Ridley, H.N. 1905. New Malayan plants. Journal Royal Asiatic Society Straits Branch. 44:201. Ridley, H.N. 1907. Palmae. Materials for a Flora of the Malay Peninsula. 2:133-221. Ridley, H.N. 1925. Palmaceae. Flora of the Malay Peninsula.5: 1-72. Scheffer, R.H.C.C. 1871. Sur quelques palmiers du groupe des Arecinees. Natuurkundig Tyjidschrift voor Ned. Indie.32:173, 176-179. Scheffer, R.H.C.C. 1876. Sur quelques palmiers du groupe des Arecinees. Ann. Jard. Bot. Buitenz. 1: 148, 150,151, pl.19. Whitmore, T.C. 1970. Taxonomic notes on some Malayan palms. Principes. 14: 125. Whitmore, T.C. 1973. Palms of Malaya: Oxford University Press, Kuala Lumpur, Malaysia. Winkler, H. 1913. Beitrage zur Kenntnis der Flora und Pflanzengeographie von Borneo. Botanische Jahrbucher. 48:89. ERRATA Gardens’ Bulletin Singapore 48 (1996) Lim, C.K. Unravelling [guanura BI. (Palmae) in Peninsular Malaysia Page 62: Acknowledgements: Third paragraph, lines 9 & 10:- “Mr S. Nadarajah” should read “Mr D. Nadarajah” “Ahmad Ismail” should read “ Ismail Ahmad” “Mohamad Noor Jamalulail” should read “Mohd Nor Jamalulail” Lim, C.K. Palms in the Farquhar Collection of Natural History Drawings Page 70: Item 7, line 2: “tunngal” should read “tunggal” Page 72: Second paragraph, line 21: “Rottl.” should read “Rottb” line 25: “Findlaysonis” should read “Findlayson’s” Page 72: Fourth paragraph, line 11: after “mystery” insert: “had been” Page 73: Plate 6: The illustration is erroneously that of “Nipah: Nipa fruiticans” Gardens’ Bulletin Singapore 50 (1998) 99-114. Four New Pinanga Blume (Palmae) Species from Peninsular Malaysia LIM CHONG KEAT Palm Search Malaysia Project 215 Macalister Road, 10450 Penang, Malaysia with contributions by JOHN DRANSFIELD', RUTH KIEW? ann SAW LENG GUAN? Abstract Four new Pinanga species, all from Johor, are described: P. jamariensis, P. johorensis, P. palustris and P. pantiensis. Introduction Since 1989, the Palm Search Malaysia project has made innumerable and repeated trips around Peninsular Malaysia, gaining important field experiences and findings of new or forgotten species. Stimulated by fresh data, the genera /guanura Blume and Pinanga Blume have been given priority for updating and revision — a process of “unravelling”, especially because of historical uncertainties, inherent in the monumental and strenuous efforts of earlier collectors including H.N. Ridley and others, in the determination of some herbarium specimens. The /gwanura_ revision has since been published (Lim, 1996). To facilitate the ongoing revision of Pinanga within Peninsular Malaysia, I decided to sort out certain vexatious aspects relating to P. patula sensu Scheffer, Beccari and Ridley non Blume (Lim, 1998), and now, for taxonomic convenience, to publish four new species, which have been in draft since 1994 or earlier. The taxa are all coincidentally from Johor: P. jamariensis C.K. Lim, P. johorensis C.K. Lim & L.G. Saw, P. palustris Kiew, and P. pantiensis J. Dransfield. Saw Leng Guan had shared the discovery of P. johorensis, for which he is co-author. We gladly decided to honour the State of Johor by the epithet, as indeed it is quite widespread there (several previous collectors ‘Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, UK *Singapore Botanic Gardens, Cluny Road, Singapore 259569 *Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Malaysia 100 Gard. Bull. Singapore 50(1) (1998) having considered it to be P. malaiana (Mart.) Scheff., e.g. Whitmore FRI 0187, SING). P. jamariensis and P. pantiensis are more localised in their known habitat, the latter being probably more seriously endangered by forest clearance at Linggiu and Gunung Panti, where an in situ conservation effort would be most desirable and urgent. P. palustris, although earlier thought to be localised to the Endau area, is now known to be quite widespread not only in Johor but also along the east coast of the Peninsula up to Terengganu. As with P. johorensis, with which it often shares its habitat, many early collections of this taxon have been labelled as P. malaiana, e.g. Tan Ah King 23 from Mawai, 1959, SING (but note: Tan Ah King 23A, SING, collected contiguously is P. johorensis). It is also one of the few Malayan Pinanga species that appears to have an affinity with Sarawak ones, in particular, P. mirabilis Becc. (1886). Note: Within this account, as in my other taxonomic papers, certain specimens (prefix: *H) currently kept in the Palm Search Malaysia collection are cited to supplement herbarium collections examined. Although it is intended eventually to deposit more specimens in the major reference herbaria, many items represent field records of the in situ conservation status, which the PSM project is in the process of monitoring. 1. Pinanga jamariensis C.K.Lim sp. nov P. auriculatae var. merguensi similis sed foliis parvidissectis et glaucis bene distincta. Typus: Johor: Bukit Jamari, 1993, C.K.Lim H1456 (holotypus SING, isotypus KEP). Plates 1-4. Solitary, stilt-rooted; stem grey-brown, erect, 3—8.5m tall, slender, 2 cm diam., internodes 5-12 cm. Crown with eight or more leaves; leaf sheath c. 28 cm, distinctly glaucous, white, tinged pink within; petiole 5 mm diam. to 30 cm long, glaucous; lamina thick and fleshy, 65 x 40 cm, glaucous, darker green above, lighter below and white to silvery, sometimes prominently whitish along nerves; blade often entire in juveniles, later divided into three or more irregular pairs of leaflets, with three to five nerves, leaves sometimes (rarely, e.g. *H/460) with serrated leaf edges. Inflorescence infrafoliar, pendent, with 3-4 branches; prophyll thin, papery, brown, often lingering though shrivelled; peduncle short c. 10 cm, 6 mm wide; rachillae slender, to 15 cm, reddish, with distichous floral pits. Staminate and pistillate Four New Pinanga Species from Peninsular Malaysia 101 flowers not seen. Fruit distichously borne, c. 24 pairs per rachilla; immature drupes light green with darker tips, ellipsoid, elongate and pointed, ripening to buff colour then blood red to black, broadening ellipsoid, c.12 x 10 mm; testa fibrous. Seedling leaf entire-bifid, acute, dark green, glabrous. Notes: This handsome and elegant palm is relatively rare, found so far in Johor from Gunung Panti (where I first saw it) to Kahang, Mersing Forest Reserve, and at its type location in Bukit Jamari (Plate 3), which its epithet identifies. Its glaucousness is indeed quite diagnostic, and the thick white Plate 1. Pinanga jamariensis C.K.Lim (holotype: 1993, C.K. Lim H1456, SING). By courtesy of SING. 102 Gard. Bull. Singapore 50(1) (1998) Plate 3. Pinanga jamariensis, solitary Plate 4. Pinanga jamariensis, young adult palm at Bukit Jamari, Johor. plant. Four New Pinanga Species from Peninsular Malaysia 103 leaves with fewer and broad leaflets (Plates 2 & 4) tell it apart from P. auriculata Becc. var. leucocarpa C.K. Lim (synon: P. patula sensu Ridley non Blume, see Lim, 1998) found in the same areas, which, however, has leaves with more numerous leaflets that are glabrous and sigmoidal in shape, and fruits that are globose and creamy white when immature, resembling those of P. limosa Ridley. Juvenile stages of the new species may indeed also look similar to the diminutive P. limosa, which occasionally has glaucous leaves, entire or dissected, and similarly thick; the eophylls are practically indistinguishable, and suggest an affinity within what might be called the ‘limosoid group’. Curiously, in these two Pinanga taxa, serrations to leaf edges beyond the apical teeth have been observed (which I have also seen in P. subintegra Ridley), although as a rare occurrence. Although compared with P. auriculata Becc. var. merguensis C.K. Lim (1998), the precedent variety, in the diagnosis (the habit and infructescence are similar), the drupes of that taxon are, however, different in colour, being distinctively shiny, wine-red, and its leaves (similarly with var. leucocarpa) are glabrous, and quite different in dissection and shape. Furthermore, their respective domains are geographically distant and disjunct. The new species is often sympatric with P. auriculata var. leucocarpa, aS mentioned above, and also with P. limosa, P. simplicifrons (Miq.) Becc. and the other new species to be described in this paper, P. Johorensis, and P. singaporensis Ridley in the Kahang area and at Gunung Panti. It may be found fruiting at less than 2 m in height, contrasting with the surprisingly tall individuals towering at over 8 m, with disproportionately slender stems, able to endure in wind-sheltered habitat at Jamari, where P. Johorensis and Johannesteijsmannia altifrons Reichenb.f. & Zoll. are also luxuriant. This new and attractive Pinanga can easily become endangered due to deforestation, as at Kahang, where it is already rare, and may require protection. Distribution: Johor: Mersing F.R., Bukit Jamari, Kahang, Gunung Panti. Habitat: lowland dipterocarp forest, to 50 m a.s.l., not common. Specimens examined: Johor: Gunung Panti, 1990, C.K.Lim *H0515, Bukit Jamari, 1991, C.K.Lim H1004 , 1992, C.K.Lim H1149 , 1993, C.K.Lim H11455, 1993, C.K.Lim H1457 (SING), C.K.Lim H1402, H1459, H1460, H1526, 1994, C.K.Lim H1674, H1682, 1995, C.K.Lim H1895, H1923 , 1996, CR W195. 104 Gard. Bull. Singapore 50(1) (1998) 2. Pinanga johorensis C.K.Lim & L.G.Saw sp. nov A P. malaiana minor, rachillis plerumque 2-ramulis, longis stolonibus bene distincta. Typus: Johor: Lenggor F.R., 1993, L.G.Saw FRI 37435 (holotypus KEP, isotypus K). Plates 5-7. Clustering, pleonanthic, monoecious palm. Stem with basal suckers forming very loose clumps, stoloniferous with distant stems up to 3 m apart; stem to 7 m tall, slender to 3 cm diam.; nodal scars conspicuous, 1 cm wide, internode to 15 cm apart; stem surface green, sometimes sparsely lepidote, with brown scales. Crownshaft to c. 65 cm long, dark green, sometimes glaucous, conspicuously swollen in developing inflorescences. Leaves six to nine in crown; leaf sheath to 32 cm long, glaucous on freshly exposed parts, prominently lepidote on older parts; leaf with sheath to 1 m or longer; petiole to 38 cm long, c. 1 cm diam., slightly channelled adaxially, round abaxially, lepidote; leaflets acuminate, often five to seven regularly arranged on each side of rachis, broad with 4-5 nerves (sometimes with 17 to 22 pairs of leaflets, each with fewer nerves), the apical leaflets broader, very prominently toothed and deeply lobed; lamina up to 122 cm long by 75 cm wide, shiny green, coriaceous, drying dull greenish brown on upper surface, darker brown on abaxial surface. Inflorescence infrafoliar, pendulous; prophyll from immature inflorescence elliptic, strongly two- keeled, pink when fresh; peduncle short to 1.5 cm long, flattened, wide at the prophyll scar; rachillae two, rarely three, with floral triads arranged distichously. Immature staminate flowers asymmetrical, sessile; calyx with three free triangular unequal lobes, c. 2 mm long; corolla with three well- developed ovate lobes, joined shortly below; stamens c. 38. Immature pistillate flower sessile; globose, calyx with three triangular, ciliate-margined lobes, about the same size as calyx lobes; staminodes absent; ovary cylindrical to ovoid, c. 1.5 x 1 cm; stigma with short style c. 0.5 mm long, 0.5 mm wide; stigma irregularly lobed and flattened. Infructescence infrafoliar strongly reflexed, up to 22 cm long. Immature fruits buff coloured with pink tips, maturing to bright red and black, with black calyx and corolla, borne on coral red rachillae. Mature fruit ellipsoid to 3 x 1.5 cm, with a distinct low collar surrounding the apical stigmatic remains; epicarp smooth; endocarp with conspicuous longitudinal fibres; seed adhering to endocarp, 1.5 x 1.2 cm, attached basally; endosperm deeply and irregularly ruminate; embryo basal. Four New Pinanga Species from Peninsular Malaysia 105 Plate 5. Pinanga johorensis C.K.Lim & Plate 6. Pinanga johorensis, note toothed L.G.Saw, stoloniferous palm at apical leaflets. Bukit Jamari. Plate 7. Pinanga johorensis, inflorescences and fruit, Kahang, Johor (*H0744). 106 Gard. Bull. Singapore 50(1) (1998) Notes: This smaller relative of P. malaiana (Mart.) Scheffer has undoubtedly been often confused with its larger kin, and perhaps many herbarium specimens still exist under that appellation. It can frequently be seen along the road from Kluang to Jamaluang, where it is under threat from forest clearance, and at Bukit Jamari (Plate 5). Apart from the two-, sometimes three-branched rachillae (Plate 7), it can be differentiated by the slender stems growing out of surprisingly distant stolons, and its fewer broad leaflets with the apical leaflets prominently toothed (Plate 6), although multi-pinnate forms with narrower leaflets can also be found. In the field, the swollen leaf sheaths have been observed to be penetrated by insects eager to ravage the inflorescence within; one rarely sees exposed flowers in anthesis. After abscission the prophyll may sometimes be erect, but are usually deflexed. It is relatively widespread in Johor, justifying its epithet. From the Lenggor F.R. to Mersing, it grows sympatrically with P.limosa, P. palustris Kiew (see below), Nenga grandiflora Fernando, and N. pumila var. pachystachya (Blume) Fernando, I. geonomiformis Griff. ex Mart., I. asli C.K. Lim, and the rattans of the area including Korthalsia echinometra Becc., and K. flagellaris Miq.. Ridley’s 1903 specimen indicates its presence in Singapore. Further research might possibly yield collections in Sumatra and the Riouw islands. Distribution: Johor, Lenggor F.R., Mersing F.R., Bukit Jamari; Singapore. Habitat: lowland dipterocarp forest, to 80 m a.s.l.; common palm. Specimens examined: Johor: Kota Tinggi, Mawai, 1959, Tan Ah King 23A (SING), Kahang, Kg. Sri Lukud, 1990, C.K.Lim *H0512, 1991, C.K.Lim H0745, H0970, H1051, H1057, Bukit Jamari, 1991, C.K.Lim H1003, 1993, C.K.Lim H1458 (SING), H1400, H1525, 1994, C.K.Lim H1683, 1995, C.K.Lim H1896, H1924, Labis F.R., 1966, T.C.Whitmore FRI 0187 (SING), 1970, T.C.Whitmore FRI 15618 (KEP), 1993, C.K.Lim H1519 , Lenggor F.R., 1993, C.K.Lim H1588, H1589 ; Singapore: Bukit Panjang, 1903, Ridley 1841 (SING). 3. Pinanga palustris Kiew sp.nov. A P. malaiana fructibus grandibus_ infructescentia erecta et interfoliacea differt. Typus: Pahang: Sg. Kinchin, 1989.R. Kiew RK2806 (holotypus KEP). Plates 8-9. Four New Pinanga Species from Peninsular Malaysia 107 Robust, clustering palm, clumps c.1 m across at the base, consisting of 10 or more stems with leafy canopy more than 3 m across. Majority of stems in clump either short and completely covered by leafsheaths, or are basal suckers with undivided leaves. Juvenile undivided leaf with lamina up to 50-80 cm by 14-17 cm with a deep apical notch, apical leaflet prominently toothed, petiole c. 65 cm long. Tallest stem in clump 1.5-3.5 m tall and 3 cm thick with whitish annuli 1-2 cm apart, and c. 1 cm wide. Individual stems with c. 3 leaves. Crownshaft c. 25 cm long, lower 1-2 leafsheaths dead and partially rotten. Leafsheath 15 cm long, reddish-brown, or stems yellow within sheath, persistent. Petiole 1.5 m long, yellowish-green, glabrous, channeled above. Lamina pinnately divided, up to 2.5 m long and 90 cm wide, with six to eight pairs, not constricted at insertion, mid- leaflets c. 60 cm by 5 cm, each with three to four veins, veins minutely furfuraceous on lower surface, distal leaflets 35-40 cm by c. 6 cm, with deeply serrate margin, with teeth 1.5—3 cm long. Inflorescences interfoliar, produced in lower leaf axils and only emerging through rotten leafsheath when in fruit, glabrous, stout, erect, peduncle flattened 2—3 cm by 1-1.5 cm, thickening and becoming 2 cm wide in infructescence; rachillae two to three (rarely four), 10 cm long and 5-10 mm wide, flattened, in infructescence yellowish with ruby red or crimson hue. Fruit scar circular, flat c. 10-12 mm across. Prophyll 9 cm by 3.5 cm, rosy red or white flushed at apex when immature. Flower triads alternate and distichous, 3-4 mm apart. Male flowers (from immature inflorescence) with perianth parts fleshy, more or less triangular, stamens 30 plus (to 44), and sessile with oblong anthers. Female flowers with three imbricate sepals, broadly ovate with minutely apiculate apex, 4 mm by 8 mm; petals three, apically valvate, basally imbricate, with acute apex, 8 mm by 5 mm, margin finely ciliate. Ovary with capitate stigma. Immature fruit ellipsoid and peachy-pink in colour, swelling when ripening, ovoid, 30-35 mm by 18 mm and rosy-red to black. Calyx persistent, black in fruit. Epicarp smooth and matt with apical “nose” 3 mm long, mesocarp spongy, endocarp fibrous. Seed deeply ruminate. Notes: In Kiew and Dransfield (1987), this species was referred to as Pinanga aff. mirabilis and was subsequently listed as such in several checklists. The current epithet denotes its characteristic swamp and wet habitat. It does resemble the Bornean P. mirabilis Becc. not only in habit and habitat, but also in the large size of fruit (up to 25 x 12 mm in the latter). The new taxon is quite different from the other large Pinanga, P. malaiana, which has taller and distinct stems, although caespitose, not clumping, and having longer pendulous infructescences with red to shiny black drupes, which are not as large. 108 Gard. Bull. Singapore 50(1) (1998) AOWERT VEG AmE NEIYFOre COFON Maa cca » - WeLetyré Uae Prats per esee m Sa. lie on Seg BE MOLETY PE Pmtepn prdecg Wis ay Cre) SS Le Lite mt a: Nad, i “Sao ovnd Low trad) se Le tong OE moe po lnitit ee Plate 8. Pinanga palustris Kiew (holotype: 1989, R. Kiew 2806 KEP). By courtesy of KEP. Four New Pinanga Species from Peninsular Malaysia 109 Plate 9. Pinanga palustris, with leafsheaths stripped, at Kahang, Johor. P. mirabilis has leaves which are usually entire, although pinnate plants are not uncommon and are sometimes sympatric (Dransfield, 1991); the stems are taller, up to 5 m, but usually 2—3 m, cleanly exposed by the abscissing leaves, which also reveal pendent or infrafoliar inflorescences, although some have been observed to be erect as at Lambir Hills; its fruits are different in colour when immature, a brownish-pink, whereas they are coral red in P. palustris. Again, the Malayan species has characteristic orange stems, when revealed by stripping the rotting leafsheaths (Plate 9), and so far, no large entire-leaf forms have been encountered, barring the juveniles. 110 Gard. Bull. Singapore 50(1) (1998) As for P. johorensis, many herbarium specimens of this new taxon have been filed under P. malaiana. Furtado in his determinations (at SING) had noticed differences, and began to suggest comparisons with P. malayana (Griff.) Scheff. var. sumatrana Becc. or with P. malayana var. baramensis (P. malaiana (Griff.) Scheff. var. barramenis Becc. in Martelli), which are Sumatran and Bornean taxa respectively, the latter since reduced under P. mirabilis by Dransfield (1991). Distribution: Johor: Ulu Endau, Lenggor F.R., Kahang, widespread, also Pahang: Rompin, Kedaik. Habitat: usually along sides of streams, or in swampy places, sometimes on hills up to 300 m as observed at Ulu Endau on sandstone, growing among Johannesteijsmannia altifrons. Specimens examined Johor: Mawai, 1935, Corner & Furtado 29240 (K, SING), 1959, Tan Ah King 23 (SING), Sg. Kayu, 1937, Keah 32423 (SING), Gunung Panti East, 1892, Ridley s.n. (SING), 1973, J. Dransfield JD3044 (SING), Ulu Endau, Dransfield JD3548 (SING), B.H. & R. Kiew RK1613 (KEP), 1977, J. Dransfield & F.W. Fong JD5040 (K, KEP), Sg. Sempanong 1985, R. Kiew RK1743 (KEP), Kuala Marong, 1985, R. Kiew RK1761 (KEP), Kahang, 1990, C.K.Lim *H0279, H0426, H0533, Lenggor F.R., 1991, C.K. Lim H0933, L.G. Saw et alia FRI 37439 (KEP), Ulu Sedili, 1991, C.K. Lim H0929; Pahang: Sg. Kinchin (see type), Endau-Rompin, 1985, C.M. Low FRI 25900 (KEP), Kedaik, 1991, C.K. Lim H1001, Lesong F.R., 1992, L.G. Saw FRI 37532 (KEP), 1993, L.G. Saw FRI 38522 (KEP). 4. Pinanga pantiensis J.Dransf. sp. nov. Inter species Malayanas rachillis luteis vel aurantiacis valde fractiflexis bene distincta, P. pachyphyllae, specie Borneensis, verosimiliter affinis sed textura et dissectione folti et rachillis fractiflexis differt. Typus: Johor: Kota Tinggi, Gunung Panti.F.R., 1973, J. Dransfield et alia JD3048 (holotypus SING). Plates 10-13. Clustering undergrowth palm to 6 m tall. Stem c. 20 mm diam., green with grey-brown leaf scars; internodes 40-50 mm long, with scattered caducous black scales when newly exposed. Crownshaft c. 35 cm long; sheaths pale green, c. 25 cm long, striate when dry, minutely dotted with small caducous Four New Pinanga Species from Peninsular Malaysia i scales. Leaves arcuate, to 2 m long; petiole c. 50 cm long, c. 5 mm diam. near base; rachis light orange-yellow when fresh; leaflets 10-16 on each side of the rachis, arcuate, diverging at angle of about 30 degrees from the rachis, the longest to 38 x 3 cm, very coriaceous, glossy green when fresh, acuminate and consisting of three to ten folds except for the apical two leaflets on each side, consisting of three to ten folds and lobed to a depth of 1 cm at the tips; transverse veins conspicuous, close, leaflet surfaces glabrous, ramenta absent. Inflorescence infrafoliar, known only in immature to mature fruiting state, to 15 cm long with three to five branches; prophyll 14 x 4x 2 cm, thick, yellow green; peduncle c. 3 cm long, c. 8-10 mm wide at the base, tapering to 2 mm wide, rachillae conspicuously zig-zag, yellow to orange; rachilla bract triangular, c. 2-4 mm, flower scar 4 mm diam. Immature fruit green, mature fruit satiny-black, ovoid, 32-35 x 15-17 mm; epicarp minutely striate, pericarp c. 4 mm thick. Seed 10 x 20 mm, endosperm deeply ruminate; embryo basal. Seedling leaf coriaceous. Notes: When Dransfield recognised this as a new taxon, after viewing herbarium records deposited by Corner as early as 1936, and from his own collections, it was thought to be localised and endemic to the unique flora of Gunung Panti, hence the epithet. The species has since been found in adjacent areas in Johor, especially at Linggiu where the recently constructed dam has diminished its population, further threatening what is undoubtedly a rare palm. On a recent collection trip to that locality, on the stems of the few residual plants the internodes were seen to vary from 13 cm at the base, to 4 cm at the upper end, providing an indication of effects on growth, perhaps due to ecological change and disturbance. In appearance the taxon resembles P. malaiana, though it is not observed to be as tall or robust. Although clustering, it usually has one or two dominant stems (Plate 11). The pinnae are usually narrower and more widely spaced (than in P. malaiana), and are characteristically tough and stiff. The inflorescence is its striking feature (Plate 12), with zig-zag rachillae, often bright yellow in colour and glossy black fruit (Plate 13). The recently described palm from Khao Sok in Thailand, P. fractiflexa Hodel (1997), has wavy but green, and not so strongly fractiflexing rachillae. Distribution: Johor: Linggiu, Kota Tinggi, Gunung Panti F.R.(east). Habit: Hill slopes, ridge top, dipterocarp forest, to 250 m a.s.l. Specimens examined: Johor: Linggiu, 1992, C.K. Lim *H1343, 1993, C.K.Lim H1530, 1998, H2001; Kota Tinggi, 1957, T.C. Whitmore 63 (SING); G. Panti, Ulu Segun (300m alt) 1936, Corner SFN 30659 (SING); G. Panti 112 Gard. Bull. Singapore 50(1) (1998) ‘Ma y, y ‘ * | I aaa Fl ee Ii bs x Plate 10. Pinanga pantiensis J.Dransfield (holotype: 1973, JD3048 SING). By courtesy of SING. Four New Pinanga Species from Peninsular Malaysia i3 Plate 11. Pinanga pantiensis, dominant Plate 12. Pinanga pantiensis, inflorescences stem in clump, at Linggiu, Johor. infrafoliar. Plate 13. Pinanga pantiensis, zig-zag rachillae, and fruit (*H/343). 114 Gard. Bull. Singapore 50(1) (1998) (forested eastern slope, 300m alt.) 1967, 7.C. Whitmore FRI 4515 (KEP), 1967, Suppiah FRI 98978 (KEP); Ulu Sedili, 1991, C.K. Lim H0926; Acknowledgements Special thanks to Dr. John Dransfield, Dr. Ruth Kiew and Dr. Saw Leng Guan for generously contributing new species and collaboration, and to the Directors of the herbaria at L, K, KEP, SING and WSRL, and to friends at KEP, Dr. Lilian Chua, Kamarudin bin Salleh, Mat Asri bin Ngah Sanah, and at SING, Dr. Chin See Chung, Mohamad Shah bin Mohamad Noor, Ali Ibrahim, to my field collectors Adong Pandak, Alus Sarip, Hamid Busu, Adnan bin Yusuf, Busu Ngah, Mohamed Noh bin Muhamad, to Dr. Tim Whitmore for his ready help and advice, and to the Minister for Primary Industries, Malaysia, Datuk Seri Dr. Lim Keng Yaik, and the Departments in his portfolio for their assistance to the Palm Search Malaysia project. References Beccari, O.D. 1886. Nuovi studi sulle palme asiatiche. Malesia. 3:126. Dransfield, J. 1991. Notes on Pinanga (Palmae) in Sarawak. Kew Bulletin. 46:697. Hodel, D. 1997. New species of palms from Thailand, part II. The Palm Journal. 136: 19. Kiew, R. and J. Dransfield. 1987. An annotated checklist of palms at Ulu Endau, Johor, Malaysia. Malayan Nature Journal. 41: 257-265. Lim, C.K. 1996. Unravelling /guanura BI. (Palmae) in Peninsular Malaysia. Gardens’ Bulletin, Singapore. 48: 1-64. Lim, C.K. 1998. Unravelling Pinanga patula (Palmae) sensu Scheffer, Beccari and Ridley non Blume. Gardens’ Bulletin. 50: 83-98. Gardens’ Bulletin Singapore 50 (1998) 115-119. ‘Some New Eastern Gingers’ — a Paper by H.N. Ridley Containing Descriptions of Four Species Overlooked since their Publication in 1900 I.M. TURNER Center for Ecological Research Kyoto University Shimosakamoto Otsu 520-01, Japan AND M.R. CHEEK Herbarium Royal Botanic Gardens Kew, Richmond Surrey TW9 3AB, UK Abstract Attention is drawn to four Malesian species of ginger (Alpinia pectinata, Alpinia celebica, Amomum terminale and Tapeinochilos koordersianus) validly published by H.N. Ridley in 1900 that do not appear in Index Kewensis. We lectotypify Alpinia pectinata Ridl., a new synonym of Alpinia eremochlamys K. Schum. Alpinia celebica Ridl. pre-dates Schumann’s use of the same combination. Introduction In the July 1900 issue of the Journal of the Straits Branch of the Royal Asiatic Society, there appeared on pages 97-99 a short, anonymous, paper entitled ‘Some New Eastern Gingers’ (Ridley 1900). The paper contained descriptions of four species named as Alpinia pectinata, Alpinia celebica, Amomum terminale and Tapeinocheilus koordersiana. The only one of these names to appear in Index Kewensis is Alpinia celebica, which is attributed to K. Schumann in a publication of 1899 (though this was a nomen nudum only validated in 1904). Therefore, if these are validly published names, they require to be circulated to prevent the unwitting adoption of later homonyms when new names or combinations are put forward. 116 Gard. Bull. Singapore 50(1) (1998) Validity of the Four Species It is first necessary to establish the author of these names. This is not difficult. At the time of the paper’s publication, H. N. Ridley was Director of the Singapore Botanic Gardens and a well-known authority on the Scitamineae (Zingiberales) of tropical Asia. The first sentence of the paper states: “The following new species of Scitamineae have passed through my hands since writing the paper published in Journal No. 32 and do not appear in Schumann’s paper.’ Ridley (1899) published a monograph of ‘The Scitamineae of the Malay Peninsula’ in the Journal of the Straits Branch of the Royal Asiatic Society No. 32, pp. 85-184, leaving no doubt that Ridley must be the author of the ‘New Eastern Gingers’. The Schumann paper referred to is probably K. Schumann’s ‘Monographie der Zingiberaceae von Malaisien und Papuasien’ (Schumann 1899). The species descriptions of the four gingers are written in English, which does not invalidate the names for a publication of this date. The descriptions are reasonably detailed, certainly sufficiently so, to make rejection of the names on such grounds untenable. Finally, there is the problem of typification. There is no specific mention of herbarium specimens directly associated with any of the species described. Instead after the diagnosis of each species there is a locality stated. For three of the species this is Celebes (the former name of Sulawesi), with the following extra information being given for Alpinia pectinata: ‘at Gunong Klabat 1300 to 1600 metres elevation, fruiting in January.’ The fourth species, Amomum terminale, is referred to as from ‘Bismarck Archipelago (Micholitz.) Flowered in Botanic Gardens, Singapore, Feb. 1900.’ Micholitz is known to have collected in the Bismarcks (van Steenis-Kruseman 1950), and presumably he sent live material to Singapore for Ridley to have cultivated in the Botanic Gardens. The final sentence of the paper may offer a clue to the identity of the collector of the specimens that Ridley was sent from Celebes. He writes: ‘I have great pleasure in associating it [the new species of Tapeinochilos| with the name of Dr. Koorders, who made such fine collections of plants in Celebes recently.’ Koorders collected on Celebes in 1894-95 and visited Gunong Klabat on 17-19 January 1895 (van Steenis- Kruseman 1950). It seems highly probable therefore that the three species from Celebes described by Ridley should be typified by Koorders’ collections. Identity of Alpinia pectinata Ridl. Unfortunately, a thorough search of the herbarium of the Singapore Botanic Gardens (SING) has failed to locate any likely material to typify any of New Eastern Gingers 147 the four names. This may be accounted for by Ridley’s opening reference to the new species as having ‘passed through my hands.’ Possibly, Koorders sent material to Ridley for naming but without sufficient duplicates, at least of some collections, for any material to be lodged at SING. The absence of any collections of the Amomum terminale grown in the Singapore Botanic Gardens is less easily explained. A visit to Herbarium Bogoriense in Indonesia was more successful. A Koorders specimen exactly matching the collecting details indicated for Alpinia pectinata by Ridley was discovered. This we select as the lectotype for the species since it is determined ‘A/pinia pectinata Ridley’ apparently in Ridley’s hand and signed and dated by him ‘Ridley 10.x.99.’ Smith (1990), in her synopsis of Alpinia, referred to Alpinia pectinata Ridl. as a nomen nudum first employed by Holthuis (in Holthuis & Lam 1942), who was given the name on material identified at Bogor. Smith identified the species as Alpinia eremochlamys K. Schum., which she claimed was only validly published in 1904, with the 1899 publication of the name representing a nomen nudum. We cannot agree with this conclusion. Name, diagnosis and type specimens are all included in the original protologue of Alpinia eremochlamys. We conclude that Alpinia pectinata Ridl. is valid, and represents a new synonym of Alpinia eremochlamys as summarized below: Alpinia eremochlamys K. Schum., Bot. Jahrb. Syst. 27 (1899) 288. Syntypes: Sulawesi; Tomohon, Sarasin 412, 6 June 1894; Tondano, A.B. Meyer s.n., May 1871; Kandari Peninsula, Beccari, May 1874. Alpinia pectinata Ridl., J. Straits Branch Roy. Asiat. Soc. 34 (1900) 97 synon. nov.. Type: “Celebes at Gunong Klabat 1300 to 1600 metres elevation, fruiting in January,” Koorders 196508, 19 January 1895 (lectotype, selected here, BO!). Three Unidentified Species Type material for the three other Ridley names has yet to be found and we cannot therefore identify the species concerned with certainty, but we conclude that all three were validly published. However, it would be premature to propose any formal name changes or new synonyms. The full citation and Ridley’s original diagnoses (with their idiosyncratic punctuation) of the three species are given below as an aid to others who may want to attempt to resolve the problem. Alpinia celebica Ridl., J. Straits Branch Roy. Asiat. Soc. 34 (1900) 98. Type: “Celebes.” [non Alpinia celebica K. Schum., Pflanzenr. 20 (1904) 362. Type: Sulawesi, Riedel s.n. (K!, lectotype, selected by Smith, 1990)]. 118 Gard. Bull. Singapore 50(1) (1998) ‘A (Hellenia) Celebica n. sp. A herb more than 18 inches tall with glabrous very long pointed lanceolate leaves, 8 inches long 1'/2 inch wide, petiole terete striolate graceful one inch long ochrea oblong truncate. Panicle graceful erect branches short many flowered five inches long. Bracts caducous. Flowers 1!/2 inch long. Calyx tubular truncate '/2 inch long. Corolla tube twice as long, lobes oblong obtuse '/2 inch long. Lip narrow shorter than corolla, deeply bifid, lobes spathulate emarginate. Staminodes narrow subulate. Stamen with a rather long filament, another [sic] oblong not crested. Style graceful.’ Amomum terminale Ridl., J. Straits Branch Roy. Asiat. Soc. 34 (1900) 98. Type: “Bismarck Archipelago (Micholitz.) Flowered in the Botanic Gardens, Singapore, Feb. 1900.” ‘Amomum terminale n. sp. Stems crowded slender about 2 feet tall, or much taller '/s inch through. Leaves dark green, elliptic lanceolate acuminate thinly coriaceous pale beneath glabrous 7 inches long, 2 inches wide, petiole very short, ocrea [sic] '/s inch long rounded. Spike terminal or basal cylindric 4 inches long */s inches through. Bracts ovate obtuse margins hairy */4 inch long '/2 inch wide red. Bracteole '/4 inch long oblong obtuse pink. Flowers in pairs. Calyx tubular dilated upwards trifid pink 7/4 inch long. Corolla tube one inch long slender white, lobes lanceolate acute '/2 inch long. Lip three lobed, two lateral lobes shorter curved outwards, acute, median obovate obscurely lobed, '/2 inch long. Anther with a broad connective rounded crenulate.’ Ridley added after the diagnosis: ‘The habit of this plant and its red bracts cause it to resemble some species of Zingiber, but it has not the long anther beak of that genus. It is abnormal among Amomums in having the spike terminal, but it.is also said to produce basal spikes from the rhizome. It is indeed difficult to refer it to any genus but I am unwilling to make a distinct genus for it alone. In some respects it may be classed with an ornamental plant known as Costus Zebrinus of gardens, which however has no relationship with the genus Costus at all.’ Tapeinochilos koordersianus Ridl., J. Straits Branch Roy. Asiat. Soc. 34 (1900) 99, sphalm. Tapeinocheilus koordersiana. Type: “Celebes.” ‘Tapeinocheilus Koordersiana n. sp. A tall plant, 25 feet tall. Leaves broadly oblong nearly four feet long 8 inches wide, subcoriaceous pubescent or glabrous narrowed at the base. Spike subcylindric 8 inches long, 4 inches wide. Bracts stiff coriaceous not woody oblong or ovate cuspidate ribbed pubescent the larger ones 2'/2 inches long and one inch wide, the inner ones lanceate cuspidate pubescent longer. Bracteoles linear narrowed acute shorter than flowers. Calyx tube one inch long narrow little enlarged above, lobes lanceolate acute quite covered with silky hairs. Corolla tube hairy but little longer, lobes narrow acute. Lip oblong rounded hairy. Anther oblong hairy. Capsule an inch long obovate warted covered with brown wool.’ After is added: ‘Another species of this grand Eastern island genus, allied to Miquel’s 7. pungens but with larger flowers and pubescent bracts.’ Acknowledgements We are grateful to the directors of the Singapore and Bogor Herbaria for allowing us access to the material under their charge. D.J. Middleton kindly New Eastern Gingers 119 checked the Rijksherbarium for potential type material. R.K. Brummitt is thanked for valuable comments he gave on a draft of this paper. References Holthuis, L. B. and H. J. Lam. 1942. A first contribution to our knowledge of the flora of the Talaud Islands and Merotai. Blumea. 5: 93-256. Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal of the Straits Branch of the Royal Asiatic Society. 32: 85-184. Ridley, H.N. 1900. Some new eastern gingers. Journal of the Straits Branch of the Royal Asiatic Society. 34: 97-99. Schumann, K. 1899. Monographie der Zingiberaceae von Malaisien und Papuasien. Botanische Jahrbucher fiir Systematik, Pflanzengeschichte und Pflanzengeographie. 27: 259-350. Smith, R. M. 1990. Alpinia (Zingiberaceae): a proposed new infrageneric classification. Edinburgh Journal of Botany. 47: 1-75. Steenis-Kruseman, M. J. van. 1950. Malaysian plant collectors and collections being a cyclopedia of botanical exploration in Malaysia. Flora Malesiana, series 1, 1: 3-606. j 4 . i] ' .« : - 4 é J eo 1 - 7 . . p owe Math oles of f ’ ; : .} ry A) ‘4 iW iv Par Oe) } oe ae | y OIE fal) j 4 ye ‘ Hin! + é ‘ | } 4 Q via ‘ .% with ’ i 4 / as. 1 i” i a} ‘ tee i. j van 7 \y Wa: i] Vor aa) , AY hice it 4 * if * ; Phd vf > i = CAA 4 Real imeid J Py ey 40) ( Ue ’ a Dar of ; a ‘ 7 ’ : : ry } ‘ iM f y ’ Gardens’ Bulletin Singapore 50 (1998) 121-122. TREE FLORA OF PASOH FOREST Kochummen, K.M. 1997. Tree Flora of Pasoh Forest. Malayan Forest Records No. 44. Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Malaysia. xv + 416. Figs. 1-20, 29-42, coloured plates; figs 21-28 b&w line drawings. ISBN 983-9592-69-6. Hardbound. Price RM60.00. Available from the Director General, Forest Research Institute Malaysia. Pasoh Forest Reserve, a patch of rain forest in Negeri Sembilan, Malaysia, has been made internationally known for the biological research carried out in it since 1970, first under the auspices of the International Biological Programme (IBP) and later by the Smithsonian Tropical Research Institute (STRI). The research station in the reserve is managed by the Forest Research Institute of Malaysia (FRIM). In 1985 a joint research programme between STRI and FRIM, based on a 50 hectare plot, took off. All trees with a diameter of 1 cm and above were enumerated. The task of identifying all the 335,240 individuals (in 814 species, 290 genera and 78 families) was carried out by K.M. Kochummen and his team. As most individuals would not be flowering or fruiting at the time of sample, they had to be identified based on vegetative characters. This required a special skill of which Kochummen is the master practitioner and in this publication he has made available a useful summary of the work done. This book, therefore, is not only the definitive account of the flora of the 50 hectare plot, but also a practical manual to the art and science of identifying southeast Asian rainforest trees using vegetative characters. In the first part of the book, vegetative characters useful for the identification of trees are defined and, in many cases, illustrated. The second part consists of three keys to species or genera. The first key identifies selected trees with “spot” characters, for example, those with large leaves at least 20 cm long and 10 cm wide, with latex from cut bark, or whose boles have thorns; the second key uses mainly bole and bark characters and the third, leaf characters. As each key is broken into several sub-groups each with a new numbering sequence, it would have been useful at the beginning of each key to have an index listing the sub-groups and the pages on which they begin. Keys two and three list the sub-groups, but do not provide the page numbers. In the key using bole and bark characters, the bole surface is divided into four types, smooth or cracking, scaly, fissured, and dippled. However, in the illustrations on bole characters there are also lenticellate and cankered types; both of which appear more smooth than scaly, fissured or dippled. Where would they go in the key? As it turns out, Parartocarpus bracteatus 122 Gard. Bull. Singapore 50(1) (1998) (lenticellate bole), keys out under the sub-group “bole smooth or cracking.” However, Pterocymbium javanicum (cankered) does not appear in the key using bole and bark characters. | The third, and major part of the book, is a description of the families, genera and species covered. Where there is more than one genus per family, a key to the genera is provided under each family. However, this is not always consistent, sometimes in place of a Key to genera is a key to “genera and species” (Anacardiaceae) and sometimes there may be more than one key to the genera (Leguminosae, one generic key based on bole characters and another based on leaf characters). Following this, there is often a “forest key” to the species of the family. Again, there are some families in which this key is not provided. Subsequently each species is keyed under the respective genus. The families, genera (except where there is only one species) and species are all described. It is explained that as there is already a Tree Flora of Malaya, this account, which covers about one quarter of that flora, gives only brief mention of flower and fruit characters. The distribution of each species in the 50-hectare plot is given. A number of typographical errors are noted in passing, but these are minor and should not detract from the main achievement of this book as a major contribution to the use of vegetative characters in the identification of rainforest trees. The only disturbing feature to the reviewer is the use of slash characters (latex, colour and texture of inner bark). Although this is not essential to identification in most cases, slashing at boles has become a standard procedure for foresters and botanists in this part of the world. Over the years most trees in a place like the 50-hectare plot (which presumably would be heavily used for teaching) would run the risk of being slashed repeatedly. The growth and health of such trees could be affected. This is a reasonably priced book that is of great practical use for students of Malesian botany and all who are interested in the identification of trees of the region. Chin See Chung Singapore Botanic Gardens Gardens’ Bulletin Singapore 50 (1998) 123-124. BRYOPHYTES IN THAILAND compiled by Renoo Sornsamran and Obchant Thaitong, Office of Environmental Policy and Planning, Ministry of Science Technology and Environment. 19 colored photographs + One Map + 274 pp (1995). Free upon request from the Office of Environmental Policy and Planning of the Ministry of Science Technology and Environment at 60/1 Soi Pibulwattana Rama VI Road, Bangkok 10400, Thailand. This nicely designed checklist of bryophytes of Thailand with good quality printing came to my attention in 1997 when an announcement of its existence was made public through the Bryonet listserve system. According to the two authors, the checklist was compiled to help researchers in Thailand who have little access to references of Thai bryophytes. Yet, for reasons beyond my comprehension, the authors limit their reference sources to publications dated from 1900 to 1979. Prof. Z. Iwatsuki and I published in 1993 an updated checklist of the mosses of Indochina, Thailand included, in Vol. 74 of the widely circulated Journal of Hattori Botanical Laboratory. Our 1993 checklist and many monographs published in the 80’s (e.g., Iwatsuki and Suzuki 1982; Mohamed and Reese 1985; Reese and Mohamed 1985) were not consulted by the compilers for nomenclatural update of their new checklist. As a result, this is the first checklist of mosses that I know which became outdated on the day it was published. Since I study mainly mosses of East and Southeast Asia, I shall confine my review to the moss part of this new publication. The 1995 listing included 644 species of mosses for the entire country while Tan and Iwatsuki (1993) counted 563 species of mosses in Thailand. The increase in the number of mosses is not due to later discovery but is the result of extensive use of old and discarded synonyms, and even basionyms. Furthermore, the compilers of the checklist under review have not been careful and accurate in their bibliographical work. In a number of instances, the same binomial has been repeated on two different pages, thus, inflating further the total number of moss species. Misspelled names of taxa are not uncommon. The new checklist of Thai bryophytes is done in a tabulated format consisting of four columns entitled Order/Family, Species, Site Found, and Reported by/year. Unfortunately, the arrangement of moss taxa is alphabetical by their ordinal names first, and then the families and genera. Most practising taxonomists, not to mention the students of bryophytes in general, may know well the genera and families of mosses, but will not know by heart the classification of various orders of mosses. This arrangement has made it a difficult reference from which to retrieve information. To aggravate the confusion, the arrangement of liverwort 124 Gard. Bull. Singapore 50(1) (1998) genera in the book is done in a different manner. The hepatic taxa are arranged alphabetically by family and genus after grouping them first under Marchantiales, and followed by Metzgeriales, Calobryales and Jungermanniales. This inconsistency of system of arrangement of taxa leaves the finding of a taxon only by way of the Index of Species that concludes the book. Because of the limitation to references only from 1900 to 1979, the locality information of each species is neither complete nor up-to-date in coverage. However, for local consumption, the compilers are correct to include the names of mountains and other specific locations where the species had been reported in literature. Compared with the latest and electronic catalogue of Thai mosses prepared by the Missouri Botanical Gardens (http://www.mobot.org/MOBOT/moss/Thailand/ thailand.htm), this 1995 checklist, with its enumeration of localities under each province reported for a species, is more useful this way. In the MO’s catalogue of Thai mosses, the locality information stops at the provincial level. One other feature that I like about the new publication is the coloured pictures of a number of Thai bryophytes identified to genus. Unfortunately, the usefulness of this Thai checklist of bryophytes is handicapped by the outdated nomenclature and limited search of literature. References Iwatsuki, Z. and T. Suzuki. 1982. A taxonomic revision of the Japanese species of Fissidens (Musci). Journal of Hattori Botanical Laboratory. 51: 329-508. Mohamed, H. and W. D. Reese. 1985. Syrrhopodon (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist. 88: 223-254. Reese, W. D. and H. Mohamed. 1985. A synopsis of Calymperes (Musci: Calymperaceae) in Malaysia and adjacent regions. Bryologist. 88: 98- 109. Tan, B. C. and Z. Iwatsuki. 1993. A checklist of Indochinese mosses. Journal of Hattori Botanical Laboratory. 74: 325-405. Benito C. Tan School of Biological Sciences National University of Singapore | " MPM: ee Pat ; y ae ae ii Ha bt | ai i } ne) fF A + te y's ? i t by ie a) y a) as (43 i tae ‘ ! ili } bY ie Le L A i, i J fhe ky, Rg de ig ea aM | Ue eee) OA et ae # hl Ay bh ae ; ve hd ks P's | wth | Wits eee m ; ve i. ye ae ; “i ‘ 4 i Pt ALAN Peet ’ 4 t wind!) ‘oe it phy : : y. ; iy ven He eH N a! A ren ran iameson a) 10 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. Numerical data should only be included if itis essential to the argument and this can be presented either in the form of tables or diagrams. 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For literature citations in taxonomic papers the following style is required: Medinilla alternifolia Blume, Mus. Bot. Lugd.-Bat. I:2 (1849) 19. Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524. Illustrations: Drawings should be done in indian ink. Authors should indicate where individual illustrations receive first mention in the text. Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid for, prior to publication. Manuscripts should be sent to: THE EDITOR, GARDENS’ BULLETIN SINGAPORE, SINGAPORE BOTANIC GARDENS, CLUNY ROAD, SINGAPORE 259569 en = = es a aie Sess os he Re Sl” an, ay ae =: —, event th Noman ee So ae Si es a moet - i a Sn meee = “sem lla a = 2 ~ -. Wy Tatham Ide acne, See ee re ° OP OU a “= hema iua’s & cagheetiedy oT 4 tom i a a a ee te nd eg ee ale « = Wel hem very ee = ) ot Sma : a em ~ The Gardens' Bulletin Singapore VOL. 50 (Part 2) December 1998 ISSN 0374-7859 NATIONAL PARKS BOARD > ce RTE va Singapore Botanic Gardens naan Road Singapore 259569 Tel: 4741165 5; Telefax: 4754295 erat THE GARDENS’ BULLETIN The Gardens’ 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. Dr Ruth Kiew (Editor) Singapore Botanic Gardens Dr T.W. Foong (Assist. Editor) Singapore Botanic Gardens Dr S.C. Chin Singapore Botanic Gardens Dr M.J.E. Coode Royal Botanic Gardens Kew, U.K. EDITORIAL BOARD Dr R.T. Corlett University of Hong Kong Hong Kong Dr M.C. Roos Rijksherbarium Leiden, 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 nor 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 contributing authors are found on the inside backcover. Phe a8 a V LDS De Pe) The Gardens' Bulfétin ® Singapore VOL. 50 (Part 2) December 1998 ISSN 0374-7859 CONTENTS Julia, S. and E. Soepadmo New species and new record of ne Blume (Fagaceae) Riad AA AM SATAW AK. WIALAV BNA) 1c ciesk) ct aessdaveenccdsnc.cencecsehsuarsdqceeveressosshecsaderaancesdtesencese 125 Chung, R.C.K. New species of Helicia Lour. and Heliciopsis Sleumer (Proteaceae) SAO RIABES ROMO Mecca a cde ec un eee at Cm ENE rere Uc key 2) eee AI ie Doce Gen zh sunt sa Uekndaeunens ilPop Kiew, R. Niche partitioning in limestone begonias in Sabah, Borneo, ATS BUCO S TE IVEY TSUTOMU S aa dha d) Jase ve silgebee mull ntdea tis Ladmadeadiedebatiedse stil 161 Choo-Toh, G.T., S.L. Chaw, C.E.Z. Chan, D.H.W. Goh and E.Y.H. Lee A survey of termites in the Singapore Botanic Gardens Rain Forest... eee ig Kiew, R. The unique elements of the limestone flora of Batu Tengar Cave aN ADEA ONAN NUM ii oe ey eho ee eta gah nese Negus eanlediadesovanienesdeacebetinbyasonte 185 Kochummen, K.M. INew species and varieties of Moraceae from Malaysia ....1....2.c.ccccesescecscseocsstaceoncesenesones 197 Hay, A. The genus Alocasia (Araceae-Colocasieae) in West Malesia BARN CUR ESI ree ees eM ee ee NL eA Ci Men eee Me Bs had LS la aileaiy ouduesms bine ndaltevemdestve vA BOOK REVIEW mierda Miabesiata series TNE WV OlUme Bio. iis eel ves i iidoculesccssdpocsdaceasecgneeoscdecsvescssernee Benito C. Tan - 335 Date of Publication: 31 December 1998 Published by National Parks Board Singapore Botanic Gardens Cluny Road Singapore 259569 Printed by Oxford Graphic Printers Pte Ltd 4 d f A a] [ i f { wh - i ) ] : AR Ls i) A ive! ne i ‘da a A we ae Carers I 10G: (ir i < 4 a, a tw a, é , i ‘Soir rs f 7 ‘ . ‘ v\4q 4 *! mE 44 .* ‘ ,.¥% Pa ae ' pee u 4 ‘ ' I ' q ' - f 9 bit oe 2 ane a a A Ren a ie adie Ac’ ; ie Saal Sor Ae 7 oY ~ MY) AIS Ry eh . an | 7 Th » * " { AF i i Lidpogetl “Leeta that iar a ‘ oT a: Ba «wt ie © ( , ‘ ' My 02 We ee = ’, | r so ie iv rh Viet « j aoe art # il ‘ aie hee : a) it j 1 7" gi ay 9 aire PY ais, 2 oy 1s : q r ua 7 a Ranks 2 in | 2 ; or ey } fe 5 L RIVE DE, Git: pA 7 sy . / ¢ q ‘ ey bal ae j = Tihs Ny ua ’ qa - f Os 1? Pra a y i. A - web tc Mile gf 7 \ "Cr - ui an j 7 ’ 6 dy - | a. a ' gi Fé 4=f « 4 : 4 j h at ¢ ra Sl aera [ shu ioe by ‘a mY no v ” eds eae dita }* re ‘7 tbe. — i ba? : oy o/h on a) b2 Arey hy Ge » | of och) a ie > < ‘ Lays oi A eb aa ed ‘ < ey ae a?! ry § di ih LS f eich ius fy Weg): ry ey iy ¥ ‘i * ry pe ye | DPipb si 4a) ge rie Gardens’ Bulletin Singapore 50 (1998) 125-150. New Species and New Record of Lithocarpus Blume (Fagaceae) from Sabah and Sarawak, Malaysia S. JULIA and E. SOEPADMO Forest Research Institute Malaysia Kepong, 52100 Kuala Lumpur, Malaysia Abstract Eleven new species and one new record of Lithocarpus are described from Sabah and Sarawak. The new species are L. brochidodromus, L. corneri, L. kalkmanii, L. keningauensis, L. kochummenii, L. melataiensis, L. muluensis, L. oblancifolius, L. sandakanensis, L. stonei, and L. tawaiensis, and the new record is L. hystrix. Descriptions of the new taxa are provided. Introduction In his account of the Malesian Fagaceae, Soepadmo (1972) recognised 49 species of Lithocarpus in Sabah and Sarawak. Upon revision of the genus for the Tree Flora of Sabah and Sarawak, eleven new species and one new record are to be added, making the number of Known species from the two eastern states of Malaysia 61. Of the eleven described new taxa, six occur in Sabah only, two in Sarawak only, two both in Sabah and Sarawak, and one in Sarawak and Brunei. Description of the New Species 1. Lithocarpus brochidodromus S. Julia & Soepadmo, sp. nov. Fig.1 (Latin, brochidodromus=loop-veined; referring to the leaves) Lithocarpo cooperto simillimus, sed foliis multo maioribus crassioribusque, venis lateralibus valide brochidodromis, glande ovoideo-globosa differt. Typus: Dewol & Lideh SAN 105591, Borneo, Sabah, Pensiangan, Sapulut, Sg. Saburan (holotypus SAN!). Tree up to 20 m tall, 15-55 cm in diameter. Bark flaky, greyish or brownish; inner bark reddish or greyish or greenish. Sapwood brownish or whitish. Twig densely tomentose with appressed stellate and simple hairs, later subglabrescent, smooth or sparsely large-lenticellate, sometimes scaly. Stipules linear or ovate, 5-15 x 2-5 mm. Leaves coriaceous, rigid, sparsely appressed yellowish tomentose with simple and stellate hairs or rarely 126 Gard. Bull. Singapore 50(2) (1998) glabrous above, densely appressed yellowish tomentose below; blades broadly elliptic-oblong or ovate, 14—22(-—32) x 5—9(-11) cm, base rounded or broadly acute, margin strongly recurved, apex acuminate, acumen 15-20 mm long; midrib broad, slightly raised above, strongly raised below, densely stellate-tomentose above, sparsely appressed, stellate-tomentose below; lateral veins thick, 7-12 pairs, lax, strongly impressed above, strongly raised below, clearly and strongly joining near the margin, forming an angle of 20°-30° with the midrib; intercostal veins scalariform or subscalariform, lax, prominent and impressed above, clearly prominent below; petioles 4-12 x 2-5 mm. Inflorescences male and female. Male inflorescences solitary in the axils of distal leaves or in subterminal, lax paniculate clusters on the new shoot, 6-10 cm long; bracts linear to acute, 1.5-3.5 x 1-2 mm; bracteoles linear, c. 1.7 x 0.6 mm. Male flowers solitary along the rachis; perianth 6-lobed, thick coriaceous, elliptic, 2.5-3 x 1-1.5 mm; stamens 12, filaments c. 2 mm long; pistillode subglobose, 1.5—2 mm in diameter. Female inflorescences in subterminal, lax paniculate clusters on the new shoot or solitary in the axils of distal leaves, 14-18 cm long; bracts acute, 2—2.5 x 1-1.7 mm; bracteoles linear-elliptic, c. 2 x 0.5 mm. Female flowers solitary along the rachis; perianth 6-lobed, thick coriaceous, acute-rounded, 0.4—0.7 mm across; staminodes 12; styles 3, conical, slightly recurved, 1.5—2.5 mm long. Cupules solitary along the rachis, c. 5-mm-stalked, deeply cup-shaped, 1.5—2.5 x 2—2.5 cm, densely appressed stellate-tomentose, scaly or set with spine-like appendages; wall bony, thick, enclosing the acorn completely or more than half of the acorn; scales distinct, sturdy, hook-like, set irregularly. Acorn ovoid-globose, 1.5-2 cm across, sparsely tomentose with simple hairs, brown, base flat, top rounded-acute; scar convex, c. 1 cm in diameter; wall woody, thin, greater parts free from the cupule. | Vernacular names: Sabah: tikalod (Dusun Ranau); Sarawak: tekalat (Murut). Distribution: Endemic to Borneo. In Sabah, recorded from Sapulut in Keningau, Ulu Tungud in Beluran, and Sg. Timbulanan in Labuk Sugut area. In Sarawak, collected from Sg. Plieran, Belaga and Lambir NP in Miri. Ecology: In primary to secondary forests, including riparian forest. Usually grow on hill slopes, up to 465 m. Notes: A species closely allied to Lithocarpus coopertus but differs by its much bigger and thicker leaves with strongly looped lateral veins, and its ovoid-globose acorn. Lithocarpus from Sabah and Sarawak 127 Figure 1. Lithocarpus brochidodromus S. Julia & Soepadmo, sp. nov. A, flowering (female) leafy twig; B, female flower; C, longitudinal section of female flower; D, part of male inflorescence; E, longitudinal section of male flower; F, infrutescence (A—C from SAN 103647, D-E from S 24089, F from SAN 106929). 128 Gard. Bull. Singapore 50(2) (1998) Specimens Examined: BORNEO. SABAH: Labuk Sugut, Sg. Timbulanan, Aban SAN 90489 (K, KEP!, L, SAN!, SAR!); Beluran, Ulu Tungud, Sg. Dual, Joseph et al. SAN 96531 (K, KEP!, KLU!, L, SAN!, SAR!); Keningau, Sapulut FR, Sg. Tibou, Sumbing & Soludi SAN 103647 (K, KEP!, L, SAN!); Nabawan, Sapulut, Sg. Saburan, RP. 474, Dewol & Lideh SAN 105591 (SAN!); Sapulut, Sg. Saburan, Fidilis & Omar SAN 106929 (SAN!); Keningau, Sapulut, East of Sg. Saburan RP. 474, Leopold Madani SAN 119228 (K, KEP!, SAN!). SARAWAK: Miri, Lambir NP, Chai S 24089 (KEP!, SAN!, SAR!); 7 Division, Belaga, Plieran Rapids, Sg. Pleieran, Murum, Lai et al. §S 67948 (SAR!); F: Division, Belaga, Plieran Rapids, Sg. Pleieran, Murum, Lai et al. S 67949 (SAR!). 2. Lithocarpus corneri S. Julia & Soepadmo, sp. nov. Fig.2 (E. J. H. Corner, 1906-1996, prominent former Professor of Tropical Botany, University of Cambridge, United Kingdom) In folii characteris Lithocarpo ruminato simillimus, sed cupula glandeque obconica, foliis venis intercostalibus densis facile distinguendus. Typus: Berhaman SAN 132620, Borneo, Sabah, Tenom District, Lumaku FR (holotypus: SAN!; isotypi KEP!, L, SAR!). Tree 10-15 m tall, 20-30 cm in diameter. Bark rough, brown. Twig sparsely tomentose, glabrescent, greyish brown, sparsely lenticellate, sometimes smooth. Stipules linear, 3-4 x 1 mm, persistent. Leaves thin coriaceous, sparsely appressed yellowish tomentose above, densely appressed yellowish tomentose below; blades elliptic, (6—)8-12 x 3-4.5 cm, base acute to broadly acute, margin recurved, apex acuminate, acumen 5-10 mm long; midrib raised on both surfaces, stronger above, sparsely appressed tomentose on both surfaces; lateral veins thin, 5-8 pairs, lax, flat above, slightly raised below, disappearing towards the leaf margin, forming an angle of 45°-60° with the midrib; intercostal veins scalariform or subscalariform, dense, obscure on both surfaces or slightly prominent below; petioles 5-10 x 2 mm. Inflorescences and flowers unknown. Cupules solitary along the rachis, sessile, obconical, 2.5—4.5 x 3.5-5.5 cm, base rounded-acute, top flat, densely appressed tomentose, lamellate; wall woody, 2-3 mm thick, completely enclosing the acorn except for an opening of 1-2 cm in diameter; lamellae distinct, entire, set in 12-15 regular lines, denser towards the top. Acorn obconical, 2-4 x 3-5 cm, densely tomentose with simple and stellate hairs, brown; wall woody, thick, greater parts adnate to the cupule. Distribution: Endemic to Sabah, Borneo. So far collected from Lumaku FR, Tenom and Mark Pang logging area, Ranau. Lithocarpus from Sabah and Sarawak 129 Figure 2. Lithocarpus corneri S. Julia & Soepadmo, sp. nov. A, leafy twig; B, stipules; C, mature cupule; D, longitudinal section of the cupule showing the acorn (A—C from SAN 132620, D from SAN 110435). 130 Gard. Bull. Singapore 50(2) (1998) Ecology: In primary to disturbed forests on hillsides. Notes: A species closely related to Lithocarpus ruminatus in leaf characters but can be easily distinguished by its obconical cupule and acorn, densely lamellate cupule, and leaves with dense intercostal veins. Specimens Examined: BORNEO. SABAH: Ranau, Mark Pang Logging area, Amin & Lideh SAN 110435 (SAN!); Tenom, Lumaku FR, Berhaman et al. SAN 132620 (KEP, L, SAN!, SAR!); Tenom, Lumaku FR, Maikin et al. SAN 132937 (K, KEP!, L, SAN!). 3. Lithocarpus kalkmanii S. Julia & Soepadmo, sp. nov. Fig.3 (C. Kalkman, 1928 - 1998, former Director of the Rijksherbarium, Leiden, the Netherlands) In characteris cupulae Lithocarpo halleri simillimus et in folii Lithocarpo pulchro, sed ab Lithocarpo halleri foliis crassis coriaceis rigidis, venis lateralibus densis costa angulis 30°40° abeuntibus, venis intercostalibus dense scalariformibus differt. Ab Lithocarpo pulchro cupula obovoideo-globosa lamellata glandem praeter aperturam c. 0.5 cm in parte apicali omnino tegenti distinguendus. Typus: Meijer & Hendry SAN 42460, Borneo, Sabah, Ranau, Kinabalu NP, W border, N of Sosopodon (holotypus SAN!; isotypi: AA, K-E). Tree up to 30 m tall, 45-60 cm in diameter. Bark large-lenticellate or cracky, brownish or greyish; inner bark reddish brown. Sapwood yellowish. Twig sparsely brownish tomentose, later glabrescent, smooth or sparsely large-lenticellate. Leaves thick coriaceous, sometimes rigid, densely appressed yellowish tomentose above, densely appressed greyish brown tomentose below; blades broadly elliptic, 11-15.5 x 5-8 cm, base broadly acute, margin recurved, apex acuminate, acumen 5-12 mm long; midrib slightly raised above, strongly raised below, sparsely appressed tomentose on both surfaces; lateral veins thick, 12-14 pairs, dense, flat above, strongly raised below, disappearing near the margin, forming an angle of 30°-40° with the midrib; intercostal veins scalariform, dense, obscure above, slightly prominent below; petioles 15-20 x 2-3 mm. Inflorescences and flowers unknown. Cupules solitary along the rachis, sessile, obovoid-globose, 5—7 x 4-5.5 cm, base rounded, top flat, glabrous, lamellate; wall woody, thick, completely enclosing the acorn or with an opening of c. 0.5 cm in diameter; lamellae distinct, entire or minutely denticulate, especially the upper-most ones, set in 6-7 regular lines. Acorn obovoid-globose, 4.5-6 x 3-5 cm, densely tomentose, dark brown; scar deeply concave, c. 1 cm in diameter; wall woody, thick, greater parts adnate to the cupule. Lithocarpus from Sabah and Sarawak 131 Figure 3. Lithocarpus kalkmanii S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, detailed of lower leaf surface; C, longitudinal section of cupule; D, young infructescence (A- B from SAN 42460, C-D from SAN 5671/4). 132 Gard. Bull. Singapore 50(2) (1998) Distribution: Endemic to Sabah, Borneo. So far only found in the Kinabalu NP and Sosopodon FR in Ranau area and one collection from Nabawan. Ecology: In primary upper hill mixed dipterocarp to submontane oak- laurel forests at altitude 1080-1500 m. Notes: A species closely related to Lithocarpus halleri in cupule characters and to Lithocarpus pulcher in leaf characters but differs from Lithocarpus halleri by its thick coriaceous and rigid leaves, its dense lateral veins forming an angle of 30°-40° with the midrib and densely scalariform intercostal veins. From Lithocarpus pulcher it can be distinguished by its obovoid-globose, lamellate cupule, which completely encloses the acorn except for an opening of c. 0.5 cm on the apical part. Specimens Examined: BORNEO. SABAH: Ranau, Kinabalu NP, W. border N. of Sosopodon, Meijer & Henry SAN 42460 (AA, K, L, SAN!); Ranau, Kinabalu, Jalan Liwagu, Francis Sadau SAN 42825 (SAN!); Ranau, Sosopodon FR, Aban Gibot SAN 56714 (KEP!, K, L, SAN!); Nabawan, Ignasius SAN 139147 (SAN!). 4. Lithocarpus keningauensis S. Julia & Soepadmo, sp. nov. Fig.4 (Of Keningau, Sabah) In folti characteris Lithocarpis meijeri luteo sericobalanoque simillimus, ab his cupula magna lignosa glandem omnino tegenti, lamellis etiam in iuventu arcte distinctis facile distinguiendus. Typus: Amin SAN 95311, Borneo, Sabah, Keningau, Bukit Kitau (holotypus SAN!; isotypi K!, KEP!, L, SAR!). Tree 15-20 m tall, 25-60 cm in diameter. Bark fissured or scaly, brown or reddish; inner bark brown or reddish. Sapwood yellowish to brownish. Twig densely appressed tomentose, later glabrescent, slightly fissured or sparsely large-lenticellate or smooth. Leaves thick coriaceous, rigid, glabrous or densely appressed yellowish tomentose above, densely appressed yellowish or brownish tomentose below; blades elliptic or broadly elliptic, 12.5-16.5(-29) x 4.5-7(-12.5) cm, base acute, margin recurved, apex acuminate, acumen c. 10 mm long; midrib broad, slightly raised above, strongly raised below, densely appressed tomentose on both surfaces; lateral veins thick, 12-14 pairs, lax, slightly raised or flat above, raised below, disappearing near the margin, forming an angle of 35°—45° with the midrib; intercostal veins scalariform, dense, obscure above, slightly prominent below; petioles 10-15 x 3-5 mm. Inflorescences and flowers unknown. Cupules solitary along the rachis, sessile, obovoid, 7-11 x 5-6.5 cm, base acute, top flat, glabrescent, lamellate; wall woody, thick, completely Lithocarpus from Sabah and Sarawak 133 ~ Figure 4. Lithocarpus keningauensis S. Julia & Soepadmo, sp. nov. A, leafy twig; B, young infructescence; C, mature cupule; D, longitudinal section of mature cupule (A & D from SAN 50208, B from SAN 92174, C from SAN 95311). 134 Gard. Bull. Singapore 50(2) (1998) enclosing the acorn; lamellae strongly distinct even in the young cupule, folded inwardly, longitudinal groove entire or wavy at the top, set in 6-10 regular lines, protruding from the cupule surface. Acorn obovoid globose, 4.5-6 x 3-5 cm, densely tomentose, dark brown; scar deeply concave, c. 1 cm in diameter; wall woody, thick, greater parts adnate to the cupule. Distribution: Endemic to Sabah, Borneo. Known only from Ulu Biah, Bukit Kitau and Keningau trail in Keningau, Sabah. Ecology: In primary to disturbed forests on hill slopes, up to 300 m, on dark brown soils. Notes: A species closely related to Lithocarpus meijeri, Lithocarpus luteus and Lithocarpus sericobalanus in leaf characters but can be easily distinguished from the three by its big and woody cupule, which is completely enclosing the acorn and by its strongly distinct lamellae even in the young cupule. Specimens Examined: BORNEO. SABAH: Papar, Keningau trail, Francis Sadau SAN 50208 (KEP!, SAN!, SAR!); Keningau, Ulu Biah, Oikawa SAN 92174 (K!, KLU!, L, SAN!, SAR!); Keningau, Bukit Kitau, Amin SAN 95311 (K!, KEP!, L, SAN!, SAR!). 