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The Gardens' Bulletin
Singapore Volume 55
Singapore Botanic Gardens 2003
© Singapore Botanic Gardens Singapore, 2003
Applications for reproduction should be made to the Singapore Botanic Gardens, 1 Cluny Road
Singapore 259569
DATE OF PUBLICATION OF THE TWO PARTS WERE:
Part | Pp. 1-172 17 September 2003 Partt2 Pp. 173-315 15 December 2003
a
Gardens’ Bulletin Singapore 55 (2003)
CONTENTS
Chew, W.-L. PTS. 1 Wer AOC Se eA Sere we whines aks wa dads s adaaiepumkee i iuda sccenouaabeeoanstvorsedl &
Chua, L.S.L and L.G. Saw A new record of Dryobalanops beccarii (Dipterocarpaceae ) from Peninsular UMMA re Net it de dao a Sete ee ee Re a piositecn us pean das vn noxteh@eunden od sar onee cag iars ntekogs sogeepaapau
Chua, L.S.L and L.G. Saw Conservation notes on Vatica yeechongii (Dipterocarpacae) from ae nae AOA erie Dh ou inh kus ook avon nine ules Oba08 Sevinoncpeplgnwws Dasennnasnetemnanwakasvodmeseand
Chung, R.C.K., E. Soepadmo and A.L. Lim. The significance of pollen morphology in the taxonomy of Grewia and
Microcos (Tiliaceae) in Peninsular Malaysia and Borneo ....................0.0sss000secceeee ee 02-239
Gusman, G.
Arisaema fimbriatum (Araceae) and its intraspecific Variation ................eseeseeeeerereee ee e201 Hassan I. Amote On ordia premnifolia Rid). (Borapinaceat) «isi. ac<cccooeetessciencdpcvadsgense so vapess ser LOD
Hay, A. and C. Herscovitch A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ...........ccceceeeeeeeees 27
Herscovitch, C. and A. Hay An unusual new species of Homalomena (Araceae) from New Guinea ...............::ceee 31
Hilliard, O.M., B.L. Burtt and M.H. Bokhari
Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from Borneo...................0068 35 Kiew, R. Begonia peninsulae (Begoniaceae), a confused species from Malesia ..................::00000 61
Kiew, R. and C. Geri
Begonias from the Bau limestone, Borneo, including a new SpeCi€s ..............:.::eeeeeeeeeees 113 Kiew, R. and I.M. Turner
te ity Plants CnieiIC 10 wai APOE 1 55.0 yd ned a escies dn tod dctence c dbdvcenereetenereescdaveen as Bansel 1S Kloet, S.P. Vander
Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ...........0:.:ccccesseeeeeees 163
Lee, S., A. Samsuri, P. Leong, Ali Ibrahim and A.-T. Gwee A botanical survey of Chek Jawa, Pulau Ubin, Singapore ..................sscscssessscccsestee se ees 271
Mabberley, D.J. New species of, and other notes on, Chisocheton and Walsura (Meliaceae)................... 189
Middleton, D.J. A new species and a new combination in Bornean Kopsia (Apocynaceae:
APOCYNOIGEAREC)..........veseaceesianniesnantbuasivniiosaicncinesieh niniiov ch Geoedeeitns tual tse
Middleton, D.J.
A revision of Dyera (Apocynaceae: Rauvolfioideae) ...............csscccecccceeeeeesssreeee ee ee
Pearce, K.G.
New species and varieties of Symplocos (Symplocaceae) from Borneo...............:::0++
Pearce, K.G. Five new Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia ..................000
Skornickova, J., M. Sabu and M.G. Prasanthkumar
A new species of Curcuma L. (Zingiberaceae) from Mizoram, India ...............eeee
Skornickova, J., M. Sabu and M.G. Prasanthkumar Curcuma codonantha (Zingiberaceae) — a new species from the Andaman Islands,
Tinta ....ceccacaccovecscvsdeccsaeescccencsxceensnvazses<wawebuuystadaes gone ee ace ne Ese ee ee eee aaa ea
Vermeulen, J.J. and P. O’Byrne
New species and new records of Southeast Asian Bulbophyllum (Orchidaceae).........
Vermeulen, J.J. and P. O’Byrne
Six new species of Bulbophyllum (Orchidaceae) from Sulawesi ............:::cceeeeeeeeeree es
Wong, C., G. Argent, R. Kiew, Ohn Set and Y.Y. Gan The genetic relations of Musa species from Mount Jaya, New Guinea, and
a reappraisal of the sections of Musa (Musaceacy Tittt.J2 is. .nceresscsensteoastenes en ate te
Zahid, M.S.
Three new species of Porterandia (Rubiaceae) from Mount Kinabalu, Borneo..........
Obituary
Mohamad Shah bin Hj Mohamad Noor - R.Kaiew............ 00. ...ccscccccececccceeeeee ee eeceee ones
Dr Chang Kiaw Lan - K.M WOmtng.......020co: oe0ss0% «050 s00nse oysodnvsxsntranaaeen ian
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THE GARDENS’ BULLETIN
The Garden’s Bulletin Singapore publishes papers on plant taxonomy (including revisions), horticulture, phytogeography, floristics, morphology, anatomy and related fields with emphasis on plants in the West Malesian region.
EDITORIAL BOARD
Dr Ruth Kiew (Editor) Singapore Botanic Gardens
Dr P.Y. Tan (Assist. Editor) Singapore Botanic Gardens
Dr S. C Chin Singapore Botanic Gardens
Dr M.J.E. Coode Royal Botanic Gardens Kew, U.K.
Dr R.T. Corlett University of Hong Kong Hong Kong
Dr M.C. Roos National Herbarium of Netherlands Leiden University Branch, Netherlands
Dr E. Soepadmo Forest Research Institute Malaysia Kepong, Malaysia
Dr W.K. Tan Singapore Botanic Gardens
The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. Neither the National Parks Board not the Editorial Board is responsible for the opinions or conclusions expressed by the contributing authors.
The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including postage. Overseas subscribers are required to make payment in the form of bank drafts or international money orders in Singapore currency payable to National
Parks Board, Singapore.
Instructions for contribiting authors are found on the inside backcover.
‘ * 0 3 2003 ‘ et acl
The Gardens’ Bulletin
Singapore
VOL. 55 (Part 1) July 2003 ISSN 0374-7859
CONTENTS
Chua, L.S.L. and L.G. Saw
A new record of Dryobalanops beccarii (Dipterocarpaceae) from Fane VSB Bs Sete fas ees tetrad! oc aes:! use 1, oily. dgee iy als CMa aweeeseceoe d
Chua, L.S.L. and L.G. Saw
Conservation notes on Vatica yeechongii (Dipterocarpaceae) from Be eRe Nh Nts os os dace pee damnns weg dees give ndeeeoebiod bani es ‘4
Chew, W.-L.
SLMS th WIRMeSIal PigeraCege Beis tihs anys vein ches iencs'ssl bite qepehgemedovenxdeeoveheers.y0004 3
Hay, A. and C. Herscovitch
A new species of Schismatoglottis (Araceae) from Sabah, Malaysia ............... 27
Herscovitch, C. and A. Hay
An unusual new species of Homalomena (Araceae) from New aN a BS a cn ae cctas as atid aanicd apuumpt nc cksen ipiscsie davcpibaiine 31
Hilliard, O.M., B.L. Burtt and M.H. Bokhari
Pleuroschisma, a new section of Cyrtandra (Gesneriaceae) from
EE re TURE Node tees et Ae ee A EP = Kiew, R.
Begonia peninsulae (Begoniaceae), a confused species from Malesia..................04. 61 Middleton, D.J.
A new species and a new combination in Bornean Kopsia AR ON NEN IN Fetes Niet nso a ia lsvannepecigeraneenasesseusvencasdncwnnese 65
Pearce, K.G.
New species and varieties of Symplocos (Symplocaceae) from Borneo................ 69 Pearce, K.G.
Five new Begonia species (Begoniaceae) from the Niah National Park,
Sarawak, Malaysia... sisccccck cited escarole Ste a 73
Skornickova, J., M. Sabu and M.G. Prasanthkumar A new species of Curcuma L. (Zingiberaceae) from Mizoram, India.................... 89
Wong, C., G. Argent, R. Kiew, Ohn Set, and Y.Y. Gan
The genetic relations of Musa species from Mount Jaya, New Guinea,
and a reappraisal of the sections of Musa (Musaceae) .............cscssecccsesseeeee eee iD] Kiew, R. and C. Geri Begonias from the Bau Limestone, Borneo, including a new species ................. 113
Vermeulen, J.J. and P. O’Byrne
New species and new records of Southeast Asian Bulbophyllum
(Orchidaceae) -.2.2i0c ssc csceceswsieattderstarresliecevave fess pest Le ee 125 Kloet, S.P. Vander
Re-examination of Vaccinium dialypetalum (Ericaceae) — Erratum ................... 163 Obituary
Mohamad Shah bin Hj Mohamad Noor ................. 00. seessseeeeeeeeeeeee RY Kiiew-165
Date of Publication: 17 September 2003
Published by National Parks Board Singapore Botanic Gardens 1 Cluny Road Singapore 259569
Printed by Print Dynamics (S) Pte Ltd
Garden’s Bulletin Singapore 55 (2003) 1-6
A New Record of Dryobalanops beccarii (Dipterocarpaceae) from Peninsular Malaysia
L.S.L. CHUA AND L.G. SAW
Forest Research Institute Malaysia, Kepong 52109, Kuala Lumpur, Malaysia
Abstract
Dryobalanops beccarii Dyer, Dipterocarpaceae, first described and recorded from Borneo, has now been documented from Johore, Peninsular Malaysia, where it grows on ridges in several compartments in the Panti Forest Reserve, Kluang Forest Reserve, and the Labis Forest Reserve. It is a large tree frequently reaching 40 m height and more than 1 m diameter.
Introduction
Two species of Dryobalanops, D. aromatica and D. oblongifolia, were recorded for Peninsular Malaysia (Ashton, 1982). The genus Dryobalanops is clearly distinguished from other dipterocarp genera by the presence of five wing-shaped, subequal fruit calyx lobes and closely parallel venation on its leaf blade. In Peninsular Malaysia, trees of both species are characterized in the field by scaly bark, peeling in large, irregular, thin flakes. In D. oblongifolia, leaves are oblong or oblong-lanceolate and the fruit calyx lobes are shorter than the nut, while in D. aromatica, leaves are oval or sub-ovate and the calyx lobes are longer.
During a plant diversity enumeration exercise in the Virgin Jungle Reserves (VJR) network in Peninsular Malaysia, Ang Khoon Cheng and Kamarudin Saleh discovered in Kluang Forest Reserve (FR) and the VJR in Labis FR, Johore, trees that had leaves that closely resembled D. aromatica but the bark was distinctly different in that the outer bark was scaly, very shallowly fissured and had a laminate inner bark. It was clear that these populations belonged neither to D. aromatica nor D. oblongifolia but their identity could not be confirmed in the absence of flowers and fruits.
A dipterocarp mast-fruiting episode occurred in Peninsular Malaysia during the second and third quarters of 2002. Taking advantage of this cyclic event, teams from the Botany and Seed Technology Units, Forest Research Institute Malaysia (FRIM), launched an extensive fruit collection exercise, targeting species that are rare and/or threatened. Fruiting trees in several of these Dryobalanops populations
i)
Gard. Bull. Singapore 55 (2003)
enabled materials for herbarium specimens and fruits for ex situ planting to be collected. The identity of the species could be confirmed and proved to be D. beccarii Dyer. A full description of the species based on characters from these populations is provided below.
Dryobalanops beccarii Dyer J. Bot. 12 (1874) 100; Ashton, Fl. Malesiana I (9) (1982) 375. Type: Malaysia, Sarawak, Matang, Beccari PB2944 (K, lecto).
Large tree to 30 m tall, 63.7 cm diam., with small buttress to 1.5 m tall. Bark shallowly fissured to scaly, flaky and peeling off in pieces on the lowest part of the trunk immediately above the buttress, greyish-white; inner bark laminate, reddish-brown; sapwood yellowish brown, exuding resin when cut. Twigs 3 mm diam., glabrous, drying almost black. Leaves thickly coriaceous, glabrous, drying brown above, dark brown below; petiole 10-17 mm long, 1 mm thick, glabrous, drying almost black; blade broadly ovate, 3.5—6.3 x 2.4—3.7 cm, apex tapering to a 7-mm long acumen, base broadly cuneate to rounded, margin thickened, midrib prominently raised below, sunken above, secondary veins numerous, closely parallel, faint on both surfaces, smelling of camphor smell when crushed. Flowers not seen. Fruit calyx glabrous, shallow cup tapering gradually and cuneate at the pedicel, lobes 5, equal, oblong- spathulate, base to 2 mm, yellowish green when fresh, 4.0-6.5 x 0.6—1.0 cm, veins conspicuously raised on both surfaces, nut ovoid, glabrous, yellowish green when fresh, to 1.4 x 0.9 cm, style remnant 1 mm long, free from calyx except the base. Vegetatively, the leaf morphology of Dryobalanops aromatica and D. beccarii is very similar and hence it is difficult to differentiate them when sterile. However, the bole character as described above can readily distinguish the two species. In addition, the fruits of D. beccarii are different from D. aromatica (Table 1).
Table 1: The bark and fruit characters of Dryobalanops aromatica and D. beccarii
D. aromatica D. beccarii Outer bark long, thin, recurved, irregular shallowly fissured to scaly scales Inner bark fibrous laminate Fruit calyx size (cm) 4.0-6.0 x 0.8-2 4.0-6.5 x 0.6—1.0 Max. nut size (cm) 3% 1 14x09
Specimens examined: Johore: Compartment 6B in Panti FR, Damahuri S. FRI 43567, 7 August 2002 (KEP), Ang K.C. FRI 43651, 13 August 2002 (KEP); Kluang FR, Ang K.C. FRI 43653, 14 August 2002 (KEP).
Dryobalanops beccarii in Peninsular Malaysia 3
Geographical distribution in Peninsular Malaysia
In Peninsular Malaysia, Dryobalanops beccarii is known only from Johore from three localities, namely in compartments 6A, 6B, 7, 8A and 8B, Panti FR; in compartment 132, Kluang FR; and in compartment 841 in VJR, Labis FR.
Prior to this discovery, D. beccarii was only known to occur in Borneo on leached sandy soils on coastal hills and inland ridges below 700 m (Ashton, 1982). Like D. aromatica and D. oblongifolia, it shows the same pattern of geographic distribution in southern Johore and west Borneo. Corner (1958) has drawn attention to the similarity in the floras of these two regions, which he termed the Riouw pocket.
Ecology
In Johore, this species is found in the lowland dipterocarp forest at about 75 m altitude, confined to ridges. They are large trees in the emergent and main canopies. On some ridges, it is co-dominant with Shorea curtisii Dyer ex King. In Kluang FR, it grows sympatrically with D. aromatica on steep hill slopes. In all the three sites, D. beccarii grows gregariously. In Borneo, the species is locally abundant on leached sandy soils on coastal hills and inland ridges below 700 m.
Germination Three seed batches of mature fruits were collected from five trees in Panti FR and six trees in Kluang FR in August 2002 and germinated with the seed wings removed in the FRIM nursery (Table 2). Germination is deemed to begin when the radicle protrudes through the seed coat. Germination is epigeal. The hypocotyl elongates to c. 1.8 cm. The emergent cotyledons are bilobed, fleshy and unequal. The first two leaves are opposite. In some seedlings, branching may begin at the first node.
Mean percentage germination was 73 + 12 s.d., the percentage varying from 62 to 86 (Table 1). All three seed batches showed a typical germination sigmoid curve (Fig. 1).
The germination period for Dryobalanops beccarii seed batches was longer than that of D. aromatica (100% within 20 days, Ng 1991, Siti Asha er al. 1995, Tamari 1976) and D. oblongifolia (100% within 10 days, Siti Asha et al. 1995 and 33 days, Ng 1991). In addition to the longer germination period, the three seed batches of D. beccarii achieved a maximum germination percentage of only 86.
Conservation notes
In Panti FR, ground checks in August 2002 indicated that the boundary of compartments 6B, 8A and 8B, which forms part of the forest reserve boundary, is adjacent to land being developed for small-scale agriculture activities. Because agriculture has a propensity for expansion, it is recommended that the forest office regularly monitors this section of the boundary to ensure its integrity and prevent future conflict. Compartments 6B, 8A and 8B had been logged in 1982/83. These
4 Gard. Bull. Singapore 55 (2003)
compartments are neither VJR nor are they above 1000 m elevation and hence have no legal protected status and may be licensed out for timber harvesting. Populations in Labis VJR and in Compartment 132, Kluang FR have a much brighter prospect as these forests are virgin and VJR and water catchment areas have protected status. Studies are currently being undertaken to determine the species’ population size and genetic structure.
In view of the above scenario, Dryobalanops beccarii in Peninsular Malaysia is given the 2001 IUCN category of Endangered EN A3dB1. D. beccarii falls within the A3 category (there will be a projected or suspected population size reduction of 250% within the next 10 years or three generations, whichever is the longer);’d’ (the reduction can be based on actual or potential levels of exploitation); and B1 (its geographic range, in the form of the extent of occurrence, is estimated to be less than 5000 km’).
Table 2. Percentage germination and survival in three seed batches of Dryobalanops
beccarii
Batch No. 2002-0502 (FRI 43567) 2002-0545 (FRI 43653) FRI 43651
Locality Panti FR, Compartment Kluang FR Panti FR,
6B Compartment
6B
Collector Damahuri S Ang KC Ang KC
Date planted 12 August 2002 16 August 2002 16 August 2002
No. sown 4] 617 78
Max. germination (%) 71 86 62
Germination period 4—32 4-3] 2-31
(days)
No. days to achieve 6 9 23
50 % germination
% seedling survival 22.8 72.8 100
after 1 month
Dryobalanops beccarii in Peninsular Malaysia 5
100
Germination (%) Nn S
Days
—e— 2()02-0502 —m— 2002-0545 —&— FRI 43651
Figure 1: Germination curve for three different seed
batches of Dryobalanops beccarii
Acknowledgements
We thank Hj. Ros and the foresters and rangers of the South Johor District Forest Office and Hj. Ahmad Fekri and staff of the Kluang District Forest Office for the permission and field assistance provided; Peter S. Ashton for confirming species identity; Kamarudin Saleh and Ang Khoon Cheng (FRIM) who first noted the presence of Dryobalanops beccarii and together with Damahuri Sabari, Mohd. Aidil Nordin, Mustapa Data and Ayau Kanik collected sufficient data for confirmation. Financial support from the Flora Malaysiana Centre Fund, Ministry of Primary Industries, Malaysia under Project 6: Conservation Monitoring System for Threatened Plants is gratefully acknowledged.
6 Gard. Bull. Singapore 55 (2003)
References Ashton, P.S. 1982. Dipterocarpaceae. Flora Malesiana. Ser. 1, 9: 237-552.
Corner, E.J.H. 1958. An introduction to the distribution of Ficus. Reinwardtia 4: 15-45.
IUCN, 2001. JUCN Red List Categories. Version 3.1. 1UCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK.
Ng, F.S.P. 1991. Manual of Forest Fruits, Seeds and Seedlings. Malayan Forest Records No. 34, Vol. 1. Forest Research Institute Malaysia. Kuala Lumpur. 400 PP.
Siti Asha A.B., K.C. Ang, and S. Zaiton. 1995. Records of germination trials of dipterocarp seeds at FRIM, Kepong, 1979-1994. Research Data No. 4. Forest Research Institute Malaysia. Kuala Lumpur. 161 pp.
Tamari, C. 1976. Phenology and seed storage trials of Dipterocarps. FRI Research Pamphlet No. 69. Forest Research Institute Malaysia. Kuala Lumpur. 73 pp.
Garden’s Bulletin Singapore 55 (2003) 7-12
Conservation Notes on Vatica yeechongii (Dipterocarpaceae) from Peninsular Malaysia
L.S.L. CHUA AND L.G. SAW
Forest Research Institute Malaysia Kepong 52109, Kuala Lumpur, Malaysia
Abstract
The conservation status of Vatica yeechongii L.G. Saw, recently discovered from Peninsular Malaysia, is described.
Introduction
Vatica yeechongii was discovered along Sungai Tekala in Sungai Tekala Recreational Forest, which is located in Sungai Lalang Forest Reserve, Selangor, Peninsular Malaysia, 3° 03.485’ N, 101° 52.373’ E, alt. 79 m asl (Saw, 2002). Several weeks after its discovery in the type locality, a population was found in Compartment 4, Setul Forest Reserve, along the upper stream of Setul River, Negeri Sembilan (2°46.937’ N, 101°55.069’ E). The population was located in the forest margin of a logged-over forest adjacent to the trunk road from Mantin to Seremban town at an altitude of c. 192 m.
Vatica yeechongii is very distinct in leaf characters and nut shape (Saw, 2002). The leaf blade is oblanceolate, 44-84 x 10—6.5 cm, thickly coriaceous, bullate above with 28—30 pairs of veins. The nut is ovoid.
Ecology
A medium-sized tree 8—15 m tall and 9-13 cm diameter, V. yeechongii occupies the dense understorey of lowland dipterocarp forest. In the type locality, nine trees were found on gentle earth banks of a free flowing stream near the public campsite. There were no trees further upstream. Details of its growth pattern are described in Saw (2002). Large dipterocarps such as Shorea leprosula Miq., S. parvifolia Dyer, S. macroptera Dyer, S. acuminata Dyer, S. pauciflora King, S. ovalis (Korth.) Blume, S. bracteolata Dyer, S. dasyphylla Foxw., Dipterocarpus crinitus Dyer and D. cornutus Dyer dominated the forest. Non-dipterocarps included /ntsia palembanica
4 Gard. Bull. Singapore 55 (2003)
Mig. (Leguminosae), Saraca cauliflora Baker (Leguminosae), Terminalia subspathulata King (Combretaceae), Pometia pinnata J.R. Forst. & G. Forst. (Sapindaceae), Canarium patentinervium Mig. (Burseraceae), Elateriospermum tapos Blume (Euphorbiaceae), Palaquium gutta (Hook.f.) Baill. (Sapotaceae), Pimelodendron griffithianum (Miull.Arg.) Benth. (Euphorbiaceae), Endospermum diadenum (Miq.) Airy Shaw (Euphorbiaceae), Dyera costulata (Miq.) Hook.f. (Apocynaceae), Artocarpus lanceifolius Roxb. (Moraceae), A. scortechinii King, Gironniera subaequalis Planch. (Ulmaceae), G. nervosa Planch., Payena lucida A. DC. (Sapotaceae), Streblus elongatus (Miq.) Corner (Moraceae), Litsea costalis (Nees) Kosterm. (Lauraceae), Gynotroches axillaris Blume (Rhizophoraceae), Cratoxylum formosum (Jack) Dyer (Guttiferae), Scaphium macropodum (Miq.) Beumée ex Heyne (Sterculiaceae) and Pternandra echinata Jack (Melastomataceae).
At Setul, the forest margin vegetation gradually gave way to a forest having a main canopy to c. 20 m tall, dominated by Shorea leprosula, S. macroptera, S. multiflora (Burck) Symington, S. acuminata, §. parvifolia, Parkia speciosa Hassk. (Leguminosae), Pometia pinnata, Campnosperma auriculatum (Blume) Hook.f. (Anacardiaceae), Artocarpus scortechinii, Pterocymbium tinctorium (Blanco) Merr. (Sterculiaceae) and Alstonia angustiloba Miq. (Apocynaceae). The understorey lacked palms and climbing rattans and the forest floor was covered with abundant dipterocarp seedlings. Herbs such as Tacca integrifolia Ker Gawl. (Taccaceae) and Costus speciosus (J. KOnig) Sm. (Costaceae) were present in scattered numbers. At least 70 V. yeechongii trees were found on steep slopes of riverbanks and on slopes away from the riverbanks. Trees here averaged 9.8 m tall (range 2—21 m), had a diameter at breast height c. 8.2 cm (range 1.6—18.8 cm) with a spreading crown. Its bark was whitish and smooth with horizontal rings. Those growing along the riverbanks had poor bole form and multiple branching was common while those further away had a better bole form.
Germination
Mature fruits were available from at least six trees in Sungai Tekala and one tree in Setul Forest Reserve from June to August 2002. Three seed batches, collected from two localities at different dates, were germinated with the seed wings removed and observed in the Forest Research Institute Malaysia, FRIM (Table 1). Germination is deemed to begin when the radicle protrudes through the seed coat. Germination of Vatica yeechongii is epigeal. The emergent cotyledons are bilobed, fleshy and equal. The hypocotyl elongates to c. 4.8 cm tall and the first two leaves are opposite. Mean percentage germination was 63.011 +31.207 s.d., the percentage varying from 42.1 to 98.9. All three seed batches showed a typical sigmoidal germination curve. Germination percentage on the first day of germination, i.e. day 5, was 38% for seed batch 2002-0510 and 18.7% for seed batch 2002-0503. Despite this promising
Vatica yeechongii 9
Table 1. Percentage germination and survival of three seed batches of Vatica
yeechongii
Batch No. 2002-0503 (FRI 46613) 2002-0510 2002-0542 (FRI 46668)
Locality Sungai Tekala, Sungai Sungai Tekala, Setul Forest Reserve, Lalang Forest Reserve, Sungai Lalang Negeri Sembilan Selangor Forest Reserve
Collector Chung R Saw LG Chan YC & Ayau K
Date planted 14 July 2002 28 July 2002 20 August 2002
No. sown 107 fie, 89
Max. germination (%) 42.1 48.1 98.9
Germination period 5-37 5-25 2-25
(days)
No. days to achieve 45 32 2
40% germination
% seedling survival 5.9 97.3 40
after 1 month
beginning, however, these two seed batches took a longer time to achieve the 40% germination mark compared to seed batch 2002-0542 (Fig. 1). Preliminary germination results suggest that seed viability is higher in the population at Setul Forest Reserve but this could be due to a variety of factors e.g., fruit maturity, seed vigour, timing of fruit collection, tree health, pest infestation on fruits, etc. In addition, population sizes could also affect gene flow; a larger population is probably more genetically diverse.