5. Lithocarpus kochummenii S. Julia & Soepadmo, sp. nov. Fig.5 (K. M. Kochummen, senior forest botanist at the Forest Research Institute Malaysia, Kepong) Lithocarpo cooperto simillimus, sed foliis maioribus crassioribus, venis lateralibus distincte brochidodromis, cupula maiore appendiculis validioribus spiniformibus differt. Typus: Bernard Lee S 38884, Borneo, Sarawak, Miri, Gunung Mulu NP (holotypus SAR!; isotypi K!, KEP!, SAN!). Tree 15-30 m tall, 10-60 cm in diameter; stilt root up to 2 m high. Bark fissured or lenticellate, reddish brown. Twig densely tomentose, later glabrescent, sparsely to densely large-lenticellate. Stipules linear, 6-10 x 1 mm. Leaves thick coriaceous, rigid, sparsely appressed tomentose or glabrescent above, densely appressed brownish or yellowish tomentose below; blades broadly or rarely narrowly lanceolate or oblanceolate, (10- )14-18(-22) x (4-)5-7(-8.5) cm, base cordate or rounded, margin strongly recurved, apex sharply acute or acuminate, acumen 15-20 mm long; midrib broad, raised on both surfaces, stronger below, glabrescent or sparsely tomentose on both surfaces; lateral veins thick, (11—)14—18(—20) pairs, dense, strongly impressed or rarely flat above, strongly raised below, clearly joining Lithocarpus from Sabah and Sarawak 135 SONY 3cm PHAN SS (i Sore Si ti A zal Nye Figure 5. Lithocarpus kochummenii S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, part of female inflorescence; C, longitudinal section of female flower; D, longitudinal section of male flower; E, top view of cupule; F, side view of cupule; G, longitudinal section of cupule showing the free acorn (A from S 4557, B—D from DK 964, E-G from S$ 38884). 136 Gard. Bull. Singapore 50(2) (1998) near the margin, forming an angle of 30°—40° with the midrib; intercostal veins thin, scalariform, rarely subscalariform, lax, obscure above, thinly prominent below; petioles 5-10 mm long. /nflorescences male, female or androgynous. Male inflorescences in lateral or subterminal, dense paniculate clusters on the new shoot or solitary in the axils of distal leaves, 5-13 cm long; bracts acute-linear, 1.5-1.7 x 0.8 mm; bracteoles oblong, c. 0.5 x 0.2 mm. Male flowers solitary along the rachis; perianth 6-lobed, thin coriaceous, elliptic, c. 1.2 x 1 mm; stamens 12, filaments 2—2.5 mm long; pistillode c. 1 mm in diameter. Female or androgynous inflorescences solitary in the axils of distal leaves or in terminal, lax paniculate clusters on the new shoots, 7-15 cm long; bracts acute, c. 1.2 x 0.6-1 mm; bracteoles acute, c. 1 x 0.8 mm. Female flowers solitary along the rachis; perianth 6-lobed, thick coriaceous, ovate-acute, c. 1.1 x 0.8 mm; staminodes 10-12; styles 3, conical, recurved, c. 2 mm long. Cupules solitary along the rachis, sessile, conical-ovoid, 2—2.5 cm across, densely tomentose with simple and stellate hairs, scaly or with spine-like appendages; wall woody, thin, enclosing the acorn completely; spine-like appendages distinct, broad and sturdy, straight or slightly recurved, set irregularly. Acorn conical, 1.5—2 cm across, densely tomentose with simple hairs, rarely glabrous, brown, base flat, top acute; scar deeply concave, 1.3-1.5 cm in diameter; wall woody, thin, greater parts free from the cupule. Vernacular name: Sarawak: salad (Murut). Distribution: Endemic to Borneo. Recorded from Gunung Mulu, Ulu Sg. Kayan in Belaga, Batu Lawi in Bario and Ulu Sg. Masia, Kota FR in Lawas, Sarawak. Also known from Bukit Tudal, Temburong, Brunei. Ecology: In submontane, kerangas and riparian forests, at 900-1280 m. Notes: Closely related to Lithocarpus coopertus but differs by its much larger and thicker leaves with clearly looped lateral veins, and larger cupule with sturdier spine-like appendages. Specimens Examined: BORNEO. SARAWAK: Kalabit Highland, foot of Batu Lawi, a tributary of Sg. Tabun, Nooteboom & Chai 2314 (K!, KEP!, SAR!). Baram, Gunung Mulu, path from Sg. Melinau Paku, Anderson S 4557 (K!, SAN!, SAR!); Limbang, Lawas, along Sg. Masia at Maligan Range, Ilias Paie S 32852 (KEP!, SAR!); Limbang, Lawas, Ulu Sg. Masia in Kota FR, Tong & Jugah S 32911 (KEP!, SAR!); Miri, Gunung Mulu NP, Martin S 38182 (K!, KEP!, SAN!, SAR!); Miri, Gunung Mulu NP, Bernard Lee S 38884 (K!, KEP!, SAN!, SAR!); Kapit, Belaga, Dulit Range, Lithocarpus from Sabah and Sarawak Lay Ulu Sg. Kayan, Dayang Awa & Yii S 46836 (K!, KEP!, SAR!); BRUNEI: Temburong subdistrict Amo, Bukit Tudal, Kirkup 964 (BRUN, K!, KEP!). 6. Lithocarpus melataiensis S. Julia & Soepadmo, sp. nov. Fig.6 (Of Bukit Melatai, Sarawak) In folii characteris Lithocarpo pusillo simillimus, petiolo longiore, cupula maiore, glande piloso differt. Typus: Yti S 48455, Borneo, Sarawak, Kapit, Batang Balleh, Bukit Melatai (holotypus SAR; isotypi: BKF, K, KEP!, KELL! L). Tree 12-27 m tall, 25—70 cm in diameter. Twig sparsely tomentose, smooth or fissured. Stipules linear, c. 10 x 1 mm. Leaves thin coriaceous, sparsely appressed brownish tomentose above, densely yellowish brown tomentose, sometimes with simple hairs below; blades narrowly elliptic, 9-13.5 x 2—3 cm, base sharply acute or cuneate, margin recurved, apex caudate or long acuminate, acumen 15-22 mm long; midrib slightly raised on both surfaces, glabrous above, sparsely appressed tomentose with simple hairs below; lateral veins thin, (8—)9-12 pairs, dense, flat above, raised below, faintly joining towards the margin, forming an angle of 30°—40° with the midrib; intercostal veins (sub)scalariform, lax, obscure on both surfaces or slightly prominent below; petioles 6-12 mm long. /nflorescences male and female. Male inflorescences in lateral or subterminal, lax to dense paniculate clusters on the new shoot or solitary in the axils of distal leaves, 5—11 cm long; bracts linear-triangular, 1-1.3 x 0.3 mm; bracteoles acute, 0.4—0.6 x 0.3-0.5 mm. Male flowers solitary along the rachis; perianth 6-lobed, coriaceous, ovate-rounded, 1—1.2 x 0.5—1 mm; stamens 12, filaments 2—2.5 mm long; pistillode globose, c. 1 mm in diameter. Female inflorescences in terminal, lax paniculate clusters on the new shoot, 9.5—12.5 cm long; bracts linear-triangular, c. 1.2 x 0.3-0.5 mm; bracteoles acute-rounded, 0.2-0.3 mm across. Female flowers solitary along the rachis; perianth 6-lobed, thick coriaceous, ovate-acute, 0.4—-0.6 x 0.2-0.3 mm; staminodes 10; styles 3, conical, straight, c. 1 mm long. Cupules solitary along the rachis, sessile, deeply saucer-shaped, 0.4—0.7 x 1.2-1.7 cm, densely tomentose with stellate and simple hairs, lamellate; wall bony, thin, enclosing less than half of the acorn; lamellae distinct, minutely denticulate, set in 6-8 regular lines. Acorn conical, 1.7—2 x 1.1-1.5 cm, densely simple hairy, brown; scar concave, 0).7— 1 cm in diameter, base flat, top sharply acute; wall bony, thin, greater parts free from the cupule. Distribution: Endemic to Sarawak, Borneo. So far only collected from Bukit Melatai and Ulu Balleh in Kapit division. 138 Gard. Bull. Singapore 50(2) (1998) Figure 6. Lithocarpus melataiensis S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, male inflorescence; C, longitudinal section of male flower: D, female inflorescence; E, longitudinal section of female flower (A, D & E from S 48455, B-C from S 48454). Lithocarpus from Sabah and Sarawak 139 Ecology: Mixed dipterocarp forests at altitude 300-880 m. Notes: A species closely related to Lithocarpus pusillus in leaf characters but differs by its bigger cupule, hairy acorn and longer petiole. Specimens Examined: BORNEO. SARAWAK: Kapit, Ulu Balleh, above Sg. Mengiong, Othman Haron S 29203 (AA, BO, K, KLU!, L, MEL, SAN!, SAR!); Kapit, Batang Balleh, Bukit Melatai, Camp. 2, Yii & Jegong S 48440 (BKF, K, KEP!, KLU!, L, SAR!); Kapit, Batang Balleh, Bukit Melatai, Camp. 2, Yii S 48454 (BKF, K, KEP!, KLU!, L, SAN!, SAR!); Kapit, Batang Balleh, Sg. Melatai, Bukit Melatai, above camp 2, Yi S$ 48455 (BKF, K, KEP!, KLU!, L, SAR!), S 48463 (BKF, K, KEP!, KLU!, L, SAR!). 7. Lithocarpus muluensis S. Julia & Soepadmo, sp. nov. Fig.7 (Of Gunung Mulu, Sarawak) Folii textura Lithocarpo rigido simillimus, folii basi late acuto ad rotundato, petiolo plerumque longiore, venis lateralibus angustioribus, glandis apice rotundato differt. Typus: Illias Paie S 15082, Borneo, Sarawak, Gunung Mulu (holotypus SAR!). Tree 12-24 m tall, 15-75 cm in diameter. Bark flaky. Twig glabrescent, fissured or sparsely lenticellate. Leaves thick coriaceous, rigid, glabrous above, sparsely appressed yellowish tomentose below; blades elliptic to broadly elliptic, (8—)11-17 x (4.5—)5.5-8.5(—9) cm, base broadly acute, margin strongly recurved, apex acute to acuminate, acumen (5—)13—23 mm long; midrib raised on both surfaces, stronger above, glabrescent; lateral veins thin, 9-15 pairs, dense or lax, slightly raised on both surfaces, faintly joining towards the margin, forming an angle of 40°—50° with the midrib; intercostal veins (sub)scalariform, lax, obscure on both surfaces; petioles (8—)10—13(- 15) mm long. Inflorescences and flowers unknown. Cupules in clusters of 2 along the rachis when young, later solitary along the rachis, sessile, saucer- shaped, 1.8—2.4 x 2—2.5 cm, sparsely stellate-tomentose, scaly; wall woody, thick, enclosing less than half of the acorn; scales distinct, broad and rigid, set irregularly. Acorn depressed subglobose, 1.8—2.4 x 2-2.5 cm, glabrous and shiny, dark brown, base flat, top rounded; scar deeply concave, 1.5—1.7 cm in diameter; wall woody, thick, greater parts free from the cupule. Distribution: Endemic to Sarawak, Borneo. So far only collected from Gunung Mulu in Miri division. Ecology: In primary hill and submontane forests, 1350-1500 m. 140 Gard. Bull. Singapore 50(2) (1998) Figure 7. Lithocarpus muluensis S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, infructescence showing cup-shaped cupules; C, acorn; D, longitudinal section of acorn (A—B from § 15082, C—D from S 15081). Lithocarpus from Sabah and Sarawak 141 Notes: A species closely related to Lithocarpus rigidus in leaf texture but differs by its broadly acute or rounded leaf base, usually longer petiole, thinner lateral veins, and acorn with rounded apex. Specimens Examined: BORNEO. SARAWAK: Gunung Mulu, path from Sg. Melinau Paku, Anderson S 4506 (AA, K, L, SAN!, SAR!, SING), S$ 4598 (AA, K, L, SAN!, SAR!, SING); Gunung Mulu, //ias Paie S 15077 (SAR!), S 15081 (BO, K, L, SAN!, SAR!, SING), S 75082 (SAR!); Miri, Mulu NP, Gunung Mulu, Camp. 3, Yi & Abu Talib S 58283 (K, KEP!, KLU!, L, SAN!, SAR!). 8. Lithocarpus oblancifolius S. Julia & Soepadmo, sp. nov. Fig.8 (Latin, oblancifolius=with reversed spear-shaped leaves) In folii characteris Lithocarpo lucido simillimus, sed cupula minore tenuiore, glande conico, venarum lateralium numero minoribus, venis intercostalibus minus prominentibus differt. Cupula cupulae Lithocarpi papilliferi similis sed folia differunt. Typus: Leopold Madani SAN 133942, Borneo, Sabah, Tawau, Tawau Hill Park (holotypus SAN!). Tree up to 24 m tall, 20-45 cm in diameter. Bark lenticellate or smooth, brown or greyish; inner bark greyish to yellowish. Sapwood whitish to yellowish. Twig sparsely tomentose, later subglabrescent, smooth or sparsely large-lenticellate. Stipules linear, c. 2 x 1 mm. Leaves thin coriaceous, glabrous above, sparsely appressed greyish tomentose below; blades oblanceolate, (6—)8-13.5 x (2.5—)3—4.5 cm, base cuneate, margin recurved, apex rounded or shortly acute; midrib raised on both surfaces, stronger below, glabrescent; lateral veins thin, 8-13 pairs, lax, flat or impressed above, raised below, joining near the margin, forming an angle of 30°-50° with the midrib; intercostal veins reticulate or subscalariform, dense, obscure above, prominent below; petioles 3-5 x 2 mm. Inflorescences male, female, androgynous or mixed. Male inflorescences in lateral, lax to dense paniculate clusters on the new shoot, 3-11 cm long; bracts linear, c. 1 x 0.3 mm; bracteoles linear, c. 5 x 0.1 mm. Male flowers solitary along the rachis; perianth 6-lobed, thin coriaceous, ovate or elliptic, 0.8-1 x 0.6—-0.8 mm; stamens 10-12, filaments c. 2 mm long; pistillode globose, 0.7—-0.8 mm in diameter. Female or androgynous inflorescences in subterminal, lax paniculate clusters on the new shoot or solitary in the axils of distal leaves, 4-12 cm long; bracts linear-acute, 1—1.2 x 0.2-0.4 mm; bracteoles acute, c. 0.5 x 0.2 mm. Female flowers solitary along the rachis; perianth 6-lobed, coriaceous, broadly acute or rounded, 0.5-0.9 x 0.5—0.7 mm; staminodes 12; styles 3, conical, recurved, 1—-1.2 mm long. Cupules solitary along the rachis, sessile, saucer-shaped, 1—1.2 x 1.6—2 cm, densely appressed tomentose 142 Gard. Bull. Singapore 50(2) (1998) Figure 8. Lithocarpus oblancifolius S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, male inflorescence; C, longitudinal section of male flower; D, female inflorescence; E, longitudinal section of female flower (A from SAN 132942, B-C from SAN 91636, D-E from SAN 111174). Lithocarpus from Sabah and Sarawak 143 with stellate and simple hairs, lamellate; wall woody, thin, enclosing less than half of the acorn; lamellae distinct, entire, set in 6-8 regular lines. Acorn ovoid-conical, 1.6-1.8 x 1.6-2 cm, glabrous, brown, base flat, top acute; scar concave, c. 1.5 cm in diameter; wall bony, thick, greater parts free from the cupule. Distribution: Endemic to Sabah, Borneo. Collected from Tawau Hill Park in Tawau and Sg. Piso in Labuk Sugut area. Ecology: Near the riverbank, rather open area at altitude c. 90 m. Notes: A species closely allied to Lithocarpus lucidus in leaf characters but differs by its smaller and thinner cupule, conical acorn, less number of lateral veins and by its less prominent intercostal veins. The cupule is similar to that of Lithocarpus papilifer but the leaves are different. Specimens Examined: BORNEO. SABAH: Labuk Sugut, Sg. Piso, Aban Gibot & Dewol SAN 91636 (K, SAN!); Tawau, Tawau Hill Park, Water Pump, Leopold Madani & Sigin SAN 111174 (K, L, SAN!, SAR!); Tawau, Tawau Hill Park, Leopold Madani et al. SAN 133942 (SAN!). 9. Lithocarpus sandakanensis S. Julia & Soepadmo, sp. nov. Fig.9 (Of Sandakan, Sabah) Lithocarpo stonei simillimus, foltis tenuioribus, cupula parum minore, glande piloso distinguendus. Typus: Wood SAN A 4697, Borneo, Sabah, Sandakan, Sepilok FR, Compartment 17 (holotypus KEP!; isotypi AA, L, MEL, SING). Tree 10-45 m tall, 15-90 cm in diameter; buttresses small. Bark smooth, greyish or brownish; inner bark yellowish. Sapwood whitish or purplish. Twig glabrescent, sparsely lenticellate. Leaves coriaceous, sparsely appressed yellowish tomentose or glabrescent above, densely appressed brownish tomentose below; blades broadly elliptic-oblong, 26-39(—46) x 7.5—11.5 cm, base acute-rounded, margin recurved, apex sharply acute to acuminate, acumen c. 15 mm long; midrib flat or slightly raised above, strongly raised below, glabrous on both surfaces; lateral veins thick, 11-14 pairs, dense, flat or impressed above, strongly raised below, disappearing towards the margin, forming an angle of 30°-45° with the midrib; intercostal veins subscalariform or reticulate, lax, thinly prominent on both surfaces, stronger below; petioles 8-18 x 3-6 mm. Male inflorescences c. 10 cm long; bracts linear-acute, 1—1.1 x 0.4 mm; bracteoles linear-acute, c. 0.6 x 0.2 mm. Male flowers solitary along the rachis; perianth 6-lobed, coriaceous, elliptic, 1.1- 144 Gard. Bull. Singapore 50(2) (1998) wie. 2B Figure 9. Lithocarpus sandakanensis S. Julia & Soepadmo, sp. nov. A, leafy twig; B, part of infructescence; C, longitudinal section of cupule and acorn (All from SAN A 4697). Lithocarpus from Sabah and Sarawak 145 1.3 x 0.6—0.8 mm; stamens 10, filaments c. 2 mm long; pistillode subglobose, 0.6-1 mm in diameter. Female inflorescences and flowers unknown. Cupules solitary along the rachis, 0.5—0.7-cm-stalked, cup-shaped, 1.5—2 x 3-3.5 cm, densely tomentose, lamellate; wall woody, thick, enclosing up to half of the acorn; lamellae distinct, minutely denticulate or wavy, set in 7-8 regular lines. Acorn depressed ovoid, 3.5—4 cm across, densely tomentose with simple hairs, dark brown, base flat, top acute; scar deeply concave, c. 2 cm in diameter; wall woody, thick, greater parts free from the cupule. Distribution: Endemic to Sabah, Borneo. All known specimens were collected from Sepilok FR, Sandakan. Ecology: Primary lowland mixed dipterocarp forest, on yellow soil, up to 15 m altitude. Notes: A species closely related to Lithocarpus stonei but can be distinguished by its much thinner leaves, slightly smaller cupule and by its hairy acorn. Specimens Examined: BORNEO. SABAH: Sandakan, 15 miles W of Sandakan, Sepilok FR, Compartment 17, Wood SAN A 4697 (AA, KEP!, L, MEL, SING); Sandakan, Sepilok FR, Jalan Batu 15, Patrick SAN 19698 (KEP!, SAN!); Sandakan, Sepilok FR, Batu 15, Meijer SAN 34286 (K, L, SAN!); Sandakan, Sepilok FR, Jalan Kantor Pos, Patrick SAN 62061 (SAN!); Sandakan, Arboretum, Mile 14, Aban Gibot, Kumin & Rakim SAN 73710 (SAN!). 10. Lithocarpus stonei S. Julia & Soepadmo, sp. nov. Fig.10 (B. C. Stone, 1933 — 1994, former Reader in Botany, Department of Botany, University of Malaya, Kuala Lumpur, Malaysia) Species cupula magna lignosa cupuliformi, glande magno glabro apice nitido rotundato, foliis crassis rigidis notata. Cupulae Lithocarpi revoluti satis similis sed folti characteri differunt. Typus: Yap & Khairuddin SAN 106051, Borneo, Sabah, Tambunan road (holotypus SAN!; isotypus KEP!). Tree 10-25 m tall, 60-100 cm in diameter. Bark fissured or smooth or lenticellate, dark grey; inner bark fibrous, yellowish brown or whitish. Sapwood whitish. Twig densely tomentose, sparsely to densely large- lenticellate. Leaves thick coriaceous, rigid, densely appressed yellowish tomentose above, sparsely yellowish tomentose below; blades broadly elliptic-oblong, 28—36(-40) x 9-14 cm, base rounded to broadly acute, margin recurved, apex acuminate, acumen c. 15 mm long; midrib strongly raised 146 Gard. Bull. Singapore 50(2) (1998) on both surfaces, stronger below, sparsely appressed tomentose on both surfaces, denser above; lateral veins thick, (9—)10-12 pairs, lax, flat or impressed above, strongly raised below, disappearing towards the margin, forming an angle of 30°-45° with the midrib; intercostal veins reticulate, lax, obscure above, prominent below; petioles 10-15 x 5-7 mm, kneed. Male inflorescences and flowers unknown. Female inflorescences solitary in the axils of distal leaves, much branched and sturdy, c. 20 cm long; bracts linear-acute, 2—2.5 x 0.2-0.5 mm; bracteoles broadly acute, c. 0.5 x 0.7 mm. Female flowers in clusters of 2—3 along the rachis; perianth 6- lobed, coriaceous, acute, c. 1 x 0.5 mm; staminodes 12; styles 3, conical, straight, c. 1.5 mm long. Cupules in clusters of 2-3 or rarely solitary along the rachis, sessile, deeply cup-shaped, 2—3 x 4-5 cm, densely tomentose, lamellate; wall woody, 3—5 mm thick, enclosing half or more than half of the acorn; lamellae strongly distinct, folded inward, wavy, set in 7-9 regular or irregular lines. Acorn depressed ovoid globose, 3—4 x 3—5 cm, glabrous and shiny, brownish, base flat, top rounded; scar deeply concave, 2—2.5 cm in diameter; wall woody, thick, greater parts free from the cupule. Vernacular name: Sarawak: saled urong (Kelabit). Distribution: Endemic to Borneo. In Sabah, collected from Crocker Range in Tenom, Gunung Alab and Tambunan road in Tambunan. In Sarawak, collected only from Sg. Marariro in Bario area. Ecology: In lower montane forest, on recent alluvium, clay soils, at 960- 1050 m. Notes: A species characterised by its big, woody and cup-shaped cupule, big, glabrous acorn with shiny and rounded apex, and by its thick and rigid leaves. The cupule is rather similar to that of Lithocarpus revolutus but the leaf characters are different. Specimens Examined: BORNEO. SABAH: Tambunan District, Crocker Range, Kota Kinabalu to Sunsuron Road, Andrews 851 (K!); Tambunan, Crocker Range, along Tambunan-Penampang road, Sugau et al. JBS 66 (AA, AAU, EDH, HAST, K, KEP!, L, PNH, SAN!, SAR!, SING); Kota Kinabalu, Tambunan Road, Yap & Khairuddin SAN 106051 (KEP!, SAN!); Tambunan, Gunung Alab, km 56, Jalan Kota Kinabalu-Tambunan, Fidilis & Sumbing SAN 121701 (K!, KEP!, L, SAN!); Tenom, Crocker Range, Meijer SAN 136522 (CHI, KY, SAN!). SARAWAK: Bario, Ulu Baram, path to Pa’Main, near Sg. Marariro, Anderson S 20067 (SAN!, SAR!). Lithocarpus from Sabah and Sarawak 147 Figure 10. Lithocarpus stonei S. Julia & Soepadmo, sp. nov. A, fruiting leafy twig; B, longitudinal section of cupule; C, female inflorescence; D, female flower; E, longitudinal section of female flower; F, part of female inflorescence (A-B from SAN 106051, C-F from SAN 136522). 148 Gard. Bull. Singapore 50(2) (1998) 11. Lithocarpus tawaiensis S. Julia & Soepadmo, sp. nov. Fig.11 (Of Bukit Tawai, Sabah) Ab generis speciebus omnibus foltis sessilibus crasse coriaceis rigidis, basi cordato, venis lateralibus fere invisibilis, inflorescentiis masculis validis differt. Typus: Berhaman et al. SAN 134267, Borneo, Sabah, Bukit Tawai (holotypus KEP!). Stunted treelet of about 1-1.5 m tall and c. 5 cm in diameter. Bark pale brown. Twig sparsely appressed tomentose, smooth. Stipules ovate-rounded, 10-13 x 10-18 mm. Leaves sessile, thick coriaceous, rigid, glabrous above, sparsely appressed yellowish tomentose or glabrescent below; blades ovate- rounded, 6.5—10.5 x 6-9 cm, base cordate, margin recurved, broadly acute; midrib raised on both surfaces, slightly stronger below; lateral veins thin, 10-12 pairs, dense, flat and almost invisible on both surfaces, faintly joining near the margin, forming an angle of 20-30° with the midrib; intercostal veins reticulate, dense, obscure above, slightly prominent below. Male inflorescences in lateral or terminal, lax paniculate clusters on new shoots or solitary in the axils of distal leaves, 6-15 cm long; bracts and bracteoles ovate-acute, 0.6—0.8 x 0.4-0.6 mm. Male flowers solitary or in clusters of 2-3 along the rachis; perianth 6-lobed, thick coriaceous, elliptic, c. 1.2 x 0.7 mm; stamens 12, filaments c. 2 mm long; pistillode globose, c. 0.5 mm in diameter. Female inflorescences and flowers unknown. Young cupule solitary along the rachis, sessile, saucer-shaped, 1-1.3 x 0.3-0.4 cm, scaly, enclosing less than half of the acorn; scales distinct, set irregularly. Acorn for the greater parts free from the cupule. Distribution: Endemic to Borneo. Twice collected from Bukit Tawai, Telupid, Sabah. Ecology: Primary hill forest on ultrabasic soil. Notes: A species differs from any other known species of the genus by its sessile, thick coriaceous and rigid leaves with cordate base and almost invisible lateral veins, and its sturdy male inflorescences. Specimens Examined: BORNEO. SABAH: Telupid, summit of Bukit Tawai, Berhaman et al. SAN 134267 (KEP!); Telupid, Bukit Tawai, Sugau et al. SAN 138832 (KEP!, SAN!). Lithocarpus from Sabah and Sarawak 149 Figure 11. Lithocarpus tawaiensis S. Julia & Soepadmo, sp. nov. A and B, flowering (male) leafy branches; C, distal twig with axillary, branched male inflorescence; D, distal part of male inflorescences with flower buds; E, cluster of 3 male flowers; F, proximal twig with young infructescence; G, cluster of 3 developing cupules and acorns; H, very young developing cupule; I, young infructescence; J, mature cupules; K, detailed venation on the leaf undersurface (A- H from SAN 134267; I-K from SAN 138832). 150 Gard. Bull. Singapore 50(2) (1998) New record Lithocarpus hystrix (Korth.) Rehder The species is previously known from Sumatra, Peninsular Malaysia and Kalimantan in Borneo (Soepadmo 1972). Recent collections extend the distribution to Sarawak and Brunei. Specimens Examined: SARAWAK: Bintulu division, Bukit Lumut, Abg. Mohtar & Yii S 65896 (BKF, K, KEP!, L, SAN!, SAR!); 7 Division, Belaga, Dulit Range, Ulu Sg. Kayan, Dayang Awa & Yii S 46731 (BKF, K, KEP!, KLU!, L, SAR!); Lundu, Pasir Tengah/Biawak, Othman Ismawi et al. S 63890 (BKF, K, KEP!, KLU!, L, SAR!); 1° division, Lundu, Sampadi FR, Syarikat Woodworking Salmas, Othman Ismawi S 37824 (K, KEP!, KLU!, L, SAR!); Kapit District, Bukit Raya, Soepadmo & Chai S 28198 (AA, BO, K, KLU!, L, SAN!, SAR!, SING); Simanggang, Ulu Skrang, path to Bukit Sadok, J/lias Paie, Banyeng & Manggi S 44907 (BKF, K, KLU!, L, SAR!); 4" division, Gunung Mulu NP, Martin S 38902 (K, KEP!, KLU!, L, SAN!, SAR!); 1st and 2™4 division boundary, Ulu Simunjan, Gunung Angkong, Martin S 36953 (K, KEP!, KLU!, L, MO, SAR!); 1* division, Padawan, Bukit Woen, Yii S 61453 (BKF, K, KEP!, KLU!, L, SAR!). BRUNEI: Temburong River Valley, Johns 7325 (BRUN, KEP!). Acknowledgements We wish to extend our gratitude for the support and encouragement extended by the Director-General of FRIM, Kepong and to the Curators of K, L, SAN and SAR herbaria for the loan of specimens. We are also deeply indebted to Dr. J.F. Veldkamp of the Rijksherbarium, Leiden, the Netherlands for providing the Latin diagnoses of the new species, and to Ms. Rosemary Wise (OFI) and Mr. Joseph Pao (SAR) for preparing the illustrations. The research was part of the Tree Flora of Sabah and Sarawak Project, funded by the International Tropical Timber Organization (ITTO), the Malaysian Government, and the Overseas Development Administration of the United Kingdom (ODA-UK). References Soepadmo, E. 1972. Fagaceae. Flora Malesiana 7, 2: 318-385. Gardens’ Bulletin Singapore 50 (1998) 151-160. New Species of Helicia Lour. and Heliciopsis Sleumer (Proteaceae) from Borneo R.C.K. CHUNG Forest Research Institute Malaysia Kepong, 52109 Kuala Lumpur, Malaysia Abstract Two new species of Helicia Lour. (H. sessilifolia and H. symplocoides) and two new species of Heliciopsis Sleumer (H. percoriacea and H. litseifolia) are described and illustrated from Borneo. Introduction In his accounts of Malesian Proteaceae, Sleumer (1955a, 1955b) recognised eight species of Helicia and two species of Heliciopsis from Borneo. The revision of the genera Helicia Lour. and Heliciopsis Sleumer (Proteaceae) for the Tree Flora of Sabah and Sarawak revealed, four new species. Helicia symplocoides and Heliciopsis percoriacea are endemic to Sabah and Sarawak respectively, while Helicia sessilifolia is known from Sabah and Sarawak only. Heliciopsis litseifolia is common throughout Borneo (except Brunei), Peninsular Malaysia and Sumatra. These species are described and illustrated below. Helicia 1. Helicia sessilifolia R.C.K. Chung, sp. nov. Fig. 1 (Latin, sessilis=stalkless, folium=leaf) Helicia sessilifolia Heliciae maxwellianae similis, sed in ramulis gracilibus, folii margine erecurvata, foliis anguste coriaceis flavido brunnescentibus in sicco, fructu stipitato ellipsoideo-apiculato castanescenti in sicco differt. Typus: Borneo, Sarawak, Limbang Division, Lawas, Ulu Trusan, Bt. Tebunan, 9 May 1986, Bernard Lee S 52436 (holotypus KEP!; isotypi K, L, MO, SAN!, SAR!). Treelet to small tree, up to 10 m tall. Twigs: youngest parts subangular, older ones terete, light brown, glabrous. Leaves spiral or subopposite; blades broadly oblong to elliptic, rarely obovate, (6—)9-17 x (4.5—)5—8.5 cm, thinly 152 Gard. Bull. Singapore 50(2) (1998) coriaceous, yellowish brown when dry, not shining, glabrous; base rounded to subcordate, margin entire, apex acute; midrib slightly raised above, prominent below; lateral veins 6-7 pairs, curving and joining near margin, prominent on both surfaces; intercostal veins reticulate, inconspicuous on both surfaces; petioles extremely short, up to 2 mm long, slightly swollen at base, dull brown, glabrous. /nflorescences racemose, axillary, solitary, c. 7 cm long, laxly flowered near the base; rhachis terete, c. 1 mm diameter, glabrous; bracts minute, less than 0.5 mm long, glabrous. Flowers: pedicels 5—6 mm long, in pairs, not winged, connate up to about 2-3 mm from the base, glabrous; perianth (12—)16-19 mm long, glabrous, limb ellipsoid, 0.8- 1.2 mm diameter; anthers 1-1.5 mm long; ovary ovoid, glabrous; style filiform, apex clavate, glabrous; stigma punctiform, terminal, stigmatic surface glandular; disk glands almost entirely connate in a crenulate ring. Fruits ellipsoid, 44.5 x 2.4-2.7 cm, slightly oblique, glabrous, chesnut- brown when dry, apiculum 1—4 mm long, contracted into a stipe of c. 3 mm long; pericarp smooth, 2.5—3 mm thick; fruit stalk unknown. Distribution: Endemic to Borneo. Rare in Sarawak and Sabah, known in Sarawak only from Bt. Tebunan, Lawas (S 52434 and S$ 52436) and in Sabah from Tambunan (SAN 60837 and SAN 111305). Not yet recorded from Brunei and Kalimantan. Ecology: Mixed dipterocarp forest, up to 900 m. Notes: This species is similiar to H. maxwelliana, from which it is distinguished by its slender twigs (stout in H. maxwelliana), non-recurved leaf-margin (curled inwards in H. maxwelliana), thinly coriaceous leaves (thickly coriaceous in H. maxwelliana) which turn yellowish brown when dry (olivaceous-yellowish to dark brown in H. maxwelliana), long-apiculate and stiped ellipsoid fruit, which turns chesnut-brown when dry (subglobose fruit, without apiculum and stipe, and black when dry in H. maxwelliana). The new species 1s apparently confined to hill mixed dipterocarp forest. In contrast, H. maxwelliana is restricted to submontane forest. Specimens Examined: BORNEO. SARAWAK: Limbang Division—Lawas, Ulu Trusan, Bt. Trusan, 9 May 1986, Bernard Lee S 52434 (K, KEP!, L, MO, SAN!, SAR!), S 52436 (K, KEP!, L, MO, SAN!, SAR!). SABAH: Pedalaman District—Tambunan, 21 July 1984, Amin & Suali SAN 60837 (AA, K, KEP!, L, SAN!, SAR!, SING!), Rafflesia FR, 7 Sept. 1985, Leopold Madani & Ismail SAN 111305 (K, KEP!, SAN!). 153 Helicia and Heliciopsis from Borneo Figure 1. Helicia sessilifolia. A, leafy twig; B, flower buds; C, base of ovary with disk glands; D, fruit; E, fruit in longitudinal section; F, seed. (A—C from S 52436, D-F from S 52434.) 154 Gard. Bull. Singapore 50(2) (1998) 2. Helicia symplocoides R.C.K. Chung, sp. nov. Fig. 2 (Greek, -oides=resembling; with leaves resembling those of Symplocos) Hac species nova a generis speciebus aliis foliis crasse coriaceis c. 10 cm longis 5 cm latis, apice emarginato vel obtuso, basi decurrenti cuneata, marginibus recurvatis, fructibus minutis ad 1.7 cm longis 1.4 cm latis differt. Typus: Borneo, Sabah, Pantai Barat District, Mt. Kinabalu, Mesilau Cave, 1 April 1964, Chew & Corner RSNB 4786 (holotypus SAN!; isotypi K, L). Tree 15 m tall, 25 cm diameter. Twigs terete, grey or greyish brown, glabrous with distinct leaf scars up to 3 mm diameter. Leaves spiral; blades obovate, 5—10 x 2.5—5 cm, thickly coriaceous, deep green above, brown below, not shining, glabrous; base cuneate, decurrent, margin entire or occasionally with 1-3 minute teeth in the upper half, recurved, apex obtuse or emarginate; midrib raised above, prominent below; lateral veins 6-8 pairs, curving near the margin and joining with next one to form looped intramarginal veins, visible below, inconspicuous above; intercostal veins inconspicuous on both surfaces; petioles 2-4 x 1.5—2 mm, swollen and wrinkled at the base, dark brown when dry, glabrous. Flowers not known. Fruits ellipsoid to broadly ellipsoid, 1.5—1.7 x 1.2-1.4 cm, oblique, glabrous, black when dry, shortly apiculate, apiculum c. 1 mm long, stipe c. 2 mm long; pericarp smooth, 0.8—1.5 mm thick; fruit stalk 5-7 x 1.5—2 mm. Distribution: Recorded only from Sabah where it is known from a single collection, Chew & Corner RSNB 4786, from Mt. Kinabalu, Mesilau Cave, on ultramafic soil. Ecology: Submontane forest at 1850 m. Notes: The leaves of the new species resemble those of Symplocos Jacquin (Symplocaceae). : Specimens Examined: BORNEO. SABAH: Pantai Barat District—Mt. Kinabalu, Mesilau Cave, 1 April 1964, Chew & Corner RSNB 4786 (K, L, SAN!). Heliciopsis 1. Heliciopsis litseifolia R.C.K. Chung, sp. nov. Fig. 3 (With leaves resembling those of Litsea, Lauraceae) Heliciopsis litseifolia Heliciopsidi montanae proxime affinis, a posteriore foliis simplicibus anguste coriaceis basi attenuata decurrenti, apice acuto vel Helicia and Heliciopsis from Borneo 155 Figure 2. Helicia symplocoides. A, leafy twig; B, fruiting leafy twig; C, fruit; D, fruit in longitudinal section. (From RSNB 4786. ) 156 Gard. Bull. Singapore 50(2) (1998) acuminato, marginibus integris, petiolis brevibus ad 2 cm longis distinguendam. Typus: Borneo, Sarawak, Kapit Division, Belaga, 2 Sept. 1958, Jacobs 5401 (holotypus SAR!; isotypi B, CANB, G, K, L, S, US). Small to medium-sized tree, 6-25 m tall, 10—25(-50) cm diameter. Twigs: youngest parts angular, older ones terete, grey-brown, glabrous. Mature leaves elliptic to broadly elliptic, 10-25 x 4-11.5 cm, thinly coriaceous, yellowish green to olivaceous brown when dry, not shining, glabrous; base attenuate, decurrent, margin entire, apex acute or acuminate; midrib slightly raised above, prominent below; lateral veins 5-6 pairs, curved upwards and joining near the margin to form loops, prominent on both surfaces; intercostal veins reticulate, faint above, typically visible below; petiole (0.5—)1—2.5 cm long, swollen at the base, black and rarely yellowish brown when dry, glabrous. Inflorescences racemose, axillary or born on older, leafless branches, solitary, 12-26 cm long, laxly flowered except for about 3 cm from the base; rhachis 1-1.5(—2) mm diameter, rufous pubescent, soon glabrescent; bracts subulate, 1-2 mm long, persistent, rufous pubescent. Flowers: pedicels 5-8 mm long, mostly in pairs, connate up to 3-5 mm from the base, rufous pubescent; perianth 8-10 mm long, rufous pubescent to glabrescent, limb clavate, c. 1.5 mm diameter; anthers 1—-1.5 mm long; ovary glabrous; style filiform, clavate towards the apex, glabrous; stigma discoid, lateral, stigmatic surface glandular, with distinct cleft; disk glands truncate, free, spaced. Fruits cylindric ellipsoid, (2.7—)3-3.5(-3.8) x (1.7—)2-2.2(-—2.5) cm, smooth, shining black when dry; exocarp leathery, c. 1 mm thick; mesocarp built up by radial, soft brown fibres c. 2.5 mm long; endocarp woody, thin; fruit stalk 10-12 x 3-4 mm. Distribution: Sumatra, Peninsular Malaysia and Borneo. Ecology: Lowland and hill mixed dipterocarp forest, up to 900 m. Notes: In Borneo, the leaf and petiole characters are rather variable. In Othman Haron S§ 29994, Sumbing Jimpin SAN 110338, and Church 173, the leaves range from 16-25 cm long and 9-11.5 cm wide, and the petioles from 2-2.5 mm in diameter. Furthermore the fruits in de Wilde & de Wilde-Duyfjes 16611 from Sumatra, are larger (c. 4.5 x 3.5 cm) than those of the Bornean specimens and the endocarp is thicker (c. 3 mm). Specimens Examined: SUMATRA. Atjeh, G. Leuser Nature Reserve, G. Mamas, 5 May 1975, de Wilde & de Wilde-Duyfjes 16611 (BO, KEP!, L). PENINSULAR MALAYSIA. TERENGGANU—Ulu Setiu FR, 4 Aug. 1977, Chan FRI 23980 (K, KEP!, L, SING!). JOHORE—Ulu Sg. Anak Endau, 3 April 1968, Cockburn FRI 8118 (K, KEP!, SING!). BORNEO. Helicia and Heliciopsis from Borneo 157 Figure 3. Heliciopsis litseifolia. A, leafy twig; B, male inflorescence; C, male flower buds; D, open female flower; E, base of ovary with disk glands; F, stigma; G, infructescence; H, fruit in longitudinal section. (A, G-H from SAN 67659, B-C from S$ 34497, D-F from Jacobs 5401.) 158 Gard. Bull. Singapore 50(2) (1998) SARAWAK: Kuching Division—Sematan, G. Pueh, 23 June 1974, James et al. S 34497 (A, K, L, MO, SAN!, SAR!). Samarahan Division—Serian, Sabal FR, 14 May 1974, Tong S 34320 (K, KEP!, L, MO, SAN!). Sibu Division—Anap, Ulu Muput Kanan, Bt. Kemantan, 12 Oct. 1963, Chai S$ 19547 (A, BO, K, L, MEL, SAN!, SAR!, SING!), Ulu Kakus, 9 March 1970, Othman Haron S 29994 (A, K, KEP!, L). Kapit Division—Belaga, Rajang R., 2 Sept. 1958, Jacobs 5401 (B, CANB, G, K, L, SAR!, SING!, US), Batu Laga, 4 Sept. 1984, Abg. Mohtar S 48191 (K, KEP!, L, MO, SAN!, SAR!). SABAH: Pedalaman District—Keningau, Pensiangan FR, 16 Oct. 1985, Sumbing Jimpin SAN 110338 (SAN!). Tawau District—Tawau, Sg. Pang Burong FR, 17 July 1969, Leopold Madani & Saikeh SAN 67659 (K, L, SAN!, SAR!), Kinabutan Kecil, 20 May 1963, Aban Gibot SAN 35872 (SAN!, SAR!). KALIMANTAN: Kalimanatan Barat—Sintang, Bt. Baka NP, 17 Oct. 1993, Church 173 (A, BO, KEP!). Kalimantan Tengah— Ulu Barito, 22 June 1990, Ridsdale PBU 595 (BO, KEP!, L). 2. Heliciopsis percoriacea R.C.K. Chung, sp. nov. Fig. 4 (Latin, per=exceedingly, coriaceus=leathery, referring to leaves) Heliciopsidi montanae similis, foliis late ellipticis, petiolis glabrescentibus, pedicellis 8-10 mm longis, periantho 12-15 mm longo limbo c. 2.5 mm diam. distinguendam. A Heliciopside litseifolia in foltis crasse coriaceis, apice obtuso, petiolis perianthiis longioribus differt. Typus: Borneo, Sarawak, Kuching Division, Lundu, G. Pueh, 4 Oct. 1985, Othman Ismawi et al. S 49967 (holotypus KEP (Sheet 1)!; isotypi K, KEP (Sheet 2)!, L, MO, SAN!, SAR!) Medium-sized tree, 21 m tall, 36 cm diameter. Twigs terete, greyish brown, rufous tomentose when young, soon glabrous. Mature leaves broadly elliptic, (10—)12-18(-21) x (7—-)8-11(-12.5) cm, thickly coriaceous, yellowish olivaceous or yellowish brown when dry, shining above, glabrous; base acute, margin entire, recurved, apex obtuse; midrib slightly raised above, distinctly prominent below, rufous tomentose, becoming glabrescent; lateral veins 7-8 pairs, curving and joining near the margin, slightly raised above, distinctly prominent below; intercostal veins reticulate, prominent on both surfaces; petioles 3.5-4.5 x 2.5-3 cm, rufous tomentose when young, glabrescent. Inflorescences racemose, solitary on older, leafless branches, 26-28 cm long, laxly flowered except for 1-2 cm from the base; rhachis terete, c. 2.5 mm diameter, rufous tomentose; bracts subulate, c. 1 mm long, persistent, rufous tomentose. Flowers (male): pedicels 8-10 mm long, in pairs, connate up to 4-6 mm from the base, rufous tomentose; perianth 12-15 mm long, rufous tomentose, limb ellipsoid, c. 2.5 mm diameter; anthers c. 2 mm long; disk glands ovate, free, slightly distant from each Helicia and Heliciopsis from Borneo 159 x 3 ni Ey \\ i YW es NY € Not oi RA 7 ‘4 Ry ‘ ye WA A Vy A =) ts SS A-B ¥ TSK 3cm Cp Koseuany WASL. Figure 4. Heliciopsis percoriacea. A, leafy twig; B, male inflorescence; C, longitudinal section of male flower. (From S 49967 (Sheet 1).) 160 Gard. Bull. Singapore 50(2) (1998) other. Flowers (female) and fruits not known. Distribution: Endemic to Sarawak, it is very rare, once collected from G. Pueh. No record from Sabah, Brunei and Kalimantan. Ecology: In Heath forest. Specimens Examined: BORNEO. SARAWAK: Kuching Division—Lundu, G. Pueh, 4 Oct. 1985, Othman Ismawi et al. S 49967 (K, KEP (2 Sheets)!, L, MO, SAN!, SAR!). Acknowledgements I wish to express my gratitude for the support and encouragement extended by the Director-General of FRIM, Kepong, the Directors of the Forestry Departments of Sabah and Sarawak. I am grateful to the Curators and Keepers of the BM, BO, BRUN, K, KEP, KLU, L, SAN, SAR and SING herbaria for loan of specimens and for permission to consult specimens at their institutions. I thank Dr J.F. Veldkamp of the Rijkksherbarium, Leiden for providing Latin diagnoses of the new species. Dr Martin Cheek of the Royal Botanic Gardens, Kew, Mr Steve Cafferty of the Natural History Museum, London and Ms Stans Kofman of the Rijsherbarium, Leiden, for providing photographs of the type specimens. Dr L.G Saw of Forest Research Institute Malaysia, Kepong, for his assistance in searching the needed specimens during his tenure of Liaison Officer for the Tree Flora of Sabah and Sarawak Project. Grateful acknowledgement is also due to Dr E. Soepadmo and Mr K.M. Kochummen of Forest Research Institute Malaysia, Kepong, for their guidance and comments in the preparation of this paper. Mr Zainal bin Mustafa, University of Malaya, Kuala Lumpur, and Ms Rosemary Wise, Oxford Forestry Institute, U.K. kindly prepared the illustrations for this account. The present investigation is part of a revision of the Proteaceae for the Tree Flora of Sabah and Sarawak Project, funded by the U.K. Overseas Development Administration (ODA), the International Tropical Timber Organization (ITTO) and the Malaysian Government. References Sleumer, H. 1955a. Studies in Old World Proteaceae. Blumea. 8(1): 1-95. Sleumer, H. 1955b. Proteaceae. Flora Malesiana. 1, 5(2): 147-206. Gardens’ Bulletin Singapore 50 (1998) 161-169. Niche Partitioning in Limestone Begonias in Sabah, Borneo, Including Two New Species RUTH KIEW Singapore Botanic Gardens Cluny Road, Singapore 259569 Abstract The begonia flora of limestone hills in Sabah is extremely biodiverse with several begonias being found at a single locality, the great majority of which have extremely local distributions. For example, on Bukit Dulong Lambu (better known as Gomantong Cave) four species co- exist. Two are new species, Begonia gomantongensis and B. postarii for which descriptions are provided. The former is endemic to Bk. Dulong Lambu, as is B. malachosticta Sands. The fourth species, B. gueritziana Gibbs is widespread on limestone, as well as on other rock types. Field observations show that niche partitioning occurs between these four species based on light conditions (that also relate to severity of water stress) and substrate. All four begonias are vulnerable to habitat changes — B. gomantongensis and B. postarii that grow in the damp and shaded conditions around the base of the hill are vulnerable to clearance or disturbance to the tree canopy, while all species are endangered by the periodic forest fires. Indeed, the summit vegetation of Bk. Dulong Lambu has still not recovered from the 1983 fires. Introduction The richness of the limestone flora in Peninsular Malaysia compared with the land area it occupies was highlighted by Chin (1977). Kiew (1991) ascribed this richness in part to the variety of microhabitats that limestone hills provide within a very confined area, from the damp dark conditions at the base of the hill, to the vertical walls that with increasing height become more exposed to sunlight, heat and water stress, to the variety of substrates - the friable soil at the base, the pocked boulders, crevices in rocks, peat- filled pockets and so on. Begonias are one group of plants that are well represented on limestone. For example in Peninsular Malaysia, nine of the 55-odd begonia species grow on limestone and six of these are restricted to limestone. Most limestone hills support at least one species (the peltate B. kingiana Irmsch. is the most widespread) and frequently a second species occurs. For example on Batu Caves, B. kingiana grows on shaded vertical cliffs near the base of the hill while B. phoeniogramma Ridl. grows on steep earth slopes in gullies. In Borneo, the genus Begonia is much more speciose and begonias 162 Gard. Bull. Singapore 50(2) (1998) are well represented on limestone. Indeed, it is common for several begonias to be found on the same hill. (There is still a great deal to be learnt about Bornean begonias as there are more species undescribed than described and field collecting continues to discover novalties). This paper examines niche partitioning of begonias on Bukit Dulong Lambu (5° 31°30”N 118° 4°15”E), a tower karst massif 229 m high in the Gomantong Virgin Jungle Reserve. This limestone hill is famous for its caves from which birds’ nests are harvested. It is better known as Gomantong Cave (Lim & Kiew, 1997). Four begonias are known from this massif, two are endemic to it (B. malachosticta Sands and B. gomantongensis, described here as a new species), one (B. postarii, also described here as a new species) is found in another two limestone localities on the Kinabatangan River, and B. gueritziana Gibbs, a widespread species, is found, not only on almost every limestone outcrop, but also on other rock types too. Populations of these four species were observed in the field to see whether differences in their habitat characteristics could be discerned to explain how they co-exist in the same locality. At the base of the hill grow B. gomantongensis and B. postarii, both confined to damp shaded conditions below intact forest canopy. Ascending the hill, these two species are replaced by B. malachosticta that grows in rocky crevices in the sheer cliffs or on jagged outcrops. It is not found in deep shade but occurs at a height where the tree canopy begins to open up. Unlike any begonia species in Peninsular Malaysia, it can grow on the exposed summit fully exposed to full sunlight, heat and presumably also water stress. At one time, its population must have been quite plentiful on the summit as it is reported as eaten as a vegetable by the birdnest collectors who camp on the summit of Bk. Dulong Lambu. Leaves of some begonias are pleasantly sourish, cf. Reza and Kiew (1998). However, it is no longer found on the highly disturbed summit except for the inaccessible shoulders. Begonia gueritziana is also not found growing in the deep shade at the base of the cliff nor is it found growing in habitats exposed to full sunlight. Unlike the other three species, it produces a compact rhizome, which grows closely appressed to its substrate. (The other three species are erect begonias, B. malachosticta produces woody cane-like stems, while B. gomantongensis and B. postarii have rather lax stems that tend to become decumbent). B. gueritziana is most frequently found in soil-filled crevices and is particularly common on lightly-shaded, humus-covered ledges. Of the other two species, B. postarii is a delicate begonia with soft leaves and it is not found in conditions exposed to either hot or dry conditions. It grows most profusely in sheltered, deeply shaded habitats at the base of the hill. Substrate does not seem critical as it grows both on Niche Partitioning in Limestone Begonias 163 soil, limestone boulders and even on the base of tree trunks (semi-epiphytic). Its distribution overlaps with that of B. gomantongensis and in one shaded area where the jagged limestone bedrock is exposed at soil level, both species grow together in profusion. However, B. gomantongensis 1s also found growing on vertical rock faces, but only in deep shade within 3 m of ground level where the cliff or boulder is covered by a black layer of algae. In contrast, B. postarii is never found on such deeply shaded vertical rock faces. This rich biodiversity of begonias is a characteristic of the Sabah limestone flora and many other hills support at least three species, one at the base, another on the summit, as well as B. gueritziana. In this respect, Sabah is richer in its limestone begonia flora than Peninsular Malaysia where no begonia is found above the tree canopy, i.e no begonia is adapted to living on the exposed summit. The niche partitioning of these four begonias illustrates the importance of protecting Bukit Dulong Lambu from habitat disturbance in order to conserve the rich biodiversity of the limestone flora. In the past, limestone hills in Sabah have been particularly vulnerable to burning in drought periods and the summit vegetation of hills such as Bk. Dulong Lambu, Gunung Madai and Batu Batangan, for example, have still not recovered from the Great Burn of 1983 and the blackened trunks of large dead trees still stand and the summit is covered by a tangle of creepers. The flora that grows at the base of the hill is particularly vulnerable to forest clearance that opens up the canopy and would expose tender species, such as B. gomantongensis and B. postarii to the drying conditions of high light and temperature. To conserve the limestone flora of these hills, it is therefore necessary to protect a broad buffer zone of primary forest around the base of the hill to prevent fire spreading from the surrounding area. In addition, Bk. Dulong Lambu and G. Madai remain particularly vulnerable to accidental fires as birdnest collectors live on or close to these hills. It is, however, in their interest to prevent fires as smoke from the burning drives away the birdnest swiftlets. While Bk. Dulong Lambu and G. Madai are protected within virgin jungle reserves, the majority of limestone localities are not adequately protected as they are not surrounded by protected forest (Lim and Kiew, 1997). In view of the extremely rich biodiversity of the limestone flora, a strategy needs to be implemented to protect key limestone localities of which Bk. Dulong Lambu is one (Kiew, in press). 164 Gard. Bull. Singapore 50(2) (1998) Two New Begonia Species Both new species belong to Section Petermannia in possessing an erect, branching habit; stamens produced on an elongated column; obovate anthers, which are about the same length as the filament; female flowers with five tepals, a trilocular ovary with three bifurcating styles, the stigma forming a continuous spiral papillose band; and trilocular fruits with three equal wings, bilamellate placentas and caducous styles. Begonia gomantongensis also possesses male flowers with two tepals, which is characteristic of Section Petermannia. However, B. postarii has male flowers with four tepals. Sands (1990) has already noted that a few species of this section, such as B. malachosticta, are anomalous in this respect. Begonia gomantongensis Kiew sp. nov. Holotype: James Awing SAN 47257 (SAN, unicate). A Begonia pryeriana Ridley petiolis longioribus laminis latioribus et fructibus verruculosis differt. Erect monoecious begonia up to 60 cm tall, sparsely branched. Young stem, petioles and lower surface of veins minutely bristly. Stems green, purplish at nodes, up to 4-5 mm thick when dry. Stipules green, narrowly lanceolate, up to 2 cm long, 5 mm at base strongly tapered towards acute apex. Leaves alternate, distant and held horizontally. Petiole deep purplish towards apex and base, 10-16 cm long. Lamina glossy, mid-green to dark green above, pale green beneath, unpatterned, glabrous above, obliquely subrotund, 13-16(-30) by 13.5—14(-23) cm, base cordate, not overlapping, basal lobe rounded 6-8 cm long, margin minutely serrulate, apex shortly acuminate, acumen 0.5 cm long; veins slightly impressed above, conspicuously prominent beneath, main vein and 4 lateral veins radiating from the petiole with an additional 2-4 veins supporting the basal lobes, veins bifurcating three times before reaching the margin. /nflorescence axillary, protogynous, with 1-2 female flowers produced from the leaf axil and many male flowers on an erect monopodial rachis up to 5 cm long with short cymose branches c. 0.5 cm long. Bracts foliaceous, broadly ovate to semi-circular, up to 25 by 17 mm, diminishing in size towards the shoot apex, margin undulate, upper bracts enveloping clusters of male flowers and overlapping with bract above. Female flowers with pale yellow-green pedicels c. 10-12 mm long, ovary oblong tapered into pedicel, 16 by 11 mm, outer surface verruculose, wings pale yellowish green c. 4 mm wide, 3-loculate, placentas axile, bilamellate with many ovules on both surfaces, tepals 5, white, margin Niche Partitioning in Limestone Begonias 165 entire, apex acute, outer tepals ovate 11 by 7 mm, inner narrower 7 by 4 mm, styles c. 3-4 mm long divided to base, bifurcating, stigma papillose forming a continuous twisted band. Male flowers with slender pedicels 3—3.5 mm long, densely bristly, tepals 2, white, rotund, 3.5—4 by 3-3.5 mm, stamens (45—)51(—53) in an obovoid cluster 1.25 by 2 mm, torus columnar c. 0.75 mm long, filament c. 0.5 mm long, anther yellow, ovoid, c. 1 by 0.5 mm, apex emarginate. Capsule pendant, pedicel 1.5—2.5 cm long, oblong, 17-20 by 13-15 mm, base broadly rounded, wings 3, equal, 4-5 mm wide, drying stiffly papery (not fibrous), wing tip abruptly truncate or rounded, 3-loculate with the locule reaching to the base, outer surface of locule flat and completely verruculose, dehiscing along suture between locule and wing, placentas axile, bilamellate with numerous minute seeds on both surfaces, styles caducous. Seeds broadly ovoid, 0.3 by 0.2 mm, base truncate, apex rounded, testa strongly reticulate. Distribution: Endemic to Sabah, known only from Bk. Dulong Lambu, Sandakan District. Habitat: Base of limestone hill in deep shade on boulders or cliff faces. Specimens examined: James Awing SAN 47257 11 Oct 1964 (SAN), S.P. Lim & Ubaldus LSP 785 6 Aug 1996 (SAN, SING), R. Kiew & S.P. Lim BDL 3 29 Oct 1996 (SAN, SING). Notes: In its inflorescence with 1—2 female flowers at the base and an erect rachis bearing male flowers and its oblong capsules 17-19 mm long, it resembles B. pryeriana Ridl., which was first described from Sandakan, Sabah. However, it would not be mistaken for this species as the leaves of B. gomantongensis are subrotund and 13-16 cm wide (those of B. pryeriana are lanceolate acuminate and about 5 cm wide) and its petioles are much longer (only 2.5 cm long in B. pryeriana). In addition, the outer wall of the locules of the ovary and capsule are remarkable in being verruculose, those of B. pryeriana are smooth. In the field, B. gomantongensis is a very striking species as its large subrotund leaves are held horizontally and look like tea plates. The leaves form a perfect leaf mosaic without any overlap. Label notes on the type specimen record flower colour as yellow. All the plants I have seen have white tepals. Begonia postarii Kiew sp. nov. Type: R. Kiew & Lim S. P. RK 422] Bukit Panggi, Kinabatangan (holo 166 Gard. Bull. Singapore 50(2) (1998) SAN, iso SING) Figure 1. A Begonia congesta Ridley petiolis longioribus, laminis brevioribus, staminibus 23 vel 35 (non 12) et fructibus brevioribus differt. Weak erect but rather straggling branched monoecious begonia to 50 cm tall becoming decumbent and rooting at nodes. Plant hispid with uniseriate, glandular, white (sometimes red) trichomes 1-3 mm long on the stem, stipules, petioles and lamina, bracts, inflorescence, outside of male and female tepals and the ovary, particularly dense and brownish on young stems, petioles and lower surface of veins, on upper lamina surface trichomes have raised bases. Stem reddish, c. 8 mm thick when dry. Stipules foliaceous, broadly elliptic with a distinct dorsal midrib, up to 2.5 by 1.5 cm, margin entire with dense fringe of hairs, apex rounded or slightly acute, persistent. Leaves alternate and distant. Petiole reddish, (4-)6-8 cm long. Lamina plain pale green above and beneath, soft and thin in life, drying thinly papery, obliquely ovate, (8.5—)10-11(-14) by 7—9(-12) cm, base cordate (not overlapping), basal lobe rounded, (3.5—)6-8 cm long, margin minutely dentate, apex cuspidate, acumen to 1.25-1.5 cm long, main veins 5, of almost equal length radiating from the petiole and bifurcating twice before reaching the margin, with 2 minor veins running into basal lobes, slightly impressed above and raised beneath. /nflorescence erect and projecting above leaves, paniculate, 13.5-18 cm long, peduncle 5.5—7-cm long, lowest 1-2 branches 1.5-3.5 cm long, each producing 2-3 female flowers, upper branches branching three times, ultimate branch 1.5—-3 cm long with many male flowers. Bracts, peduncle and rachis pale green and fleshy, upper bracts tipped red and ultimate branches of rachis reddish. Lower bracts broadly oval up to 1.5 by 1 cm, margin minutely toothed; ultimate bracteoles broadly ovate, c. 3 by 4mm with dentate margin each tooth tipped by long trichome. Female flowers with reddish pedicels c. 7-10 mm long, ovary broadly ovate, 8 by 9.5-10 mm, wings crimson, c. 2 mm wide, 3-loculate, placentas axile, bilamellate with many ovules on both surfaces, outer surface smooth, tepals 5, pink, margin entire, apex rounded, outer broadly ovate 8 by 4 mm, inner narrower 4.5 by 2.5 mm, slightly tapered to base, styles 3 mm long divided to base, bifurcating, stigma papillose forming a continuous twisted band. Male flowers with pedicels up to 8 mm long, tepals 4, outer 2 white tinged pink merging to cerise towards centre, elliptic 7-8 by 4.5-5 mm, apex rounded, conspicuously hirsute outside, inner pair glabrous, narrowly lanceolate, 7 by 1.3-2 mm, stamens 23-35 in a conical cluster c. 3 by 1.7 mm, torus columnar c. 2 mm long, filament c. 0.5 mm long, anthers yellow, 1 by 0.5 mm, apex emarginate. Capsule pendulous on Niche Partitioning in Limestone Begonias 167 pe vga’ if), Vit RY NN SYS AS top tly p tN ae 4 / Witte t Np len Ye Figure 1. Begonia postarii A Habit (x 0.4), B Anther (x 8), C Male flower (x 2), D Androecium (x 4.8), E Branchlet with male flowers (x 1.6), F Male bud (x 2.8), G Capsule (x 1.2). 168 Gard. Bull. Singapore 50(2) (1998) a thin thread-like stalk c. 1.5—2 cm long, broadly ovate, (8—)10 by (9-)13 mm, hirsute outside, 3-loculate, locules bulging, wings 3 equal, narrow 3 mm wide, distinctly rounded at base, truncate distally, tip sometimes acute, drying thin and papery and dehiscing along suture between locule and wing, styles caducous leaving a scar. Seeds ovoid, apex and base truncate, c. 0.4 by 0.3 mm, testa strongly reticulate. Distribution: Endemic to Sabah, known only from two limestone hills along the Kinabatangan River (Bukit Dulang Lambu and Bukit Panggi) and from a low unnamed outcrop in the Subak Estate on opposite side of the Kinabatangan River from Bukit Garam. Habitat: At base of the limestone hill (but not on cliff faces) at about 100 m a.s.l., growing in deep shade in sheltered, damp habitats on soil, low limestone boulders or base of tree trunks. Specimens examined: Bukit Dulong Lambu Joseph B. et al. SAN 122763 21 Jan 1988 (SAN), Lim S.P. & Ubaldus LSP 802 7 August 1996 (SAN, SING), R. Kiew & Lim S.P. BDL4 29 Oct 1996 (K, SAR, SAN, SING); Bk. Panggi R. Kiew & Lim S.P. RK4221 19 Sept 1996 (SAN, SING); Subak Estate, Lower Kinabatangan River J. Dransfield et al. JD5770 17 Oct 1979 (SAN). Notes: This softly hairy begonia is quite unlike any other limestone species in Sabah. Neither are any of the Sarawak limestone begonias as hairy. B. congesta Ridl. from limestone in Sarawak is scantily hairy on the young shoots, leaf margins and lower vein surface but B. postarii is distinct from this species in leaf indumentum, shape and margin (the lamina of B. congesta is longer and more oblong ranging from 15-23 cm in length, is more than 1.5 times longer than wide and its margin is undulate), longer petiole c. 7.5 cm long, glabrous male flowers and fruits, and a longer oblong fruit 2.5 by 1.25 cm and, according to Ridley, only 12 stamens. This begonia is named in honour of Postar Jaiwit, tree climber at SAN, whose sharp eyes and interest in plants led him to discover this species on Bk. Panggi. | Acknowledgements This work forms part of the collaborative project between Universiti Putra Malaysia and Forest Research Centre, Sabah Forest Department on “The Biodiversity and Conservation of the Limestone Flora of Sabah’. The author Niche Partitioning in Limestone Begonias 169 thanks the Ministry of Science, Technology and the Environment, Malaysia, for funding the project under IRPA Grant No. 52858; A. Berhaman and Lim S.P. for organising field excursions; herbarium staff at SAN for help in the field and in processing herbarium specimens; and Mdm Yap P. H. for the botanical drawing of Begonia postarii. The author particularly appreciates the constant and enthusiastic support of the project by Wong Khoon Meng and Robert C. Ong. Mark J.E. Coode is thanked for correcting the latin. References Chin, S. C. 1977. The limestone hill flora of Malaya. I. Gardens’ Bulletin Singapore. 30: 165-219. Kiew, R. 1991. The limestone flora. In: R. Kiew (ed.) The State of Nature Conservation in Malaysia. Malayan Nature Society, Kuala Lumpur, Malaysia. pp. 42-50. Kiew, R. (in press) Towards a limestone flora of Sabah. In: K.M. Wong (ed.). Proceedings of Stone Memorial Symposium, Malaysian Nature Society, Kuala Lumpur, Malaysia. Lim, S.P. and R. Kiew. 1997. Gazetteer of limestone localities in Sabah, Borneo. Gardens’ Bulletin Singapore. 49: 111-118. Reza Azmi and R. Kiew. 1998. Begonia lazat (Begoniaceae), a new culinary begonia from Borneo. Gardens’ Bulletin Singapore. 50: 43-48. Sands, M.J.S. 1990. Six new begonias from Sabah. Kew Magazine. 7: 57-85. r i i , ' e r 4 , ; he be ah 1. i ey ne 4 4 A Mol . 7 bd . (eh a1 a a tee ine Ao Vw 1 : ” r Able i R me i totals nd wi aide ioe my * uh if Ma m ays / " ties i mae i : vig? ney a © ut 198 Pm", a hee Ph aa ee | a | Ah 1 4 SAG N VEAL rer ij 0 I ? ' ‘4 ¥ b ' | i ke, 4 f i 1 | . | up ect qc Lie we { ae Dr eretagts) we pie ‘as ene a, } y tT. * ni hh i Pi Nia it , thn’ alae hy ev ait oA et tal te i * J } ris, i } . & \ 4 "il ‘ ‘ : ee L a vy >) a ‘ 4 ; es Aw i uta aa Thats Pith) oe id o a i i ne f Bi ib Pau we Nerstichir Kt: Las avai ; . ; a ra, ir Uys ¥ wor 4 Suay , ’ 7, »' ih A Vix od 2 r f y 4 > ph) i Piva i i ; re hin j . hy bis 1) oe i Be Aids i Wit 17 i ‘ iv? raw ee ; ? Par er ae a ‘ mire GH % 5. Fi ig ; fi) Ge PG AAT | PS ui Na at ve Oh wre Ly nes Res 1 rely ; iy: 7 ae i ny ’ J A Ve ‘ tye } oy hy) ree ORY: at iN c he a ! i ’ i ry ; : . ' PF i? os 7 : 2 ra y ii r Ava?) ere ‘ 4 P] iT \ ity Oh « ahr. f «! 4 7 7] a ive ’ ha I , ‘* 1 ‘ \s ia st r ' eM brat apie Pettit é ri vee » nf Lite AM Recs, ey) UP f rea i | ty ( SEA 7 d at i] 44 in f Ww imi yi ) « ik ' I di LS cs rae 7 il il sleet ae wi j } ‘ { me 7 ci 7 P . i 5 i] PA ' ¢ i) «@ ; ‘ Gardens’ Bulletin Singapore 50 (1998) 171-183. A Survey of Termites in the Singapore Botanic Gardens Rain Forest G.T. CHOO-TOH!, S.L. CHAW!, C.E.Z CHAN’, D.H.W. GOH? AND 0) io EN ad he ‘Singapore Botanic Gardens, Cluny Road, Singapore 259569 ?Anglo-Chinese Junior College, 25 Dover Close East, Singapore 139745 Abstract A survey on termites in the Singapore Botanic Gardens Rain Forest included termite collection and quantitative assessment of vegetation, dead trees and wood litter in 15 random sample plots, which covered 7.5% area of the forest. Termite infestation was high in the northern zone, moderate in the central and light in the southern zones. The abundance of termite-infested microhabitats shows a positive relationship with the number of big trees, dead standing trees and ground timber, and a negative relationship with the number of herbs. A total of 22 termite species were found. Three genera of gallery-forming termites identified were Bulbitermes, Nasutitermes and Microcerotermes. The major fungus-growing genera were Macrotermes, Odontotermes and Microtermes. The ground-nesting genera included Termes, Dicuspiditermes, Hospitalitermes and Prohamitermes. The other genera found were Coptotermes, Schedorhinotermes, Subulitermes and Procapritermes. Three new records for Singapore are Bulbitermes borneensis, B. constrictus and Microcerotermes crassus. Microcerotermes and Nasutitermes are involved in the self-pruning of trees. The Bulbitermes, Macrotermes and Odontotermes are dominant in ground timber. The different species richness and uneven distribution of termites in different parts of the forest is attributed, not only to the differences in forest structure and flora, but also the degree to which the forest floor space has been depleted for visitor and horticultural activities. Introduction The Singapore Botanic Gardens (SBG) includes a 6.3-ha plot of forest, which according to Corlett (1992), no longer resembles primary rain forest in structure or flora. The insect fauna is similarly depleted as exemplified by the ants (Murphy, 1973). There are few publications on the vegetation in the early years although Corner (1935) did make an interesting study of higher fungi. Turner et al. (1996) surveyed the vegetation and compared it with historical records of herbarium collections made since 1893 and revealed that the forest has suffered a significant loss of species. They attributed this to the rampant growth of climbers, dense undergrowth of exotic and clonal herbs and a loss of animal disposal agents, which have led to a low density of saplings and seedlings and hence no recruitment of the high forest species. Calophyllum ferrugineum (Guttiferae) is an exception being numerically the dominant species making up 26.2% of the 172 Gard. Bull. Singapore 50(2) (1998) trees. The death of trees, caused either by lightning strike, disease or other factors, has given rise to a great variety of fallen and standing dead timber in all stages of decay. The aim of the survey of the termite fauna in this disturbed rain forest was to record termite species diversity and their distribution and also the roles the different species play in the decomposition of the dead timber, as well as the effects the different vegetation life forms have on termite distribution. Methods of Study Sampling methods A map of SBG rain forest by the Nature Conservation Branch was used for the selection of sampling points. The map has grid lines spaced 40 m apart, which give rise to 45 points evenly spaced over the entire area. Using a random table (Snedicor & Cochran, 1969), one third of these intersection points, 1.e. 15, were picked as the sample points for the termite survey. These sample points, i.e. points 5, 6, 8, 15, 16, 17, 20, 24, 28, 31, 34, 40, 41, 42, 45, were located and pegged on site (Fig.1). Around each point, a sample plot of radius 10 m was cordoned off for the survey. The 15 sample plots covered a total area of 7.5% of the 6.3-ha rain forest. The sampling of termites and the quantitative assessment of the vegetation and wood litter were conducted once to three times a week from end May to end August 1998. Termite Collection and Identification Within each sample plot, the following microhabitats were searched: ground nests, inside of dead logs, fallen branches, leaf litter under rotten logs and galleries on logs and tree trunks up to a height of 2 m above ground level. Termites in soil, high canopy, tree stumps and fallen macro-timber equal or greater than 30 cm diameter were not surveyed. From each infested microhabitat, one series of termites comprising soldiers and workers was collected for each species found. The soldiers were identified to species level as far as possible using taxonomic keys (Thapa, 1977; Tho, 1992; Holmgren, 1913). The location of each infested microhabitat was mapped. Quantitative assessment of vegetation and wood litter Each sample plot was divided into quarters and a systematic count of vegetation categorised into trees, shrubs, herbs and climbers was made. To correctly categorise the plants, especially the seedlings, to their life forms, the identity of the plants was confirmed with assistance of SBG Herbarium Termites in SBG Rain Forest we ry aa SB wa ! o = os Sof 2 a 3 4 Plot 5 Plot 6 6 = eos \ N Highly Infested ~~ [-\\ 2 & Vv Northern Zone A 8 15 16 7S aierapte plots as — No. of infested microhabitats Moderately Infested Central Zone B Sample plots on =~ | : oT 16 TS w 4 £8 12 2 2 io 4 ee, pa te Ze + 24 By Sample plots os infested Southern Zone Ay AN is lo we a | 4 a ne) neo Figure 1. The three zones of SBG Rain Forest with different termite infestation levels. The locations of the termite infested microhabitats within the sample plots are marked by black dots, while their abundance is depicted by the histograms for A. northern zone, B. central zone and C. southern zone. 174 Gard. Bull. Singapore 50(2) (1998) staff and checked to be consistent with previous studies (Turner et al., 1996). The trees were further categorised into big trees (=>0.1 m dbh), the remainder into smaller trees and seedlings (below 1.5 m tall). Because the shrubs were very low in number, they were grouped into the same category as herbs during data analysis. Palm trees were assessed as a separate category as they were mostly in seedling stages and the mature ones, if present, were in very small numbers. The abundance of the wood litter on the forest floor was assessed by counting the number of logs and fallen branches above 1.5 cm in diameter. | Results Distribution and infestation level of termites For each sample plot, the locations of individual termite-infested microhabitats (e.g. a nest, a log or a living tree, etc.) were mapped and their abundance was depicted (Fig. 1). Termites were found in 98 microhabitats in only 13 of the 15 sample plots. Of these infested microhabitats, 65 (1.e.66.5% ) were in the northern part of the rain forest, 25 (1.e.25.5%) in the central part and 8 (i.e.8.2%) in the southern part. Using F Test followed by LSD tests (Snedicor & Cochran, 1969), the number of termite infested microhabitats in the northern zone was found to be significantly higher than that in the southern zone (P=0.01) and in the central zone (P=0.05). Based on the abundance of the infested microhabitats, which reflects the infestation level of the termites, the SBG Rain Forest can therefore be divided into three zones: a) the highly infested northern zone, b) the moderately infested central zone and c) the lightly infested southern zone (Fig. 1). The influence of vegetation and wood litter on termite infestation The abundance of termite-infested microhabitats, the different types of vegetation and logs/branches in the sample plots are depicted in Fig. 2A- G. The abundance of termite-infested microhabitats (Fig. 2A) follows almost the same trend as the abundance of the wood litter (Fig. 2B), big trees => 0.1 m dbh (Fig. 2C) and dead standing trees (Fig. 2D), that is, high in the northern zone, moderate in the central zone, and low or absent in the south. On the other hand, a comparison of Fig. 2A and Fig. 2E shows that the abundance of termite-infested microhabitats was in contrast to the abundance of herbs and shrubs in the sample plots. Fig. 2F and 2G show that the abundance of palm trees and climbers in the sample plots do not have any influence on the abundance of the termite-infested microhabitats. Termites in SBG Rain Forest 75 A E ; 8 1000 + B 2. | = 800 } ee: % 600 or . S 400 + = cz : 5 200 4 za : . : By Osea Wom ith 520424) 28 St) 38'40) 411142 .45 | 5 6 8 15 16 17 20 24 28 31 34 40 41 42 45 Sample plots Sample plots {pial oe 8 B 3 300 S 250 o 7) 5 200 : E Oo 50 6 5 6 8 15 16 17 20 24 28 31 34 40 41 42 45 Sample plots Sample plots Cc G 40 500 3 2 | © 30 2 ae Fe E 300 2 : = 200 5 TE : 2?) = 0 BREESE SSR Es : : ds : 0 + : cS Aare re ee ree rere 5 6 8 15 16 17 20 24 28 31 34 40 41 42 45 Sample plots Sample plots No. of dead trees 15 16 17 20 24 28 31 34 40 41 42 45 Sample plots Figure 2. Characteristics of the sample plots A. No. of infested microhabitats; B. No. of logs and fallen branches; C. No. of mature trees (=> 0.1 m dbh); D. No. of dead trees; E. No. of herbs and shrubs; F. No. of palm trees; G. No. of climbers. 176 Gard. Bull. Singapore 50(2) (1998) The number of logs and fallen branches and the various vegetation life forms were plotted against the corresponding number of termite-infested microhabitats (Fig. 3A-F). As expected, the figures indicate that the abundance of termite-infested sites has a positive relationship with the abundance of wood litter (Fig. 3A), big trees (Fig. 3B) and dead trees (Fig. 3C), and a negative relationship with the abundance of herbs and shrubs (Fig. 3D). There is no relationship with the presence of palms and climbers (Fig. 3E and 3F). A D rs] “ 900 cd 2 s 2 800 a 5 700 “ s 8 600 = 4 500 7) © 400 8 = m0 : A S Z 100 = 0 0 5 10 15 20 No. of termite-infested microhabitats No. of termite-infested microhabitats B E No. of big trees 0 5 10 15 20 0 E 10 15 20 No. of termite-infested microhabitats No. of termite-infested microhabitats No. of dead trees @ No. of climbers On fA OD 0 5 10 15 20 0 5 10 15 20 No. of termite-infested microhabitats No. of termite-infested microhabitats Figure 3. The relationship between the abundance of termite-infested microhabitats and the various types of vegetation and wood litter. A. Logs and fallen branches; B. Big trees =>0.1 m dbh; C. Dead trees; D. Herbs and shrubs; E. Palms; F. Climbers. Termites in SBG Rain Forest aa The termite fauna a) Species diversity A total of 102 series of termites were collected from the 98 microhabitats (4 of the microhabitats had series of two co-existing termite species). These termites were identified to 22 species. The species diversity of the northern part of the forest was distinctly different from those of the other two parts. The northern zone had 20 of the 22 species, the central zone 10 species, and the southern zone only 6 species. All but 2 species in the central and southern zones were found in the northern zone. b) The habitats, feeding and nesting behaviour Table 1 indicates the occurrence of the 22 species of termites, which are described below: 1) The wood-feeding cum gallery-forming termites As shown in Table 1, Bulbitermes was the commonest and most widespread genus, which made up 29.4% of the 102 series collected and occurred in 69.2% of the 13 infested plots. It was represented by three species: B. borneensis, B. constrictus and B. kraepelini. The latter was rare, being collected only once. B. constrictus was slightly more widespread than B. borneensis. The Bulbitermes consumed mainly the sapwood and sometimes the heartwood of dead fallen branches. They frequently formed brown, crumbly galleries on the branches and logs where they were found. Two other genera that form galleries are the Nasutitermes, which are closely related to Bulbitermes, and Microcerotermes. The latter made up 15.7% and Nasutitermes 10.8% of the series collected. Although Microcerotermes was second in frequency of collection to Bulbitermes, it was comparatively very localized in its occurrence, as reflected by the lower percent incidence of 46.2%. Likewise, Nasutitermes was also very localized in its occurrence, being found in only 38.5% of the plots studied. Of the two Nasutitermes species present, N. matangensiformis was rare being only collected once. N. havilandi formed galleries on both dead and live trees and was confined mainly to the northern zone. It was highly concentrated south of sample plots 5 and 6. The Nasutitermes fed on the dead branches of both live and dead trees. On dead fallen timber, they consumed both the sapwood and heartwood, often after these had been invaded and deserted by other species such as the Macrotermes. They were found mainly inside small stems or branches as small as 1.5 cm diameter. Microcerotermes was represented by three species: M. serrula, M. havilandi and M. crassus. M. serrula was found once, in a nest at the base of a live tree. M. havilandi formed galleries at two plots. M. crassus, the 178 Gard. Bull. Singapore 50(2) (1998) most common species, was collected from many different infested microhabitats including seven around a ground nest in plot 15. It formed characteristic small and hard galleries on both live and dead trees as well as fallen branches. It attacked the sapwood of dead branches as small as 1.5 cm diameter. Table 1. Termite species collected from the Singapore Botanic Gardens Rain Forest Species No. of % of No. of plots % of plots Sample plots where series series with termites | with termites termites are found collected | collected | Bulbitermes) 1) yo) WN) M308 || ORR VS A eS ane | bormeensiss | NO MOE LE Ie ee 2. ee) De ae eS OR ee ee i ES as a | Miioroceratermess.). Vile. V6 \s jy MGB A vids Boies see a ORT Cg i 2 a EE TE IG AOE ER Po Micarat 2 SPSREEL TIS TALE AT RESET ea |e A Te Meee aE eee nn ec eT mn eee yr ll 10.8 BBS ual lain Ms eee havilandi US Jee Se PY spncatcanpersiforsmis 0 TOUT» AEA MY (iS GSR AA eee ae) | Macrotermes |. \ 5.) ASAI) 8 Se es OA eC eee ae en ee AR eee Te | paclaccensis 97 NT | | Odontotermes” 5) NN SA 8 SET ag aa Me ee Se Mn ie a Soa se Te) ee OTR ey NLT WT RI Eee er Dermestids a Li A ATR AD Nila is gh icy, Wisp ee em SMe MSE eS See | Dicuspiditermes (0.4 3 ).2 2 |wwe 2eO cody pa GN yal ae Al fen a 2 : RE a Hospratitermes ©) 2 2.0" ee Tae aa umbrinus ae PO eee Atevonermes 220 2 aA STG EY PE | Schedorhinotermes °F) 20 NG | So, SARIS a) a a ae Te A AE RS :Copiotermes 0 eve 1 R00) 20.050 as i eee ee A RE es Procapritermes i. iN ipdiyie dowd. alu co U Lei ee gee LT a A RAs ANAL RGD WS AI yA a Prohamitermes RES GER MR ET ME SE ES mirabilis i as ee PYeudiermes AO | ivlidensipedep NET Vian ece tna ies aad Termites in SBG Rain Forest 179 ii) The wood-feeding cum leaf-litter feeding termites The two common and widespread genera found belong to Macrotermes and Odontotermes (Macrotermitinae). These are capable of growing fungi in their underground nests (Collins er al. 1983; Collins, 1981). Typical signs of their presence are the clayey soil refill inside the burrowed dead branches or stems. They were found inside branches or in the soil among leaf litter just beneath the branches. It was observed that both genera burrowed into the sapwood and sometimes heartwood under the bark of branches greater than 4 cm diameter. Of the two Macrotermes species, M. malaccensis and M. gilvus, M. malaccensis was more widespread (Table 1). A nest of M. gilvus contained fungus gardens. Of the two Odontotermes, O. denticulatus was collected from only two northern plots, and O. oblongatus was present in eight plots all over the rain forest. Microtermes was the other Macrotermitine found, with only two collections of M. pallidus from one plot. iii) The ground-nesting termites Species of Termes were most prevalent with eight series collected from five plots. Two series of 7. comis were found but no nest could be located. On the other hand, 7. rostratus was found inside two hard ground nests and in another nest freshly made with black soil refills inside a piece of rotted log. The nests were fairly easily broken into soil crumbs. The three other species, Dicuspiditermes nemorosus, Hospitalitermes umbrinus and Prohamitermes mirabilis, formed prominent ground nests. Their percent incidence was comparatively low, especially for P. mirabilis. Dicuspiditermes made raised, ball-like nests. Two Hospitalitermes nests were found, one being free-standing while the other leaned against the base of a large tree. iv) Other soil and wood-feeding termites The other species made up only 1-2% of the series collected and included Coptotermes travians and Schedorhinotermes medio-obscurus of the Rhinotermitidae, an unidentified species of the Subulitermes complex and Procapritermes augustignathus of Termitidae. Discussion The distribution and infestation levels of termites in the SBG Rain Forest, as in other ecosystems, are dependent on the availability of their food sources. These include the wood and leaf litter on the forest floor, the 180 Gard. Bull. Singapore 50(2) (1998) standing dead trees and the dead branches of live trees. The northern part of the forest has a higher population of live trees than the central and southern parts and larger amounts of ground and standing dead timber as evidenced by our quantitative assessments. Having greater amounts of food for termites explains why the northern part of the rain forest is significantly more heavily infested than the central and southern parts. In the central and southern parts, where herbaceous undergrowth thrives, there is a lower density of trees and saplings. The abundance of herbaceous plants, which are often accompanied by the presence of ants, has a negative relationship with the presence of termites. Ants have often been considered as enemies of termites (Eisner et al., 1976) and strong negative associations between ants and termites had been attributed to competition between the two groups (Fowler et al., 1980). The total of 102 series of 22 termite species collected within three months is high when compared with the 70 species collected over several years from the much larger Bukit Timah and Central Catchment Nature Reserves (Murphy, 1997). Due to the short time frame of the study, the collection though fairly intensive is by no means exhaustive. There was probably under-collection of the fungus-growing Macrotermitinae, which reportedly consumed over 75% of total organic matter taken by termites in Peninsular Malaysian rain forests (Collins et al., 1983) and 95% of leaf litter in Nigerian savanna (Collins, 1981). Other than Macrotermitinae, the termite species inhabiting the soil and the macro-timber are probably also under-represented. According to the lists of termite species compiled for Singapore’s nature reserves (Murphy, 1997) and Malaysia’s Pasoh Forest Reserve (Jones & Brendell, 1998), the soil-inhabiting species and the wood- feeding Rhinotermitidae together constitute about 40% to 50% of all species recorded. Further studies need to focus on these soil and wood-feeding termites to cover the entire termite fauna in the rain forest. Of the species collected, three are of particular interest because they have not been reported by earlier collectors and are not in the most recent list of termites species reported from Singapore (Murphy, 1997). These are the Bulbitermes borneensis, B. constrictus and Microcerotermes crassus. The northern zone of the rain forest contains some species that are common in the actively regenerating secondary forests of the Central Catchment Nature Reserves (Murphy, 1997). These include the gallery- forming Microcerotermes and Nasutitermes, which are actively involved in the self-pruning of younger trees, and the wood-feeding Bulbitermes. However, unlike the nature reserves, the nest of Bulbitermes was not found. Murphy (1997) reported the widespread occurrence of Macrotermes and Odontotermes in the more established secondary forests and the primary forests. This holds true for both genera in the northern and central zones Termites in SBG Rain Forest 181 but not in the southern zone, which has very little ground timber. Separate studies have been initiated to check whether the fallen timber in the northern zone are attacked by Kalotermitidae (dry wood termites) including species of Cryptotermes, Glyptotermes and Neotermes, which reportedly had a high incidence of attack in the nature reserves’ secondary forests. Eggleton et al. (1995) found that species diversity of termites was greatly reduced in severely disturbed plots of forests, but increased marginally in slightly disturbed regenerating plots. The different termite species diversity of the three zones of the rain forest reflects to a certain extent the different degrees to which the forest has been disturbed. The study of the forest vegetation also reveals that different parts of the forests differ in their structure and regeneration. For instance, much of the northern zone of the forest has high forest species of Calophyllum and Garcinia (Guttiferae) and Syzygium (Myrtaceae), which are common in established secondary forests (Wong et al., 1994). Other than having a large gap caused by trees struck by lightning, and an adjacent patch with a large number of small trees and rapid turnover of saplings, the northern zone is not much disturbed in most parts. This could account for its relatively high termite diversity. In contrast, the central and southern parts of the forest which have lower mature tree populations but contain more specimens of the larger sized trees e.g. dipterocarps, are much more disturbed. Here the faunal diversity, forest structure and tree regeneration are clearly affected by the rampant growth of climbers, dense undergrowth of herbs and ornamentals, and large spaces created by paved roads criss-crossing the forest, the encroachment of buildings and horticultural facilities and the proximity of the forest to public roads. The three zones were markedly different in the amount of space utilised for human activities. Assuming that plots of radius 10 m are drawn around each of the 15 or so transect points within each zone, the northern zone would have 7 plots trespassed mainly by 1.3m wide paths; whereas the central zone would have 9 plots trespassed by 2 m, 4 m and 6 m wide roads, while the southern zone would have 12 out of 14 plots depleted of floor space by 2 m, 4m, 5.5m and6m wide roads, as well as by a gazebo and potting yard. In conclusion, the reduction of space depletion of this rain forest and removal of the herbaceous undergrowth and the tree-smothering climbers will do much to minimise the loss of its termite species. Acknowledgements We are most grateful to Professor D. H. Murphy for his guidance in the project and particularly in his assistance in identifying the termites found. 182 Gard. Bull. Singapore 50(2) (1998) We also wish to thank the Chief Executive Officer of National Parks Board and the Director of Singapore Botanic Gardens for their encouragement, to the staff of SBG Herbarium for plant identification and Nature Conservation Branch especially Chew Ping Ting and Cheryl Chia for supplying the map of SBG Rain Forest and for giving information about the Biological Survey of Singapore’s nature reserves. We would also like to thank the staff of SBG Management in providing assistance with some of the field work. References Collins, N. M. 1981. The role of termites in the decomposition of wood and leaf litter in the Southern Guinea savanna of Nigeria. Oecologia. 51: 389-399 Collins, N. M., S. L. Sutton, T. C. Whitmore, and A. C. Chadwick. 1983. Termite populations and their role in litter removal in Malaysian rain forests. In: S. L. Sutton, T. C. Whitmore, and A. C. Chadwick. (eds.) Tropical Rain Forest: Ecology and Management, Blackwell, U.K. pp. 311-3235. Cores BT, 1992... The ecological transformation of Singapore, 1819- 1990. Journal of Biogeography. 19: 411-420. Corner, E. J. H. 1935. The seasonal fruiting of agarics in Malaya. Gardens’ Bulletin, Straits Settlements. 9: 79-88. Eggleton, P., D. E. Bignell, W .A. Sands, B. Waite, T. G. Wood and J. H. Lawton. 1995. The species richness of termiites (Isoptera) under differing levels of forest disturbance in the Mbalmayo Forest Reserve, southern Cameroon. Journal of Tropical Ecology. 11: 85-98. Eisner, T., I. Kriston and D. J. Aneshansley. 1976. Defensive behaviour of a termite (Nasutitermes exitiosus). Behavioural Ecology and Sociobiology. 1: 83-125. Fowler, H. G., B. L. Haines and P. Jaisson. 1982. Species diversity in leaf- cutting ants and mound-building termites in relation to the succession of vegetation in Paraguayan grasslands. In: Jaisson, P. (ed). Social Insects in the Tropics. pp. 187-201. Holmgren, N. 1913. Termitenstudien 4. Versuch einer systematishen Monographie der Termiten der orientalischen Region. Kungliga Svenska vetenskapsakademiens handlingar. 50: 1-276 Termites in SBG Rain Forest 183 Jones, D. T. and M. J. D. Brendell. 1998. The Termite (Insecta: Isoptera) Fauna of Pasoh Forest Reserve, Malaysia. The Raffles Bulletin of Zoology. 46: 79-91. Murphy, D. H. 1973. Animals in the forest ecosystem. In: Chuang, S. H., (ed). Animal Life and Nature in Singapore. Singapore University Press. pp. 53-73. Murphy, D. H. 1997. Faunistic Survey Part 2 — The Isoptera. Unpublished technical report of a survey commissioned by the National Parks Board, Singapore. Snedicor, G. W. and W. G. Cochran. 1969. Statistical Methods. The lowa State University Press. U.S.A. Thapa, R. S. 1977. Termites of Sabah. Sabah Forest Records. 12: 1-374 Tho, Y. P. 1992. Termites of Peninsular Malaysia. Malayan Forest Records.No. 36. Forest Research Institute Malaysia, Kepong. Malaysia. rent Woo kK. Ss. Chua, J. 5S. Y. Ong, BoC. Soong and H. ‘T. W.. Tan. 1996. A century of plant species loss from an isolated fragment of lowland tropical rain forest. Conservation Biology. 10: 1229-1244. Wong, Y. K.; P. T. Chew and Ali Ibrahim. 1994. The tree communities of the Central Catchment Nature Reserves. Gardens’ Bulletin, Singapore. 46: 37-78. Gardens’ Bulletin Singapore 50 (1998) 185-196. The Unique Elements of the Limestone Flora of Batu Tengar Cave (Segarong), Sabah, Malaysia. RUTH KIEW Singapore Botanic Gardens, Cluny Road, Singapore 259569 Abstract Unique elements of the flora of the Batu Tengar Cave include Begonia keithii (Begoniaceae), a new species endemic to this hill, and the phytogeographic affinities of its flora, which are not only with the nearby Madai limestone hills, but also with limestone in Kalimantan (Borneo) and the Philippines. Notes on species of special interest, Euphorbia lacei (Euphorbiaceae), Jmpatiens winkleri (Balsaminaceae) and Paraboea madaiensis (Gesneriaceae) are given. Introduction Batu Tengar Cave (4° 42’ 00”N 118° 00’ 30”E) is located within the Segarong Protected Forest Reserve (Lim & Kiew, 1997), adjacent to the Segarong River. Commercial quantities of bird nests are still collected from the cave. The earliest botanical collections from this locality (called ‘Segarong’ on herbarium labels) date from 1938 when Symington with Agama and then Keith collected a handful of plants. These few collections are, however, extremely interesting as they included the first collection of a balsam from limestone in Sabah, the succulent Euphorbia lacei (Euphorbiaceae) and a begonia with decorative leaves. In May 1997, as part of the botanical survey for “The Biodiversity and Conservation of the Limestone Flora of Sabah Project’, this hill was revisited to relocate these species and to make a general collection of fertile material (Kiew, in press). The hill is now a 25-minute journey from Semporna by speedboat to the jetty built by the birdnest concessionaire on the Segarong River. It is then a short walk through cultivated land to the hill base that is surrounded by a narrow strip of forest. The summit of the hill (about 200 m a.s.1.) closest to the river shows signs of burning being bare of vegetation apart from dead tree trunks that are still standing (Fig. 1). However, the way to the cave on a well-beaten track is through primary forest that provides the damp shaded conditions that support lush vegetation. Once the cliff base is reached, the path ascends a narrow gully, where on the large scattered limestone boulders, Impatiens winkleri (Balsaminaceae) and Epithema 186 Gard. Bull. Singapore 50(2) (1998) dolichopodum Hillier & Burtt (Gesneriaceae) are encountered in abundance. As the path ascends, it becomes steeper and the canopy is more open and here Begonia keithii (the new species described below) is first encountered. Higher up on the exposed shoulders of the summit it is abundant. It is an extremely decorative species with glossy cardinal red stems that look lacquered, narrowly scolloped leaves and bright red male flower buds. The summit vegetation is remarkable being completely dominated by a cactus-like forest of Euphorbia lacei with a candelabrum-like crown (Fig. 2). This is unique to Batu Tengar Cave as no other limestone hill in Sabah has this type of forest on the summit. On the sheer cliffs, Paraboea madaiensis (Gesneriaceae) 1s abundant. Phytogeographic Affinities of the Flora Apart from the begonia, which is endemic to Batu Tengar Cave itself, two species (Epithema dolichopodum and Paraboea madaiensis) are endemic to Sabah, while /mpatiens winkleri and Euphorbia lacei, while not endemic, are rare in Sabah being known from a few other localities. Impatiens winkleri was first described from south Kalimantan and in Sabah is known from just four other limestone localities all in the south east (Fig. 3). Discounting common species that are everywhere widespread, it is the only rare species from Sabah limestone that is known to occur in both Sabah and Kalimantan. Euphorbia lacei is rare in Sabah being known from just three localities (all close to Semporna, but the other two are not limestone). However, elsewhere it is a widespread species ranging from Myanmar, though Thailand and IndoChina to the Philippines where it reaches Palawan. Three of these species are also known from Gunung Madai. Paraboea madaiensis is only known from Gunung Madai and Batu Tengar Cave, Impatiens winkleri is known also from Baturong, Batu Belas and Tempadong, while Epithema dolichopodum is known from six other localities, which are all located in the southeast, Batu Tengar Cave, Baturong, G. Madai and Tempadong (Fig. 3), except for one population on limestone on Pulau Balambangan and one collected by Elmer (Elmer 20569) from Tawau but without a specific locality. Termites in SBG Rain Forest 179 ii) The wood-feeding cum leaf-litter feeding termites The two common and widespread genera found belong to Macrotermes and Odontotermes (Macrotermitinae). These are capable of growing fungi in their underground nests (Collins et al. 1983; Collins, 1981). Typical signs of their presence are the clayey soil refill inside the burrowed dead branches or stems. They were found inside branches or in the soil among leaf litter just beneath the branches. It was observed that both genera burrowed into the sapwood and sometimes heartwood under the bark of branches greater than 4 cm diameter. Of the two Macrotermes species, M. malaccensis and M. gilvus, M. malaccensis was more widespread (Table 1). A nest of M. gilvus contained fungus gardens. Of the two Odontotermes, O. denticulatus was collected from only two northern plots, and O. oblongatus was present in eight plots all over the rain forest. Microtermes was the other Macrotermitine found, with only two collections of M. pallidus from one plot. iil) The ground-nesting termites Species of Termes were most prevalent with eight series collected from five plots. Two series of 7. comis were found but no nest could be located. On the other hand, 7. rostratus was found inside two hard ground nests and in another nest freshly made with black soil refills inside a piece of rotted log. The nests were fairly easily broken into soil crumbs. The three other species, Dicuspiditermes nemorosus, Hospitalitermes umbrinus and Prohamitermes mirabilis, formed prominent ground nests. Their percent incidence was comparatively low, especially for P. mirabilis. Dicuspiditermes made raised, ball-like nests. Two Hospitalitermes nests were found, one being free-standing while the other leaned against the base of a large tree. iv) Other soil and wood-feeding termites The other species made up only 1-2% of the series collected and included Coptotermes travians and Schedorhinotermes medio-obscurus of the Rhinotermitidae, an unidentified species of the Subulitermes complex and Procapritermes augustignathus of Termitidae. Discussion The distribution and infestation levels of termites in the SBG Rain Forest, as in other ecosystems, are dependent on the availability of their food sources. These include the wood and leaf litter on the forest floor, the 180 Gard. Bull. Singapore 50(2) (1998) standing dead trees and the dead branches of live trees. The northern part of the forest has a higher population of live trees than the central and southern parts and larger amounts of ground and standing dead timber as evidenced by our quantitative assessments. Having greater amounts of food for termites explains why the northern part of the rain forest is significantly more heavily infested than the central and southern parts. In the central and southern parts, where herbaceous undergrowth thrives, there is a lower density of trees and saplings. The abundance of herbaceous plants, which are often accompanied by the presence of ants, has a negative relationship with the presence of termites. Ants have often been considered as enemies of termites (Eisner et al., 1976) and strong negative associations between ants and termites had been attributed to competition between the two groups (Fowler et al., 1980). The total of 102 series of 22 termite species collected within three months is high when compared with the 70 species collected over several years from the much larger Bukit Timah and Central Catchment Nature Reserves (Murphy, 1997). Due to the short time frame of the study, the collection though fairly intensive is by no means exhaustive. There was probably under-collection of the fungus-growing Macrotermitinae, which reportedly consumed over 75% of total organic matter taken by termites in Peninsular Malaysian rain forests (Collins et al., 1983) and 95% of leaf litter in Nigerian savanna (Collins, 1981). Other than Macrotermitinae, the termite species inhabiting the soil and the macro-timber are probably also under-represented. According to the lists of termite species compiled for Singapore’s nature reserves (Murphy, 1997) and Malaysia’s Pasoh Forest Reserve (Jones & Brendell, 1998), the soil-inhabiting species and the wood- feeding Rhinotermitidae together constitute about 40% to 50% of all species recorded. Further studies need to focus on these soil and wood-feeding termites to cover the entire termite fauna in the rain forest. Of the species collected, three are of particular interest because they have not been reported by earlier collectors and are not in the most recent list of termites species reported from Singapore (Murphy, 1997). These are the Bulbitermes borneensis, B. constrictus and Microcerotermes crassus. The northern zone of the rain forest contains some species that are common in the actively regenerating secondary forests of the Central Catchment Nature Reserves (Murphy, 1997). These include the gallery- forming Microcerotermes and Nasutitermes, which are actively involved in the self-pruning of younger trees, and the wood-feeding Bulbitermes. However, unlike the nature reserves, the nest of Bulbitermes was not found. Murphy (1997) reported the widespread occurrence of Macrotermes and Odontotermes in the more established secondary forests and the primary forests. This holds true for both genera in the northern and central zones Termites in SBG Rain Forest 181 but not in the southern zone, which has very little ground timber. Separate studies have been initiated to check whether the fallen timber in the northern zone are attacked by Kalotermitidae (dry wood termites) including species of Cryptotermes, Glyptotermes and Neotermes, which reportedly had a high incidence of attack in the nature reserves’ secondary forests. Eggleton et al. (1995) found that species diversity of termites was greatly reduced in severely disturbed plots of forests, but increased marginally in slightly disturbed regenerating plots. The different termite species diversity of the three zones of the rain forest reflects to a certain extent the different degrees to which the forest has been disturbed. The study of the forest vegetation also reveals that different parts of the forests differ in their structure and regeneration. For instance, much of the northern zone of the forest has high forest species of Calophyllum and Garcinia (Guttiferae) and Syzygium (Myrtaceae), which are common in established secondary forests (Wong et al., 1994). Other than having a large gap caused by trees struck by lightning, and an adjacent patch with a large number of small trees and rapid turnover of saplings, the northern zone is not much disturbed in most parts. This could account for its relatively high termite diversity. In contrast, the central and southern parts of the forest which have lower mature tree populations but contain more specimens of the larger sized trees e.g. dipterocarps, are much more disturbed. Here the faunal diversity, forest structure and tree regeneration are clearly affected by the rampant growth of climbers, dense undergrowth of herbs and ornamentals, and large spaces created by paved roads criss-crossing the forest, the encroachment of buildings and horticultural facilities and the proximity of the forest to public roads. The three zones were markedly different in the amount of space utilised for human activities. Assuming that plots of radius 10 m are drawn around each of the 15 or so transect points within each zone, the northern zone would have 7 plots trespassed mainly by 1.3m wide paths; whereas the central zone would have 9 plots trespassed by 2 m, 4 m and 6 m wide roads, while the southern zone would have 12 out of 14 plots depleted of floor space by 2 m, 4 m, 5.5 m and 6 m wide roads, as well as by a gazebo and potting yard. In conclusion, the reduction of space depletion of this rain forest and removal of the herbaceous undergrowth and the tree-smothering climbers will do much to minimise the loss of its termite species. Acknowledgements We are most grateful to Professor D. H. Murphy for his guidance in the project and particularly in his assistance in identifying the termites found. 182 Gard. Bull. Singapore 50(2) (1998) We also wish to thank the Chief Executive Officer of National Parks Board and the Director of Singapore Botanic Gardens for their encouragement, to the staff of SBG Herbarium for plant identification and Nature Conservation Branch especially Chew Ping Ting and Cheryl Chia for supplying the map of SBG Rain Forest and for giving information about the Biological Survey of Singapore’s nature reserves. We would also like to thank the staff of SBG Management in providing assistance with some of the field work. References Collins, N. M. 1981. The role of termites in the decomposition of wood and leaf litter in the Southern Guinea savanna of Nigeria. Oecologia. 51: 389-399 Collins, N. M., S. L. Sutton, T. C. Whitmore, and A. C. Chadwick. 1983. Termite populations and their role in litter removal in Malaysian rain forests. In: S. L. Sutton, T. C. Whitmore, and A. C. Chadwick. (eds.) Tropical Rain Forest: Ecology and Management, Blackwell, U.K. pp. 311-325. Corleti.. R..-T;. 1992.) The ecological transformation of Singapore, 1819- 1990. Journal of Biogeography. 19: 411-420. Corner, E. J. H. 1935. The seasonal fruiting of agarics in Malaya. Gardens’ Bulletin, Straits Settlements. 9: 79-88. Eggleton, P., D. E. Bignell, W .A. Sands, B. Waite, T. G. Wood and J. H. Lawton. 1995. The species richness of termiites (Isoptera) under differing levels of forest disturbance in the Mbalmayo Forest Reserve, southern Cameroon. Journal of Tropical Ecology. 11: 85-98. Eisner, T., I. Kriston and D. J. Aneshansley. 1976. Defensive behaviour of a termite (Nasutitermes exitiosus). Behavioural Ecology and Sociobiology. 1: 83-125. Fowler, H. G., B. L. Haines and P. Jaisson. 1982. Species diversity in leaf- cutting ants and mound-building termites in relation to the succession of vegetation in Paraguayan grasslands. In: Jaisson, P. (ed). Social Insects in the Tropics. pp. 187-201. Holmgren, N. 1913. Termitenstudien 4. Versuch einer systematishen Monographie der Termiten der orientalischen Region. Kungliga Svenska vetenskapsakademiens handlingar. 50: 1-276 Termites in SBG Rain Forest 183 Jones, D. T. and M. J. D. Brendell. 1998. The Termite (Insecta: Isoptera) Fauna of Pasoh Forest Reserve, Malaysia. The Raffles Bulletin of Zoology. 46: 79-91. Murphy, D. H. 1973. Animals in the forest ecosystem. In: Chuang, S. H., (ed). Animal Life and Nature in Singapore. Singapore University Press. pp. 53-73. Murphy, D. H. 1997. Faunistic Survey Part 2 — The Isoptera. Unpublished technical report of a survey commissioned by the National Parks Board, Singapore. Snedicor, G. W. and W. G. Cochran. 1969. Statistical Methods. The lowa State University Press. U.S.A. Thapa, R. S. 1977. Termites of Sabah. Sabah Forest Records. 12: 1-374 Tho, Y. P. 1992. Termites of Peninsular Malaysia. Malayan Forest Records.No. 36. Forest Research Institute Malaysia, Kepong. Malaysia. Morden. i M.. hs. Chua, JS. Y. Ong, B.C. Soong and H. T. W. Tan. 1996. A century of plant species loss from an isolated fragment of lowland tropical rain forest. Conservation Biology. 10: 1229-1244. Wong, Y. K.; P. T. Chew and Ali Ibrahim. 1994. The tree communities of the Central Catchment Nature Reserves. Gardens’ Bulletin, Singapore. 46: 37-78. cal co A iv Oh ed P e. YF neh ‘ vm | it. % : ¥ PD ih f Na ay ve A r Fi me if Kv ee. a M, ” s ; OP we aT Wg ey > aes Ys pe Ghd hed Wea pensar F aye eae Sa RAM NS ta ; 14 F : ‘ i a rn j rm) Ve hee fn « Baye i 7 ' { ‘ it aT, ye ‘ > . ; u is ath FIP a ATA BOT dP .s ’ are A NM 1 Phat te ae iil i , PP pth i" 7 | dace Maa \ ae i ; yh f ' | r] 7 ie, ~ e t rs baled gt e f att ty ; Mi " 1 6 4 “ sa Al . oe -_ : Le vee A | f p é j F 5 S , } 1 7 y * ‘ A Jj . t .f 5 | lt i! ad q at ‘ ms | a Sth hi A fi : ey ¥ : ? ® 4 a : 1a) oe i c a ialic 2 cat 5 J oF: Pe AM ‘ aky % é ¢ 4) } F ‘ ' }. ( - hdbe aie ‘ > Tr . : : i Ta 4 fr at f i o , v3, A i" wk 1 citvins Hs vanes { 7s ’ ‘ Lee se Sy hea, ne ‘) iat oN aus i" he ™* 4 tie Pie drt the eae p iN tal y i oat) par. = GEE RINE OED PSTN a By eae ad ‘ y . : ye! rae OU LA, Lavy i Pht - a ¢ ; 7" } hag \ ffs , ] 7 ; rit : ies ae - E's ; 4) ee | Na, ys es ok rr, ae Nit - ; we) Ad v/ ,) i 7 7 ily), r ; Wi a tied ae - uf 4 ie } : Aah aoa, a oe, 7 bag 17.9 ‘art uw Ss jo. Ke Ppa Laser he ys hig eS es ieee wee Gardens’ Bulletin Singapore 50 (1998) 155-196. The Unique Elements of the Limestone Flora of Batu Tengar Cave (Segarong), Sabah, Malaysia. RUTH KIEW Singapore Botanic Gardens, Cluny Road, Singapore 259569 Abstract Unique elements of the flora of the Batu Tengar Cave include Begonia keithii (Begoniaceae), a new species endemic to this hill, and the phytogeographic affinities of its flora, which are not only with the nearby Madai limestone hills, but also with limestone in Kalimantan (Borneo) and the Philippines. Notes on species of special interest, Euphorbia lacei (Euphorbiaceae), Jmpatiens winkleri (Balsaminaceae) and Paraboea madaiensis (Gesneriaceae) are given. Introduction Batu Tengar Cave (4° 42’ 00”N 118° 00’ 30”E) is located within the Segarong Protected Forest Reserve (Lim & Kiew, 1997), adjacent to the Segarong River. Commercial quantities of bird nests are still collected from the cave. The earliest botanical collections from this locality (called ‘Segarong’ on herbarium labels) date from 1938 when Symington with Agama and then Keith collected a handful of plants. These few collections are, however, extremely interesting as they included the first collection of a balsam from limestone in Sabah, the succulent Euphorbia lacei (Euphorbiaceae) and a begonia with decorative leaves. In May 1997, as part of the botanical survey for “The Biodiversity and Conservation of the Limestone Flora of Sabah Project’, this hill was revisited to relocate these species and to make a general collection of fertile material (Kiew, in press). The hill is now a 25-minute journey from Semporna by speedboat to the jetty built by the birdnest concessionaire on the Segarong River. It is then a short walk through cultivated land to the hill base that is surrounded by a narrow strip of forest. The summit of the hill (about 200 m a:.s.l.) closest to the river shows signs of burning being bare of vegetation apart from dead tree trunks that are still standing (Fig. 1). However, the way to the cave on a well-beaten track is through primary forest that provides the damp shaded conditions that support lush vegetation. Once the cliff base is reached, the path ascends a narrow gully, where on the large scattered limestone boulders, /mpatiens winkleri (Balsaminaceae) and Epithema 186 Gard. Bull. Singapore 50(2) (1998) dolichopodum Hillier & Burtt (Gesneriaceae) are encountered in abundance. As the path ascends, it becomes steeper and the canopy is more open and here Begonia keithii (the new species described below) is first encountered. Higher up on the exposed shoulders of the summit it is abundant. It is an extremely decorative species with glossy cardinal red stems that look lacquered, narrowly scolloped leaves and bright red male flower buds. The summit vegetation is remarkable being completely dominated by a cactus-like forest of Euphorbia lacei with a candelabrum-like crown (Fig. 2). This is unique to Batu Tengar Cave as no other limestone hill in Sabah has this type of forest on the summit. On the sheer cliffs, Paraboea madaiensis (Gesneriaceae) is abundant. Phytogeographic Affinities of the Flora Apart from the begonia, which is endemic to Batu Tengar Cave itself, two species (Epithema dolichopodum and Paraboea madaiensis) are endemic to Sabah, while Jmpatiens winkleri and Euphorbia lacei, while not endemic, are rare in Sabah being known from a few other localities. Impatiens winkleri was first described from south Kalimantan and in Sabah is known from just four other limestone localities all in the south east (Fig. 3). Discounting common species that are everywhere widespread, it is the only rare species from Sabah limestone that is known to occur in both Sabah and Kalimantan. Euphorbia lacei is rare in Sabah being known from just three localities (all close to Semporna, but the other two are not limestone). However, elsewhere it 1s a widespread species ranging from Myanmar, though Thailand and IndoChina to the Philippines where it reaches Palawan. Three of these species are also known from Gunung Madai. Paraboea madaiensis is only known from Gunung Madai and Batu Tengar Cave, Impatiens winkleri is known also from Baturong, Batu Belas and Tempadong, while Epithema dolichopodum is known from six other localities, which are all located in the southeast, Batu Tengar Cave, Baturong, G. Madai and Tempadong (Fig. 3), except for one population on limestone on Pulau Balambangan and one collected by Elmer (E/mer 20569) from Tawau but without a specific locality. Limestone Flora of Batu Tengar Cave 187 Figure 1. Batu Tengar Cave, Segarong Protected Forest Reserve. (Burnt summit visible to right) Figure 2. Euphorbia lacei Craib on the summit of Batu Tengar Cave. 188 Gard. Bull. Singapore 50(2) (1998) Segama River ry O SO? ths Figure 3. Distribution in Sabah of rare species from Batu Tengar Cave.. (@ localities; ® Euphorbia lacei; 0 Impatiens winkleri (also from Kalimantan limestone), + Paraboea madaiensis; & Epithema dolichopodum (endemic to Sabah, also from Pulau Balambangan and Tawau). Conservation Status of Batu Tengar Cave Recently, the hill was proposed as a commercial source of limestone (Robert C. Ong, pers. comm.). In view of its endemic begonia species, the four rare species (Fig. 3), two of which are endemic to Sabah, and the remarkable ‘cactus forest’ of Euphorbia lacei, the like of which is not known from any other limestone hill, and the possibility of other new and endemic species becoming known as research on the collections from the hill continues (e.g. the Cyrtandra RK4319 (Gesneriaceae) is possibly a new species), Batu Tengar Cave is one of the eight most important limestone sites (Kiew, in press) from the point of view of the conservation of plant biodiversity and it is recommended for permanent legal protection. Protection of the vegetation will also help to maintain conditions suitable for the cave swiftlets to nest which, if harvested sustainably, are a renewable economic resource for local people. Limestone Flora of Batu Tengar Cave 189 Notes on Species of Special Interest 1. Begonia keithii Kiew sp. nov. (Begoniaceae) Typus: R. Kiew, S. Anthony & S.P. Lim RK4327 Batu Tengar Cave, Semporna F.R., Sabah (holo SING, iso K, KEP, L, SAN, SAR). Figure 4. A Begonia amphioxus foliis non peltatis tepalis feminibus 5 liberi et ovari triloculari differt. Erect bushy, cane-like, monoecious, glabrous begonia forming clumps, producing short slender branches along the length of the stem. Stems woody, up to 1.5 m tall and c. 1 cm thick at base, shiny, deep crimson becoming brown at base, older nodes swollen with a conspicuous leaf scar. Stipules pale green suffused red, broadly elliptic, 10-18 by 8-9 mm, margin entire, apex setose, early caducous. Leaves alternate. Petiole slender, crimson, 3-7(-15) mm long. Lamina mid- to dark green above, sometimes mottled silver-grey, usually green beneath but sometimes suffused crimson, margin crimson, succulent and brittle, markedly unequal sided with the narrow side scarcely developed, narrowly lanceolate, 7.5 by 2.5 cm to 10 by 1.5 cm, attenuating to acute apex, main vein of base sometimes almost in line with the midrib but frequently at a 135° angle to midrib, base very variable in shape ranging from attenuated to acute point (matching the apex) and up to 6 cm long or bluntly rounded to truncate and 1.25-—2.5 cm long or base scarcely developed and 0.75 cm long, margin scolloped becoming distantly serrate towards apex and base, main vein reaching to apex with a pair running almost parallel, the other lateral veins radiating into the basal lobe. Leaves decreasing markedly in size towards the stem apex. Female flowers solitary from lower axils, up to 3 produced before the male inflorescences develop. Pedicel pale reddish, 10-12 mm long; bracteoles absent; ovary pale greenish with wings suffused red towards margin, 3-loculate, ovoid 9-17 mm long and 7-8 mm wide narrowing to 1.5 mm below style, wings 3, equal, placentas axile, bilamellate with many ovules on both surfaces; tepals 5, free, whitish suffused reddish; glabrous, oval, 7-8 mm long, margin entire, apex rounded, outer tepals 5-6 mm wide, inner 2.5-3.5 mm wide, style and stigma pale yellow green, styles 3, 1-2.5 mm long, branching from base, spreading and bifid with a continuous twisted papillate stigmatic band. Male flowers produced on erect, many- flowered, twice-branched cymose inflorescences from the upper axils, 4.25-9.5 cm long; bracts not persisting; peduncle carmen, 2—4.25 cm long, branches thread-like and slightly zigzag; bracteoles reddish, lanceolate 190 Gard. Bull. Singapore 50(2) (1998) Figure 4. Begonia keithii Kiew. A Habit (x 0.4), B Male flower bud (x 1.2), C Male flower (x 1.2), D. Stamen (x 8), E Androecium (x 4), F Terminal flowers of male inflorescence (x 0.8), G Male inflorescence (x 0.4), H Female flower (front view) ( x 1.2), I Styles (x 4), J Female flower, side view (x 0.8), K Capsule (x 0.8), L T.S. capsule (x 1.6). Limestone Flora of Batu Tengar Cave 19] 1-3 mm long, apex acuminate, soon falling; male buds small, c. 4 mm long and carmen outside; pedicels carmen, slender, 1-5 mm long; male flowers with 4 glabrous tepals, outer two with inner surface scintillating white with carmen showing through, almost rotund, 5-7 by 4-7 mm, inner two completely white, narrowly lanceolate, 2-5 by 1-2 mm, apex rounded, margin entire; stamens c. 55-65, pale yellow green (matching exactly the colour of stigmas), forming a spherical cluster on columnar torus 1.5—1.7 mm long, filament c. 0.5 mm long, anther obovoid, 0.5-0.7 by 0.5 mm, apex deeply emarginate. Capsule dangling on slender thread-like stalk 2-3 cm — long, ovoid and narrowed to pedicel, 18—25 by 15—25 mm; locules 3, c. 14 mm long, not reaching to pedicel or apex; wings 3, isomorphic, broader distally and 8-12 mm wide, wing tip slightly rounded or sometimes acute, becoming dry and papery and dehiscing along the junction with locule, styles caducous. Seeds broadly ovoid, c. 0.35 by 0.2 mm, testa strongly reticulate. Distribution: Endemic to Batu Tengar Cave (Segarong Cave), Semporna Protected Forest Reserve, Sabah, Borneo. Habitat: In light shade to full sun, growing in rock crevices on the tower karst limestone hill, dominating the exposed shoulders of hill where it forms thickets with its woody cane-like stems. Specimens examined: Symington & Agama 9315 20 July 1938 (K, SING), Keith A9416 26 Aug 1938 (K, SING), Kiew et al. RK4327 9 May 1997 (K, KEP, L, SAN, SAR, SING). Notes: A most decorative begonia, it has polished crimson stems that appear lacquered, dainty scolloped leaves with a crimson margin and the many tiny, carmen, heart-shaped buds of the male flowers set on slender sprays. It has yet to be found on other limestone hills in Sabah or elsewhere. In possessing an erect, bushy habit, female flowers with 5 petals, 3 bifid styles and a 3-loculate fruit with 3 equal wings, it conforms to Section Petermannia, except that its male flowers have 4 tepals. (Section Petermannia is characterised by 2 tepals). In its cane-like habit, narrow leaves with the pointed apices and serrate margin, solitary female flowers, male flower with 4 tepals, it most resembles B. amphioxus Sands. Sands (1990) had already noted this similarity remarking that the two species ‘may be at least very closely allied’. The two species are, however, readily distinguished by the suite of characters listed in Table 1. This begonia is named for H.G. Keith, in 1925 Assistant Conservator of Forests in the then British North Borneo, rising in 1931 to Conservator. 192 Gard. Bull. Singapore 50(2) (1998) He survived internment during the war returning to Sandakan in 1946. It was he who in 1938 collected from Batu Tengar Cave this begonia, the first collection of a balsam from Sabah limestone and the Euphorbia. Table 1. Characters that distinguish Begonia keithii from B. amphioxus. Character keithii amphioxus Max. stem length (cm) 150 75 Stem colour glossy crimson green Petiole length (mm) 3-7(-15) (10—)15-55(-60) Petiole colour crimson light green Leaf not peltate peltate Leaf pattern unpatterned or grey mottled red spotted Tepals female flower 5; free 3-4(—5), joined No. styles 3 2(-3) No. locules 3 2 Capsule shape ovoid columnar — Capsule size (mm) (18—)20-25 x 15-25 (9-)10-13 x (5—)6-7 Length capsule stalk (mm) 20-30 2(-2.2) Wings 3, equal 2(—-3), third shorter Length male inflorescence (cm) 4.25-9.5 up to 3 Colour of male buds carmen white 2. Euphorbia lacei Craib (Euphorbiaceae) Shaw (1975) listed this Euphorbia as one of the seven species of Bornean Euphorbiaceae that are confined to limestone. He also expressed uncertainly as to its identity recording it as E. sp. cf. lacei. Now that more specimens are available and field observations could be made, its identity is confirmed as typical E. lacei, which differs from the other Malesian limestone succulent Euphorbia, E. antiquorum L. in its longer spines, more distant spine shields with deep sinuses between them. Euphorbia lacei is indeed a remarkable plant, which on the summit of Batu Tengar forms a “cactus” forest, unique for limestone in Malaysia. It grows to 4-5 m tall with a stem diameter of 4.5—5 cm, the lower trunk is bare but the upper branches curve outward giving the plant a candelabrum- like appearance (Fig. 2). However, in Sabah it is not confined to limestone. One of the localities quoted by Shaw is Mt Sidungol (correctly spelt Sirongol), which is one of the rocky peaks of Timbun Mata Island, which is of volcanic conglomerate. Staff of the Forest Department in Semporna have also collected it from the summit of Bohey Dulang Island (also of volcanic conglomerate) and are successfully growing it as a pot plant at their office. Limestone Flora of Batu Tengar Cave 193 In Borneo, E. lacei is presently known only from these three dry rocky summits close to the coast near Semporna (Fig. 3). Outside Malaysia it is widespread. It is recorded from Myanmar and Laos, from limestone in southern Thailand and also from the Philippines. Merrill (1923) recorded it (under the name E. trigona Roxb.) from Luzon, Mindoro and Palawan describing its habitat as ‘in thickets and on limestone cliffs, usually along the seashore ascending to 300 m’. The local names that Merrill cites (suda- suda and tuba) are the same as Keith recorded in the Kedayan language — tuba suduh (Keith 9415). E. antiquorum in Peninsular Malaysia is also called sudu-sudu in Malay on account of its leaves that are spoon-shaped. The name tuba suggests that this species may be used as fish poison. Specimens examined: Semporna District: Batu Tengar Cave Keith A9415 26 Aug 1938 (K), Kiew et al. s.n. 9 May 1997 (SING). Mount Sidungol Keith A9337 18 July 1938 (K, KEP). 3. Impatiens winkleri Hook f. (Balsaminaceae) Impatiens winkleri was first collected from Sabah by Keith in 1938 from Segarong F.R. Our survey confirms that it grows on limestone on Batu Tengar Cave and also that it is more widespread growing on several limestone hills in S.E. Sabah, namely Bukit Baturong, Gunung Madai, Batu Belas and Tempadong (Fig. 3). It was originally collected from south Kalimantan by Hubert Winkler from Batu Babi but he did not record whether from limestone. More recently it was recollected from south Kalimantan from limestone on Gunung Serempaka. It therefore appears to be confined to limestone. It has not been collected from Sarawak. Its habitat is typical of limestone balsams (Kiew, 1991) in that it grows only on limestone boulders and ledges close to the cliff base in particularly damp and deeply shaded conditions. Where conditions are suitable, e.g. in narrow gullies, it forms thickets. As forest surrounding the base of the limestone hills is cleared for agriculture, this habitat is particularly vulnerable to exposure to light and drying out, which will endanger the continued existence of this species. Already the undergrowth around the base of the hills at G. Madai and Batu Tengar Cave is disturbed by birdnest collecting activities. Impatiens winkleri is a giant balsam up to 2 m tall. The fleshy dark green trunk ranges from 4.5 to 6 cm in diameter at the base and bifurcates or trifurcates at about 1.5 m to product a spreading crown of succulent branches. As the branches grow, they bend under their weight and many erect branches are then produced from the upper side of this horizontal branch (Rauf’s tree architecture model). The thick fleshy branches are brittle and frequently break off to be replaced by many, more slender stems. 194 Gard. Bull. Singapore 50(2) (1998) This life form is not shown on herbarium specimens as only the terminal portion of the branch with its bunch of leaves and flowers fits onto the sheet, misleadingly giving the impression it is a small herbaceous species. Indeed, Hooker (1910) described the stature as ‘humilis’, i.e. low growing, and the stem as at least 1 m tall and 2-3 cm thick. Shimizu (1970) identified specimens as J. scortechinii Hook. f., a herbaceous species from Peninsular Malaysia that grows to about one meter tall. Among all the limestone balsams in Malaysia, [. winkleri in its tall ‘trunked’ habit most resembles J. mirabilis Hook. f. from limestone in the extreme north of Peninsular Malaysia and S. Thailand. However, it differs from I. mirabilis as its trunk base is not swollen (J. mirabilis is called the ‘gouty balsam’ because of its swollen base). Its habit, broad leaves and white flowers make J. winkleri distinct from all other Bornean balsams. Limestone balsams in Peninsular Malaysia, are very biodiverse with about ten species, of which six are endemic to Peninsular Malaysia, three ~ are endemic to the region spanning northern Malaysia and peninsular Thailand, and one is shared with Sumatra. In Sarawak, there are at least four species (most still unnamed) collected from the Bau area and the Mulu National Park and at least three of these are endemic to Sarawak. However, in Sabah there is just J/mpatiens winkleri, which also occurs in S. Kalimantan. This mirrors a pattern seen in several other genera of herbs, e.g. Chirita, Monophyllaea and Paraboea in the Gesneriaceae, where biodiversity decreases from Peninsular Malaysia, to Sarawak, to Sabah where few or no species are found (Kiew, in press). Specimens examined: Borneo: South Kalimantan — Batu Babi et Lumnovia H. Winkler 2866 (type, drawing at K), G. Serempaka J. Dransfield JD2318 (L); Sabah — Batu Tengar Cave Keith A9329 20 July 1938 (K), R. Kiew et al. RK4327 (SAN, SING), Baturong Lim S.P. et al. LSP713 12 June 1996 (SAN, SING), R. Kiew RK s.n. 12 June 1996 (SING); G. Madai Mansus et al. SAN 117117 (K, SAN), Lim S.P. et al. LSP671 9 June 1996 (SAN, SING), Batu Belas, Segama River J.H. Beaman et al. 10131 13 June 1984 (K), Tempadong, Segama River J.H. Beaman et al. 10091 (K). 4. Paraboea madaiensis Xu & Burtt (Gesneriaceae) Previously recorded only from Gunung Madai (Xu & Burtt, 1991), it is abundant on the exposed summit and cliffs of Batu Tengar Cave. Our local guides collected plants for medicine. However, since a sizeable population grows on inaccessible sheer cliffs, it is unlikely that harvesting this species for personal use will endanger the population. Limestone Flora of Batu Tengar Cave 195 Our informants (a local villager, a birdnest collector and a forest guard) used it in several remedies: 1. Used fresh, the leaves are put in hot water and the vapour is inhaled to reduce fever in adults, The smoke from burning leaves is used to: 2. Quieten crying babies, and 3. In adults, to clear the eyes if they are cloudy. 4. The plant can be used after childbirth but our informants were unable to give precise information on how it is used. Ackowledgements The author is extremely grateful to the Ministry of Science, Technology and the Environment, Malaysia, for funding the collaborative project on ‘The Biodiversity and Conservation of the Limestone Flora of Sabah’ between Universiti Putra Malaysia and Forest Research Centre, Sabah Forest Department, under IRPA research grant 52858; to Wong Khoon Meng and Robert C. Ong for their strong support, to S. Anthonysamy, Lim S.P. and the staff at SAN for assistance in the field and herbarium, to Mdm P.H. Yap for the illustration of the begonia, to Pius Primus District Forest Officer, Semporna, who made available the speedboat and accommodation, and to the Curators of the herbaria at K, KEP, L and SAN for permission to examine specimens in their care. References Hooker, J.D. 1910. XIII Decades Kewensis Plantarum Novarum. Kew Bulletin. p. 75. Lim, S.P. & R. Kiew. 1997. Gazetteer Of limestone localities in Sabah, Borneo. Gardens’ Bulletin Singapore. 49: 111-118. Kiew, R. 1991. The limestone flora. In: R. Kiew (ed.). The State of Nature Conservation in Malaysia. Malayan Nature Society, Kuala Lumpur, Malaysia. pp. 42-50. Kiew, R. (in press) Towards a limestone flora of Sabah. In: K.M. Wong (ed.) Proceedings of Stone Memorial Symposium. Malaysian Nature Society & Rimba IImu, University of Malaya, Kuala Lumpur, Malaysia. Merrill, E. D. 1923. Enumeration of Philippine Plants. p. 464. 196 Gard. Bull. Singapore 50(2) (1998) Sands, M.J.S. 1990. Six new begonias from Sabah. The Kew Magazine. 7: 57-85. Shaw, H.K.A. 1975. The Euphorbiaceae of Borneo. Kew Bulletin. Additional Series. 4: 111, 223-224. Shimizu, T. 1970. Contributions to the Flora of Southeast Asia. II. Impatiens of Thailand and Malaya. South East Asian Studies. 8: 206. Xu, Z. & B.L. Burtt. 1991. Towards a revision of Paraboea (Gesneriaceae). Edinburgh Journal Botany. 48: 4. Gardens’ Bulletin Singapore 50 (1998) 197-219. New Species and Varieties of Moraceae from Malaysia K. M. KOCHUMMEN Forest Research Institute of Malaysia, Kepong 52100 Kuala Lumpur, Malaysia Abstract Three new species and one new variety of Artocarpus and fourteen new species of Ficus and seven new varieties are described. All the new species and varieties are from Sabah and Sarawak except F. ngii, which is from Peninsular Malaysia. The new species and varieties are Artocarpus corneri, A. jarrettiae, A. primacki, A. anisophyllus var. sessilifolius, Ficus ashtonii, F. borneensis, F. chaii, F. chewii, F. corneri, F. dulitensis, F. gamostyla, F. ilias-paiei, F. kerangasensis, F. longistipulata, F. ngii, F. pseudotarennifolia, F. sabahana, F. soepadmoi, F. cereicarpa var. ashtonii, F. deltoidea var. recurvata, var. subhirsuta, F. obscura var. lanata, F. oleifolia var. calcicola, and var. impressicostata, and F. sundaica var. impressicostata. Descriptions of the new taxa are provided. Introduction In her revision of the genus Artocarpus of the Malesian region, Jarrett (1959, 1960, 1975) recognised 19 species from Sabah and Sarawak. Corner (manuscript 1972) recorded 20 species. In his checklist of Ficus in Asia and Australasia and in additional publications, Corner (1965, 1970, 1972) recognised 128 species of Ficus for Sabah and Sarawak. Completion of the study of these two genera in Sabah and Sarawak has added three new species and one new variety of Artocarpus and thirteen new species and seven new varieties of Ficus making a total of 23 species of Artocarpus and 141 species of Ficus known in Sabah and Sarawak. Of the newly described species of Artocarpus, one is recorded only from Sarawak. Similarly of the thirteen new species of Ficus, four are found only in Sabah, seven only in Sarawak, and two in Sabah and Sarawak. Description of New Taxa 1. Artocarpus corneri Kochummen sp. nov. (E.J.H. Corner, 1906-1996, prominent Professor of Tropical Botany, University of Cambridge, UK, who undertook extensive studies of the Malesian Moraceae) Artocarpo lanceifolio simillimus, sed in foliis in sicco badiis supra scabridis, marginibus integris, syncarpiis cylindricis differt. Typus : Dayang Awa & 198 Gard. Bull. Singapore 50(2) (1998) Yii P.C. S 46878, Borneo, Sarawak, Dulit Range, Belaga (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Tree to 22 m tall, 50 cm diameter. Twigs 7-9 mm thick, dark brown, closely ridged, very sparsely short-hairy, with distinct ring-like stipular scars. Stipules lanceolate, to 3 cm long, densely covered with long hairs. Leaves obovate or oblong, (12—)17—28 x (5.5—)7-12 cm, base cuneate, margin faintly wavy, apex acuminate or acute; both surfaces glabrous to the naked eye but with short rough hairs on the midrib and lateral veins, drying reddish brown to chocolate brown, upper surface sandpapery to touch; midrib raised above; lateral veins 12-15 pairs, raised below, faint above; intercostal veins scalariform, faintly visible below, invisible above; petioles 3-5 cm long, with short sparse hairs. Inflorescences axillary, solitary. Male not seen. Syncarp (immature) green when fresh, cylindric, 5-7 x 3.5-4 cm; tepal densely hairy, style distinctly bifid; peduncles 5.5—8 cm long. Vernacular name: Sarawak: talun (Murut). Distribution: Endemic to Borneo, rare, known only from Sarawak. Ecology: Submontane forest between 820-1000 m altitude. Notes: Closely related to A. lanceifolius in section Duricarpus of subgenus Artocarpus but differing in the reddish brown dry leaves with scabrid upper surface, entire leaf margins and in the cylindric syncarps. Specimens Examined: BORNEO. SARAWAK: Lawas, Maligan Range, Ilias Paie S 32879 (BO, CGE, K, KEP!, L, SAR!); Belaga, Dulit Range, Ulu Sg. Kayan, Dayang. Awa & Yii P.C. S 46878 (CGE, K, KEP!, L, SAN!, SAR!). 