In situ Conservation
Vatica yeechongii is so far known only from these two localities. Preliminary observations on population size suggest that this species is a rare endemic. The population at Sungai Tekala lies in recreational forest. Under the National Forestry
10 Gard. Bull. Singapore 55 (2003)
100 80
S60
=
—
= 40
=
5
Oo 2 0
12.3.4. 5 6.7, 8,9 1011 12.13)14 15.16.17, 18,19-98 2128 Days
—o— 20()2-0542 —a— 2002-0503 —a— 2002-0510
Figure 1: Cumulative germination percentage for different seed batches of Vatica yeechongii
Act, no harvesting of resources is permitted in such forests as they are strictly meant for recreation. Population loss will only take place if the Forest Reserve or parts of the reserve, through de-gazettement, are converted to non-forestry land use. For Sungai Tekala, such action is, however, highly unlikely as the area is in the vicinity of the water catchment for the Semenyih Dam, a principal dam providing water supply to Kuala Lumpur. In terms of man-made threats to the reduction in population size and area of occupancy, there are no immediate foreseeable pressures either from habitat loss or degradation or timber harvesting. We note that the practice of clearing undergrowth in areas designated for recreational use may affect regeneration and survival of populations. Careful observa tions on the individuals here suggest that the species has no potential commercial value, as trees do not attain significant harvestable size at maturity and has a poor bole form. The Forest Department of Central Selangor District has been alerted to the presence of the new species and steps are being taken to conserve the population.
The population at Setul faces a different scenario. Although it lies in the forest reserve, it is adjacent to the main road. It has some form of protection by virtue of it being in Forest Reserve, but the fact that it lies very close to the road could pose
Vatica yeechongii ‘il
future conservation conflicts when road expansion is required. It is therefore recommended that more stringent protection measures be provided for this population. In view of the above scenario, Vatica yeechongii should be given the 2001 IUCN category of Critically Endangered CR A2cB1. It falls within the A2 category (inferred or suspected population size reduction of >80% over the last 10 years or three generations, whichever is longer); ‘c’ (a decline in area of occupancy, extent of occurrence and/or quality of habitat) and B1 (its geographic range, in the form of the extent of occurrence and area of occupancy, is estimated to be less than 100 km?). An attempt to estimate the population size of the species in each locality is currently being undertaken.
From the viewpoint of conservation, the discovery of a new dipterocarp species in Selangor and Negeri Sembilan is remarkable because in comparison to other states, these two states encounter higher conflict between socio-economic development and conservation. Both populations face an acute need for non-forestry related land development and hence there exist tremendous pressures leading to habitat loss and degradation. The fact that a new species was discovered in this current era of diminishing forest areas merely shows that we do not know enough about our plant diversity and that long-term field observations are necessary. This can only take place with concerted botanical research programmes having full national, financial and personnel support.
Ex situ Conservation
One month after potting, 215 seedlings from the three batches survived, each bearing at least two pairs of leaves. Distribution to other gardens and arboreta is envisaged in the near future.
Acknowledgements
We thank Azid Adam, Mohd. Jantan, Borhan Mat Saad and Abu Kasim Omar, Selangor District Forest Office, Cheras, Kuala Lumpur and Hj. Mohamad Hj. Ismail of the Negeri Sembilan Barat District Forest Office for the permission and field assistance provided; Chan Yee Chong who discovered the species; Damahuri Sabari, Mohd. Aidil Nordin, Markandan Moorthy and the support staff of Botany Unit, FRIM, for the field collection and ex situ maintenance of plants. Financial support from the Flora Malaysiana Centre Fund, Ministry of Primary Industries, Malaysia under Project 6: Conservation Monitoring System for Threatened Plants is gratefully acknowledged.
12 Gard. Bull. Singapore 55 (2003)
References
IUCN, 2001. JUCN Red List Categories. Version 3.1. IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge, UK.
Saw, L.G. 2002. A new species of Vatica (Dipterocarpaceae) from Peninsular Malaysia. Gardens’ Bulletin Singapore. 54: 247-251.
Garden's Bulletin Singapore 55 (2003) 13-25
Studies in Malesian Piperaceae 3!
WEE-LEK CHEW
12 Robinson Street Chatswood NSW Australia
Abstract
Thirteen species, mostly from New Guinea, are dealt with in this paper, of which 12 other species previously considered distinct are reduced to synonymy. No new taxon is proposed.
Introduction
This is part of a series of papers, albeit variously titled, resulting from continuing work initiated some 35 years ago. As mentioned previously (vide Chew, Journal Arnold Arboretum 53 (1972) 1-25), the work began by way of an undertaking to identify the collection of Piper made by the Royal Society Expedition to the Solomon Islands in 1965. This necessitated reference to the New Guinea species of Piper which then extended to an assessment of the taxonomic status of the more than 150 species described by Casimir de Candolle from that region and surrounding islands.
I was encouraged to this much larger task by the fact that the type materials from the former German New Guinea held in the Berlin Herbarium are still extant, having been removed to safety prior to the destruction of the Herbarium, and assisted further by the availability of a vast collection of high quality specimens made since about 1945.
Thus, I was enabled to determine (at the least to assess, in a few cases) the taxonomic status of some 108 of de Candolle’s species and to identify over 1,000 collections from New Guinea and the surrounding region. This resulted in the reduction of 75 Candollean names to synonyms. The assignment to synonymy of this large number of de Candolle’s species should not detract from the otherwise excellent work accomplished by this great taxonomist: he did not have the advantage of the availability of the vast range of contemporary collections that I have.
Apart from the floristic account of the alpine species by van Royen in The Alpine Flora of New Guinea 3 (1982) 1267-1287, I am not aware of any major study having been undertaken on the New Guinea species since commencement of my work.
‘Following: Blumea 20 (1972) 145-149 and Blumea 37 (1992) 159-164.
14 Gard. Bull. Singapore 55 (2003)
There still remains quite a number of Candollean species requiring determination of their taxonomic status. Additionally, large quantities of unnamed materials in many herbaria await identification, some of which may represent undescribed species. As this is a large undertaking, it may need the assistance of future botanists to continue the work.
Unless otherwise stated, all collections cited in this paper have been seen and studied by me personally.
1. Piper abbreviatum Opiz in Presl., Rel. Haenk. 1 (1828) 157; Quisumbing, Philip. J. Sci. 43.1 (1930) 58, fig. 24 & 25; Chew, J. Arn. Arb. 53 (1972) 1. Type: Philippines, Luzon: Haenke s.n. (holotype PR).
Synonyms: Piper minus K. Schum. & Lauterb., Fl. Schutzgeb., (1900) 258, syn. nov. Type: New Guinea, Ramufluss, Lauterbach 2611 (holotype B).
Piper subnudilimbum C.DC., Bot. Jahrb. 55 (1918) 205, syn. nov. Type: New Guinea, Djamu, Schlechter 17347 (holotype B, iso K).
Piper quintuplinervium C.DC., Candollea, 1 (1923) 203, 204; Candollea, 2 (1925) 224 syn. nov. Type: New Guinea, sine loc., Lauterbach 605 (isotype WRSL).
Dioecious glabrous climber. Lamina shortly petiolate, elliptic, ovate to broadly ovate, c. 10 x 5 cm, length:breadth ratio c. 2:1, glabrous on both surfaces, apex acute to acuminate, base cuneate to rounded, symmetrical; lateral veins 4 pairs, the lowermost pair very short, arising from the base, the 2nd and 3rd usually long, arising from the midrib a little above the base, the distal 4th pair very short, often absent, arising from the midrib near the apex. Stipules to 1.5 cm long, often 1 cm long, usually as long as petioles. /nflorescences shorter than leaves, peduncle c. 1.5 cm long, usually as long as petioles; males thin, to 6 cm long, bracts orbicular, peltate, subsessile; females shorter and thicker, c. 2 x 0.8 cm, bracts orbicular, peltate, sessile. Male flowers 2-staminate; pedicel very short stout, hirsute; stamens c. 1 mm long, anthers reniform to subglobose, 2-valved, slightly shorter than filaments. Female flowers sessile; stigmas 3- or 4-fid, sessile. Fruits sessile, entirely concrescent at maturity.
Distribution: Philippines, Java, Sulawesi, Maluku, New Guinea, Bismark Archipelago and Solomon Islands.
Notes: Piper minus is a smaller-leafed version of P. abbreviatum and there are no essential differences that can justify their being kept apart. Kajewski 2047 has leaves that vary greatly from the narrowly ovate c. 4 x 1.5 cm to broadly ovate c. 6 x 4 cm.
In Piper subnudilimbum, the type materials contain immature inflorescences. The leaves are large, much like those of the Philippine populations of P. abbreviatum, but the leaf bases are like the typical New Guinea plants with three pairs of lateral
Malesian Piperaceae 3 15
veins. This difference is, however, not significant and cannot be relied on to distinguish P. subnudilimbum from P. abbreviatum.
As for Piper quintuplinervium, the isotype shows two leaf forms: the narrowly ovate, small ones borne on end of free branches and the large pentagonal leaves borne at the nodes of the climbing parts of the plant. Other than small differences between P. quintuplinervium and P. abbreviatum, there is nothing significant to keep the two species apart.
Specimens examined: NEW GUINEA: Djamu Distr., Schlechter 17347; Ramufluss, Lauterbach 2611; sine loc., Lauterbach 605. SOLOMON ISLANDS: Kajewski 2047 (A).
2. Piper arfakianum C.DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 127. Type: New Guinea, [rian Jaya: Arfak Mts., Angi Lake, L.S. Gibbs 5525 (isotypes K, L).
Synonym: Piper pilosulinodum C. DC. in Gibbs, Phyt. Fl. New Guinea. (1917) 128, syn. nov. Type: New Guinea, Arfak Mts., Koebre Ridge. L.S. Gibbs 5624 (holotype BM; isotype K).
Climber, usually glabrous, rarely hairy. Leaves with 1—2 cm-long petioles. Lamina symmetrically ovate, c. 9 x 4 cm; lateral veins 3 pairs, arising from the base; apex long acuminate; base rounded, not auriculate. /nflorescence longer than leaves; peduncle longer than petiole. Bracts peltate, circular. Flowers sessile, crowded. Stigma 3-fid, reflexed, subsessile.
Distribution: New Guinea.
Notes: Piper pilosulinodum, also from the Arfak Mountains, is conspecific with P. arfakianum. Having compared the type materials of these two species and with other recent collections, | am unable to keep the species apart. Of the collections cited above, Kanehira & Hatusima 13708 is the closest match to the type material of P. arfakianum and van Royen & Sleumer 7456 is a good match to the type material of P. pilosulinodum.
Piper arfakianum is closely related to P. macropiper Pennant from which it can be distinguished by the above enumerated combination of characters, especially the petiole being much longer than in the latter species and the lamina base not auriculate.
Specimens examined: IRIAN JAYA: Arfak Mt., alt. 2200 m, Kanehira & Hatusima 13708, April (A, BO); alt. 2400 m, L.S. Gibbs 5525 (BM, K); Koebre Ridge, L.S. Gibbs 5624 (BM, K); Mt. Nettoti, alt. 1920 m, van Royen & Sleumer 7456, Nov (A,
16 Gard. Bull. Singapore 55 (2003)
BRI, CANB, L, LAE). PAPUA NEW GUINEA: Central Division, Mt. Tafa, alt. 2300 m, Brass 4047 (BRI), 5029 (BRI), 5/76 (BRI); Chimbu Distr., Mt. Wilhelm, alt. 2600 m, Borgmann 292, Aug. (LAE); Milne Bay Distr., Mt. Dayman, alt. 2200 m., Brass 22534, May (A, CANB, LAE); Morobe Distr., Edie Creek, Ridsdale NGF 30266, Nov (LAE); alt. 2000 m, Womersley & Thorne NGF 12837, Jun. (BRI, CANB, LAE); Mt. Kaindi, alt. 2200 m, Brass 29732, May (LAE); alt. 2438 m: Coode & Katik NGF 32863, Jun. (BRI, CANB, LAE); alt. 2300 m, Millar NGF 23627, Aug. (BRI, CANB, LAE); alt. 2100 m, Streimann NGF 30867, Aug (BRI, LAE, NSW); Southern Highland Distr., Anga, alt. 2000 m, Schodde 1556, Jul. (A, CANB, LAE); Spreader Divide, alt. 2255 m, Streimann & Kairo NGF 45446, Nov (CANB, NSW).
3. Piper austro-caledonicum C.DC. in DC. Prodr. 16.1 (1869) 346; Schlechter, Bot. Jahrb. 39 (1906) 92: Moore, J. Linn. Soc. Bot. 45 (1921) 381; Guillaumin, Flore analytique et synoptique de la Nouvelle-Caledonie, Phanerogames. (1948) 90. Type: sine loc., Forster s.n. (syntype: BM). sine loc., Vieillard 1227 (syntype: P).
Synonyms: Piper paitense Schltr., Bot. Jahrb. 39 (1906) 92, syn. nov. Type: New Caledonia, Mt Paita. Schlechter 14964 (isotypes L, P, WRSL).
Piper comptonii S.Moore, J. Linn. Soc. Bot. 45 (1921) 381, syn. nov. Type: New Caledonia, Mt Panie. Compton 1805 (isotype BM).
Piper peekelii C.DC., Bot. Jahrb. 57 (1922) 354, syn nov. Type: Neu-Mecklenburg, Namatanai, P. Peekel 322 (holotype B).
Glabrous climber. Leaves petiolate. Lamina of lower leaves broadly ovate to somewhat rounded to cordate, c. 9 x 7 cm, those of the upper leaves usually smaller; apex shortly blunt-acuminate; base rounded to peltate, slightly unequal, glabrous on both surfaces. /nflorescences slender, as long as the leaves, peduncle as long as the petiole: bracts pedicellate. Male flowers 3-staminate, stamens subsessile. Female flowers with 3-fid stigmas.
Distribution: New Guinea, Bismarck Archipelago, Solomon Islands and New Caledonia.
Notes: Forster f. misidentified his collection from New Caledonia as Piper siriboa Forst. f. (auct. non L.). I have seen both Forster’s and Vieillard’s collections annotated by C.DC. and am confident that the collections belong to P. austro-caledonicum. Based on examination of the type specimens, P. paitense, P. comptonii and P. peekelii are all conspecific with this species because I cannot differentiate their type specimens from specimens of P. austro-caledonicum. They are therefore reduced to synonymy. Piper austro-caledonicum is closely related to P. elbertii C.DC. of Indonesia and P.
Malesian Piperaceae 3 17
retrofractum Vahl of the Philippines. Further study is needed to assess their relationship.
Specimens examined: NEW GUINEA: Torricelli, Schlechter 14381] (BO). BISMARCK ARCHIPELAGO: New Ireland (Neu-Mecklenburg), Namatanai, P. Peekel 322 (B). SOLOMON ISLANDS: Guadalcanal, Wanderer Bay, BS/P 12247 (LAE); Kolombangara, Kokove area, BS/P 7579 (LAE); New Georgia, Kibukibu River, BSIP 4819 (LAE); Ranongga Island, BS/P 15572 (LAE); Santa Ysabel, Jejevo River, BSIP 4700 (LAE), Sigana, Brass 3451 (A, BRI), Vuranimala village, BSJP 1540] (A, LAE). NEW CALEDONIA: Aoui, Mei, Baumann-Bodenheim 10156 (P), 10176 (A, BRI, P); Bois du Mois de Mai, R. Blanche, Baumann-Bodenheim 14071 (A, BRI,P), 15179 (A, BRI, P); Col de Vulcan, Baumann-Bodenheim 8259 (A, BRI, P); Dambea, Franc 3 (P), Franc 3A (L); Ermitage stream, Compton 149 (P); Mt. Paita, alt. 300 m, Schlechter 14964, Oct 1902 (L, P, WRSL); Mt. Panie, Compton 1805, Aug 1914 (BM); Noudoure: MacKee 2670 (A, P); Yaouhe, Schlechter 14786 (P). SINE LOC: Forster s.n. (BM); Vieillard 1227 (P); Schlechter 14787 (L, WRSL).
4. Piper bosnicanum C.DC. in Gibbs, Phyt. Fl. New Guinea, (1917) 207; Candollea 1 (1923) 221, as ‘bosniakanum’. Type: New Guinea, Schouten Islands, Bosnic, L.S. Gibbs 6277 (holotype K; isotype L).
Glabrous climber. Lamina ovate, to 15 x 10 cm, with 3 pairs of lateral veins arising directly from the base. Petiole to 2 cm long. Inflorescence to 5 cm long, shorter than leaves; peduncular stalks c. 2 cm long, bracts oblong with sinuous margin. Female flowers subsessile, spirally arranged; stigmas 3-4-fid, sessile, strongly reflexed. Fruits subsessile, obovoid to ovoid, c. 10 x 6 mm broad, crowded.
Distribution: New Guinea and the Solomon Islands.
Notes: Piper bosnicanumiis very close to P. buruanum Migq. of Maluku: further study is needed to assess their relationship.
Specimens examined: IRIAN JAYA: Schouten Islands, Bisnik, L.S. Gibbs 6277, Jan (K; L); Vogelkop Peninsula, Aifact River, van Royen & Sleumer 7260, Nov (L). PAPUA NEW GUINEA: Sepik Distr., Aitepe Subdistr., along Pieni River, Darbyshire & Hoogland 8007, Jun (CANB). SOLOMON ISLANDS: Bougainville Island, north of Buin, Craven & Schodde 256, Aug (CANB, LAE); Guadalcanal, Wanderer Bay, BS/P 9104 (LAE); Kolombangara, Bambari area, BSJP 751] (LAE); Malaita, Takwa, BS/P 10785 (LAE); New Georgia Group, Roniane Lagoon, BS/P 2918 (LAE); Wagina Island, BSIP 5493 (LAE).
5. Piper lessertianum C.DC., J. Bot. 4 (1866) 164; Quisumbing, Philip. Jour. Sci.
18 Gard. Bull. Singapore 55 (2003)
43.1 (1930) 36, fig. 10, including var. oblongibaccum (C.DC.) Quisumbing. Type: Philippines, Luzon, sine loc. Cuming 1343 (‘1342’), 1841 (holotype BM).
Synonyms: Chavica lessertiana Miq., Syst. Piperac., (1843) 270 non Piper lessertianum C.DC. (1866): Piper pseudo-chavica C.DC. in DC., Prodr. 16.1 (1869) 351, nom. superfl. Type: Philippines: Luzon, Cagayan Province, Cuming 1343 (holotype G, not seen; isotype K).
Piper biroi K.Schum. & Lauterb., Nachtr. Fl. Schutzgeb., (1905) 238, syn. nov. Type: New Guinea, near Malanaku, Biro 32, Oct. (holotype B).
Piper oblongibaccum C.DC. in Leafl. Philip. Bot. 3 (1910) 777, Type: Philippines: Oriental Negros Prov., E/mer 9456 (not seen).
Piper lineatipilum C.DC., Nova Guinea Bot., 8.6 (1914) 1009, syn. nov. Type: New Guinea, Mt. Hellwig, Rémer 962 (holotype L).
Piper viridibaccum Trel., J. Arn. Arb., 9 (1928) 150, syn. nov. Type: New Guinea, Mowabula, L./J. Brass 1370 (holotype A, isotype BRI).
Climber, dioecious, glabrous to villose. Leaves shortly petiolate, petiole to 1 cm long. Lamina narrowly ovate, ovate to broadly cordate, 12—32 x 5—17 cm; base unequally rounded, one lobe acute to narrowly rounded, the other auriculate, lateral veins 34 pairs, the distal pair arising about 2 cm from the midrib, the others almost directly from the base; apex often very long acute to acuminate. /nflorescences 5—12 cm long, less than 5 mm thick; peduncle slender, 3—8 cm long, much longer than the petiole and the floriferous part of the inflorescences. Male flowers with 2 subsessile stamens, with pedicellate peltate bracts. Female flowers with 3-fid rounded sessile stigmas. Fruits oblong, to 2.5 x 1.5 mm, borne crowded but not concrescent at maturity.
Distribution: Philippines, Sulawesi, New Guinea (Irian Jaya and Papua New Guinea).
Notes: This species is attributable solely to C.DC. who published it in 1866 basing on Cuming 1342 (in herb. BM) without reference to Miquel’s earlier species Chavica lessertiana which is based on Cuming 1343 (in herb. G). I am grateful to have the advice of J.F. Veldkamp of Leiden (pers. comm., 2003) that Cuming 1342 in C.DC.’s protologue of 1866 should read Cuming 1343 because Cuming 1342 refers to a species of Gramineae. Nevertheless, the authority of Piper lessertianum (1866) remains attributable solely C.DC. without reference to Miquel.
Later, in DC. Prodr. 16.1 (1869) 351, when C.DC. combined Miquel’s Chavica lessertiana (1843) with his own Piper lessertianum (1866), he created the name Piper pseudo-chavica for the combined species. By contemporary rules, the name Piper pseudo-chavica is superfluous because his own Piper lessertianum (1866) is perfectly legitimate and should have been used instead.
Malesian Piperaceae 3 19
Our Malesian species is not to be confused with Piper lessertianum (Miq.) C.DC. in DC. Prodr. 16(1): 258 (1869) based on Ottonia lessertiana Migq., which refers to an American species and, which binomial is a later homonym of our species.
Piper lessertianum C.DC. (1866) is readily recognised by the petioles being very short and the peduncles very long, often more than five times longer than the petiole and the floriferous part of the inflorescence. Yet, it is most variable in leaf form with lamina ranging from very narrowly ovate with a long attenuate apex through ovate ob-pentagonal to broadly cordate or heart-shaped. Besides, leaves borne on the climbing parts are usually small and broadly cordate to heart-shaped while those borne on the freely hanging branches tend to be ob-pentagonal to very narrowly ovate with long attenuate apices.
All these leaf forms have now been observed to occur on the same plant as evidenced by Vink & Schram BW 8&7]. In the circumstances, I have to assign to synonymy Piper biroi, Piper lineatipilum and Piper viridibaccum as they are distinguished purely on these leaf forms. For the same reason and after examination of collections authenticated by Quisumbing, I am unable to maintain var. oblongibaccum (C.DC.) Quisumbing.
Specimens examined: PHILIPPINES: Luzon, Cagayan Prov., Cuming 1343 (BM, K); Mindoro, Mt. Halcon, Ramos & Edano 40676, Mar (NSW); Panay, Capiz Prov., Mt. Bulilao, Martelino & Edano 35706, Jun (NSW); Jaminden, Ramos & Edano 30861, 31272, May-Apr (NSW); Libacao, Martelino & Edano 35395 (NSW), 35442, May- Jun (NSW); Mt. Macosolon, Ramos & Edano 30776, Apr-May (NSW). SULAWESI: Kabaena Island, Elbert 3393 (BO). IRIAN JAYA. Wati, Mapon Distr., /jiri & Niimura 674, Apr (L); Wissel Lakes, Doglia, Vink & Schram BW 8&71, May (CANB, LAE). PAPUA NEW GUINEA: Milne Bay, near Mayu Island, Streimann NGF 28829, Apr (L, NSW): Morobe Distr., Asubazo, Kairo NGF 24403, Jul (L); Northern Division, Sibium Range, R. Pullen 5924, Sept (CANB).
6. Piper longipilum C.DC., Bot. Jahrb. 55: 216 (1918); Candollea 1: 206 (1923). Type: New Guinea, Sepik, alt. 1-200 m, Ledermann 7565 (holotype B, isotype B).
Climber, dioecious, lightly pilose. Leaves shortly petiolate, petiole to 2.5 cm long. Lamina ovate, c. 28 x 18 cm; base unequally cordate, lateral veins 5—6 pairs, the basal pair arising c. 1 cm from the base; apex acuminate. Male inflorescences (immature) c. 8 cm long, c. 3 mm thick; peduncle slender, c. 1.7 cm long. Male flowers with pedicellate peltate glabrous bracts. Female unknown.
Distribution: New Guinea.
Notes: The type material of this species consists of two sheets: the one with field
20 Gard. Bull. Singapore 55 (2003)
notes and annotation by C.DC in 1917 is the holotype, the other is the isotype. A cursory examination of the type material suggests an alliance with Piper decumanum L. of which it might represent a hairy variant. As the type seems to be the only material there is of this species, I am unable to determine its status until I see more collections, especially of female plants.
7. Piper melula Trel., J. Arn. Arb. 9 (1928) 148. Type: Papua New Guinea, Vaitata River, L.J. Brass 1130 (holotype A; isotype BRI).
Dioecious glabrous climbers. Leaves moderately petiolate, petioles 2-4 cm long, much longer than peduncles. Lamina irregularly ovate, broadly ovate to pentagonal, c. 12 x 8 cm; apex bluntly shortly acuminate; base rounded to shallowly cordate; lateral veins 3 pairs, the distal pair arising alternately from the midrib about 0.5—1 cm from the base, the other 2 pairs directly from the base; only the distal pair reaching to the apex, the 2" pair reaching to 1/2 the length of the lamina, the basal pair only to 1/3 of the length of the lamina. /nflorescences shorter than the leaves, males slightly longer than females. Male floriferous part to 6 cm long on a peduncle to 7 mm long; female floriferous part to 4 cm long on a peduncle to | cm long. Bracts circular, margin sinuate, peltate, very shortly pedicellate. Male flowers with strongly reniform anthers, dehiscing by long apical slits. Female flowers sessile, crowded; stigmas 3- to 4-fid, sessile, stigmatous part reflexed. Fruits almost entirely concrescent, free only at the apices.
Distribution: Papua New Guinea.
Notes: Piper melula has the same leaf venation pattern as P. plagiophyllum K.Sch & Laut., and in some cases even the lamina shape of that species, but it differs in being a climber amongst other characteristics.
The infructescences of Piper melula appear similar to those of P. pachyarthrum K.Sch., but these species differ in other respects.
Piper melula appears also to be related to P. elbertii C.DC. of the Lesser Sunda Islands and P. austro-caledonicum of the Solomon Islands and New Caledonia.
There is a collection, Kellman ANU 1597 (CANB) from Mindanao in the
Philippines, which is remarkably similar to Piper melula. However, as this collection has a very young inflorescence, I hesitate to identify it as this species.
Specimens examined: PAPUA: Gulf distr., Tauri River, Schodde & Craven 4700, Mar (CANB); Vailala River, L.J. Brass 1130, Mar (A, BRI); Milne Bay distr., Kwagira River, Peria Creek, L.J. Brass 23997 (A, CANB, LAE), 24006, Aug (A, CANB, LAE).
Malesian Piperaceae 3 a |
8. Piper novo-guineense Warb., Bot. Jahrb., 13 (1891) 284, K.Schumann & Lauterbach, Fl. Schutzgeb., (1900) 258. Type: New Guinea, Sattelberg bei Finnschlafen, Warburg 20740 (holotype B; isotype A).