2. Artocarpus jarrettiae Kochummen sp. nov. (J. Francis Jarrett who revised the genus Artocarpus for the Malesian region) Prope Artocarpum rigidum, in syncarpio cylindrico, perianthio pilis longis glandulosis basi inflatis differt. Typus : Amin & Francis SAN 120933, Borneo, Sabah, Ranau (holotypus SAN; isotypi K, KEP, L). Small tree to 15 m tall, 50 cm diameter. Twigs 6-7 mm thick, dark brown, closely ridged. Stipules lanceolate, up to 5.5 cm long, with reflexed edges, densely long hairy outside. Leaves elliptic or oblong, 16.5—23 x 8.5-12 cm, base cuneate, apex obtuse or acute; scabrid on both surfaces, upper sparsely New Species and Varieties of Moraceae 199 and lower densely hairy; midrib raised above; lateral veins 11-13 pairs, raised below, flat or faintly sunken above; intercostal veins scalariform, visible below, faint above; petioles 3-5 cm long, covered with short hairs. Inflorescences axillary. Male not seen. Syncarp (immature) yellowish green when fresh, cylindrical, c. 4.5 x 3 cm; perianth covered with glandular hairs with swollen bases; style exserted, simple; peduncles 4.5—5.5 cm long, rough hairy. Vernacular name: Sarawak: tekalong (Iban). Distribution: Endemic to Borneo, very rare, only known from Sabah and Sarawak. Ecology: Lowland forest by streams. Notes: Near to A. rigidus in section Duricarpus of subgenus Artocarpus but differing in the cylindric syncarp and in the perianth with long glandular hairs with swollen bases. Specimens Examined: BORNEO. SABAH : Ranau, Langanan, Amin & Francis SAN 120933 (K, KEP!, L, SAN!). SARAWAK : Sarikei, km 6, Bernard Lee S 54906 (CGE, K, KEP!, L, SAN!, SAR!). 3. Artocarpus primackii Kochummen sp. nov. (Richard B. Primack of Boston University, U.S.A., author of Foresters’ Guide to the Moraceae of Sarawak) Artocarpo glauco simillimus in sectione Pseudojaca, in capitulo masculo globoso, perianthio praeter apicem lobatum connato differt. Typus : Aban Gibot SAN 99596, Borneo, Sabah, Kota Merudu (holotypus SAN, isotypus KEP). Small to medium-sized tree, rarely to 33 m tall and 150 cm diameter. Bole with tall buttresses to 3.3 m high; bark grey brown or orange brown, smooth to cracking. Twigs 5-7 mm thick, covered with short rough hairs. Stipules ovate, c. 4 mm long, hairy outside. Leaves oblong or obovate, 12-33 x 7.5-19 cm, base cuneate or rounded, apex with 1 cm long sharp tip; upper surface glabrous except midrib, lower surface rough hairy; midrib flattened above; lateral veins 10-16 pairs, prominently raised below, finely sunken above; intercostal veins scalariform, raised below, faint above, reticulation prominently raised below; petioles 2—5.5 cm long, short-hairy, often glaucous. Inflorescences solitary, axillary. Male head globose, c. 1.8 cm 200 Gard. Bull. Singapore 50(2) (1998) diameter, sessile or with 5 mm long hairy peduncle; tepal 3-lobed, covered with short hairs; stamen one, exserted with stout filament; bracts numerous, with peltate heads. Female flowers with exserted 3-lobed styles. Syncarp sessile or shortly peduncled, subglobose, surface warty, c. 6 cm across, velvety hairy, pale yellow with pink flesh when fresh. Vernacular names: Sabah : beruni ; Sarawak : dadah (Iban). Distribution: Endemic to Borneo. Most collections are from Sabah with only one record from Sarawak. Ecology: Lowland and hill forest to 600 m altitude. Notes: Closely allied to A. glaucus in series Peltatae of section Pseudojaca in subgenus Pseudogaea, but differing by its globose male head and united perianth except for the lobed apex. It also differs from A. tomentosulus, (under which some of the collections mentioned below were included by Jarrett), by its rough hairy, large leaf with sunken veins, small ovate stipules, and in the sessile or shortly stalked syncarp. Specimens Examined: BORNEO. SABAH : Keningau, Sook Road, T. Oikawa SAN 92157 (SAN!); Kota Merudu, Aban Gibot SAN 99596 (KEP!, SAN!); Tenom, Agriculture Station, W. Meijer SAN 120622 (SAN!); Kinabalu National Park, S. Kokawa et al. 5223 (SAN!); Beaufort, Lumat Estate Reserve, Stephen Madius SAN 50064 (K, SAN!); Sandakan, Sepilok F.R., C. Charington SAN 21181 (K, L, SING!); Lahad Datu, Silam, Heya et al. SAN 61681 (K, KEP!, L, SAN!, SAR!, SING!); Lahad Datu, Silam F.R. Block 8, Agam Ambullah SAN 31491 (K, L, SAN!); Tawau, G.H.S Wood A 3684 (A, KEP!, L, SING!); Tawau, Tinagat F.R., J. Singh et al. SAN 48990 (KEP!, K, L, SAR!, SAN!); Tawau, Mile 9 Apas Road, F. Krispinus SAN 86649 (KEP!, SAN!, SAR!, SING!); SARAWAK : Mulu National Park, Melinau Gorge, R.B. Primack S 43309 (A, K, KEP!, L, SAN!, SAR!). 4. Artocarpus anisophyllus var. sessilifolius Kochummen var. nov. (Latin, sessilis=sessile, folius=leaved, i.e. the sessile leaflets) A varietate typica in ramulis laevibus foliolisque sessilibus differt. Typus : G.H.S. Wood SAN 16549, Borneo, Sabah, Sepilok F.R. (holotypus SAN). Vernacular name: Sabah : terap ikal (Malay). Distribution: Widely distributed in Sabah, but known only from a single New Species and Varieties of Moraceae 201 collection from Sarawak. Notes: This new variety differs from the typical variety by the smooth twigs and sessile leaflets. Specimens Examined: BORNEO. SABAH : Terintidon, Aban Gibot SAN 99508 (SAN!); Kota Merudu, Aban Gibot SAN 100079 (SAN!); Sg. Sapi, Beluran, Ag. Ahmad & Chiba SAN 124521 (SAN!); Kinabatangan, Austin Cuadra A 2133; (KEP!, SAN!, SING!); Sandakan, Sepilok F.R., G.HS. Wood SAN 16549 (A, BO, BRI, K, KEP!, L, SAN!, SING!); Sandakan, Labuk Road, Tamiji & Laurence SAN 47090 (SAN!). SARAWAK : Kuching, Hewitt 177 (SAR!). 5. Ficus ashtonii Kochummen sp. nov. (P.S. Ashton, sometime Forest Botanist in Brunei and Sarawak) Species prope Ficum hookerianum seriei Orthoneura, subgen. Urostigma, sed bracteis basalibus non connatis cupuliformibus sunt. In foliis crassis eis Fici stupendae similis, sed costa supra impressa. Typus : Dayang Awa & B. Lee S 47846, Borneo, Sarawak, Limbang (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Tree up to 28 m tall, 60 cm diameter. Twigs yellowish brown, irregularly ridged. Stipules ovate—lanceolate, pointed, c. 2.5 cm long, caducous. Leaves thickly leathery, elliptic, 14.5-17.5 x 6.5—9 cm, base broadly cuneate, apex pointed; midrib sunken above; lateral veins 6-7 pairs, curving and joining near margin, trinerved, basal pair reaching to more than '/3 of blade, raised below, faintly raised above; intercostal veins reticulate, distinct below, invisible above; petioles 2.2—-3.7 cm long, wrinkled on drying. Syconia from leafy twigs, axillary, orange turning deep red, sessile, oblong, c. 2 x 1 cm, apex flat with disc-like bracts; basal bracts large, with rounded apex, persistent. Male flowers with long stout stalk; tepal not distinct; stamen one with short filament. Female flowers sessile; perianths 3, lanceolate acuminate, up to the lower half of style; ovary elliptic, brown, slightly ridged, style lateral, long, stigma clavate. Gall flowers similar to female flowers, but with short pedicels. Distribution: Endemic to Borneo; known only from the Bario and Limbang districts in Sarawak. Ecology: Submontane forest at 1020 m altitude. Notes: A species near to F. hookeriana of Series Orthoneura in Subgen. 202 Gard. Bull. Singapore 50(2) (1998) Urostigma but the basal bracts are not united and cup-like. The thick leaves resemble those of F. stupenda but the midrib is impressed above. Specimens Examined: BORNEO. SARAWAK : Bario, Pa Ukat, 4" Division, Peter Sie S 35394 (A, CGE, K, L, MO, SAR!); Limbang, G. Pagon Periok, Dayang Awa & B. Lee S 47846 (CGE, K, KEP!, L, SAN!, SAR!). 6. Ficus borneensis Kochummen sp. nov. (Of Borneo) In subsectione Dictyoneuron prope Ficum delosyce, sed ficorum bracteis truncatis vel planis differt. Typus: W. Meijer et al. SAN 131862, Borneo, Sabah, Telupid, Bukit Tangkunan F.R. (holotypus KEP). Strangling fig. Young twigs yellowish, grooved. Stipules ovate—lanceolate, c. 15 x 5 mm, glabrous or hairy. Leaves leathery, drying chocolate brown; elliptic or oblong, 6.5—10.5 x 1.5—5 cm, base broadly cuneate, apex pointed; midrib sunken above, lateral veins 4-6 pairs, trinerved, basal pair reaching !/3 of blade, curving and joining to form a looped intramarginal vein, distinct below, invisible to faintly visible above, intercostal veins reticulate, visible below, invisible above; petioles 1-2:cm long, drying black. Syconia axillary, solitary or in pairs, ripening red, sessile, oblong, 10-12 x 7 mm, surface rugose and with distinct ridges, apex truncate with disc-like bracts; basal bracts triangular with acute tip, persistent. Male flowers stalked; tepals 3; stamen one, almost sessile. Gall flowers sessile; tepals 3; ovary reddish on one side, with short subterminal style. Female flowers shortly pedicelled; style lateral, stigma clavate; seed covered with mucilage. Distribution: Endemic to Borneo, very rare, recorded from Sabah and Sarawak. Ecology: Lowland to submontane forest to 1350 m altitude on ultrabasic soil. Notes: Somewhat like F. pellucido-punctata but the leaves are thicker and the figs are without perforation. Within the Subsection Dictyoneuron of section Conosycea of subgenus Urostigma , F. borneensis 1s near to F. delosyce but differs by its truncate apex and flat apical bracts of the figs. Specimens Examined: BORNEO. SABAH: Ranau, G. Mikil SAN 38549 (SAN!, SING!); Telupid, Bukit Tangkunan F.R., SAN 131862 (KEP!); Mount Kinabalu, J. & M.S. Clemens 29170 (SING!), 31275 (SING!). New Species and Varieties of Moraceae 203 SARAWAK : Kapit, Ulu Balleh, [lias Paie S 28556 (A, CGE, E, K, L, SAR!); G. Mulu National Park, Sg. Mentawi Paul P.K. Chai S 39749 (CGE, K, KEP!, L, SAN!, SAR!). 7. Ficus chaii Kochummen sp. nov. (Paul P.K. Chai, sometime Forest Botanist, Forest Department, Sarawak). Hac species prope Ficum ixoroidem sectionis Sycocarpus subgenus Ficus, sed in foltis distincte dentatis venarum lateralium paribus multis differt. Typus: Paul P.K. Chai S 36002, Borneo, Sarawak, Sg. Kapit (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Treelet to 1 m tall. Twigs reddish brown, angled. Stipules lanceolate, up to 10 mm long, caducous, finely hairy outside. Leaves narrowly oblong, drying to greenish yellow, 13.5—23.5 x 1.5—2.2 cm, base cuneate, apex pointed with 1-2 cm long tip, margin distantly toothed; midrib raised above; lateral veins 13-21, distinctly curving and joining near margin, trinerved, basal veins short, visible below, very faint above; intercostal veins reticulate, visible below only; petioles 1-1.5 cm long, yellowish on drying. Syconia from leaf axils, solitary or in pairs, pear-shaped, 5-8 x 3-4 mm, green ripening to orange; peduncles to 3 mm long; basal bracts tiny. Gall flowers shortly stalked; tepals 3, lanceolate, transparent; ovary globose, whitish, style short, terminal. Male flowers not seen. Distribution: Endemic to Borneo, very rare, known from a single collection from Sarawak. Ecology: Lowland forest by river side. Notes: This species is near to F. ixoroides of Section Sycocarpus Subgenus Ficus, differing in the distinctly toothed leaves and many pairs of lateral veins. Specimens Examined: BORNEO. SARAWAK. Kapit, Sg. Kapit, Paul P.K. Chai S 36002 (CGE, K, KEP!, L, SAN!, SAR!). 8. Ficus chewii Kochummen sp. nov. (W.L. Chew, formerly taxonomist, Singapore Botanic Gardens) Species as sectionem Conosycea subgenus Urostigma pertinet Ficus sumatrana similis, sed ficis oblongis apice truncato differt. Typus: Chew et al. RSNB 1972, Borneo, Sabah, Mount Kinabalu (holotypus SAN; isotypus SING). 204 Gard. Bull. Singapore 50(2) (1998) Strangling fig. Twigs greyish yellow, finely ridged. Stipules ovate—lanceolate, pointed, 15—17 mm long, hairy outside, caducous. Leaves thickly leathery, shiny above, elliptic or oblong, (2.7—)6—-11 x 3.5—5.5(-1.5) cm, base rounded or broadly cuneate, apex acute; midrib raised above with distinct central groove; lateral veins 6—8, with short intermediate veins, trinerved, basal veins reaching to middle of blade, curving and joining near margin, faintly visible on both surfaces; intercostal veins reticulate, faint below, invisible above; petioles 7-17 mm long, drying black. Syconia axillary, solitary or in pairs, sessile, oblong, c. 10 x 6 mm, apex truncate with disc-like apical bracts, surface irregularly wrinkled; basal bracts persistent, with rounded apex. Male flowers with 3 spathulate tepals; stamen one, sessile. Female flowers sessile; tepals 3; ovary with red markings, style long, subterminal, stigma clavate. Distribution: Endemic to Borneo; very rare, known only from Mount Kinabalu in Sabah. Ecology: Submontane forest at 1290 m altitude. Notes: F. chewii belongs to Section Conosycea of Subgenus Urostigma and is close to F. sumatrana but differs in the oblong figs with truncate apex. Specimens Examined: BORNEO. SABAH : Mount Kinabalu, Ulu Liwagu and Ulu Mesilau, W.L. Chew et al. RSNB 1972 (SAN!, SING!). 9. Ficus corneri Kochummen sp. nov. Species ad sectionem Leucogyne subgen. Urostigma pertinet foliis sine venis intercostalibus distinctis sed venis intermediis multis brevibus venis principalibus aequiliter prominentibus, ficis globosis sessilibus bracteis basalibus minutis absconditis, ovario albido notata. A Fico ngii in staminibus sessilibus periantho rubello marginibus albis differt. Typus: Fidilis & Asik SAN 119744, Borneo, Sabah, Keningau (holotypus KEP; isotypus SAN). Climber. Twigs yellowish brown, strongly grooved. Leaves yellowish green on drying, elliptic to oblong, 12—20 x 4—-6.5 cm, base cuneate, apex pointed, margin curled inwards; midrib raised above with distinct central sunken groove; lateral veins 5—7 pairs, with many short veins in between each pair, trinerved, basal pair reaching almost to half the length of blade, looping and joining near margin to form a looped intramarginal vein, visible below, very faint above; petioles 2.5-4 cm long, wrinkled on drying. Syconia yellowish when fresh, drying to reddish brown, from leaf axils, solitary or New Species and Varieties of Moraceae 205 in pairs, sessile, globose, 12-15 mm across, surface smooth, apex slightly sunken and closed by 2 bracts; basal bracts 3, small, concealed by base of syconium which is thickened and ring-like with white edge. Male flowers pedicelled, pedicels dark brown; tepals 4, dark brown, united; stamen one, sessile. Gall flowers similar to male flowers, ovary smooth, style lateral. Female flowers very few, sessile, with 4 narrow tepal lobes; style lateral; seeds smooth, subglobose. Distribution: Endemic to Borneo. Recorded from Sabah and Sarawak and also from Brunei. Ecology: Lowland and hill forests to 790 m altitude. Notes: A species belonging to Section Leucogyne of Subgen. Urostigma, it is characterised by its leaves, which are without distinct intercostal veins but with many short intermediate veins that are as equally prominent as are the main veins, by its globose sessile figs with small concealed basal bracts and by the whitish ovary. It differs from F. ngii in the sessile stamen and the reddish perianth with white edges. Specimens Examined: BORNEO. SABAH : Sipitang, Y.F. Lee & Dewol SAN 68975 (K, KEP!, L, SAN!, SAR!); Ranau, Tempurungon, Amin & Jarius SAN 115969 (KEP!, SAN!); Keningau, Lanas, Fidilis & Asik SAN 119744 (KEP!, SAN!). SARAWAK : Limbang, Bukit Pagon, Yahud Hj. Wat S 4763 (CGE, K, KEP!, L, SAN!, SAR!). BRUNEI : Temburong, Bukit Belalong, G.T. Prance et al. 30602 (K, KEP!). 10. Ficus dulitensis Kochummen sp. nov. (Of Mount Dulit in Sarawak) Hac species prope Ficum binnendykii var. coriacea sectionis Conosycea subgenus Urostigma, sed in flore masculo perianthiis 2, pedicello infundibuliformi differt. Typus: Dayang Awa & Yii P.C. S 46743, Borneo, Sarawak, Belaga, Dulit Range (holotypus KEP; isotypus SAR). Strangling fig. Twigs dark brown. Stipules oblong, pointed, c. 1 cm long, glabrous, caducous. Leaves leathery, elliptic to oblong, 4-6.5 x 2-3.5 cm, base broadly cuneate, apex acute; midrib sunken above; lateral veins 3-4 pairs, raised below, invisible above, trinerved, basal pair reaching more than half the length of blade; intercostal veins reticulate, visible below, invisible above; petioles 0.5—1 cm long. Syconia from leaf axils, yellowish when ripe, subglobose, c. 8 mm diameter, sessile, with 3 large basal bracts. Male flowers with obconic pedicels; tepals 2; stamen 1 with distinct filament 206 Gard. Bull. Singapore 50(2) (1998) which broadens towards the apex. Female flowers sessile; tepals 3, lanceolate; ovary brownish, reddish at stylar side, smooth, style long, terminal, stigma club-shaped. Gall flowers similar to female flowers but the style is shorter. Distribution: Endemic to Borneo. Very rare, known only from one collection from Sarawak. Ecology: Submontane forest at 820 m altitude. Notes: This species is near to F. binnendykii var. coriacea of Section Conosycea, Subgenus Urostigma but differing in the male flower with 2 tepals and in the obconic pedicel. Specimens Examined: BORNEO, SARAWAK : Belaga, Dulit Range, Dayang Awa & Yii P.C. S 46743 (KEP!, SAR!). 11. Ficus gamostyla Kochummen sp. nov. (Greek, gamo=united; with united styles of adjoining flowers) Ad Fico disticha sectionis Rhizocladus subgenus Ficus vergens, ab hac specie in ficorum pedunculis longioribus (10-13 mm), floris foemini stylo longo apicaliter florum aliorum stylis in fici cavitate adnatis. Typus: Amin et al. SAN 107123, Borneo, Sabah, Tongod (holotypus SAN). Climber. Twigs dark brown, hollow. Stipules lanceolate—acuminate, c. 4 mm long, caducous. Leaves obovate or elliptic, 7-11 x 2.5-5.5 cm, base cuneate, apex pointed, margin recurved; midrib flattened above; lateral veins 5-6 pairs, with short intermediate veins, trinerved, basal veins short, raised below, very faint above, intercostal veins fine, reticulate, distinct below, invisible above; petioles 7-10 mm long. Syconia arising from leafless twigs and branches, in clusters, greenish, ripening red, subglobose, c. 5 mm across, apex umbonate with slight depression in the centre; peduncles 10-13 mm long; basal bracts persistent. Female flowers pedicelled; tepals 4, dark brown, oblong; ovary oblong, pale brown with white edges, style lateral, long, joined up near the apex forming a white ring-like mesh; centre of syconium hollow. Male and gall flowers not seen. Distribution: Endemic to Borneo, very rare, known only from one collection from Sabah. Ecology: Lowland forest. New Species and Varieties of Moraceae 207 Notes: Near to F. disticha of Section Rhizocladus Subgenus Ficus, from which it differs in the longer (10-13 mm) peduncles of the syconia, and in the long style of female flower, which is united at the tip with the styles of other flowers within the fig cavity. Specimens Examined: BORNEO. SABAH : Tongod, Ulu Sg. Pinaggah, Amin et al. SAN 107123 (SAN). 12. Ficus ilias-paiei Kochummen sp. nov. (Ilias Paie, the collector of the type specimen) Hac species ad serie Apiocarpae, sectionis Kalosyce subgenus Ficus pertinet, prope Ficum warburgii, sed in hac specie fici subglobosae pedunculis 0-4 mm longis sunt. Typus : Ilias Paie S 42527, Borneo, Sarawak, Path to Gunong Silantek (holotypus SAR; isotypus CGE). Climber. Twigs reddish brown. Stipules semi-persistent. Leaves elliptic, 10-12 x 2.5-4 cm, base tapered, apex pointed; midrib raised above, lateral veins 5-6 pairs, very faintly visible on both surfaces with distinct areolate reticulation below, not trinerved; petioles 1.7—2 cm long. Syconia borne on older, leafless branches, arising on stout finger-like branches, greenish with white dots when ripe, elliptic, c. 7 x 2.5 cm, apex pointed, base tapered and stalk-like, about 1 cm long; peduncles 2.5-3 cm long; basal bracts tiny. Male flowers with pedicels; tepal united with 3 lobes; stamen one, exserted. Gall flowers with pedicels; tepals linear, lanceolate, brownish in the centre with white edges; ovary dark red, style lateral, stigma funnel-shaped. Distribution: Endemic to Borneo, very rare, known only from one collection from Sarawak. Ecology: Mixed dipterocarp forest at 200 m altitude. Notes: This species belongs to Series Apiocarpeae Section Kalosyce, Subgenus Ficus, and is near to F. warburgii but which has subglobose figs with a 0-4 mm-long peduncle. Specimens Examined: BORNEO. SARAWAK : Ulu Sg. Silantek, 85" Mile Simanggang Road, Ilias Paie S 42527 (CGE, SAR!). 13. Ficus kerangasensis Kochummen sp. nov. (Of kerangas forest) 208 Gard. Bull. Singapore 50(2) (1998) Species prope Ficum tristaniifolii subgenus Urostigma, sed a hac specie in ficis oblongis, folii costa immersa differt. Typus : Ilias Paie S 38595, Borneo, Sarawak, Sabal FR, (holotypus KEP; isotypi CGE, L, SAN, SAR, SING). Climber. Twigs blackish. Stipules lanceolate, pointed, 10-14 mm long, caducous. Leaves obovate, drying dark brown or blackish, 6—7.5 x 2.5-4 cm, base cuneate, apex rounded, margin curved inwards; midrib sunken above, lateral veins 6-8 pairs with short intermediate veins, curving and joining near margin, trinerved, basal pair extending up to '/3 of blade, faintly visible below, invisible above; intercostal veins reticulate, faintly visible below, invisible above; petioles 10-15 mm long, drying black. Syconia axillary, in pairs, yellowish green when fresh, sessile, oblong, 8-10 x 7 mm, surface rugose, apex truncate with distinct disc; basal bracts with blunt apex, persistent. Male flowers pedicelled; tepals 3, brown with white edge; stamen 1. Female flowers sessile; tepals 3; ovary oblong, reddish brown, longitudinally ridged, style lateral, stigma broad. Gall flowers similar to female flowers; interfloral bracts abundant. Distribution: Endemic to Borneo. Known only from one collection from Sarawak. Ecology: Kerangas forest. Notes: This species is near F. tristantifolia of section Conosyce in Subgenus Urostigma. It differs from that species in the oblong figs and in the sunken midrib of the leaf. Specimens Examined: BORNEO. SARAWAK : Sabal F.R., Simungan, Ilias Paie § 38595 (CGE, KEP!, L, SAN!, SAR!, SING!). 14. Ficus longistipulata Kochummen sp. nov. (Latin, /ongistipulatus=with long stipules) Hac species prope Ficum globosam in sectione Conosycea subgenus Urostigma. A hac species in stipulis multo longioribus, ficis minoribus differt. Typus : G. Argent et al. 441987, Borneo, Sabah, Palum Tambun (holotypus SAN). Strangling fig. Twigs brownish, irregularly ridged. Stipules lanceolate, pointed, c. 3.5 cm long, drying pinkish. Leaves elliptic to oblong, 11.5—13.5 x 5-6 cm, base rounded or broadly cuneate, apex pointed; midrib flattened above, drying to pinkish below; lateral veins 12-15 pairs with short intermediate veins, curving and joining near margin, trinerved, basal veins New Species and Varieties of Moraceae 209 very short, distinct below, visible above; intercostal veins reticulate, faint below, invisible above; petioles 2-3 cm long. Syconia in clusters on twigs below leaves, green when fresh, subglobose, 5—7 mm across, apex swollen and prominently umbonate, basal bracts small, persistent; peduncles c. 1 cm long, sparsely hairy. Male flowers in the centre of the syconium; pedicels stout; tepal brownish, united, with 3 lobes, stamen 1, anthers oblong, sessile. Female flowers sessile; tepal shorter than ovary, brownish, lanceolate; ovary whitish with red brown dots, subglobose, with faint ridges, style dark brown, subterminal. Gall flowers similar to female flowers. Distribution: Endemic to Borneo; very rare, known only from one collection from Sabah. Ecology: Lowland forest at 150 m altitude. Notes: This species is near to F. globosa in Section Conosycea Subgenus Urostigma. It differs from that species in the much longer lanceolate stipules and in the smaller figs. Specimens Examined: BORNEO. SABAH : Palum Tambun, Argent et al. 441987 (SAN!). 15. Ficus ngii Kochummen sp. nov. (F.S.P. Ng, former Forest Botanist, Forest Research Institute Malaysia) Prope Ficum corneri sectionis Leucogyne subgenus Urostigma, staminibus filamento valido, perianthiis albis, folii basi etrinervosa differt. Typus : Saw L. G. FRI 44887, Peninsular Malaysia, FRIM, Kepong (holotypus KEP). Strangling fig when planted, becoming independent tree up to 15 m tall with multiple stems, few aerial roots, and a spreading bushy dark green crown. Twigs c. 3 mm thick, covered with pale white lenticels. Stipules lanceolate, c. 8 mm long, with long white hairs on the outside. Leaves elliptic to oblong, 10-16 x 3—5.3 cm, base cuneate, apex pointed with tip c. 1 cm long; midrib flattened to slightly impressed above; lateral veins 8-11 pairs, not trinerved, thin, curving and joining near margin to form looped intramarginal vein, distinct below, very faint above, with equally prominent intermediate veins and reticulations; petioles 1-2 cm long. Sapling leaves oblong, to 3 cm broad, with sharp midrib below. Syconia from leaf axils, solitary or in pairs, sessile, green ripening yellow to deep red, subglobose to slightly obovoid, with scattered tubercles on the surface, 8-15 mm across, apex slightly depressed, closed by 3 apical bracts; basal bracts 3, concealed 210 Gard. Bull. Singapore 50(2) (1998) by the basal part of the synconium. Male flowers with short pedicels; tepals 3, white; stamen one with stout filament, anthers crescent-shaped. Gall flowers pedicelled; tepal united at base with 3-4 lobes; ovary subglobose, slightly angled, style lateral, short. Female flowers with three tepals, free; ovary white, almost globose, style lateral, stigma brownish; seed smooth. Distribution: Endemic to Peninsular Malaysia, rare, known only from two collections. Ecology: Limestone forest. Notes: This species belongs to Section Leucogyne Subgenus Urostigma, which until now was known from only two species: F. amplissima, a species of India, Sri Lanka and the Maldives, and F. rumphii, a species widely distributed in the Indo-Malesian region. F. ngii differs from these two species in leaf shape and the distinct stamen with a stout filament. F. ngii is near to F. corneri Kochummen, another newly described species of Section Leucogyne, but that species has red tepals with white edges and sessile stamens. In addition, F. corneri has leaves with a distinctly trinerved base. The type specimen was collected from a planted tree at the Forest Research Institute Malaysia, Kepong. It was grown from a cutting collected in 1982 from limestone forest near Ipoh in Perak by Dr. F. S. P. Ng. It is a very fast growing species; figs appear 2-3 times a year. Specimens Examined: PENINSULAR MALAYSIA : Perak, Cave Temple north of Ipoh, F.S.P. Ng FRI 27361 (KEP!); Forest Research Institute of Malaysia, Kepong, Selangor, planted tree, Saw L.G. et al. FRI 44887 (KEP!). 16. Ficus pseudotarennifolia Kochummen sp. nov. (Latin, pseudo=false, like F. tarennifolia) Fico tarennifolio sectionis Sycocarpus subgenus Ficus in subserie Tuberculifasciculatae similis, floribus cecidiophoris pedicellis gracilibus, foliis multo angustioribus differt. Typus : Dayang Awa S 51027, Borneo, Sarawak, Bario, Sg. Mengalio (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Small tree to 6 m tall. Twigs reddish brown, angled. Stipules lanceolate, pointed, 15-17 mm long, caducous. Leaves opposite, narrowly oblanceolate, oblong or elliptic, 9.5-22.5 x 1-3.5 cm, base cuneate, apex pointed, margin recurved, faintly wavy; midrib raised above; lateral veins 8-13, often with glands in axils of veins, not trinerved, curving and joining near margin, visible below, very faint above; intercostal veins reticulate, visible below, New Species and Varieties of Moraceae 201 almost invisible above; lower surface with few scattered glands, sometimes in axils of lateral veins; petioles 1—2.5 cm long. Syconia borne in clusters on older branches and stems, pear-shaped, c. 2 x 1.2 cm, apex depressed; peduncles to 1.2 cm long; basal bracts tiny, persistent. Gall flowers with long slender pedicels; tepal whitish, cup-shaped with irregularly shaped lobes covering 3/4 of red ovary, style short, lateral. Male flowers not seen. Distribution: Endemic to Borneo; rare, recorded only from Sarawak. Ecology: Lowland to submontane forests up to 950 m altitude, by streams. Notes: Similar to F. tarennifolia of Section Sycocarpus Subgenus Ficus Subseries Tuberculifasciculatae, but differing in the gall flowers with slender pedicels and in the much narrower leaves with faintly wavy recurved margins. Specimens Examined: BORNEO. SARAWAK : Bario, Sg. Menalio, Dayang Awa S 51027 (CGE, K, KEP!, L, SAN!, SAR!); Kakus-Pandare primary forest, Tatau, E.F. Brunig S 11929 (SAR!). 17. Ficus sabahana Kochummen sp. nov. (Of Sabah, one of the states of Malaysia) Hac species prope Ficum sagittatam sectionis Rhizocladus subgenus Ficus. A hac species in foliis glabris subcordatis infra cystolithis scabris tectis differt. Typus : Ashik Mantor SAN 114907, Borneo, Sabah, Pandewan, Mesopo River (holotypus SAN). Root climber. Twigs flattened, greyish yellow. Stipules ovate lanceolate, pointed, c. 2 cm long, subpersistent. Leaves ovate-lanceolate, greyish yellow on drying, undersurface sand papery to touch, 12.5-18.5 x 7-10 cm; base cordate or subcordate, apex pointed, margin recurved; midrib flattened above; lateral veins 3-4 pairs, trinerved, basal veins reaching more than half the length of blade, prominently raised below, flattened or faintly raised above; intercostal veins scalariform-reticulate, distinct below, faint above; petioles 1-2 cm long, sparsely hairy. Syconia borne in clusters in the leaf axils, globose, 6-11 mm across, surface smooth on drying, apex depressed, base narrowed to a cylindrical stalk; peduncle absent; bracts densely long-hairy. Male flowers shortly pedicelled; tepals 3, dark red; stamens 2, filaments united, stout. Gall flowers shortly pedicelled; tepals 3, dark red; ovary globose with dark red spots, style short, lateral. Female flowers not seen. 212 Gard. Bull. Singapore 50(2) (1998) Distribution: Endemic to Borneo. Rare, known only from Sabah. Ecology: Lowland forest in disturbed areas. Notes: This species is near to F. sagittata of Section Rhizocladus Subgenus Ficus. It differs from that species in the glabrous cordate leaves covered with rough cystoliths on the undersurface. Specimens Examined: BORNEO. SABAH : Nabawan, Keningau, Dewol Sundaling SAN 83817 (KEP!, SAN!, SAR!, SING!); Pandewan, Mesopo River, Ashik Mantor SAN 114097 (SAN!). 18. Ficus soepadmoi Kochummen sp. nov. (E. Soepadmo, Collaborator and Chief Editor of the Tree Flora of Sabah and Sarawak Project) A subsectione Dictyoneuron sectionis Conosycea subgenus Urostigma pertinet. Ad Ficum sumatranum vergens, Sed ficis carinatis et folii venatione differt. Typus : Ilias Paie & Yeo Eng Teck S 38376, Borneo, Sarawak, Lambir Hill National Park, Ulu Sg. Lebau (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Climber. Twigs dark brown, angled. Stipules lanceolate, c. 15 mm long, appressed hairy outside. Leaves drying greenish yellow; oblong or oblanceolate, 9-13 x 2.8-3.7 cm, base cuneate, apex pointed, margin recurved; midrib sunken above; lateral veins 5-7 pairs with a number of short intermediate veins, trinerved, curving and joining near margin to form looped intramarginal vein; intercostal veins reticulate, venation visible below, very faint to inconspicuous above; petioles 1—-1.5 cm long, drying black, sparsely hairy. Syconia axillary, sessile, in pairs, pale green when ripe; subglobose, 6-8 mm across, apex faintly sunken and closed by 2 apical bracts; surface rugose with five prominent ridges from apex to base; basal bracts persistent, with rounded apices. Male flowers sessile; tepals 3, stamen 1, sessile. Female flowers sessile; tepals 3, oblong, reddish; ovary oblong, irregularly lobed, dark brown on one side, style lateral, long, stigma dark brown, lobed. Gall flowers similar to female flowers but style short; interfloral bracts abundant. Distribution: Endemic to Borneo; very rare, known only from the type collection from Sarawak. Ecology: Submontane forest by river side at 1400 m altitude. New Species and Varieties of Moraceae 213 Note: Belonging to Subsection Dictyoneuron Section Conosycea Subgenus Urostigma, it comes near to F. sumatrana but the ridged figs and leaf venation are different. A sterile collection from Brunei, BRUN 5332 from S. Belalong, a juvenile stage collection, probably belongs here. Specimens Examined: BORNEO. SARAWAK : Ulu Sg. Lebau, Lambir Hill National Park, Ilias Paie & Yeo Eng Teck S 38376 (CGE, K, KEP!, L, SAN!, SAR!). 19. Ficus cereicarpa Corner var. ashtonii Kochummen var. nov. A varietate typica in ramulis crassioribus, foliis maximis cordatisque, ficis longe pedunculatis differt. Typus : P.S Ashton S 17806, Borneo, Sarawak, Balleh, Ulu Selentang (holotypus KEP; isotypi A, BO, CGE, K, L, SAR, SING). Small tree to 6 m tall. Twigs 10-18 mm thick, glabrous and prominently ridged with prominent stipular scars, young twigs covered with patent brown hairs. Stipules to 5 cm long covered with patent brown hairs. Leaves elliptic or oblong, 32-43 x 20-24 cm, lower surface sparsely brown hairy on the midrib and lateral veins, base distinctly cordate, apex pointed, lateral veins 16-20 pairs, palmately veined at base; petioles 9-14 cm long, covered with patent brown hairs when young, becoming glabrous. Syconia borne at base of bole, pear-shaped, 2.5-3 x 2.5—3 cm, surface hairy, with thick lateral bracts, peduncle 3 cm long. Distribution: Endemic to Borneo; recorded only from Sarawak. Ecology: Lowland forest by streams. Notes: Differs from the typical variety in the stouter twigs, very large cordate leaves and in the long-peduncled figs. Specimens Examined: BORNEO. SARAWAK : Balleh, Ulu Selentang, P.S. Ashton S 17806 (A, BO, CGE, K, KEP!, L, SAR!, SING!); Lambir National Park, Sg. Riam Libau, Rena George S 40302 (CGE, E, K, KEP!, L); Belaga, Sg. Iban, Bernard Lee S 45512 (CGE, K, L, MO, SAN!, SAR!). 20. Ficus deltoidea Jack var. recurvata Kochummen var. nov. (Latin, recurvatus=curved backwards; the leaf margin) Ab varietatibus aliis Fici deltoideae in foliis maioribus crasse coriaceis marginibus valde recurvatis differt. Typus : Othman, Yii et al. S 48969, 214 Gard. Bull. Singapore 50(2) (1998) Borneo, Sarawak, Tubau (holotypus KEP; isotypi SAN, SAR). Epiphyte. Leaves thickly leathery, deltoid, 7.5-12 x 4-10 cm, base tapered, apex rounded, margin curled inwards; midrib forked, sunken above on the lower half; lateral vein 1 pair, trinerved; intercostal veins reticulate, visible below, invisible above; petioles 1-3 cm long, channelled above. Syconia green with brown tip when fresh, becoming black on drying; subglobose, c. 12 mm across, apex umbonate, peduncle short. Gall flowers with irregularly lobed ovary and short style; tepals 3 in male and gall flowers. Female flowers not seen. Distribution: Endemic to Borneo; rare, recorded from Bintulu, Semengoh, Simanggang and Tubau in Sarawak. Ecology: Lowland swamps to submontane forest. Notes: Differs from other varieties of F. deltoidea in the larger thickly leathery leaves with distinctly recurved margins. Specimens Examined: BORNEO. SARAWAK : Simanggang, E.W.F.O. Brunig S 4807 (SAR!); Arboretum, Semengoh F.R., J/.A.R. Anderson S 12930 (SAR!); Tubau, Dataran Tinggi Merurong, Othman Yii et al. S 48969 (K, KEP!, L, SAN!, SAR!); Ulu Sg. Kemena, Selah, Tatau, Rantai Jawa S$ 65653 (SAR!). ; 21. Ficus deltoidea var. subhirsuta Kochummen var. nov. (Latin, sub=somewhat, hirsutus=rough hair covering; the figs) A varietatibus aliis Fici deltoideae in ficis pubescentibus et foliis venis lateralibus in 4 vel 5 paribus differt. Typus : Yii P. C. S 48452, Borneo, Sarawak, Batang Balleh, Bukit Melatai (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Epiphyte. Twigs dark brown, scaly. Leaves in life yellowish on the under surface; obovate, 2.2-3.5 x 1.7-2 cm, base tapered, apex rounded; midrib forked near the apex; lateral veins 4-5 pairs, curving and joining near margin, distinct below, faint above; petioles 3-6 mm long. Syconia axillary, red when ripe, solitary, elliptic, c. 3 x 2 mm, surface rough hairy; peduncles c. 2 mm long; basal bracts persistent, hairy. Distribution: Endemic to Borneo, very rare, known only from Sarawak. Ecology: Hill forest at 870 m altitude. New Species and Varieties of Moraceae 215 Note: Differs from other varieties of F. deltoidea in having hairy figs and leaves with 4—5 pairs of lateral veins. Specimens Examined: BORNEO. SARAWAK : Batang Balleh, Bukit Melatai, Yii P. C. S 485452 (CGE, K, KEP!, L, SAN!, SAR!). 22. Ficus obscura Blume var. lanata Kochummen var. nov. (Latin, /anatus=woolly) A varietate typica in pilis lanuginosis in ramulis foliis ficisque differt. Typus: Henry T. Sinanggul SAN 57361, Borneo, Sabah, Semporna, Kuala Kalumpang (holotypus KEP; isotypus SAN). Twigs with woolly reddish brown hairs. Leaves: petioles and undersurface of leaves woolly hairy, uppersurface rough to touch. Syconia from leaf axils and on twigs below leaves, in pairs or in clusters, sessile or with peduncles, covered with woolly hairs, subglobose, 8-10 mm broad. Distribution: Endemic to Borneo, frequent in Sabah, very rare in Sarawak. Ecology: Lowland and hill forest, often by streams, at Serian on limestone. Note: Differs from the typical variety in having woolly hairs on the twigs, leaves and figs. Specimens Examined: BORNEO. SABAH : Lahad Datu, Kennedy Bay Timber Co. Road, G.H.S. Wood SAN 16080 (A, BO, BRI, K, KEP!, L, SAN!, SING!); Semporna, Kuala Kalumpang, Henry T. Sinanggul SAN 57361 (KEP!, SAN!); Ranau, Kampung Takutan, G. Shea & Aban SAN 77208 (A, K, KEP!, L, SAN!, SAR!, SING!); Kalabakan, Hap Song logged area, Fedilis & Sumbing SAN 91428 (BO, K, KEP!, L, SAN!, SAR!, SING!); Kalabakan, Fedilis Krispinus SAN 94809 (KEP!, SAN!, SAR!), SAN 95899 (KEP!, SAN!); Ranau, Poring, Amin et al. SAN 121496 (KEP!, SAN!). SARAWAK : Serian, S 28102 (SAR!). 23. Ficus oleifolia King var. calcicola Kochummen var. nov. (Referring to the limestone habitat) A varietatibus aliis Fici oleifoliae in venatione indistincta et habitione calcarea differt. Typus : Bernard Lee S 38626, Borneo, Sarawak, Gunung Doya (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Small tree to 4.5 m tall. Twigs grey brown. Stipules linear, c. 4 mm long. 216 Gard. Bull. Singapore 50(2) (1998) Leaves elliptic, drying to greenish yellow; base cuneate, apex pointed, margin recurved; midrib raised above, lateral veins and intercostal veins very faint to inconspicuous; petioles 7-10 mm long. Syconia ellipsoid to subovoid, c. 3 mm diameter, apex umbonate; peduncles 3-4 mm long. Distribution: Endemic to Borneo; recorded only from the 1” Division in Sarawak, locally frequent. Ecology: Limestone forest. Note: Differing from other varieties of F. oleifolia in the indistinct venation and its limestone habitat. Specimens Examined: BORNEO. SARAWAK : Bau, G. Jebong, Banyeng Ludong S 38535 (CGE, K, KEP!, L, SAN!, SAR!); G. Doya, Bernard Lee S 38626 (CGE, K, KEP!, L, SAN!, SAR!); G. Jambuan, Bernard Lee S$ 38604 (CGE, K, KEP!, L, SAN!, SAR!); G. Majar, Tebekang, Yii & Othman S 46283 (CGE, K, KEP!, L, SAN!, SAR!); Bukit Jambusam, Bau, Yii et al. S 50349 (CGE, K, KEP!, L, SAN!, SAR!). 24. Ficus oleifolia King var. impressicostata Kochummen var. nov. (Latin, impressus=sunken, costa=midrib; midrib impressed above) Prope Ficum oleaefoliam var. memecylifoliam, sed in foliis costa immersa differt. Typus : Ilias Paie S 40961, Borneo, Sarawak, Kapit, Melinau (holotypus KEP; isotypi CGE, K, L, SAN, SAR). Epiphyte. Leaves elliptic, base cuneate, apex pointed; midrib sunken above, lateral veins and intercostal veins very faint to invisible; petioles short, 2-3 mm long. Syconia oblong to subglobose, 3-4 mm wide, on 5—8 mm long slender peduncles. Distribution: Endemic to Borneo; reported only from Sarawak, rare. Ecology: Hill and submontane forest between 700-1300 m altitude. Notes: Very close to var. memecylifolia but differing in the leaves with the midrib sunken. Specimens Examined: BORNEO. SARAWAK : Ulu Melinau, Hose Mountains, Paul Chai et al. S 37304 (CGE, K, KEP!, L, MO, SAN!, SAR!); Anap, Bukit Mersing, Sibat ak Luang S 21943 (A, BO, K, KEP!, L, MEL, P, SAN!, SAR!, SING!); Kapit, Melinau, Jlias Paie S 40961 (CGE, K, KEP!, L, SAN!, SAR!). New Species and Varieties of Moraceae 217 25. Ficus sundaica Blume var. impressicostata Kochummen var. nov. A varietatibus aliis Fici sundaicae a folii pagina superiore costa immersa distinguendam. Typus : Talip Bidin SAN 80664, Borneo, Sabah, Papar, Mandahan (holotypus KEP; isotypi K, L, SAN, SAR, SING). Twigs grey brown, ridged. Stipules glabrous, ovate—lanceolate, 1—-1.5 cm long. Leaves elliptic to narrowly obovate, 6.5-11 x 2.8-5.5 cm; drying chocolate brown; base broadly cuneate, apex cuspidate with short tip, margin wavy, recurved; midrib sunken above; lateral veins 5-6 pairs with 3-4 intermediate veins between each pair, looping and joining near margin to form an intramarginal vein, trinerved, basal pair reaching more than half the length of blade, very faint on both surfaces; intercostal veins reticulate, very faint below, almost invisible above; petioles 1.5—2 cm long, distinctly channelled above. Syconia axillary, sessile, oblong, 15-22 x 12-20 mm, yellowish when fresh, irregularly wrinkled on drying, apex almost flattened; basal bracts large, ovate, c. 10 x 8 mm, persistent. Distribution: Endemic to Borneo. Common and widely distributed in Sabah and Sarawak. Ecology: Lowland forest on sandy soils, in kerangas and peat swamp forests. Notes: It is distinguished from the other varieties of F. sundaica by the midrib, which is impressed on the upper surface. Specimens Examined: BORNEO. BRUNEI: Bukit Pasir Puteh, Ladi anak Bikar BRUN 5116 (BRUN, KEP!); Seria, B.E. Smythies et al. S 5864 (BRUN, KEP!, SAR!); Temburong, S. Atkins et al. 498 (BRUN, K, KEP!); Belait, Badas, M.J.E. Coode et al. 6470 (BRUN, K, KEP!), D. Kirkup et al. 387 (BRUN, K, KEP!); Tutong, Pasir Puteh, M.J.E. Coode et al. 6850 (BRUN, K, KEP!); Tutong, Tanjong Maya, D.A. Simpson et al. 2188 (BRUN, K, KEP!); Tutong, Bukit Pasir, K.M. Wong 161 (BRUN, K, KEP!). SABAH : Papar, Kimanis F.R., Aban Gibot SAN 49382 (SAN!); Papar, Bongawan F.R., Dewol & Talip Bidin SAN 80345 (A, KEP!, OX, SAN], SAR!, SING!); Papar, Mandahan, Talip Bidin SAN 80664 (K, KEP!, L, SAN!, SAR!, SING!); Beaufort, Hindian F.R., Talip Bidin SAN 84555 (KEP!, SAN!, SAR!); Sipitang, Malalia F.R., R.A. Marsal & Heya SAN 86202 (KEP!, SAN!); Sipitang, Maritinetaman Forest Area, Ag. Amin & Heya SAN 86485 (KEP!, SAN!, SAR!, SING!); Membakut, Sg. Damit, Ag. Amin SAN 103271 (K, KEP!, L, SAN!, SAR!); Weston, Sianggu F.R., Ag. Amin SAN 105981 (KEP!, SAN!); Sipitang, Mengalong F.R., L. Madani SAN 111407 (KEP!, SAN!); Sipitang, Melakis F.R., Ag. Amin SAN 114863 218 Gard. Bull. Singapore 50(2) (1998) (KEP!, SAN!). SARAWAK : Miri, Lambir Hills, Joseph Au S 17259 (A, BO, CGE, K, L, MO, SAN!, SAR!); Balingian, Begruh, Bawan, Paul Chai S 19474 (A, BO, CGE, K, L, SAN!, SAR!, SING!); Baram, Mt Dulit, Sylvester Tong S 34898 (CGE, K, KEP!, L, MO, SAR!); Ulu Simunjan, G. Buri, Bernard & Ilias § 36850 (CGE, K, KEP!, L, MO, SAR!); 99 mile Sri Aman, Kampung Gum, J/ias Paie S 42734 (CGE, K, KEP!, L, SAN!, SAR!); 99 mile Sri Aman, Ilias Paie S 42739 (CGE, K, KEP!, L, MO, SAN!, SAR!); Batang Balleh, Bukit Melatai, Yi P. C. S 48453 (CGE, kK KPIS SAN!, SAR!). Acknowledgements I wish to extend my gratitude for the support and encouragement of the Director-General of FRIM, Kepong and to the Curators of K, L, SAN, SAR and SING herbaria for the loan of specimens. I am also deeply indebted to Dr. J.F. Veldkamp of the Rijksherbarium, Leiden, the Netherlands, for providing the Latin diagnoses of the new taxa. The research was part of the Tree Flora of Sabah and Sarawak Project, funded by the International Tropical Timber Organization (ITTO), the Malaysian Government, and the Overseas Development Administration of the United Kingdom (ODA-UK). References Corner, E.J.H. 1965. Check-List of Ficus in Asia and Australasia with keys to identification. Garden’s Bulletin Singapore 21: 1-186 Corner, E.J.H. 1970. Ficus Subgen. Ficus: Two rare and primitive pachycaul species. Philosophical Transactions Royal Society, London. Ser. B 259 No. 831: 353-381. Corner, E.J.H. 1972. New taxa of Ficus (Moraceae). Blumea. 20: 427-432. Corner, E.J.H. 1972. Moraceae of Malesia (Unpublished manuscript). Jarrett, J.M. 1959. Studies in Artocarpus and allied genera, III. A revision of Artocarpus subgenus Artocarpus. Journal Arnold Arboretum. 40: 113-155. Jarrett, J.M. 1960. Studies in Artocarpus and allied genera, IV. A revision of Artocarpus subgenus Pseudojaca. Journal Arnold Arboretum. 41: 73-140. New Species and Varieties of Moraceae 219 Jarrett, JM. 1975. Four new Artocarpus species from Indo-Malesia (Moraceae). Blumea. 22: 409-410. nARDWce “AM eX i wee 4 xh pyrite ils a nae o bse Le “ We Mie Ae » ye f ‘, ane ty * Ny 1 a, m a ME ie ioe 4, Lae / ihe were Re cas A has i} Gardens’ Bulletin Singapore 50 (1998) 221-334. The Genus Alocasia (Araceae-Colocasieae) in West Malesia and Sulawesi A. HAY Royal Botanic Gardens Sydney, Mrs Macquaries Road, Sydney 2000, Australia Abstract Alocasia (Schott) G. Don is revised for West Malesia and Sulawesi. Thirty one species are recognised, including one extremely variable species, A. longiloba Miq. s. /., in which seven incompletely delineable informal entities are further recognised. Ten species are new to science. The history, geography, ecology and morphology of the genus and conservation status of its species are discussed and foci for further study are briefly delineated. A key to species is provided. Approximately 25% of names are epi- or neotypified owing to lack of adequate original material - a situation deriving mainly from the horticultural history of the genus. New synonyms include A. margaritae L. Linden & Rodigas, A. ovalifolia Ridl., 7A. crassinervia Engl. = A. puber (Hassk.) Schott; A. imperialis L. Linden, A. guttata N.E. Br., A. villeneuvei L. Linden = A. scabriuscula N.E. Brown; A. porphyroneura Hallier f. = A. princeps W. Bull; A. grandis N.E. Br. = A. macrorrhizos (L.) G. Don; A. nobilis Hallier f, = A. inornata Hallier f.; A. bantamensis Koord., A. crassifolia Engl. = A. alba Schott; A. lowii Hook., A. korthalsii Schott, A. denudata Engl., A. putseyzii N.E. Br., A. eminens N.E. Br., A. watsoniana Mast., A. curtisii N.E. Br., A. cuspidata Engl. = A. longiloba Mig. s. l.. Alocasia perakensis Hemsl. is reinstated. Indian Alocasia montana (Roxb.) Schott is considered a synonym of A. macrorrhizos. Contents Peete eee melanin OAS 5522 ehh I in RR I, LG gas ds 222 Ree tee hte eet ae, SUT eet EE OL ls DAR Ee eh Peer 223 SAE Sire ert AME ed I er oh GEES L004) Sag tis ATM hee eae ded 225 ss ok ae 1s) 77 1 (Mie) 1G [ea 00rd aa eae SE Sia pe et CR 226 OTIS DV 1 ae ae Sey a Zee ee 226 De Ue Ce RMmNOIO ny LAN ures. Le. LU eo Ze Diraiguishine vocasia mom Colocasia 3!) BIRO ae 233 eter Se Cnt LE on A ANA lode 234 Ae EE Ts ME aa 0 SE ls ao eae a eee 2355 CAAT SAIS Mo SIMS 1) ER ALA MRS Ue hyn 0) Se de 0 ke, A 236 ine ye sPeGlcs anid Species COMPIEKES hs. A I a. 238 oh 100 ae B30 OF nn ae nag ei Ola ERs aes 8 I aaa 242 Peers MLL CREM nanos oey teats al VEEN: Le Odean Me Pine 252 Oe Coon) (ge Ss Sates ee i co Ra ef Sl 261 Ped EO RII AON RONMD oh hte ea Bee Uh el a 282 UU Glo T GUO a eas A OR Ue Ree On RU 298 222 Gard. Bull. Singapore 50(2) (1998) Cuprea Group ii cassived age Dated dag A aaa ae ae 316 Inadequately KmOwi- Spe cues oie) a on 15/4 civtn piste Ruel nice SEM oie ue aay Doubtful species and records 05)... 2: - ds. se ea oet ns are a 328 Acknowledgements. iiii ios Soh si ceuse ano ce Apes Aue ae 329 References .oi.ic.5s cise d bade oa sere ob hdc t dn tess wih a Wien Us ea 330 Brace x €0SPCCHES iss oid ed als w's eateeate Oy ac ae elo aregs cae aaa cea eT 334 Aims, Scope and Limitations The present review is, as with previous papers on this genus covering other parts of Malesia (Hay & Wise, 1991; Hay, in press), limited to an ‘alpha- taxomonic’ review of the species as a precursor to a treatment for Flora Malesiana (see Hay, 1994b) and aims to bring understanding of the genus to a basis point for more intensive phylogenetic, molecular, biogeographic and ecological analyses. Alocasia species are widely cultivated as ornamental plants and often abundant in the herb layer and gap phase of forests, and there is an urgent need for the means of identification of species. Defining species through standard herbarium-based methods is difficult, as the plants are generally unpleasant to collect and unsuited to herbarium preservation, often having enormous leaves and rhizomes, soft parts, often complex and bulky synflorescences, ephemeral inflorescences and irritant sap, and they are phenotypically variable. Moreover, they are highly attractive to herbarium beetles and the percentage of specimens with well preserved floral parts is minute. Field work and the assembling of a living collection, in which sterile field-collected plants can be brought to flower, has been an essential prerequisite to this revision. While this has enabled many species to be described from fresh material, there still remain a number known only from herbarium specimens, and a number in which variability has to be interpreted almost solely from the inadequate resource of dried material. Many cultivated accessions have been cited, from which further preserved material will be distributed to relevant herbaria in due course. The convention I have used in citing these is to give the RBG Sydney Accession Number as well as the original collection number. Wild-collected vouchers bear only the original collection number, while preserved specimens made from the cultivated plants bear both numbers. Infrageneric species groups have been recognised informally, as it is beyond the scope of the present work to engage in the rigorous analysis of species relationships that would justify formal infrageneric nomenclature. Comment on the coherence or otherwise of the informal groups may be found under each in the body of this paper. The division of the genus into two formal sections based on stigma shape, as proposed by Engler & ET ge —_—— -e— Alocasia in West Malesia and Sulawesi 223 Krause (1920) is plainly unsatisfactory. It splits the Longiloba Group, for example, presumably on misobservation, and aligns, in Sect. Ensolenanthe, A. zebrina Schott ex van Houtte (Philippines) with A. cuprea and members of the A. longiloba complex - a very probably heterogeneous assemblage. History The first Malesian Alocasia species were attributed, as were many monoecious aroids before the advent of Schott, to the genus Arum. In the pre-Linnean period, Rumphius (1747) illustrated two ‘species’ (within which he recognised further ‘species’ in his discussion), Arum indicum sativum and Arum sylvestre. The former was the widespread Alocasia macrorrhizos (L.) G. Don, for which, as the genus becomes better understood in the wild state, there is a growing body of evidence that it is a cultigen associated almost entirely with human habitation and agriculture as an inferior starch crop. Arum indicum sativum was taken up by Loureiro (1790) as Arum indicum Lour., the basionym of Alocasia indica (Lour.) Spach, now recognised as a synonym of A. macrorrhizos. Arum sylvestre has remained of obscure identity, though Hasskarl (1844) considered it perhaps identical with West Malesian Colocasia pubera Hassk. (= Alocasia puber (Hassk.) Schott). That interpretation seems incorrect, however, and it seems more plausible that Arum sylvestre is A. aequiloba N.E. Br., an East Malesian species. The generic name Alocasia was first attributed to Necker by Rafinesque (1837: 64) for some aroid species now in Arisaema. The name was earlier used by Schott (1832), the first significant specialist in the taxonomy of Araceae, as a section in Colocasia. That concept of Alocasia was raised to generic rank by Don in Sweet (1839), with the Indochinese A. cucullata (Lour.) G. Don as the type. The name has been used in Schott’s and Don’s sense ever since, with Nicolson (1963) proposing it be conserved over Alocasia Rafinesque. Early species now in the genus were described or combined in Colocasia for some time after Don with Kunth (1841) transferring Arum indicum Lour. into Colocasia and Hasskarl (1842) misapplying Colocasia odora Roxb. to (probably) Alocasia macrorrhizos and Colocasia montana (Roxb.) Kunth to Alocasia flemingiana (a species newly described here). In 1844 he called one of the earliest described (post-Linnean) Malesian endemic Alocasia species Colocasia pubera Hassk. Hasskarl further recognised several forms of Alocasia macrorrhizos (now best regarded as cultivars) in Colocasia. However, from Miquel (1855) onward, Alocasia has been consistently treated by botanists of Malesian aroids, including Hallier, Ridley, Hemsley, Koorders, and more recently Backer & Bakhuizen and Hotta as a genus distinct from Colocasia, though 224 Gard. Bull. Singapore 50(2) (1998) confusion persisted among those describing species in the horticultural arena. Curiously, van Alderwerelt van Rosenburg, who wrote a series of papers on Malesian Araceae (Alderwerelt 1920, 1922), did not deal with Alocasia at all. Much earlier, the great botanist of the Malesian flora Blume took a particular interest in Araceae (1837) but did not deal with any Alocasia species either, though in his Catalogus (1823: 102) he listed Arum maximum (presumably Alocasia macrorrhizos). He also determined certain of his herbarium collections as Caladium pubigerum (= Alocasia puber), though the name was apparently never published. Caladium had been erected by Ventenat (1801) for the ornamental, peltate-leaved neotropical C. bicolor. The genus is superficially similar to Alocasia (though on molecular evidence no longer considered very closely allied - French et al., 1995; Mayo, Bogner & Boyce, 1997), and the earliest post-Linnean Malesian Alocasia - Philippine A. heterophylla (Presl) Merr. first apeared in it (Caladium heterophyllum Presl, 1835). By the mid-19th Century, in spite of the development of Schott’s generic concepts (1853-1860), some ornamental peltate-leaved Alocasia species, coming to be known in the stream of exotic plants being brought into cultivation in Britain and Europe, were still first described in Caladium, including Alocasia cuprea (C. Koch & Bouche) C. Koch) and Alocasia veitchii (= A. longiloba Migq.). Included in the history of Alocasia is the recognition of four segregate genera: Ensolenanthe Schott (1861), proposed without any binomials and later used by Engler & K. Krause at infrageneric rank; Schizocasia Engl. for a group of species with deeply divided leaf blades, now understood to be heterogeneous (Bunting, 1962; Nicolson 1968); Xenophya Schott, now regognised as an East Malesian species group within Alocasia (Hay & Wise, 1991); and monotypic Panzhuyia Z.Y. Zhu (1985), conspecific with the type of Alocasia, A. cucullata. The role of British and Continental nurserymen in the discovery of Malesian (particularly Bornean and Philippine) Alocasia species in the latter part of the 19th century has been considerable, notably the firms of Veitch, Sander, Bull and the Compagnie Continentale d’Horticulture Gand. Some spectacularly beautiful plants were introduced and named, principally by Lucien Linden, Masters and N.E. Brown, through the horticultural route. However, a legacy has been left of species named in the absence of any understanding of variation in the wild - since evidently only the most striking forms were introduced on the basis of their commercial potential, of protologues consisting of short entries in retail catalogues or rhapsodic accounts of horticultural qualities neglecting botanical detail, fragmentary or non-existent type specimens or second-rate and barely interpretable illustrations, and provenances inaccurate, vague and sometimes quite Alocasia in West Malesia and Sulawesi 225 probably deliberately misleading to commercial competitors. This has resulted both in a high level of new synonymy, and, together with the WWII destruction of the herbaria at Vienna and Berlin, where the two great figures in Araceae taxonomy, Schott and Engler, worked, a particularly large number of cases of difficulty in interpreting the application of names in this genus, requiring the neotypification or epitypification of no less than a quarter of them. The genus was last revised in toto by Engler & Krause (1920). Nicolson (1987) revised the Sri Lankan species, Hay & Wise (1991) revised the East Malesian and Australian species, Noltie (1994) revised Himalayan species and Hay (in press) has revised the Philippine species. Ecology Alocasia species are predominantly lowland tropical plants of ‘ever-wet’ areas. A few species, such as A. perakensis and A. kerinciensis are montane elements, though few occur above about 1200 m alt. and none above about 2000 m alt. in the region under study here. Some species, such as A. princeps, have wide altitudinal ranges. Several species, such as A. sarawakensis, A. minuscula and A. puber show a preference for or restriction to swampy habitats, while others, such as A. princeps and A. beccarii, are restricted to well-drained sites. Several are facultative or obligate lithophytes, including A. longiloba ‘watsoniana’, A. longiloba ‘lowii’, A. principiculus, A. puteri, A. pangeran, A. ridleyi, A. venusta and A. reversa. Detailed information is often lacking, but some species are associated with or confined to particular substrates. Alocasia melo is confined to ultramafic areas. A. reversa, A. venusta, A. ridleyi, A. puteri, A. pangeran, A. principiculus are confined to limestone areas. A. minuscula is known only from peat swamp forest. Several species do not appear to be much influenced by substrate - for example, A. cuprea is found on sandstone, limestone and in ultramafic areas, and A. princeps occurs on a wide variety of substrates, limestone, sandstone, shale and so on. The genus can be broadly divided into gigantic species, which are associated with open sites - gap phase of forests, landslips, river banks, open swamps, road-sides and plantations - such as Alocasia robusta, A. sarawakensis, A. alba, A. puber, and smaller elements that are generally found within forest. Alocasia scabriuscula and A. inornata, and to a lesser extent, some elements of the A. lJongiloba complex can be found both within forest and in open conditions. 226 Gard. Bull. Singapore 50(2) (1998) Geography and Endemism Few Malesian Alocasia species are widespread, other than those distributed directly by human activity, such as A. macrorrhizos. Borneo is the main node of diversity, endemism and richness in Alocasia, with a second node in New Guinea/Australia which, though slightly less speciose than the Philippines, is taxonomically distinct (Hay & Wise, 1991; Hay, 1994a). The ratios of endemic species to total species (calculated excluding A. macrorrhizos and including entities of the A. Jongiloba complex) for the main land masses in Malesia are as follows: Malay Peninsula - 1:8; Sumatera - 3:8; Java: 2:4; Borneo - 20:23; Philippines - 14:14; Sulawesi - 3:4; Papuasia - 12:12; [Australia - 1:1]. Of the informal infrageneric groups recognised here, the Puber Group group has four species, two Bornean endemics, one Philippine endemic and one species from Java and the Malay Peninsula, possibly also represented in Sumatera. This group appears to be closely allied to the Scabriuscula Group, which is endemic to, richly represented and abundantly common in Borneo, including two highly variable complex entities - Alocasia scabriuscula and A. princeps - each with several narrowly defined localised segregates. The Longiloba Group is predominantly West and Central Malesian, extending from the central highlands of Vietnam to the Philippines and Sulawesi. With the exception of Philippine and Sulawesi segregates, it is treated here as one species including localised, sometimes sympatric ‘topospecific’ entities that merge globally into a single presently intractable complex. It is most diverse in Borneo and Sumatera. The Cuprea Group is mainly Bornean, with a single endemic representative in each of Sumatera and the Malay Peninsula. Remaining species have been very provisionally placed in an alliance with A. macrorrhizos as the Macrorrhizos Group, which occurs from mainland Asia and thoughout the Malesian range of the genus except Borneo. However, this grouping may prove heterogeneous on further study. Conservation Status One outcome of a taxonomic study is to highlight species that are known from very few collections or that are known from only a few localitites and which may therefore be rare and in need of protection. However, before listing those species, it should be emphasised that conservation status of course requires verification on the ground, preferably by local botanists. Nevertheless, existing herbarium records are a start in evaluating how abundant or localised the species may be. Alocasia in West Malesia and Sulawesi ZIT The following species are Known from very few collections and/or localities: Sumatera - Alocasia kerinciensis, Borneo - A. melo, A. minuscula, A. pangeran, A. principiculus, A. puteri, A. reginae, A. reginula, A. venusta and Sulawesi - A. suhirmaniana and A. celebica, the latter known only from its type, collected over a century ago in Sulawesi. The following wide-ranging species are known from few collections in certain main subdivisions of the Malesian Archipelago while being more frequent elsewhere: Alocasia puber in the Malay Peninsula and Sumatera, and A. longiloba ‘watsoniana’ in the Malay Peninsula and Borneo. Many of these, especially Alocasia melo, A. principiculus, A. reginae, A. reginula, A. suhirmaniana, A. venusta and A. longiloba ‘watsoniana’ are highly ornamental (as indeed are less rare species such as A. reversa, A. cuprea and certain forms of A. inornata, A. longiloba, A. perakensis, A. princeps and A. scabriuscula). On the one hand they are potentially threatened by unsustainable unscrupulous collecting from the wild, and on the other they are open to ex situ conservation (in a broad sense) through the medium of ornamental horticulture sustained by tissue culture. Indeed, a number of species have been successfully micropropagated, especially in the U. S. A., and are available for sale over the internet. With increasing ease of developing and implementing tissue culture protocols for micropropagation, there is great potential for commercially developing these plants within their countries of origin at the same time as relieving collecting pressure on limited wild plant populations. A horticultural account describing a large number of cultivars in some detail was given by Burnett (1984), though there is a pressing need to stabilise the nomenclature of the cultivars and to align it, where appropriate, with the botanical nomenclature. Structure and Terminology The stem, typically of most Araceae, is a physiognomically unbranched sympodium. The number of foliage leaves per module is variable between and within species and individuals, but during flowering episodes in some species it may be reduced to one. In some species, e.g. A. kerinciensis, foliage leaves alternate with cataphylls within a module. In such instances the cataphyll performs the role of protecting the subsequent emerging leaf. That role in other species is performed by the sheath of the next oldest foliage leaf. Those species with regularly interspersed cataphylls typically have very short leaf sheaths, while those without interspersed cataphylls have longer sheaths. A prophyll and usually at least one cataphyll is always associated with the initiation of a new vegetative module. The vascular system of the petiole divides, as it runs into the blade, 228 Gard. Bull. Singapore 50(2) (1998) into three principal veins - the anterior costa (midrib) and two posterior costae which support the anterior and posterior lobes of the blade respectively. The shape of the posterior lobes of the leaf is sometimes of diagnostic importance. Terms used here to describe them are for the most part self-explanatory, such as ‘acute’, ‘obtuse’, etc. The posterior lobes are assymetric, the outer sides being united with the anterior lobe, while the inner sides (i.e. those that face each other across the sinus) are free (unless the leaf is peltate). In some instances, such as the Scabriuscula Group, the shape of the piece of lamina on the ‘inside’ of each posterior lobe may need to be used for identification purposes. For these parts I have used terms such as ‘lanceolate’, ‘ovate’, etc. even though the posterior lobe is not symmetrical about the posterior costa. Thus, ‘inner side of posterior lobe lanceolate’ means that the inside piece of lamina is shaped like a longitudinally bisected lanceolate leaf (Fig. 1). Figure 1. Diagram of Alocasia leaf blade A. anterior lobe; B. posterior lobe; C. anterior costa; D. posterior costa; E. primary lateral vein; F. subsidiary vein (with axillary gland); G. secondary vein; H. sector with interprimary collective vein formed by meeting of secondary veins; I. sector with interprimary collective vein not formed; J. axillary gland; K. intramarginal vein; L. inner side of posterior lobe; M. petiole; N. sinus. Alocasia in West Malesia and Sulawesi 229 Primary veins run pinnately off both sides of the anterior costa and pedately off the outer (anterior) side of each posterior costa. Glands, of unknown function, are found in the axils of the primary veins on the abaxial surface of the leaf, and may also occur scattered over the surface of the petiole. Secondary venation arises direct from the costae and from the primary lateral veins and is typically colocasioid: secondary veins arising from the primary lateral veins typically run initially at a wide angle from the primary venation and are then deflected towards the margin of the blade. In some species the secondary veins unite between the primary veins into more or less sinuous interprimary collective veins. These may be very well developed and distinct, and while they are a useful feature for distinguishing some species, the state intergrades with a complete absence of interprimary veins and some species evince a variety of intermediate states. In some species (e.g. Alocasia suhirmaniana), some secondary veins are intermediate in thickness between the normal secondary venation and the primary veins, and they may even bear glands in their axils like the primary venation. These are termed subsidiary veins. The primary and secondary veins run into a marginal vein, or in some species a distinct intramarginal vein (e.g. A. peltata). The primary and secondary venation patterns in those species with interprimary collective veins is highly suggestive of derivation by connation of leaflets or segments of a pinnate or pinnatifid leaf. Indeed, more or less deeply pinnatifid leaves occur sporadically in the genus in different species groups, e.g. Philippine A. sanderiana W. Bull (Longiloba Group), Papuasian A. brancifolia (Schott) A. Hay (Xenophya Group) and Philippine A. portei Schott (Macrorrhizos Group), and (radiately) divided leaves are found in the allied relict Seychelles endemic genus Protarum Engl. (Hay & Mabberley, 1991). Seedling leaves, where known, are peltate with partially to completely connate posterior lobes, a condition which may or may not persist into various stages of subadulthood and which may be retained altogether in the Cuprea Group. Alocasia reversa is variable in this respect and may have all peltate leaves, no peltate leaves or mixed states as adult plants. In most, if not all species, the rhizome produces at or below soil level a number of short slender branched or unbranched stolons terminating in more or less globose cormels. These remain dormant for protracted periods, often until the stolons that bear them have decayed. Conditions that stimulate either their production or their release from dormancy are unclear, and while cormels are often produced in large quantities, the plants are typically solitary, though very dense, apparently clonal, populations may occur. Except in the most diminutive species where the inflorescence is solitary (e.g. Alocasia minuscula), the vegetative module is terminated by i) wy) =) Gard. Bull. Singapore 50(2) (1998) a synflorescence composed of pairs of inflorescences forming bimodular synflorescence subunits (Fig. 2). Each consists of a cataphyll subtending a terminating inflorescence (i.e. peduncle with spathe and spadix) and a second inflorescence arising in the axil of the cataphyll and itself subtended by a bicarinate prophyll. A relay axis develops from the axil of the leaf (or leaf homologue) immediately below the first cataphyll of the bimodular subunit. It bears first a bicarinate prophyll and then, depending on whether or not the relay module is initially vegetative, a foliage leaf (which may or Le ey ~ Coady i oe Tite as aaa / Ph NX ale Kak U iS A oS ¥ \ Q : \\ \ \ \ , ¥ \ V \ I \ l ‘O / a / Sa Zieh a ph tery TW Reh ay ' ms ’ \ Y 4 Yh i kK, g/ \ V, { f) : Vy { J 4 J ) ' ; : / if Sarr: 6 4 ~ * at ¢ ae SS Nx a \ F Figure 2. Schematic representation of synflorescence construction in Alocasia Left: synflorescence subunits of cataphyll, inflorescence, prophyll and inflorescence (contained in dotted outlines), interspersed with foliage leaves. Right: synflorescence subunits repeated - with the foliage leaves substituted by cataphylls - making a compound synflorescence, followed by resumption of vegetative growth. /= foliage leaf; |= cataphyll; = inflorescence; ’ = prophyll. Note that in the two-dimensional diagrams, some prophylls are followed by leaves or cataphylls that appear superposed and all other organs appear distichous. This is not the case in life, where all organs are produced in three-dimensional phyllotactic spirals. Organ numbers within the dotted lines appear constant throughout the genus (except in species with solitary inflorescences); with respect to outside the dotted lines, a prophyll is a constant feature at the beginning of modules, but additional cataphylls may be present in either case between the prophyll and the first cataphyll of the synflorescence subunit. Alocasia in West Malesia and Sulawesi 21 may not be preceded by one or more vegetative cataphylls) or another bimodular synflorescence subunit. If a foliage leaf has been produced, the module may continue to produce foliage leaves and the flowering episode has consisted simply of two inflorescences - the usual case, for example, in A. reversa. Alternatively, the foliage leaf may be followed immediately by another pair of inflorescences and another relay axis with a single foliage leaf and so on, so that the flowering episode consists of a compressed sympodium of bimodular flowering units displaced into physiognomically lateral positions and interspersed with foliage leaves, as in A. macrorrhizos. If, however, the first relay axis is not initially vegetative, a second pair of inflorescences follows upon the first, and a third and fourth and so on, each arising from the axil of the cataphyll subtending the previous subunit. Eventually the flowering episode ceases and the last relay axis bears a foliage leaf, which emerges from the centre of a larger or smaller sympodial cluster of inflorescence pairs - up to about 20 in robust species such as A. sarawakensis. After rapid resumption of vegetative growth, the stem may bear a ring of physiognomically lateral infructescences below the leaf crown of the new vegetative module, e.g. in A. robusta. The spathe is divided into a convolute thicker lower portion - the ‘lower spathe’ - housing the female zone of the spadix and persisting into fruiting, and a thinner, ephemeral open limb (this part is also convolute and persistent in some East Malesian species of the Xenophya Group). The two portions of the spathe are differentiated by a constriction so that the lower part is globose to ovoid. The spadix, as is the general case in Araceae, is protogynous and at the time of stigma receptivity the spathe constriction loosens, providing pollinators access to the pistils; simultaneously the inflorescence may emit a detectable scent - highly fragrant to an odour of decay - which appears to be produced from the inside of the lower spathe and not from the floral organs themselves (e.g. A. alba, A. robusta) or from the appendix (e.g. East Asian A. odora (Lodd.) Spach). At this time the spathe limb is generally erect. At the end of female anthesis, the spathe constriction closes and grips the spadix and scent production ceases. There is a sterile zone between the male and female zones of the spadix and typically the spathe constriction is level with this so that after female anthesis the female zone is isolated from the male zone. Male anthesis then occurs. The pollen is mealy and drops to collect between the lip of the limb and the spadix or in a trough-like annular fold that has developed at the base of the limb. Pollination has not been observed in detail (but see van der Pijl (1933) who described pollinator attraction by deceit in A. puber), though one might speculate that in order to leave the lower spathe chamber, insects must force their way between the spadix and the spathe constriction 232 Gard. Bull. Singapore 50(2) (1998) and pick up pollen as they leave. However, while many species have the sterile zone of the spadix and the spathe constriction aligned, this is not universal. The Scabriuscula Group, for example, usually has part or even all of the male zone contained within the lower spathe chamber. What the implications of this are for pollination mechanisms and breeding systems is unknown. The female zone of the spadix consists of naked pistils. The sterile zone, or interstice, is partly or entirely covered with truncate neuter organs (synandrodia), which often but not always appear to be of two types. The lower whorl(s) (with respect to the spadix) may be composed of smaller, often more prominent structures than the upper ones and they commonly react differently (remaining white) in alcohol to the wider upper ones which closely resemble the male flowers except for the absence of pollen thecae (Plate 1). This differentiation of the neuter organs is much clearer os { Plate 1. Inflorescence structure in Alocasia A. Whole spadix of (extra-Malesian) Alocasia odora (Lodd.) Spach [RBG Sydney Acc. No. 940137, voucher NSW], ca. 25 cm long, showing, from the base, female zone, large sterile interstice, male zone of regularly hexagonal synandria and terminal appendix; lower spathe containing female zone and most of sterile interstice, constriction corresponding with upper part of interstice and limb subtending the male zone and appendix. B. Detail of lower part of same spadix showing interstice with basal free staminodial neuter organs, then connate staminodial organs and upper synandrodial neuter organs. C. Spadix of A. principiculus [Hay et al. 12162|,ca.5 cm long, with reduced sterile interstice of a few lax lower staminodial neuter organs and above, whorls of sterile synandria; remaining fertile synandria with pollen thecae visible as dark dots; both fertile zones contained within the lower spathe chamber. Pollen thecae not visible in A. odora as they are overtopped by the synconnective. Alocasia in West Malesia and Sulawesi 233 in Alocasia odora, and sometimes in A. alba, where the lowermost neuter organs are not in connate groups, but instead partially encircle the uppermost pistils clearly in the positions of staminodes. The next whorl (with respect to the spadix) of neuter organs consists of united ‘staminodes’ with a central hole, seemingly where the pistil would be. There is then one or more similar whorls followed by an abrupt transition to structures resembling sterile synandria. This situation closely resembles that in the relict Protarum, which differs in having all the pistils regularly subtended by staminodes. Comparison with Protarum suggests that the organs of the sterile interstice in Alocasia are differently derived in the lower and upper parts of that zone - in the lower part by connation of staminodes, in the upper by sterilisation of synandria. The rhombohexagonal synandria - fertile male flowers - are generally 4—6-merous and consist of connate truncate stamens. The body of the male flower is here termed the synconnective and the vertical pollen thecae are attached throughout their length to its flanks. Typically the thecae reach the top of the synandrium and open through apical pores. However, in some species the synconnective is expanded over the top of the thecae which release pollen from apical slits into the spaces between the synandria. The upper part of the spadix forms a well-developed sterile appendix, which is at least sometimes thermogenic (as may be the male zone). The appendix surface is occasionally smooth, but is more usually sinuously, longitudinally and finely channelled - apparently formed of irregular elongate compressed synandrodia. After anthesis, only the female part of the spadix and the lower spathe remain, the rest rotting and falling away. As the fruits develop and expand, the peduncle generally elongates and the lower spathe enlarges, sometimes becoming conspicuously coloured (e.g. A. balgooyi A. Hay). When the fruits ripen, the fruiting spathe dehisces to reveal them, analogous to arillate seeds in a capsule. The fruits are orange to red, odourless as far as 1s known, fleshy and contain one to several seeds a few millimetres in diameter. Distinguishing Alocasia from Colocasia Traditionally, these two genera, which are undoubtedly closely allied and frequently confused with one another, have been separated on the basis of ovule number and placentation - many ovules on parietal placentas in Colocasia, few on basal placentas in Alocasia (e.g. Mayo, Bogner & Boyce, 1997: 90). These rather academic states are not really of practical use in field identification. However, they translate in the fruiting plants into i) os) pl ewe Gard. Bull. Singapore 50(2) (1998) markedly different dispersal syndromes apparently (though not observed in West Malesia) involving birds in Alocasia, in marked contrast to the mammal dispersal syndrome of Colocasia where the fruits are smelly and inconspicuously coloured with many tiny seeds in slimy mucilage (see Hay, 1996). In respect of synflorescence architecture, Alocasia may be readily distinguished from Colocasia by its bimodular synflorescence subunits. Inflorescence multiplication in Colocasia is achieved in such a way that the whole synflorescence is equivalent to one bimodular unit in Alocasia. Where the inflorescence terminating the vegetative module has only one further inflorescence in the axil of its subtending cataphyll in Alocasia (with the synflorescence being built up by relay axes), in Colocasia the second inflorescence has a third in the axil of its prophyll and so on up to ca. 8 in Colocasia gigantea. The relay axis in Colocasia is vegetative and thus the whole synflorescence is displaced to a quasi-lateral position on one side of the shoot. Foci for Further Study The iterative taxonomic process in the genus as a whole needs additional data - from further collections throughout the range but especially from Kalimantan, Aceh (Sumatera) and Sulawesi, and from additional data sources, such as macromolecular analysis. Three areas stand out as potentially fruitful subjects for intensive analysis - the problem of circumscribing two species groups and of developing biological species concepts in the genus. First, the Alocasia longiloba complex: this is treated here as an ochlospecies - a taxon where locally discrete entities coexist but globally merge. Such taxa are being recognised under whatever label with increasing frequency in the Malesian flora. Whether this example is ontologically ‘real’ or whether the term is a smoke screen for the taxonomic ‘too hard basket’ (cf. Gentry, 1990), the A. Jongiloba complex is extremely interesting not only because of the enormous amplitude of its apparent continuum of variation, but also because of the large and fragmented land area, which the threads of the continuum pervade. On present understanding, forms may discreetly coexist in Sabah, for example, but merge in Sarawak, or discretely coexist in both Borneo and the Malay Peninsula, but merge in Sumatera. The precise nature and history of these patterns, uncovered at geological, ecological, morphological and molecular levels would doubtless provide valuable insights to speciation processes. Second, the Scabriuscula Group: confined to Borneo, this group tae emai Alocasia in West Malesia and Sulawesi aad appears to be in active and recent speciation. There are two wide-ranging highly variable species, A. scabriuscula and A. princeps, rather narrowly differentiated from one another, each with a small set of diminutive, geographically confined and sometimes edaphically specialised segregates. Again, how the elements are related and what processes and historical events may be driving speciation and maintaining their differentiation are intriguing questions relating to the evolution of diversity in Borneo. Third, the development of biological species concepts in the genus: Alocasia inflorescences are structurally and behaviourally complex and next to nothing is known in detail about how they work, beyond observations of spathe behaviour during the phases of anthesis (see above under ‘Structure’). Mechanisms for biological species differentiation may reside in some or all of the following parameters (or indeed others) and their impact on pollinators and breeding systems: e differing flowering times e differing synflorescence architecture differing colours differing odours differing patterns of odour production during anthesis differing sources of odour in the inflorescence differing patterns of thermogenesis during anthesis (at least partly independent of odour production) e differing sites of thermogenesis ¢ corresponding differing patterns of infra-red radiation and, perhaps, visibility e differing proportions of staminodial and synandrodial neuter organs in the insterstice e differing proportions of the fertile male zone within the lower spathe chamber. Now that a preliminary taxonomic framework exists, the way is open for those resident in the region to be making comparative systematic studies of reproductive biology in the genus. A Note on Types Several types designated in this paper consist of specimens occupying more than one herbarium sheet or of paintings executed on more than one sheet. The ICBN clearly defines types as specimens (or images) and not sheets, bottles or any other incidental object whose status as an entity is defined merely by mechanics of curation and not by the biological nature of the specimens concerned. Many specimens of Araceae, and, of course, 236 Gard. Bull. Singapore 50(2) (1998) many other taxa, such as palms, are each preserved in numerous sheets, bags, bottles or combinations of those. This is a necessity arising from the size of the parts of many of the species, which are quite incompatible with standard herbarium methods of preservation based in the neatness of temperate twigs and wildflowers. While there may be some good reason for preferring that types be single objects, it would plainly be ludicrous in plants where only a fragment of a whole leaf can be mounted on a single herbarium sheet or where leaf and inflorescence require separate accommodation, to elevate one fragment of the same specimen to higher status than another or to altogether disqualify a complete and maximally informative specimen from serving as a type. It is beneficial to indicate the number of sheets, so that it is clear how many sheets should be examined. ALOCASIA Alocasia (Schott) G. Don in Sweet, Hort. Brit. Ed. 3 (1839) 631; Schott, Oesterr. Bot. Wochenbl. 2 (1852) 59; Migq., Fl. Ind. Bat. 3 (1855) 205; Schott, Syn. Aroid. (1856) 44; Schott, Prodr. Syst. Aroid. (1860) 144; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 497; Hook. f., Fl. Brit. Ind. 6 (1894) 524; Engl. in Koord., Meded. Lands Plantentuin 19 (1898) 299; Ridl., Mat. Fl. Mal. Pen. 3 (1907) 16; Koord., Exkurs.-Fl. Java 1 (1911) 261-261; K. Krause in Engl., Pflanzenr. 71 (IV.23E) (1920) 71; Merr., J. Straits Br. Roy. Asiat. Soc. special number (1921) 104; Koord., Fl. Tjibodas 6 (1922) 36; Ridl., Fl. Mal. Pen. 5 (1925) 97; Henderson, Mal. Wildfl. Monoc. (1954) 224; Backer & Bakh.f., Fl. Java 3 (1968) 118; A. Hay & Wise, Blumea 35 (1991) 503; nom. cons., non Alocasia Necker (1790), see Nicolson (1963) - Colocasia Schott sect. Alocasia Schott, in Schott & Endl., Melet.Bot. (1823) 18. - Type: Alocasia cucullata (Lour.) G. Don. (Arum cucullatum Lour.). [Ensolenanthe Schott, Bonplandia 9 (1861) 368 - no binomials were proposed]. Xenophya Schott, Ann. Mus. Lugd.-Bat. 1 (1863) 124; Nicolson, Blumea 16 (1968) 116. Type: Xenophya brancifolia Schott (‘brancaefolia’) [= Alocasia brancifolia (Schott) A. Hay]. Schizocasia Schott ex Engl., Bot. Jahrb. Syst. 1 (1880) 186; K. Krause in Engl., Pflanzenr. 71 (I1V.23E) (1920) 115; Bunting, Baileya 10 (1962) 112. Type: Schizocasia acuta Engl. [ Alocasia brancifolia]. Panzhuyuia Z.Y. Zhu, J. Sichuan Chinese Med. School 4 (5) (1985) 49 - Alocasia in West Malesia and Sulawesi 237] Type: Panzhuyuia omeiensis Z.Y. Zhu [= Alocasia cucullata (Lour.) G. Don]. Massive, sometimes arborescent, to small erect, decumbent or creeping terrestrial or lithophytic herbs with irritant juice; stems sympodial, sometimes bearing multiple cataphylls; /eaves glabrous to scabrid or pubescent, solitary to multiple; petiolar sheath persistent to deliquescent; leaf blade simple, deeply pinnatifid to entire, sagittate to hastate to rarely almost lanceolate and then basally auriculate, peltate or not, sometimes strikingly coloured and/or bullate, membranous to strongly coriaceous or subsucculent; primary lateral veins pinnate, usually with conspicuous glands in their axils on the lower leaf surface; secondary venation reticulate, arising along the primary veins and costae, often uniting between the primary veins to form interprimary collective veins, or these ill-defined or absent; inflorescences sweet- to foul-smelling, rarely solitary, usually in pairs orientated parallel to the circumference of the stem, the pairs sometimes in series interspersed with leaves (with inflorescences appearing lateral) or interspersed with cataphylls (with inflorescences appearing as a terminal cluster); peduncles usually short or hidden within subtending leaf sheath or cataphyll, occasionally subequalling the petioles; spathe constricted; the limb persistent to deciduous, variably coloured purple to green, yellow or white, sometimes spotted or streaked; spadix stipitate or not, shorter than to subequalling the spathe; female zone free or sometimes partly adnate to the spathe; pistils naked; ovaries unilocular to (usually incompletely) plurilocular; stigma button-like to stellate, sessile or not; male and female zones separated by a sterile interstice of at least one (very rarely incomplete) whorl of sterile male flowers (synandrodia), more usually of several whorls, the lower morphologically and ?physiologically differentiated from the upper, the interstice usually but not always attenuate and corresponding to spathe constriction; male zone of mostly 4-6-sided male flowers composed of united stamens (synandria); appendix well developed, pointed to blunt, smooth to somewhat rugose; fruits mostly red to orange berries, contained within the persistent spathe base; fruiting spathe dehiscing at maturity; seeds ca. 3-5 mm diam., albuminous. Distribution: Indomalesia, ca. 65 species with A. macrorrhizos (L) G. Don now naturalized pantropically; in Malesia 57 indigenous species and one widely cultivated (A. cucullata); 31 species in West Malesia and Sulawesi. Habitat: Primary and secondary forests, early regrowth and open swamps, sometimes lithophytic, rarely rheophytic; primarily in everwet conditions, but some species tolerant of quite strong seasonality; predominantly in the lowlands, extending from sea level to lower and mid-montane zones. i) 2 ioe) la. 1b. 2a. 2b. 3b. 4a. Ab. Eee Sb. 6a. 6b. Ja. Tb. Sa. 8b. Gard. Bull. Singapore 50(2) (1998) Key to Species and Species Complexes Leaf blades:not peltate ic... 5.is.0scssgodietecustes tonne tie eeeiyren nee are ae eee zg Leaf blades distinctly (shallowly to completely) peltate in adult plants Secondary venation distinctly prominent abaxially and forming well- defined interpnimary collective veins: 20. een, eee 3 Secondary venation not prominent abaxially, or, if prominent, then not forming well-defined interprimary collective vein .............ceeeeeee 5 Petiole and abaxial leaf blade pubescent (Java, ?Sumatera, Malay Pominisiala) 40 ee See ee ee ne 1. A. puber Petiole and abaxial leaf blade glabrous in adult plants (beware juvenile AS SOTOWURETISIS) <5. Seeiedathcnns nobaceas an Ae ee 4 Lower spathe green; spathe constriction level with sterile interstice FAVA Ys cvs ccane'sc oolagaun saeioenel estar aa e teem ee rene teers anne 18. A. alba Lower spathe ivory, marked red-purple; spathe constriction within male Zone (BOMEO) newex ceca ee eee eee 2. A. sarawakensis Leaf blade membranous, often immense, abaxially waxy-glaucous (Bormeo, Natuma tslands e250) se enaceey nae nawin aera 3. A. robusta Leaf blade of various sizes and textures, not waxy-glaucous (though soinetimes abaxially srey-oneem). .. 4.) yao ecu cee ee 6 Leaf blade thickly coriaceous; posterior costae not naked in the sinus; all secondary venation obscure in the dry state; petiole finely pubescent f SEM ANVESED iu asin gsi nije sears A aera tre tia eat areecaee te mea ee Mei ae 23. A. celebica Not. this combinatidn..!.,...d4alek ashi posnenay oak een eee toe eee A Male zone of spadix completely exserted from lower spathe chamber (not Borneo except.A. macrorrhizos)ipiiaidiy oes ek ee 8 Male zone of spadix partly or wholly within lower spathe chamber (Bomieo) .cciesseieia dete WARE eee ee eee 13 Thecae of synandria not overtopped by synconnective (pores visible on surface of male zone); stigma stellate (with pointed lobes) (S. Malay Peninsula,’ Sumatera) 2.00... ncaa 22. A. longiloba ‘denudata’ Thecae of synandria overtopped by synconnective; stigma not lobed or lobes rounded {200.0.000)3 R00 a ees tek ee ee 9 Alocasia in West Malesia and Sulawesi 239 9a. oF 10a. 10b. hha: 11b. ia: 12b. 134: 13b. 14a. 14b. ioav 15b. 16a. Posterior costae (usually) with lamina to the sinus; spathe limb coriaceous; synandria very numerous and small (ca. 1 mm diam., dry); iruiting spathe (usually) red (Sulawesi), ................... 19. A. balgooyi Posterior costae naked in the sinus; spathe limb more or less membranous; synandria ca. 2 mm diam. or more (dry); fruiting spathe ie MEME INNER tn ict RN eI EUR ails diotebia alts Oaibield Uae Lo bade « 10 Inflorescence pairs in a central cluster; leaves distinctly leathery coriaceous, either (rarely) purple throughout or with the petiole apically purple (Malay Penimsula, Sumatera)! 0400. il yelecs ee. 17. A. inornata Inflorescence pairs interspersed with foliage leaves; leaves more or [ese mcm Dranous and mot inus Coloured wii see ial Secondary venation very fine but distinctly darker than abaxial ground. colour, forming interprimary collective veins (Sumatera) JO TG Ge GULL Ree a adie Aad a eee 21. A. arifolia Massive plants with leaf blades ca. 80 cm long (or more); primary venation not gathered into a distinct intramarginal vein (widespread in assoctationi with People)! /iins sisi sees hola. 16. A. macrorrhizos Smaller plants with leaf blades ca. 35 cm long; primary venation running into a distinct intramarginal vein (Java) ............... 20. A. flemingiana Leaf blade narrowly to broadly ovato-sagittate, nearly always stiffly (Oe ie ab SAP rated [21 aan a IU eh 14 Leaf blade hastato-sagittate, triangular in outline, mostly rather thinly TG FCI Gilat AND toon Bie 0 are Se IU on UE Se Gar El me Sere ao eee ae ae ea 18 Adaxially leaf blade grey-green and distinctly dark green about main OU ae loos. hy elec v ELA eto Re Ee? oe an ee eee i) Abaxially leaf blade purple; anterior costa with ca. 6 primary lateral veins on each side, with conspicuous subsidiary veins (Sarawak, SSIS TUT CL ANAM Ane a ne oe Re 31. Alocasia sp. A Abaxially leaf blade not purple; anterior costa with 2-3 primary lateral veins on each side; subsidiary veins absent (S. Sarawak) LOTT Re cae VERB G SRI CR A AOE UE ee ds Ee 14. A. reversa Inflorescence pairs solitary and secondary venation adaxially impressed OO STD TSR LT. ie T6115) C1 ne ead ee 5. A. reginae 240 16b. 17a. 17b. 18a. 18b. 19a. 19b. 20a. 20b. Pate 21b. 22a Gard. Bull. Singapore 50(2) (1998) Inflorescence pairs clustered or if solitary then secondary venation NOL MBPTESSEM 5.0.0620 705s OAL A ee 17 Posterior lobes ca. half or more the length of the anterior; blade stiffly leathery (S. Sarawak - lithophytic on or terrestrial in close association with limestone) sis cee cua ts eae ncek ee receeee ee aes 8. A. ridleyi Posterior lobes less than to ca. half the length of the anterior; blade thickly coriaceous to subsucculent; (widespread in Borneo - terrestrial and not especially associated with limestone) ......... 4. A. scabriuscula Wenrestrgal 900.518 Shue Gu NO CR ee 19 Lithophytic.on limestone s33.)040 Us ee ee 20 Petiole mottled with wavy oblique zones of dense brown lines, occasionally scabrid; spathe dusky brownish mauve, the limb darker; lower spathe narrowly ovoid; limb mostly narrowly lanceolate (Sabah) aio osiged Was carpe Sotovate oa ide bral RANR GAR O0 OE EVN SE AS ae eae ga ae 10. A. wongii Petiole variously and more or less haphazardly marked with lines and/ or dots, smooth or occasionally faintly bumpy (glands), but not scabrid; spathe mostly ivory to yellowish ivory, variously marked or not with pink to purple, and/or purple-margined; lower spathe broadly ovoid; limb more or less oblong (widespread in Borneo) ...... 9. A. princeps Male zone of spadix completely within lower spathe chamber; leaf blades distinctly grey-green adaxially (Sabah, E. Kalimantan) wee Ma tc pm bend ducati w'e wicts Scie lelopta aha ep mene eT ele gne 13. A. principiculus Male zone of spadix partly exserted from lower spathe chamber; leaf blades dark ito bright sreen adaxially (Sabah),.....2.525.2.0 ce tse mee Za Leaf blade bright green adaxially; inner side of posterior lobe ovate; male zone more or less adjunct to female zone or interstice short, not attenuate, formed of 1-2 whorls of synandrodia resembling synandria sgdipitidgie de cp uine Wek We wie'y even ba cie'veldammateenaeelth, ace Ane ee eee 12. A. puteri Leaf blade dark green adaxially; inner side of posterior lobe elliptic to narrowly ovate; interstice elongate, partly naked, with neuter organs resembling staminodes below and resembling synandria above seip DUBE Songs orecetun dee inte dosed de Joh sean gt one aie MEER st nena 11. A. pangeran Leaf blades more or less membranous and pendent, often solitary or only 2-3 together, often adaxially dark green with whitish major veins (sometimes adaxially concolorous), often purple-backed, shallowly to deeply peltate; stigma stellate with pointed lobes; insterstice Alocasia in West Malesia and Sulawesi ‘ 241 22b. 23a; 23b. Dag, 24b. 25a. 2b. 26a. 26b. IN 2/0. 28a. 28b. 29a. 290: corresponding with spathe constriction and male zone completely Fare pe CINI oe era ATED ress EVN ioe 8d Dad ke cieblda «dL edgdeneeedidiiace.s 23 Leaf blades variously coriaceous, pendent. or not, few to several together, mostly not variegated, deeply to almost completely peltate; stigma not lobed or lobes rounded; interstice and part or all of male zone within lower spathe chamber (except A. kerinciensis) .......... 24 Petiole glabrous; spathe limb greenish white (widespread) AC Re SUR eee EPR ey nee Me A Obs a 22. A. longiloba complex Petiole minutely and densely pubescent; spathe limb purple-black PASM MINES Hi ht ines eMC ate RG Educ 24. A. suhirmaniana Leaf blades metallic greenish brown adaxially, bullate between primary veins; lower primary veins diverging at first at more than 90° (Sabah) PN ea ies x Bb sails seiti sche adieu paar aUAatiiveclad ai-bigiais «> it 30. A. cuprea fare eae IRIOR TEV AOL OE 200 ads an eases mune ber HEME aif fil aidaieeieie gene nin a 2D Adaxial leaf surface rugose with the tertiary venation raised (Sabah) Re OPE Mn icy Solid ena kL hUaM aan ume bony CAsdhea wea? 7. A. melo Adaxial leaf surface smooth or with secondary venation impressed ooh ts Se ee IH rane Teen Ome | er 91 The ie a meme ae 26 Adaxial leaf blade dark green with whitish impressed primary and secondary venation; spadix with appendix reduced (origin unknown) ee CORR Ly one TAREE TORRY MOREE MUD PEE OeRtant anT A DEO a ae 6. A. reginula Leaf blade not variegated, or if variegated then main veins and neighbouring blade darker than the rest; appendix well developed Beret See Paes Thais) ch cast acl Was a Sadia SR EMU LINE ha tole uefa) sd mbied sais app pa Blade broadly ovate to ovate, with the base rounded; male zone exserted from lower spathe (S.W. Sumatera) ......... 29. A. kerinciensis Blade broadly to narrowly elliptic, with the base cuneate; male zone Mitmd te lower spathe BOrme@) .... 26.6.6) 28 Loess. 28. A. peltata Primary lateral veins numerous, 8-10 on each side of midrib; secondary venation striate; (Sarawak - in peat swamp forest) PRG ee UCAS SOLE AMER hae nC ARRAS eke 27. A. minuscula Primary lateral veins much fewer; secondary venation clearly colocasioid, but not forming interprimary collective veins ............ 30 242 Gard. Bull. Singapore 50(2) (1998) 30a. Leaf blade (ovate to) narrowly ovate to oblanceolate; connate posterior lobes attenuate; male zone of spadix within lower spathe chamber 30b. Leaf blade oblong-elliptic; connate posterior lobes cuneate; male zone only partly included within the lower spathe chamber (Sarawak - on limestone} 272 ete a Se 32 31a. Leaf blade (coriaceous to) thickly coriaceous to subsucculent, 14 x 6 to 34 x 12 cm; spathe ca. 6 cm long. (Malay Peninsula, usually above AG) Up ti MOM MentINbires Ure, eeangbeiaReeabeb a SA Tata phuderagsiecaay by it 25. A. perakensis 31b. Leaf blade coriaceous, 9 x 2.2 — 18 x 6 cm; spathe ca. 4 cm long; (N.W. Bormeo; usually below 1000 my! cen nay Se aes 26. A. beccarii 32a. Leaf blade dark green throughout and somewhat darker around mid- veins; inflorescences to ca. 6 together; stigma mostly tri-lobed (Niah, SABER WEIS) 60 hie ois cu acdactocded con pene ee ert naa ent any a te ee 15. A. venusta 32b. Leaf blade grey-green and dark blue-green around veins; inflorescences solitary to paired; stigma mostly bi-lobed (S. Sarawak) ... 14. A. reversa Puber Group Species 1—3 Massive arborescent or decumbent plants; /eaves several together, often hairy; inflorescences numerous in succession not interspersed with foliage leaves, usually with horizontal elliptic often dark red glands on the lower spathe; male zone of spadix held partly within lower spathe chamber. | Note: The group includes the three west Malesian species described below and Philippine A. maquilingensis Merr. The species are elements of gap phase, secondary forests and open swamps. 1. Alocasia puber (Hassk.) Schott Alocasia puber (saepe ‘pubera’) (Hassk.) Schott, Syn. Aroid. (1856) 47; Mig., Fl. Ind. Bat. 3 (1856) 209; Schott, Prodr. Syst. Aroid. (1860) 146; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 500; Koord., Exkurs.-FI. Java 1 (1911) 261; K. Krause & Engl., Pflanzenreich 71 (IV.23E) (1920) 79; Koord., Exkurs.-Fl. Java 4 (1923) 197, fig. 398; Ridl., Fl. Mal. Pen. 5 (1925) 99; Backer & Bakh. f., Fl. Java 3 (1966) 119. - Colocasia pubera Hassk., Cat. Bog. (1844) 302. - Neotype: Indonesia, Java, Djampangs, Cicurug, 1914, Backer 17192 (BO, designated here). Alocasia in West Malesia and Sulawesi 243 Alocasia margaritae L. Linden & Rodigas, Ill. Hort. 33 (1886) 155; Anon., Kew Bull. (1888) 92; Engl. & K. Krause, Pflanzenr. 71 (I1V.23E) (1920) 92. - Type: L. Linden & Rodigas, Il. Hort. 33 (1886) t. 611. Epitype: Java, Desa Ciomas, Kampung Pabuaran, Serang, West Java, 12 Dec 1997, Yuzammi 297017 (BO, NSW, designated here - see below). Alocasia ovalifolia Ridl., J. Straits Br. Roy. Asiat. Soc. 41 (1904) 47; Ridl. Materials Fl. Mal. Pen. 3 (1907) 18; Engl. & K. Krause, Pflanzenr. 71 (TV.23E) (1920) 109; Ridl., Fl. Mal. Pen. 5 (1925) 99. - Type: Malaysia, Perak, Sungei Larut, July 1888, L. Wray Jr. 2457 (SING, lecto, selected here; K, isolecto; see discussion below). ?Alocasia crassinervia Engl., Pflanzenr. 71 (IV.23E) (1920) 82. - Type: Indonesia, cult. Hort. Bogor, Jan/Feb 1906, A. Engler 4101 (B!, holo) - see note under A. sarawakensis. [Caladium pubigerum Bl., mss| Robust to massive herb; stem erect to decumbent, to ca. 10 cm diam.; leaves several together; petiole to ca. 1.1-1.5 m long, sheathing in the lower ca. !/3, green to dark red, sparsely glandular, it and abaxial lamina venation sparsely to densely hairy; hairs straight, short, ca. 0.5 mm long, colourless fresh, becoming yellowish brown when dry; blade usually sagittate, occasionally broadly ovato-sagittate, to ca. 80 cm x 70 cm; anterior lobe ca. 60 cm long, widest at about the base; anterior costa with ca. 10 primary lateral veins on each side, diverging at ca. 50°, with rather conspicuous glands in their axils; primary veins distally often bearing subsidiary veins (with glands in their axils) running the same course as the secondary venation; secondary veins rather prominent abaxially, numerous and close- spaced, running into well defined interprimary collective veins; posterior costae diverging mostly at ca. 90°, naked in the sinus for up to 4 cm; posterior lobes ca. 30 cm long, more or less triangular, rarely rounded; inflorescences appearing as clusters in the centre of the leaf crown, up to ca. 14 together not interspersed with foliage leaves; peduncle hardly exserted from subtending bracts; spathe ivory white, suffused purple, especially near the base of the limb margins, with scattered purple horizontal elliptic glands especially near the junction of spathe and peduncle, 9-18 cm long; lower spathe narrowly ovoid, somewhat angular in cross section, differentiated from the limb by an initially rather weak constriction about 1/3 of the way from the base of the spathe and corresponding to about midway along the length of the male zone of the spadix; limb more or less cucullate at first, leathery-membranous, broadly lanceolate with the tip 244 Gard. Bull. Singapore 50(2) (1998) obtuse, sometimes conspicuously apiculate, initially sharply reflexed at the base, forming an annular trough, the rest erect, then entirely reflexed; spadix slightly shorter than the spathe, ca. 6-15 cm long, very shortly stipitate; female zone 1-2 cm long, subcylindric; ovaries very pale green, subglobose, close-packed, ca. 1.2 mm diam., style short, slender, ca. 1 mm long; stigma ivory, cap-like, weakly 3—-4-lobed; sterile interstice hardly more slender than the fertile zones, ca. 2 whorls of flat white synandrodia ca. 2 mm diam.; male zone yellowish ivory, half within and half exserted from the lower spathe chamber, ca. twice as long as female zone; synandria mostly rhombohexagonal and ca. 2 mm diam., sometimes united into irregular horizontal bands; synconnective impressed, not overtopping the thecae; appendix pale apricot, about half the length of the spadix, about as thick as the male zone, slightly narrowed at its base and distally tapering to a pointed tip; fruiting spathe ca. 4 cm long. Habitat: In open swampy areas, and wet places in open forests, sea level to ca. 1000 m alt. Distribution: West to central Java, ?southern Sumatera, Peninsular Malaysia. Notes: 1. No Hasskarl material has been located that might be the type of A. puber. In the protologue, Hasskarl cited Arum sylvestre Rumph. (Herb. Amb. 5: 310, t. 107) as a synonym, but with doubt indicated by ‘an’. Rumphius’ concept of Arum sylvestre included more than one ‘species’, though only one was illustrated. The plate, which does not appear to be of a Javan plant at all, bears insufficient resemblance to A. puber in the sense of Schott and subsequent authors, to warrant using it as the type. It gives no indication of the characteristic features of pubescence on the leaves and horizontal red markings on the lower spathe that Hasskarl mentions for C. puber and, which indicate that Hasskarl’s and Schott’s concepts are almost certainly of the same species. Moreover, it shows a configuration of inflorescences paired amongs the leaves, which is also not characteristic of this species. It is probable that Rumphius’ plate is of the east Malesian Alocasia aequiloba N.E. Br. Hence there is no alternative but to designate a neotype. There are very few fertile Javan collections to select from, Backer 17192 being the most complete. 2. Alocasia margaritae L. Linden & Rodigas was described from Javan material cultivated in Europe. No herbarium material has been located. The illustration in the protologue is of a sterile, immature plant, but the description includes reference to the puberulent petioles typical of A. puber and the depicted leaf shape is not incompatible with that species. A. margaritae appears to be no more than a particularly strongly red-brown Alocasia in West Malesia and Sulawesi DAS coloured variant of A. puber and it is epitypified accordingly to remove all doubt. 3. The identity Ridley (1904) intended for Alocasia ovalifolia is obscured by various muddles. The description in the protologue is scant and sloppy. The leaf is said to have 20 pairs of ‘nerves’, which I understand to be primary lateral veins. I know of no species of Alocasia, with the exception of the gargantuan Bornean Alocasia robusta that has this many. The spathe is said to be up to six inches long, while the dimensions given for its components, the lower part and the limb, add up to four and a half inches. On top of these incongruities, the description records no distinctive features by means of which it could be matched with known Malay Peninsula species. Four syntypes were cited thus: ‘Johore, base of Gunong Panti; Selangor, Tras Route at the 15th mile Pahang Track (Ridley 8487), Ginting Peras, Bukit Kutu; Perak, Sungei Larut (Wray 2457), Larut Hills; Penang, Moniots Road (Curtis).’ All of these are missing from or perhaps misplaced in the Singapore herbarium and are not duplicated elsewhere, with the exception of Wray 2457. That specimen is of Alocasia puber, but it is not annotated by Ridley with any name and by no-one as A. ovalifolia. I suspect therefore that it was cited in error; indeed it was later cited as A. puber in Ridley’s Flora, while other previously cited material remained under A. ovalifolia (Ridley 1925: 99). Other specimens, not syntypes but which might guide interpretation of A. ovalifolia, are equivocal. There are two collections made by Ridley and annotated by him with this name. One is Ridley s.n. (SING) with the locality ‘Thaiping Hills’; this is Colocasia esculenta. The other is Ridley 13392 (SING), collected at Batu Caves in 1908 (but not cited in the Flora). This latter is of acommon Malay Peninsula species (see A. inornata Hallier f.) which seems very unlikely to have been overlooked by Ridley as a species distinct from other Alocasia in the Malay Peninsula. It seems probable to me, in spite of his bad description, his citation of Wray 2457 and his misdetermination of a plant in another genus as A. ovalifolia, that the plant in Ridley 13392 is what Ridley intended A. ovalifolia to be. Nevertheless, the ICBN demands lectotypification from among the syntypes (Art. 9.9). An alternative is not to lectotypify, and simply leave the issue open in the hope that the remaining syntypes turn up one day. However, since the name Alocasia ovalifolia has never been taken up by subsequent authors [Henderson and Furtado, for example, both misapplied other names to this species (i.e. A. inornata) on herbarium sheets, which suggests they had never seen the other syntypes either] and cannot be said to be in current use, there seems to be no pressing need to do other than simply dispose of the name on the basis of the identity of the one syntype that is available. Alocasia ovalifolia is therefore reduced to the synonymy of A. puber. 246 Gard. Bull. Singapore 50(2) (1998) 4. Backer & Bakh. f. (1968: 119) pointed out that the epithet ‘pubera’ is grammatically incorrect, and amended it to puber without elaboration. It is evident that Hasskarl intended the epithet to refer directly to the hairiness of the leaves and peduncles, for in the description he used the adjectival ‘puberis’ (in the ablative plural) referring specifically to those parts of the plant that are hairy. It would appear that he used the word as though it were a ‘group A’ adjective such as glaber, which it is not. Puber (or pubes) is used in mediaeval and classical latin as a noun referring to adolescence, and puber was used in classical latin specifically of plants as an adjective referring to juiciness (which Hasskarl also described in the protologue of this plant, but without using this word). In botanical latin puber (or preferably pubes) could be used as a substantive epithet for hairiness, and puberula as an adjectival epithet for hairy [see Oxf. Lat. Dict., ed. P.G.W. Clare, 1982]. Since Bakh. f. has already chosen one of these alternative corrected forms, I follow him. 5. Alocasia puber is apparently very rare in the Malay Peninsula, having been collected there only four times, and only once at all recently (Chua FRI 26675). Although Javan representatives have been collected more frequently, there is very little fertile material from which to judge intraspecific variability, but the Malay Peninsula element seems to have a rather smaller inflorescence. The collections made by Corner also show rather widely rounded posterior lobes to the leaf blade in contrast to the more typical triangular shape in Java, and it may be that the Malay Peninsula element could be recognised as a segregate subspecies once it is better known. However, it evidently has the same habitat preference as A. puber in Java. 6. A single collection from Sumatera (Praetorius s.n.) determined by Schott as A. macrorrhizos, is apparently of this or a closely allied species, being quite densely hairy on the leaf underside, with rather prominent secondary venation forming well defined interprimary collective veins. The specimen, from Palembang, is sterile Other specimens seen: PENINSULAR MALAYSIA: Terengganu, Besut, Sg. Kemia, foothills of G. Lawit, Chua FRI 26675 (KEP); Terengganu, Kemaman, Ulu Kajang, Corner 30138 (SING); Johore, Mawai Rd., Corner s.n. (SING); Johore, Jason Bay, Mile 4, Block 1, Sinclair 10869 (E, SING); Perak, Sg. Larut, Wray 2457 (K, SING). SUMATERA: Palembang, Praetorius s.n. (L). JAVA: Tjidadapi Tjibeber, Preanger, Cadas Kabang, Bakhuizen 2484 (BO); Blume s.n. (L) & 792 (L); Bidara Tjina, Edeling s.n. (BO); Tjiloewar, nr Bogor, Hallier s.n. (BO); Danau Situgunung, Hay & Yuzammi 14001 (NSW); Djapara Ngarongan, Koorders 34996b (BO, L); Bantam, Lebakkidoel, G. Kantjana, Koorders 41042b (BO); Kuhl & van Hasselt s.n. (L); Djasinga, 45 km W of Bogor, Nicolson 938 (BO); Tjitjadas, Batavia, van Steenis 5364 (BO); Cult. Bogor, Wigman s.n. (BO); Blok Cimanuk, Rawa Alocasia in West Malesia and Sulawesi 247 Danau Natural Reserve, Serang, West Java, Yuzammi 297013 (BO, NSW), Yuzammi 297016 (BO, NSW); Desa Ciomas, Kampung Pabuaran, Serang, West Java, Yuzammi 297007 (BO, NSW), Yuzammi 297017 (BO, NSW); Zollinger 472 (K). 2. Alocasia sarawakensis M. Hotta Alocasia sarawakensis M. Hotta, Acta Phytotax. Geobot. 22 (1967) 159, fig. 6, G-L. - Type: Malaysia, Sarawak, Mardi, along Sungei Melinau, 14 Mar 1964, M. Hotta 1439 (KYO, holo; n.v.) [? Alocasia crassinervia Engl., Pflanzenr. 71 (IV.23E) (1920) 82. - Type: Indonesia, cult. Hort. Bogor, Jan/Feb 1906, A. Engler 4101 (B, holo) - see note below. | [Alocasia puber (‘pubera’) sensu auct. non (Hassk.) Schott: Hotta, Acta Phytotax. Geobot. 22 (1967) 158.] Massive arborescent herb; stem more or less erect, to ca. 15 cm diam., to 70 cm tall; /eaves several together with the blades erect to oblique; petiole to 130 cm long, sheathing in the lower !/3—2/5, pale dull green, very slightly rough, with numerous glands mainly in the sheathing portion, these ellipsoid, ca. 4 mm long, aligned along the axis of the petiole, red at first, later turning yellow, in juveniles often ringed with blackish purple; blades somewhat glossy mid-green above, paler below, glabrous in adult plants, abaxially hairy in juveniles, cordato-sagittate, ca. 90 cm x 80 cm; anterior lobe ca. 60 cm long, with the margins slightly undulate; posterior lobes ca. 35 cm long, rounded, held somewhat above the plane of the anterior lobe; posterior costae diverging at ca. 80-90°, naked in the sinus for ca. 2 cm; primary lateral veins 10-12 on each side of the anterior costa diverging at ca. 45°, their distal portions often emitting subsidiary veins, especially on side facing posterior lobes; secondary veins forming very well-defined interprimary collective veins; costae and primary veins whitish abaxially, green adaxially; primary and secondary venation very prominent abaxially, more or less flush adaxially; glands conspicuous in axils of primary veins and very large at junction of petiole with costae, yellowish green; inflorescences very numerous, to ca. 40 crowded in the centre of the leaf crown from within which the relay axis eventually appears, paired and subtended by somewhat persistent (thence marcescent-deliquescent) ca. 30 cm long lanceolate cataphylls bearing glands; peduncles ca. 30-40 cm long, mostly hidden within the cataphylls, with a few scattered glands, pale dull green, ca. 2 cm diam.; spathe ca. 19 cm long; lower spathe 7 cm x 2.5 cm and somewhat flattened, white with a basal ring of confluent glands, these at first shiny white, becoming purple, the remainder of the lower 248 Gard. Bull. Singapore 50(2) (1998) spathe with scattered ellipsoid glands aligned transverse to the long axis of the spathe and somewhat clustered at about 2/3 of the way up the lower spathe; spathe limb white, to 12 cm long, erect at female anthesis, then sharply reflexed and rolled back at male anthesis, broadly lanceolate, to 5 cm wide, horizontally wrinkled abaxially; spadix to ca. 16 cm long, stipitate for ca. 5 mm, stipe white; female zone 2.5 cm long ca. 1.5 cm wide at base, distally somewhat tapering; ovaries whitish ivory, sub-globose, ca. 1.5 mm diam; style very slender, ca. 1 mm long; stigma abruptly wider than style, + rounded and inconspicuously 2—3-lobed, like the style, ivory; sterile interstice ca. | x 1 cm, hardly attenuate; synandrodia flat-topped, ivory, rhomboid, ca. 3 mm long; male zone 3 cm x 1 cm, partly within the lower spathe chamber; synandria + hexagonal, opening by apical pores not overtopped by synconnective, 6—8-merous, ivory; appendix apricot coloured, 9 cm long, 1.5 cm diam., tapering to a point, the surface covered with horizontally elongate, sinuous staminodes; fruiting spathe white, dehiscing longitudinally; fruits red. Distribution: Endemic to Borneo; in Sabah and Sarawak, with one doubtful record from Kalimantan. Habitat: Common in open swampy places; often seen in roadside ditches; encountered in swampy places in forest as a hairy juvenile; from sea level to ca. 1200 m altitude. Notes: 1. This species is easily distinguished from A. robusta and A. macrorrhizos, which sometimes all occur together and resemble each other in the very large broad leaves and preference for open habitats, by the very prominent venation on the abaxial side of the leaf blade, forming well defined interprimary collective veins. It can be distinguished further from A. robusta by having the posterior costae naked in the sinus and the abaxial side of the lamina not glaucous. However, occasional specimens are intermediate (e.g. Agama & Valera 9887): they suggest that some hybridization may take place where the two species occur together. 2. Alocasia sarawakensis is far more common that the meagre number of herbarium collections would suggest, perhaps because it (and A. robusta) is mistaken for A. macrorrhizos and threrefore not considered worth collecting. Aside from the leaf venation characteristics, it is amply distinguished from A. macrorrhizos by the basically white spathe with red markings, and by the arrangement of the inflorescences in a large central cluster, where the spathe of A. macrorrhizos is green and yellow, and the inflorescence pairs are interspersed with foliage leaves. Moreover, while Alocasia in West Malesia and Sulawesi 249 A. sarawakensis (and A. robusta) are found widely in disturbed places, A. macrorrhizos is hardly ever encountered more than a short distance from human habitation, suggesting that it is not a native Bornean plant at all, while A. sarawakensis and A. robusta are Bornean endemics (the latter also in the Natuna Islands). 3. Alocasia crassinervia Engl. was described from a seedling cultivated at Bogor, said to have been from Borneo. The type consists of a single early juvenile pubescent leaf, which could be from A. sarawakensis. However, it could equally be a juvenile of A. puber, which grows in the vicinity of Bogor. In the event that it could be demonstrated clearly that the type of A. crassinervia is of the Bornean species, I would recommend conserving the name Alocasia sarawakensis. 4. I have not seen the Bornean specimens determined by Hotta (loc. cit.) as A. puber (Hotta 14175 and 14231), but it seems highly probable that these hairy sterile plants are juveniles of A. sarawakensis. The only collection of this species from Kalimantan (Burley et al. 596) is identified with some doubt, as the usually characteristic persistent horizontal glands on the lower spathe appear to be lacking. The specimen is in fruit. The leaf matches this species more than any other known Bornean element. Other specimens seen: SABAH: Semporna, Timbun Mata F.R., Agama & Valera 9887 (K, SING); Kinabatangan Besar, Kori Timber Camp, Cuadra A2144 (all seedlings: BO, KEP mixed - see also A. robusta, K, L, SING); Sepilok, Forest Research Centre grounds, Hay 10010 (SAN, spirit only); Lahad Datu Rd, ca. 10 mi ex jnct Sandakan-Kota Kinabalu Rd, Hay 10013 (SAN); Cult. RBG Sydney Acc. No. 950366 ex Ulu Dusun, Hay 10029 (NSW); Cult. RBG Sydney Acc. No. 950374, Madai Falls, Hay 10037 (NSW); Cult. RBG Sydney Acc. No. 960547 ex Tibau Forest Station, Kinabatangan, Hay 12138 (NSW); Cult. RBG Sydney Acc. No. 960577 ex 2.5 km above Maliau Falls, G. Rara F.R., Hay 12056 (NSW); Cult. RBG Sydney Acc. No. 960597 ex Kinabatangan, Kalabakan Virgin Jungle Reserve, Hay 12017 (NSW); Semporna, Timpun Mata F.R., Mapat R., Keith BNB 7419 (A, KEP, K, L). KALIMANTAN: Headwaters of Sg. Kahayan, 5km N.E. of Haruwu Vill., Burley et al. 596 (KEP, K, SING). 3. Alocasia robusta M. Hotta Alocasia robusta M. Hotta, Acta Phytotax. Geobot. 22 (1967) 159, fig. 6, A-F; A. Hay in Plant Talk 5 (1996) 25, un-numbered photo. - Type: Malaysia, Sarawak, Bintulu, at the foot of Bukit Kana, 22 Nov 1963, M. Hotta 15502 (KYO, holo - n.v., L, iso). Colocasia gigantea sensu auct. non (BI. ) Hook.f: Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178 & Merr., Enum. Bornean PI. (1921) 107. 250 Gard. Bull. Singapore 50(2) (1998) Gigantic arborescent herb to palmiform tree to 6 m tall with clear sap that almost instantly turns orange on contact with air; stem erect to decumbent, 15-30 cm diam. (the base swelling to ca. 40 cm ), smooth; leaves several together; petiole smooth, somewhat glaucous, pale green to pinkish brown, ca. 1.5-3.5 m long, sheathing in the lower 1/3, with scattered flat + circular glands; blade ovato-sagittate, ca. 1.5-4 m long, ca. 80 cm to 2.5 m wide, slightly glossy mid-green adaxially, abaxially waxy glaucous, the margin entire to slightly sinuate; anterior lobe up to ca. 3.2 m long, widest at the base; anterior costa with 10-18 primary lateral veins diverging at ca. 70-80°; secondary venation not prominent, not forming interprimary collective veins; posterior lobes to ca. 90 cm long, rounded; posterior costae diverging at ca. right angles, bearing lamina right into the sinus (in juveniles peltate until leaf blades are ca. 30 cm long); inflorescences ca. (10—)30-40 crowded together in the centre of the leafy crown, subtended by robust broadly lanceolate deliquescent to marcescent cataphylls; peduncles stout, hardly exserted from the cataphylls, ca. 15-20 cm long; spathe to ca. 20 cm long, constricted at ca. 4-5 cm from the base; lower spathe pale greenish ivory with few to numerous shiny colourless to (?0r becoming) dark red horizontal glandular markings, ovoid, ca. 2.5 cm diam.; limb cowl-like, by male anthesis the base sharply reflexed then the rest erect, forming a pollen-filled trough at the bottom, deep purplish pink to greyish ivory faintly suffused pink, ca. 15 cm long x 6 cm wide; spadix somewhat shorter than the spathe, ca. 15-19 cm long, sessile to very shortly stipitate; female zone subpyramidal, trigonous, 2—3 cm long, ca. 2 cm diam at base, tapering to ca. 1.5 cm diam. distally; pistils ivory; ovary subglobose, faintly longitudinally ridged, ca. 1.5 mm diam.; style slender, ca. 0.5 mm long or absent; stigma button-like to faintly 3-4-lobed; sterile interstice slightly wider than female zone, reduced to 1-2 whorls of small yellow synandrodia with the staminodes incompletely fused; male zone ca. 4cm long, much expanded (ca. 2 cm diam.) in the part distal to the spathe constriction, below the spathe constriction ca. 1.2 cm diam; synandria rhombohexagonal, 1.5-2 mm diam., ivory, with the thecae overtopped by the synconnective; appendix dirty pale yellow to ivory, slightly constricted at base, narrower than distal part of male zone, tapering gradually to a fine point; fruiting spathe whitish, sometimes borne quasi-laterally in a ring on the trunk below the leaf crown and then pendulous, the spathe dehiscing longitudinally; fruits reddish orange, rather small, ca. 4 mm diam., mostly single-seeded. Distribution: Widespread in Borneo, though mainly in the northwest, and with one record from the Natuna Islands. Alocasia in West Malesia and Sulawesi 251 Habitat: Wet but well-drained open disturbed places, road sides, plantations, river banks, land slides, canopy gaps in lowland to lower montane forest. Notes: 1. A magnificent arborescent herb with, in the biggest examples, certainly the largest undivided leaf of any herbaceous plant (not counting the habitually tattering leaves of the biggest Musaceae) and challenging the few palms with large undivided leaves. It is strikingly glaucous on the underside. It is common and widespread in northern Borneo at low elevations, and it is remarkable that it was not described until 1967. From a distance it resembles, but is much larger than, A. macrorrhizos, with a rather similar subtriangular ovato-sagittate leaf outline, which may account for it having been overlooked. Vegetatively it may be distinguished by the posterior costae with lamina to the sinus and the glaucous abaxial side to the lamina. The seedlings, even when quite large, have peltate leaves with the undersides glaucous and the whole glabrous (cf. A. sarawakensis). The inflorescences are very different from those of A. macrorrhizos, both in their clustered arrangement, and the thick marcescent/deliquescent cataphylls (bits of which often adhere to the open spathe), the ivory horizontally red-marked lower spathe and the thick greyish to purplish limb. 2. Though they are morphologically almost identical, plants I have observed in the living state in Sarawak and Sabah differ in the spathe limb colour and inflorescence odour. In the former, the spathe limb is purple and the inflorescence smells of decomposing beef extract; in the latter, the spathe limb is sometimes greyish, slightly suffused with purple, and sweetly fragrant. This may suggest that they are biologically differentiated by having differing pollinators, and warrants further investigation. In plants at Sepilok, I observed that the odour is produced from the inside of the lower spathe, not the spadix. 3. The sap turns orange on exposure - a feature not common in Alocasia, but known in allied Colocasia gigantea. 4. Possible hybridity with A. sarawakensis was noted under that species. Hay 10039 (cult. RBG Sydney Acc. No. 950376) appears in the vegetative state to be intermediate between A. robusta and A. wongii. At the Madai Falls, Sabah, the two putative parents grow together. 5. Ridley’s Bornean record of Colocasia gigantea (reiterated by Merrill, loc. cit.), was based on a sighting at ‘Byte Estate’ near Sandakan, with no 252 Gard. Bull. Singapore 50(2) (1998) specimen preserved. Colocasia gigantea is unknown in Borneo, but bears some resemblance to Alocasia robusta - particularly in the large size and glaucousness of the leaf blade. It seems probable that Ridley, who was interested in Araceae, would have encountered Alocasia robusta, which is a common plant in northern Borneo, and I can only assume that this record is his misidentification of it. Other specimens seen: NATUNA ISLANDS: Pulau Bunguran, E of G. Ranai, van Steenis 1117 (BO); SARAWAK: Cult. RBG Sydney Acc. No. 940569 from Kubah National Park, Matang, Hay et al. 9416 (NSW). BRUNEI: Temburong Prov., Batu Apoi F.R., Poulsen et al. 43 (E, K). SABAH: Mt Kinabalu, E shoulder, Chew et al. 574 (K); Lahad Datu Rd, ca. 10 mi from jnct Sandakan-Kota Kinabalu Rd, Hay 10012 (SAN); Cult. RBG Sydney Acc. No. 950378 ex Madai Falls, Hay 10041 (NSW); Elopura, Leila Rd., Sandakan, Kadir A2698 (KEP, SING). KALIMANTAN: Central East Borneo, W. Koetai, No 45 Kombeng, Endert 5234 (BO, K, L). Scabriuscula Group Species 4—7 Small to very robust lithophytic to terrestrial herbs; /eaves coriaceous, leathery to subsucculent; petioles glabrous to scabrid, occasionally pubescent, often ornamented with purple lines, dots and circles; secondary venation generally flush with the lamina abaxially and adaxially, sometimes impressed adaxially, sometimes prominent abaxially; inflorescences (2—)several to many together not interspersed with foliage leaves; spathes typically with the ground colour white to ivory or yellowish, occasionaly brownish pink throughout, with purple dots and/or margins, sometimes the limb wholly suffused purple to brown, usually constricted at a level well above the sterile interstice of the spadix, sometimes level with the interface of the male zone and the appendix; spadix typically ivory throughout - including the ovaries; stigmas typically with two drop-shaped lobes suberect; fruiting spathe generally white. Notes: 1. This group is endemic to Borneo and appears closely allied to the more widespread but less speciose Puber group, sharing the multiple inflorescences, white, often purple-marked spathes constricted in the male zone. The Puber group is relatively different ecologically, however, evidently associated with open sites, disturbed places and the gap phase of forest, while this group generally is associated with more shaded situations (though A. scabriuscula itself is sometimes found in open sites). 2. The Scabriuscula Group is taxonomically difficult, with, as currently understood, two widespread highly variable species, Alocasia scabriuscula Alocasia in West Malesia and Sulawesi 253 and A. princeps, and a number of local segregates, which may be difficult to differentiate morphologically at the extremes of their variation. Interpreting herbarium material is exceptionally difficult in this group. There is a strong need for more field-based data on their ecology, phenology and pollination biology. Only Alocasia melo, A. principicula and A. reginula are very clearly defined morphospecies, the latter known only from cultivated plants. 3. A curious feature of the Scabriuscula Group, noted already for the Puber Group, is the common positioning of the spathe constriction above the base of the male zone of the spadix such that all or part of the male zone is held within the lower spathe chamber. More usually in this genus, the spathe constriction corresponds in position with the sterile interstice and tightening of the constriction between female and male anthesis appears to be a mechanism that precludes self-fertilisation, preventing pollen from falling into the lower spathe chamber. It might seem that the arrangement of (some of) both fertile zones within the lower spathe chamber could allow self-fertilisation, which might in turn promote the formation of local entities through inbreeding. However, self fertilisation does not appear to take place in this group, as cultivated plants do not set seed spontaneously (though in the field there does seem to be an extremely high level of fruit production). Moreover, the A. longiloba complex (q.v.), which also has highly complex variation patterns, has the more usual relationship between the sterile interstice and the spathe constriction. Whatever the explanation, both these groups appear to be undergoing active evolution. 4. Alocasia scabriuscula N.E. Br. Alocasia scabriuscula N.E. Br., Gard. Chron. 12 (2) (1879) 296; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 79. - Type: Cult. RBG Kew ex N.W. Borneo, Aug 19 1879, N.E. Brown s.n. (K, holo, 3 sheets). Alocasia imperialis L. Linden ex N.E. Br., Gard. Chron. n.s. 21 (May 1884) 711 - Alocasia guttata var. imperialis (L. Linden ex N.E. Brown) N.E. Br., Ill. Hort. 31 (Dec 1884) 185, t. 541. - Type: Cult. Comp. Cont. d’Hort., Gand, 10 Sep 1884, Anon. s.n. (K, holo). Alocasia guttata N.E. Br., Ill. Hort. 31 (Dec 1884) 185, syn. nov. - Type: Cult. Hort. Veitch ex Lawas R., N.W. Borneo, Jan 1879, N.E. Brown s.n. (K, holo, 2 sheets). 