Glabrous climber. Lamina narrowly elliptic, narrowly ovate to broadly ovate or broadly pentagonal, 17—19 x 5—9 cm; apex shortly blunt acuminate to long acute; base cuneate to rounded, very slightly asymmetrical, not auriculate; lateral veins 3 pairs, the distal pair arising slightly alternately from the midrib c. 1—2 cm from the base, the other 2 pairs arising directly from the base, sometimes another faint short pair arising from the base; the veins on the underside, especially at the basal portion, often covered with minute reddish brown scales, petiole to 1 cm long. Male inflorescences thin, as long as the females, peduncle as long as the males. Male flowers with bracts as in the females; stamens subsessile with broadly crescent-shaped anthers. Female inflorescences to as long as the leaves, sometimes longer, usually to 15 cm long; peduncular stalk c. 2 cm long. Female flowers sessile, stigma 3-4-fid. Fruits sessile, oblong, crowded; bracts rounded peltate.
Distribution: New Guinea.
Notes: Piper novo-guineense is reported to be common in primary forest and older secondary forests, between 500 and 2000 m altitude. At higher altitudes, the plants tend to have smaller leaves.
This species is closely related to Piper macropiper Pennant and especially to P. truncatibaccum C.DC.
Specimens examined: IRIAN JAYA: Lake Habbema, L.J. Brass 10299, Oct (A). PAPUA NEW GUINEA: Morobe Distr., Boana Mt. Clemens 8467 (A); Gurokar, L.J/. Brass 29499, May (CANB, LAE); Huon Peninsula, Mt. Rawlingson, Hoogland 9054, Jun (A, BRI, LAE); Sattelberg, Warburg 20740 (B, A); Wantoat area, Kikiepa, Womersley & Thorne NGF 11875, Jun (BRI, LAE); Selileo, Hellwig 580, Apr (B); Sepik, near Kilifas village, Forman & Kumul NGF 48261, Mar (LAE).
9. Piper pallidilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1009. Type: New Guinea, Mt. Hellwig, alt. 2600 m, von Romer 1316 (holotype L).
Synonym: Piper peracutilimbum C.DC., Nova Guinea Bot. 8.6 (1914) 1005, syn.nov. Type: New Guinea, Mt. Hellwig alt. 2600 m, von Rémer 1252 (holotype L).
Climber, glabrous. Leaves shortly petiolate. Lamina ovate, 6-8 x 24 cm; base rounded, apex acuminate. Lateral veins 2-3 pairs, distal pair arising from the midrib a little
22 Gard. Bull. Singapore 55 (2003)
above the base. /nflorescence shorter than leaves. Peduncular stalk 7-10 mm long, about as long as petiole. Males up to 4 cm long, females c. 2 cm long. Flowers borne crowded, sessile. /nfructescence congested, to 12 mm diam. as in Piper abbreviatum Opiz.
Distribution: New Guinea.
Notes: The type material of Piper peracutilimbum is indistinguishable from specimens of P. pallidilimbum. Therefore I see no reason to keep the two species apart.
Piper pallidilimbum is very close to P. abbreviatum Opiz from which it differs in the lamina being furnished with fewer lateral veins, one of which issues from the midrib, and also by the peduncular stalk being somewhat filamentous.
Specimens examined: IRIAN JAYA: Mt. Hellwig, alt. 2600 m, von Romer 1316 (L), 1252 (L); sine loc., Pulle 700 (L), 737 (L), 834 (L).
10. Piper pseudamboinense C.DC., Bot. Jahrb. 55 (1918) 206. Type: New Guinea, Keneyia, alt. 150 m, Schlechter 15445 (holotype B).
Climber. Leaves very shortly petiolate. Lamina pentagonal to oblong, commonly c. 24 x 11 cm; apex bluntly acuminate, acumen to 1.5 cm long, base asymmetrically cordate, one side often forming a large lobe, the lobes often covering the node; lateral veins usually 5 pairs, the distal pair alternate, arising from the midrib c. 2—3 cm from the base, the 2° pair often arising also from the midrib but very close to the base; the other 3 pairs, often very short, issuing directly from the base; intercostals very prominent; upperside of lamina glabrous, undersides with short brown pubescence on the veins. Female inflorescences to as long as leaves, 20—24 cm long; the floriferous portion as long or to only slightly longer than the peduncular stalk; peduncle glabrous. Bracts circular, peltate, long pedicellate. Female flowers sessile, congested. Stigma 2-lipped, spreading, subsessile. Fruits sessile, free.
Distribution: New Guinea.
Notes: A lowland climber, Piper pseudamboinense 1s a little known species. Of the two collections identified as this species, NGF 25678 appears to be the best match to the holotype, the other being a more hairy variant.
Piper pseudamboinense is very closely similar to P. amboinense (Miq.) C.DC. and from which it differs in the more prominent lateral veins and its stigma being 2-lipped. It is also closely similar to P. lessertianum C.DC. from which it can be distinguished by the 2-lipped stigma in addition to lamina and inflorescence characters.
Malesian Piperaceae 3 23
Specimens examined: PAPUA NEW GUINEA: Morobe District, Buso River, NGF 25678 Aug (CANB, LAE); Oomsis Creek, T.G. Hartley TGH 10564, July (A, CANB, LAE); Keneyia, alt 150 m, Schlechter 18445 (B).
11. Piper reinwardtianum (Miq.) C.DC., DC. Prodr. 16.1 (1869) 354. Merrill, Inter. Rumph. Herb. Amb., (1917) 182. Basionym. Macropiper reinwardtianum Miq.., Linnaea 21 (1848) 481. Type: Indonesia, Maluku, Nussa Laut, Reinwardt, July 1821 (holotype L?).
Synonym: Chavica reinwardtiana (Migq.) Miq., Ann. Mus. Bot. Ludg. Bat.1 (1863) 136.
Dioecious shrub, glabrous to lightly pilose. Leaves with petiole 2.5—5 cm long. Lamina broad ovate to rounded, c. 16 x 15 cm, slightly asymmetrical; apex shortly acute, base asymmetrically truncate-cordate, lateral veins usually 34 pairs, upperside of lamina glabrous, undersides lightly pilose on the veins. /nfructescences to 11 cm long, peduncle glabrous to 2.5 cm long. Female flowers sessile, congested. Stigma 3, rarely 4, short thickset. Fruits sessile, somewhat obovoid, concrescent on the lower half.
Distribution: Indonesia, Maluku.
Notes: I have not seen any type material of this species which is probably in the Leiden Herbarium to which Reinwardt donated his collections from the region. The collections of Robinson and Teysmann cited herewith are all that I have seen of this species; and I must of neccesity rely on C. de Candolle and Merrill in the interpretation of this apparently rare species.
In general growth habit, Piper reinwardtianum recalls that of P. sundaicum Blume, also a shrub from the same region. However, it has infructescences similar to P. mestonii Bailey of New Guinea and Australia from which it can be distinguished by the stigma being 3- or 4-fid, not bifid, and the plant a shrub, not a climber. In these respects, P. reinwardtianum appears to be an interesting species, and further investigation may yield insight into the phytogeograpical relationship between the three species.
Specimens examined: INDONESIA: Maluku, Ambon, Robinson 60 (L). Ceram,
Teysmann s.n. (L).
12. Piper subvirosum C.DC., Bot. Jahrb. 55 (1918) 215. Type: New Guinea, Alexishafen, Wiesenthal 68 (holotype B).
24 Gard. Bull. Singapore 55 (2003)
Climber, twigs minutely pubescent. Lamina broadly ovate, 15—20 x 10-12 cm, upper surface glabrous, lower surface sparsely covered with short stiff hairs especially on lateral veins on basal part of the lamina; apex shortly acuminate, acumen 1.5—2 cm long; base broadly cuneate, rounded to truncate; lateral veins 3 pairs, distal pair alternate, arising from midrib a little above the base, the others directly from the base; intercostals prominent. Petiole to 2 cm long. Female inflorescence to 27 cm long, peduncle to 3 cm long, floriferous part to 24 cm long; inflorescences usually longer than leaves, peduncle longer than petiole; mature inflorescence to 7 mm diam. Female flowers sessile, not concrescent, with 3- or 4-fid stigmas.
Distribution: New Guinea.
Notes: Piper subvirosum is related to P. macropiper Pennant from which it can be distinguished by the much larger leaves and inflorescences, at least one pair of the lateral veins arising from the midrib, the lamina base not auriculate, and the peduncle not excessively longer than the petiole.
Not much is known about this species. The two recent collections cited here are the closest match to the holotype.
Specimens examined: IRIAN JAYA: Star Mts., Sibil Valley, alt. 1200-1500 m, Kalkman & Nicholas 4160, May (L). PAPUA NEW GUINEA: Central Division, Mt. Tafa, L.J. Brass 4127 (A); Alexishafen, Wiesenthal 68 (B).
13. Piper sundaicum Blume, Verh. Bat. Genoots. 11 (1826) 187, fig. 14; C.DC., DC. Prodr. 16.1 (1869) 352; Koorders, Verh. Kon. Ak. Wet. Amsterdams. 2, 14.4 (1908) 42, reprint; Koorders, Exkursionsflora van Java. 2 (1912) 26; Koorders- Schumacher, Exkursionsflora Atlas. 4.2 (1924) 445, fig. 718G. Type: Maluku, Ambon. Blume s.n., Leiden Herb. 908.363-43 (lectotype L, here designated).
Synonyms: Chavica sundaica (Blume) Mig. Syst. Piperac., (1843) 274; Fl.Ind. Bat 1(2) (1859) 446, exclusive of the synonym P. a/bum Vahl.
Piper amplilimbum C.DC., Candollea 1 (1923) 213; Schroeder, Candollea 3 (1926) 135; Chew, Blumea. 20 (1972) 145, fig. 1, syn nov. Type: Maluku, Ambon, Teysmann s.n. 1905 (holotype BO).
Dioecious scandent shrubs, entirely glabrous. Twigs 5—8 mm diam.; internodes smooth; stipular scars prominent. Lamina symmetrically ovate, (23—)25—27 x 14-19 cm, somewhat coriaceous; apex acuminate, acumen to 2 cm long; base rounded; margin entire; lateral veins 34 pairs, very prominent, the distal pair arising alternately from the midrib at 2-3 cm from the base, the others directly from the base; intercostals
Malesian Piperaceae 3 25
numerous, not straight, quite prominent. Petioles c. 2 cm long, smooth. Stipules to 8 x c. 1 cm, apparently not adnate to petioles. /nflorescences: male long, thin; female 11-13 x 0.7 cm, the peduncular stalk c. 1.5 cm long, glabrous. Bracts somewhat irregularly circular, peltate. Female flowers sessile; ovary pyramidal; stigmas 3-fid, short, sessile, slightly reflexed. Fruits ovoid, sessile, c. 3 cm long: crowded, not concrescent.
Distribution: Indonesia, Maluku.
Notes: Blume recorded this species as also occurring in Java. Koorders (1908, 1912) doubted the Javanese record as he failed to find any collection from that island. I have also been unsuccessful in finding any material of the species from Java. This is probably a very rare species: the type materials are all that is known of this species.
I have compared the type materials of Piper sundaicum with that of P. amplilimbum C.DC., and I am unable to distinguish the one from the other.
Piper sundaicum appears to have some relationship with Piper reinwardtianum from the same region (see above).
Piper sundaicum has been identified with P. album Vahl by previous botanists. The two species are however, quite distinct: P. album Vahl is a climber, our species is not.
Specimens examined: MALUKU: Ambon, Blume s.n. (L): Teysmann s.n. 1905 (BO).
Acknowledgements
I thank the curators of the following herbaria (identified by their acronyms) for loan of their collections, including type materials, for my study at the Herbarium of the Royal Botanic Gardens, Sydney where I held tenure as Senior Research Scientist from 1970 to 1975: A, B, BO, BRI, CANB, K, L, LAE, P, PR, PRC and WRSL.
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Garden’s Bulletin Singapore 55 (2003) 27-30
A New Species of Schismatoglottis (Araceae) from Sabah, Malaysia
A. HAY AND C. HERSCOVITCH Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia
Abstract
Schismatoglottis tahubangensis A. Hay & Herscovitch is described, illustrated and inserted into the latest key to Bornean Schismatoglottis species.
Schismatoglottis tahubangensis A. Hay & Herscovitch, sp. nov.
A Schismatoglottis platystigma M. Hotta spadice tenuiore, staminodiis inter pistilla, connectivo longiore differt. - Typus: Malaysia, Sabah, just outside Kinabalu Park, 100 m to Tahubang River along route to Marai-Parai, 11,000 m alt. [sic], 12 Sep 1993, J. Nais, Dolois, Harry & Masius SP 05312 (KEP, holo).
Terrestrial herb c. 30 cm high. Stem creeping or decumbent, rooting between the nodes, c. 1 cm diam., with internodes c. 5 mm long. Leaves 6 together; petiole 10-17 cm, sheathing in the lower 1—2 cm, wings of sheath extending into a papery narrowly triangular ligule to 2.5 cm; blade 11—15 cm long, 3.5—5 cm wide, narrowly obovate, somewhat asymmetric, base truncate to acute, somewhat asymmetric, inserted into adaxial face of petiole, apex acute; midrib pronounced and minutely puberulent abaxially, with c. 10 primary lateral veins on each side diverging at c. 45°, interprimary veins somewhat conspicuous abaxially, obscure adaxially, fine. /nflorescence solitary, subtended by an inconspicuous papery cataphyll c. 6 mm long; peduncle 3 cm long. Spathe pinkish, 7 cm long, slender, weakly constricted; lower spathe 1.5 cm long, 3mm diam. Spadix very slender, cylindric for most of its length, 5.3 cm long, 1.5 mm diam.; female zone 1.2 cm long, (not clear whether adnate to spathe at base); pistils closely packed, subglobose, 0.3 mm diameter; stigma discoid, almost as wide as the ovary; interpistillar staminodes conspicuous, round and flat-topped or concave, drying pale yellow, of the same height and diameter as pistils; sterile interstice c. 2 mm long comprised proximally of 2 irregular whorls of looser packed, 0.5 mm diam., flat-topped staminodes, differentiated from interpistillar staminodes by larger size, slightly convex top, and dark yellow colour on drying, and distally of 2—3 irregular whorls of sterile
28 Gard. Bull. Singapore 55 (2003)
stamens drying dark brown; male zone 1.2 cm long; stamens subsessile, more or less oblong ovate from above, more or less irregularly paired and oriented along long axis of spadix, c. 0.8 mm wide; thecae round, opening by a single apical pore; connective robust, columnar, flat-topped and somewhat impressed, slightly exceeding the thecae in height and about 1.5 times their diameter, rounded triangular from above with ends indented above thecae; appendix 2.8 cm long, tapering to a fine naked tip, composed of flat-topped, more or less rounded or irregularly polygonal staminodes c. 0.3 mm diam. similar to those of sterile interstice. Fruit unknown.
Distribution: Malesia — Borneo, Sabah, Kinabalu (known only from the type).
Habitat: Ona steep slope in unspecified vegetation type at c. 1,100 m alt. [Note that the label on the type records the altitude as 11,000 m, which 1s, of course, impossible].
Notes: This species somewhat resembles Schismatoglottis platystigma (Brunei) in having discoid stigmas and relatively small thecae on the edge of a broad connective. However, it is readily distinguished from that species by the presence of interpistillar staminodes, the subsessile stamens with a much higher connective and the more slender spadix.
Schismatoglottis tahubangensis may be inserted into the key to Bornean Schismatoglottis provided by Hay & Yuzammi (2000) as follows:
17a. Inflorescence solitary or two together; anthers with broadly dilated connective
Fe ee EO RES re ree ree Teeter en ee 18 17b. Inflorescences c. 4-6 together; anthers dumb-bell-shaped, with the connective a i a a ee a ERE NITED
18a. Interpistillar staminodes absent; appendix 1—2 cm long, blunt-tipped; Temburong, BOE? 2A ANE. RU TER G.. RUA ASS ORAS Se
18b. Interpistillar staminodes numerous; appendix c. 3 cm long, tapering to a fine CIN, PA MNANAIEL, DOIN. os os nog om eee i rmadtaesapasenn esata as <email S. tahubangensis
19 etc = 18 etc in Hay & Yuzammi (2000).
Schismatoglottis tahubangensis 29
Figure 1. Schismatoglottis tahubangensis Herscovitch & A. Hay
A. habit; B. ligule at base of petiole; C. detail of leaf venation; D. midrib detail; E. inflorescence with part of spathe removed: F. pistils with interpistillar staminodes; G. sterile interstice with staminodes and
sterile stamens; H. stamens; I. staminodes of appendix. J. Nais, Dolois, Harry & Masius SP 05312.
30 Gard. Bull. Singapore 55 (2003)
Acknowledgements
We thank Josef Bogner for drawing our attention to the specimen on which this species is based, Dan Nicolson for commenting on the ms and and Lesley Elkan for the illustration.
Reference
Hay, A. & Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I — Schismatoglottis. Telopea. 9: 1-177.
Garden’s Bulletin Singapore 55 (2003) 31-34
An Unusual New Species of Homalomena (Araceae) from New Guinea
C. HERSCOVITCH AND A. HAY
Royal Botanic Gardens, Mrs Macquarie’s Road, Sydney, NSW 2000, Australia
Abstract
Homalomena impudica Herscovitch & A. Hay is described, illustrated and inserted into the latest key to Homalomena species of New Guinea.
Homalomena Schott was revised for New Guinea by Hay (1999). An overlooked specimen on loan to NSW from LAE represents a very distinctive undescribed species.
Homalomena impudica Herscovitch & A. Hay sp. nov.
Homalomena tenuispadix Engl. simulans sed inflorescentia femina spatham aequanti, inflorescentia mascula clavata tota exserta differt. - Typus: Papua New Guinea, West Sepik Province, Vanimo, Ossima, 28 Jan 1969, H. Streimann & A. Kairo NGF 39239 (holo, LAE).
Erect multi-stemmed herb to c. 30 cm high. Stem condensed, 1.5 cm diam. Leaves to 8 together; petiole longer than blade, 17—32 cm long, glabrous, slender, sheathing in lower 6—7 cm; blade dull green above, green below, glabrous, sagittate to ovate- cordate,14-19 cm long, anterior lobe 12—16.5 cm, c. 8 cm wide, posterior lobes triangular to rounded, 2.5—4 cm long; midrib more prominent abaxially, with c. 8 pairs of lateral veins diverging at c. 45 °. Inflorescences to 9 together, each subtended by a narrow triangular papery cataphyll to 4 cm long; peduncle erect, later declinate, 4.5—6 cm long, longer than spadix. Spathe slightly constricted in bud, then narrow cylindrical, 2.0—2.8 cm long, 5 mm diam., with a conspicuous rigid darker sometimes reflexed apiculum 2—3 mm long. Spadix 2.7—3.6 cm long, much exceeding the spathe at anthesis, stipitate for c. 5 mm; female zone equalling to exceeding the spathe, 2.0—2.4 cm long; pistils very widely scattered along axis of female zone, becoming loosely and irregularly whorled distally, globose, c. 1 mm diam., infrapistillar staminodes absent; stigma button-like, sessile, 0.3 mm across; male zone narrow oblong-ovoid, 0.7—1.1 cm
32 Gard. Bull. Singapore 55 (2003)
long, whitish, fully exserted from spathe; male flowers (-)2-merous; stamens more- or-less oblong to irregularly polygonal from above, c. 1 x 0.3-0.5 mm, the thecae overtopped by connective. Fruit unknown.
Distribution: Malesia— New Guinea, Papua New Guinea, West Sepik Province (known only from the type locality).
Habitat: In a dry stream bed in lowland rain forest at c. 30 m alt.
Notes: This species resembles Homalomena tenuispadix Engl., but differs in the greatly elongated and lax female zone which equals to exceeds the spathe, and in the completely exserted and somewhat clavate male zone. The inflorescence at first resembles that of Schismatoglottis after the male zone has been exposed by the fall of the spathe limb, until it is noted that the spathe, as typical for Homalomena, 1s fully persistent.
While some New Guinea species of Homalomena, such as H. davidiana A. Hay and others (see Hay 1999), characteristically have the male zone partially exserted ~ from the spathe after anthesis, this species extends the entire male zone beyond the apex of the spathe prior to anthesis. This characteristic, as distinct from the state where the male zone is exposed by the spathe being wide open ms anthesis, appears not to be recorded for any other species in the genus.
Field notes describe the flower (sic) as white and the fruit green. The specimen includes fragments of declinate fruiting peduncle, but the infructescence is not present. It is unclear whether or not the colour observations refer to the spathe at different stages.
The specific epithet, meaning lewd or shameless, alludes to the fully exposed male zone of the spadix.
Homalomena impudica can be inserted into the key provided by Hay (1999) as follows:
10a. Spathe tip + hood-forming; spadix contained within the spathe after anthesis; peduncle tice Sorter tian tite PetiOles .2-. tcc cce cee ree ececee se H. lauterbachii
10b. Spathe with the tip reflexed; spadix exserted from the spadix during and/or after anthesis; peduncle c, 1/3—1/2 the leneth of (he petioles |. na beriecekes Gannann een 11
lla. Spadix with the male zone clavate and completely exserted from the spathe at anthesis). A SS a ee eae See H. impudica
1 1b. Spadix with the male zone cylindric-tapering, not exceeding the spathe at anthesis, partially exserted from spathe after anthesis ...................ssseeeseeeH. Cenuispadix
12 etc. = 11 etc. in Hay (1999)
Homalomena impudica 33
Figure 1. Homalomena impudica Herscovitch & A. Hay. A. habit; B. inflorescence with part of spathe removed; C. tip of spathe; D. inflorescence in bud; E. pistils; F. stamens. Streimann & Kairo NGF 39239.
34 Gard. Bull. Singapore 55 (2003)
Acknowledgements
We thank LAE for the loan of the specimen on which this species is based and Catherine Wardrop for the illustration.
Reference
Hay, A. 1999. Revision of Homalomena (Araceae-Homalomeneae) in New Guinea, the Bismarck Archipelago and Solomon Islands. Blumea. 44: 41-71.
Garden’s Bulletin Singapore 55 (2003) 35-60
Pleuroschisma, a New Section of Cyrtandra (Gesneriaceae) from Borneo
O.M. HILLIARD’, B.L. BURTT! AND M.H. BOKHARI? ‘Royal Botanic Garden Edinburgh, EH3 5LR, U.K.
*Multan, Pakistan
Abstract
Twelve Bornean species of Cyrtandra constitute a new section, Section Pleuroschisma, the name derived from an important diagnostic character, namely the unique fruit with a median septicidal split on each side (not reaching either apex or base of the fruit). Nine of the species and two varieties are newly described, C. angustielliptica Hilliard & B.L.Burtt, C. coacta Hilliard & B.L.Burtt, C. insolita Hilliard & B.L.Burtt, C. linauana B.L.Burtt, C. seganica Hilliard & B.L.Burtt, C. tesselata Hilliard & B.L.Burtt, C. tunohica Hilliard & B.L.Burtt, C. pendulifera Kraenzl. var. grossipilosa Hilliard & B.L.Burtt and C. sarawakensis C.B.Clarke var. Jongipilosa Hilliard & B.L.Burtt. Two species recognized remain nameless because the material is inadequate to typify a name.
Introduction
There are already 40 sectional names in Cyrtandra and the limits and interrelationships of the groups so designated are far from being understood; the addition of yet another sectional name to the list may seem somewhat irresponsible. Nevertheless, the fruit characters that are used to diagnose this section - the smooth fruit wall (due to a thin layer of sclerenchyma beneath the epidermis, Fig. 3), and the septicidal splits with inrolled margins (Fig. 1) form a unique combination in this small group. It is the lateral splits in the fruit-wall that form the basis for the sectional name Pleuroschisma; these splits, it should be noted, are septicidal and there is some reason to suppose that septicidal dehiscence may have been the primitive condition in the family as it is found in Ramonda, Haberlea, in the type species of Corallodiscus, all on the northern fringe of the subfamily Cyrtandroideae, and in the tribe Coronantherae, a basal tribe of subfamily Gesnerioideae found in New Caledonia, E. Australia, New Zealand and southernmost S. America. Its presence elsewhere in the family (e.g. in Leptoboea) may be secondary. It must be noted, however, that the lateral splits in this new section of Cyrtandra do not reach from apex to base of the fruit (Fig. 1) and do not give rise to free fruit-valves, as in a fully septicidal fruit. These features set section smaller.
36 Gard. Bull. Singapore 55 (2003)
Figure 1. Mature fruit of Cyrtandra sarawakensis The two halves remain held together at apex and base. Burtt B4¢71]2 x4.
Pleuroschisma apart from all the other sections.
As so often in herbarium work on Cyrtandra, we have been much hampered by lack of flowers on many specimens, although mentioned in the collector’s field notes. Nevertheless, it is not the flowers that provide the diagnostic features of the Section: it is the fruits. We have therefore felt justified in naming C. tesselata (no. 6), C. seganica (no. 7) and C. insolita (no. 11), which we know only in fruit. In the eight species where we have seen flowers, the major floral characters are uniform: calyx divided almost to the base into five deltoid lobes, corolla c. 35-40 mm long, white, cream or palest yellow with two orange/yellow bars in the throat, puberulous outside, disc cupular, occasionally deeply excavated on one side, ovary crowned with a coma of relatively long hairs, otherwise so minutely gland-dotted or pustulate as to appear glabrous on casual inspection, style puberulous, stigma distinctly bilobed, lobes relatively large, clavate in outline.
Leaf-venation provides a useful character in distinguishing some of the species, both the number of lateral veins on each side of the midrib, and the pattern described by the tertiary venation, of which three main types have been distinguished; for these we use the terms subscalariform, reticulate and pinnate (Fig. 2, A,B,C,D). In some species, the tertiary venation is invisible (e.g. C. insolita).
Cyrtandra sect. Pleuroschisma af
Figure 2. Types of tertiary venation found in Cyrtandra sect. Pleuroschisma A. subscalariform (C. sarawakensis, Burtt B1947, about actual size). B. finely reticulate (C. tesselata,
Hotta 1417, slightly enlarged). C. coarsely reticulate (C. sp. nov. no. 12, Burtt B2658, x2). D. pinnate (C. coacta, Beaman 10656, slightly enlarged).
38 Gard. Bull. Singapore 55 (2003)
The distinctive leaf anatomy, with tracheoids in the hypodermis, has already been illustrated (Bokhari & Burtt, 1970, Plate 2, G,H): the tracheoids may have either parallel or reticulate bars of thickening (Fig. 4). In some species (e.g. C. hoseana), similar tracheoids are found in the mesocarp of the fruit; spheroidal sclereids are commonplace and abundant (Fig. 3), making fruit-sectioning very difficult. In several species, polymorphic sclereids occur in the mesophyll of the leaf (C. tesselata, C. seganica, C. hoseana, C. angustielliptica); in C. penduliflora, they may be present or absent. The other species in section Pleuroschisma lack sclereids. (C. insolita and sp. no. 12 have not been examined).