254 Gard. Bull. Singapore 50(2) (1998) Alocasia villeneuvei L. Linden & Rodigas, Ill. Hort. 34 (1887) 59; Anon., Kew Bull. (1888) 92 (‘villaneuvii’); Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 86; Merr., Enum. Bornean Pl. (1921) 106 (‘villanuevei’). - Type: Ill. Hort. 34 (1887) t. 21. Epitype: Brunei Darussalam, Temburong, Gunong Retak, 13 Mar 1991, R.J. Johns 6721 (K, designated here). [Alocasia porphyroneura auct. non Hallier f:: Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 100, p.p. quoad specim cit. Hallier 328] [? Alocasia reginae sensu auct. non Linden ex N.E. Br.: M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156 (‘regia’)| Robust herb ca. 0.5—1.2 m tall; rhizome ca. 5-10 cm thick; leaves several together; petioles typically spreading and proportionately rather short - about equalling the length of the blade, ca. 40-100 cm long, smooth to scabrid or sparsely to densely minutely pubescent, rarely plain pale grey- green, usually ornamented with irregular sparse to dense purple-brown dots, circles and longitudinally aligned broken fine lines, sheathing in the lower ca. 1/3; blade ovato-sagittate to broadly ovato-sagittate, ca. 40-85 cm long, adaxially dark to light grey-green and sometimes conspicuously darker along the main venation, abaxially pale grey-green to rich purple, occasionally flushed purple on both surfaces, very thickly leathery to almost succulent; anterior lobe widest at ca. 1/4 of the way distal to the petiole insertion, the apex acute to obtuse; anterior costa. with (4-)5-8 primary lateral veins on each side diverging at 45—60° and with conspicuous green, purple or purple-ringed axillary glands, these sometimes also present in the axils of the larger secondary veins; secondary venation + flush with the lamina to somewhat impressed on both surfaces (depending on thickness of blade), often abaxially obscure or conversely sometimes conspicuous through pigmentation of the bordering lamina, forming more or less well- defined interprimary collective veins; posterior lobes ca. 1/3—!/2 the length of the anterior, usually acute, sometimes rounded, the inner sides usually narrowly to very narrowly oblanceolate, sometimes wider becoming ovate; posterior costae diverging at 60-120°, naked in the sinus for ca. 2-4 cm, rarely with lamina to the sinus, but never peltate as adult plants; inflorescences several together in a tight low cluster; peduncle hardly or not exserted from the subtending cataphylls and leaf sheaths; cataphylls ovate, rather fleshy, often marked like the petiole; spathe greenish to yellowish white to ivory, often speckled purple, sometimes suffused purple throughout or the limb purple, (7—)9-10 cm long, constricted (2—)2.5—3(-5) cm from the base; lower spathe thick, narrowly to broadly ovoid; limb Alocasia in West Malesia and Sulawesi 20 oblong to ovate, 2-3 cm wide, eventually completely reflexed, the tip apiculate to acuminate for 1.5 cm; spadix very shortly stipitate for ca. 1.5 mm, (5.5—)6-7 cm long; female zone (1—)ca. 1.5 cm long, subcylindric to slightly conic, squat, (0.8—)1.2-1.5 cm wide at base; ovary pale green to cream, ovoid, 1—2 mm long; style slender, 0.5—1 mm, facing diagonally out- and up-wards; stigma cream (turning yellow in spirit) mostly 2-lobed; sterile interstice (4—)7-8 mm long, somewhat narrower than female zone at base and ca. 3-4 mm diam., narrowly obconic, basally ca. 2 whorls of somewhat lax white subcylindric synandrodia ca. 1.5 mm diam., distally composed of ca. 3 whorls of synandrodia ca. 2-3 mm diam., closely resembling synandria; male zone ivory, ca. 1.5—-2 cm long, 5-7 mm diam., often somewhat constricted level wth spathe constriction, (1/4—)!/3—-!/2 (—2/3) within the lower spathe chamber; synandria 2—3 mm diam., rhombohexagonal to somewhat irregular; thecae not overtopped by synconnective; appendix ivory, (2—)2.5-3 cm long, tapering to slightly spindle shaped, 5-7 mm diam. at base, the tip usually slightly obtuse; fruiting spathe ca. 4.5 cm long broadly ovoid, white, usually speckled purple. Distribution: Endemic to Borneo, widespread. Habitat: Lowland forest to hill forest to ca. 1200 alt., often in disturbed areas, in swampy to well-drained sites, river banks, occasional on roadsides and in plantations. Notes: 1. As conceived here, this is a highly variable species. Typical forms have ovatsagittate and thick, dark to mid grey-green leaves with the inner sides of the posterior costae very narrowly oblanceolate. These forms conform with the type of A. guttata N.E. Br. Rather less common are large forms with well developed, broad posterior lobes, which conform with the type of A. scabriuscula. There are intermediates. There is also very considerable variation in the thickness of the leaf blade, from distinctly though rather thinly leathery, to blades almost 5 mm thick and virtually succulent. The petiole varies considerable in texture from finely pubescent to scabrid to smooth. 2. Alocasia scabriuscula s. |. is only narrowly distinguished morphologically from A. ridleyi (q.v.), which has relatively longer posterior lobes, longer peduncles, a more slender inflorescence and is a limestone element confined ‘to SW Sarawak. 3. Of the specimens cited below, Beaman et al. 7451 is tentatively ascribed to this species. It differs in the rather long posterior lobes, long peduncles 256 Gard. Bull. Singapore 50(2) (1998) and paired infructescences. Hay 938/ has unusually thinly textured leaves and atypically small inflorescences, with the spathe only ca. 7 cm long, but conforms in other respects. Poulsen 235 is evidently taken from an exceptionally robust plant, as the lower spathe is ca. 5 cm long. 4. There are a number of colour variations, including specimens with red- purple leaf undersides or the leaves entirely suffused with purple (e.g. Afriastini 406). 5. No original material has been found that could serve as a type of Alocasia villeneuvei. The protologue, though singing the praises of this plant in horticulture, contains almost no botanical information. However, it is attributed to Borneo. Dimensions are not given, but the plant is clearly robust. The description does mention brown spotting on the petioles, which is clearly apparent in the illustration. This feature is found in some forms of the A. princeps complex, and to varying intensity in A. sarawakensis, A. reginae and A. scabriuscula. The illustration is not of high quality, but A. sarawkensis can be eliminated as there is no sign at all in the plate of that species’ conspicuous interprimary collective veins. Moreover, it is hardly ornamental. The leaves are too broad and the petioles relatively too short for any larger forms of the A. princeps. Alocasia reginae is small and characteristicaly has the inner side of the posterior costae narrowly lanceolate, while the plant depicted has broadly developed lamina there, which matches the type of A. scabriuscula. Furthermore, the upper side of older leaves is depicted dark dull grey-green which is also typical of this species. Alocasia villeneuvei is therefore epitypified accordingly. The epitype, Johns 6721, has broad posterior lobes, matching well those apparent in the original illustration of A. villeneuvei. : 6. Alocasia imperialis and A. reginae may have been first described by Linden, just prior to N.E. Brown, in an 1883 or 1884 catalogue of the Compagnie Continentale d’Horticulture Gand. However, I have not been able to locate the relevant issues. The plants were exhibited by the Compagnie in May 1884 at the International Exhibition of the Imperial Society of Horticulture at St Petersburg, on which Brown reported in the Gardener’s Chronicle in the same year, describing (validly but unintentionally) Alocasia imperialis. There he also mentioned and briefly described A. reginae, but said it was the same species as A. imperialis, thus invalidating it. Other specimens seen: SARAWAK: Santubong, Carrick & Enoch JC/191 (SING); Cult. RBG Sydney Acc. No. 940505 ex Niah Caves area along track from Niah town, Hay et al. Alocasia in West Malesia and Sulawesi 2 9354 (NSW); Cult. RBG Sydney Acc. No. 940471 ex 2.5 km past Kemena R. bridge on Bintulu-Sibu Rd., Hay et al. 9317 (NSW); Cult. RBG Sydney Acc. No. 940531 ex Kg Sentah, nr Kuching, Hay et al. 9381 (NSW); Kuching, Hewitt 6 (SING); Semunsan Wildlife Sanctuary, Kiew RK&85 (KEP); Anap, Ulu Muput Kanan, path to Bukit Kemantan, Paie S19507 (US); Matang, Ridley s.n. (SING); Santubong, Ridley s.n. (SING). BRUNEI: Temburong Distr., Bukit Biang, Forman 897 (K) & 931 (K); Temburong, Batu Apoi F.R., Sg Baki, Poulsen 108 (K); Temburong, Batu Apoi F.R., Sg Temburong, Poulsen 235 (K) SABAH: Ranau Distr., Pinosuk Plateau, W Mesilau R. at waterworks dam, Beaman et al. 7451 (K, US); Cult. RBG Sydney Acc. No. 960516 ex 2.5 km above main Maliau Falls, G. Rara F.R., Hay et al. 12045 (NSW); Cult. RBG Sydney Acc. No. 960520 ex Kinabatangan, Kalabakan VJR, Hay et al. 12006 (NSW). KALIMANTAN: Central Kalimantan, sei Sampit, Kualakuayan, Afriastini 406 (BO); Pulau Lemoehoetan, Hallier 328 (L). 5. Alocasia reginae L. Linden ex N.E. Br. Alocasia reginae L. Linden ex N.E. Br., [Gard. Chron n.s. 21 (1884) 711 (‘regina’; invalidly published - not accepted by author)]; Ill. Hort. 32 (1885) 11, t. 544; Engl. & K. Krause, Pflanzenr. 71 (1V.23E) (1920) 82 (‘regina’). - Type: Cult. RBG Kew ex Hort. Compagnie Continentale d’Horticulture Gand, Nov 1883, N.E. Brown s.n. (K, holo). Small herb 20-30 cm tall; rhizome ca. 1.5 cm diam.; leaves 2—4 together; petiole 10-25 cm long, sheathing in the lower !/4—!/3, sparsely to densely minutely pubescent, green to densely spotted purple; blade broadly ovato- sagittate to ovato-sagittate, 13-25 cm long, thickly leathery, dark green, paler abaxially or flushed purple throughout; margin entire to slightly and irregularly sinuate; anterior lobe widest ca. !/3 of the way from the base, the apex acute to obtuse and then apiculate; anterior costa with 3-4 abaxially pubescent primary lateral veins on each side diverging at 60° (proximal)—45° (distal) and with small axillary glands; secondary venation adaxially impressed, abaxially flush with the lamina to somewhat impressed (dry), conspicuous and apparently broad (ca. 1 mm) owing to pigment in the bordering lamina, forming mor-or-less well-defined interprimary collective veins at least in the outer part of the blade; posterior lobes 1/3-2/5 the length of the anterior, the inner sides very narrowly oblanceolate; posterior costae naked in the sinus for 4 mm to 1 cm, basally diverging at ca. 90-100° then somewhat to abruptly back-turned; inflorescence pairs solitary; peduncles ca. 1/3 the length of the petiole at anthesis (not or hardly elongating in fruit), subtended almost throughout their length by very narrowly lanceolate cataphylls; spathe ca. 5-7 cm long, constricted ca. 1.5—2 cm from the base; lower spathe ovoid, white, sometimes spotted purple; limb narrowly ovate, white and spotted to suffused purple; spadix distinctly shorter than to subequalling the spathe, sessile, ca. 5 cm long; female zone 1 cm long; pistils flask-shaped; style short, ca. 0.5 mm; stigma 2—3-lobed; interstice 3-5 mm long, partly naked to covered with synandrodia; 258 Gard. Bull. Singapore 50(2) (1998) male zone subcylindric, 9-15 mm long ca. !/4-3/4 held within the lower spathe chamber; synandria rhombohexagonal, ca. 1.5 mm diam.; thecae not overtopped by synconnective; appendix 1-2 cm long tapering to somewhat obtuse, ca. 3 mm diam (dry); fruiting spathe ovoid, ca. 2.5 cm long. Distribution: Endemic to Borneo, known from only three-localities in Sarawak and Central Kalimantan. Habitat: On the floor of primary lowland rain forest: Jermy 13579 on alluvial soil with residual limestone Karst; ca. 40-270 m altitude. Notes: 1. This species is evidently closely allied to the variable Alocasia scabriuscula, differing in the smaller size overall, inflorescences in single pairs, relatively much longer peduncle and the long, narrow inflorescence cataphylls. 2. The description is pieced together from four incomplete collections. The type differs from the only other flowering specimen, Jermy 13579, in having a larger proportion of the male zone exserted from the lower spathe and in having the interstice covered with synandrodia. | 3. Hotta (1967: 156) cited several specimens from Brunei and Sarawak as this species. I have not seen any of the collections concerned, but it would appear from his notes (loc. cit.), in which he gives significantly larger dimensions, that these specimens are probably of A. scabriuscula. 4. Alocasia reginae has been in cultivation in the U.S.A. for some years, under the name Alocasia Elaine. An image may be found at http://www.skg.com/alocasia4.html, and the cultivar is also illustrated in Burnett (1984: 77, fig. 4). Other specimens seen: SARAWAK: Limbang Distr., Sg. Terikan, Chew 1183 (GH, K, L, SING); G. Mulu National Park, around Sg. Berar Camp, Jermy 13579 (K). KALIMANTAN: Headwaters of Sg. Kahayan, 5 km NE of Haruwu Village, Burley et al. 527 (GH). 6. Alocasia reginula A. Hay, sp. nov. Ab Alocasia melo folii lamina atrovirida haud rugosa, nervis albidis impressis, spathae lamina et appendice valde brevioribus differt. - TYPUS: Cult. U.S.A., Florida, Jul 1998, D. Fisk s.n. (NSW, holo). Small herb to ca. 25 cm tall; leaves several together; petioles ca. 18 cm long, Alocasia in West Malesia and Sulawesi 259 sheathing in the lower ca. !/3; blade elliptic to ovate, ca. 15 cm long, 8-11 cm wide, thickly coriaceous, almost completely peltate save for a shallow retuse notch between the tips of the connate posterior lobes, apically acute to obtuse and mucronate for ca. 1 cm, adaxially very dark matt green, abaxially paler and flushed purple; anterior costa with 2-3 primary lateral veins on each side, diverging at ca. 90° (proximal ones) to 45° (distal ones); primary veins adaxially whitish, somewhat impressed, abaxially with inconspicuous axillary glands; secondary venation somewhat impressed adaxially, more or less flush with the lamina abaxially (the larger ones somewhat prominent abaxially in the dry state and adaxially whitish), forming interprimary collective veins towards the margin only; posterior lobes about 2/5 the length of the anterior, with the posterior costae diverging at ca. 30°; inflorescences paired, subtended by short, broad cataphylls; peduncle very short, hidden within cataphyll; spathe ca. 5* cm long, white with scattered purple flecks on the lower part; lower spathe ca. 2.5 cm long, ovoid to subcylindric, separated from limb by a rather weak constriction; limb much reduced, erect even after anthesis, broadly lanceolate, ca. 2 cm long; spadix sessile, somewhat shorter than spathe, ca. 4.5* cm long; female zone about '/4 of the length of the spadix; sterile interstice a single whorl of close-packed synandrodia, not attenuated; male zone cylindric, about half the length of the spadix, about 2/5 as wide as long, ivory; synandria with the thecae not overtopped by synconnective; appendix about !/4 of the length of the spadix, much reduced, narrowly conic; infructescence unknown. Distribution: ?Borneo (see notes). Habitat: Unknown. Notes: 1. This plant is known only in cultivation, but it is clearly quite distinct from any known species. It exhibits few if any characteristics which might suggest that it is of hybrid origin, and is therefore described here as a new species. The description is based on dried leaves sent to me by Mr Dewey Fisk (Florida) and on images posted on the internet (http://ul netgate.net/~kk/Araceae/Alocasia/Black_Velvet.html and http://www.skg.com/alocasia2.html). Since there is no scale on the images, the dimensions above indicated ‘*’ are estimates. Alocasia reginula is well- established in cultivation under the U.S.A. trade-mark cultivar name Black Velvet, and has been successfully micropropagated (see http://www.agristarts.com/tech.htm). 2. The origin of Alocasia reginula is unclear. Scott Hyndman (pers. comm.), 260 Gard. Bull. Singapore 50(2) (1998) who named the cultivar, obtained material from Lyon Arboretum in Hawaii purportedly having been in turn obtained from a Japanese collector in Borneo. In aspect it appears to belong in the Bornean A. scabriuscula group. 3. The specific epithet means ‘little queen’, following the several regal epithets that have been used in this group. 7. Alocasia melo A. Hay, P.C. Boyce & K.M. Wong Alocasia melo A. Hay, P.C. Boyce & K.M. Wong, Curtis’s Bot. Mag. 14 (1997) 82, pl. 315. - Type: Cult. RBG Kew from material collected in Sabah, Malaysia by Mrs S. Collenette s.n., Accession No. 1960 - 443 (holotype K, isotype K spirit coll. no. 22427!). Small herb ca. 25-35 cm tall; stem to ca. 3 cm diam., erect, short; /eaves to ca. 4 together, their bases overlapping; petiole ca. 14-19 cm long, pale green, glabrous, smooth, sheathing and sparingly purple-spotted in the lower 1/5; wings of sheath rather broadly triangular; leaf blade very broadly ovate to sub-orbicular, 18-25 cm x 15 cm, rugose and bullate and deep somewhat bluish green adaxially, smooth and pale greenish white abaxially, coriaceous, almost completely peltate; anterior lobe ca. 12.5-16 cm long, the tip broadly acute to obtuse and then shortly acuminate for ca. 1 cm and/or apiculate; posterior lobes to 8.5 cm long, united for 75-90% of their length; posterior costae diverging from one another at ca. 20-30°, poorly developed and not or hardly differentiated in size from the primary venation arising from the anterior costa; primary lateral veins 3-4 on each side of anterior costa, diverging at 90° (most proximal) to 45° (most distal), adaxially deeply impressed, abaxially more or less flush with lamina and dark green, irregularly bearing veins intermediate in thickness between primary and secondary venation, running the same course as the latter; secondary venation deeply impressed adaxially, abaxially somewhat raised and concolorous with abaxial lamina, arising at a wide angle (c. 80°) from the primary venation and running to form quite well-defined interprimary collective veins; tertiary venation strongly raised adaxially into an irregular honeycomb pattern, abaxially imperceptible; inflorescences paired, subtended by conspicuous broadly lanceolate persistent cataphylls ca. 8-10 cm long; peduncle ca. 5 cm long; spathe ivory-white, 9-16 cm long, constricted slightly less than half way from the base; lower spathe purple- spotted, especially towards the insertion of the peduncle, ovoid, ca. 1.5 cm wide; spathe limb broadly lanceolate, strongly reflexed by male anthesis, the tip acuminate for ca. 1 cm, margins translucent, the entire limb swiftly withering and marcescent (in cultivation; probably deciduous or deliquescent Alocasia in West Malesia and Sulawesi 261 in nature); spadix shortly stipitate, much shorter than spathe, ca. 5 cm long, with the male and female zones enclosed within the lower spathe; female zone 1.2 cm long, 1.3 cm wide at base, tapering distally; ovaries pale green, ca. 1.5—2.2 mm diam., ovoid; style virtually none, stigma 2-3- lobed, orange-brown; sterile interstice much narrower than female zone, ca. 5 mm long, 1.2 mm thick, bearing a few somewhat distant white synandrodia; male zone cylindric, pale ivory-white, 1.3 cm long, 4-6 mm thick; synandria rhombohexagonal, 1—2 mm across; thecae not overtopped by synconnective, opening by apical pores; appendix ivory-white to pinkish, cylindric- subclavate, 1.7 cm long, 4 mm thick; fruiting peduncle ca. 12 cm long; fruiting spathe ovoid, ca. 4 cm long; ripe fruit unknown. Distribution: Endemic to Sabah, Borneo. Habitat: Rain forest on ultramafic rock: in rock crevices and on thin soil along steep banks of fast-flowing streams, 120-400 m. Notes: 1. This species is the only one so far known to be confined to ultramafic substrate, though Sulawesi Alocasia balgooyi (qv) shows a rather strong association with it. 2. The peculiar finely and strongly rugose adaxial leaf surface appears to be unique in the genus, and A. melo would appear to have potential for horticultural exploitation. Other specimens seen: SABAH: Labuk, Sg. Porog, Collenette 502 (K); Cult. Royal Botanic Gardens Sydney, Acc. No. 960489 ex Tongod, G. Tingkar, Hay & Wong 12001, (NSW); Beluran, Porog, W side of Bidi Bidu nr Kubar Labuk, Meijer 41241 (K, L, SAN); cult. Royal Botanic Gardens Sydney, Acc. No. 950381 ex Tongod, G. Tingkar, Radin s.n. (NSW, sterile). Alocasia princeps Complex Species 8—15 Very robust to small terrestrial or lithophytic herbs; petioles smooth to asperous, sheathing in the lower !/9-!/3, concolorous and bright mid-green to purple-brown or variously marked with oblique zones of close-spaced longitudinal brownish lines or with scattered longitudinal thin or thick purple-black lines and dots; blades triangular to narrowly triangular in outline, hastato-sagittate to sagittate, rarely ovato-sagittate, mostly rather thinly leathery to strongly coriaceous, but not subsucculent, usually very dark green adaxially, occasionally grey-green; secondary veins mostly flush 262 Gard. Bull. Singapore 50(2) (1998) with the lamina adaxially and abaxially, occasionally abaxially somewhat prominent and adaxially slightly impressed, not forming interprimary collective veins, or these poorly formed and then only towards the blade margin; posterior lobes about !/3 to subequalling the length of the anterior, acute, the inner side ovate to very narrowly lanceolate (lamina rarely almost entirely lacking on inner side of poserior costa); posterior costae naked in the sinus; inflorescences (2—)several— ca. 10 together, not interspersed with foliage leaves; spathe usually white to ivory to pale yellow throughout, marked with purple dots on the lower spathe, the limb often with a thin purple margin, or spathe dirty pinkish brown and then sometimes sparsely mottled darker and with the limb darker in colour throughout, or spathe greenish; spadix sessile, somewhat shorter than the spathe, pistils usually somewhat acroscopic, ovoid, close-packed to barely touching one another, mostly ivory to pale yellow-green throughout (including stigma); style slender; stigma 2(-3)-lobed, the lobes drop-shaped and suberect to spreading; interstice sometimes partly naked or with the synandrodia loosely packed, or densely covered with synandrodia; male zone subcylindric, often partly to wholly within the lower spathe chamber, ivory; synandria with the thecae not overtopped by synconnective and opening by apical pores; appendix ivory to very pale violet; fruiting spathe (ovoid to) globose, more or less the same colour as the flowering lower spathe, 2-5 cm diam., dehiscing longitudinally from the top. Distribution: Endemic to Borneo; there widespread, but with a concentration of variation in northwest Borneo. Alocasia princeps ‘itself is widespread (though it is interpreted here rather broadly, particularly with respect to material from Kalimantan), while A. ridleyi, A. wongii, A. pricipiculus, A. puteri and A. pangeran are localised segregates. Notes: 1. This complex consists of a number of very similar, closely related elements and presents the greatest difficulties for interpretation of species limits in the genus. Members of the complex cohere in their relatively long, erect petioles, narrowly triangular leaf blades, which are thinly leathery to leathery but not subsucculent (cf. A. scabriuscula), relatively elongate inflorescences with tapering appendices (cf. A. scabriuscula). In other respects, the group is very similar to A. scabriuscula, and Alocasia ridleyi is intermediate, approaching it in its stiffly leathery leaves and relatively short, squat inflorescences. Nevertheless in aspect that species is fairly clearly nearer to A. princeps than to A. scabriuscula. Within the limitations imposed by the material that there is to work with and the timetable for this Flora Malesiana precursor, the treatment of this group, more than any other, should be regarded as no more than a basis for further studies, not Alocasia in West Malesia and Sulawesi 263 only in the way of further collecting and field observation, particularly in Kalimantan, but also studies both at the lower, molecular level, and at the higher, ecological level of population and pollination biology. It is not unreasonable to speculate that this group is in active speciation. 2. Where I have been able to study the plants in the field and in cultivation, it has been possible to recognise species segregated from A. princeps s.l. Given the difficulties of interpreting herbarium material in this group, I suspect that study of living plants in or from other parts of the range would enable the recognition of further segregates - particularly in Kalimantan Selatan, and further refinement of the concept of A. princeps. 3. Members of the Alocasia princeps complex have been frequently misidentified as Alocasia denudata (see A. longiloba ‘denudata’), which is restricted to southern Malay Peninsula and Sumatera, and which is not closely related to A. princeps. They can be quickly differentiated on the basis of the spathe, which in A. /ongiloba ‘denudata’ is green in the lower part and yellowish in the limb, while in this group the spathe is generally ivory coloured with various patterns of purple, and by the stigmas, which are stellate and 3-4-lobed in the A. /ongiloba group and generally bi-lobed with drop-shaped lobes in the A. princeps complex. A. longiloba has distinctly membranous cataphylls, becoming papery and fibrous on drying, whereas those of this group are comparatively leathery in the fresh state. 8. Alocasia ridleyi A. Hay, sp. nov. Ab Alocasia scabriuscula in habitu calcicola, folii lamina plus producta, minus crassa, superne atro-viridi, spatha valde albida vel rosea differt. - TYPUS: Cult. RBG Sydney Acc. No. 940541 ex Malaysia, Sarawak, Bau, Hay et al. 9388 (NSW, holo; iso, K, KEP, L, SAR, SING (to be distributed)). [? Alocasia denudata auct. non Engl.: Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178.] Small to robust herb ca. 45-ca. 1 m tall; rhizome 3-6 cm thick; leaves several together; petioles 35-50(-85) cm long, smooth or rarely scabrid, suberect to somewhat spreading, pale green and unmarked or with few to dense purple-brown dots and lines to flushed purple-brown throughout, sometimes with scattered small circular glands, sheathing in the lower ca. !/5; blade narrowly ovato-sagittate to ovato-sagittate, ca. 15-30(-50) cm long; stiffly leathery, dark green adaxially and shining when young, becoming dull, paler abaxially; anterior lobe widest ca. 2-6 cm distal to petiole insertion; anterior costa with 3—4(-6) primary lateral veins, diverging 264 Gard. Bull. Singapore 50(2) (1998) Morion We stnacott Figure 3. Alocasia ridleyi A. Hay RBG Sydney Acc. No. 940541 - A. habit; B. venation; C. inflorescence with part of spathe removed. - Scale: A, bar = 4 cm; B, bar = 2 cm; C, bar = 8 mm. Alocasia in West Malesia and Sulawesi 265 at ca. 70° (proximal ones) to 50° (distal ones); axillary glands conspicuous (when petiole heavily pigmented) or not (when petiole not so); secondary venation flush on both surfaces, or very slightly prominent abaxially, not or hardly forming interprimary collective veins (these sometimes present in robust specimens); posterior lobes more than 1/2 to subequalling the length of the anterior, the inner sides narrowly to very narrowly (ob)lanceolate; posterior costae diverging at 60-90°; inflorescences several together; peduncle ca. 8-15 cm long, somewhat exserted from the cataphylls; spathe white to pink, sometimes with purple spots, 7—10(-13) cm long; lower spathe ovoid to pyriform, 2—4 cm long; limb oblong-lanceolate, erect then completely reflexed; spadix somewhat shorter than the spathe, very shortly stipitate for 1.5 mm, 6—9 cm long; female zone 1.5—2 cm; ovaries subglobose, close-packed, ca. 2 mm diam.; style very short, 0.25—0.5 mm long; stigma 2—3-lobed, turning yellowish in spirit; interstice 0.5—1 cm long, slightly attenuate, 4-5 mm diam., composed of 2—3 whorls of more or less close-packed rhombo-hexagonal synandrodia ca. 1.5-2 mm diam.; male zone 1.5—2 cm long, !/4—!/2 within the lower spathe chamber, usually slightly constricted level with the spathe constriction; synandria ivory, rhombo- hexagonal, ca. 2 mm diam.; appendix tapering, 2—3 cm long, slightly narrower than male zone, 5-6 mm diam. at base; fruiting peduncle elongating, to ca. 13 cm long; fruiting spathe broadly ovoid, white, sometimes with red spots; longitudinally dehiscent; fruits orange-red. Distribution: Restricted to S.W. Sarawak. Habitat: In forest on limestone at low elevation. Notes: 1. Alocasia ridleyi is named for H.N. Ridley, who took a special interest in Aroids of Borneo (Ridley, 1905), and who was the first to collect this species. 2. Alocasia ridleyi is evidently very closely allied to A. scabriuscula, which also has ovato-sagittate leathery leaf blades and narrowly lanceolate posterior lobes. It differs in the very dark green adaxial leaf surface, relatively longer petioles, relatively narrower leaf blades and somewhat more elongate posterior lobes. The leaves are stiffly coriaceous, but are not subsucculent as those of A. scabriuscula often are. The spathe is recorded as white to pink, whereas that of A. scabriuscula is generally greenish white to yellowish, speckled with purple (though the latter feature is sometimes present in A. ridleyi, as indeed it is in several species in this group). In all these respects, this species is somewhat intermediate between A. scabriuscula and A. princeps. These differences seem quite trivial, but 266 Gard. Bull. Singapore 50(2) (1998) they are correlated with a restricted geographic distribution and association with limestone substrate. Even within its restricted distribution, A. ridleyi exhibits a high level of variability; Nicolson 1285 is exceptionally robust. Other specimens seen: SARAWAK: Seburan, Bau, Anon. 14596 (K); Ist Division, Bukit Rawan, Tebakang area, Awa & Paie $45245 (K); nr Bau, Bogner 1433 (US); Bau, Brooke 9895 (L), 10797 (BM, L); Bidi Cave, Clemens & Clemens 21920 (BO, K); Cult. RBG Sydney Acc. No. 940545 ex G. Gading, Lundu, Hay et al. 9392 (NSW); Bukit Krian [?= G. Kerian], Madison 7344 (K); Bau limestone hills, G. Setiak (SE of G. Doya), Martin $38666 (K); vicinity of Bau, Nicolson 1285 (L, US); Bau, Purseglove P4467 (GH, K, L, SING); Bau, Ridley 11715 (K, SING); Cult. RBG Sydney Acc. No. 942741 ex Bau, Vogel s.n. (NSW). 9. Alocasia princeps W. Bull Alocasia princeps W. Bull, Retail List (1888) 7; ?N.E. Brown, Kew Bull. (1889) 76. - Neotype: Malaysia, Sabah, Mt Kinabalu, Dallas, 27 Aug 1931 Clemens & Clemens 26213 (BM, neo; K, SING, isoneo; designated here - see below). Alocasia porphyroneura [Engl. ex Hallier f., Bot. Jahrb. Syst. 25 (1898) 25, nom. nud. & Notizbl. Koenigl. Bot. Gart. Berl. 2 (1898) 185, nom. nud.] Hallier f., Bull. Herb. Boiss. ser. 2, 1 (7) (June 1901) 671, nom. illeg. (incl. — A. princeps W. Bull); Engl. & K. Krause, Pflanzenr. 71 (I1V.23E) (1920) 100; M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156. Robust to very robust herb ca. 0.8-1.8 m tall; rhizome ca. 5-10 cm diam.; leaves ca. 4 together; petioles suberect, to ca. 1.6 m long, sheathing in the lower !/4—!/3, smooth to slightly rough but not scabrid, dark brownish green, very faintly mottled with an oblique wavy pattern, paler distally, varying to thickly and densely, haphazardly marked with longitudinally aligned purple- brown lines and dots, occasionally with few scattered slightly raised circular glands ca. 2 mm diam.; blade to ca. 55 cm long, leathery but not subsucculent, dark green and shining at least when young, paler and sometimes more or less faintly flushed purple beneath, hastato-sagittate, triangular to narrowly triangular in outline, the margin entire to slightly sinuate and undulate; anterior lobe widest at base; anterior costa with 3(—5) primary lateral veins on each side diverging at ca. 60°, often purple-tinged, with conspicuous axillary glands; secondary venation flush on both surfaces, fine and usually purple-tinged, not or hardly forming interprimary collective veins; posterior lobes subequalling the anterior, narrowly lanceolate to ovate; posterior costae diverging at ca. 90°; inflorescences several (ca. 6) to numerous together, subtended by somewhat leathery marcescent green to pinkish or chocolate brown cataphylls marked similarly to the petioles; spathe white Alocasia in West Malesia and Sulawesi 204 ‘ i es: @ » ~~@ ‘a0 a5 7a Y 9g Figure 4. Alocasia princeps W. Bull RBG Sydney Acc. No. 950357 - A. habit; B. venation; C. petiole ornamentation; D. inflorescence with part of spathe removed. - Scale: A, bar = 5 cm; B, bar = 2 cm; C, bar = 4 mm; D, bar = 8 mm. 268 Gard. Bull. Singapore 50(2) (1998) to yellowish ivory, ca. 11 cm long, constricted at ca. 3 cm; lower spathe ovoid, somewhat to densely spotted purple; limb oblong lanceolate, usually with purple margins, occasionally pink-tinged to bright purple throughout, reflexed and rolled back, the tip acuminate for 1-2 cm; spadix ivory throughout or the female zone very pale green, distal parts sometimes suffused rose, very shortly stipitate for ca. 2 mm, ca. 8 cm long; female zone ca. 1.5 cm long, subcylindric, ca. 1 cm diam.; ovaries subglobose, ca. 1.5 mm diam.; style ca. 0.5 mm, slender; stigma mostly bi-lobed, yellowing in spirit; interstice ca. 5 mm long, slightly attenuate, ca. 4 mm diam.; synandrodia lax and ca. 1 mm diam. in the lower 2 mm, the remainder 2-3 dense whorls of more or less rhombohexagonal synandrodia; male zone ca. 2 cm long, !/2 within the lower spathe chamber, subcylindric, somewhat constricted level with the spathe constriction, ca. 5 mm diam.; synandria rhombohexagonal, ca. 1-2 mm diam. (larger in the lower part of the zone), ivory; appendix 3.54 cm long, slightly narrower than the male zone, 4-5 mm diam., tapering gradually to a point, ivory to flushed pink; fruiting peduncle to ca. 20 cm long; fruiting spathe broadly ovoid, ca. 3-4 cm diam, white, sometimes spotted purple. Distribution: Widespread and common in Borneo. Habitat: In rain forest generally on well-drained slopes and ridgetops, on a variety of substrates including basalt and limestone, from more-or-less sea level to ca. 1200 m altitude. Notes: 1. Alocasia princeps was described from a sterile plant from ‘the Malay Archipelago’ in an 1888 retail horticultural catalogue produced by the British nurseryman William Bull. The description is by no means exhaustive but nevertheless is quite detailed about leaf texture, shape and colour, and it does not seem to fit anything other than the species defined here. The following year the description appeared in Kew Bulletin, very slightly re-worded, in an anonymously compiled list of the previous year’s new plants. Since N.E. Brown was the aroid authority at Kew at the time, it seems quite probable that he contributed at least those plants to that list, if not compiled the list entirely himself. Hallier (loc. cit.) had seen A. princeps growing at Kew (though not until 1897) and considered it the same as his A. porphyroneura. Hallier evidently did not think A. princeps had been properly published, but since it had been, and since he placed it, albeit attributed to Brown and not Bull, in the synonymy of A. porphyroneura, 1 conclude that the latter is superfluous. Although Brown usually preserved material of new aroids in cultivation at Kew, no identifiably original material nor illustration exists, which fixes the application of A. princeps or can directly assist in its Alocasia in West Malesia and Sulawesi 269 interpretion. The leaf is described as having ‘deeply sinuate’ margins, which is extreme for this species, in which the leaf margin is usually more or less entire, sometimes somewhat undulate and occasionally shallowly sinuate (Hallier described the leaf margin of A. porphyroneura as repand-sinuate and strongly undulate, which fits better the states I have observed). However, ‘deeply sinuate’ is perhaps open to a variety of interpretations, and the state may even have been somewhat exaggerated by Bull for marketing purposes. Although, as has been said, there is no original material bearing this name at K, a sterile specimen nevertheless exists there obtained from Bull, allegedly originating from the Philippines, dated Aug 11 1887 and with Bull’s number 4454/5. It is simply annoted ‘Alocasia’ in Brown’s hand. It matches Bull’s catalogue description of A. princeps closely (except that the leaf margin is shallowly sinuate), and is clearly not Philippine, conforming exactly to Alocasia princeps in the sense here. It seems likely that this specimen is from the original plant Bull introduced and perhaps only subsequently called Alocasia princeps. Since Hallier had seen what is presumed to be the original plant and considered it the same as A. porphyroneura, which he had described in considerable detail and of which he had preserved material, I consider that A. princeps should be neotypified accordingly. This interpretation is to some extent further supported by the only preserved material I have come across originally annotated with this name: a plant cultivated at the Peradeniya Gardens, Ceylon, collected by Alston in 1926 (Alston 794, K!) and which clearly falls within the present concept of A. princeps. The designated neotype collection is one of few that is fertile, of known wild provenance and that exists in duplicate. Other specimens seen: SARAWAK: Betong, Saribas F.R., Anderson 8507 (L); Hose Mts, Mujong, Ulu Temalad, Ashton S17639 (L); Simpang Tiga, Ulu Mayeng, Kabu, Chai $19219 (US); Baram Distr., G. Api foothills, Chew 1099 (GH, K, SING); Lingga, Hullett s.n. (SING); 4th Div., Garden of Eden, Mulu National Park, Primack S42403 (K). BRUNEI: Temburong Distr., Sg. Temburong at Kuala Belalong, Boyce et al. 384A (K); Temburong, headwaters of Sg. Temburong to NE Guning Retak, Johns 6733 (K); Temburong Prov., Batu Apoi F.R., Sg. Belalong, Poulsen 18 (K); Temburong Prov., Batu Apoi F.R., Sg. Enkiang, Poulsen 264 (K). SABAH: Kudat Distr., Pulau Balembangan, NE inner side of Telok Lung, BCS-EFA-LM et al. (sic) SAN 86716 (L); Mt Kinabalu, Ulu Liwagu and Ulu Mesilau, Chew et al. 1954 (K, SING); Mt Kinabalu, Dallas, Clemens & Clemens 26134 (BO) & 26268 (BM, SING); Penibukan, Clemens & Clemens 30519 (BM, SING) & 31161 (BM, K, SING) & 32095 (BM, BO, GH, L, SING); Mabul Isl., Creagh s.n. (K); Moyog, along rd Kota Kinabalu-Tambunan, 24 mi SE of Kota Kinabalu, Croat 53102 (MO); Kinabatangan Besar, Kori Timber Camp, Cuadra A2153 (BO, K, KEP, L, SING); Nungkok Mt, Darnton 443 (BM); Kinabalu, Tenompok, Clemens & Clemens 29148 (BM, BO, GH, K, L) & 29149 (BM, SING) & 29150 (SING) ); Cult. RBG Kew EN 441-63 ex Sapong estate, nr Tenom, Giles & Woolliams s.n. (K); Cult. RBG Sydney Acc. No.950357 ex Kinabatangan, Gua Batu Puteh, Hay et al. 10015 (NSW; fertile voucher SAN); Cult. RBG Sydney Acc. No. 960560 ex 2.5 km above main Maliau Falls, G. Rara F.R., Hay et al. 12039 (NSW). Cult. 270 Gard. Bull. Singapore 50(2) (1998) RBG Sydney Acc. No. 960467 ex Tibau Forest Station, Kinabatangan, Hay et al. 12140 (NSW); Lahad Datu, Selangan Island F.R., Semporna, Keith A1519 (K, KEP, L) & 7658 (SING) & 928] (KEP, SING) & 9286 (KEP); Bajau, Mt Sidungol, Keith 9295 (K, KEP, SING) Keringau Distr., Camp C area Tiulan, Maikin Lantoh 102054 (K); Ranau, ca. 3 mi NW of Kg Pinawanti, SAN 76856 (K). KALIMANTAN: Sungei Utung, Amdjah 356 (BO); East Kalimantan, Balikpapan, PT. ITCI area, Darnaedi 431 (BO); Kalimantan Selatan, G. Halauhalau (G. Besar), Pegunungan Meratus, Barabai, Dransfield 2870 (BO); Kalimantan Selatan, Djaro Dam, Muara Uja, Dransfield & Saerudin 2210 (BO) & 2217 (BO); Kalimantan Selatan, 2km S of Djaro Dam, Muara Uja, Dransfield & Saerudin 2296 (BO); Central East Borneo, W. Koetai, Endert 2702 (BO, L); Cult. Hort. Bogor, Hallier s.n. (BO); Lombok Utan, Hallier 358 (BO, L); Amai Ambit, Hallier 3455 (BO); East Kalimantan, Loa Haur, W of Samarinda, Kostermans 6810 (BO, K); East Kalimantan, Nanukan Island, Kostermans 8784 (BO); Kutei, G. Beratus, piek van Balikpapan, Meijer 653 (BO); East Kalimantan, Nunukan, N of Tarakau, Meijer 2033 (BO) & 2168 (BO) & 2300 (BO) 10. Alocasia wongii A. Hay, sp. nov. Ab Alocasia princeps inflorescentia tenuiore, spatha brunneo-rosacea, spathae lamina fusca, inflorescentia mascula minus exserta, petiolo Alocasia longiloba ‘denudata’ simulans sed tenuiore saepe aspero differt. TYPUS: Cult. RBG Sydney Acc. No. 960457 ex Madai Caves, Hay et al. 12180 (NSW, holo: iso, K, KEP, L, SAN — to be distributed). Terrestrial herb ca. 60(—90) cm tall; rhizome ca. 2.5-4 cm diam.; petioles smooth or asperous, slender, erect, grey-green, tinged pinkish at the base, densely mottled in an oblique pattern of crowded longitudinally aligned fine dark brown lines, sheathing in the lower !/5—!/s; blade ca. 3/4 the length of the petiole, hastato-sagittate, narrowly triangular, dark (grey-)green above, paler below; anterior lobe widest at the base, the margin sometimes slightly undulate; anterior costa with 2—3 primary lateral veins on each side diverging at ca. 45-80(-90)°; axillary glands inconspicuous, sometimes conspicuous at the junctions of costae and petiole; secondary venation flush with the lamina to somewhat prominent abaxially and sometimes markedly divaricating, forming ill-defined interprimary collective veins towards the margin or these absent; posterior lobes rather slender, subequalling the anterior, the inner sides lanceolate to ovate; posterior costae diverging at ca. 100°, naked in the sinus for 2-4 cm; inflorescences to ca. 6 together; cataphylls membranous, mottled pinkish brown and green, marcescent; peduncle ca. 10 cm long, pale dirty pinkish; spathe ca. 7-8 cm long; lower spathe pear-shaped to ellipsoid, ca. 3 cm long, pale brownish pink, the mottling often forming vertical streaks; constriction usually oblique; limb narrowly (to broadly) lanceolate, distally tapering, brownish pink, distinctly darker than the lower spathe; spadix 5—7 cm long, very shortly stipitate for ca. 1 mm; female zone pale green; pistils somewhat distant - barely touching; ovaries subglobose; style slender, ca. 0.5 mm Alocasia in West Malesia and Sulawesi oar | Morion Weshnacot Figure 5. Alocasia wongii A. Hay RBG Sydney Acc. No. 960478 - A. habit; B. venation; C. petiole ornamenation; D. inflorescence with part of spathe removed; E. pistils and two staminodes. - Scale: A, bar = 8 cm; B, C, bar = 2em; D bar = $.mm:E, bar ='1 mm. 272 Gard. Bull. Singapore 50(2) (1998) long, abruptly differentiated from the ovary; stigma 2-lobed; interstice ca. 5 mm, partly naked, slightly attenuate, 3-5 whorls of synandrodia, the lower ones lax; male zone ca. 1.2-1.5 cm long, ca. 4 mm diam., cylindric, somewhat constricted at level of spathe constriction, 3/4 to almost entirely within lower spathe chamber; synandria rhombo-hexagonal, ca. 2 mm diam., not overtopped by synconnective; appendix ca. 3 cm long, somewhat constricted at base, ca. 4 mm diam. near base, tapering to a blunt tip; fruiting spathe ovoid, to 4.5 cm long, off white to dusky pink with vertical darker streaks. Distribution: Northeastern Sabah. Habitat: Terrestrial in lowland mixed dipterocarp forest on well-drained and occasionally inundated sites at low elevation. Note: A fairly well-defined and geographically coherent element distinguished from Alocasia princeps s.s. by the generally more slender stature, the petiole with markings distinctive in the complex but almost identical to those typically found in Alocasia longiloba ‘denudata’. The spathes are suffused a rather dirty purple-brown or violet-brown, with the limb somewhat darker than the lower spathe. The lower spathe in flower and the fruiting spathes are less rotund, and the limb shorter and more slender than in Alocasia princeps. The spadix is more slender with the pistils and synandrodia rather loosely arranged and the male zone 1s less exserted. Populations in the vicinity of Sandakan have distinctively asperous petioles and the secondary venation is somewhat prominent abaxially and, towards the leaf margin, forms rather widely and loosely divaricating interprimary collective veins. Elsewhere in the range, e.g. at Madai, the petioles are smooth, the secondary venation is flush with the lamina and only faintly forms interprimary collective veins if at all. Alocasia wongii is named in honour of Dr Wong Khoon Meng, who has contributed much to knowledge of the botany of Sabah. Other specimens seen: SABAH: Tongod, Ulu Sg. Pinangah, Amin & Ismail SAN 107302 (K); Bettotan, nr Sandakan, Boden Kloss 19095 (SING); Tongod, ridge of Bukit Mengalas- Kalas, Dewol Sundaling SAN 93149 (GH, K); Cult. RBG Sydney Acc. No. 950375 ex Madai Falls, Hay et al. 10038 (NSW; fertile voucher SAN); Cult. RBG Sydney Acc. No. 950358 ex Ulu Dusun, Hay et al. 10021 (NSW); Cult. RBG Sydney Acc. No. 960478 ex 2.5 km above main Maliau Falls, G. Rara F.R., Hay et al. 12068 (NSW); Kabili, Sandakan, Keith 4969 (K, SING) & 9951 (K); Sepilok F.R., Kiew RK 757 (KEP); Tawau Dist., Gemok Hill F.R., Madani & Sigin SAN 111570 (KEP); Sandakan, Ramos 1580 (K, US). Alocasia in West Malesia and Sulawesi MEGS 11. Alocasia pangeran A. Hay, sp. nov. Ab Alocasia principicula folio fusco-viridi, inflorescentia majore, interstitio valde attenuato partim nuda, inflorescentia mascula dimidium exserta differt. - TYPUS: Cult. RBG Sydney Acc. No. 960509 ex Malaysia, Sabah, Lahad Datu, Madai Caves, Hay et al. 12175 (NSW, holo; iso, K, KEP, L, SAN, SING - to be distributed). Lithophytic herb to ca. 60 cm tall; /eaves ca. 4 together; petioles to ca. 60 cm long, often somewhat spreading, smooth, dark green, sheathing in the lower ca. !/7; leaf blade variable, sagittate to hastate, triangular in outline, dark green adaxially, paler abaxially, ca. 20-35 cm long; anterior lobe widest at the base, the tip acute and slightly acuminate; anterior costa with 3 primary lateral veins on each side, diverging at 45-60°; axillary glands inconspicuous; secondary venation flush with the lamina on both surfaces, not forming interprimary collective veins, or these sporadic, ill- defined and only in the outer part of the blade; posterior lobes more than !/2 to subequalling the anterior, widely spreading or not, acute to narrowly rounded, the inner sides elliptic to narrowly obovate; posterior costae diverging at ca. 120°, straight (when leaf hastate) or somewhat back-curved (when leaf sagittate); inflorescences to 6 together; peduncle ca. 15 cm long, subtended by lanceolate cataphylls 8-14 cm long; spathe ca. 11 cm long, constricted ca. 2.5 cm from the base; lower spathe ovoid, greenish ivory; limb ca. 8.5 cm long, lanceolate, acuminate for ca. 2 cm, ivory; spadix subsessile, stipitate for 1.5 mm, 8 cm long; female zone 1 cm long; pistils facing diagonally upward, ovaries subglobose, ca. 2 mm diam.; stigma more or less sessile, bilobed; interstice 8 mm long, naked and attenuate in the lower 5 mm with scattered small synandrodia, in the upper part with 1-2 whorls of close-packed synandrodia; male zone ivory, 1.5 cm long, 6 mm diam., !/2 exserted from the lower spathe chamber and constricted level with the spathe constriction; synandria ca. 2 mm diam., rhombo-hexagonal; thecae not overtopped by synconnective; appendix subcylindric, isodiametric with male zone, distally tapering; fruiting peduncle about half the length of the petiole; fruiting spathe broadly ovoid, white. Distibution: Endemic to Sabah, known only from Madai Caves, where it is abundant. Habitat: In soil and humus pockets on limestone outcrops and boulders in mixed lowland dipterocarp forest at ca. 400 m alt. Note: This species is distinguished from Alocasia princeps by its calcicolous lithophytic habit, smaller stature, more slender inflorescence, relatively 274 Gard. Bull. Singapore 50(2) (1998) a atl Marion Weslmacott 3g Figure 6. Alocasia pangeran A. Hay RBG Sydney Acc. No. 960509 - A. habit; B. venation; C. inflorescence with part of spathe removed; D. pistils; E. lower neuter organs; F. synandria. - Scale: A, bar = 4 cm; B, bar = 2 cm; C, bar = 1 cm; D, E, F, bar = 2 mm. Alocasia in West Malesia and Sulawesi DIE hs, longer lower spathe and lax interstice. It coexists with the terrestrial A. wongii and the lithophytic A. puteri (q.v.). The specific epithet is a princely title in Malay, alluding to the relationship of this species to Alocasia princeps. For further discussion see under Alocasia puteri. 12. Alocasia puteri A. Hay, sp. nov. Ab Alocasia pangeran A. Hay lamina folii claro-viride, lobis posticis intus ovatis, inflorescentia breviori, interstitio neutro brevissimo crasso vel nullo, inflorescentia mascula minus exserta, stigmate trilobato differt. - TYPUS: Cult. RBG Sydney Acc. No. 960603 ex Malaysia, Sabah, Lahad Datu, Madai Caves, Hay et al. 12178 (NSW, holo; iso, K, KEP, L, SAN, SING - to be distributed). Lithophytic herb to ca. 60 cm tall; rhizome ca. 3 cm diam.; leaves ca. 3 together; petioles more or less erect, to ca. 50 cm long, bright green, very faintly mottled slightly darker green, sheathing in the lower !/6; blades hastato-sagittate to slightly ovato-sagittate, ca. 30 cm long, bright green adaxially, somewhat paler abaxially, thinly leathery; anterior lobe widest at or slightly above the base, the apex acute to obtuse and apiculate for ca. 1 cm; anterior costa with 2-3 primary lateral veins on each side diverging at 45—70°; axillary glands very inconspicuous; secondary venation flush with the lamina on both surfaces, faint, forming poorly defined interprimary collective veins in the outer part of the blade; posterior lobes acute, more or less equalling the anterior, the inner sides ovate, peltate even in subadult plants but eventually free; posterior costae naked in the sinus for ca. 2 cm, diverging at ca. 60°; inflorescences ca. 4 together; cataphylls lanceolate, to ca. 7 cm long; peduncles exserted, to 15 cm long; spathe ca. 8 cm long, constricted 2 cm from the base; lower spathe ovoid, greenish ivory; limb lanceolate, green; spadix subequalling the spathe, minutely stipitate; female zone ca. 6 mm long; pistils subglobose, ca. 2 mm diam., pale greenish ivory, more or less outward-facing; style short, ca. 0.5 mm, sharply differentiated from the ovary and stigma; stigma (2—)3-lobed; sterile interstice more or less lacking, the male zone adjunct to the female with some of the lowermost synandria incompletely fertile, or one to two whorls of close-packed synandrodia resembling the synandria, not or hardly attenuated; male zone ivory, cylindric, ca. 1.8 cm long ca. 4 mm diam., 2/3 within the lower spathe chamber, somewhat constricted level with the spathe constriction; synandria rhombo-hexagonal, ca. 2 mm diam., the thecae not overtopped by synconnective; appendix ivory, basally isodiametric with male zone, tapering; infructescence unknown. 276 Gard. Bull. Singapore 50(2) (1998) “f Marion Westmacot- Figure 7. Alocasia puteri A. Hay RBG Sydney Acc. No. 960603 - A. habit; B. venation; C. inflorescence with part of spathe removed. - Scale: A, bar = 4 cm; B, bar = 2 cm; C, bar = 4 mm. Alocasia in West Malesia and Sulawesi 277 Distribution: Endemic to Sabah, known only from Madai Caves where it is uncommon. Habitat: In soil pockets on limestone outcrops and boulders in mixed lowland dipterocarp forest, at ca. 400 m altitude. Notes: 1. The specific epithet, Malay for ‘princess’, alludes to the relationship with Alocasia princeps and to this species’ coexistence with A. pangeran. 2. Alocasia puteri occurs together with and in the same habitat as A. pangeran. They can be differentiated by the former’s bright green leaf colour (unusual for this entire species group) vs darker green, the broader posterior lobes, more upright petiole, tendency for the leaves to be peltate in sub-adult plants, virtually lacking sterile interstice of the apendix vs a long partially naked interstice of lax synandrodia, and the stigma being usually 3-lobed vs generally 2-lobed. Moreover, at the time of original collecting both of these species at Madai Caves, plants of A. pangeran were extensively in fruit, while no plants of A. puteri were in either flower or fruit, suggesting they may be isolated by flowering time. 13. Alocasia principiculus A. Hay, sp. nov. Ab Alocasia princeps W. Bull stature valde minore, habitu calcicola lithophytica, pedunculo longo, inflorescentia mascula haud exserta differt. - TYPUS: Cult. Royal Botanic Gardens Sydney Acc. No. 960576 ex Malaysia, Sabah, Kinabatangan Distr., Gomantong Caves, originally collected 24 Apr 1996, Hay et al. 12162 (NSW, holo; iso, K, L, KEP, SAN, SING - to be distributed). Small more or less lithophytic herb; rhizome ca. 1.5—2 cm diam.; leaves several together; petioles ca. 20-30 cm long, sheathing in the lower !/7-!/o, grey-green to dark purple-brown, becoming paler distally, somewhat spreading, smooth; blade ca. 20(—25) cm long, hastato-sagittate, narrowly triangular in outline, occasionally somewhat ovato-sagittate, sometimes some subadult leaves peltate, dull to slightly glossy, distinctly grey-green above, paler below, very thinly leathery; anterior lobe ca. 9-14 cm long, widest at base; anterior costa with 2(—3) primary lateral veins on each side, diverging at ca. 45—60°; secondary venation flush with the lamina, forming poorly defined interprimary collective veins towards the margin; axillary glands inconspicuous; posterior lobes ca. 2/3 the length or subequalling the anterior, the inner sides lanceolate; posterior costae diverging at ca. 60—90°; inflorescences mostly 2 together (1-4); peduncles much extended 278 Gard. Bull. Singapore 50(2) (1998) a Fae sees. z € CIO Marton Weslmocalt: Figure 8. Alocasia principiculus A. Hay RBG Sydney Acc. No. 960576 - A. habit; B. venation; C. inflorescence with part of spathe removed. - Scale: A, B, bar = 2 cm; C, bar = 8 mm. Alocasia in West Malesia and Sulawesi 279 from the cataphylls, ca. !/3-2/3 the length of the petioles; cataphylls lanceolate, ca. 7 cm long; spathe ca. 6—7 cm long; lower spathe ovoid, ivory to yellowish ivory, ca. 2 cm long; limb lanceolate, pale green faintly suffused brown; spadix ca. 5 cm long; female zone ca. 1 cm long; pistils not or hardly touching one another, pale green; ovaries ovoid, ca. 1.5 mm diam., facing diagonally up, pale greenish yellow; style short, ca. 1 mm; stigma white, bilobed; interstice ca. 4 mm long, attenuate, 2-3 whorls of loosely arranged synandrodia (sometimes partly naked); male zone ca. 8 mm to 1| cm long, pale ivory, completely within the lower spathe chamber; appendix ivory, ca. 2.5-3.5 cm long, 4 mm diam., slightly constricted at junction with male zone corresponding to spathe constriction; fruiting peduncle subequalling petioles. Distribution: Endemic to Borneo: Sabah and East Kalimantan. Habitat: Lowland rain forest, on and among limestone boulders; sea level to 600 m altitude. Notes: 1. The substantive epithet, meaning ‘little prince’, is the diminutive of princeps, alluding to the small stature and relationship of this species. 2. Alocasia principiculus differs from other members of the A. princeps complex in its small overall size, grey-green, thinly leathery leaves, relatively long peduncles and in the male zone being held entirely within the lower spathe chamber. Other specimens seen: SABAH: Sandakan Forest District, Elopura, Gomantong, Cuadra A1489 (BO, BRI, K, KEP, L, SING); 20 mi S of Sandakan, Gomantong Caves Hill, Wood A4602a (L). KALIMANTAN: East Kalimantan, Berouw, Mt Bungaan, Kostermans 13781 (L). 14. Alocasia reversa N.E. Br. Alocasia reversa N.E. Brown, Gard. Chron. 8 (2) (1890) 38; Hook.f. in Curtis’s Bot. Mag. Ser. 3, 52 (1896) t. 7498; Usteri, Beitr. Kenntn. Philipp. (1905) 130; K. Krause & Engl., Pflanzenr. 71 (I1V.23E) (1920) 95; Merr. Poum,, Philipp. Bl Pl. 1-(1922), 185...- Type: Cult: RBG Kew ex Hort. Sander, Apr 1890, N.E. Brown s.n. (K, holo). Small herb to ca. 35 cm tall (often less); rhizome short, condensed, erect to decumbent, ca. 3 cm diam.; /eaves several together, irregularly interspersed with marcescent brown lanceolate cataphylls to ca. 7 cm long; petioles ca. 10-20(-30) cm long, sheathing in the lower ca. 10-15%; blades thinly 280 Gard. Bull. Singapore 50(2) (1998) leathery, in adult plants all peltate or (usually) mixed peltate and non- peltate, occasionally none peltate, ovate to rather narrowly ovato-sagittate, 14 x 6 cm to 22 x 10 cm, widest at or slightly distal to junction of petiole, glossy dark green about the midrib and primary veins adaxially, the rest grey-green; anterior lobe acute, with the tip abruptly acuminate for ca. 1 cm; anterior costa with 3-4(—5) primary lateral veins on each side, diverging at ca. 90° proximally, the distal ones at ca. 60°, running almost straight to the margin and joining a submarginal vein; axillary glands not conspicuous; secondary venation fine, not forming interprimary collective veins, or these very poorly defined, flush with the lamina; posterior lobes about !/2-2/s the length of the anterior, united for 50-90% of their length or free, when maximally united the base of the lamina rounded except for an acute notch; posterior costae diverging at ca 15° when blade peltate, or at ca. 90° when not peltate but then soon curved back (so posterior lobes not widely divergent) and naked in the sinus for ca. 1 cm; inflorescences 1-2 together, subtended by lanceolate marcescent brown cataphylls; peduncle ca. 10 cm long at anthesis, often exceeding the petioles at fruiting; spathe ca. 6 cm long; lower spathe ca. 2 cm long, ovoid, pale green; limb paler green to ivory edged purple with the colour extending into the constriction ventrally, at first erect, then sharply deflected, oblong lanceolate, mucronate for ca. 6 mm; spadix somewhat shorter than the spathe, white except for bright green ovaries, stipitate for ca. 3 mm; female zone 1 cm long; pistils somewhat loosely packed, flask-shaped, ca. 1.5 mm diam., facing obliquely upwards; style slender, ca. 0.5 mm long; stigma 2-lobed; sterile interstice ca. 4 mm long, not much attenuated and situated within the chamber of the lower spathe, lower synandrodia irregular in shape, ca. 2 mm diam., upper ones rhombo-hexagonal; male zone ca. 1 cm long, 4 mm diam., subcylindric, slightly constricted ca 2/3 from the base corresponding to spathe constriction; synandria rhombo-hexagonal ca. 2 mm diam., 4—-6-merous, the thecae opening by apical pores, synconnective not expanded; appendix about isodiametric with male zone, ca. 2 cm long, gradually tapering to a blunt point, faintly longitudinally channelled; fruiting spathe broadly ovoid, ca. 2 cm long, eventually reflexed, pinkish to orange; berries bright orange to red. Distribution: Borneo, endemic to Sarawak Habitat: In forest over limestone, often on boulders, to 300 m altitude. Notes: 1. This species, first described from cultivation in Britain following introduction by Sander & Sons, was originally attributed to the Philippines and consequently cited by some later authors as a Philippine plant. No Alocasia in West Malesia and Sulawesi 281 authentically Philippine material of this species has been found however, and it must be assumed that Sander’s attribution was a mistake, or deliberately misleading - a practice, according to Burnett (1984), of competing nurserymen of the time. 2. This and the following species appear in their inflorescence structure to belong in the Scabriuscula Group. They differ in their (usually) peltate leaves and the frequent (but ?irregular) cataphylls among the foliage leaves, which suggests relationship with the Cuprea Group. 3. A clone of Alocasia reversa is traded in Singapore under the cultivar name Alocasia ‘Hana’. An image may be found at http:// www.springleaf.com.sg/hana.html. Other specimens seen: SARAWAK: Kuching Distr., Tiang Bekap, Padawan Rd, Anderson 10095 (K, L, SING); Cult. RBG Edinburgh Acc. no. 67.1612 ex Bukit Serapat, ca. 13 mi from Kuching on Semenggang Rd, Burtt & Martin 4745 (E); Kuching Distr., Tiang Bekap, Mt Mentawa, Chew 670 (GH, K, L, SING), 1292 (GH, K, SING); Along Kuching-Padawan Rd, 10 mi SW of main Kuching-Serian Highway, Croat 53183 (MO); Bukit Manok, Padawan, 38 mi from Kuching, Erwin & Paul $.27408 (K, US); Zuab (CHECK), Hewitt s.n. (K); Bidi, Hewitt 4 (SING); Bukit Mentawa off 32nd Mile Padawan Rd, Kuching, Ist Div., Mamit S.32673 (K, L, SING, US); Gat, ‘Native Collector’ D106 (E); Cult. RBG Sydney Acc. no. 942723 ex cult. Hort. Leiden Acc. no. 933071 ex N of Padawan, Vogel s.n. (NSW); Bukit Payang, 10 km Tebakang-Tebedo Rd, Ist Div., Yii & Othman S.46241 (K, KEP). 15. Alocasia venusta A. Hay, sp. nov. Ab Alocasia reversa planta robustiore, folio tenuiter oblongo-ovato atro- viride, inflorescentibus 4—6 in synflorescentia confertis, pedunculo breviore, inflorescentia majore, stigmate trilobato differt. - TYPUS: Cult. RBG Sydney Acc. No. 940504 ex Niah National Park, path between Niah town and Niah caves, Hay et al. 9346 (NSW, holo). Lithophytic herb to 45 cm tall; rhizome condensed, to 3 cm thick; eaves to 6 together, interspersed with marcescent cataphylls ca. 5 cm long; petiole to ca. 40 cm long, sheathing in the lower !/9; blade ca. 27-35 cm long, completely peltate, stiffly leathery, shiny very dark bluish green adaxially and somewhat darker about the main veins, somewhat paler abaxially, oblong-ovate, ca. 9-12 cm wide; anterior lobe widest at the base, the apex acute, acuminate for 1.5 cm; anterior costa with 3 primary lateral veins on each side, diverging at ca. 60—80°, drying flush with the lamina, with very inconspicuous axillary glands; secondary venation fine, not forming interprimary collective veins, almost striate; combined posterior lobes cuneate, somewhat elevated, ca. 8 cm long, ultimately with a shallow retuse notch between the posterior costae; posterior costae diverging at ca. 15°; 282 Gard. Bull. Singapore 50(2) (1998) inflorescences 4—6 together not interspersed with foliage leaves, subtended and equalled to exceeded in length by lanceolate cataphylls to 10 cm long; peduncle ca. 6 cm long at anthesis; spathe ivory, suffused and spotted purple, ca. 9 cm long, constricted at ca. 3 cm from the base; lower spathe ovoid; spadix ca. 7 cm long, stipitate for 1-4 mm with the stipe obliquely inserted; female zone 1-1.5 cm long, subcylindric, slightly conic; ovaries ovoid, ca. 1.5 mm diam; style ca. 1 mm long; stigmas mostly 3-lobed with drop-shaped diagonally upward-facing lobes; sterile interstice 3-4 mm long, partly naked, with 1-2 whorls of synandrodia; male zone 1.5 cm long, narrowly conic, ca. 4 mm diam. at base, 3/4 held within lower spathe chamber; synandria ivory, rhombo-hexagonal, ca. 2 mm diam.; thecae not overtopped by synconnective; appendix subcylindric, slightly narrowed at the base, then more or less isodiametric with top of male zone, tapering, ca. 4 cm long; infructescence unknown. Distribution: Endemic to Borneo, known only from the Niah Caves area, northern Sarawak. Habitat: Lithophytic on limestone in swamp-forest at ca. 200 m altitude. Notes: 1. This species is evidently closely allied to Alocasia reversa. It differs in the more robust habit, the more complex synflorescence (inflorescences solitary to paired in A. reversa), shorter peduncles, more densely arranged pistils and trifid stigmas. The leaves are all completely peltate and narrowly oblong-ovate, whereas they are often mixed peltate and non-peltate in A. reversa, especially in more robust forms approaching A. venusta in size. The leaf blades of A. reversa are conspicuously variegated. Alocasia venusta 1s geographically disjunct from A. reversa, which is known only from SW Sarawak. 2. The specific epithet alludes to the plant’s highly ornamental qualities. It does not appear yet to have entered commercial horticulture, in which it has potential. Macrorrhizos Group Species 16—21 Moderately to very robust herbs; inflorescence pairs mostly interspersed with foliage leaves, occasionally clustered; spathe constriction corresponding with sterile interstice of the spadix; thecae of synandria overtopped by synconnective. Alocasia in West Malesia and Sulawesi 283 The species in this group are of somewhat doubtful affinity, but the group extends throughout Malesia except Borneo (other than A. macrorrhizos itself). 16. Alocasia macrorrhizos (L.) G. Don Alocasia macrorrhizos (saepissime ‘-rrhiza’) (L.) G. Don in Sweet, Hort. Brit. ed. 3 (1839) 631 (‘-rrhizon’); Schott in Oesterr. Bot. Wochenbl. 4 (1854) 409; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 502; Engl. in Koord., Meded. Lands Plantentuin. 19 (1898) 300; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178; Koord., Exkurs.-Fl. Java 1 (1911) 261; Merr., Interpr. Herb. Amboin. (1917) 130; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 84, fig. 15; Furtado in Gard. Bull. Straits Settlem. 11 (1941) 244-257- Arum macrorrhizon L., Sp. Pl. (1753) 965; - Colocasia macrorrhiza (L.) Schott in Schott & Endl., Melet. Bot. (1832) 18. - Type: Arum maximum macrorrhizon zeylanicum Herm., Parad. Bat. (1698) t. 73 (lecto; selected by Furtado, 1941). Arum indicum Lour., Fl. Cochinch. (1790) 536; ed. Willd. (1793) 655; Roxb., Fl. Ind. ed. 2, 3 (1832) 498. - Colocasia indica (Lour.) Kunth, Enum. Pl. 3 (1841) 39. - Alocasia indica (Lour.) Spach, Hist. Nat. Veg. Phan. 12 (1846) 47; Schott, Oesterr. Bot. Wochenbl. 4 (1854) 410; Engl. in A. DC, Monogr. Phan. 2 (1879) 501 & Bot. Jahrb. Syst. 25 (1898) 23; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178; Koord., Exkurs.-Fl. Java 1 (1911) 262; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 87; Backer & Bakh.f., FI. Java 3 (1968) 119. - Caladium indicum C. Koch, Berl. Allg. Gartenz. (1857) 136. - Type: Arum indicum sativum Rumph., Herb. Amboin. 5 (1747) t. 106 (lecto; selected by Nicolson, 1979). Colocasia indica var. atroviridis Hassk., Flora 25(2), Beibl. 1 (1842) 8 = Alocasia macrorrhizos (L.) G. Don. - Type: None designated; based on Rumphius’ very brief description of a second ‘species’ of Arum indicum sativum Rumph., Herb. Amb. 5 (1747) 308. [Colocasia odora (‘odorata’) sensu auct. non (Roxb.) Brongn.: Hassk., Flora 25 (2), Beibl. 1 (1842) 9; see Hay (1996)]| Arum montanum Roxb., FI. Ind. 3 (1832) 497. - Colocasia montana (Roxb.) Kunth, Enum. Pl. 3 (1841) 40. - Alocasia montana (Roxb.) Schott, Oesterr. Bot. Wochenbl. 