Protection of the apical bud in Cyrtandra is achieved in several ways. In section Pleuroschisma the bud is hidden between the erect petioles of the two fully developed uppermost major leaves on the stem. The following observations were made on herbarium material, and observations are needed on living plants to determine time-intervals in leaf development: when the developing leaf in C. /inauana measured 38 mm, the young apical bud was only 2.5 mm long. We examined the stem apex of C. linauana, removing the tip of the stem and rehydrating it. The very young major leaf of the apical bud is hidden by the conduplicate base of the minor (stipule-like) leaf of the uppermost pair of fully developed leaves. It is plane, measured 38 x 8 mm including the petiole (scarcely developed at this stage), and both surfaces are thickly clad in long silky hairs. The minor leaf has similar indumentum and 1s well developed, being 20 mm long. In contrast, the fully developed leaves below the apex show no trace of the juvenile indumentum except over the midrib and lateral veins on the lower surface. There is the same abrupt transition from a very small juvenile leaf to a fully developed one in five other species where we had stem-apices suitable for inspection (C. penduliflora, C. sarawakensis, C. coacta, C. angustielliptica, C. insolita). These species in section Pleuroschisma are all strongly anisophyllous. A comparable condition in isophyllous species was noted by Burtt (2001, p.399).
Field observations by one of us (B.L.B.) found that in C. hoseana and C. penduliflora, as the new apical leaf begins to grow out, the petiole is held erect but the young lamina is pendulous, the hairs in C. hoseana being silvery white, while in C. penduliflora they are dark red. As the leaf blades enlarge, the hairs become more spread out, and are eventually shed, resulting in the blade of the mature leaf being glabrous above. Whether or not these young leaves perform any attractive function is not known, but they certainly catch the eye of the plant-hunter.
Cyrtandra section Pleuroschisma Hilliard & B.L.Burtt sect. nov. ab sectionibus omnibus ad huc descriptis fructibus duris laevibus (strato annulari subepidermali schlerenchymatis uni- vel bi-seriato), parietibus in medio longitudinis fissuris duabus septicidalibus marginibus involutis notatis, foliis cellulis tracheoidalibus hypodermalibus praeditis facile distinguenda.
Cyrtandra sect. Pleuroschisma
WN |
|
39
Ay 4 4 ie!
=
~ i a> Ss
——
——— ~~ — ~
Figure 3. Transverse sections of part of fruit wall and septum
A. Cyrtandra sarawakensis; B. Cyrtandra penduliflora. b. scl. = brachysclereid; fib. 1. = fibrous layer; m. scl. = macrosclereid; scl. p. = sclerified parenchyma [sclerenchyma]; sp. scl. = spheroidal sclereid; tr. = tracheoid [with reticulate thickening]. M.H. Bokhari del.
Figure 4. Sclereid and tracheoid types. A. polymorphic sclereids. B. tracheoid with reticulate bars. C. tracheoid with parallel bars. M. Mendum after pencil sketches by M.H. Bokhari.
40 Gard. Bull. Singapore 55 (2003)
Species typica: C. sarawakensis C.B.Clarke Distribution: Endemic to Borneo.
Simple-stemmed perennial herbs (C. insolita bushy), leaves usually strongly anisophyllous, tracheoids, as far as is known, in hypodermis, inflorescence a dichasial cyme, bracts free, bracteoles present or absent, calyx divided almost to base, corolla medium-sized, white to palest yellow ground colour, disc cupular, ovary crowned with a coma of relatively long hairs, style pubescent, stigmatic lobes large, clavate, fruit with two septicidal grooves (later splits).
Key to Species la. Lateral veins in largest mature leaves up to 10 each side of midrib ............... 2 ib, ‘Lateral veins 14-20 on each side of madrid... a eee ~
2a. Hairs (lower leaf surface) on midrib and lateral veins very inconspicuous, short (to | mm), strongly appressed, pedicels c. 20 mm long,calyx lobes 10-15 tats... eae ee a ee ee 8. C. linauana
2b. Hairs (lower leaf surface) on midrib and lateral veins long, silky, strongly appressed, sometimes matted together to produce a papery surface,
pedicels c. 5—8 mm, calyx lobes to'8 mii) ree eee 3 3a. Largest leaves c. 60-165 mm broad, petioles 75-135 mm, bracts c. 35— 40 x
10—16 mm, conspicuous, stronely veined. ::... 2.25.2... ae 3. C. coacta 3b. Largest leaves c. 36-60 mm broad, petioles 40-45 mm, bracts c. 5.5—7 x
1.8—2 mm, inconspicuous, only midrib visible.............. 10. C. angustielliptica 4a. Hairs (lower leaf surface) on midrib spreading: .... 200, clk. a. ..s0ek-seeeeesct mean 5 4b. Hairs (lower leaf surface) on midrib strongly appress................::sssecceeeeeeeees 10
5a. Peduncle c. 45-240 mm long, inflorescence lax, trailing..............eeeeeseeeeeeeeeeeeee Pie: omy te 1. C. penduliflora var. grossipilosa
5b. Peduncle c. 2-30 mm long, inflorescence neither lax nor trailing.................. 6 6a. Bracts c. 20-50 x 7-25 mm, 3-5 veins, clearly visible, calyx lobes
2A MAREE TOM ees Sead St ae a a leek ae eee 7 6b. Bracts c.15—18 x 2—3 mm, only midrib visible, calyx lobes
©. 6-10 TM TOTS oie occ eccce deed, Din, babdnetdaetesncs eee ttt. Aa state ge en ee 9 7a. Inflorescence a several- to many-flowered cyme, both
bracts and bractéolés present... ...050..0. 00. 82. ee eee 8 7b Flowers 1-3, clustered and, at least when young, hidden by two
foliaceous bracts, bracteoles wanting. ...............cssssssseeeeeeeeeeeeeeees 11. C. insolita
8a. Petioles 80-150 mm long, peduncles 5—30 mim...............cccceecesceeeeeeeeeeeeeeeeeeeeeeeees sauaind adboonvivesniNootecd seed dadana easel pl ee ee 2. C. sarawakensis var. longipilosa
Cyrtandra sect. Pleuroschisma 4]
8b. Petioles 30—70 mm long, peduncles 0—2 mm............... 5. Cyrtandra sp. nov. 9a. Largest leaves c. 26-34 mm broad, lower surface displaying finely
reticulate tertiary venation, petiole c. 20—25 mm long ............ 6. C. tesselata 9b Largest leaves c. 85—110 mm broad, lower surface displaying coarsely
reticulate venation, petiole c. 45—50 mm long........... 12. Cyrtandra sp. nov. 10a. Peduncles c. 50-175 mm long, inflorescence lax, trailing...
MPT eisintex. 0k). neo) er, SL. BAO... ae ee 1. C. penduliflora 10b. Peduncles c. 1-30 mm long, inflorescences neither lax nor
mbit os Be es, a EO) 3 be ade RL a ail WR 11 ibaai ise. 60-20 ann booed cece UA heck coe tO RR ES 12 RAG ) aeaeee ond 1S bread a3 41h. een ahs; ae id is en NII ‘3
12a. ‘Tertiary venation on lower leaf surface subscalariform, hairs on stem, leaves (including petioles), bracts, bracteoles, pedicels and calyx
appressed, all: hairs. separatel ncveli icc egl. wisn Ab sdae des 2. C. sarawakensis 12b. Tertiary venation more or less pinnate, hairs on all parts (as above) matted together giving a papery look to the surface................... 3. C. coacta
13a. Stem silky-villous, tertiary venation (lower leaf surface) coarsely reticulate, almost invisible, largest mature leaves c. 85—95 mm broad,
foisbengthiz breath wo 0e3G: beniys. hd enh Qed 9. C. hoseana 13b. Stem pubescent, tertiary venation more or less subscalariform, leaves c. 35-60 mm broad, ratio length: breadth 4—-5.7:1........ eee 14
14a. —_ Largest leaves c. 35-50 mm broad, lateral veins 13—15 on each side
of midrib, bracts 20—30 x 4—7.5 mm, at least midrib clearly
ie aa oe eet hires. Dada. heed Sao r9 tate. a RSI 4. C. tunohica 14b. — Largest leaves c. 55-60 mm broad, lateral veins 18 on each side
of midrib, bracts c. 20 x 4 mm, midrib scarcely
ae ry ee ie Behar tit osiencssisvedass oonssanccsievcassse 7.C. seganica
1. Cyrtandra penduliflora Kraenzl., Mitt. Inst. Allg. Bot. Hamburg 7:100 (1927); B.L. Burtt, Notes Roy. Bot. Gard. Edinb. 30:36 (1970).
Lectotype (Burtt, 1970): Kalimantan, Bukit Obat [c.0°56'N 113°20'E], 150m, Winkler 1328 (lecto HBG, isolecto E).
Unbranched herb, stem to c. 600 mm long, 9-16 mm in diam., base often decumbent, rooting, prop roots present, young parts strongly appressed-pubescent. Leaves opposite, strongly anisophyllous, few forming a fan at apex of stem, reduced leaves stipule-like, c. 20-46 x 5-14 mm, lanceolate, acuminate, strongly appressed- pubescent, largest developed leaves c. 210-300 x 150—200 mm, very broadly elliptic, apex abruptly acute, base cuneate, very narrowly decurrent, briefly or to c. 40 mm, margins entire to obscurely to more distinctly irregularly serrate, lateral veins 1 1—13 on each side of midrib, tertiary venation more or less pinnate, upper surface glabrous
4? Gard. Bull. Singapore 55 (2003)
at maturity, coarsely pitted, lower surface with fine appressed hairs on veins and blade; petiole 130-210 mm long, hairy as midrib. /nflorescence an axillary very lax dichasial cyme, loosely branched, trailing, tending to spring from axils of fallen leaves, peduncle c. 50-175 mm long, pubescent. Bracts (lowermost pair) c. 15—57 x 4-18 mm, lanceolate, acuminate, finely pubescent. Pedicels 6-20 mm long, puberulous. Calyx 5-lobed almost to base, lobes subequal, c. 6-14 x 1.5 mm, deltoid, outside puberulous, inside minutely gland-dotted. Corolla c. 37 mm long (no complete flowers seen, only a big bud), white with an orange-yellow blotch or two bars in throat, puberulous all round mouth. Stamens inserted c. 15 mm above base of tube, filaments c. 5 mm long, glabrous, anthers 3.5 x 1.8 mm, cohering face to face by a small ligature, connective glabrous or a very few glandular hairs. Disc 1 x 1.8 mm, cupular. Ovary 6 x 1 mm, minutely pustulate, coma of hairs at apex. Sty/e 12 mm, glandular-puberulous. Stigmatic lobes c. 1.5 x 1 mm (will enlarge), clavate. Fruit c. 20-45 x 5~7 mm, pericarp smooth. Seeds c.0.3 x 0.25 mm, testa red-brown.
Notes: Cyrtandra penduliflora is a plant of the forest floor, often on steep banks, the stem supported by prop roots. It is distinctive by virtue of its trailing inflorescences, the peduncles mostly very long, the cyme always very lax, far laxer than in any other species of this group. The leaves are broad in relation to length, those of the type specimen, collected in southern Kalimantan, measuring roughly 300 x 200 mm, and exactly matched by those of Burtt 2610 from Sarawak collected 1° of latitude north of the type collection and at very nearly the same degree of longitude (1°56’N 113°06’E versus 0°56’N 113°20’E), but nearly all specimens have the leaves very broad in relation to their length.
Tracheoids with parallel bars of thickening occur in the hypodermis; polymorphic sclereids are often present in the spongy mesophyll but they may be absent.
Other specimens examined: Sarawak. Miri distr., Niah Forest Reserve, [c. 3°52'N 113°44'E], Anderson $31663 (E). Bintulu distr., Ulu Sungai Sinonok, 20-60 m, Hotta 14247 (E, KYO). Tatau, Batang Anap, Ulu Sg. Kana [c. 2°54'N 112°50'E], Mokhtar & Jugah 41779 (E). Sungai Kakus [c. 2°48'N 112°42'E], 30-80 m, Hirano & Hotta 220 (E, KYO). Ulu Kakus [c. 2°48'N 112°42'E], Anap, Haron S29980 (K, E). Ulu Segan [c. 2°30'N 113°E], N Setungan, c. 50 m, Ashton $22019 (E). Pelagus Rapids on Rajang [‘Rejang’], c. 2°10'N 113°E, Burtt & Woods B2559 (E). Sungai Bena [c. 1°56'N 113°06'E], a tributary of Sungai Sut, Burtt B26/0 (E). Sungai Jelok, near Bukit Sengkajang [1°14'N 111°31'E], Lanjak-Entimau P.F, c. 2100 ft, Chai S34034 (E, K). Bukit Ubah Ribu, Ulu Sungai Kaup, Lubok Antu [1°03'N 111°50'E], c. 200 ft, Chai S33761 (E).
Cyrtandra penduliflora var. grossipilosa Hilliard & B.L.Burtt var. nov. a planta
Cyrtandra sect. Pleuroschisma 43
typica pilis longis grossis patentibus in caulibus petiolis et subtus in costis (nec pilis tenuibus valde appressis) filamentorum apicibus et antherarum connectivo dense glanduloso-puberulo (nec glabro vel fere glabro) differt.
Type: Sarawak, ascent to Gunung Mulu [4°O1'N 114°52'E], 13 vi 1962, Burtt & Woods B2060 (holo. E).
Unbranched herb, stem up to | m long, 15 mm diam., base often decumbent, rooting, young parts villous, hairs coarse, spreading. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, largest developed leaves 190-370 x 85-160 mm, elliptic, apex acute to acuminate, base cuneate, very narrowly decurrent, margins entire to irregularly toothed, lateral veins 12—18 on each side of midrib, upper surface glabrous at maturity, coarsely pitted, lower surface with coarse spreading hairs scattered among short appressed ones on the midrib, similar appressed hairs on the blade; petiole 100—150 mm long, hairy as midrib. /nflorescence as in the typical plant, peduncle c. 45—240 mm long. Bracts c. 24-45 x 4-13 mm, finely pubescent. Calyx lobes c. 6-9 mm long, outside puberulous, inside minutely gland-dotted. Corolla (only one flower seen) 41 mm long, white to pale yellow with 2 orange-yellow bars in throat, outside puberulous, mouth glandular-puberulous inside, tube 27mm long, cylindric in lower part, expanded above, upper lobes c. 8 x 8 mm, lower lip c. 14 x 22 mm, median lobe 9 x 8 mm, all lobes suborbicular. Stamens inserted 22 mm above base of tube, filaments c. 5 mm long, glandular-puberulous in upper part, strongly twisted post anthesis, anthers 3 x 2 mm, cohering face to face by a small apiculus, connective densely glandular-puberulous. Disc 1 x 1.5 mm, cupular. Ovary 6 x 1.2 mm, very minutely pustulate, coma of hairs at apex. Style 13 mm long, glandular- pubescent. Stigmatic lobes 3 x 2.5 mm, clavate. Fruit c. 25-35 x 5 m, pericarp smooth. Seeds c. 0.4 x 0.2 mm, testa red-brown.
Notes: This plant differs from typical Cyrtandra penduliflora in the coarse spreading hairs on the stem and along the midrib on the undersurface of the leaf (versus fine strongly appressed ones). The tertiary venation also differs subtly : in the typical plant, the tertiary veins are not clearly visible on the lower surface of the leaf and they are more or less pinnate in arrangement; in the variety they are clearly visible, subscalariform, with a fine reticulum of lesser veins between them. The leaves closely resemble those of C. sarawakensis, distinguished at once by its compact inflorescence. The label on Sands 5390 (cited below) described the leaves as ‘paler beneath with reddish-brown veins’; Church 2210 ‘whitish-green beneath with purple veins distinctly raised’. There is also a difference in the shape of the leaves, those of the typical plant being broader in relation to length than in the variety, ratio of length to breadth being 1.2—1.6:1 versus 1.7—2.5:1; in absolute terms c.150—200 mm versus 85—160 mm (largest leaves).
44 Gard. Bull. Singapore 55 (2003)
Anatomically there is some difference in the thickenings of the tracheoids in the hypodermis: parallel bars in the typical plant, reticulate thickenings in the variety often mixed with others with parallel bars.
The anther-connective in the typical plant is glabrous or nearly so, that in the variety densely glandular-puberulous.
From what is known of the distribution of the species, the variety tends to be more northern and eastern than the typical plant though the few collections from Kalimantan indicate close proximity.
Other specimens examined: Brunei. Temburong river at Wong Nguan rapids, 4°31'N L15°15'E, 120m alt., Coode 6649 (E). Temburong distr., Sungai Belalong, 4°30'N 115°10°'E, c. 100 m, Argent et al. 9/1 (E). Slopes above Temburong headwaters NE of Gunung Retak, 4°22'N 115°17'E, Sands 5309 (E). Tutong, Ladan Hills Forest Reserve, upstream from Belabau on E bank of Tutong river, 4°26'N 114°46'E, 20 m, Coode 6409 (E). Sungai Tokat, a branch of Sungai Batu-Apoi, 50-100 m, Hotta 13759 (E). Sarawak. Limbang [4°45'N 115°E], Ulu Medamit [4°27'N 114°55'E], Sungai Ensungei, Tanjong Long Amok, George et al. $4¢2368 (E). Gunong Api, Ulu Melinau, Tutoh, 4°07'N 115°15'E, 400 ft, Anderson S31784 (E). Melinau Gorge pathway, c. 4°05'N 114°50'E, Burtt & Woods B2239 (E). Gunong Mulu National Park, Gua Rusa, c. 4°02'N 114°50'E, Argent et al. 809 (E). Ulu Belaga, Sungei Semawat, 3°N 113°54'E, c. 250 m, Hansen 623 (E). Dulit Range [c.3°16'N 114°14'E], Sungai Sirui, 200 m, Awa & Yii $46629 (E). Valley of Sungai Keyan Linau-Balui divide, Sungai Nawai, c. 2°26'N 114°10'E, c. 2600 ft, Burtt B1 1456 (E). South of Punan Lusong, c. 2°28'N 114°12'E, Burtt B] 1294 (E). Kalimantan. Sintang, Bukit Baka National Park, Sungai Ella and environs, 0°38'S 112°17'E, 320m, Church 280 (A, E, HBG). Serawai, Uut Labang, 750 m, 0°36'6"S 112°38'56'E, Church 2210 (E).
2. Cyrtandra sarawakensis C.B. Clarke, DC., Monog. phan. 5:209 (1883); B.L. Burtt, Notes Roy. Bot. Gard. Edinb. 30:38 (1970).
Lectotype (Burtt 1970): Sarawak, Kuching, July 1865, Beccari 153 (lecto FI, isolecto K, P).
Unbranched herb, stem c. 110—230 x 7-20 mm, erect or decumbent and rooting, young parts densely appressed-pubescent, hairs c. 2 mm long. Leaves opposite, strongly anisophyllous, c. 3-8 forming a fan at apex of stem, reduced leaves stipule-like, c. 30 x 3 mm, lanceolate, acuminate, closely appressed-pubescent; largest developed leaves c. 225-300 x 57-125 mm, elliptic, apex very acute, base cuneate, very narrowly decurrent, margins subentire to serrate, lateral veins 16—20 on each side of midrib, tertiary veins subscalariform, upper surface glabrous at maturity, coarsely pitted, lower surface appressed-pubescent on blade, long (to 3mm) delicate appressed hairs on midrib and lateral veins (tawny or purple in life); petiole c. 80-150 mm, hairy as
Cyrtandra sect. Pleuroschisma 45
midrib. /nflorescence a several-flowered congested cyme, borne mainly in axils of fallen leaves, peduncie c. 5—30 mm long, stout, puberulous. Bracts persistent, c. 20-35 x 7-20 mm, lanceolate, acuminate, margins entire or toothed, strongly veined, both surfaces appressed-pubescent; bracteoles similar but smaller. Pedicels c.10—12 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes subequal, 4.5—8 x 1—1.5 mm, narrowly deltoid, patent-puberulous outside, inside minutely gland-dotted. Corolla to c. 44 mm long, palest yellow, 2 orange-yellow bars on palate, outside puberulous, hairs acute, tube c.27 mm long, narrowly cylindric below, abruptly expanded above, two upper lobes c. 9 x 9 mm, lower lip 3-lobed, c.14 x 24 mm, median lobe c. 8 x 8 mm, all lobes suborbicular, glandular-puberulous inside, with patch of longer hairs adjacent to sinus of upper lobes. Stamens inserted 20-25 mm above base of tube, c.6mm long, strongly twisted after anthesis, minute glandular hairs fringing connective; lateral staminodes c. 2 mm long, posticous staminode c. 1 mm. Disc c.2x 2mm, cupular. Ovary c.7 x 1.4mm, minutely gland-dotted, coma of hairs at apex. Stigmatic lobes c. 3.5 x 1.2 mm, spathulate, conspicuous, stigmatic papillae relatively long. Fruit c. 20-25 x 4-5 mm, pericarp smooth. Seeds c. 0.25 x 0.2 mm, testa bright red-brown, reticulate.
Notes: Cyrtandra sarawakensis was originally collected by Beccari at Kuching. Beccari had a hut on Mt. Matang, not far from Kuching, and this is possibly the type locality; Burtt & Woods B1947 (cited below) precisely matches the isolectotype specimen in Paris. The other collections seen came from the Semengoh Forest Reserve a few miles south of Kuching. Many species of Cyrtandra seem to have a very limited distribution and this may be the case here. The distinguishing features of C. sarawakensis are the many lateral veins, tertiary veins subscalariform and clearly visible especially on lower surface, hairs on veins (lower surface) strongly appressed, peduncles c. 5—30 mm long, bracts broad, strongly and conspicuously veined. It is a plant mainly of the forest floor.
Tracheoids with parallel bars occur in the 1—2-layered hypodermis of the leaf, and there are no sclereids in the spongy mesophyll. Stomatal turrets are well developed.
Other specimens examined: Sarawak. Mt. Matang [1°36'N 110°11'E], Burtt & Woods B1947 (E); Matang, Ulu Sungai Rayu, Lee $54099 (E); Semengoh Forest Reserve, 12 miles S of Kuching [1°28'N 110°22'E], Burtt & Martin B4712 (E); Semengoh Forest Reserve, Burtt & Woods B2487 (E).
Cyrtandra sarawakensis var. longipilosa Hilliard & B.L.Burtt var. nov. a planta typica primum pilis in caule et costa grossis et patentibus (nec tenuibus et valde appressis) et calycis lobis plerumque longioribus 4.5—8 mm (nec 2.54 mm) et apicibus foliorum longe acuminatis (nec acutis) differt.
46 Gard. Bull. Singapore 55 (2003)
Type: Sarawak [c.2°N 114°10'E, Hose Mts.], Melinau Community Forest, near base camp at Nanga Tunoh, Burtt & Martin B4772 (holo. E).
Notes: The variety has been collected north and east of Kuching. It resembles Cyrtandra sarawakensis in nearly all features, including habit, number of lateral veins in leaves, subscalariform tertiary venation, bracts and other floral characters, but it differs in its long-acuminate (not merely acute) leaf tips, hairs on stem, petiole and midrib on under surface of leaves being coarse and spreading (not fine and closely appressed) and possibly also in length of calyx lobes, 2.54 mm (not 4.5—8 mm).
The coarse spreading hairs are brightly coloured in life, and in the dried state the colour often persists in the cross walls of the hairs; the hairs on C. sarawakensis may also be coloured, but, being so much finer, the colour is more difficult to see in dried specimens. The occurrence of a variety with long coarse spreading hairs as opposed to appressed ones is paralleled in C. penduliflora, a species with tertiary venation indistinguishable from that of C. sarawakensis; C. penduliflora differs markedly in its lax inflorescence, and many specimens have fewer lateral veins in the leaves.
Other specimens examined: Sarawak. Tatau, path to Bukit Buan, 300 ft. [c.2°56'N 112°55'E], Purseglove 5469 (E). Tatau, Ulu Anap [c. 2°56'N 112°49'E], Mokhtar S448]5 (E). Bukit Kana [2°42'N 112°54'E], 600-850 m, Hirano & Hotta 1299 (KYO); Bukit Kana, 50-150 m, Hirano & Hotta 15480 (E). Ulu Anap, Bukit Mersing [2°30'N 113°06'E], c.1000 m, Ashton $17679 (E). Hose Mts., Bukit Temedu [2°24'N 113°41'E], c. 1100 m, Ashton $19007 (E). Pelagus Rapids on Rajang River [‘Rejang’], also on Bukit Raya nearby, c. 2°10'N 113°E, Burtt & Woods B2544 (E). Bukit Raya, Smith § 27719 (E). SE end Hose Mts., Bukit Nibong, c. 2°6'W 113°42'E, Burtt & Martin B4845 (E). Sungai Bena, a tributary of Sungai Sut, c. 1°55'N 113°5'E, Burtt B2609 (E). Lundu distr., Gunung Gading [1°44'N 109°50'E], Burtt & Woods B2685 (E). Bukit Lumut, Ulu Amau [1°27'N 112°04'E], c. 950 m, Ashton $21264 (E). Bukit Goram, Ulu Sungai Kapit [1°34'N 112°45'E], Chai S36172 (E).
3. Cyrtandra coacta Hilliard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke venis lateralibus foliorum 9-15 (nec 15-20), venis tertiariis plus minusve pinnatis (nec subscalariformibus) pilis in bracteis et in foliorum pagina inferiore papyraceo- coactis (nec pilis singulatis manifestis) distinguenda.
Type: Sabah, Ranau distr., road from Lohan to Mamut Copper Mine near Tank 47,6°O1'N 116°41'E, 1100 m, 9 vii 1984, Beaman 10656 (holo E).