4 (1854) 410. - Type: India, Orissa, Circars, Roxburgh drawing No. 248 (CAL, K). Plate 2. 284 Gard. Bull. Singapore 50(2) (1998) FCOVES NORDUNCHIANE Norang Cron Cae gee el pelt ; Plate 2. Type of Arum montanum Roxb. Icones Roxburghianae 248 (K, copyright R.B.G., Kew). [Alocasia grandis N.E. Brown, Gard. Chron. n.s. 25 (2) (1886) 390, nom. illeg., non Alocasia grandis Clemenc., Rev. Hort. 40 (1868) 380 (see below)]. Massive pachycaul with the stem decumbent or erect, to 4 m tall; petioles to 1.3 m long, sheathing in lower !/3-!/2; blades ovato-sagittate, bluntly triangular in general outline, held more or less erect, with the margin entire to very slightly; anterior lobe ca. 70 cm to over 1 m long, ca. 60- 90 cm wide at base, with ca. 9 rather distant primary lateral veins on each side of the anterior costa diverging at ca. 60°; glands in axils of primary veins on abaxial side distinct; secondary venation flush with the lamina or but slightly raised abaxially, not forming interprimary collective veins or these poorly defined; posterior lobes ca. 1.3—!/2 the length of the anterior, somewhat rotund, often overlapping; inflorescences paired among the leaf bases, subtended by membranous cataphylls; peduncles barely exceeding the cataphylls at anthesis; spathe rather variable in length, ca. 13-35 cm long, constricted about !/6th of the way from the base; lower part green, Alocasia in West Malesia and Sulawesi 285 ovoid; limb broadly oblong-lanceolate, cowl-like at anthesis, later reflexed, then deliquescent, membranous, pale yellow; spadix slightly shorter than the spathe, shortly stipitate; female zone 1-2 cm long, ca. 1.5 cm diam.; ovaries pale green, ca. 3 mm diam.; stigma sessile, 3—5-lobed, the lobes conic, yellow; sterile interstice slightly shorter than to equalling the female zone, whitish, very slightly narrowed corresponding to the spathe constriction, composed of rhombo-hexagonal synandrodia ca. 2.5 mm diam., the lower ones paler, incompletely connate or with a central hole, the upper ones resembling synandria; male zone cylindric, ca. 3-7 cm long, ca. 2 cm diam., whitish; synandria rhombo-hexagonal, convex-topped due to cap-forming synconnective, ca. 2 mm diam; appendix yellowish, slightly thicker than the male zone at the base, thence tapering, equalling to considerably exceeding half the length of the spadix, staminodial; fruiting spathe ca. 8 cm long, longitudinally dehiscent, green; berries scarlet. Distribution: IndoMalesia to Oceania. It is not clear where, if anywhere, this species occurs wild. It has evidently been distributed widely in tropical Asia in prehistoric times as a subsistence crop and is now pantropical by introduction as an ornamental. Habitat: Road sides, waste places, gardens, mostly in wet sites at low to medium elevation. Notes: 1. A number of ornamental varieties have been recognised, which were discussed by Furtado (1941). As cultivated plants they should be named as cultivars within the domain of the International Code of Nomenclature for Cultivated Plants. One element, Alocasia macrorrhizos var. rubra (Hassk.) Furtado, was discussed further by Bunting & Nicolson (1963), who considered it sufficiently distinct from other varieties as to be recognised as a distinct species - Alocasia plumbea van Houtte (FI. des Serres 21 (1875) t. 2206). While differing somewhat from other forms of Alocasia macrorrhizos in dimensions and leaf shape, it is not known except in cultivation, and I doubt that it is more than a sport of A. macrorrhizos. Alocasia grandis N.E. Br., introduced to cultivation in Britain from ‘the East Indian Islands’ appears to be the same element. 2. This species does not appear to be a wild plant at all in Malesia, though the situation in the Philippines, where there is indirect evidence for natural hybridisation with A. portei Schott, is unclear. No collections exist that are from places unequivocally away from human habitation, and I have never encountered it in the field except closely associated with human settlement. Arguably Alocasia macrorrhizos is a cultigen. 286 Gard. Bull. Singapore 50(2) (1998) 3. Alocasia grandis N.E. Brown, based on a specimen Brown took from a plant cultivated at Kew ex Hort. Bull, 10 Sep 1886 (K!), is evidently the same or very similar to Alocasia macrorrhizos var. rubra. It is, however, illegitimate due to the pre-existence of Alocasia grandis Clemenc. The identity of the latter is not clear. 4. Discussion of the identity of Alocasia montana (Roxb.) Schott can be found under Alocasia flemingiana. The type is reproduced in PI. 2. Other specimens: PENINSULAR MALAYSIA: Negeri Sembilan, Kuala Pilah Rd., Burkill 1648 (SING); Kelantan, Khota Baru, Gimilette s.n. (SING); Selangor, Genting Simpah F.R., Nicolson 1145 (US), Telok Gading, Md Nur 3002 (SING). SINGAPORE: Botanic Gardens, Sinclair 8340 (K, US). SABAH: Sandakan, Kadir A2698 (K, L); Kelawat, Kota Belud, Keith 6787 (SING); Kuala Napagun, Keith 9345 (SING); JAVA: Buitenzorg cult., Bakhuizen 8169 (L); Java, Buitenzorg, Hallier 1893 (L); Java, Horsfield s.n. (K); Java, Batavia, Junghuhn s.n. (L); Java, Kuhl & van Hasselt s.n. (L); Java, Zollinger 472 (L). 17. Alocasia inornata Hallier f. Alocasia inornata Hallier f., Meded. Herb. Leiden 26 (1915) 7; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 86. - Neotype: Indonesia, Sumatera, Riau Province, Tigapulu Mts., 5 km W of Talanglakat on Rengat-Jambi Road, Bukit Karampal area, 13 Nov 1988, J.S. Burley, Tukirin et al. 1444 (K, neo; KEP, SING, US, isoneo; designated here). Alocasia nobilis Hallier f., Meded. Herb. Leiden 26 (1915) 6. - Type: Indonesia, Java, Cult. Hort. Bogor ex Deli, Sumatera, 14 Jan 1896, H. Hallier s.n. (L, holo) Robust herb to 1.3 m tall; stem erect to decumbent, to ca. 8 cm diam; leaves several together, glabrous to minutely pubescent on the petiole and abaxial lamina; petiole ca. 50 cm — 1 m long, sheathing in the lower 1/3-1/2, pale green to pale chocolate brown with the apical 1-6 cm deep purple, or the whole purple throughout, otherwise more or less unmarked; blade sagittate to broadly ovato-sagittae, distinctly but not very thickly leathery, to ca. 50 (-80) cm long and up to 40 (—50) cm wide, slightly olive mid-green to suffused brownish chocolate to deep purple throughout and slightly shining adaxially, paler and dull abaxially; anterior lobe widest at or slightly above the base, the tip mostly obtuse and apiculate for ca. 1 cm; anterior costa with 4~7 primarly lateral veins on each side diverging at ca. 65—45°, with prominent glands in their axils abaxially; secondary venation flush with the lamina abaxially and adaxially, forming not very well-defined slightly sinuous to almost straight interprimary collective veins; posterior lobes ca. 1/2 the length of the anterior, rounded to acute, the inner side Alocasia in West Malesia and Sulawesi 287 lanceolate to narrowly ovate to subrhomboid; posterior costae naked in the sinus for ca. 1.5 cm; inflorescences rather slender, (2—)several together, forming a cluster in the centre of the crown and not interspersed with foliage leaves; peduncle mostly hidden within leaf sheaths and cataphylls, pubescent if the leaves are; cataphylls semi-persistent; spathe ca. 12 cm long; lower spathe rather narrowly ovoid, green, ca. 2 cm long, separated from the limb by a rather weak constriction level with the basalmost part of the male zone of the spadix or with the sterile interstice; limb to ca. 4 cm wide, as first erect and cucullate with the basalmost part reflexed, then wholly reflexed, pale greenish yellow to white, deciduous after anthesis; spadix subequalling the spathe, stipitate for ca. 3 mm; female zone ca. 1 cm long, ca. 7 mm diam.; ovaries flattened subglobose, somewhat lobed, green, ca. 1.5-2 mm diam.; stigma sessile, about 1.2 mm wide, of 3—5 rounded lobes, white; sterile interstice basally as broad or broader than female zone, distally attenuate, ca. 7 mm long, composed of numerous white to pale apricot synandrodia, the lowermost sometimes with the staminodes incompletely united, the rest closely resembling the synandria; male zone cylindric, ca. 1.5—2 cm long, ca. 7 mm diam., white; synandria hexagonal, 1.5 mm diam., slightly convex-topped; thecae overtopped by synconnective and opening by short apical slits; appendix to 7.5 cm long, slender, basally isodiametric with male zone, tapering very gradually to a point, pale orange- yellow to yellowish ivory; fruiting spathe green, to ca. 6 cm long, ovoid. Distribution: Southern parts of Peninsular Malaysia (not recorded from Singapore) and Sumatera. Habitat: This species has wide ecological amplitude and is found in disturbed places in forest, scrub, swampy areas, river banks, sometimes on limestone, from sea level to ca. 1200 m altitude. Notes: 1. Alocasia inornata was described from a sterile plant cultivated at Bogor, originally collected by Jaheri from Deli, Sumatera. Preserved material neither of the cultivated plant nor of the original field collection has been located. However, Hallier’s description includes reference to the leaves being pubescent and the petiole being purple at the apex, and there can be little doubt that this name is applicable to the species defined here. The neotype here designated is weakly pubescent, reference is made in the notes to the petiole being purple at the top (this feature becomes obscured on drying) and the specimen includes a well-preserved inflorescence. While the colour of the petiole apex may seem trivial, purple pigmentation is present at that point in all living plants I have seen of this species, both in Sumatera and in Peninsular Malaysia, and appears to be a diagnostic feature. 288 | Gard. Bull. Singapore 50(2) (1998) The exception is cases where the entire leaf is suffused with purple - the condition of the type of A. nobilis. Pubescence on the other hand is not a constant feature of Sumateran specimens and apparently is absent from Peninsular Malaysian ones. 2. Alocasia nobilis was also described from cultivation at Bogor and the cultivated plant also originated from a collection by Jaheri from Deli. The description includes reference to the inflorescence, though this does not appear to have been preserved. The leaf, besides being suffused purple throughout, is at the extreme of narrowness overall and reduction of the posterior lobes for this species, but it connects with rather than falls outside the rest of the morphological range, Lorzing 5061 being an intermediate example. As might be expected if the leaf is suffused purple, the spathe is described as being so too, where typically in A. inornata it is green in the lower part and pale greenish yellow to white in the limb. 3. It was possible that the name Alocasia ovalifolia Ridl. had priority, if syntype material of that species could have been located that is conspecific with A. inornata. The issues around this uncertainty are discussed under A. puber (q.v.) in the synonymy of which A. ovalifolia has been placed on the basis of the identity of the only syntype so far located and on which A. ovalifolia has been lectotypified in order to dispose of the name. 4. The distinguishing combination of features of this species includes, in addition to the purple-topped petiole, the slender spadix, the spathe constriction corresponding with the basalmost part of the male zone or the interstice, the narrowly tapering appendix, the large, sessile, rounded 3-5- lobed stigma and syandria with the thecae overtopped by synconnective. Other specimens seen: PENINSULAR MALAYSIA: Pahang, Pulau Tioman, Burkill s.n. (SING), Cult. RBG Sydney, Acc. No. 893742 Goodwin 16 (NSW), Md Nur 18890 (BO, SING); Selangor, Forest Research Institute, Kepong. Croat 53294 (K); Cult. RBG Kew Acc. No. 472-82.05004 ex Selangor, Ulu Gombak, Hay s.n. (K); Cult RBG Sydney, Acc. No. 940101 ex Selangor, Batu Caves, Hay 9057 (NSW); Cult. RBG Sydney, Acc. No. 940338 ex Pahang, Semangoh Pass, Hay 9271 (NSW); Pahang, Fraser’s Hill, Henderson 11389 (BO); Perak, Gua Badak, Lenggong, Henderson 23836 (BO, SING); Selangor, Batu Caves, Ridley 8168 (K, SING), 13392 (SING); Selangor, Anak Takun, Templer Park, Sang JS33 (KEP). SUMATERA: Atjeh, NE of Kroengloeas, E of Troemon, Asdat 74 (L); G. Koerintji, Biinnenmeijer 9222 (BO) & 10354 (BO, L); NE Sebessi Island, Docters van Leeuwen-Reijnvaan 5139 (BO); East Coast, Karo Highlands, E of Berastagi, Hamel & Rahmat si Toroes 631 (L); Cult. RBG Sydney Acc. No. 970531 ex Lembah Anai Nature Researve, Hay & Yuzammi 13105 (NSW); Sibolangit, Lérzing 5061 (BO); Karohoogvlakte bij Lingga, Lérzing 6274 (BO); South Sumatera, Enggano, Liitjeharms 3856 (BO, GH, K, L) & 4745 (BO, K, L); W. Sumatera, Pesisir selatan Kerintji, Mt Kerintji, nr Pondok Patjet, Meijer 6219 (L); Batoe Island, Raap 8 (BO). Alocasia in West Malesia and Sulawesi 289 18. Alocasia alba Schott Alocasia alba Schott, Oesterr. Bot. Wochenbl. 2 (1852) 59; Miq., Fl. Ned. Ind. (1855) 210; Schott, Syn. Aroid. (1856) 48; Schott, Prodr. Syst. Aroid. (1860) 149; Engl., in A. & C. DC., Monogr. Phan. 2 (1879) 500; Hook. f,, Fl. Brit. India. 6 (1894) 528; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 84. - NEOTYPE: Alocasia alba Schott, Icones no. 86-88 (W, lecto; designated here) [fiche 68: d6-d8 in the microfiche edition]. Alocasia bantamensis Koord., Bull. Jard. Bot. Buitenzorg HI, 1 (1918) 162, figs 11-13; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 84; Koord., Exkurs.-Fl. Java 4 (1923) 195-196, fig. 395-396; Backer & Bakh.f., Fl. Java 3 (1968) 119. - TYPE: Indonesia, Java, Bantam, Danoe-moeras, 26 May 1912, Koorders 41445b (L, lecto; selected here). Alocasia crassifolia Engl., Pflanzenr. 71 (IV.23E) (1920) 82; Koord., FI. Tjibodas 6 (1922) 36; Burnett, Aroideana 7 (1984) 134, fig. 101. - TYPE: Indonesia, Java, cult. Hort. Bogor, Jan-Feb 1906, Engler 4101 (B, holo). [Alocasia macrorrhizos (‘-rrhiza’) sensu auct. non (L.) G.Don: Backer & Bakh-f., Fl. Java 3 (1968) 119]. Massive pachycaul, stout, up to 2 m tall; /eaves several together, held erect; petiole greenish, whitish at the sinus, with scattered yellowish glands, up to 170 cm long, sheathing for about !/3 its length, wings of sheath persistent, straight to recurved; blade thick, tough, usually slightly bullate, green above, light yellowish-green below, broadly ovato-sagittate to cordato-sagittate, margin entire; anterior lobe ca. 80 cm long, ca. 75 cm wide at base, the tip shortly acuminate; anterior costa with up to 11 primary veins diverging at - an angle of about 40°-60°, prominent on both surfaces, with conspicuous small flat glands in the axils on the abaxial side; secondary veins sunken adaxially, prominent abaxially, interprimary collective vein well-defined; submarginal vein 1-2 mm from margin; posterior lobes obtuse, ca. 45 cm long from the sinus; inflorescences in groups of up to ca. 10 at a centre of leaf crown, not interspersed with foliage leaves (but occasionally pairs produced singly); peduncle up to 38 cm long, like the petiole with scattered small broadly elliptic glands; spathe to ca. 17 cm long, constricted at level of sterile zone of spadix, lower spathe broadly ovoid-cylindric, ca. 5 cm from the base, green, the limb reflexed between male zone and sterile zone, thinly leathery, greenish yellow to greenish white; spadix cylindrical, ca. 15 cm long, sessile to very shortly stipitate; female zone ca. 1.7—2.2 cm long, 1-1.4 cm wide, with ca. 60-100 close-packed pistils; ovary green, ovoid to subglobose, 2-3 mm in diam.; style abruptly-differentiated from 290 Gard. Bull. Singapore 50(2) (1998) ovary and c.. 1 mm long, to lacking; stigma white, 2-3-lobed, the lobes rounded; sterile interstice ca. 1-1.6 cm long, with ca. 5—6 whorls of rhombo- hexagonal synandrodia, the lowermost whorls isodiametric with female zone and resembling connate staminodes, the upper portion attenuate and resembling sterile synandria; male zone white, ca. 2.5-3.5 cm long, ca. 1-1.5 cm wide; synandria white, swollen-topped, rhombo-hexagonal, 2 mm diam., thecae overtopped by synconnective, opening through apical slits; appendix ivory, ca. 5.5—-8 cm long, tapering, smooth to faintly rugose and composed of irregular sinuous staminodes, basally isodiametric to or slightly narrower than the male zone; fruiting peduncle to ca. 25 cm long; fruiting spathe broadly ovoid, to 6 cm long: fruit ellipsoid, orange, 5 mm. Distribution: Java, widespread at low to medium elevation. Plants sighted by me in 1996 near Telukbetung along the Palembang road in SE Sumatera may also be of this species. Circumstances prevented my collecting specimens. Habitat: In open spots in forest and beside roads and fields, mainly in swampy sites, but also on well drained soils. Notes: 1. Schott evidently described this species from cultivated material — known in horticultural circles of the time as Colocasia alba and Homalomena alba (the former apparently never validly published, the latter not in the sense of H. alba Hassk.). Schott did not know the origin of the plant, though he indicated it was probably Malesian (‘verosimiliter in insulis Archipelagi Indiae orientalis’). No collector or collection was cited in the protologue, but illustrations were prepared which are here designated as the neotype. They show the distinctive venation of this species, and give the clear impression of its characteristically coriaceous and slightly bullate leaves. That the plant illustrated was very probably the same one as that described in the protologue is suggested by the illustration of the ovaries (in Ic. 87) which appear abortive and which were described by Schott in the protologue as ‘ovariis (in spadicibus omnibus speciminis nostri) rudimentariis’. This appears to be a teratum: the pistils of other specimens of this species are normal. 2. Engler (1879: 500) attributed Alocasia alba to Sri Lanka, on the basis of a Burmann specimen at G. Although I have not seen this specimen, the attribution of A. alba to Sri Lanka appears to be erroneous, an opinion shared implicitly or explicitly by Brown (1884: 870), Hooker (1894: 528) and Trimen (1898: 360), who attributed it to Java. Moreover, Nicolson (1987: 55) did not include Alocasia alba in his treatment for Sri Lanka, Alocasia in West Malesia and Sulawesi 291 where the only species approaching it is the amply distinct A. macrorrhizos. 3. Backer & Bakhuizen (1968: 119) misapplied the name Alocasia macrorrhizos to what, from their description, specimen annotations and synonymy, is clearly this species. For the species correctly named A. macrorrhizos, they used its synonym A. indica (Lour.) Spach. What lead them to make the misapplication is not apparent. 4. The function of the distinctive glands on the petiole and peduncle, which resemble those in the axils of the primary veins, is not clear. At female anthesis the inflorescence is sweetly fragrant with the scent produced from the inside of the lower spathe. Other specimens seen: JAVA: Buitenzorg, Boerlage s.n. (L); W. Java, Batavia, Solear, Tjisoka, Eyma s.n. (L); Cult. RBG Sydney Acc. No. 892944 ex Bogor, Hay 4087 (NSW); W. Java, Curug Sawer, Hay & Yuzammi 14002 (NSW); Bawean, G. Tunggangan, Karta 30 (BO, L); Djapara, Ngarengan, Koorders 34996 (L); Hort. Bogor, Koorders 42804 (L); Preanger, Tasikmalaya, Pendjaloe, Koorders 44346 (L); Kuhl & van Hasselt s.n. (L); Mousset 589 (BO); Bantam, Danoe Moeras, van Steenis 10513 (L); Hort. Bogor, Teijsmann s.n. (L); Kediri, Pandan, Djeroek, Thorenaar 290 (BO). 19. Alocasia balgooyi A. Hay, sp. nov. Ab Alocasia macrorrhizos (L.) G. Don costis posticis haud vel vix nudis, spathae lamina coriacea breviore, inflorescentia femina et interstitio neutro longioribus tenuioribus, synandriis minoribus, spatha fructifera rubra differt. - TYPUS: Indonesia, South Sulawesi, Soroako, Malili Road, 29 Jun 1979, M. van Balgooy 3812 (BO, holo; GH, K, L, iso). Robust to massive herb 1-3 m tall; rhizome stout (?diam.), clothed in fibrous leaf base remains; /eaves several together; petiole ca. 70-100 cm (?0r more) long, glabrous, sheathing in the lower !/3-!/2; blade broadly ovato-sagittate to cordato-sagittate, ca. 50-100 cm long, membranous; anterior lobe widest slightly above the base, the apex obtuse and apiculate; anterior costa with 5 (?or more) primary lateral veins on each side diverging at 45—70°; axillary glands inconspicuous; secondary venation not or hardly raised abaxially, not or hardly forming interprimary collective veins; posterior lobes 1/2-3/4 the length of the anterior, obtuse, the inner sides ovate to more or less rhomboid; inflorescences in pairs ?interspersed with foliage leaves; peduncle ca. 30 cm long; spathe ca. 17—20 cm long, constricted 3—4 cm from the base with the constriction corresponding with the sterile interstice of the spadix; lower spathe narrowly ovoid to ovoid, thick; limb oblong-lanceolate, coriaceous, erect and later reflexed and somewhat persistent, thence deciduous, greenish cream; spadix somewhat shorter than 292 Gard. Bull. Singapore 50(2) (1998) to subequalling the spathe, ca. 14 cm long, stipitate for ca. 4 mm and the stipe inserted obliquely; female zone 2 cm long, ca. 5 mm wide; pistils close-packed, globose-cuboid, ca. 2 mm diam.; stigma sessile, bluntly 3- lobed, ca. 1 mm diam.; sterile interstice 1-1.5 cm long, distinctly attenuate, ca. 3 mm thick, composed of numerous small synandrodia resembling the synandria; male zone 3-4 cm long , cylindric, 5-6 mm diam., cylindric; synandria very numerous, small, ca. 1 mm diam.; thecae overtopped by synconnective; appendix 4—5.5 cm long , slightly narrower than the male zone, deeply longitudinally channelled, tapering to a point; fruiting peduncle to ca. 50 cm long; fruiting spathe broadly ovoid to spindle-shaped, 6-12 cm long, creamish green, becoming bright red. Distribution: Endemic to Sulawesi. Habitat: Low to mid-elevation forests, sometimes in swamp forest (Kjellberg 2396), or on slopes, often in disturbed places and on ultrabasic soils; sea level to 1200 m altitude. Notes: 1. The description is pieced together from fragmentary dry specimens. This species appears at least superficially similar to A. macrorrhizos, though clearly differing in the leaves with the posterior costae usually not naked | in the sinus, the thick and relatively short spathe limb, longer peduncle, the relatively longer and narrower female and sterile zones of the spadix, the smaller and more numerous synandrodia and synandria, and the red fruiting spathe. 2. Alocasia balgooyi is named for Dr Max van Balgooy, who first drew attention to this new species. Other specimens seen: SULAWESI: Menado, Poso, above baroega S. Malei, Eyma 1670 (BO, L); nr Malino, Eyma 3460 (BO, L); Malili, Kjellberg 2396 (BO) & 2116 (BO); Matamo Lake nr Soroako, NE of Malili, Meijer 11120 (BO); Larona, W of Towuti lake, E of Malili, Meijer 11298 (BO); N Sulawesi, Dumoga Bone National Park, Gorontalo Distr., Sg. Olama below G. Gambuta, Millikin 976 (K); Tangguma, Poli-polia, Kolaka, Prawiroatmodjo & Maskuri 1530 (BO); N shore of Lake Matano, E of Nuha, de Vogel 5840 (K). 20. Alocasia flemingiana Yuzammi & A. Hay, sp. nov. Ab aliis speciebus javanicis statura minore, folii lamina adulta haud peltata nervo intramarginale praedita, interstitio neutro et spathae constrictio congruentibus, stigmate lobato sessili, synconnectivo expanso differt x TYPUS: Indonesia, West Java, Ciseeng, 25 km NNW of Bogor, 28 Jan 1961, Nicolson 848 (L, holo; BO, US, iso). Alocasia in West Malesia and Sulawesi 293 [Colocasia montana auct. non (Roxb.) Kunth: Hassk., Pl. Jav. Rar. (1848) 148. ] [Alocasia montana auct. non (Roxb.) Schott: Miq., Fl. Ned. Ind. 3 (1855) 209, p.p., quoad loc. ‘Java’; Engl., in A. & C. DC., Monogr. Phan. 2 (1879) 499, p.p., quoad loc. ‘Java’; Hook.f, Fl. Brit. India 6 (1894) 525, p.p., quoad loc. ‘Java’; Koord., Exkurs.-Fl. Java 1 (1911) 261; K.Krause & Engl., Pflanzenr. 71 (I1V.23E) (1920) 77, p.p., quoad loc. ‘Java’; Koord., Exkurs.-Fl. Java 4 (1923) 194, fig. 393, non fig. 394 - i.e. Alocasia montana s.s. (= Alocasia macrorrhizos)| [Alocasia heterophylla sensu auct. non (Presl) Merrill: Backer & Bakh.f,, Fl. Java 3 (1968) 120.] Small herb, ca. 50 cm tall; rhizome to ca. 3.5 cm diam.; leaves several together; petiole green, sometimes mottled reddish purple, sometimes purple-streaked, 25-55 cm long, sheathing in the lower about !/4—1/3 of its length; blade mid-green adaxialy, green-yellowish abaxially, sagittate to broadly ovato-sagittate, thin, membranous, glabrous on both surfaces; anterior lobe ca. 25 cm long, ca. 19 cm wide, the tip acuminate, ca. 1 cm long, anterior costa with 3 or 4 primary veins on each side, diverging at ca. 40°—60°, prominent abaxially; primary veins often bearing small flat glands in the axils abaxially, running to a distinct submarginal vein ca. 1-3 mm from margin; secondary veins flush to lamina, interprimary collective veins absent or poorly differentiated; posterior lobes acute, up to ca. 16 cm long, inner sides elliptic to obovate; posterior costae diverging at ca. 90-110°, naked in the sinus for 0-1 cm; inflorescences in pairs interspersed with foliage leaves; peduncle, ca. 2/3 the length of the petiole at anthesis, elongating in fruit, up to ca. 31 cm long; spathe white to greenish white, ca. 10-15 cm long, lower spathe ovoid, ca. 2-4 cm long, constricted level with top of sterile zone of spadix (to half way along male zone); limb narrowly- oblong and falling after anthesis; spadix somewhat shorter than to subequalling the spathe, ca. 8-11 cm long, slender, stipitate for ca. 0.5 cm; female zone to 1 cm long, with ca. 40 pistils; ovary globose, ca. 2 mm diam.; stigma sessile, ca. 1 mm diam., 2—3 lobed, the lobes bluntly pointed; sterile interstice hardly to somewhat attenuate, ca. 5 mm long, ca. 3 whorls of rounded to rhombo-hexagonal synandrodia; male zone 1.5 cm long, 5 mm wide; synandria rhombo-hexagonal to rhomboid, 1 mm diam., thecae somewhat displaced to overtopped by synconnective, opening through apical slits; appendix 6.5 cm long, tapering, cream; fruiting spathe becoming white. 294 Gard. Bull. Singapore 50(2) (1998) Figure 9. Alocasia flemingiana Yuzammi & A. Hay RBG Sydney Acc. No. 980045. A. habit; B. venation; C. inflorescence with part of spathe removed; D. pistils; E. neuter organs; F. synandria. - Scale: bar to A, B = 8 cm, to C = 32 mm, to D, E, F = 6 mm. Alocasia in West Malesia and Sulawesi 295 Distribution: Endemic to Java; widespread in West Java and sporadic in Central Java. Habitat: Found in teak-forest, swamp-forest, disturbed forest, on volcanic soils, sometimes over limestone, from sea level to ca. 1000 m altitude. Murata et al J-2042 (BO) recorded that this species was found on ‘rocky sea coast’. This seems an unlikely habitat for Alocasia, and possibly there has been an error in labelling of this specimen. Notes: 1. The rationale behind the application of Colocasia montana or Alocasia montana, both based on Arum montanum - which Roxburgh had coined for a plant from the Northern Circars in the Indian state of Orissa - to Javan material remains obscure, doubly so since the identity of Arum montanum is itself unclear. The earliest misapplication appears to have been that of Hasskarl (loc. cit.) who made the connection on the basis that his Javan plant and Arum montanum were both ‘stemless’, followed by Miquel and, much later, Koorders. Hooker (1894) also noted in his Flora of British India that A. montana extended to Java. That opinion was not followed by Backer & Bakh. f. (1968) - though they in turn misapplied Alocasia heterophylla (Presl) Mertr. to this species. Arum montanum Roxb. first appeared in Hortus Bengalensis (1814: 65) as a nomen nudum, and the first valid publication was in Roxburgh’s Flora Indica 3 (1832: 497). There, he wrote: ‘T long considered this to be A. |Arum] macrorrhizon, but changed my opinion on observing that Forster, who must have seen and examined that species in its recent State, says, the flowers are hermaphrodite; there being six sessile, twin anthers surrounding each germ, and that the stigma is orbicular. There are no traces of stamina, anthers or glands round the germs of my [Roxburgh’s] plant; and the stigma is regularly three or four-lobed. In short, a very perfect Arum, or Caladium according to Ventenat’. Roxburgh did not record from where he took Forster’s observation, but it was probably his dissertation on esculent plants (Forster, 1786), where he said of Arum macrorrhizon |I am indebted to Dan Nicolson for this]: ‘Fructificatio a charactere generico aliquanto recedit, floculis in spadice omnibus et singulis hermaphoditis...COR. nulla.’ STAM. Filamenta nulla. Antherae sex, spadice adnatae, didymae, singulo stylos cingunt./ PIST. Germen subrotundum. Stylus solitarius, brevis, crassiusculus, apice depressus. Stigma maculata orbiculata in apice styli....’ 296 Gard. Bull. Singapore 50(2) (1998) Arum macrorrhizon L. is the basionym of Alocasia macrorrhizos (L.) G. Don, and there is little doubt that either Forster’s observation, or the way Roxburgh interpreted it, or the identification of the plant Forster observed, was incorrect. That being so, the probability is raised that A. montana is identical with A. macrorrhizos, as Roxburgh had originally thought. Neither Kunth, Schott, nor Engler and Krause, nor authors of floristic accounts of Alocasia in India have cited any additional specimens when treating Alocasia montana, simply reiterating descriptions obviously based on Roxburgh’s protologue and the incomplete and rather naive illustration which, unless pertinent preserved material comes to light (Forman’s (1997) account of Roxburghian species does not include A. montanum), will form the type. In the absence of evidence to the contrary, it would seem reasonable to treat Alocasia montana (Roxb.) Schott as an Indian synonym of A. macrorrhizos (q.v.), at least provisionally; however, epitypification of Arum montanum will be dealt with elsewhere. Suffice to say that, even without unequivocally disposing of the name Alocasia montana (Roxb.) Schott, there appears no tangible basis for applying the name of a ‘species’ known from a bare description and a poor painting of a plant from Orissa and resembling Alocasia macrorrhizos, to plants of a species apparently endemic to Java and bearing little resemblance to A. macrorrhizos, on the sole basis of ‘stemless’ habit - a characteristic of juveniles of most if not all species of Alocasia. 2. Alocasia flemingiana was incorrectly identified as Alocasia heterophylla (Presl) Merrill by Backer & Bakh. f. (1968). However, these species are readily distinguished by the following features: A. flemingiana has the lamina ovato-sagittate, with the margin entire, the leaf is only peltate in juvenile plants, and the sterile zone has relatively small synandrodia; Alocasia heterophylla has the lamina narrowly (hasto-) sagittate, sometimes deeply peltate in adult plants, with the margin sometimes undulate and the sterile zone with large synandrodia filling the upper part of the lower spathe chamber. 3. The specific epithet acknowledges Conrad D. Fleming’s generous sponsorship of field work on Malesian Araceae, including Yuzamm1’s field work in Java during December 1997. Other specimens seen: JAVA: Hort. Bogor, Adelbert 400 (L); Banjumas, Tjilatjap, Backer 21009 (BO); Java, Batavia, Backer 34987 (BO); Java, Preanger, Tjadas Malang, Bakhuizen. 1379 (BO); Preanger, Tjadas Malang, Tjidadap, Tjibeber, Bakhuizen 1930 (BO); Madjalengka, Cirebon, Beumée 1753 (BO); Pekalongan, Margasari, Beumée 1767 (BO); Banjumas, Beumée 4845 (BO); Batavia, Krawang, Beumée 5397 (BO); Bidara Tjina, Edeling s.n. (BO); Tjiampea bij Buitenzorg, Koorders 30810b (BO); Pelabuhan Ratu, Koorders Alocasia in West Malesia and Sulawesi 297 34459b (BO), Koorders 34463b (BO), Nicolson 935 (BO, US), Nicolson 957 (BO); Cultuur Sepakaeng, Oengaran, Telomojo, Ambarawa, Semarang, Koorders 35968b (BO); Depok, Buitenzorg, Koorders 42817b (BO, L); Kranhau, West of Pelabuhan Ratu, W. Java, Murata, Kato, Mogea J-2042 (BO); Depok, W. Java, van Ooststroom 12609 (L); Tjilatjap, Riviere s.n. (L). Java, Preanger, Tjibodas, Sapiin 85 (BO); Tjirebon, Telaga Erang, Vermeulen 6 (BO); Miramere, Pamempek, West Java, Yuzammi 297042 (NSW); Miramere, Pamempek, West Java, Yuzammi 297043 (NSW). 21. Alocasia arifolia Hallier f. Alocasia arifolia Hallier f., Bull. Herb. Boiss. Sér 2, 1 (1901) 670, fig. 11; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 79. - Type: Cult. Hort. Bogor ex Sumatera, Deli, Permandian, 1 Apr. 1896, H. Hallier 105d, (BO, holo; Herbarium Bogoriense sheets 100563 and 100564). Herb to 70 cm tall; stem erect to decumbent, to ca. 30 cm long, 2-4 cm diam.; /eaves several together; petiole ca. 20-50 cm long, finely but densely puberulent to finely scabrid to glabrous, sheathing in the lower ca. !/4—!/3, mid-green to purplish brown; blade sagittate to hasto-sagittate, somewhat glossy mid/dark green adaxially, paler below, membranous to thinly coriaceous, to ca. 30 cm long; anterior lobe to ca. 22 cm long, widest at base, to 16 cm wide; anterior costa with 3—5(-—6) primary lateral veins on each side, diverging at ca. 45-60°, with inconspicuous axillary glands, sometimes abaxially puberulent; secondary venation flush with lamina when fresh, somewhat prominent abaxially when dry, fine but abaxially conspicuous, forming undulating to zig-zag interprimary collective veins and running to a rather conspicuous submarginal vein ca. 1.5—2 mm from the margin; posterior lobes acute, to ca. 15 cm long; posterior costae diverging at obtuse to very obtuse angles, naked in the sinus for up to 3 cm; inflorescences paired, the pairs interspersed with foliage leaves; peduncle to ca. 6 cm; spathe 5.5—-9 cm long; lower spathe 1—1.5 cm long, separated from the limb by an abrupt constriction, ovoid; limb oblong to oblong lanceolate, apiculate for up to 1 cm, greenish white, sometimes suffused purple-brown, erect, then reflexed; spadix subequalling the spathe, 4.5—-8 cm long, shortly stipitate; stipe white, ca. 3 mm long; female zone ca. 8 mm long; ovaries subglobose, green, ca. 1.5 mm diam.; style very short, ca. 0.5 mm long; stigma white, 3—4-lobed, the lobes rounded; sterile interstice ivory, hour-glass shaped, corresponding with spathe constriction, ca. 7 mm long, ca. 3 whorls of rhombo-hexagonal synandrodia; male zone short, about equalling female zone, ca. 6 mm diam.; synandria ivory, rhombo- hexagonal, 2-3 mm diam.; thecae opening by apical pores overtopped by synconnective; appendix somewhat constricted at base, the rest slightly narrower than male zone, finally tapering to a point, pale apricot; fruiting peduncle to ca. 20 cm long; fruiting spathe ovoid, ca. 2.5 cm long. 298 Gard. Bull. Singapore 50(2) (1998) Distribution: Endemic to Sumatera, recorded from few, scattered localities. Habitat: Low montane forest 400-1300m alt., often near streams, but not rheophytic. Notes: 1. The two sheets that comprise the type, as interpreted here, consist of a leaf and an inflorescence respectively. However, it is not entirely clear that they represent the same collection (the second is undated and has no collection number), though they are both collected from the Bogor Botanic Garden. I am in no doubt that they are of the same species and they are both determined as A. arifolia by Hallier f. It would be desirable to designate an unambiguous epitype, but, as yet, complete, authentically provenanced material is not available. 2. Relationships of Alocasia arifolia are not readily apparent, but the arrangement of inflorescences and the synandria with the synconnective overtopping the thecae suggest alliance to Javan A. flemingiana. Other specimens seen: SUMATERA: Cult. RBG Sydney Acc. No. 942737 ex cult. Hort. Leiden Acc. No. 940819 ex Sumatera, Vogel s.n. (NSW); Cult. RBG Sydney Acc. No. 970498 ex West Sumatera, G. Gadut, Hay et al. 13069 (NSW); Mentawi Islands, Siberut Island, Jboet 26 (BO); Lampung Prov., Mt Tanggamus, Jacobs 8265 (L); Aceh, Gajolanden, Bivak Aer Poetih waterfall, nr Pendeng, van Steenis 9283 (BO); Longiloba Group Species 22—24 Growth pattern strongly rhythmic, with a pronounced delay between flowering and resumption of leaf-production; cataphylls thinly membranous, degrading into rather sparse fibres; /Jeaves solitary or few together, the blades mostly peltate and pendulous, often purple-backed and/or with white major venation; interprimary collective veins absent to very pronounced and zig-zagged; spathe limb opening wide and deciduous; spadix stipitate; stipe white; ovaries green with white to yellowish stellate stigmas; sterile interstice attenuate and corresponding with spathe constriction; male zone ivory; thecae not overtopped by synconnective; appendix pale orange-pink to yellow, occasionally ivory. Distribution: About four species from Indochina to West Malesia, the Philippines and Sulawezi. Note: This group coheres on the basis of its highly uniform inflorescence Alocasia in West Malesia and Sulawesi 299 morphology, strongly rhythmic growth and membranous, fibrous cataphylls, and leaves with usually peltate, often dark green, purple-backed and white veined blades. There is nevertheless a great deal of variation in leaf blade shape, colour and venation. It is represented by three morpho- geographically circumscribable species - Philippine A. sanderiana W. Bull and A. boyceana A. Hay ined. and Sulawesi A. suhirmaniana Yuzammi & A. Hay. The fourth ‘species’ is a taxonomically intractable complex centred on West Malesia, extending into mainland Asia to the north and Sulawesi to the east, here treated as the Alocasia longiloba complex. Seventeen names have been proposed for Malesian elements within this complex. 22. Alocasia longiloba Miq. Alocasia longiloba Miq., Bot. Zeit. 14 (1856) 561 & FI. Ind. Bat. 3 (1856) 207; Schott, Prodr. Syst. Aroid. (1860) 153; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 506; Hook.f, Fl. Brit. India 6 (1894) 527; Hallier f., Bull. Herb. Boiss. 7 (1898) 607; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179; K. Krause & Engl., Pflanzenr. 71 (IV.23E) (1920) 103; Backer & Bakh.f., Fl. Java 3 (1968) 118); M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156. Type: Indonesia, Java, Tjikoja, 15 Aug 1856, Zollinger 601 (L, holo; B, BM, K, P iso). Caladium veitchii Lindley, Gard. Chron. (1859) 740. - Alocasia veitchii (Lindley) Schott (‘veitchi’), Ann. Mus. Lugd.-Bat. 1 (1863) 125; Koord., Exkurs.-Fl. Java 1 (1911) 261. - Alocasia lowii var. veitchii (Lindley) Engl. in A. DC., Monogr. Phan. 2 (1879) 508; Engl. & K. Krause, Pflanzenr. 71 (IV .23E) (1920) 107. Neotype: Java, without date, L sheet nos. 898. 87 115 &.898. 87 116, Kuhl & van Hasselt 12 (L, designated here). Caladium? lowii Lem., Ill. Hort. 10 (Jan 1863) descr. ad t. 360. Type: III. Hort. 10 (1863) t. 360. Alocasia lowii Hook., Curtis’s Bot. Mag. (May 1863) descr. ad t. 5376; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 508; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178 (‘lawii’); Ridl., Mat. Fl. Mal. Pen. 3 (1907) 18; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 106; Ridl., Fl. Mal. Pen. 5 (1925) 98; M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156. - Type: Cult. RBG Kew ex Hort. Low, Anon. s.n. (K!, holo). [Not based on Caladium lowii; see Hay in Hay et al., Blumea Suppl. 8 (1995) 16]. Alocasia korthalsii Schott in Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (Nov 1863) 124; Engl. in A. & C. DC., Mongr. Phan. 2 (1879) 509 & Bot. Jahrb. Syst. 300 Gard. Bull. Singapore 50(2) (1998) 25 (1898) 25; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 108, fig. 25; M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156. - Type: Indonesia, Kalimantan, G. Sakumbang, Korthals s.n. (L, holo). Alocasia singaporensis Linden, Gartenfl. 14 (1865) 252. - Neotype: Cult. R.B.G. Kew, 12 Feb 1879, N.E. Brown s.n. (K; designated here - see below). Alocasia lowii var. picta Hook.f., Curtis’s Bot. Mag. (1865) descr. ad t. 5497. - Type: Bot. Mag. (1865) t. 5497. Alocasia thibautiana Mast., Gard. Chron. 9 (1878) 527; N.E. Br., Ill. Hort. 28 (1881) 72, t. 439; N. E. Br., Gard. Chron. ser. 3, 17 (1895) 485, fig. 68. Neotype: Cult. RBG Kew ex Hort. Veitch, 25 Jan 1879, N.E. Brown s.n. (K, designated here). Alocasia amabilis W. Bull, Retail List 143 (1878) 9. - Neotype: Cult. RBG Kew ex Hort. Bull, 20 Sep 1878, N.E. Brown s.n. (K, designated here). Alocasia denudata Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 507; Hook.f., Fl. Brit. Ind. 6 (1893) 525; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 178; Ridl., Mat. Fl. Mal. Pen 3 (1907) 17; Engl. & K.. Krause, Pflanzenr. 71 (IV.23E) (1920) 100; Ridl., Fl. Mal. Pen. 5 (1925) 97; Corner, Gard. Bull. Sing. Suppl. 1 (1978) 73, 100. Type: Singapore, Gaudichaud 106 (G, holo, n.v.; P iso). Alocasia putzeysii (‘putzeysi’) N.E. Br., Ill. Hort. 29 (1882) 11; N.E. Br., Gard. Chron. ser. 2, 19 (1883) 501, fig. 75; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 108. - Type: Cult. RBG Kew ex Hort. Linden, Dec 1881, N.E. Brown s.n. (K, holo). Alocasia eminens N.E. Br., Gard. Chron. ser. 3, 1 (1887) 105; anon., Kew Bull. (1888) 91. - Type: Cult. RBG Kew ex Hort. W. Bull (no. 3954), 10 Nov 1886, N.E. Brown s.n. (K, holo). Alocasia watsoniana Mast., Gard. Chron. ser. 3, 13 (1893) 442; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 109; Burnett, Aroideana 7 (1984) 128, figs. 93-94. - Type; Gard. Chron. ser. 3, 13 (1893) 569, fig. 83. Epitype: Cult. RBG Kew ex Hort. Sander, April 1893, N.E. Brown s.n. (2 sheets) (K, designated here). Alocasia curtisii N.E. Br., Kew Bull. (1894) 347; Engl. & K. Krause, Alocasia in West Malesia and Sulawesi 301 Pflanzenr. 71 (I1V.23E) (1920) 106. - Type: Cult. RBG Kew ex Malaysia, Penang, 16 July 1894, N.E. Brown s.n. (K, holo). Alocasia cuspidata Engl., Bot. Jahrb. Syst. 25 (1898) 25; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 102. - Type: Indonesia, Kalimantan, Dusson Timor, Telang, Grabowski s.n. (B, holo). Alocasia denudata var. elongata Engl., Pflanzenr. 71 (IV.23E) (1920) 100, fig. 21, A-E. - Type: Singapore Botanic Garden, December 1905, Engler 3803 (B, holo; presumed destroyed). Neotype: Singapore, Seletar forest behind Rifle Range, 29 Aug 1948, J. Sinclair 5020 (E, designated here - see below). Small to robust herbs ca. 40-150 cm tall; terrestrial to lithophytic; rhizome generally elongate, erect to decumbent, often completely exposed, sometimes swollen and sub-cormous, ca. 8-60 cm long, 2—8cm diam., usually bearing remains of old leaf bases and cataphylls; growth markedly rhythmic with renewal growth delayed after flowering; vegetative modules often unifoliar (to 3-leaved), subtended by conspicuous lanceolate papery- membranous often purplish-tessellate cataphylls degrading to papery fibres; petioles glabrous, purple-brown to pink to green, often strikingly obliquely mottled chocolate brown, ca. 30-120 cm long, sheathing in the lower ca. 1/4 or less; blades often pendent, shape and venation extremely variable, peltate (except ‘denudata’), plain mid-green throughout to adaxially dark green and abaxially rich purple, often adaxially with the major venation white to pale grey-green and sometimes with the lamina bordering the main veins grey-green, hasto-sagittate and triangular in outline to ovato- sagittate and shield-shaped, 27—65(-—85) cm long x 14~ca. 40 cm wide, with the widest point anterior to the petiole insertion to near the tips of the posterior lobes; anterior costa with 4-8 primary lateral veins on each side, the proximal ones diverging at ca. 60—100°, the angle decreasing in distal veins and the course more or less straight to the margin to markedly acropetally deflected; axillary glands conspicuous abaxially at the junctions of the main veins and costae; secondary venation obscure to conspicuous abaxially, mostly arising from the primary veins at a wide angle then sooner or later deflected towards the margin, forming variously well-defined interprimary collective veins or these absent, concolorous with the abaxial lamina or sometimes markedly paler; interprimary collective veins when present weakly undulating to strongly zig-zagging at broadly acute angles; posterior lobes ca. 3/4 to 1/2 the length of the anterior lobe, when peltate united for (S—)10-66% of their length; posterior costae straight to pedately incurved; inflorescences (solitary to) paired, with up to 4 pairs in succession 302 Gard. Bull. Singapore 50(2) (1998) ESSN R Figure. 10. Alocasia longiloba Miq. s. |. Leaf blades showing a continuum of variation, some exemplifying entities recognised in the text. - A. ‘denudata’; D. ‘thibautiana’; F, ‘longiloba’; K, ‘lowi1’; M, Romhalse : = putseyzi is Te Go ne poets * (not all drawn to exactly the same scale). A. 901379 - Hay s.n., Singapore; B. 940485 - Hay 9332, Bintulu, Sarawak, C. 940102 - Hay 9058, Batu Caves, Se 82112 aes ms eee hea ‘ stones 9069. Fraser’s Hill, Pahang; F. 942735 - Vogel s.n., Sarawak; G. 950372 - Hay 10035, Kebun Cina, Sabah; H. 940138 - Hay 9142, Bukit Tanka, Negeri Sembilan; I. 904647 - Boyce 384, Tem Te ease ean Bsa Bukit Larut, Perak; K. 940064 - Hay 9019, Kaki Bukit, Perlis; L. 940047 - Hay 9001, Kangar, Perlis; M. 940462 - Hay 9308, Bintulu, Sarawak; N. 970528 - Hay 13002, Lembah Anai, West Sumatera; O. 942718 gel s.n., Bau, Sarawak; P. 970474 - Hay 13036, Kerinci Seblat NP, Jambi; Q. Lérzing 4804, Sibolangit, Sumatera; R. 920745 - Dearden s.n., Long Jowe, Sarawak. Six-figure numbers are all Royal Botanic Gardens Sydney Accession numbers with vouchers at NSW. without interspersed foliage leaves; peduncles ca. 8-18 cm long, usually resembling the petioles in colour and markings, erect at first, then often declinate, elongating and then erect in advanced fruit, subtended by a series of progressively larger cataphylls resembling those of the vegetative phase; spathe ca. 7-17 cm long, abruptly constricted ca. 1.5-3.5 cm from the base; lower spathe green, ovoid to subcylindric; limb pale green, membranous, lanceolate, canoe-shaped and longitudinally in-curved, eventually reflexing after male anthesis; spadix somewhat shorter than to subequalling the spathe, ca. 6-13 cm long, stipitate, with the stipe conic, whitish, to 5 mm long; female zone 1-1.5 cm long; ovaries subglobose, ca. 1.5—2 mm diam., green; stigma subsessile or on a slender style to ca. 0.5 mm long, white, acutely and conspicuously 3—4-lobed, the lobes pointed, more or less spreading; sterile interstice 7-10 mm long, narrower than the fertile zones, corresponding with the spathe constriction; lower synandrodia often with incompletely connate staminodes, the rest elongate rhombo- hexagonal, flat-topped; male zone subcylindric, somewhat tapered at the base, 1.2-2.5 cm long, 4.5-8 mm thick, ivory; synandria more or less hexagonal, ca. 2 mm diam., 4-6-merous; thecae opening by apical pores not overtopped by synconnective; appendix 3.5—9 cm long, about the same thickness as the male zone and demarcated from it by a faint constriction, subcylindric, distally gradually tapering to a point, faintly rugose to rugose in the lower part, very pale orange to bright yellow; fruiting peduncle to 25 cm long; fruiting spathe ovoid, ca. 4-7 cm long; fruits orange-red. Distribution: Indochina to West and Central Malesia excluding the Philippines. Habitat: In rainforest and swamp-forest floor, regrowth, on boulders in forest and on exposed cliffs and ravines at low to medium elevation. Notes: 1. In developing a classification that reflects what is at present known of this complex, I have avoided reducing its elements blithely to a single specics without a clear qualification that it is not equivalent to a ‘simple’ species of low variation content. However, the complex cannot have forced onto it overstated and simplistic hierarchical discontinuities that would be implied in recognising separate species and/or infraspecific taxa within it on the basis of currently available evidence (cf. Gentry, 1990). Like many Alocasia species, the elements of this complex freely hybridise in cultivation, but within this group hybrid series involving three or four ‘species’ are recorded (see Burnett, 1984), confirming their close relationship. Many of the formally named parent forms are highly 304 Gard. Bull. Singapore 50(2) (1998) ornamental and striking, and the horticultural community may. be irritated by their disappearing as named species. However, there is no reason why these names cannot be transposed into the nomenclature for cultivated plants as cultivars or cultivar groups. The background to the description of ‘species’ within this complex has been largely horticultural, through the introduction of the finest and most striking forms to European stove culture in the 19th Century. These forms represent what I have called the ‘peak variants’ in the complex, and I have used their types and associated nomenclature as a framework for the informal infraspecific classification proposed here. The entities proposed cannot at present be regarded as more than peaks in an overall continuum of variation and so not all specimens encountered can be categorically accomodated in this classification. Nevertheless, there are perceptible but incompletely resolved geographical patterns to the variation, and some ecological variation, which is somewhat but incompletely correlated with morphological variation and geographic pattern, discussed under the relevant ‘peak variants’. The key provided to these variants must be regarded as a guide only to ‘typical’ forms, and no pretence is made that by using it all specimens encountered can be unequivocally identified. 2. This complex can be considered an ochlospecies since, while there is a continuum of variation globally, at particular localities sharply differing forms may coexist and evidently behave locally as discrete sympatric ‘topospecies’. A rigorous analysis of variance might reveal statistically significant narrow discontinuities, which could form the basis for species distinctions at the global level. However, at present, carrying out such an anlysis is impeded by the inadequate and very uneven sampling over the range, exacerbated by the incompatibility of this genus with standard herbarium preservation methods. As a consequence of this incompatibility, collections consist of conveniently sized leaves or fragments which cannot be deemed comparable between individuals when the plants are known to show considerable plasticity of form depending on age and environmental factors. The local coexistence of distinct forms presupposes the existence of local reproductive isolating mechanisms regardless of whether or not these might translate into the defintion of species within the complex globally. The percentage of specimens preserved directly from the wild that bear inflorescences is extremely low and inadequate to form a basis for detecting isolation mechanisms based on flowering time. Moreover, there is an almost total dearth of ecophysiological data on finer aspects of phenology, which are known to be complex in this genus involving intricate patterns of flux in thermogenesis (Leick, 1915 - n.v., cited in Grayum, 1990) possibly Alocasia in West Malesia and Sulawesi 305 associated with scent production and the differentiation of pollinator prelerences. 3. Lindley described Caladium? veitchii from a cultivated plant obtained from Borneo. No original material has been located, and Lindley did not illustrate it. When Schott (1863) made the combination in Alocasia, he cited a collection by Kuhl and van Hasselt, from Java, in addition to noting the original provenance Borneo. Since Schott was the world authority on aroids at the time and had established an enormous collection of living plants, it is highly probable that he knew Caladium veitchii first hand. I therefore consider it enough to be guided by Schott’s interpretation and designate the Kuhl & Hasselt specimen as the neotype. 4. The above-cited illustration of A. watsoniana is of a sterile plant, though it shows the distinctive bullate blade and in-curved posterior costae. It does not actually accompany the protologue, having appeared in an issue of Gardeners’ Chronicle one month later. However, the caption includes direct reference to the protologue. Whether or not direct connection may be inferred between the illustration and the protologue, application of the name requires to be established more firmly. The above-selected epitype is preserved from a flowering cultivated plant of, evidently, the same clone introduced by Sander & Co, which Masters described. 5. No material was preserved of Alocasia thibautiana when it was first described, nor was it illustrated. The designated neotype consists of two sheets annotated by N.E. Brown ‘from the type plant’. Both consist of leaf only. A third sheet, dated 12 November 1881 consists of a dried inflorescence. 6. Linden’s description of Alocasia singaporensis is extremely scant, reading, from the German, ‘From Singapore. The large leaf is arrow-shaped, with large spreading basal lobes and of dark green colour’. Assuming the provenance Singapore alludes to origin from the wild, this description can only match Alocasia longiloba ‘denudata’. Material preserved by Brown at Kew under the name A. singaporensis is indeed of that entity. 7. Alocasia denudata var. elongata Engl. was differentiated from the typical variety by the narrower lobes of the leaf blade. The designated neotype well exemplifies this state. The illustration that accompanied the protologue is not good enough to serve as the type in the absence of the material it was apparently based on. There are no details of leaf venation and the lower part of the spadix is stylised. 306 Gard. Bull. Singapore 50(2) (1998) 8. Alocasia amabilis W. Bull was validly published in the above-cited retail list, and is neotypified with material preserved by Brown at Kew from a plant obtained from Bull under that name. KEY TO THE PEAK VARIANTS la. Leaf blade of adult plant not distinctly pendent, not peltate (S. Malay Penmeula. E. Sumateray. 33... 404 ivan slain a ee ae ‘denudata’ 1b.. Leaf blade.of adult plant pendent; pellate 07...) eco ao eee 2 2a. Leaf blade narrowly triangular in general outline (ca. 3 times as long as broad) or sometimes slightly hastate (throughout range of species) wis sah cab ahis S paints vk b nliot Huy Geka dace ied adie ened ae aaa a ‘longiloba’ 2b. Leaf blade broadly triangular to broadly oval in general outline (ca. 2 times.aslong as. broad) ;notiastate cc Bic cccucescea ss aden eee ae 3 3a. Posterior lobes of leaf united for at least half their length ............... + 3b. Posterior lobes of leaf united for less than half their length ............ 5 4a. Interprimary collective veins zigzagging at acute angles (Peninsular Malaysia; Sumateta, BOMmeeyh ue tigate ee fon cote tes ‘watsoniana’ 4b. Interprimary collective veins more or less undulate (Borneo) wb: Btcgte n'a diay aceip 5 b-slgha'g Xone lean Rey thats et en ara ata & een ee ae ae ‘korthalsii’ Sa. Interprimary collective veins zigzagging at ca. right angles; secondary venation paler than blade ground colour (Sumatera) ......... ‘putzeysii’ 5b. Interprimary collective veins weakly formed to indiscernible; secondary venation concolorous with leaf blade ..... Saishaln cant iy Ialeae steams ae 6 6a. Blade broadly triangular (Peninsular Malaysia, NW Borneo) ........ ‘lowii 6b. Blade broadly ovato-sagittate (northern Peninsular Malaysia, ?7NW BOrmco) gi gan sessile pelea tance eee eas ae een ‘thibautiana’ a. ‘denudata’ - Alocasia singaporensis Linden - Alocasia denudata Engl. - Alocasia denudata var. elongata Engl. Terrestrial herb to ca. 1m tall; eaves 1-3 together; petiole to 80 cm (often less), mostly rather densely obliquely mottled chocolate, the ground colour occasionally bright pink; blade green, sagittate, usually not pendent, with the primary venation not or barely of a different colour from that of the Alocasia in West Malesia and Sulawesi 307 lamina; posterior lobes subequalling the anterior; posterior costae naked in the sinus for up to 7 cm; interprimary collective veins absent to weakly formed and then only slightly undulating. Distribution: Southern Malay Peninsula and E . Sumatera. Habitat: In rain forest and regrowth understorey at low elevation. Note: This entity is qualitatively distinct from others on the basis of its non-peltate leaves, which are not as makedly pendulous as they are in the rest of the complex. However, even quite advanced sub-adult plants have peltate leaves, which are identical to A. longiloba ‘lowi1’. It is geographically fairly coherent, but intergrades in Sumatera, through very shallowly peltate forms, with ‘longiloba’ (e.g. Docters van Leeuwen-Reijnvaan 11790 (BO)) and ‘putzeysii’ (e.g. Meijer 6859 (L)) variants. Were this entity to be recognised as a separate species, A. singaporensis would have priority over A. denudata. However, A. denudata ha been a name in wide use, while A. singaporensis has never been taken up to any significant extent, and I would recommend that A. denudata be proposed for conservation. Selected other specimens seen: PENINSULAR MALAYSIA: Malacca, Pulau Nangka, Burkill 2641 (K); Selangor, Kuala Langat, nr Klang, Burkill 4102 (SING); Johore, Jason Bay, Corner s.n. (SING); cult. RBG Sydney Acc. no. 940260 ex Johor, G. Panti via Kg Lukit, Hay et al. 9192 (NSW); Johore, Kluang F.R., Holttum 9235 (K, SING); Johore, 20th Milestone, Kota-Tinggi - Jemalung Rd, Nicolson 1229 (US); Sembilan Islands, Pulau Rembia, Sinclair 76443 (KEP); Dindings, Lumut, Ridley s.n. (SING). SINGAPORE: Jurong Rd, Burkill 255 (SING); Mandai Rd., Burkill 11432 (SING); Singapore Botanic Garden, Croat 53241 (B, K); Cult. RBG Sydney Acc. no. 901379 ex Singapore Botanic Garden Rain Forest, Hay s.n. (NSW); Singapore Botanic Garden Jungle, Nicolson 1007, 1120, 1121 (all US); Bukit Timah, Upper Fern Valley Ravine, Nicolson 1106 (US); ‘Woodlands’, Ridley s.n. (SING); Chan Chu Kang, Ridley s.n. (SING), Changi, Ridley s.n. (SING); Tuas, Ridley s.n. (SING); Bukit Timah, Ridley s.n. (K, SING), Kranji, Ridley s.n. (SING); Kg. Pulau Damar Darat, Sinclair 6874 (E, US). SUMATERA: Siberut Island, Boden-Kloss 11438 (SING); Banka, P. Pinang, G. Mangkol, Biinnenmeijer 2124 (BO); Riau Archipelago, Belobang, Biinnenmeijer 7696 (BO, L); Cult. RBG Sydney Acc. no. 970448 ex Jambi Prov., 100km along rd Jambi-Palembang, Hay et al. 13007 (NSW); Mentawi Islands, Siberut Island, Iboet 26 (BO, L, SING); SE Bangka, Lobok Besar, Kostermans 266 (BO); Malacca Straits, Pulau Bukala, Sinclair 76419 (KEP). b. ‘longiloba’ - Alocasia longiloba Miq. s. str. - Alocasia cuspidata Engl. - ?Caladium veitchii Lindl. - Alocasia veitchii (Lindl.) Schott - Alocasia lowii var. veitchii (Lindl.) Engl. - Alocasia amabilis W. Bull. 308 Gard. Bull. Singapore 50(2) (1998) Generally terrestrial (occasionaly lithophytic) sometimes robust herb, to ca. lm (1.5m) tall (usually ca. 60 cm); leaves solitary to 3 together; petiole to 80 cm tall (usually ca. 40 cm), mottled dark green to chocolate; blade hasto-sagittate, rather narrowly triangular, dark to very dark green, usually with the major venation grey-green adaxially, posterior lobes 1/2-2/3(-3/4) the length of the anterior, peltate for (5S—)10-30% of their length, acute; secondary venation initially widespeading, then sooner or later deflected towards the margin; interprimary collective veins absent to weakly formed and zig-zag at widely obtuse angles. Distribution: Central Vietnam and Thailand to Peninsular Malaysia, Sumatera, Borneo, Java and Sulawesi. Habitat: In rain forest and regrowth understorey, in swampy areas and well drained slopes, occasionaly on rocks, at low to medium elevation; in Sulawesi from sea level to ca. 2000 m altitude. Note: This form is very widespread. In Borneo and Peninsular Malaysia it intergrades with A. longiloba ‘lowi’, which typically has broader leaf blades and is generally lithophytic. In Sumatera it intergrades with A. longiloba ‘putzeysil’ (e.g. Hay et al. 13080). All the collections from Sulawesi are sterile. Those plants have extremely shallowly peltate leaves. Selected other specimens seen: PENINSULAR MALAYSIA: Perak, Taiping Waterfall, Furtado s.n. (SING); Cult. RBG Sydney Acc. no. 940165 ex Perak, Thaiping, Bk Larut, Hay et al. 9069 (NSW); Selangor, Ulu Langat, Millard 1866 (SING). SUMATERA: Asahan, Silo Maradja, Bartlett 6441 (US); Aceh, G. Leuser Nature Resrve, Upper Mamas R.., ca. 15 km W of Kutacane, de Wilde & de Wilde-Duyfjes 19001 (L); Cult. RBG Sydney Acc. no. 970459 ex Jambi Prov., 120 km along rd Sungei Penuh - Bangko, Hay et al. 13018 (NSW); Cult. RBG Sydney Acc. no. 970509 ex West Sumatera, Padang, Gunung Gadut, Hay et al. 13080 (NSW). JAVA: West Java, Bogor, Boerlage s.n. (L); Bantam, Lebak Kidoel, G. Kancana, Koorders 40970b (L); Preanger, Tasik Malaja, Pendjalu, Koorders 44348b, (L); West Java, W of Djasinga, Djankapa forest reserve, Meijer 2947 (BO); Batavia, Wanajasa, Wisse 1237 (L). KALIMANTAN: Pulau Lampei, Korthals s.n. (L); East Borneo, Berau distr., Kostermans 21838 (L); West Kalimantan, Pontianak, S. Raja, Mondi 15 (L). SARAWAK: Matang F.R., 10 mi W of Kuching, Nicolson 1271 (US); Bako National Park, 20 mi NE of Kuching, Nicolson 1307 (US); Kuching, Ridley 12250 (SING); Binatang, Pulau Bruit, Sanusi bib Tahir 9219 (L). BRUNEI: Temburong Distr., Sg. Temburing at Kuala Belalong, Boyce 359, 384 (both K); Belait Distr., Sg. Liang Arboretum, Foreman & Blewett 1082 (K); Bangarmassing, Motley 1131 (L); Belait District, Rasau, van Niel 4247 (L). SABAH: Cult. RBG Sydney Acc. no. 950372 ex Sandakan, Kebun Cina, Hay et al. 10035 (NSW); cult. RBG Sydney Acc. no. 960481 ex Sepilok F.R., Hay et al. 12152 (NSW); Cult. RBG Sydney Acc. no. 960512 ex G. Rara F.R., ca. 2.5 km above main Maliau Falls, Hay et al. 12050 (NSW). SULAWESI: Masamba, Takala-Teboro, Eyma 1460 (BO); E Central Sulawesi, Morowali Prov., Grimes 1906 (K); Central Celebes, Mt Nokilalaki, Meijer 9859 (L); Makassar, Malinoboren, Rant 440 (BO); Enrakang Distr., Latmojong Mts, Bunteh Tjejeng, Sands 193 (K); | Alocasia in West Malesia and Sulawesi 309 c. ‘putzeysii’ - Alocasia putzeysti N.E. Br. Terrestrial (?always) herb to ca. 90 cm tall; /eaves 1-3 together; petiole to ca. 80 cm, brown-mottled or more or less concolorous brownish purple; blade narrowly ovato-sagittate, rather shallowly peltate (to ca 20% of the depth of the posterior lobes), nearly always purple-backed; secondary venation conspicuous, forming weakly zig-zag interprimary collective veins. Distribution: Sumatera. Habitat: On rainforest floor, usually on slopes at low to medium elevation. Note: This element is distinguished from ‘watsoniana’ by the straight posterior costae diverging at a wider angle, the more triangular leaf outline and the less deeply peltate posterior lobes. The interprimary collective vein is mostly less markedly zig-zag in course, though Hay et al. 13102 comes from a population in which some individuals have the venation pattern typical of ‘watsoniana’, though the leaf shape is of ‘putzeysu’. The secondary venation is typically paler in colour than the ground colour of the lamina on either the adaxial side or both sides. This element links ‘watsoniana’ with ‘longiloba’. Meijer 6859 resembles ‘denudata’ in its long posterior lobes distinctly elliptic on the inner sides, but matches ‘putzeysii’ in other respects. The type of Alocasia putzeysii is of a leaf only, but it is quite distinctive. The shape is narrowly ovato-sagittate with the posterior lobes 2/3 the length of the anterior and peltate for ca. 20% of their length. The interprimary collective veins form a weakly zig-zag pattern and the secondary venation is paler than the ground colour. In the protologue, A. putzeysii was attributed to Java, however, this form matches Sumateran material, not Javan. That the attribution to Java was in error appears to be confirmed by N.E. Brown’s notes on the type specimen, where he states the provenance as Sumatera, ‘Atchin’ (?= Aceh) Province. Specimens seen: SUMATERA: Cult. RBG Sydney Acc. No. 970528 ex W. Sumatera, Lembah Anai, Hay et al. 13102 (NSW); W. Sumatera, Taram, E of Pajakumbuh, Meijer 6859 (L); ‘West Coast’, Micholitz s.n. (K); d. ‘watsoniana’ - Alocasia watsoniana Mast Mainly lithophytic but also terrestrial moderately robust herb to ca. 1.25 m 310 Gard. Bull. Singapore 50(2) (1998) tall; /eaf usually solitary (—3 together); petiole purplish, not or faintly mottled; blade ovato-sagittate, shield-shaped, to ca. 60 (—-85) cm long, adaxially dark green with strikingly whitish major venation, abaxially purple, sometimes shallowly undulate on the margin, often bullate with long narrow wrinkles running between and more or less perpendicular to the primary veins and arranged more or less concentrically around the insertion of the petiole; proximal primary venation diverging at a very wide angle (to over 90°), distal primary veins diverging at ca. 45°; secondary venation rather dense, arising at a very wide angle and uniting into an interprimary collective veins very strongly zig zagging at acute angles; posterior lobes rounded acute, united for over half their length; posterior costae diverging at ca. 45—90° then somewhat incurved in the manner of the posterior rhachises of a pedate leaf. Distribution: Peninsular Malaysia, Sumatera, Borneo. Habitat: Terrestrial and on cliffs and on boulders in forest, sometimes on limestone, from sea level to ca. 700m. Notes: In Sumatera this element closely approaches, in its blade shape and secondary venation pattern, and occasionally intergrades with, ‘putzeysii’, but that typically has less deeply peltate leaves, straight posterior costae and blades that are not bullate. The two evidently differ to some extent ecologically, A. longiloba ‘putzeysi’ found usually on forest floor, while A. longiloba ‘watsoniana’ appears generally lithophytic, at least in Borneo and Peninsular Malaysia. Unfortunately in those Sumateran specimens most closely conforming to ‘watsoniana’, habitat details are not clear, though Hay et al. 13036 is terrestrial. Within the Bornean and Peninsular Malaysian part of the range, it appears sharply distinct morphologically from other elements of the A. longiloba complex. If ecological differentiation between the Sumateran and these other repesentatives of ‘watsoniana’ could be demonstrated more clearly, the Bornean and Peninsular Malaysian element should perhaps be regarded as a distinct (and in that case, new) species, and Sumateran ‘putzeysil’ and ‘watsoniana’ might be more usefully recognised as a single, though still fuzzily circumscribed, local variant of the A. longiloba complex. Selected other specimens seen: PENINSULAR MALAYSIA: Perak, Kuala Dipang, Curtis s.n. (SING); Perak, Kampar, G. Tempurong, Ng FRI 5834 (FRIM, L). SUMATERA: Cult. RBG Sydney, Acc. No. 970474 ex Jambi, Kerinci Seblat National Park, above Lempur Vill., Hay et al. 13036 (NSW); Sibolangit, Lérzing 4804 (BO). KALIMANTAN: [without locailty] Amdjah 165 (BO); Kalimantan Timur, foot of G. Batukenye, along Sg. Belayan, Alocasia in West Malesia and Sulawesi Sit NW of Tabang, Murata et al. 1519 (BO); Bidang Menabei, Winkler 1064 (E, L). SARAWAK: Cult. RBG Sydney, Acc. No. 920745 ex Long Jowe, Dearden s.n. (NSW); SABAH: Cult. RBG Sydney, Acc. No. 960609 ex Kinabatangan, Kalabakan Virgin Jungle Reserve, Hay et al. 12012 (NSW). e. ‘korthalsii’ - Alocasia korthalsti Schott Moderately robust terrestrial herbs, rhizome rather slender, to ca. 2.5 cm diam; /eaf usually solitary (-4 together); blade ovatosagittate, shield-shaped, to ca. 40 cm long x 17 cm wide, widest ca. 3 cm anterior to the petiole insertion, plain mid-green on both sides to deep purple abaxially and then deep green adaxially, not usually with contrastingly pale major venation; anterior costa with 3-4 primary lateral veins, the proximal ones diverging at ca. 85°, the distal at ca. 45°; secondary venation arising at a wide angle, thence deflected towards the margin and forming rather weakly undulating interprimary collective veins; posterior costae diverging at ca. 45° or less; posterior lobes peltate for ca. 60% of their length, the free part rounded to rounded-acute; inflorescences mostly at the smaller end of the size range for this complex, with the spathe limb rather markedly cucullate; appendix ivory to yellow. Distribution: Borneo. Habitat: Terrestrial on rainforest floor mainly at low elevation (Purseglove & Shah P4749 at ca. 1200 m). Notes: This form is distinguishable from the other shield-shaped leaved member of this complex, ‘watsoniana’, by the smaller rhizome and inflorescence, more cucullate spathe limb, undulating interprimary collective veins and (in Borneo) terestrial habit. A. Jongiloba ‘korthalsii’ intergrades with A. longiloba ‘lowiv’ in Sarawak. In Sabah it is found, e.g. at Sepilok, mixed and not intergrading with A. /ongiloba ‘longiloba’. The population I have seen near Bintulu, Sarawak, is mostly of unifoliar individuals with dark green, purple-backed leaves, while the above-mentioned population at Sepilok is of multifoliar individuals with plain green leaves. Other specimens seen: KALIMANTAN: Kalimantan Timur, 10-20 km N of Sebulu, Murata et al. 703 (BO). SARAWAK: Cult. RBG Sydney Acc. no. 940462 ex 2.3 km from Kemena R. bridge towards Sibu, Hay et al. 9308 (NSW); G. Pueh, Purseglove & Shah P4749 (SING); Tambusan, Ridley s.n. (SING). SABAH: Cult. RBG Sydney Acc. no. 960519 ex Sepilok F.R., Hay et al. 12153 (NSW); Danum, Lambert TBO (E). 342 Gard. Bull. Singapore 50(2) (1998) f. ‘thibautiana’ - Alocasia thibautiana Mast. - Alocasia curtisii N.E. Br. Robust, often lithophytic, herb usually with the rhizome somewhat swollen, subcormescent; /eaves 1-several together; blades broadly ovato-sagittate, to ca. 50 cm long, peltate for ca. 25% of the length of the posterior lobes, plain mid green throughout to dark green, red-backed and with whitish primary adaxial venation; secondary venation not or hardly forming interprimary collective veins. Distribution: Peninsular Malaysia, ?Sarawak. Habitat: Terrestrial or on limestone rocks at low elevation. Note: Intergrades with ‘low’. Leaves of juveniles of this form strongly resemble ‘korthalsi’. I recognise this entity around its extreme form, which has not only rather shallowly peltate, ovato-sagittate leaves with no or weak interprimary collective veins, but also a distinctive swollen and abbreviated corm-like stem. The type of A. thibautiana 1s alleged to be from Borneo. Some wild-collected specimens from Borneo are intermediate between this entity and ‘lowii’, though I have seen no authentically Bornean material, which corresponds directly with ‘thibautiana’. Other specimens seen: PENINSULAR MALAYSIA: Penang, Waterfall, Curtis s.n. (SING); Cult. RBG Sydney Acc. no. 940064 ex Perlis, Kaki Bukit, Gua Kelam, Hay et al. 9019 (NSW); Pahang, Pulau Tioman, Pulau Tulai, Henderson 18506 (SING); g. ‘lowii’ - Alocasia lowii Hook. f. - Alocasia lowii var. picta Hook.f. - Alocasia eminens N.E. Br. Robust, often lithophytic, herb ca. 70cm —1.5 m tall; eaves (1—-)2+4 together; petiole usually obliquely mottled dark green or chocolate, sometimes unmottled; blade sagittate, rather broadly triangular in outline, usually dark green with contrastingly paler major venation adaxially, sometimes purple-backed, sometimes concolorous green throughout; anterior lobe sometimes slightly ovate, ca. 30-70 (—90) cm long, generally widest about level with or slightly distal to the insertion of the petiole (occasionally widest almost at the tips of the posterior lobes); anterior costa with ca. 4 primary lateral veins on each side, the proximal ones diverging at ca. (100—)80°, the distal ones at ca. 45°; secondary venation arising from the primary at a wide angle then soon deflected towards the margin and forming Alocasia in West Malesia and Sulawesi 413 ill-defined interprimary collective veins or these absent; posterior costae straight, diverging at ca. 45—90°; posterior lobes shallowly peltate - for 10-15% of their length, acute. Distribution: Peninsular Malaysia, NW Borneo. Habitat: In forest, often on rocks including limestone at low to medium elevation, extending into quite markedly seasonal areas. Notes: 1. This element represents little more than a robust aspect of A. longiloba ‘longiloba’, with which it intergrades, indeed ascription of a considerable number of collections to one or other of these is somewhat arbitrary. The anterior lobe is typicaly relatively wider and the posterior lobes relatively longer than in typical ‘longiloba’, and in this respect it approaches ‘denudata’. Alocasia veitchii (Lindley) Schott, whose basionym is earlier than A. /owii, falls in between. If it was classed in the same group as specimens conforming to ‘lowii’, it would of course formally have priority, however, it seems to me marginally closer to ‘longiloba’ and since the nomenclatural framework used here is anyway informal, rules of priority need not apply in the event that another worker interpreted A. veitchii as falling within the ‘lowi’ variant. The epithet ‘lowi’ is more widely used by both botanical and horticultural collectors. In Borneo A. longiloba ‘loww also intergrades with ‘korthalsi’ in a few instances, for example Chew 709 and Jacobs 5476 (see below). Selected other specimens seen: PENINSULAR MALAYSIA: Kedah, Langkawi Is, P. Bumbon Besar, van Balgooy 2293 (L); Perak, Kuala Kangsar, logging road up G. Bubu from Manong, Boyce 706 (KEP); Malacca, Pulau Nangka, Burkill 2641 (SING); Kelantan, Kota Bahru, Gimlette 5962 (SING); Cult. RBG Kew ex Negeri Sembilan, Pasoh F.R., Hay 2005 (K); Cult. RBG Kew ex Kedah, Langkawi Is., Pulau Dayang Bunting, Hay 2032 (K); Cult. RBG Sydney, Acc. no. 940047 ex Perlis, Kangar, Bukit Lagi, Hay et al. 9001 (NSW); Cult. RBG Sydney Acc. no. 940102 ex Selangor, Batu Caves, Hay et al. 9058 (NSW); Cult. RBG Sydney Acc. no. 940138 ex Negeri Sembilan, Bukit Tangga, nr Jelabu, Hay et al. 9142 (NSW); Cult. RBG Sydney Acc. no. 940355 ex Pahang, Bukit Fraser, Hay et al. 9288 (NSW); Selangor, Sg. Tinggi, Md Nur 34111 (A); Pahang, Ulu S. Krau, NE G. Benom, Whitmore FRI 3135 (K). BRUNEI: Belait Melilas, Kuala Ingei, Melilas side of Belait R., Thomas 216 (K); Temburong Distr., Bukit Belalong, Wong 1417 (K). SARAWAK: Kuching district, Tiang Bakap, Mt Maja, Chew 709 (L); Ist division, 30 km SW of Kuching, Sebuaran Bau, Jacobs 5476 (L). 23. Alocasia celebica Engl. Alocasia celebica Engl. in Koord., Meded. s’Lands Plantentuin 19 (1898) 299; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 106. - Type: Indonesia, Sulawesi, Minahassa Prov., Ratatotok, 25 Mar 1895, S. H. Koorders 161628 (B, holo). 314 Gard. Bull. Singapore 50(2) (1998) Herb ca. ?1.5 m tall; rhizome ca. 4 cm diam., clothed in old cataphyll bases; leaves two together subtended by papery fibrous marcescent cataphylls; petiole ca 35 cm long, sheathing in the lower 1/4, densely and minutely pubescent, mottled with an oblique zig-zag pattern; blade somewhat ovato- sagittate, 36 cm long, rather thickly coriaceous; anterior lobe widest ca. 3 cm above the petiole insertion, the apex acute; anterior costa with 6 primary lateral veins on each side, diverging at 80-60° and running almost straight to the margin; axillary glands inconspicuous; secondary venation obscure; posterior lobes about half the length of the anterior, acute, distally slightly out-turned, the inner sides oblanceolate; posterior costae diverging at ca. 60°, not naked in the sinus (leaf blade very slightly peltate? - sinus obscured on holotype); inflorescence unknown. Distribution: Endemic to Sulawesi, known only from the type collection. Habitat: Unknown; the type was collected at 200 m altitude. Notes: 1. The affinities of this species are not clear from the type, which is sterile. However, the papery-fibrous cataphylls, mottled petioles and (almost) peltate leaf suggest the Longiloba Group. The obscurity of the secondary venation is due to the thickness of the leaf blade, which is not a feature of any other member of this group. Koorders and Engler & Krause overlooked the pubescence on the petiole, which also occurs, among Sulawesi species, in A. suhirmaniana (q.v.), from which A. celebica is amply distinct. 2. The holotype has three Koorders numbers on it: the field number 2587 attached to the specimen; the Herb. Koordersianum number /6/62f on a label dated 25 Mar 1895 giving the provenance Ratatotok (as in the protologue), and a Museum Botanicum Berolinense label dated 3 Jul 1895 with the number Koorders 19750 and the provenance Ratahan. 24. Alocasia suhirmaniana Yuzammi & A. Hay Alocasia suhirmaniana Yuzammi & A. Hay, Telopea 7 (1998) 303, fig. 1. - Type: Cult. Kebun Raya Bogor ex SE Sulawesi, Kabupaten Kolaka, 23 Jun 1997, Yuzammi s.n. (BO, holo; NSW, photo). Terrestrial herb to ca. 65 cm tall; rhizome 13-15 cm long, ca. 3 cm diam.; leaves 1-3 together; petiole to ca. 60 cm long, sheathing in the lower !/ 5—!/4, yellowish green, densely longitudinally and obliquely mottled purple- Alocasia in West Malesia and Sulawesi ei i, brown, minutely and densely puberulous, subtended by papery-membranous cataphylls; blade broadly ovato-sagittate, to ca. 55 cm long, peltate, pendent, thinly leathery, with the margin somewhat undulate, glossy dark green adaxially with the major venation pale grey-green, dark purple abaxially; anterior lobe widest about 1/4 of the way from the base, the tip broadly acute to obtuse, shortly apiculate; anterior costa with up to 8 primary lateral veins on each side, diverging at 80-45°, with conspicuous purple glands in their axils abaxially; subsidiary veins frequent in the outer part of the blade; secondary venation otherwise inconspicuous, forming undulating interprimary collective veins; posterior lobes acute, about !/2-2/3 the length of the anterior, united for !/2-2/3 of their length; posterior costae more or less straight, diverging at ca. 35—45°; inflorescence pairs solitary (?always), subtended by papery membranous cataphylls to ca. 11 cm long; peduncle to 24 cm long, minutely puberulent in the upper part, purple-brown; spathe ca. 12 cm long, deep purple, slender, glabrous, abruptly constricted at ca. 2 cm from the base; lower spathe subcylindric; limb narrowly lanceolate; spadix somewhat shorter than the spathe, ca. 10 cm long, slender, very shortly stipitate for 4 mm, stipe ivory; female zone ca. 1.2 cm long; ovaries greenish yellow; stigma bluntly 2-4-lobed, subsessile, yellow; sterile interstice ca. 0.5 cm long, attenuate, level with spathe constriction; lowermost synandrodia strongly lobed, the rest rhombo-hexagonal, ca. 1.5 mm diam.; male zone ca. 2 cm long, 1 cm diam., cylindric; synandria rhombo-hexagonal, the tops impressed, ca. 2 mm diam., yellowish ivory; thecae opening by apical pores somewhat laterally displaced by overgrowth of the synconnective; appendix ca. 6 cm long, 8 mm diam. at base, slightly constricted at junction with male zone, the rest cylindric, then tapering in the upper !/3, yellowish, somewhat rugose in the lower half; infructescence unknown. Distribution: Endemic to SE Sulawesi. Habitat: In damp shady spots in lowland rain forest on slopes, sometimes over limestone. Note: This species is distinguished from other members of the Longiloba Group by its puberulent petioles, blackish-purple spathe and somewhat marginally expanded synconnectives. Other specimen seen: SULAWESI: SE Sulawesi, Tolala, Kjellberg 2428 (BO). 316 Gard. Bull. Singapore 50(2) (1998) Cuprea Group Species 25—30 Leaves more or less completely peltate, interspersed with cataphylls; spadix generally distinctly shorter than the spathe; male zone often mostly or completely within the lower spathe. Note: This group includes six species, four Bornean, one from each of the Malay Peninsula and Sumatera. The vegetative characteristics that define it do not seem to be matched by distinctive reproductive features and it is not clear that this group is natural. It may be linked to the Scabriuscula group via A. reversa and the A. princeps complex. 25. Alocasia perakensis Hemsl. Alocasia perakensis Hemsl., J. Bot. 25 (1887) 205. - Type: Malaysia, Perak, Birch’s Hill, Wray 29 (K, holo; iso K, SING). [Alocasia beccarii sensu auct. non Engl.: Hook.f., Fl. Brit. Ind. 6 (1893) 527; Ridl., Mat. Fl. Mal. Pen. 3 (1907) 17; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 95, pro parte quoad A. perakensis in synon. et specim. cit. Mal. Pen.; Ridl., Mat. Fl. Mal. Pen. 3 (1907) 16 & Ridl., Fl. Mal. Pen. 5 (1925) 97 pro parte quoad specim cit.; Henderson, Mal. Wildfl. Monoc. (1954) 225, excl. fig 134 (i.e. A. beccarii s.s.)]. Herb to ca. 75 cm tall (often smaller); stem creeping to decumbent, somewhat elongate - the internodes as long as or longer than wide, ca. 2.5 cm diam.; /eaves several along the stem, (?)irregularly interspersed with lanceolate cataphylls to 8 cm long and drying red-brown; petiole to ca. 40 cm long, sheathing in the lower !/4, grey-green to purple-brown; blades dark green to grey-green, ovate to elliptic, peltate, coriaceous to thickly coriaceous and subsucculent, 14 x 6 — 34 x 13 cm; anterior lobe widest ca. 2-4 cm distal to insertion of petiole, the tip broadly acute, acuminate for ca. 1.5 cm, the margin mostly entire, occasionally somewhat sinuous in the lower part; anterior costa with 2—3(-4) primary lateral veins on each side, diverging at ca. 45—-60°, running to a submarginal vein ca. 1 mm from the margin; secondary venation not forming interprimary collective veins, mostly inconspicuous, but, like primary venation, adaxially impressed in dry state in thickly coriaceous leaves; posterior lobes completely united except for a shallow retuse notch, rarely with an acute notch to ca. | cm deep, together cuneate to slightly attenuate, !/3—!/2 the length of the anterior lobe; posterior costae subparallel; inflorescence solitary to paired; peduncle about half to Alocasia in West Malesia and Sulawesi 5L7 subequalling the length of the petioles; spathe greenish yellow to white, ca. 6 cm long; lower spathe ovoid, ca. 3 cm long; limb narrowly ovate, at first erect, then reflexed; spadix shorter than spathe, ca. 5 cm long, stipitate for 4 mm; female zone ca. 7 mm long; pistils few - ca. 15, rather large - ca. 4 mm long; ovary globose, 2.5 mm diam.; style 1.5 mm long; stigma prominently 2—3-lobed; sterile interstice ca. 2.5 mm long, a single whorl of synandrodia; male zone 1.5 cm long, entirely within and filling the upper half of the lower spathe chamber, conic - ca. 8 mm diam. at base narrowing to 5 mm at apex corresponding with spathe constriction; synandria relatively large - 4 mm diam., more or less hexagonal, 3—5-merous; thecae opening by apical pores not overtopped by synconnective; appendix narrowly cylindric, ca. 2.5 cm long, 4 mm diam., deeply grooved, white to yellowish; fruiting peduncle subequalling the petioles; fruiting spathe ovoid, ca. 4 cm long, the spathe dehiscing longitudinally; berries bright red. Distribution: Endemic to Peninsular Malaysia. Habitat: In montane forests, in leaf litter and on rocks, mostly at 1100-1525 m altitude. Ridley (ll. cc.) noted it as low as 650 m (2000 feet), probably based on his collection from Kuala Teku. Notes: Although Alocasia perakensis has not been accepted as an entity distinct from A. beccarii by any author since its first description, they are readily distinguishable, though evidently closely related, allopatric species. Alocasia perakensis is on the whole much more robust, the leaves are generally more leathery - sometimes almost succulent, and the connate posterior lobes are cuneate rather than attenuate, the stem is more elongate, the bracts between the leaves are less frequent, the inflorescence, though structurally very similar to that of A. beccarii, is about twice the size, and the spathe is greenish yellow to whitish. With the exception of two high altitude collections from Mt Kinabalu (Sabah), doubtfully attributed here to A. beccarii (q.v.), it has a higher altitudinal range than that species. Although quite a number of collections have been made, this species is poorly known in flower and the description of the inflorescence is prepared from a single spirit collection (Hay et al. 9280). Other specimens seen: PENINSULAR MALAYSIA: Pahang, Cameron Highlands, Batten Pool s.n. (SING); Perak, Larut, trail from Bk. Larut to G. Hijau, Boyce 681 (K, KEP); Perak, G. Hiyau, Burkill & Haniff 12769 (SING); Negeri Sembilan, Ladang Gadis, Carrick 692 (SING); Selangor, along old abandoned rd to Genting Highlands, Croat 53321 (K); Perak, Genting Highlands, Croat 53338 (K); Perak, G. Hijau, Mohd. Haniff & Mohd. Nur 2350 (K, SING); Cult. RBG Sydney Acc. No. 940347 ex Pahang, Bukit Fraser, Hay et al. 9280 (NSW); Pahang, No. 5 Camp, Cameron Highlands, Henderson FMSM 11666 (BO); Perak, Larut Hill, Thaiping, Long 6 (K); Selangor, top of Fraser’s Hill, along path from 318 Gard. Bull. Singapore 50(2) (1998) Red Cross to Wray’s Cottage, Nicolson 1175 (US), 1178 (US); Pahang, Cameron Highlands, along S path to G. Beremban, Nicolson 1194, 1201 (both SING, US); Pahang, Fraser’s Hill, Mohd. Nur 10548 (SING); Pahang, Boh Plantation, Cameron Highlands, Mohd. Nur s.n. (SING); Pahang, Fraser’s Hill, below Methodist Mission, Purseglove P.4283 (GH, K, L, SING); Pahang, Kuala Teku, Ridley s.n. (K); Perak, Scortechini s.n. (K, SING); Perak, Maxwell’s Hill, path to G. Hijau, Mohd. Shah & Sidek 1071 (K, SING); Kelantan, G. Stong, Symington 37727 (KEP). 26. Alocasia beccarii Engl. Alocasia beccarii Engl., Bull. Soc. Tosc. di Ort. 4 (1879) 300 & in Becc, Malesia 1 (1882) 293, t. 16, figs 1-4; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179, pro parte excl. specim. cit. Ridley, Matang (i.e. Alocasia peltata M. Hotta); Engl. & K. Krause, Pflanzenr. 71 (I1V.23E) (1920) 95, fig. 19, A-D, pro parte excl. A. perakensis in synon. et. specim. cit. Ridley, Matang); Mayo, Bogner & Boyce, Genera of Araceae (1997) pl. 104(i), M (non pl. 104(i1), i.e. Alocasia kerinciensis A. Hay). - Type: Borneo, Sarawak, Matang, O. Beccari PB 1674 (FI, holo). Small herb 12-28 cm tall; stem slender, 5-10 mm diam., condensed with the internodes usually somewhat wider than long; /eaves several together, irregularly but frequently interspersed with lanceolate cataphylls to 5 cm long and drying red-brown; petioles green, sometimes flecked pale mauve, 6-16 cm long, sheathing in the lower !/7 or less; blades narrowly elliptic to ovate to narrowly obovate, mid-green above, paler below, coriaceous, 9 x 2.7-18 x 6 cm; anterior lobe widest usually ca. !/4 of the way distal to petiole insertion, occasionally level with petiole insertion, occasionally 1/2 way distal to petiole insertion; margin occasionally somewhat sinuate; anterior costa with 2-3 primary lateral veins on each side, diverging at ca. 45-60° and running to a submarginal vein 0.5—1 mm from the margin; axillary glands inconspicuous; secondary venation not forming interprimary collective veins; posterior lobes almost completely connate save for a shallow retuse notch, (1/4)!/3—2/s the length of the anterior lobe, together attenuate; posterior costae subparallel; inflorescence solitary to paired; peduncle subequalling the petioles; spathe whitish, ca. 4 cm long, constricted ca. 1.5-2 cm from the base; lower spathe narrowly ovoid, distally somewhat curved adaxially; limb narrowly oblong-lanceolate; spadix shorter than the spathe, very shortly stipitate; female zone 4 mm long, a few loosely packed large pistils, or reduced to a single whorl; pistils ca. 3 mm long; ovary globose-ovoid, 2.5 mm diam.; style ca. 0.5-1 mm long, slender; stigma prominently ?2-lobed; sterile interstice a single whorl of synandrodia ca. 1.5 mm diam., or reduced to a single synandrode and the rest naked; male zone conic, 5-8 mm long, entirely within and filling the upper spathe chamber; synandria large, ca. 3 mm diam., 3—4-merous, thecae not Alocasia in West Malesia and Sulawesi 319 overtopped by synconnective; appendix pale apricot, narrowly cylindric, 1.3-2 cm long, ca. 3 mm diam.; fruiting peduncle hardly longer than flowering peduncle; fruiting spathe ovoid, ca. 2 cm long; ripe fruit orange to orange- red. Distribution: Endemic to N.W. Borneo. Habitat: In forest on slopes at low elevation - to ca. 850 m, possibly to 1500 m on G. Kinabalu (but see note below), often among or on boulders, often over sandstone. Notes: 1. This name has been used, in the literature and/or on herbarium sheets, for five West Malesian species of rather small plants sharing various manifestations of a distinctive more or less elliptic entirely peltate leaf blade - A. beccarii s.s., A. kerinciensis, A. minuscula, A. peltata and A. perakensis. Of these, A. peltata and A. kerinciensis have very conspicuous intramarginal veins and are montane species, A. minuscula has distinctive striate venation, very large synandrodia and is restricted to lowland peat swamp-forest, and A. perakensis is much more robust than A. beccarii and is a montane element restricted to Peninsular Malaysia. A. beccarii itself is distinguished by the combination of absence of intramarginal leaf vein, more or less condensed stem, secondary venation arising from the costae and primary veins, small size compared to A. perakensis and occurrence at low elevation in non-swampy sites. Further discussion of its distinction from A. perakensis can be found under that species. 2. The two collections from G. Kinabalu cited below differ from A. beccarii in the strict sense in having relatively broader posterior lobes with the tips less markedly acute, less completely joined and slightly out-turned at the tips. The leaf texture appears to be more membranous. The venation is nevertheless typical for A. beccarii. The altitude from which they were collected (4000-5000 ft) is significantly higher than collections of A. beccarii s.s., and it is possible that they represent another species in this group. Neither collection is in flower. Brooke 8680, from Keranji, Sarawak, is anomalous in having oblanceolate leaves, with very reduced posterior lobes. Other specimens seen: SARAWAK: Kuching, Brooke 8318 (L); Keranji, Brooke 8680 (L). BRUNEI: N. Temburong, Bukit Biang, Ashton A172 (K); Belait Distr., Ulu Ingei, Bukit Batu Patam, Boyce et al. 274 (K); Temburong Distr., Bangar, Bukit Patoi, Boyce et al. 350 (K); Temburong, Batu Apoi, Bukit Gelagas, Simpson & Marsh 2271 (K). SABAH: Kinabalu, Penibukan, Clemens & Clemens 31548 & 50499 (both SING); Sipitang Distr., W slope of G. Lumaku, Wood 798 (K). 320 Gard. Bull. Singapore 50(2) (1998) 27. Alocasia minuscula A. Hay, sp. nov. Ab A. beccarii lamina folii tenuiora, venis striatis, venis primariis duplo numerosis, sylvam palustrem incolenti differt. TYPUS: Borneo, Sarawak, Betong Distr., Saribas Forest Reserve, 14 Aug 1957, J.A.R. Anderson 8364 (L, holo; BO, K, iso); Diminutive herb 10-20 cm tall; stem suberect, ca. 1 cm diam, condensed, rooting along its length and clothed in old leaf bases and marcescent cataphylls; leaves several to 9 together, interspersed with papery- membranous cataphylls to ca. 5.5 cm long (these occasionally bearing reduced petiole and blade); petiole 5-10 cm long, sheathing in the lower ca. '/7; blade narrowly ovate to oblanceolate, 8 x 2-13 x 3 cm, peltate, coriaceous, pale abaxially; anterior lobe 7-10.5 cm long, the tip acuminate for ca. 1 cm; anterior costa with 8-10 primary lateral veins on each side, diverging at 60—45° then somewhat up-curved and joining a marginal vein; primary lateral veins much darker than blade abaxially in dry specimens and the majority not visibly reaching the midrib; secondary venation obscure on both sides of the blade, striate, arising from the midrib; posterior lobes almost completely united save for a ca. 2 mm incision at the extreme base of the leaf; combined posterior lobes attenuate, 1-2 cm long; inflorescence solitary; peduncle about the same length as the petioles at anthesis, later extending somewhat; spathe 3.5-4 cm long; lower spathe narrowly ovoid, 1.5—2 cm long, separated from limb by a weak constriction; limb ca. 2 cm long, lanceolate, colours unknown; spadix shorter than the spathe, ca. 2 cm long, stipitate for ca. 3 mm, the fertile zones entirely within the lower spathe; female zone 3 mm long; pistils few, ca. 10, bottle-shaped, more or less acroscopic, 1.5 mm long, style ca. 0.5 mm long; stigma small, weakly 23-lobed; sterile interstice ca. 2 mm long, the thickest part of the spadix, ca. 2.5 mm diam.; synandrodia inflated, more or less rhomboid, ca. 2 mm diam.; male zone 5 mm long, subcylindric, 2.2 mm diam.; synandria few, ca. 12, irregular, ca. 1.5 mm diam., more or less 4-merous, mainly composed of loosely adherent thecae, the synconnective not well developed; appendix ca. 6 mm long, 1 mm diam., cylindric; fruiting spathe ca. 1.5 cm diam.; berries red-orange. Distribution: Endemic to Sarawak. Habitat: In lowland peat swamp forest. Notes: The specific epithet derives from the fact that this is the smallest presently known species in the genus. Alocasia minuscula can be readily distinguished from A. beccarii and A. peltata, which it closely resembles, in Alocasia in West Malesia and Sulawesi aA its narrow peltate leaf shape and reduced posterior lobes, and in its diminutive stature, by the distinctive pattern of leaf venation. The primary veins are much more numerous, and characteristically some appear, in the dried state, not to reach the midrib. The secondary venation is obscure on both sides of the blade, but it appears on the abaxial side that the secondary venation arises hardly or not at all from the primary veins, nearly all the secondary veins running directly into the midrib - a condition more usually associated with striate-veined genera such as Schismatoglottis. Moreover, Alocasia minuscula appears restricted to swamp forest, while A. beccarii and A. peltata are hill and montane forest species. The description is based entirely on dried material. Other specimens seen: SARAWAK: Tuso Peninsula, Anderson 2129 (SING); Sibu, Naman F.R., Anderson 9299 (K, L); Simanggang, Brooke 10764 (L). 28. Alocasia peltata M. Hotta Alocasia peltata M. Hotta, Acta Phytotax. Geobot. 22 (1967) 156, fig. 5, A- E. - Type: Borneo, Sarawak, Bintulu, eastern ridge of Bukit Kana, 20 Nov 1963, M. Hirano & M. Hotta 1464 (KYO, holo, n.v.) Alocasia peltata var. muluensis M. Hotta, op. cit.: 158, fig. 5, F. - Type: Borneo, Sarawak, Mardi, Gunung Mulu, 16 Mar 1964, M. Hotta 14513 (KYO, holo, n.v.). [Alocasia beccarii sensu auct. non Engl.: Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179 & 49 (1907) 48; Engl. & K. Krause,,Pflanzenr. 71 (IV.23E) (1920) 95, pro parte quoad specim. cit. Ridley s.n., Sarawak, Matang.] Small herb to ca. 30 cm tall; stem more or less elongate, slender, sprawling, with internodes to 2 cm long; /eaves several along the stem, regularly alternating with papery membranous lanceolate cataphylls to 4 cm long; petiole to 16 cm long, sheathing in the lower !/10; blade narrowly elliptic to oblong ovate, 12 x 3 — 28 x 10 cm, peltate, somewhat to thickly leathery, glossy green or suffused purple adaxially, paler abaxially, drying with the venation somewhat to markedly impressed adaxially; anterior lobe 9-12 cm long, widest more or less level with petiole insertion, the tip acuminate for 1.5 cm; anterior costa with two primary lateral veins on each side (subopposite) diverging at ca. 60° and running straight or somewhat upcurved into a conspicuous intramarginal vein (2—)3-6 mm from the margin; secondary venation not forming interprimary collective veins, inconspicuous to invisible in thickly leathery forms; posterior lobes completely united or with a slight retuse notch, 3-6 cm long, together 322 Gard. Bull. Singapore 50(2) (1998) cuneate, ultimately truncate; inflorescence solitary; peduncle about half as long as to equalling the petiole; spathe ca. 5 cm long, green; lower spathe narrowly ovoid, ca. 2 cm long; limb lanceolate, ca. 3 cm long, separated from the lower spathe by a weak constriction; spadix somewhat shorter than the spathe, to 3 cm long, stipitate for 2 mm; female zone 4 mm long; pistils few - ca. 12, 2 mm long, flask-shaped, more or less acroscopic; style distinct, almost 1 mm long; stigma weakly 2-—3-lobed; sterile interstice a single inconspicuous whorl of synandrodia, not attenuate; male zone fully within the lower spathe, ca. 9 mm long, ca. 3 mm diam. at base, tapering to ca. | mm diam. at junction with appendix and corresponding with spathe constriction; synandria 3-merous, with anthers only dorsally, not laterally connate; thecae opening by apical pores not concealed by synconnective; appendix white, 1.5 cm long (much reduced in Burtt & Woods 2121), narrowly spindle-shaped, ca. 2 mm wide at widest; fruiting spathe obovoid, with the peduncle elongating; fruits red-orange. Distribution: Borneo, scattered localities in Sarawak, Brunei and central Kalimantan. Habitat: In mossy forest floor on ridges at ca. 1200 m altitude. Notes: 1. This species rather closely resembles Sumateran A. kerinciensis (q.v.), Sharing the pronounced intramarginal vein and regularly alternating foliage leaves and cataphylls, and the (usually) elongate stem with internodes longer than wide. It differs in the more slender leaves and spathe, the more elongate appendix, the male zone entirely within the lower spathe, the less robust synandria and longer pistils. 2. Hotta (loc. cit.) distinguished the variety muluensis on the basis of slightly smaller leaf size and longer peduncle. The material he described was in various stages post anthesis, and it appears that the peduncle continues to elongate as the fruits ripen. I am doubtful that the lower and upper leaf length extremes of 21 cm and 19 cm that he cites respectively for the typical and segregate varieties can be viewed as sufficiently significant to warrant their recognition. Other specimens seen: SARAWAK: Bakelalan, Brooke 10559 (US); 4th Div., G. Mulu, Burtt & Woods 212] (E); Matang, Ridley s.n. (SING). BRUNEI: Temburong, valley N of Pagon Ridge, Wong & Weber Booth 1903 (K). KALIMANTAN: Central Kalimantan, Bukit Raya, SE side, ca. 10 km NNW of Tumbang Tosah, Mogea 3856 (BO, K, KEP). 29. Alocasia kerinciensis A. Hay, sp. nov. Ab A. perakensis Hemsl. caudice producto tenuiore, -foliis et cataphyllis Alocasia in West Malesia and Sulawesi 525 alternantibus, lamina minus incrassata, venis intramarginalibus valde conspicuis, inflorescentia parviore, inflorescentia mascula exserta differt. TYPUS: Indonesia, Sumatera, Gunung Kerinci, 16 Apr 1920, Biinnenmeijer 9511 (L, holo; BO iso). [Alocasia beccarii sensu auct. non Engl.: Mayo, Bogner & Boyce, Genera of Araceae (1997) fig. 104(ii).] Small terrestrial herb; rhizome elongate, slender, stiff, decumbent-creeping, epigeal, ca. 1 cm diam., to ca. 40 cm long, with internodes to 5 cm long; leaves several, alternating with cataphylls, with leaf-cataphyll internodes subequalling cataphyll-leaf internodes; cataphylls membranous, narrowly oblong-lanceolate, to 8 cm long, drying red-brown; petiole to ca. 25 cm long, sheathing in the lower '/sth or less, wing of sheath basally broad, membranous, like the cataphylls in colour and texture; blades stiffly membranous, dull mid-green, broadly to narrowly ovate, peltate with the posterior lobes almost completely joined, ca. 13 x 6 -16 x 9 cm, widest ca. 1 cm distal to petiole insertion, the tip broadly acute to obtuse and shortly acuminate, base rounded with a retuse notch; anterior costa with 2-3 primary lateral veins on each side diverging at up to 100° (proximal) to 45° (distal) and running into a conspicuous intramarginal vein ca. 3-5 mm from the margin; secondary venation forming ill-defined interprimary collective veins; venation more or less flush with the lamina abaxially and adaxially; inflorescence solitary, rarely paired; peduncle subequalling the petioles; spathe 5-6.5 cm long; lower spathe ovoid, ca. 1.5 cm long, separated from limb by a pronounced constriction; limb broadly lanceolate; spadix shorter than the spathe, 3—4.5 cm long, shortly stipitate; female zone ca. 7 mm long; ovaries globose, ca. 1 mm diam., expanding to ca. 3 mm diam while inflorescence still fully intact; style very short apically expanded into a 3-lobed stigma; sterile interstice ca. 7 mm long, narrowed above corresponding to spathe constriction; synandrodia more or less rhomboid, 1.2-3 mm long; male zone ca. 1 cm long, subcylindric, ca. 4 mm diam.; synandria 4—5-merous, rhombo-hexagonal; synconnective somewhat inflated; thecae opening by apical pores; appendix about isodiametric with male zone at base, subcylindric, tapering in upper third, ca. 1.2—2 cm long, white; fruiting spathe ovoid, ca. 2 cm long; berries globose, ca. 4 mm diam. Distribution: Sumatera, known only from an area between G. Kerinci to Lake Kerinci straddling the border of West Sumatera and Jambi Provinces. Habitat. On montane forest floor at 1500-2000 m altitude. 324 Gard. Bull. Singapore 50(2) (1998) Morion Westmacot. Figure 11. Alocasia kerinciensis A. Hay Biinnenmeijer 9511 - A. habit; B. venation; C. inflorescence with part of spathe removed. - Scale: A, B, bar = 2 cm; C, bar = 4 mm. Alocasia in West Malesia and Sulawesi 325 Other specimens seen: SUMATERA: W. Sumatera, G. Kerinci, Bunnenmeijer 9106, 9308, 9416, 10128 (all BO); 9195, 10321 (both BO, L); Cult. RBG Sydney Acc. no. 970481 ex Jambi Prov., Kerinci Seblat National Park, above Lempur Village, Hay et al. 13046 (t, no voucher); 30. Alocasia cuprea (C. Koch & Bouché) C. Koch Alocasia cuprea (C. Koch & Bouché) C. Koch, Wochenschr. Vereines Befoerd. Gartenbanes Koenigl. Preuss. Staaten 4 (1861) 141; Engl. in A. & C. DC., Monogr. Phan. 2 (1879) 509; Ridl., J. Straits Br. Roy. Asiat. Soc. 44 (1905) 179; Engl. & K. Krause, Pflanzenr. 71 (IV.23E) (1920) 110; Merr., Bibliogr. Enum. Bornean Pl. (1921) 104; Merr., Pl. Elmer. Born. (1929) 26; Burnett, Aroideana 7 (1984) 76, figs 2 & 3. - Caladium cupreum C. Koch & Bouché, Ind. Sem. Hort. Berol., Appendix (1854) 6. Type: Not located, presumed destroyed at B. Neotype: Cult. RBG Kew ex Borneo, N.E. Brown s.n., May 11th 1876 (K; designated here). [Gonatanthus cupreus C. Koch, Wochenschr. Vereines Befoerd. Gartenbanes Koenigl. Preuss. Staaten 4 (1861) 141 - nom. in synon.] [? Caladium metallicum Ed. Otto, Hamburger Garten- Blumenzeitung (1853) 517, nom. subnud.; Koch, Berlinen. Allg. Gartenzeitung. 1 (1857) 1351; [Colocasia cuprea Engl., Araceae Exsiccatae et Illustratae No. 253 [date not ascertained, see Hay et al. (1995:174)]. - ?sphalm. pro Alocasia cuprea}. [Alocasia metallica Schott, Oesterr. Bot. Wochenbl. 4 (1854) 410, nom. nud.; Schott, Syn. Aroid. (1856) 46 (nom. superfl. pro Caladium cupreum); Hook., Bot. Mag. 86 (1860) t. 5190; Lemaire, Ill. Hort. 8 (1861) pl. 283; van Houtte, Fl. des Serres & Jardins 21 (1875) t. 2208-9]. Herb to ca. 80 cm tall; rhizome decumbent, to ca. 6 cm diam.; /eaves several together, each (?always) subtended by two marcescent reddish brown cataphylls, the first ca. 1/4 and the second ca. !/2 the length of the petiole; petiole to ca. 70 cm long, green, faintly mottled brown or greenish brown throughout, sheathing in the lower !/sth; blades coriaceous, hanging, ovate, bullate between the main veins, to ca. 60 cm long x 40 cm wide, adaxially glossy bronze-green, darker near the primary veins, abaxially deep purple, with a hyaline colourless margin ca. 1.5 mm wide; anterior lobe with the tip obtuse and abruptly and shortly acuminate; anterior costa with 8-11 primary lateral veins on each side, proximal ones diverging at ca. 100° then arching forward and outward to join a submarginal vein - more distal primary veins diverging at ca 60°; all primary veins with very 326 Gard. Bull. Singapore 50(2) (1998) conspicuous axillary glands abaxially; secondary veins forming well-defined undulating interprimary collective veins; posterior lobes completely united except for a shallow retuse notch, rounded; posterior costae diverging at ca. 20°; inflorescences paired, not forming multiple series, subtended by cataphylls similar to those subtending the leaves; peduncle similar to the petiole, to ca. 22 cm long; spathe green to greenish purple, ca. 10 cm long; lower spathe oblong ovoid, ca. 4.5 cm long ca. 2 cm diam; limb about equalling the lower spathe, at first erect and cucullate, then sharply deflexed, separated from the lower spathe by an abrupt constriction; spadix considerably shorter than the spathe - ca. 6 cm long, very shortly stipitate, cylindric except appendix; female zone narrowly cylindric, ca. 2 cm long x 8 mm wide; ovaries subglobose, longitudinally 3-4-ribbed; stigma raised on a very short slender style, conspicuously 2—4-lobed; sterile interstice not attenuate, isodiametric with male and female zones, ca. 2 whorls of rhomboid synandrodia; male zone cylindric, 2/3rds or all within the lower spathe, 2 cm long; synandria rhomboid, 4—6-merous, with the synconnective raised above but not overcapping the thecae; thecae opening by apical pores; appendix white, spindle-shaped, blunt, faintly irregularly channelled, ca. 2 cm long, constricted at union with male zone; fruit unknown. Distribution: Borneo, endemic to Sabah. Habitat: On slopes in rain forest, over a wide variety of substrates including ultramafics, sandstone and limestone, ca. 1000-1500 m altitude. Notes: 1. Confusion around the use of the epithet ‘metallica’ was discussed by Bunting and Nicolson (1963). Because of historical confusion over the identity of Caladium cupreum, the epithet metallica has been applied botanically both to what is here called A. cuprea (e.g. Hooker, loc. cit.) and to a form of Alocasia macrorrhizos |A. indica var. metallica Schott = A. macrorrhizos var. rubra (Hassk.) Furtado (which in turn, if to be regarded as a species separate from A. macrorrhizos, should be called Alocasia plumbea van Houtte)]. Assuming they are synonymous, the priority of Caladium cupreum is based on the paucity of the description in Otto (loc. cit) such that the earlier Caladium metallicum Otto is to be regarded as invalid. When Schott (loc. cit.) first published Alocasia metallica, he included Caladium cupreum as a synonym, thus rendering A. metallica superfluous. It can be clearly inferred that Schott intended A. metallica to be applied to, and interpreted Caladium cupreum as applicable to, a species different from what is currently called Alocasia cuprea. This is evident from Schott’s later work, when A. metallica was reduced to varietal status Alocasia in West Malesia and Sulawesi 327 under Alocasia indica (= A. macrorrhizos) (Schott, 1860: 145). Caladium cupreum was still a synonym in Schott’s view. It appears that Koch considered his A. cuprea and Schott’s A. metallica different species, though as the type of Caladium cupreum has not been found, it is not possible to prove the correct application of this name. Confounding matters, there is at K an outline, drawn by N.E. Brown, of a specimen from Koch’s herbarium, allegedly the type of Caladium cupreum, but resembling Alocasia macrorrhizos - hence implying that Alocasia metallica Schott and Caladium cupreum may be conspecific, as Schott had indicated. However, material of A. cuprea in the sense here, preserved at K, has the annotation by N.E. Brown - ‘A specimen of this was sent by me to Carl Koch, for comparison with his type of A. cuprea, & in reply he stated that it was certainly his A. cuprea & not A. metallica Schott’. This, together with the fact that Engler, who would almost certainly have seen the type at Berlin, applied the name A/locasia cuprea to this species, leads me to conclude that this application is correct. Moreover, Alocasia cuprea has been and currently is widely used, both botanically and horticulturally, in the sense used here. The accordingly designated neotype is the sheet annotated by N.E. Brown as above. 2. Alocasia cuprea has long been recognised as one of the most spectacular and bizarre foliage plants in the genus and is a parent of several interspecific horticultural hybrids (see Engler & Krause, 1920: 112; Burnett, 1984: 142). Its occurrence in the wild is sporadic, but it sometimes occurs in very densely abundant local populations (K.M. Wong, pers. comm.). Other specimens seen: SABAH: Kinabalu, N of Mesilau Camp, Allen AK 66-38 (SING); Cult. RBG Sydney Acc. no. 912634 ex cult. RBG Edinburgh Acc. no. 19852175 ex Kinabalu, Marai Parai, Argent s.n. (NSW); Kinabalu, Penibukan, nr Dahobang R., Clemens & Clemens 40588 (SING); Elphinstone Prov., Tawao, Elmer 20471 (BO, GH, K, L, SING); Cult. RBG Sydney Acc. no. 960584 ex Maliau Basin, G. Rara F.R., 2.5 km above main Maliau Falls, Hay et al. 12092 (NSW, voucher SAN); Kinabalu, S$. Dahobang, Holttum s.n. (SING); Cult. RBG Sydney Acc. no. 841539 ex Tenom, Kallang Falls, Wallace 84/206 (no voucher); Inadequately Known Species 31. Alocasia sp. A. Herb to ca. 40 cm tall; Jeaves several together, glabrous; petiole ca. 30 cm long, sheathing in the lower ca. !/3, green, spotted purple; blade ovato- sagittate, to ca. 30 cm long, coriaceous, somewhat bullate, adaxially grey- green, dark green about the main veins, abaxially purple; anterior lobe widest somewhat above the base, the tip acute to obtuse and apiculate; 328 Gard. Bull. Singapore 50(2) (1998) anterior costa with 5-6 primary lateral veins on each side, diverging at 80-50°, with conspicuous axillary glands abaxially; secondary venation forming abaxially and adaxially conspicuous subsidiary veins themselves forming interprimary collective veins in the outer part of the leaf blade, the remaining secondary venation obscure abaxially, faint adaxially; posterior lobes acute, about !/2 the length of the anterior, the inner sides very narrowly lanceolate; posterior costae diverging at ca. 90°, naked in the sinus for ca 1 cm; inflorescence unknown. Distribution: Sarawak. Habitat: Reported from forest floor among limestone rocks. Notes: 1. Plants of this species are cultivated in the Semenggoh botanic garden, near Kuching, and are said to have been collected from the wild locally (P. Boyce, pers. comm.). 2. This highly ornamental plant is traded in the U.S.A. under the name Alocasia guttata var. imperialis or Alocasia guttata Imperialis. An image may be found at http://www.skg.com/alocasia3.html. This species is evidently allied to A. scabriuscula (which includes A. guttata) and A. reginae, differing from both in the variegated leaf blade, and from the former in its smaller stature and bullate blade, and from the latter in being glabrous and more robust with a larger number of primary lateral veins. | would suggest to the horticultural community that either an altogether new cultivar name is formally proposed or that the plant be called Alocasia Imperialis and that a standard be designated and preserved to fix the application of the cultivar name Imperialis to this particular clone, so that there is no longer any ambiguity about whether or not the plant is the same as A. guttata var. imperialis. Doubtful Species and Records Alocasia pallida C. Koch & Bouché, Ind. Sem. Hort. Berol., App. (1854) 5. If a type ever existed of this, it was presumably destroyed in the bombing of Berlin. Koch & Bouché described it from sterile material without provenance, compared it with Alocasia montana, which itself appears to be a synonym of A. macrorrhizos, and distinguished it (trivially) on the basis of the ‘stemless’ habit of A. montana. They further compared it with A. alba Schott, noting that the latter differed in its slightly peltate leaves Alocasia in West Malesia and Sulawesi 329 (which it does have as a juvenile, like most species in the genus). Engler (1879) and Engler & Krause (1920) placed A. pallida in the synonymy of A. alba. However, since Engler’s interpretation of A. alba appears to have been associated with material only from Sri Lanka (where that species in the strict sense does not naturally occur), there is some doubt about their determination. It seems likely that this is a synonym and variant of A. macrorrhizos. Alocasia warburgii Engl., Bot. Jahrb. Syst. 25 (1898) 25 (= Alocasia heterophylla (Presl) Merr.). Engler cited Warburg 15723 from Sulwesi in the protologue; the specimen (and a Philippine syntype) is presumed destroyed at B. Engler later concluded that A. warburgii was conspecific with A. heterophylla, a distinctive Philippine species of which no other Sulawesi material has been found (Hay, in press). Alocasia wavriniana Mast., Gard. Chron. 21 (1898) 241, fig. 98 (= Alocasia lauterbachiana (Engl.) A. Hay). This was originally attributed to Sulawesi, but no material with this provenance authenticated has been found. Alocasia lauterbachiana is from New Guinea and the Bismarck Archipelago (Hay & Wise, 1991). Acknowledgements I am greatly indebted to Conrad D. Fleming for his generious financial support of field work in Peninsular Malaysia, Borneo, Java and Sumatera over the last four years. For invaluable assistance in the field I am very grateful to: Prof. Ruth Kiew, Mr Anthonysamy and Roy Banka (Universiti Pertanian Malaysia) in 1994, Yahud bin Hj. Wat, Saupel B. Atot and Chang Ban Sem (Sarawak Forestry Department) in 1994, Dr Wong Khoon Meng, Mr A. Berhaman, Ubaldus, Postar and Ahmat (Forest Research Centre, Sepilok) in 1995 and 1996, and Dr Ir. Suhirman, Dra Yuzammi and Pak Tahrodin (Kebun Raya Bogor) in 1996 and 1997. The Directors or Curators of the following herbaria made their material available: B, BM, BO, BRI, E, GH, K, KEP, L, MO, SAN, SAR, SING, UPM, US. I thank Prof. David Mabberley for his invaluable advice on matters of typification and nomenclature, and Peter Boyce and Ken Hill - the latter during his tenure as Australian Botanical Liaison Officer at K - for generous assistance with library matters. I am grateful to Sylvia Fitzgerald, Chief 330 Gard. Bull. Singapore 50(2) (1998) Librarian at K for permission to reproduce the type of Arum montanum. | also thank Dewey Fisk for providing very informative material of Alocasia in his cultivated collection. I would also like to thank Ian McLellan, Randy Sing and nursery staff of the RBG Sydney for cultivating the living plants cited here, Lesley Elkan and Marion Westmacott for the botanical drawings, Suzanne Bullock for the photographs in Plate 1, and Clare Herscovitch for stoic technical assistance and curation of the living Aroid collection at Sydney. Peter Boyce, Ruth Kiew and Dr Dan Nicolson (US) provided constructive criticism of the manuscript. References Alderwerelt van Rosenburgh, C.R.W.K.van, 1920, 1922. New or noteworthy Malayan Araceae I. Bulletin Jardin Botanique Buitenzorg. Il, 1: 359- 389. - Ditto II. Ibid. 4: 163-229. - Ditto III. Ibid. 4: 320-347. Backer, C.A. & R.C. Bakhuizen van den Brink, 1968. Flora of Java, Vol. Il. Wolters-Noordhof N.V., Groningen. Blume, C.L. 1823. Catalogus der gewassen ‘s Lands Plantentuin. Batavia. Blume, C.L. 1836/7. Collectanea ad monographiam Aroidearum. Rumphia. 1: 71-154. Brown N.E., 1884. Supplement to G.W. Johnson (ed.), The Gardeners’ Dictionary. G. Bell & Sons, London. Bunting, G.S. 1962. The genus Schizocasia (Araceae). Baileya. 10: 112- 120. Bunting, G.S. & D.H. Nicolson, 1963. The Alocasia plumbea confusion. Baileya. 11: 142-146. Burnett, D. 1984. The cultivated Alocasia. Aroideana. 7: 67-163. Engler, A. 1879. Araceae. Pp. 1-681 in A. & C. de Candolle (eds.), Monographiae Phanerogamarum. Vol. 2. Masson, Paris. Engler, A. & K. Krause 1920. Araceae-Colocasioideae. Das Pflanzenreich. 71 ([V.23E): 1-139. Engelman, Leipzig. Forman, L.L 1997. Notes concerning the typification of names of William Roxburgh’s phanerogams. Kew Bulletin. 52: 513-534. Forster, J.G.A. 1786. De plantis esculentis insularum oceani australis commentatio botanica. Haude & Spener, Berlin. Alocasia in West Malesia and Sulawesi 331 French, J.C., M. Chung & Y. Hur, 1995. Chloroplast DNA phylogeny of the Ariflorae. Pp 255-275 in P. Rudall et al. (eds.), Monocotyledons: Classification and Evolution. Vol. 1. Royal Botanic Gardens Kew. Furtado, C.X. 1941. Alocasia macrorrhiza and its varieties. Gardens Bulletin Singapore. 11: 244-257. Gentry, A.H. 1990. Herbarium taxonomy versus field knowledge - is there an attainable solution? Flora Malesiana Bulletin. Special Vol. No. 1: 31- BD. Grayum, M.H. 1990. Evolution and Phylogeny of the Araceae. Annals Missouri Botanical Garden. 77: 628-697. Hasskarl, J.K. 1842. Plantarum genera et species novae aut reformatae javenses. Flora. 25 (2), Beibl.1: 1-16. Hasskarl, J.K. 1844. Catalogus Plantarum in Horto Botanico Bogoriensi Cultarum, alter. Batavia. Hay, A. 1994a. Alocasia simonsiana - a new species of Araceae from New Guinea. Blumea. 38: 331-333. Hay, A. 1994b. Araceae - an internationally co-ordinated major family for the accelerated Flora Malesiana Project. Flora Malesiana Bulletin. 11: 5- 8. Hay, A. 1996. A new Bornean species of Colocasia Schott (Araceae- Colocasieae), with a synopsis of the genus in Malesia and Australia. Sandakania. 7: 31-48. Hay, A. (in press). A revision of the genus Alocasia (Schott) G. Don (Araceae-Colocasieae) in the Philippine Islands. Nordic Journal Botany. Hay, A., J. Bogner, P.C. Boyce, W.L.A. Hetterscheid, N. Jacobsen & J. Murata, 1995. Checklist and Botanical Bibliography of the Aroids of Malesia, Ausralia and the Tropical Western Pacific. Blumea. Suppl. 8. Rijksherbarium/Hortus Botanicus, Leiden University. Hay, A. & D.J. Mabberley, 1991. ‘Transference of Function’ and the origin of aroids: their significance in early angiosperm evolution. Botansiche Jahrbiicher fiir Systematik. 113: 339-428. Hay, A. & R. Wise, 1991. The genus Alocasia (Araceae) in Australasia. Blumea. 35: 499-545. 332 Gard. Bull. Singapore 50(2) (1998) Hooker, J. D. 1894. The Flora of British India. Vol. 6. Reeve & Co., London. Hotta, M. 1967. Notes on Bornean plants. Il. Acta Phytotaxonomica et Geobotanica. 22: 153-162. Koorders, S.H. 1911. Exkursionsflora von Java .Vol. 1. G. Fischer, Jena. Kunth, C.S. 1841. Enumeratio Plantarum. Vol. 3. J.G. Collae, Stuttgart & Tiibingen. Leick, E. 1915. Die Erwarmungstypen der Araceen und ihre bliitenbiologische Deutung. Berichte der Deutschen Botanischen Gesellschaft. 33: 518-525. Loureiro, J. de, 1790. Flora Cochinchinensis. Lisbon. Mayo, S.J., J. Bogner & P.C. Boyce, 1997. The Genera of Araceae. Royal Botanic Gardens, Kew. Miquel, F.A.W. 1855/1856. Flora van Nederlandsch Indié. Vol. 3. Van der Post, Amsterdam. Nicolson, D.H. 1963. Nomina conservanda proposita: Alocasia (Schott) G. Don. Taxon. 12: 208-209. Nicolson, D.H. 1968. The genus Xenophya Schott. Blumea. 16: 115-118. Nicolson, D.H. 1979. Araceae. Pp. 438-460 in A.C. Smith (ed.), Flora Vitiensis Nova, vol. 1. Pacific Tropical Botanic Garden, Lawai. Nicolson, D.H. 1987. Araceae. Pp. 17-101 in M.D. Dassanayake & F.R. Fosberg (eds.) Flora of Ceylon, Vol. 6. Amerind, New Dehli. Noltie, H.J. 1994. Araceae. Pp. 121-210 in Flora of Bhutan, Vol. 3, Part 1. Royal Botanic Garden Edinburgh. Pijl, L. van der, 1933. Welriekende vliegenbloemen bi Alocasia pubera. Tropische Natuur. 22: 210-214. Rafinesque, C.S., 1837. Flora Telluriana. Vol. 3. Philadelphia. Ridley, H.N. 1904. Some new Malayan plants. Journal Straits Branch Royal Asiatic Society. 41: 31-51. Ridley, H.N. 1905. The aroids of Borneo. Journal Straits Branch Royal Asiatic Society. 44: 169-188. Alocasia in West Malesia and Sulawesi 333 Ridley, H.N. 1925. Flora of the Malay Peninsula. Vol. 5. Reeve & Co., London. Roxburgh, W. 1814. Hortus Bengalensis. Mission Press, Serampore. Roxburgh, W. 1832. Flora Indica. (Ed. 2, Carey). Thacker & Co., Calcutta. Rumphius, G. 1747. Herbarium Amboinense. Vol. 5. Amsterdam. Schott, H.W. 1832. Aroidearum synopsis. Pp. 15-22 in H.W. Schott & S.L. Endlicher, Meletemata Botanica. Gerold, Vienna. Schott, H.W. 1853-1857. Aroideae. Gerold, Vienna. Schott, H.W. 1856. Synopsis Aroidearum. Mechitarists’ Press, Vienna. Schott, H.W. 1858. Genera Aroidearum Exposita. Holzel, Vienna. Schott, H.W. 1860. Prodromus Systematis Aroidearum. Mechitarists’ Press, Vienna. Schott, H.W. 1861. Aroideologisches. Bonplandia. 9: 367-369. Schott, H.W. 1863. Araceae. Annales: Museum Botanicum Lugduno- Batavum. Lugd.-Bat. 1: 122-131. Sweet, R. 1839. Hortus Brittanicus. [Ed. 3, G. Don]. Ridgeway, London. Trimen, I.H. 1898. A Handbook to the Flora of Ceylon. Part IV (ed. J.D. Hooker). Dulau & Co., London. Ventenat, E.P. 1801. Description des plantes nouvelles....dans le jardin de J.M. Cels. Paris. Zhu, Z.Y., 1985. Panzhuyuia Z.Y. Zhu - a new genus of Araceae from Emeishan China. Journal Sichuan Chinese Medical School. 4 (5): 49-52. Index to Species Alocasia LSTA Sn ERE SRR DEI alte TNS ge OR ONE PASE SR oP 289 EE EARN ESS eS CSE EAU NCEE A Ce OA Oe 297 TEE SEIT) SORUTAESR SSR ES OE MOA BON NB Sk Se A DER ee a 291 ee crt ee ee eee Woes ran Miia al Ni A toe Nila a 318 Grist Cen Pek man is ie ee LUNN Ae aa eh COL outed iat 9 SS BERANE NC. a ACRE ah A ERR ae OOM eae ROR FOE Bae 334 Gard. Bull. Singapore 50(2) (1998) flemingionay,..v. sue AN AR ae eee 292 EFROVTIDUEGE 6. bi dace ass sh bald adds Pa eten hale eee Col ay Sate cee 286 KerlACLENSIS : 6 hos olbass ole adn Wed as cada Le ee eee 322 longiloba).. 2. UOT atte code cere ne tee ET Ree oe ne eee ee 299 INACTOVIPUZOS 6 i) Vas pene tian ng git det sind ths Med ey ee ee 283 PCL Oj vnininsvnrhe os beco eae eee hes Daa bce Nee Sed alde apse aa eee 260 TAINUS CURE) EU i Bah dle va ee cl Aa i ee 320 DOMId| os eA dele ya Anaad beg eh NC ae faa te | ieee ae aS 328 JECT el 601 | ELGG or ORD MME Minar Ay babiocssahait shay \be cette yyh ede Ve POM hs oN Ue a adas ots ld mem eer Ram rege eae nea 32] DETAKEASIS: 1) cD PEN sae wld tdele's sudo aes Ae ae ee Ae ee 316 PTIRGCEDS SANS. SINR ton ea Ble Rh 266 Prifecipianluse 2) Mead ask: RAR ee en es ee 277 DUDET eo Sie i aac e doc le pace big YAN a a) ee 242 PUL C TE rea i ilece oa 0d Wn bab ojo Pe Oe aA De OSE eR ee a ea 275 VOQUUAR SE Si eich ola Gren bin plallh bie d Sale a/Maille mA hala ele ucla aaa ern ieee a 25% POCO Nolen chs ovaie 20, bale) et buialt ed added eats Sie Rea ee kee ee 258 POVEF SOAR le ba La clea goth ocala wba MN dae reel ATO leat ae Ang ec 279 PILE SEN ais lino! UO areas Oe 2 263 FOWOUSTAN ANE. ok los aiesiad wade Band pean eee eee ea enon ae 249 SOTO WAR EHSTS oo ois salunaien ce oedithove Slee ak chai ued ck oaene vee on 247 SCERATUUSOCTLEE: i.e ss v'ald sin tigigueia Gala WU ane NA Ad Ube Oesecs Mie pe meee tire oleate eee 253 ye NING Dani Nek MMR ILE RE ee at Bicceamih canne tein aance abel ah eee enans 327 SUELALIUPUGILUCIUG si8is)2 i 00 sce ote be eR sia sea marshal ek a 314 VETS EN RU iA ROD OE AOR 281 WORDED Gale dike coils kes die vd pata ciate aeRO ee poet a cs See 329 WAV FUERA ON Pos va ais a dn ae RIE ee UD le aie es 329 WOPTRTE JEL Sosa Nols va 5 ws weles eed ete oteleton ha te tears oh ae aa te a 270 Gardens’ Bulletin Singapore 50 (1998) 335-336. FLORA MALESIANA SERIES II - FERNS AND FERN ALLIES, VOLUME 3 (Polypodiaceae, Davalliaceae, Azollaceae, Cheiropleuriaceae, Equisetaceae, Matoniaceae, Plagiogyriaceae), edited by C. Kalkman and H. P. Nooteboom, vi + 1-334 pp. (1998). Rijksherbarium, P. O. Box 9514, 2300 RA Leiden, The Netherlands. This new volume of series II of Flora Malesiana on Ferns and Fern Allies contains revised treatments of seven families of Malesian pteridophytes by various authors. The families revised are as follows: Polypodiaceae by P. H. Hovenkamp et al., pp. 1-234; Davalliaceae by H. P. Nooteboom, pp. 235-276; Azollaceae by R. M. K. Saunders, pp. 277-284; Cheiropleuriaceae and Equisetaceae by J. E. Leferriere, pp. 285-286 & 287-288; Matoniaceae by M. Kato, pp. 289-294; and Plagiogyriaceae by X.-C. Zhang and H. P. Nooteboom, pp. 295-316. A two-page abstract precedes the volume and an index to the scientific plant names concludes it. The format of the new pteridophyte volume is similar to that of the new volumes of seed plants in series I. Compared to the old format, the layout of paragraphs in the new volume runs across the entire page width, instead of forming two columns. The font size of the print is also bigger, which makes the reading of the text easier. For each family, one sees a general description, followed by a concise discussion of the distribution, morphology, habitat and ecology, chromosomes and taxonomy, and at times, economic importance. The same categories of information are repeated for the genera and species. Well-constructed, dichotomous keys to genera and species within the family, extensive synonymy, taxonomic bibliography and accurate illustrations are also provided. For speciose and difficult genera, such as Microsorium and Selliguea, separate keys to the species known from a large island and country, or from an island group, are provided. Because of the number of included taxa, the family Polypodiaceae, with 18 genera and 183 species, easily becomes the main feature of this large volume, to be followed by Davalliaceae (3 genera and 31 species) and Plagiogyriaceae (one genus and 7 species), in terms of family size. The rest of the families treated are either monotypic or oligotypic. Important taxonomic ideas put forth in the new volume include the generic -fusion between Humata and Davallia, Pyrrosia and Drymoglossum, Crypsinus and Selliguea, and also, Phymatosorus and Microsorum. The supportive arguments justifying a broad concept for these genera have been published previously by the authors and are not repeated in volume 3. On the other hand, several traditionally accepted small genera, such as Photinopteris (=Aglaomorpha), Thayeria (=Aglaomorpha), Merinthosorus (=Aglaomorpha), Schellolepis (=Goniophlebium) and Araiostegia 336 Gard. Bull. Singapore 50(2) (1998) (=Davallia) are not recognized by the authors of this volume. No taxonomic novelty is described in the new volume. Aglaomorpha acuminata ( Willd.) Hovenkamp is published as a nomenclatural novelty. The present volume, with its updated revision, is truly a handy source of taxonomic information for the seven families of Malesian ferns treated. Being an occasional student of Malesian fern taxonomy, I find the discussion on the family morphology and relationship, as well as the many taxonomic comments scattered through the pages, very educational and enlightening. However, I miss the selected distribution maps of plant taxa so elegantly reproduced in early volumes of this series. To me, these range maps provide an effective visual aid to our understanding of the dispersal and evolution of the Malesia flora. Future volumes of series II should perhaps consider the inclusion of distribution maps of selected pteridophytes to illustrate the biogeographical highlights of Malesian fern flora. Undoubtedly, the usefulness of a flora revision lies in its inclusiveness of the taxa found locally and its workability with the specimens collected from the area. Judging by the text presentation, especially the wording of the dichotomous keys and the illustrations, the new volume appears to be another excellent and useful guidebook to the correct identification of Malesian ferns. Both the authors and the editors of this new volume are to be congratulated for a difficult job well done. Benito C. Tan Department of Biological Sciences National University of Singapore va é { | vt M ef re me Ni ; itive iat a ‘i ra ; ops ae sai ieee hi Paha ie | ing Th my Ae Wr Cua Fe ia ‘ ae eho i My 2 UY a - ere Pee i ' Bee 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. Numerical data should only be included if it is essential to the argument and this can be presented either in the form of tables or diagrams. Once the papaer is accepted, a diskette with the final version is required. Titles and authors: The title should give a concise description of the content of the paper. The name(s) and affiliation(s) of author(s) must be given below the title. Lengthy papers and those of a complex nature must have the contents listed at the beginning of the paper. Scientific names: The complete scientific name - genus, species, authority, and cultivar where appropriate - must be cited for every organism at time of first mention. The generic name may be abbreviated to the initial thereafter except where intervening references to other genera with the same initial could cause confusion. Tables: All tables should be numbered and carry headings describing their content. These should be comprehensive without reference to the text. Abbreviations: Standard chemical symbols may be used in text (e.g. IAA, IBA, ATP), but the full term should be given on the first mention. Dates should be cited as: 3 May 1976. Units of measurement should be spelled out except when preceded by a numeral where they should be abbreviated in standard form: g, mg, ml, etc. and not followed by stops. Literature citations: Citations in the text should take the form: King. and Chan (1964). If several papers by the same author in the same year are cited, they should be lettered in sequence (1964a), (1964b), etc. When papers are by three or more authors they should be cited as e.g., Geesink et al. (1981). All references must be placed in alphabetic order according to the surname of the (first) author and the journal title be given in full, as in the following example: Stone, B.C. 1994. Additional notes on the genus Glycosmis Correa (Rutaceae). Gardens’ Bulletin Singapore. 46: 113-119. References to books and monographs should be cited according to the following form: Ridley, H.N. 1930. The Dispersal of Plants Throughout the World. L.Reeve, Ashford, Kent. For literature citations in taxonomic papers the following style is required: Medinilla alternifolia Blume, Mus. Bot. Lugd.-Bat. I:2 (1849) 19. Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) 524. Illustrations: Drawings should be done in indian ink. Authors should indicate where individual illustrations receive first mention in the text. Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid for, prior to publication. Manuscripts should be sent to: THE EDITOR, GARDENS’ BULLETIN SINGAPORE, SINGAPORE BOTANIC GARDENS, CLUNY ROAD, SINGAPORE 259569 =! 5 Sanne Poe ale aaa aren eee ty lla go Fe LO ee ae ets an we chengipa—te-e lety tt oe ay abate. See eee oe eo eee es Sage ere pees et 2 Soeeegaacs icone = A ¥ i J wr bo ihe i A er ie ; 2 na # ‘) { , Py € 4 dics ay Ag * Li : = »* a 6: ’ 1 5 ¥ 14) } ay Pp) i 4 a te ' 4 o> 2 } s P| , 6 f a \ 4 i" 4 : ; : { - “ 4 « ~ i ¥ Se, } ;

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