Cyrtandra sect. Pleuroschisma 47
Unbranched herb to c. 800 mm tall, stem horizonal at base then curving upwards, clad in a papery indumentum of matted hairs, leafy and floriferous on erect part. Leaves opposite, strongly anisophyllous, reduced leaf stipule-like, c. 25 x 2 mm, lanceolate, enveloped in papery indumentum; developed leaf c.180—360 x 60—110(—-165) mm, elliptic to broadly elliptic, apex acute to almost acuminate, base cuneate, very shortly decurrent, margins entire to obscurely or distinctly remotely serrate, lateral veins 9-15 on each side of midrib, tertiary veins more or less pinnate, upper surface glabrous, finely and closely pitted, lower surface papery, any loose hairs very delicate, toc.3mm long; petiole 75—135 mm, enveloped in papery indumentum. /nflorescence a several- flowered, very congested, axillary cyme, almost sessile. Bracts persistent, c. 35-40 x 10-16 mm, lanceolate, strongly veined, enveloped in papery indumentum, bracteoles similar but smaller. Pedicels c.5 mm. Calyx 5-lobed almost to base, lobes subequal, c. 3.5 x 1 mm, narrowly deltoid, tube c. 0.5 mm, outer surface with papery indumentum, inside minutely gland-dotted. Corolla c. 40 mm long, white or cream with 2 orange-yellow bars in throat below the two sinuses of lower lip, outside minutely puberulous, hairs acute, tube c. 25.5 mm, cylindric below for c.12mm then abruptly expanded, two upper lobes c. 9 x 6.5 mm, lower lip 3-lobed, c.14.5 x 12 mm, median lobe c. 9 x 7.5 mm, all lobes more or less suborbicular, glandular-puberulous inside and down tube to insertion of filaments. Stamens inserted c.19mm above base of tube, filaments c. 5 mm, strongly coiled post anthesis, glandular-puberulous at apex, anthers c. 2 x 1.5 mm, cohering face to face by a prominent ligature, connective densely glandular-puberulous; lateral staminodes c. 2 mm, posticous staminode c.lmm. Disc 1.5 x 2 mm, cupular. Ovary c. 8 xX 2 mm, minutely pustulate, coma of hairs at apex. Style c. 15 mm, glandular-puberulous. Stigmatic lobes c. 3 x 1.5 mm, clavate, conspicuous, stigmatic papillae relatively long. Fruitc. 20 x 6mm (Mendum & Lamb 234A), pericarp smooth. Seeds c. 0.5 x 0.25 mm, testa red-brown.
Notes: This species has the aspect of Cyrtandra sarawakensis but is at once distinguished by the indumentum on the vegetative parts: on stem, leaves (including petioles), bracts, bracteoles, pedicels and calyx, the hairs are matted together (coactum meaning felt), giving a papery look to the surface; in C. sarawakensis, hairs on the corresponding parts are appressed, the individual hairs all separate. The very striking difference is easily seen along the midrib on the lower surface where the matted hairs of C. coacta form a papery-looking skin in sharp contrast to the strongly appressed but distinctly separate hairs in typical C. sarawakensis and the long coarse spreading hairs in var. Jongipilosa. The indumentum of C. hoseana and C. angustielliptica is not unlike that of C. coacta but both species differ, inter alia, in their narrow bracts lacking conspicuous venation.
In C. coacta, there are 9—15 lateral veins on each side of the midrib, in C. sarawakensis 15—20, while the tertiary veins are more or less pinnate in C. coacta,
48 Gard. Bull. Singapore 55 (2003)
subscalariform in C. sarawakensis. Furthermore, the leaves differ in anatomical detail: the tracheoids in the hypodermis of C. coacta have reticulate bars and there are no stomatal turrets; in C. sarawakensis the tracheoids have parallel bars and there are prominent stomatal turrets.
The stamens may differ too, but too few flowers have been seen to be certain: in C. coacta, the connective in the anther and the top of the filament are densely glandular-puberulous; in C. sarawakensis both are glabrous or very minutely glandular.
Cyrtandra coacta is known mainly from Sabah and the northern part of Sarawak (Yii et al. S5/691 came from the upper reaches of the Baleh River, in east central Sarawak), while C. sarawakensis is found in the southern part.
Other specimens examined: Sabah. Sinsuron road, 850—900 m, 5°40'N 116°22'E, Mendum & Lamb 23 (E), Sinsuron road, Mendum & Lamb 23A (E); Lahad Datu distr., Research Centre, Ulu Sungai Segama, 200m, Argent et al. SAN 108203 (E). Mt. Kinabalu, eastern shoulder, 6°05'N 116°36-40'E, 3400 ft, Chew et al. 963 (E, K), Chew et al. 647 (E, K). Sarawak. Seventh division, Ulu Belaga, Sungai Semawat, 3°N 113°54'E, c. 250 m, Hansen 715 (E). NE slope Bukit Kama [2°42'N 112°54'E], c. 450 m, Hirano & Hotta 1239 (E, KYO). Ulu Sungai Balan, Bukit Batu Tiban, Yii et al. SS1691 (E).
4. Cyrtandra tunohica Hilliard & B.L.Burtt sp.nov. a C. seganica Hilliard & B.L.Burtt foliis minoribus (maxime 170—200 mm, nec 300 mm longis) et pro rata angustioribus (4—-5.7:1, nec 2.5-4.5:1), bracteis plerumque majoribus (20-30 x 4-7.5 mm prope basin, nec 20 x 4 mm), costa et venis lateralibus praecipue in pagina inferiore visibilibus (nec costa vix, lateralibus haud visibilibus), calycis lobis 2-4 x 1 mm (nec 4.5—-6 x1.5 mm) distinguenda.
Type: Sarawak, Kapit distr., Sungai Melinau [Hose Mts.], Nanga Tunoh, c.2°3'40"N 113°38'20"E, c.450ft, Burtt B12664 (holo E).
Unbranched herb, stem c. 5—6 mm in diam. near apex, height unknown, base decumbent and rooting, erect part toc. 150 mm long, upper part finely pubescent, soon glabrescent. Leaves opposite, crowded near apex of stem, slightly to strongly anisophyllous even on one stem, highly reduced leaves, when present, to c. 65 x 2mm, almost filiform, major developed leaves 170-200 x 35-50 mm, narrowly elliptic, apex acuminate, base narrowly cuneate, very narrowly and shortly decurrent, margins obscurely serrate to crenulate, lateral veins 13-15 (as few as 10 in minor leaves), tertiary venation subscalariform, upper surface glabrous at maturity, pitted, lower surface thinly hairy on blade, more densely so on veins, all hairs to c. 1 mm long, appressed; petiole 50— 70 mm long, hairy as midrib. /nflorescence: few-flowered congested cymes, axillary,
Cyrtandra sect. Pleuroschisma 49
extending from near apex of stem to well down below the leaves, almost sessile. Bracts persistent, 20-30 x 4—-7.5 mm, lanceolate, long-acuminate, both surfaces puberulous, midrib and lateral veins clearly visible especially on lower surface; bracteoles similar but much smaller. Pedicels c. 1-8 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes subequal, 2-4 x 1 mm, narrowly deltoid, outside and margins closely appressed-pubescent, inside minutely gland-dotted. Corolla c. 30 mm long, cream with yellow marks on palate, outside puberulous, hairs acute, tube c. 20 mm long, lower half narrowly cylindric, upper part expanded, two upper lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe c. 5 x 3.5 mm, all lobes subrotund, glandular-puberulous inside, hairs extending down into throat. Stamens inserted c.14 mm above base of tube, filaments c. 4 mm long, strongly twisted, anthers 2 x 1.2 mm, cohering face to face by a small ligature, both anthers and filaments glabrous; lateral staminodes c. 2 mm, posticous staminode c. 0.5 mm. Disc 1—1.2 x 1.2—1.8 mm, cupular. Ovary c. 3.5—7 x 0.8—2 mm, very minutely gland-dotted, crowned with coma of relatively long hairs. Style 5-6 mm long, puberulous. Stigmatic lobes 3—4 x 1-2 mm, clavate. Fruit 10-18 x 4-6 mm, pericarp smooth, deeply grooved longitudinally. Seeds 0.3-0.5 x 0.2—0.3 mm, testa red-brown.
Notes: The outstanding feature of Cyrtandra tunohica as we know it is variation in the degree of anisophylly; in the rest of the species in this section of Cyrtandra, the minor leaf is always reduced to a stipule-like outgrowth (with the caveat that several species are ill-known). In C. tunohica, the size of the leaves in each pair ranges from subequal (when the stem is much more leafy than is usual in this section as a whole) through degrees of reduction culminating in an extraordinarily long, almost filiform, stipule-like leaf; this occurs on a single stem. As the species is known from only one locality, the taxonomic value of this character remains to be assessed. The plants (very few) were found on the steep shady earth bank of a stream gulley, and were seen only once, despite further fieldwork in the area on this and other occasions.
In its relatively narrow leaves, C. tunohica resembles both C. seganica and C. angustielliptica; it is easily distinguished from both by its bracts, 4-7.5 mm broad near the base (not 1.8—4 mm) and with both midrib and lateral veins raised on the lower surface and thus clearly visible (in contrast to nearly or quite invisible). It is further distinguished from C. seganica by its leaves, the largest 170-200 mm long (not c. 300 mm) and relatively narrower (ratio of length to breadth 4—5.7:1 versus 2.5—4.5:1), short (to c.1mm long) appressed hairs along the midrib on the lowersurface (not long coarse spreading hairs), and c.13—15 (not 18) lateral veins on each side of the midrib.
Not only do the bracts differ from those of C. angustielliptica, but so do the leaves in several details: lateral veins 13-15 in major leaves (not 7-10), hairs on undersurface of leaves up to c.1mm long, sparse on the blade (in contrast to silky-villous, hairs c. 2 mm long, the blade eventually glabrous, hairs persistent on midrib), tertiary veins Clearly visible (not invisible or very nearly so).
50 Gard. Bull. Singapore 55 (2003)
The leaf of C. tunohica has tracheoids with reticulate thickening in the hypodermis and lacks sclereids in the mesophyll; C. seganica and C. angustielliptica have similar tracheoids but differ in having polymorphic sclereids in the mesophyll.
Other specimen examined: Sarawak. Kapit distr., Sungai Melinau [Hose Mts.], Nanga Tunoh, c. 2°3'N 113°38'E, Burtt B12665 (E).
5. Cyrtandra sp. nov.
Unbranched herb, stem very stout, c. 12 mm diam., c. 300-600 mm tall, young parts villous, lower parts glabrous. Leaves opposite, few crowded near apex of stem, strongly anisophyllous, reduced leaves stipule-like, c. 40 x 2 mm, narrowly linear-lanceolate, villous; largest developed leaves c. 195-260 x 80-90 mm, elliptic or oblanceolate, apex abruptly acuminate, base narrowly cuneate, very narrowly decurrent, margins entire, lateral veins 16—18, tertiary veins finely reticulate, upper surface glabrous, coarsely pitted (at maturity; densely pubescent when juvenile), lower surface densely pubescent, veins pilose; petiole c. 30-70 mm long, villous. /nflorescence a many-flowered, highly congested dichasial cyme, axillary, forming bracteate clusters borne on the leafless part of the stem; peduncle very stout, to 2 mm long. Bracts persistent, primary pair c.20x7mm, lanceolate, acute, midrib and lateral veins prominent, outside densely pubescent, inside hairs mainly near apex and base; bracteoles similar but narrower, a pair subtending each pedicel. Pedicels c.5 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes c. 4 x 1 mm, outside puberulous, hairs patent, inside minutely gland-dotted. Corolla c. 30 mm long, white, 2 orange-yellow bars on palate, outside puberulous, hairs acute, tube c. 20 mm long, lower part narrowly cylindric, upper part expanded, two upper lobes c. 5 x 3.5 mm, lower lip 3-lobed, median lobe c.5 x 4mm, all lobes subrotund, glandular-puberulous on lower lip, hairs extending down floor of tube, a median tuft of eglandular hairs at base of each upper lobe, a few glandular hairs towards outer margins. Stamens inserted c.17 mm above base of tube, filaments c. 4-5 mm long, twisted once near base, anthers 2.2 x 1 mm, cohering face to face by a conspicuous ligature, connective minutely glandular, glands extending briefly down back of filaments; lateral staminodes c. 2 mm, apex strongly hooked, tip globular, pale yellow (as anthers), posticous staminode c.1 mm. Disc 1.8 x 2 mm, almost cupular (deeply notched on one side). Ovary 7 x 2 mm, glabrous except for conspicuous coma. Style 10 mm (at anthesis), pubescent, hairs acute. Stigmatic lobes c. 2 x 1.5 mm (post anthesis), spathulate, stigmatic papillae conspicuous. Fruit 23 x 6 mm (one seen), pericarp smooth, split longitudinally, only apex and base cohering. Seeds c. 0.25 x 0.2 mm, testa red-brown.
Notes: This species is allied to Cyrtandra tunohica and C. sarawakensis var.
Cyrtandra sect. Pleuroschisma 34
longipilosa. It is distinguished from C. tunohica by the relatively long patent indumentum on stems, petioles and backs of leaves (versus hairs short, appressed), leaves c. 80-90 mm broad (not 35-50 mm) and many- (not few-) flowered inflorescence. It differs from C. sarawakensis var. longipilosa by its shorter petioles (30-70 mm, not 80—150 mm) and almost sessile inflorescence. Also, the pubescence on the undersurface of the leaf blade is dense and patent (not appressed). This is obvious in Haron $29978 (K), but not so well marked on the duplicate (E); it is also obvious in Jugah Kudi S23712 (K) where the finely reticulate tertiary venation shows up well.
The two collections seen are not good: Haron S29978 was obviously pressed when badly withered; the sheet at Kew is particularly bad, but this is the one with a nearly complete corolla and several buds, from which the floral details were extracted. Only an old fruit survives on the sheet in Edinburgh, and the base of a fruit on Jugah Kudi S23712 appears to have eight loculi! The need for good collections before the species can be named is obvious.
The leaves have a 1—2-layered hypodermis, and tracheoids with parallel bars in the hypodermis.
Specimens examined: Sarawak, Anap, Ulu Kakus [c.2°30'N 113°E], Othman Haron $29978 (E, K); Balingian, Ulu Sungai Arip, Bukit [ju [roughly 2°40'N 112°40'E], Jugah Kudi §23712 (K, L).
6. Cyrtandra tesselata Hilliard & B.L.Burtt sp. nov. a C. tunohica (etiam folia anguste lanceolata gerente) caulibus et pagina foliorum inferiore villosis (nec breviter pubescentibus), petiolo 20—25 mm longo (nec 50—70 mm) villoso (nec pubescente), bracteis c.15 x 1.8—2 mm venatione invisibili (nec 20-30 x 4-7.5 mm venatione elevata conspicua), calycis lobis c.6—8 mm longis (nec 2.4 mm) differt.
Type: Sarawak, Bintulu distr., en route (survey route) from Sungai Mah to Sungai Shinonok, Ulu Sungai Minah, alt. 40-150 m, Hotta 1/4117 (holo KYO).
Herb, probably unbranched, height unknown, stem collected 50 x 10 mm, villous, hairs 3-4 mm long, leafy throughout, internodes c. 10 mm long. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 30 x 3 mm, narrowly lanceolate,long-acuminate, major mature leaves 150-230 x 26-34 mm, narrowly elliptic, apex gradually very acute, base narrowly cuneate, margins very obscurely serrulate, lateral veins 12—14 on each side of midrib, upper surface glabrous at maturity, pitted, lower surface villous, longest hairs (2-3 mm) on midrib, shorter on lesser veins, tertiary venation finely reticulate, all veins raised, hairs brownish (dried state) making venation very conspicuous; petiole 20—25 mm long, villous. Inflorescence a few-flowered axillary cyme, peduncle c. 4mm, very stout. Bracts c. 15x 1.8—2 mm, narrowly lanceolate, densely pubescent, venation obscure; bracteoles similar but
52 Gard. Bull. Singapore 55 (2003)
Pedicels c.10mm (in fruit), pubescent. Calyx 5-lobed almost to base, tube c. 0.5 mm, lobes 6-8 x 1 mm in fruit, narrowly deltoid, outside and margins puberulous, hairs patent, inside minutely gland-dotted. Corolla not seen, only very young buds present, puberulous outside, hairs acute, lobes minutely glandular inside (tube scarcely developed). Stamens: filaments scarcely developed, anthers 2 x 0.8 mm, minutely glandular on margins of connective. Disc cupular. Ovary and style very young, style pubescent. Fruit c. 35 x 5 mm, pericarp smooth, traces of coma still visible at apex. Seeds c. 0.25 x 0.2 mm, testa bright red-brown.
Notes: This species is so distinctive that lack of flowers should not preclude its formal recognition. The leaves are closely set down the 150 mm length of the piece of stem on the herbarium sheet, in contrast to most of its allies where the leaves appear to be quickly deciduous leaving a few, forming a fan, at the apex of the stem. Unfortunately, the collector gave no notes either on habit or habitat of his plant. The major leaves are short-petioled, narrowly elliptic, and at maturity glabrous above but villous below, with midrib, lateral and tertiary veins raised (the latter forming a fine reticulum) and conspicuous, partly owing to the dark hairs clothing them. Similar tertiary venation has not been seen in any other species in this section.
Cyrtandra tesselata is diagnosed above against C. tunohica, which has similarly narrowly lanceolate leaves, but the indumentum on stem, petioles and leaf undersurface in C. tesse/ata is villous (not merely shortly pubescent), the petioles are remarkably short (20-25 mm versus 50-70 mm), the bracts are c. 15 x 1-2 mm, venation invisible (not 20-30 x 4—7.5 mm, venation raised and conspicuous), calyx lobes c. 6-8 mm long (not 24 mm). In sharp contrast to that of C. tesselata, the tertiary venation in the leaves of C. tunohica is rather indistinct, and tends to be subscalariform.
The leaf anatomy of C. tesselata is unlike that of C. tunohica: vertically long tracheoids with parallel bars in the hypodermis, polymorphic sclereids in the mesophyll; the tracheoids in C. tunohica have reticulate bars and there are no sclereids in the mesophyll.
7. Cyrtandra seganica Hillard & B.L.Burtt, sp. nov. a C. sarawakense C.B.Clarke bracteis tantum c. 4 mm latis (nec 7-20 mm) sine venatione conspicua (nec costa et venis lateraliibus prominentibus) et folia angustiora prorata longitudinis (5.5:1 nec 2.54.5:1) facile distinguenda.
Type: Sarawak, Bintulu distr., Ulu Segan [c. 2°30'N 113°E], rocky stream bank, 750 ft., Jlias Paie §27215 (holo E).
Cyrtandra sect. Pleuroschisma 53
Unbranched herb, height unknown, stem c. 8 mm diam. near apex, apex densely pubescent, rest of stem quickly glabrous. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, c. 25 x 3 mm, lanceolate, acuminate, densely pubescent, hairs spreading; developed leaves c. 300 x 55—60 mm, elliptic, acute, base narrowly cuneate, very narrowly decurrent for c. 35 mm, margins entire, lateral veins c. 18 on each side of midrib, tertiary venation subscalariform, upper surface glabrous, pitted, lower surface with very small fine strongly appressed hairs on blade, coarse much longer (c. 1 mm) hairs on midrib and lateral veins, probably appressed and probably purplish in life (specimens well glued); petiole c. 55-70 mm long, hairy as midrib. /nflorescence: few-flowered congested cymes borne mainly in axils of fallen leaves, peduncle c. 4-8 mm long, very stout, puberulous. Bracts persistent, c. 20 x 4mm, narrowly lanceolate, acuminate, margins entire, both surfaces puberulous, hairs spreading, midrib scarcely visible; bracteoles similar but smaller. Pedicels c. 8-10 mm. Calyx lobed nearly to base, lobes subequal, 4.5—6 x 1.5 mm, narrowly deltoid, tube c. 0.5 mm, outside and margins puberulous, inside minutely gland-dotted. Corolla not seen. Fruit c. 18—36 x 4—5.5 mm, pericarp smooth. Seeds immature.
Notes: Cyrtandra seganica is currently known with certainty only from the type collection. In the venation of the leaves, including the subscalariform arrangement of the tertiary veins, it resembles C. sarawakensis, but the leaves are narrower in relation to their length (ratio of length to breadth 5.5:1 versus 2.5-4.5:1). There is a striking difference in the bracts, only c. 4 mm broad near the base and with the midrib but faintly visible in C. seganica, in contrast to bracts c. 7-20 mm broad with prominent venation in C. sarawakensis. The leaves are similar anatomically in having tracheoids with parallel bars in the hypodermis, but differ in the presence of many polymorphic sclereids in the mesophyll of C. seganica whereas the leaves of C. sarawakensis lack sclereids.
Two collections from further south and west resemble C. seganica in leaf-venation and in narrow bracts. They appear to differ in the shape of the leaves (oblanceolate) and also in the ratio of length to breadth (3.25—-4.6:1), this being within the range of C. sarawakensis. The leaves also appear to differ anatomically, in that the tracheoids have reticulate bars, not parallel bars; polymorphic sclereids occur in the mesophyll. These two collections come from Sarawak from Datu Permanent Forest [Bukit Datu? at 1°29'N 110°50'E], 420 m alt., on the ridge between two streams, Lee S§41904 (E) and 17" mile Bau/Lundu road, Gunong Undan (1° 27'N 109° 59'B), stream bank, 50m, Yii S45942 (E).
Much more needs to be known about all three plants described here.
54 Gard. Bull. Singapore 55 (2003)
8. Cyrtandra linauana B.L.Burtt, species nova ex affinitate C. sarawakensis C.B.Clarke et C. hoseanae B.L.Burtt, ab ambabus nervis lateralibus foliorum 7-8 (nec 12—20) et calycis lobis 10-15 mm longis (nec usque ad 8 mm) differt.
Type: Sarawak, Belaga distr., hill just N of Long Linau [c.2°40'N 114°E], Burtt 11478 (holo E; iso BO, L, SAR).
Herb, stem simple, c.100 mm tall, 8 mm diam., puberulous, hairs very closely appressed. Leaves opposite, strongly anisophyllous, 5 pairs crowded near apex of stem, lower ones fallen, reduced leaf stipule-like, c. 35 x 5 mm, lanceolate, long-acuminate; developed leaf c. 210-300 x 82-83 mm, elliptic, somewhat falcate and eccentric, apex rather abruptly acuminate, base cuneate, very shortly and narrowly decurrent, margins entire, lateral veins 7-8 on each side of midrib, tertiary venation coarsely reticulate, upper surface mottled silver (living material), glabrous, pitted, lower surface with blade glabrous, midrib and lateral veins strongly appressed-pubescent, hairs very inconspicuous; petiole 70-80 mm, thickly appressed-pubescent. J/nflorescence an axillary dichasial cyme, flowers few, branches of cyme very short (3-4 mm), bearing only 4 flowers (apical bud on each branch suppressed), peduncle c. 5-8 mm, very stout. Bracts persistent, c. 20 x 2 mm, narrowly lanceolate, acuminate, both surfaces strongly appressed-pubescent, bracteoles similar but smaller, a pair subtending each pedicel. Pedicels c.20 mm, appressed-pubescent. Calyx 5-lobed almost to base, tube c.0.5 mm, lobes 10—15 mm, narrowly linear-lanceolate, outside appressed-pubescent, inside minutely gland-dotted. Corolla 26-30 mm long, white, throat yellow shading to orange, outside minutely puberulous, hairs acute, red, tube 17—18 mm long, cylindric in lower third, abruptly expanded above, two upper lobes 7—8 x 5 mm, lower lip 3-lobed, c. 9-12 mm long, median lobe 7—8 x 5 mm, all lobes oblong-elliptic, glandular- puberulous inside. Stamens inserted 8.5 mm above base of tube, filaments 2.5 mm, glabrous, anthers 2.2x 1.2mm, cohering face to face by a prominent ligature, connective fringed with stout red hairs; lateral staminodes 3 mm, posticous staminode 1.2 mm. Disc 1.5 x 2 mm, cupular but deeply excavated on one side. Ovary 5 x | mm, minutely pustulate, coma of hairs at apex. Style 7mm, glandular-puberulous. Stigmatic lobes 4x 1.8 mm, clavate, conspicuous. Fruit 24 x 3.8 mm, pericarp smooth.
Notes: Cyrtandra linauana resembles both C. sarawakensis and C. hoseana in the size, shape and stance of its leaves but differs from both in having only 7-8 lateral veins on each side of the midrib (not 12—20). The tertiary venation also differs: in C. linauana it is rather indistinct and very coarsely reticulate though the veinlets may be more subscalariform towards the margins; in C. sarawakensis the veins are prominent and closely subscalariform; in C. hoseana they are very nearly invisible.
Cyrtandra sect. Pleuroschisma 55
The tracheoids in the hypodermis have reticulate bars as in C. hoseana but there are no sclereids in the spongy mesophyll. In C. sarawakensis the tracheoidal bars are parallel and, as in C. linauana, there are no sclereids in the spongy mesophyll; C. hoseana has polymorphic sclereids.
The inflorescences of all three species differ: in C. /inauana, the bracts are only c. 2 mm broad and the venation is not or scarcely visible; pedicels c. 20 mm long, making the inflorescence rather lax. In C. sarawakensis, the bracts are very conspicuous, being broad, venation prominent, pedicels c. 9-13 mm long, inflorescence congested. In C. hoseana, the bracts are c. 34 mm broad, venation inconspicuous, pedicels c. 5-10 mm long, inflorescence congested. Stout red hairs fringe the connective in C. linauana; in C. sarawakensis and C. hoseana, the connective is either glabrous or glandular.
Cyrtandra linauana is known with certainty only from the type collection made at c. 2°40'N 114°E. There is another collection, made at c. 2°48'N 112°59'E, that is clearly very close to C. linauana, namely Hirano & Hotta 88] (KYO, E) from Bukit Keyan, Ulu Sungai Kakus, alt. 100-300 m. It differs in several details: leaves not falcate and with 10 lateral veins, otherwise identical with those of C. linauana, flowers fascicled in the leaf axils, pedicels c. 15 mm long, calyx lobes 6.5—9 mm. The most significant difference is the fascicled flowers in contrast to the shortly pedunculate dichasial cyme of C. linauana, where the long pedicels produce a very lax inflorescence.
The leaves of C. angustielliptica have a deceptive similarity to those of C. linauana. That species is, however, distinguished by the short (to c. 1 mm) appressed and inconspicuous hairs on stem, petiole and midrib of leaves on lower surface (not 2—3 mm long, silky, appressed hairs matted together and easily seen), bracts c. 20 mm long (not 5—8 mm), connective of anthers fringed with stout red hairs (versus glandular puberulous ones all over). The inflorescence of C. angustielliptica is a nearly sessile, very congested, cyme.
9. Cyrtandra hoseana B.L.Burtt, Notes Roy. Bot. Gard. Edinb. 30:33 (1970). Type: Sarawak, Lambir Hills, c. 4°7'N 113°55'E, sandstone banks, Burtt & Woods B2365 (holo E, iso SAR).
Unbranched herb, stem 20-150 cm long, c. 8 mm diam. near apex, hanging from sandstone cliffs and banks, sometimes supported by prop roots, young parts silky villous, hairs strongly appressed. Leaves opposite, strongly anisophyllous, few arranged in a fan at stem apex, reduced leaves stipule-like, c. 20 x 10 mm, broadly lanceolate, appressed silky-villous; largest developed leaves c. 255-320 x 85-95 mm, elliptic, apex abruptly acute, base cuneate, shortly and very narrowly decurrent, margins entire to obscurely serrulate, lateral veins 12-15 on each side of midrib, tertiary venation coarsely reticulate, upper surface glabrous at maturity, coarsely pitted, lower
56 Gard. Bull. Singapore 55 (2003)
surface with long, strongly appressed, silky hairs on midrib and lateral veins, blade glabrous or almost so at maturity; petiole c. 45-60 mm, hairy as midrib. /nflorescence an axillary, very congested, cyme, mainly in axils of fallen leaves, few-flowered, peduncle 2-6 mm long. Bracts c. 15-18 x 34 mm, persistent but inconspicuous, lanceolate, acuminate, closely appressed-puberulous, only midrib visible on lower surface. Pedicels c. 5—10 mm long, puberulous. Calyx 5-lobed almost to base, lobes subequal c. 2.2-3.5 x 1.25 mm, deltoid, outside appressed-pubescent, inside minutely gland-dotted. Corolla c. 32 mm long, white or creamy white outside, 2 orange-yellow bars in throat, outside puberulous, hairs acute, red, tube c. 20 mm long, lower half narrowly cylindric, abruptly expanded above, two upper lobes c. 5 x 7 mm, lower lip 3-lobed, c.12.5 x 20 mm, median lobe c. 6 x 7 mm, all lobes subrotund, glandular- puberulous inside and for a short way down tube. Stamens inserted c. 16 mm above base of tube, filaments c. 5 mm, curved and twisted once, minute globular glands at apex, anthers c. 3 x 1.6 mm, cohering face to face by a small ligature, connective with a few glandular hairs and globular glands; lateral staminodes c. 1.2 mm, posticous staminode c.lmm. Disc 1 x 1.8 mm, cupular. Ovary 8 x 1.6 mm, very minutely gland-dotted, coma of hairs at apex. Style 10mm, glandular-puberulous. Stigmatic lobes not fully developed, c. 2 mm long, clavate. Fruit c. 27-45 x 5 mm, pericarp smooth. Seeds c. 0.4 x 0.25 mm, testa red-brown.
Notes: So far, C. hoseana is known with certainty only from the Lambir Hills. Now that more material is available, it is clear that Burtt & Woods B2212 (E, SAR) from the Melinau Gorge, which was cited in the original description, is a distinct species, described here as C. angustielliptica.
It is worth noting again that ‘the youngest leaf is very densely covered with shining silky hairs and it hangs downwards at the top of the shoot, like a tiny silvery flag’ (Burtt, 1970). In this species, the tertiary veins are scarcely discernible on the lower surface of the leaf, but form a coarse reticulum that may just be visible on the upper surface (compare the tertiary veins of C. sarawakensis).
Tracheoids with reticulate bars occur in the 1—2-layered hypodermis, and a few polymorphic sclereids in the mesophyll.
Other specimen examined: Sarawak. Lambir National Park, Sungai Lapoh and nearby, Burtt 11546 (E).
10. Cyrtandra angustielliptica Hilliard & B.L.Burtt sp. nov. a C. hoseana B.L.Burtt foliis 36-60 mm latis (nec 85-90 mm) apice acuminato (nec abrupte acuto), venis lateralibus utrinque costae 7-10 (nec 12-15), calycis lobis 5-8 mm longis (nec 2.2—3.5 mm) distinguenda.
Cyrtandra sect. Pleuroschisma S¥
Type: Sarawak, Sungei Melinau Gorge, c. 4°5'N 114°50'E, sandstone hillock, Burtt & Woods B2212 (holo E).
Unbranched herb, stem of unknown height, c. 6-7 mm in diam. near apex, young parts silky-villous, hairs strongly appressed, matted together. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 18 x 6 mm, lanceolate, silky-villous as stem, developed leaves up to c.10, largest 170-275 x 36-60 mm, elliptic, apex acuminate, base cuneate, margins entire to obscurely serrulate but teeth made conspicuous by apical tuft of hairs, lateral veins 7—10 on each side of midrib, tertiary venation obscure, upper surface glabrous at maturity, coarsely pitted, lower surface with long, strongly appressed, silky hairs on midrib and lateral veins, these also on blade but this eventually glabrous; petiole c. 40-45 mm, hairy as midrib. /nflorescence an axillary very congested cyme, flowers few, peduncle c. 2 mm. Bracts c. 5.5—7 x 1.8—2 mm, persistent but inconspicuous, lanceolate, acuminate, both surfaces densely appressed-pubescent. Pedicels c. 8 mm, puberulous. Calyx 5-lobed almost to base, tube c. 0.5 mm long, lobes subequal, 5—8 x 1.2 mm (in flower and fruit), deltoid, outside densely appressed-pubescent, inside minutely gland-dotted. Corolla c. 30-35 mm long, white or creamy white outside, 2 orange-yellow bars in throat, outside puberulous, hairs acute, tube c. 22—28 mm, lower half cylindric, upper part expanded, two upper lobes c. 5 x 5 mm, lower lip 3-lobed, median lobe c. 6 x 7 mm, all lobes subrotund, glandular-puberulous inside and short way down tube. Stamens inserted c. 16 mm above base of tube, filaments c. 5 mm, curved and twisted once, glandular-puberulous at apex, anthers c. 2 x 1.5 mm, cohering face to face by a small ligature, connective glandular-puberulous; lateral staminodes c. 2 mm, posticous staminode not seen. Disc c.1.5x 1.8mm, cupular. Ovary c.6x 1.2 mm, very minutely papillose, coma of hairs at apex. Style c.6 mm elongating to c.10 mm, glandular-puberulous. Stigmatic lobes c.3x 2mm, clavate. Fruit 25-48 x 5 mm, pericarp smooth. Seeds c. 0.3 x 0.2 mm, testa red-brown.
Notes: In his original description of Cyrtandra hoseana, Burtt cited Burtt B2212 (above) as that species, but in the herbarium he later segregated it and further collections as differing in their narrower leaves. Not only are the leaves narrower, they are also shorter (170—275 x 36-60 mm versus 255-320 x 85-95 mm) and there are significantly fewer lateral veins than in C. hoseana (7-10 versus 12-15); also, the leaf tips are acuminate (not abruptly acute). They may differ further in the length of the calyx lobes (5—8 mm versus 2.2—3.5 mm, but see comments below). As in C. hoseana the tertiary veins are scarcely visible on the lower surface of the leaf; anatomically the leaves are similar, having tracheoids with reticulate bars in the hypodermis and polymorphic sclereids in the mesophyll.
Cyrtandra angustielliptica is known only from the Melinau Gorge (in Gunung Mulu National Park) at c.100 m above sea level. A plant with leaves exactly like those of C. angustielliptica in all morphological details has been collected several
58 Gard. Bull. Singapore 55 (2003)
times on the extreme headwaters of the Balleh, at c.1000—1700 ft altitude on a ridge between Sungai Balang and Sungai Balleh [1°35'N 114°30'E at 1200 ft by Anderson & Paie S28329 (E); at c.1000 ft by Anderson & Paie 28739 (E); at c.1000 ft by Anderson $2874] (E) and at 1700 ft by Paie S28409 (E, L)]. However, these specimens differ anatomically in having no sclereids in the spongy mesophyll (a character possibly of little taxonomic significance), and the calyx lobes are 2.5—5 mm long (not 5—8 mm), corolla 20—25 mm (not 30—35 mm), filaments and anthers glabrous (not glandular), and the disc unilateral (not cupular).
Other specimens examined: Sarawak. Gunung Mulu National Park, Melinau Gorge, Burtt B8274 (E); Melinau Gorge, c. 100 m, Nielsen 537 (E).
11. Cyrtandra insolita Hilliard & B.L.Burtt sp. nov. nullae arcte affinis; pedunculo longo axillari bracteis duabus flores breviter pedicellatos 1—3 involucrantibus facile recognoscenda.
Type: Sarawak, Kapit, Upper Rajang [“Rejang’] River, 1929, Clemens 21236 (holo K).
Shrub (fide collector), 3 stems seen, 170-250 mm long, woody, leafy only at apex, young parts densely pubescent. Leaves opposite, strongly anisophyllous, reduced leaves stipule-like, c. 20 x 3 mm at base, linear-lanceolate, villous; developed leaves few, c.160—190 x 55 mm, elliptic, apex long-acuminate, base cuneate, decurrent down petiole, margins obscurely serrulate, lateral veins c. 20 on each side of midrib, tertiary veins invisible, upper surface silky villous at first, soon glabrescent, coarsely pitted, lower surface densely pubescent, hairs very delicate, c.4mm long on midrib, shorter on blade; petiole c.10mm long, villous. /nflorescence mainly in axils of fallen leaves, 1—3-flowered, flowers springing from apex of peduncle and enfolded by bracts, peduncle 15-45 mm long, villous. Bracts 2, persistent, foliaceous, c. 35-45 x 10—22 mm, lanceolate, long-acuminate, 5-nerved, both surfaces villous; bracteoles wanting. Pedicels 4-5 mm, pubescent. Calyx 5-lobed almost to base, lobes subequal, 2 x 1.5 mm, deltoid, outside pubescent, inside gland-dotted. Corolla not seen. Anthers (in very young bud) 1.5 x 1.1 mm, glabrous. Gynoecium very young, but stigma clearly bilobed, disc cupular. Fruit c. 30 x 5 mm, pericarp smooth (young fruits retain an apical coma). Seeds c. 0.3 x 0.25 mm.
Notes: Cyrtandra insolita (insolitus = extraordinary) is highly distinctive among other species in this Section, from which the pair of foliaceous bracts enfolding one to three flowers at the apex of the long peduncle immediately mark it out. The other species all have few- to many-flowered cymes. Also, these species are unbranched herbs of the forest floor; Mrs Clemens described her plant as ‘shrub in thickets’.
Cyrtandra sect. Pleuroschisma aby
12. Cyrtandra sp. nov.
Unbranched herb, stem woody, height unknown, 10 mm diam., pilose, hairs coarse, spreading, red-purple, as on other vegetative parts. Leaves opposite, strongly anisophyllous, few at apex of stem, reduced leaves stipule-like, c. 75 x 6 mm at base, linear-lanceolate, long-acuminate, pilose; largest developed leaves 300-350 x 85-110 mm, elliptic, apex acuminate, base narrowly cuneate, decurrent, margins finely and remotely toothed (teeth being extensions of the veins beyond the margins), lateral veins 15—16 on each side of midrib, tertiary veins coarsely reticulate, upper surface glabrous, pitted, lower surface with short, fine appressed hairs on blade, long coarse spreading hairs on midrib; petiole c. 45—50 mm long, hairy as midrib. /nflorescence a congested several-flowered cyme, solitary in axils of fallen leaves, peduncle c. 6-12 mm long, pubescent. Bracts persistent, c. 18 x 3 mm, lanceolate, acuminate, midrib visible on lower surface, both surfaces densely pubescent; bracteoles similar but smaller, a pair subtending each pedicel. Pedice/l c. 20 mm long (in fruit), densely puberulous. Calyx 5-lobed almost to base, tube c. | mm long, lobes subequal, c. 10 x 1.5 mm (in fruit), narrowly deltoid, outside densely puberulous, inside minutely gland-dotted. Corolla not seen. Fruit c. 30 x 5.5 mm, pericarp smooth, remains of coma visible at apex. Seeds not fully ripe.
Notes: This plant needs to be re-collected. Gunung Gaharu lies c. 96 km SE of Kuching on the road to Serian. The specimen seen is a unicate, in fruit but without flowers. It much resembles Cyrtandra sarawakensis, particularly var. longipilosa, but is at once distinguished by the reticulate, not subscalariform, tertiary venation of the leaves, and by its much narrower bracts.
Specimen examined: Sarawak, Gunung Gaharu, c. 2000 ft., Burtt 2658 (E).
Acknowledgements
We thank the curators of the herbarium at Royal Botanic Gardens, Kew (K), Botany Dept., Kyoto University (KYO), the Nationaal Herbarium Nederland, Universiteit Leiden Branch (L) for the invaluable loan of specimens. We are also grateful to several of our colleagues: Frieda Christie, electron microscope unit; Mary Mendum (Bates) for art work; Adele Smith for stepping into the breach in a typing crisis; and Debbie White, photographic unit. We also thank the Regius Keeper, Steve Blackmore, working facilities at the Garden.
60 Gard. Bull. Singapore 55 (2003)
References
Bokhari, M.H. and B.L. Burtt. 1970. Studies in the Gesneriaceae of the Old World XXXII: Foliar sclereids in Cyrtandra. Notes Royal Botanic Garden Edinburgh. 30: 11-21.
Burtt, B.L. 1970. Studies in the Gesneriaceae of the Old World XX XIII: Some species of Cyrtandra chiefly from Borneo. Notes Royal Botanic Garden Edinburgh. 30: 23-42.
Burtt, B.L. 2001. A survey of the genus Cyrtandra (Gesneriaceae). Phytomorphology Golden Jubilee Issue 2001: Trends in Plant Sciences. pp. 393-404.
Garden’s Bulletin Singapore 55 (2003) 61-63
Begonia peninsulae (Begoniaceae), a Confused Species from Malesia
RUTH KIEW Singapore Botanic Gardens, Singapore 259569 Abstract
Begonia peninsulae Irmsch. subsp. peninsulae from Trengganu, Peninsular Malaysia, is synonymous with B. rajah Ridl. and B. peninsulae subsp. tambelanense Irmsch. is raised to specific rank as B. tambelanense (Irmsch.) Kiew.
Introduction
Irmscher (1929) recognised two subspecies within his Begonia peninsulae: ssp. peninsulae from Trengganu and ssp. tambelanense from the Tambelan Islands. These were both based on a mixture of plants cultivated in the Singapore Botanic Gardens and the Penang Botanic Garden. The ones collected from the Singapore Botanic Gardens in Ridley’s time (one from Trengganu; the other from the Tambelan Islands) have a clear provenance. However, later collections are without provenance and, in the case of those from Penang, are from batches that include many exotic ornamental begonias. Indeed, there is no evidence to suggest that they are plants of wild-collected origin. In addition, these two subspecies are clearly not conspecific as one has a cordate leaf and the other a peltate one.
Subspecies peninsulae
Irmscher cited one specimen from Trengganu (which was spelt ‘Tringganu’ on the label), which is without a number or information on the date and collector. The botanical name is written as ‘Begonia raja’. ‘Ridley’ has been added as the collector in a different hand after Irmscher’s publication.
This specimen matches those of Begonia rajah Ridl. in characters of the leaf, inflorescence and fruit. Despite the leaf being exceptionally large (11.5 x 16 cm as opposed to 5.5—7.5 x 5.5—7.5 cm in wild plants of B. rajah), the leaf base, like B. rajah, is deeply cordate and, even in the dried state, the variegation is conspicuous with the veins lighter than the rest of the lamina. This specimen might be one of the
62 Gard. Bull. Singapore 55 (2003)
plants originally collected in Trengganu (which Ridley spelt as Tringganu) or its progeny that was cultivated in the Botanic Gardens Singapore from where it was introduced into the ornamental trade in England. B. peninsulae ssp. peninsulae is therefore reduced to synonomy with B. rajah.
The other specimens of ssp. peninsulae cited by Irmscher are very different in that they are all peltate (B. rajah never has peltate leaves). These specimens, Burkill Begonia No. 16 July 1917 (SING), Burkill 6114 31 Aug 1920 (SING), were cultivated in the Penang Botanic Garden and have no provenance. In leaf size, shape, colour, surface (‘coppery and bullate’) and margin they are identical to Begonia goegoensis N.E. Brown from Sumatra, which is one of the few Asian begonias that is easy to grow under local conditions.
Begonia rajah Rid.
Gard. Chron. II, 16 (1894) 213 & Fig. 31. Type: Native Collector s.n. 1892 “Tringganu’ (Trengganu) (holo K ex SING’)
Synonym nova: Begonia peninsulae Irnscher ssp. peninsulae, Mitt. Inst. Allg. Bot. Hamburg. 8 (1929) 98. Type: ‘Tringganu’ without collector, number or date (K ex SING’).
Subspecies tambelanense
Irmscher described this subspecies (a begonia with peltate leaves) from a collection by Ridley made in 1895 from a plant that was cultivated in the Botanic Garden Singapore. This, in turn, was originally collected by Pereira from the Tambelan Islands, NW of Pontianak, Indonesian Borneo.
It is very different from the specimen of his subsp. peninsulae (=B. rajah) from Trengganu as not only are the leaves peltate with a long acumen but the petiole and lower surface of the veins are conspicuously hairy. (Those of B. rajah are deeply cordate with a short acumen and are more or less glabrous). Again, Irmscher included plants of unknown provenance collected from both Gardens — Nur Begonia No. 2 (SING), Nur Begonia No. 3 (K, SING) both dated March 1917 from the Botanic
‘When Irmscher monographed the begonias of Peninsular Malaysia he borrowed the entire collection of the Singapore Botanic Gardens (SING) and these he annotated and cited in his monograph. He did not consult the collection at Kew (K) and none of their specimens is cited by him. A considerable number of unicate types are no longer to be found at SING but are now at K. We have no record of how they came to be transferred but they are definitely the Singapore types as they bear Irmscher’s original annotations. These are cited as K ex SING in this account.
Begonia peninsulae 63
Gardens Singapore; and Burkill Begonia No. 1 (SING) July 1917, Burkill 6115 (SING) 31 Aug 1920 from the Penang Botanic Garden. These are among a gathering of begonias by Burkill that included ornamental begonias made over 20 years after the original Pereira specimen was introduced into Singapore. It is unlikely that the Tambelan plants had survived so long.
These garden plants differ from the original collection by Ridley in their more densely pilose petioles and distinctly toothed leaf margin. Irmscher had already raised the possibility that these were pot plants (Topfpflanzen) and that they differed from the Tambelan plant in leaf size (6.5—17 x 8.5—24 cm), margin and acumen length. The Tambelan plant is therefore regarded as a distinct species and is here raised to specific rank.
To add confusion, Irmscher also recognised a hybrid between the two subspecies, Burkill 6113 31 Aug 1920 (SING), because he considered it was intermediate between the two subspecies. This specimen is less hairy and has a less conspicuously toothed margin than Burkill 6/15 but it does not resemble B. goegoensis (Burkill 6114) at all.
Begonia tambelanense (Irmsch.) Kiew, stat. nov.
Basinym: Begonia peninsulae Irmsch. ssp. tambelanense Irmsch. Mitt. Inst. Allg. Bot. Hamburg. 8 (1929) 100. Type: Ridley s.n. 31 Aug 1895 Botanic Gardens Singapore originally collected by Pereira from Tambelan Is. (holo K ex SING’).
Reference
Irmscher, E. 1929. Die Begoniaceen der Malaiischen Halbinsel. Mitteilung aus dem Institut fur allgemeine Botanik in Hamburg. 8: 86—160.
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Garden’s Bulletin Singapore 55 (2003) 65—68&
A New Species and a New Combination in Bornean Kopsia (Apocynaceae: Apocynoideae)
DAVID J. MIDDLETON
Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138, U.S.A.
Abstract
A new species, Kopsia rajangensis D.J.Middleton, is described and a new combination, Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, is made.
Introduction
As preparation for an account of the Apocynaceae for the Tree Flora of Sabah and Sarawak, it is necessary to describe a new species of Kopsia and reduce Kopsia mitrephora Sleesen to a variety of Kopsia pauciflora Hook.f.
Kopsia rajangensis D.J.Middleton, sp. nov.
Frutex 1.2-4.6 m altus, ramis glabris. Folia elliptico-oblonga, 13.4—32 x 3.8—9.6 cm, nervis 9—25 paribus. Corollae tubus 21.5—25 cm, lobis 11-14 x 2.7-4.7 mm. Typus: Borneo, Sarawak, Kapit, Upper Rejang River, Clemens & Clemens 21221 (holo MO; iso A, BM, BO, NY, SAR)
Small tree or shrub, 1.2—-4.6 m tall. Branchlets glabrous, not or sparsely lenticellate, terete. Leaves: petiole 4-9 mm long, glabrous; blade papery to subcoriaceous, elliptic or oblong, apex caudate, base obtuse to cuneate, 13.4—32 x 3.8—9.6 cm, 2.64.3 times as long as wide, midrib shallowly sunken or raised and with a central groove above, secondary veins 9—25 with 6—25 mm spacing, 40—60° from midrib, prominent or flat above, prominent beneath, clearly discernable from tertiary venation above and beneath, straight or slightly ascending near margin, tertiary venation prominent above, prominent or flat beneath, reticulate or sub-perpendicular to midrib and oblique to secondary veins, intramarginal vein looped and inset from margin, glabrous above and beneath. Inflorescences dichasial and then with cincinnate branches, 4-15 cm long with axes 14-140 x 1.4—2.1 mm, glabrous to densely puberulent; peduncle 2—76
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x 1.7—3.7 mm; pedicel c. 4 mm long; bracts persistent in inflorescence. Sepals ovate, apex rounded, 1.5—1.7 x 1.1—1.4 mm, 1.2—1.4 times as long as wide, ciliate, otherwise glabrous outside and inside. Corolla white; tube 21.5—25 x c. 2.3 mm, pubescent around stamens and slightly beneath inside, throat pubescent, glabrous outside, 1.8—2 times as long as lobes, 12.6—16.7 times as long as calyx; lobes elliptic or oblong, apex obtuse or acute, 11—14 x 2.7-4.7 mm wide, 34.5 times as long as wide, not ciliate, glabrous outside and inside. Stamens 17-21 mm from corolla base which is 0.79-0.84 of corolla tube length; anthers 1.7—2 x 0.6—0.8 mm, 2.5—2.8 times as long as wide, 1.1—2 mm from corolla throat; filaments c. 0.7 mm long. Ovary 0.9 mm high, glabrous to densely pubescent; style 20 mm long; pistil head 0.8 mm long. Fruits falcate with small blunt projection near the base, 15—16 x 4 x 6.5—7 mm; projection 34 mm long.
Distribution: Borneo - SARAWAK: endemic to the Rajang River area. Habitat: In hill forest and disturbed primary forest on rich clay soil.
Notes: Merrill (1921) compared a specimen of this species, Haviland 3042, to Kopsia macrophylla Hook.f. but offered no further identification. Timmerman- Van der Sleesen (1959) was the first to note the peculiarity of the material and compared it to Kopsia lapidelecta Sleesen and Kopsia tenuis Leenh. & van Steenis, again without making a recommendation as to what it might be or describing a new species. Anderson (1980) and Ashton (1988) called this species Kopsia larutensis King & Gamble, a species known only from Peninsular Malaysia. Kopsia rajangensis differs from K. larutensis in its larger inflorescences, much larger flowers and relatively wider corolla lobes. Markgraf (1972) cited the specimens under Kopsia arborea Blume. As can be seen, there has been remarkably little agreement on the identification of the species around Kapit on the Rajang River, which the Clemenses spelt ‘Rejang’. Part of the problem has been the lack of good material.
This new species is probably related to Kopsia lapidelecta and Kopsia sleeseniana Markgr. It differs from the former in its secondary venation, intramarginal vein and larger flowers, and from the latter in its smaller flowers and also its venation.
Specimens examined: Sarawak: Kapit, Upper Rajang River, Clemens & Clemens 21211 (SAR), Clemens & Clemens 21221 (A, BM, BO, MO, NY, SAR); Pelagus Rapids, Ashton $17797 (L); Rajang River, Haviland 3042 (BM, SAR, SING); Bukit Raya, Smith S27738 (SAR).
Kopsia pauciflora Hook.f. var. mitrephora (Sleesen) D.J.Middleton, comb. nov.
Basionym: Kopsia mitrephora Sleesen, Blumea 10 (1960) 136. Type: G.H.S. Wood SAN 16118. Borneo, Sabah, Lahad Datu, Path between Sungai Sabahan and Sungai
Bornean Kopsia 67
Dok (holo L; iso BRI).
Notes: Kopsia mitrephora was formally described by Timmerman-Van der Sleesen (1960) following on from her earlier, but not validly published, account of the species (1959). In the earlier work she distinguished K. mitrephora on the basis of its stamens inserted in the lower part of the corolla tube, a character unusual in the genus, and on its delicate leaves. However, with the many more specimens available now, it is clear that the vegetative characters in this species are extremely variable and closely parallel the vegetative variation found in K. pauciflora. These variations include the size of the wings or angles on the stems, the thickness of the leaves, the length of the petiole and the degree of prominence of the venation. Despite long attempts to put some sort of order to this variation, and indeed there are character states concentrated in some geographical areas, no further taxa could be distinguished in either species. In addition, it became increasingly clear that the only character to distinguish K. pauciflora and K. mitrephora was the site of insertion of the stamens and that otherwise both species were remarkably similar, including in fruit characters. Therefore, varietal status is appropriate for Kopsia pauciflora var. mitrephora.
Acknowledgments
The study was partly carried out under the sponsorship of The Tree Flora of Sabah and Sarawak Project and funded by the Ministry of Science, Technology and Environment of Malaysia through IRPA Funding No. 30—08—01—01-005. The curators of the herbaria at BM, BO, L MO, NY, SAR and SING are thanked for either a loan of material or for their hospitality whilst the material was consulted.
References
Anderson, J.A.R. 1980. A Checklist of the Trees of Sarawak. Forest Dept. Sarawak, Malaysia.
Ashton, P.S. 1988. Manual of the Non—dipterocarp Trees of Sarawak. Forest Dept. Sarawak, Malaysia.
Markgraf, F. 1972. Florae Malesianae Praecursores LIII. Apocynaceae II. 6. Urnularia, 7. Willughbeia, 8. Kopsia. Blumea. 20: 407-425
Merrill, E.D. 1921. A Bibliographic Enumeration of Bornean Plants. Fraser & Neave, Singapore.
Timmerman-Van der Sleesen, E.H.L. 1959. Preliminary revision of the Malaysian species of the genus Kopsia (Apocynaceae). Flora Malesiana Miscellaneous Records. 1: 1-15
68 Gard. Bull. Singapore 55 (2003)
Timmerman-Van der Sleesen, E.H.L. 1960. In C.G.G.J van Steenis, Miscellaneous Botanical Notes X. Blumea. 10: 136—139
Garden’s Bulletin Singapore 55 (2003) 69-72
New Species and Varieties of Symplocos (Symplocaceae) from Borneo
K.G. PEARCE
32, Lorong Kumpang 4, Kuching, Sarawak, Malaysia
Abstract
A new species of Symplocos, S. buxifolioides K.G.Pearce and two varieties, S. iliaspaiensis Noot. var. pedunculata K.G.Pearce and S. laeteviridis Stapf var. alabensis K.G.Pearce are described from Borneo.
Introduction
In his account of the Symplocaceae for the Flora Malesiana region, Nooteboom (1975, 1977, 1984, 1986, 1989) recognised 30 species with 19 varieties of the genus Symplocos Jacq. in Borneo. In the process of revising and preparing the manuscript for the Tree Flora of Sabah and Sarawak, one new species, Symplocos buxifolioides, and two varieties, S. iliaspaiensis var. pedunculata and S. laeteviridis var. alabensis are recognised. Descriptions and brief notes on the geographical distribution and morphological characters distinguishing these new taxa from those of previously known are given.
1. Symplocos buxifolioides K.G.Pearce, sp. nov.
Symplocos buxifoliae Stapf similis a qua differt ramulis distincte modo gallice et anglice ‘zigzag’ et margine foliae revoluto praeter interdum ad apicem et e basi foliorum non altae obscure denticulatae. Typus: Chew & Corner RSNB 5882, Borneo, Sabah, Mt. Kinabalu, Mesilau (holo SING; iso SAN).
Tree to 10 m tall, 10 cm diam. Twigs weakly, irregularly curved, smaller ones distinctly zig—zag with nodes closely set, slender, up to 4 mm diam., dark brown to black; young parts longitudinally ridged and sparsely appressed long—pilose; older parts becoming horizontally cracked, glabrescent, without prominently raised leaf scars. Leaves spirally arranged, coriaceous, glabrous above, hairy to virtually glabrous below, drying olive—brown above, yellow—green below; blade broadly ovate—elliptic, 2.54.5 x 1-3
70 Gard. Bull. Singapore 55 (2003)
cm, base shortly attenuate, margin revolute except sometimes at apex, shallowly, obscurely toothed from near base, apex acute, shortly apiculate; midrib narrowly channelled above; /ateral veins (6—)7-& pairs, joining to form an intramarginal vein; intercostal venation reticulate, raised above and prominent below; petiole 3.5—5 mm long, sharply flanged to base. Inflorescences racemous, borne in the axils of upper leaves and below on leafless twigs; axis 2—5(—9) mm long, shortly, moderately dense appressed—pilose; bracts early caducous. Flowers 1-3; pedicels to 0.5 mm long; calyx tube c. 0.3 mm long, lobes broadly ovate, chartaceous, ciliate almost to apex, moderately dense to sparsely appressed—hairy, c. 1 mm long; corolla white, glabrous, 3.5—4 mm long; ovary sparsely to densely appressed hairy, 1.5 mm high. Fruits solitary on a pedicel to 9 mm long, ovoid to curved ellipsoid, 1-1.5 x 0.5—0.7 cm, hardly to shallowly ridged, sparsely appressed—pilose to glabrescent, drying yellowish brown, with persistent and erect calyx lobes.
Distribution: Endemic in Borneo. Known only from Mt. Kinabalu, Sabah. Ecology: Upper montane forest at 2400-3660 m altitude.
Other specimens examined: SABAH. Mt. Kinabalu: Hotta 3840 (SAN), Chew & Corner RSNB 5911 (SAN), Mikil & Aban SAN 46593 (SAN) and Aban et al. SAN 54254 (SAN).
Notes: Symplocos buxifolioides is similar to S. buxifolia Stapf but differs from the latter in its slender twigs, which are distinctly and shortly zig—zag between the closely set nodes; young parts of the twigs sparsely appressed pilose; broadly ovate—elliptic leaves (not obovate—elliptic as in S. buxifolia) with revolute and closely, obscurely denticulate margin from near the base, acute to shortly apiculate apex (not obtuse to rounded), and 6—8 pairs of lateral veins (not 4—6); inflorescences borne on leafless twigs as well as in the axils of upper leaves; calyx tube c. 0.3 mm long, ciliate almost to the apex (not sometimes ciliate at base), and moderately densely to sparsely appressed—hairy (not glabrous); and sparsely to densely appressed—hairy ovary (not glabrous). This species is named for its close morphological similarity to S. buxifolia.
2. Symplocos iliaspaiensis Noot. var. pedunculata K.G.Pearce, var. nov. A var. iliapaiense differt racemis base ramosis ad 6.5 cm longis floribus (fere) sessilibus ad 2 mm pedicellatis. Typus: Fedilis SAN 95642, Borneo, Sabah, Kalabakan
district (holo SAN; iso K, KEP, L, SAR, SING).
Tree to 13 m tall. Leaves 7.5—20 x 2.5—6 cm, base cuneate, apex with acumen to 1.4 cm long. Racemes basally branched, to 6.5 cm long. Flowers virtually sessile or
Symplocos from Borneo al
with short pedicel to 2 mm long.
Distribution: Endemic in Borneo. Sabah (Kalabakan, Sandakan and Tawau districts) and Brunei Darussalam.
Ecology: In primary and logged—over mixed dipterocarp forest at altitudes to 80 m, on flat or undulating lands and ridges.
Other specimens examined: SABAH. Sandakan district, Sigin SAN 56798 (KEP, SAR, SING), Leopold & Kodoh SAN 81396 (SAN, SAR), Leopold & Kodoh SAN 81424 (SAN, SAR); Lahad Datu Sinanggul SAN 56991 (SAN, SAR); Kinabatangan Meijer SAN 136314 (SAN), Pensiangan district Goh SAN 141166 (KEP, SAN). BRUNEI: Belait Wong WKM 1079 (SAN).
Notes: This new variety may reach 13 m tall (not 3.3 m tall as in the typical variety). It is vegetatively very similar to var. iliaspaiensis but can be distinguished by its basally branched raceme or unbranched spike or fascicle of racemes 1.5 to 6.5 cm long bearing virtually sessile to shortly pedicellate (to 2 mm long) flowers. In contrast, var. iliaspaiensis has fascicles of sessile flowers with the axis entirely absent. This variety is named for its pedunculate inflorescences.
3. Symplocos laeteviridis Stapf var. alabensis K.G.Pearce, var. nov.
Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. similis praeter ramunculos graciliores cum paginis infernis foliorum omnino glabros, foliis olivaceis, parvioribus, lanceolatis, ad 6.3 cm longis et 2.2 cm latis. Typus: Madani & Majawat SAN 119252. Borneo, Sabah, Penampang district, Gunung Alab (holo SAN, iso KEP).
Glabrous treelet to 2 m tall. Twigs slender, glabrous. Leaves drying olive green; blade glabrous below, lanceolate, to 6.3 x 2.2 cm, base cordate; lateral veins 10 pairs; petiole c. 1 mm long.
Distribution: Endemic in Borneo. Sabah, known only from Gunung Alab (the type specimen and Kamaruddin KMS 1431 (SAN, UKMS).
Notes: The new variety is similar to Symplocos laeteviridis Stapf var. mjobergii (Merr.) Noot. except that the twigs and lower leaf surface are entirely glabrous and it has more slender twigs and smaller (to 6.3 x 2.2 cm) lanceolate leaves that dry olive green. It is named for the locality where it was collected.
7? Gard. Bull. Singapore 55 (2003)
Acknowledgements The work was carried out under the Tree Flora of Sabah and Sarawak Project and funded by IRPA Research Grant 30—08—01—01—005. The curators of the herbaria at K, KEP, SAN, SAR and SING are thanked for permission to examine specimens in their care as is H.P. Nooteboom for translating the diagnoses into Latin.
References
Nooteboom, H.P. 1975. Revision of the Symplocaceae of the Old World, New Caledonia excepted. Leiden Botanical Series. 1: 33-335.
Nooteboom, H.P. 1977. Symplocaceae. Flora Malesiana. I, 8: 205-274. Nooteboom, H.P. 1984. Symplocos (Symplocaceae) from Bukit Raya. Blumea. 30:73-74. Nooteboom, H.P. 1986. Additions to Bornean Symplocaceae. Blumea. 31: 277-279.
Nooteboom, H.P. 1989. Addenda, corrigenda et emendanda. Flora Malesiana. 1,10: 719.
Garden’s Bulletin Singapore 55 (2003 ) 73-88
Five New Begonia species (Begoniaceae) from the Niah National Park, Sarawak, Malaysia
KATHARINE G. PEARCE
DANIDA/SWMPI Project, Sarawak Forest Department, Kuching, Malaysia
Abstract
Five new Begonia species, Begonia kachak K.G.Pearce, B. kasutensis K.G.Pearce, B. niahensis K.G.Pearce, B. stichochaete K.G. Pearce and B. subisensis K.G.Pearce, were collected from limestone habitats in the Niah National Park. A key, descriptions and illustrations are provided
Introduction
Sarawak’s varied geology includes a number of isolated limestone outcrops dating from a range of geological epochs and varying in size. In north Sarawak the large Melinau formation falls partly within the Gunung Mulu National Park, while in west Sarawak there are a number of limestone outcrops around Bau and Serian and south towards Tebedu. Other exposures are scattered 1n the interior of Sarawak, for example, at Bukit Sarang (Tatau), Ulu Kakus and in the Middle Baram. Between Bintulu and Miri, in the Niah area about 16 km from the coast, there is an isolated limestone massif known as the Subis massif. In 1974, this massif and surrounding forest was constituted as the Niah National Park.
The Bau limestone outcrops, being relatively accessible from Kuching, have been a favourite destination for plant collectors since the mid 19" Century and the Mulu limestone was the focus of intensive botanical collection during the Sarawak Forest Department and Royal Geographic Society Expedition in 1978. In contrast, the Subis massif was, until recently, somewhat neglected by plant collectors. G.D. Haviland, Curator of the Sarawak Museum, Kuching, and C. Hose, District Officer in Baram and Sibu, together made the first plant collections there in 1894. In 1932, P.M. Synge, a member of the 1932—-33 Oxford University Expedition to Sarawak, visited Subis, while in 1954, W.M.A. Brooke and Ahmad independently made collections in the area under the Sarawak Museum label. In 1962, B.L. Burtt with P.J.B. Woods and Chew W.L. collected there as did H.P. Fuchs in 1963 and G.
74 Gard. Bull. Singapore 55 (2003)
Alphonso and Samsuri Ahmad of the Singapore Botanic Gardens in 1965. The Botany Unit staff, Sarawak Forest Department, made collecting trips to Niah on a number of occasions from the early sixties onwards, including J.A.R. Anderson, who had a particular interest in the limestone flora of Sarawak (Anderson, 1965) and who collected there in 1966 and 1972. Their collections included specimens of the five begonia species described below but which until now remained unnamed.
All five species are endemic to the Subis limestone.
The highest point of the Subis limestone massif is G. Subis. Bukit Kasut and Gua Pangomah are located on the west of the massif. G. Brangin and G. Bekajang are two discrete hills to the north and northeast of the massif, respectively. The latter is the site of the Niah Great Cave and the Painted Cave, which are important archaeological sites.
The botanical field investigations in Niah National Park were carried out as part of the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life Master Plan Implementation through the Improved Management of Totally Protected Areas in Sarawak, Malaysia’ and enabled the five Begonia species to be re-collected from the limestone habitats and for detailed observation to be made.
Key to Begonias in the Niah National Park
la, Leat- ovate, oblong-ovate of Obovate = oo... 22.6 seaeese cena teen eee ib.) Leafwzeniform:to:sub+orbicular, sejcnic. 4 nell \enlineg beta eee 2a... ~Leaf 1.5 times.or moreastlong:as broad auiiiwe elim lab nehenel cee 2b. Leaf less than 1.5 times as long as broad ...................5..2.....+. 2. B. niahensis 3a. Leaf with lines of stiff hairs between veins ...................... 3. B. stichochaete 3b. Leaf without lines of stiff hairs between veins ...................... 5. B. subisensis 4a. Leaf more than 7 cm long and wide .....d28. Jena! dole edi.od bee 4b. Leaf less than 7 cm long and wide ...............ssccsseeeeeeee 4. B. Kasutensis
I. Begonia kachak K.G.Pearce, sp. nov.
Section Petermannia
A Begonia speluncae Ridl. foliis majoribus ob internodia longiora non in fasciculis dispositis nec peltatis nec glabris differt. Typus: Great Cave, Niah, Miri District /.A.R. Anderson, S. Tan & E.Wright S26074 (holo SAR).
Begonias from Niah National Park WS
Figure 1. Begonia kachak
A Habit, B Detail of leaf margin, C Male bud, D Male flower, E Androecium, F Stamens, G Female flower, H Style and stigma. (from $26074).
76 Gard. Bull. Singapore 55 (2003)
Figure 1
Creeping herb or low root climber, roots fibrous, produced at the nodes in contact with the substrate. Stem and petiole with minute appressed to oblique brown hairs; distal end of young petiole, main veins on abaxial surface, margins of young leaf and main vein of stipule with stiff pink-brown trichomes to | mm long. Leafy stem 6-28 cm long, 3 mm diam., little branched, internodes 1—5 cm long. Stipules broadly triangular with a distinct midrib, 4.5 x 7 mm, margin entire, prolonged into an acuminate apex, persistent. Leaves alternate; petiole 1.8 cm in upper leaves, to 17 cm in lower leaves; lamina minutely papillose above and below, upper surface somewhat lustrous, olive- green to yellowish green especially near margins, lower surface pale green or in some specimens brownish pink with pale green veins, oblique, reniform to sub-orbicular, 7.5—12.5 x 7.5—15 cm, base cordate with the basal lobes hardly overlapping, rounded, 0.24.7 cm long, margin irregularly and indistinctly serrate, teeth setose, apex acute to acuminate, venation pinnate-palmate, main veins 4 pairs, branching about half way to the margin, with 2 more veins in the broader and one in the narrower basal lobe, sunken above, prominent beneath. /nflorescences axillary, protogynous, obliquely erect cymose panicles, 8.5—29 cm long, of which peduncle is up to 14 cm long, with one or two female flowers proximally and many male flowers distally; bracts pink, similar to stipules, 6.5 x 3 mm, persistent, bracteoles pale pink-brown, broadly ovate, 4 x 3 mm, apex setose, persistent. Male flowers with pink pedicel to 7.5 mm long, tepals 2, white translucent margins pink before anthesis, minutely papillose, orbicular, 7x 5.5 mm, margin entire, apex rounded, stamens yellow, c. 29, in a spherical cluster, joined in a torus c. 3 mm diam., filaments to 1 mm long, anthers pale yellow, obovate, c. 0.7 mm long, emarginate. Female flowers on pedicels 5.5 mm long, minutely hairy; ovary pink, oblong-obovoid, 10.5 x 7 mm, wings pink, 3, subequal, c. 1.5 mm wide; tepals 5, white translucent or palest pink, with scattered minute, deeper pink hairs, unequal, oblong with rounded apex, to 9.5 x 5.5 mm, margin entire; styles 3, free, each 2 mm to bifurcation from where papillose stigmatic surface forms a continuous twisted band. Fruits with decurved pedicel, 1—1.9 cm long; capsule asymmetric, broadly oblong-elliptic, to 1.9 cm x 1.6 cm, minutely hairy, papillose, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base and apex, two narrower wings to 5 mm wide, broader wing to 7 mm wide, style caducous. Seeds cylindrical, c. 0.3 x 0.2 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (G. Brangin, Niah Great Cave and Gua Pangomah).
Habitat: Locally abundant in the shelter of overhangs in somewhat dry areas or
Begonias from Niah National Park Vl
creeping up rocks at cave mouths, infrequently at level ground in deep shade adjacent to limestone outcrops.
Notes: This is a conspicuous begonia with large shiny leaves, commonly bearing a cluster of delicate, pale pink male flowers. It is easily observed on limestone outcrops along the Plank Walk to the Niah Great Cave and around the cave mouth. It takes its name from kachak, the Malay word meaning handsome.
Specimens examined: SARAWAK: Gunung Subis, Niah B.L. Burtt & P.J.B. Woods B2010 (SAR), Mohidin S2 1603 (SAR); Great Cave K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude $78536 (SAR), Jemree Sabli S89062 (SAR); G. Brangin, Ulu Sg Subis Yii Puan Ching S40168 (SAR), Gua Pangomah J.A.R. Anderson S31691 (SAR, SING), Near Sg Subis K.G. Pearce $89463 (SAR); Niah Caves Alphonso & Samsuri A217 (SING), A222 (SING), A248 (SING).
2. Begonia niahensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia congesta Ridl. habitu erecto vel effuso (non cauli longo), capsula oblongo- ovoidea (non longa oblongaque) differt. Typus: Niah Caves B.L. Burtt & P.J.B. Woods B 2009 (holo SAR).
Figure 2
Herb with stems to 75 cm long, branching near base with some spreading horizontally, some vertical and some drooping and then curving upward. Young stems and petioles with minute, erect translucent hairs. Older stems brown and semi-woody, to 1 cm diam., young stems brownish pink, internodes 2—8.5 cm long. Stipules green flushed brownish pink at centre, plicate, broadly ovate, 14 x 7 mm, margin entire with translucent hairs at base, apex acute, persistent. Leaves spirally arranged; petiole from c. 5 cm long in the upper leaves to 22 cm in the lower leaves, deep brownish pink; lamina medium green above, paler below, with veins on both surfaces deep pink at base and at the point where they branch, pinkish between, minutely papillose with a velvety appearance above, translucent hairy beneath, oblique, broadly ovate, 15—20 x 12—16.6 cm, base cordate with basal lobes not or hardly overlapping, rounded, to 9 cm long, margin irregularly and indistinctly serrate, translucent hairy, teeth with translucent setae, apex acute, venation pinnate-palmate, main veins 4—5 pairs, branching near base, with 2 or 3 smaller veins in broader basal lobe and 1 or 2 in narrower basal lobe. /nflorescences axillary, protogynous, obliquely upright, cymose panicles 3—8 cm long, of which the peduncle is 2.5—-6 cm long, with | or 2 pairs of female flowers at the base with many male flowers tightly congested distally, bracts
78 Gard. Bull. Singapore 55 (2003)
plicate, pale green, broadly ovate, to 2—2.3 x 1.2 cm, persistent. Male flowers opening one at a time, pedicel obscured by congested bracts and male flower buds, tepals 2, white translucent, margins entire, orbicular, c. 6 x 7 mm, apex rounded; stamens yellow > 30, in hemispherical cluster, joined in a torus c. 1 mm diam., filaments c. 0.5 mm long, anthers yellow, emarginate, 0.8 mm long, opening by apical slits. Female flowers with pedicel c. 9 mm long; ovary yellow-green to pink, obovoid, 19 x 7 mm, wings 3, pale green, subequal, 1.5 mm wide; tepals 5, white translucent, subequal, broadly elliptic-obovate, 13 x 7 mm, margins entire except at fringed, obtuse apex, outer surface and margins minutely translucent hairy; styles 3, c. 1 mm long; stigmas bifurcated, stigmatic surface a minutely papillose, orange yellow spiral band. Fruits with a stiff, decurved pedicel, c. 1.5 cm long, pale green flushed pink at base; capsule oblong, to 3.2 x 1.4 cm, sparsely translucent-hairy, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, subequal, thin, broadest at apex, c. 4 mm wide, style caducous. Seeds cylindrical to obovoid, c. 0.3 x 0.2 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo - SARAWAK: endemic in the Subis limestone (G. Brangin, Niah Great Cave and Painted Cave areas).
Habitat: Below cave mouths, rooted in pockets of soil on ledges and in crevices on steeply sloping limestone rock or in light shade under canopy of small trees.
Notes: This robust and handsome begonia has large, asymmetric, velvety hairy leaves that sometimes have a bluish cast. The leaves are positioned more or less vertically against the limestone rock face in somewhat sheltered sites. It can be observed on limestone outcrops along the Plank Walk to the Painted Cave. It has been named for the type location - the Niah Caves. Its habit of drooping and upcurving stems and handsome, velvety leaves held vertically, give this species potential as an ornamental plant.
Specimens examined: SARAWAK: G. Brangin Yii Puan Ching $40166 (SAR); Great Cave J.A.R. Anderson, S. Tan & E. Wright S26075 (SAR, SING), K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude §78537 (SAR); Near Sg Subis K.G. Pearce S89460 (SAR).
3. Begonia stichochaete K.G.Pearce, sp. nov.
Section Petermannia
A Begonia congesta Ridl. foliis angustioribus seriebus setarum inter venas gerentibus, capsulis longioribus oblongis (non oblongo-ovoideis) differt. Typus: Subis, Niah Caves Ahmad 14 (holo SAR; iso SING).
Begonias from Niah National Park
Figure 2. Begonia niahensis
A Habit, B Detail of leaf undersurface, C Detail of leaf margin and upper surface, D Male bud, E Male flower, F Androecium, G Stamens, H Female flower, I Style and stigma from above, J Style and stigma. (A—C from Burtt 2009, D-I from $89460).
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80 Gard. Bull. Singapore 55 (2003)
Figure 3
Erect herb rooting at the base of the stem and at nodes along the horizontal rhizome which creeps below the soil surface and gives rise to shoots at intervals. Indumentum of stiff trichomes to 2.2 mm long, each on a raised conical base, pale brown to pinkish brown to deep brownish red on stem, young petiole and veins on lower and upper lamina surfaces, dense on young stem and petiole, less so on stipule margin and sparse on outer surface of stipule midrib and on the main veins of the upper lamina surface, in a row or rows between each pair of veins and on and between the teeth of the dentate margin, moderately dense on the main veins of the lower leaf surface. Upright stem to c. 42 cm long, 5—9 mm diam., unbranched, internodes 2.5—5 cm long to 8 cm long at base of stem. Stipules oblong-ovate with a distinct midrib, 17 x 11 mm, margin entire, apex setose, caducous. Leaves spirally arranged; petiole 0.7 cm in upper leaves, to 2.5 cm in lower leaves; lamina at first yellowish green then medium green on upper surface (rarely dark blackish-green), lower surface paler yellowish green or olive-green or deep pink or red, asymmetric, irregularly obovate, 11-19 cm long, narrower side to 2.5 cm wide towards apex, curving inwards towards base and hardly or not lobed at base, broader side 6—7.5 cm wide with a cordate base and a rounded basal lobe to 17 mm long, margin irregularly serrate (appearing almost praemorse), each tooth setose, with several setae between each pair of teeth, apex toothed, venation pinnate, c. 4 pairs of veins, branching with another 1-3 veins in basal lobes, impressed above, prominent beneath. /nflorescences axillary, protogynous, erect cymose panicles to 3.4 cm long, of which the peduncle is 2.3 cm, with 1—2 female flowers at base with distally many male flowers crowded in a terminal cluster; bracts similar to stipules, deep red, 13 x 10 mm, persistent; bracteoles pale brown to deep pink, broadly ovate, to 12 x 12 mm, apex setose, persistent. Male flowers witha green to deep pink pedicel with deep pink bristles, 7 mm long; tepals 2 or 4, the outer pair translucent white flushed pink at margin and base, the inner pair translucent white, at first finely hairy on outer surface near base, broadly elliptic, c. 4 x 3 mm, margin entire, apex rounded; stamens pale yellow, c. 32, in an ovoid cluster, joined in a torus c. | mm diam., filaments to | mm long, anthers obovate, to 1.5 mm long, shortly apiculate, opening by terminal pores which develop into slits. Female flowers solitary or in pairs; pedicels 5 mm long, medium pink with minute appressed to slightly oblique hairs; ovary pale pink tinged green with deep pink bristles, oblong-obovoid, to 24 x 16 mm, wings 3, deep pink to translucent white with margins flushed pale pink, subequal, c. 7 mm wide; tepals 5, translucent white flushed pale pink at margins or palest pink, deeper pink at base and on marginal teeth with scattered deeper pink hairs, on outer surface sparsely hispid, oblong with rounded apex, inner two larger, c. 6 x 2-4 mm, margin toothed from near base to fringed apex; styles 3, free to base, 3 mm long, bifurcating; stigmas forming a continuous twisted band, yellowish-green to bright yellow. Fruits with stiff, decurved pedicels, 5—9 mm long; capsule oblong,to 29 x 18 mm,
81
Begonias from Niah National Park
es ARES or fad ony RS EDR Sag z
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82 Gard. Bull. Singapore 55 (2003)
sparsely hispid, minutely papillose, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, subequal, thin, narrowed to base, truncate distally, two narrower wings to 5 mm wide at midpoint and 6 mm at apex, broader wing to 6 mm wide at midpoint to 7 mm at apex, style caducous. Seeds cylindrical-spherical, c. 0.3 x 0.25 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo —-SARAWAK: endemic in the Niah National Park (Kuala Subis, the Sekaloh area and near the trail to Bukit Kasut).
Habitat: This species occurs both in limestone habitats and undisturbed Mixed Dipterocarp Forest on loamy soil or brown clay as well as in secondary forest on alluvial soil. On limestone, it grows on low outcrops with little soil and in soil at the base of limestone cliffs in deep shade and damp conditions, where it is locally frequent.
Notes: This erect begonia is characterised by its bristly leaves bearing rows of stiff hairs between the veins. It grows in shady sites, both on or near limestone outcrops and in forest away from the limestone. The species shows striking variation with the upper leaf surface ranging from mid-green to dark blackish-green. It is named for its bristly leaves.
Specimens examined: SARAWAK: Niah National Park Bernard Lee S40075 (SAR), Kuala Niah near confluence with Sg Subis K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude S78539 (SAR); Route to Bukit Kasut K.G. Pearce & Narawi b. Johari S75597 (SAR); Sekaloh Jemree Sabli S8S9014 (SAR), K.G. Pearce S589267 (SAR).
4. Begonia kasutensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia conipila Irmsch. ex Kiew trichomatibus brevioribus sine basibus conicalibus, foliis suborbicularibus (non valde asymmetricis), punctis maculisque praesentibus, inflorescentiis contra folia insertis differt. Typus: Great Cave, Gunong Subis, Niah, Miri District J.A.R. Anderson S31940 (holo SAR; iso SING).
Figure 4
Creeping herb with fibrous roots produced at the nodes. Stem and petiole with white minutely appressed hairs on distal end of petiole; main veins, lamina undersurface and margin with scattered pale orange-brown hairs to 0.7 mm long, more or less regularly arranged. Leafy stem toc. 15 cm long, 1.5 mm diam., not or hardly branched, internodes to 4 cm long. Stipules oblong-ovate with a distinct midrib, to 9 x 4mm,
Begonias from Niah National Park 83
Figure 4. Begonia kasutensis
A Habit, B Detail of leaf margin, C Leaf showing variegation, D Male bud, E Male flower, F Stamens. (A & B from $31940, C from $27269, D-F from $3/940).
84 Gard. Bull. Singapore 55 (2003)
margin entire, apex setose, caducous. Leaves spirally arranged; petiole 1.5 cm in upper leaves, to 3.8 cm in lower leaves; lamina minutely papillose above and beneath, variegated, upper surface slightly lustrous, olive to dark green with irregular spots and blotches of pale grey-green between veins, lower surface reddish, sometimes light green towards base, slightly asymmetric, sub-orbicular, 4.7—6.2 x 3.9-5.5 cm, base cordate with rounded basal lobes not overlapping, to 8 mm long, margin irregularly, indistinctly and distantly serrulate, apex cuspidate, venation palmate, main veins 2 pairs, the two ‘middle’ ones branching near mid-point or towards margin, with 1—2 veins in each basal lobe, sunken above, slightly prominent beneath. Inflorescences \eaf-opposed, protogynous, racemes of cymules, to 11.5 cm long, including peduncle to 5.7 cm, with a single female flower at base and many male flowers distally; bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale green flushed pink, broadly ovate, 3.5 x 2 mm, apex setose, caducous. Male flowers with pink pedicel to | cm long; tepals 2, palest pink translucent, suborbicular, c. 7 x 4 mm, margin entire, apex rounded; stamens yellow, c. 20, ina more or less hemispherical cluster, joined in a torus 0.3 mm diam., filaments to c. 1 mm long, anthers bright yellow, c. 0.5 mm long, opening by pores, obovate, emarginate. Female flower not seen. Fruits with stiff, decurved pedicel, 7 mm long; capsule suborbicular, 1.6 x 1.2 cm, locules 3, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base, truncate distally, the two narrower wings c. 1.5 mm wide, broader wing c. 2.5 mm wide, style caducous. Seeds cylindrical, c. 0.3 x 0.25 mm, surface areolate, areoles more or less equal sided at base and apex and elongate along seed axis.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great Cave, Bukit Kasut and G. Subis).
Habitat: Infrequent, on limestone rock faces in shaded and dry crevices or in soil between limestone rocks on slopes of limestone hills up to where the canopy cover starts to thin out.
Notes: This decorative begonia is characterised by its diminutive, sub-orbicular variegated leaves that have a pink undersurface, and by its delicate inflorescences. Its small size and elegant features give it potential as an ornamental species. It has been named for Bukit Kasut, one of the locations where it occurs.
Specimens examined: SARAWAK: Subis Ahmad No. 3 (SAR, SING); Niah Cave Ahmad No. 65 (SING); Gunung Subis Jemree Sabli S89049 (SAR); Southern slopes of G. Subis, near Sekaloh River S. Tan & E. Wright § 27269 (SAR, SING); Route to Bukit Kasut K.G. Pearce & Narawi b. Johari S78596 (SAR).
Begonias from Niah National Park 85
5. Begonia subisensis K.G.Pearce, sp. nov.
Section Petermannia
A Begonia pendula O. E. Schulz habitu majore caulibus erectis rigidisque, foliis setis inter dentes setiferos carentibus, fructibus latioribus oblongis differt. Typus: G. Subis ‘in Sekaloh River’, Niah, Miri District /.A.R. Anderson, S. Tan & E. Wright S27574 (holo SAR; iso SING).
Figure 5
Cane-like, semi-herbaceous begonia with woody stems ascending, then held at 45°, drooping at apex, arising at intervals from a woody, branched rhizome c. 7-9 mm diam. bearing roots at the nodes, some stem bases bearing adventitious roots. Stem and petiole with minutely appressed brown hairs. Stems to >76 cm long, from 6 mm diam. at base to 1.5 mm at apex, bearing leaves only near apex, unbranched or with few branches, internodes 1.2 to >5 cm long. Stipules lanceolate, without a distinct midrib, 5.5 x 1.5 mm, margin entire, apex setose, caducous. Leaves spirally arranged; petiole 1—2.5 cm; lamina minutely papillose above and below, upper surface with a reflective sheen, medium to olive-green (those of young vegetative shoots variegated, olive-green with irregular spots of pale grey-green between veins), lower surface dull green or pinkish between veins to deep pink near veins, asymmetric, oblong-ovate, to c. 11.3 x 3.64.1 cm, lower half of leaf somewhat inwardly curved, base cordate with a rounded basal lobe to 1.3 cm long on one side, cuneate or slightly lobed at base on the other side, margin irregularly serrate, apex acute, venation pinnate-palmate, veins 3-4 pairs, branching except for the outermost two or three, flat or very slightly raised above, deep pink and prominent beneath. /nflorescences axillary, protogynous, cymose panicles held obliquely above the subtending leaf, axis pale to deep brownish pink, to 7.5 cm long with | or 2 female flowers at the base and up to 10 male flowers distally or sometimes, in the lower leaf axils, by reduction, only the female flowers present; bracts similar to stipules, 4 x 2.5 mm, caducous; bracteoles pale green flushed pink, lanceolate, 1.5 x 6 mm, apex setose, caducous. Male flowers with pink pedicel to 9 mm long; tepals 2, translucent white to palest pink, some flushed deeper pink at base, elliptic, 6.5 x 4mm, margin entire, apex rounded; stamens yellow, to 26, in a more or less obovoid cluster, joined in a 1 mm-diam. torus, filaments c. 1 mm long, anthers yellow, obovate, emarginate. Female flowers on a peduncle to 3 mm long; pedicels deep pink to 2.5 cm long, glabrous; ovary oblong, truncate, 1.6 x 2 cm, wings 3, subequal, c. 6 mm wide; tepals 5, palest pink flushed deeper pink at base, ovate, the outer two smaller than the inner three, c. 9 x 5 mm, margin entire, apex obtuse, glabrous; styles 3, bifurcated, 2-3 mm long to bifurcation, stigmatic surface orange with fine colourless papillae forming a continuous, twisted band. Fruits with flexuous pedicels, to 3 cm long; capsule oblong, 14 x 23 mm, locules 3, equal, each with 2 placentae, dehiscing between wing and locule, wings 3, unequal, thin, narrowed to base, truncate distally, two narrower wings 10 mm wide, broader wing 11.5 mm wide, style caducous.
86 Gard. Bull. Singapore 55 (2003)
Seeds cylindrical, c. 0.4 x 0.25 mm, surface areolate, areoles more or less equal sided at base and distally, and elongate along seed axis above base.
Distribution: Borneo — SARAWAK: endemic in the Subis limestone (Niah Great Cave and Bukit Kasut).
Habitat: Locally frequent on limestone rock faces and the steep slopes of limestone hills in deep litter layer or between limestone rocks where the canopy cover starts to thin out.
Notes: The leaves of this attractive cane begonia have a reflective sheen on the upper surface and are deep pink below. The young plant has variegated leaves. The inflorescence 1s delicate and, in fruit, the capsule dangles on a long, flexuous pedicel. It has been named for the type locality, G. Subis.
Specimens examined: SARAWAK: Great Cave J.A.R. Anderson $31939, Bukit Kasut K.G. Pearce, Bibian Diway, Saupel Atot & Dami Jude S78538 (SAR).
Acknowledgements
This paper is based on botanical field investigations at the Niah National Park carried out under the Sarawak Forest Department/DANIDA Project ‘Support to Wild Life Master Plan Implementation through the Improved Management of Totally Protected Areas in Sarawak, Malaysia’ (SWMPI). The author is most grateful to Julian T. Inglis, Chief Technical Advisor of the SWMPI Project, who has taken a lively interest in the collection and description of these begonias; the paper would not have been written without the inspiration and support provided by Dr Ruth Kiew, whose knowledge of and interest in begonias prompted a closer look at the species occurring at Niah National Park and who also checked the types at K and SING; and to Dr M.J.E. Coode for translating the diagnoses into Latin, and to Dr A. Radcliffe-Smth for suggesting ‘stichochaete’ as the appropriate name for that species. The author also thanks the staff of the Botany Unit, Forest Research Centre, and the Niah National Park, National Parks and Wildlife Department, Sarawak Forest Department, for assistance during collecting trips; the curator of SAR herbarium for permission to examine specimens; and Joseph Pau for his excellent botanical illustrations.
Begonias from Niah National Park
Figure 5. Begonia subisensis
A Habit, B Juvenile shoot with variegated leaves, C Fruit. (A & C from $27574, B from S78538).
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Reference
Anderson, J.A.R. 1965. Limestone habitat in Sarawak. Symposium on Ecological Research in the Humid Tropics. Kuching, Sarawak. Sarawak Government and UNESCO. pp. 49-57.
Garden’s Bulletin Singapore 55 (2003) 89-95
A New Species of Curcuma L. (Zingiberaceae) from Mizoram, India
J. SKORNICKOVA, M. SABU AND M.G. PRASANTHKUMAR
Department of Botany, University of Calicut, 673 635 Kerala, India.
Abstract
Curcuma rubrobracteata Skornickova, M.Sabu & Prasanthkumar sp. nov. is described from Mizoram, India.
Introduction
Exploration for gingers in Mizoram, NE India, carried out by the authors for the research project ‘Revision of Indian Zingiberaceae’ has brought an interesting species of Curcuma to light.
The genus Curcuma L. naturally occurs in tropical and sub-tropical Asia with a few species extending to Australia and the South Pacific, but economically important or ornamental species are introduced elsewhere in the tropics. According to the latest records, Curcuma has the largest number of species in India, Thailand, Indonesia and China. The total number of species for this genus is still quite uncertain but is estimated at around 100 species (Sirirugsa, 1996; Larsen et al., 1998). From India, 29 species have so far been reported and accepted (Karthikeyan et al., 1989; Mangaly and Sabu, 1993). Several new species of Curcuma L. have been described in last 20 years from India (Skornickova and Sabu, 2002). While working on Indian Zingiberaceae in August and September 2002, the authors visited Mizoram, which is considered as the least surveyed corner of India, where an undescribed species of the genus Curcuma L. subg. Curcuma (Eucurcuma K. Schum.) was collected.
Curcuma rubrobracteata Skornickova, M. Sabu & Prasanthkumar, sp. nov.
Curcumae roscoeanae similis in coma inconspicua, spica c. 10 cm supra terram per vaginam protrudenti, bracteis floriferibus rubris, floribus luteo-aurantiacis, calyce trilobato, rhizomate repenti aromatico, lamina elliptico-lanceolata basi cuneata, infra glabra, supra pubescenti in venis principalibus elevatis differt. Typus: India, Mizoram, Lawngtlai District, on the way to Ngengpui Wildlife Sanctuary — Khomoi, 22° 30'N,
90 Gard. Bull. Singapore 55 (2003)
92° 46'E. 10.1X.2002. Skornickova & Prasanthkumar 86241 (holo MH; iso CALI, K, SING).
Figure 1, Plate 1.
Rhizomatous herb, up to 1.5 m high. Rhizome creeping, slender, 10—30 cm long, c. 1 cm diam., tan outside, scales triangular, papery, light brown, underground quickly decaying, whitish yellow inside, aromatic, smell resembling Kaempferia galanga L., taste very bitter, sessile tubers absent, root tubers 2 x 1.5 cm, white inside, distanced c. 5 cm away from main rhizome on 2 mm thick roots. Leafy shoot up to 1.5 m long, leaves 4—6, pseudostem green, c. 40 cm long. Leaves petiolate, ligule 1 mm long, light green, translucent, petiole green, glabrous 10-45 cm long (lower leaves with shorter petioles). Lamina elliptic-lanceolate 35—60 x 10-16 cm, adaxially green, pubescent along the raised veins, abaxially pale green, glabrous, base attenuate, tip acuminate 2 cm long, midrib green, glabrous. /nflorescence terminal, but protruding through the base of the pseudostem through lateral slits c. 3-10 cm above ground. Peduncle 5—10 cm long, 7 mm diam., whitish, without vegetative bracts. Spike 10 x 7-9 cm. Coma absent. Bracts 20-26 per spike, all fertile, bright red or light red, yellowish or yellowish green towards the base, bract 3.5 x 3.5 cm, glabrous, margin 0.3 mm hairy, hairs 0.1 mm long, subtending a cincinnus of 5—6 flowers, usually only 1—3 flowers per spike open at the same time. Bracteoles one per flower, 2—3.5 x 1-3 cm, hyaline, glabrous, white, translucent with reddish dots on the apical part of the biggest bracteoles. Flowers 6 cm long, yellow-orange, exserted, 1.5—2 cm longer than the bracts. Calyx 1.2 cm, white, translucent, hyaline, glabrous, 3-toothed. Corolla tube 3.74 cm, light orange, glabrous. Corolla lobes light yellow-orange, dorsal lobe 14 x 8 mm fringed by red on upper mucronate portion; lateral lobes 12 x 7 mm. Labellum 15 x 17 mm, periphery yellow-orange, centre deep yellow-orange, obscurely 3-lobed, middle lobe split about 2 mm. Lateral staminodes | x 1.1 cm; yellow-orange, hooded over the anther. Stamen c. 9 mm long, anther versatile. Anther 6 x 2.5 mm, orange. Anther spurs 3 mm long, incurved, orange. Anther thecae white, 5 mm long. Filament yellow-orange, 5 mm long, constricted, 4 mm at base, 2 mm at upper part. Ovary trilocular, 2 x 2.5 mm, pubescent with 0.6 mm long hairs; ovules many. Stigma white, closely appressed within the anther lobes. Epigynous glands 2, pale orange, 4 mm long, 0.9 mm diam. Fruits not seen.
Flowering: August — September.
Distribution: Hitherto known only from the type locality. From personal communication with other specialists we suspect this species to be identical with unidentified plants growing along the Thai-Burmese border.
91
Curcuma rubrobracteata
1
oe ee mee es ee -—
i
Figure 1. Curcuma rubrobracteata
A. Habit; B. Inflorescence; C. Detail of leaf venation (adaxial side); D. Flower and subtending bract; E. Bracteole; F. Part of flower showing lateral staminodes, dorsal corolla lobe and anther; G. Part of flower showing labellum and anterior corolla lobes; H. Flower (side view); I. Calyx; J. Lateral staminode; K. Anther (front); L. Anther (side); M. Stigma; N. Epigynous glands and ovary; O. Ovary (cross section). Based on the type material Skornickova & Prasanthkumar 86241. del. J. Skornickova.
92 Gard. Bull. Singapore 55 (2003)
Habitat: Growing in undergrowth in teak plantations and along roadsides.
Etymology: This gorgeous species takes its name from the striking red bracts of its inflorescence.
Other specimens examined: India, Mizoram, Lawngtlai District, on the way to Ngengpui Sanctuary — Khomoi, 22° 30' N 92° 46' E, 10.[X.2002, Skornickova & Prasanthkumar 86239 (CALI).
Notes: The most obvious characters of this species are the creeping rhizomes, bright red bracts with no distinct coma and, in particular, the position of its inflorescence. Inflorescences with no distinct coma are reported for a few species e.g. Curcuma roscoeana Wall., C. albiflora Thwaites, C. ceratotheca K.Schum. and the recently described C. rhomba J. Mood & K. Larsen, which also possesses a small rhizome with linear growth (Mood and Larsen, 2001). The unique character of C. rubrobracteata is the position of its inflorescence, which is so far unknown in the genus Curcuma.
Many of the earlier taxonomists made attempts to classify the genus into sections based on the position of the inflorescences. Roxburgh (1820) recognised two sections based on the lateral or central inflorescence, while Horaninow (1862) distinguished three sections, namely: I. Exanha (always lateral), I. Mesantha (inflorescence invariably terminal), and III. Amphiantha (inflorescences both terminal and lateral). However, the new species produces an inflorescence about 3—10 cm above ground, which breaks through the pseudostem through lateral slits. This feature has also been observed in the Zingiberaceae in the genus Plagiostachys Ridl., where the inflorescence is borne on a peduncle and projects from the side of the leafy stalk (Ridley, 1899). Smith (1985) pointed out that the inflorescence of Plagiostachys species, although pushed out laterally, is actually terminal on the leaf shoot and so shows affinity to the genus Alpinia Roxb., in which a few species, particularly A. hansenit R.M.Smith and A. havilandii K.Schum., have similar apparently laterally produced inflorescences.
Recently, Kress et al. (2002) carried out molecular studies on phylogeny and classification of the family Zingiberaceae. The results based on analysis of DNA sequences of the nuclear internal transcribed spacer (ITS) and plastid matK regions suggest that the genus Curcuma, as it is accepted nowadays, is paraphyletic with
Plate 1. Curcuma rubrobracteata
1. Detail of flower and spike — seen from above; 2. Detail of flower and dorsal corolla lobe — front view; 3. Detail of flower and bracts — lateral view; 4. Habit; 5. Detail of spike protruding through the pseudostem; 6. Creeping rhizome and root tubers. All photographs are of the type material Skornickova & Prasanthkumar 86241. Photo J. Skornickova.
Curcuma rubrobracteata
mh
94 Gard. Bull. Singapore 55 (2003)
genera Hitchenia, Stahlianthus and Smithiatris, which also share cone-like inflorescences of few flowered, congested bracts, and that genus Curcuma is itself paraphyletic with three groups of species. Since the analysis was done with a limited number of samples (six Curcuma species), the authors recommended that more species of such large genus should be included in future molecular studies to shed more light on the difficult question of generic boundaries and allied genera.
Acknowledgements
The authors thank the Department of Science and Technology, Govt. of India for financial support (Order No. SP/SO/A-20/99 dt. 09.11.2001). We are also indebted to staff of Forest Department of Mizoram for their hospitality and help to reach otherwise inaccessible areas. The senior author thanks the Indian Council for Cultural Relations New Delhi, India, and the Ministry of Education of the Czech Republic for awarding a research fellowship, and to Singapore Botanic Gardens for providing facilities during her repeated visits. We also thank Dr. J. F. Veldkamp, National Herbarium of Netherlands, Leiden University Branch, and Dr. M. Svrcek, National Museum, Prague, for the help with the Latin diagnosis.
References
Horaninow. 1862. Prodromus Monographiae Scitaminearum. Petropoli (St. Petersburg), Russia.
Karthikeyan, S., S.K. Jain, M.P. Nayar and M. Sanjappa, 1989. Florae Indicae. Enumeratio Monocotyledoneae. Botanical Survey of India, Calcutta. Pp. 289-299.
Kress, J.W., L.M. Prince and K.J.Williams. 2002. The phylogeny and a new classification of the gingers (Zingiberaceae): evidence from molecular data. American Journal Botany. 89: 1682-1696.
Larsen, K., J.M.Lock, H. Maas and P.J.M. Maas. 1998. Zingiberaceae. In: K. Kubitzk1. (Ed.) The Families and Genera of Vascular Plants. 4: 474-495.
Mangaly, J.K. and M. Sabu. 1993. A taxonomic revision of the South Indian species of Curcuma L. (Zingiberaceae). Rheedea. 3: 139-171.
Mood, J. and K. Larsen. 2001. New Curcumas from South East Asia. The New Plantsman. 8: 207—217.
Curcuma rubrobracteata 95
Ridley, H.N. 1899. The Scitamineae of the Malay Peninsula. Journal Straits Branch Royal Asiatic Society. 32: 85-184.
Roxburgh, W. 1820. Monandria monogynia. Flora Indica. Serampore, India. Pp. 1-84.
Sirirugsa, P. 1996. The genus Curcuma of Thailand. In: T. L. Wu, Q. G. Wu and Z. Y. Chen (Eds.) Proceedings 2" Symposium on Family Zingiberaceae. Zhongshan University Press, China. Pp. 39-46.
Skornickova, J. and M. Sabu, 2002. The genus Curcuma L. in India: Resumé and Future Prospects. In: A.P. Das (EIndiad.) Perspectives of Biology. Bishen Singh, Mahendra Pal Singh, Dehradun, India Pp. 45-51.
Smith, R.M. 1985. A review of Bornean Zingiberaceae: 1 ( Alpinieae p.p.). Notes Royal Botanic Garden Edinburgh. 42: 261-314.
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Garden’s Bulletin Singapore 55 (2003) 97-111
The Genetic Relations of Musa Species from Mount Jaya, New Guinea, and a Reappraisal of the Sections of Musa (Musaceae)
CAROL WONG!', GEORGE ARGENT?’, RUTH KIEW’, OHN SET? AND YIK YUEN GAN!
'National Sciences Academic Group, National Institute of Education, Nanyang Technological University, 1 Nanyang Walk, 637616 Singapore
"Royal Botanic Garden, Edinburgh, EH3 5LR, U.K.
3Singapore Botanic Gardens, | Cluny Road, 259569 Singapore
Abstract
Molecular analysis using amplified fragment length polymorphism (AFLP) confirms the status of Musa banksii F. Muell. as a subspecies of M. acuminata Colla and shows M. johnsii Argent to be genetically more similar to the Australian M. jackeyi W. Hill than to M. lolodensis Cheesman from New Guinea. In addition, AFLP analysis supports only two sections as genetically distinct, namely Sect. Musa (including Sect. Rhodochlamys) and Sect. Callimusa (including Sect. Australimusa). No material for Sect. [gentimusa was available for study. However, for practical purposes of grouping banana species, four informal groups are recognised — the ‘acuminata’ and ‘ornata’ groups within Sect. Musa, and the ‘coccinea’ and ‘textilis’ groups within sect. Callimusa. A key to the sections and informal groups is provided.
Introduction
Wild bananas are found throughout Asia and Malesia extending into Australia and the Pacific. They have become an increasingly conspicuous element of the vegetation as they invade forest margins along logging roads and openings in forest. As roads penetrate deeper into forests, new banana species are coming to light.
Much still remains to be discovered about the relationships between wild banana species, which have been grouped into five sections (Cheesman, 1947; Argent, 1976) based on chromosome number and morphology. However, some of the species recently described do not fit comfortably into these sections (Argent, 2000, 2001), which calls into question their taxonomic validity. Also, molecular studies indicate that not all the sections are genetically uniform and distinct from each other (Jarret &
98 Gard. Bull. Singapore 55 (2003)
Gawel, 1995; Wong et al., 2002).
Bananas present a challenge to the collector who wishes to turn them into herbarium specimens, which unless accompanied by spirit material, detailed field notes and colour pictures, are often almost worthless. Molecular techniques are therefore particularly useful in this genus and have been used to study genetic diversity at the subspecific (Wong et al., 2001a), specific (Wong et al., 2001b) and sectional levels (Jarret & Gawel, 1995, Wong et al., 2002). AFLP is a DNA fingerprinting technique, which was developed by Vos et al., (1995). It is based on selective PCR amplification of DNA restriction fragments under stringent conditions. It can be used for DNA of any origin and complexity and is reported to be both reproducible and reliable (Vos et al., 1995). AFLP combines the reliability of RFLP with the power of PRC.
In this study, AFLP (amplified fragment length polymorphism) was used to analyse the genetic relations of two species collected from Mt Jaya, Papua (formerly Irian Jaya), New Guinea: (a) to study the genetic relationship of Musa johnsii Argent, which is unique in the genus in having a fruit with a sterile mucilaginous pith chamber that occupies the distal third of the fruit (Argent, 2001); and (b) to reassess the status of M. banksii F. Muell., which was reduced to a subspecies of M. acuminata Colla by Simmonds (1956) but is still considered to be a distinct species by Argent (1976).
In addition, the taxonomic status of the five sections of Musa is reviewed in the light of molecular studies.
Materials and Methods
Plant Materials
Twelve samples were used for the present study (Table 1). These included Musa species from sections Musa, Rhodochlamys, Callimusa and Australimusa and a species of the related genus Ensete as an outgroup taxon. Voucher specimens were deposited in the herbaria of the Singapore Botanic Gardens and the Royal Botanic Garden Edinburgh.
DNA Extraction
Leaf tissue was used for AFLP analysis and prepared using a procedure from Zhang et al. (1997). Plant DNA was extracted using the CTAB method according to Reichardt and Rogers (1993) as outlined in Wong et al. (2001b).
AFLP analysis
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99
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to generate template DNA for amplification. PCR was performed in two consecutive reactions. The template DNA generated was first pre-amplified using AFLP primers each having one selective nucleotide. The PCR products of the pre-amplification reaction were then used as template after dilution 5-fold in sterile water, for selective amplification using two AFLP primers, each containing three selective nucleotides. A total of eight primer combinations were used in this study (Table 2). The final PCR products were run on a 6% denaturing polyacrylamide gel in 1X TBE buffer. The EcoRI primers used were not radioactively labelled as in the original protocol. Instead, a modified silver staining method was used (Loh et al., 1999).
Table 2 The sequence of primers used in the AFLP analysis.
Name / Abbreviation
GYY 101/ EA+ GYY 102/ EA-
GYY 103/ MA+ GYY 104/ MA-
GYY 105/E-A
GYY 107/ E-AAC GYY 108/ E-AAG GYY 109/ E-ACA GYY 110/ E-ACT GYY 111/ E-ACC GYY 112/ E-ACG GYY 113/ E-AGC GYY 114/ E-AGG
GYY 106/ M-C
GYY 115/ M-CAA GYY 116/ M-CAC GYY 117/ M-CAG GYY 118/ M-CAT GYY 119/ M-CTA GYY 120/ M-CTC GYY 121/M-CTG GYY 122/ M-CTT
Enzyme
EcoRI EcoRI Msel Msel
EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI EcoRI
Msel Msel Msel Msel Msel Msel Msel Msel Msel
Type
Adapter + Adapter - Adapter + Adapter -
Primer +1 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3
Primer +1 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3 Primer +3
Sequence (5’-3’)
CTCGTAGACTGCGTACC AATTGGTACGCAGTCTAC GACGATGAGTCCTGAG TACTCAGGACTCAT
GACTGCGTACCAATTCA
GACTGCGTACCAATTCAAC GACTGCGTACCAATTCAAG GACTGCGTACCAATTCACA GACTGCGTACCAATTCACT GACTGCGTACCAATTCACC GACTGCGTACCAATTCACG GACTGCGTACCAATTCAGC GACTGCGTACCAATTCAGG
GATGAGTCCTGAGTAAC
GATGAGTCCTGAGTAACAA GATGAGTCCTGAGTAACAC GATGAGTCCTGAGTAACAG GATGAGTCCTGAGTAACAT GATGAGTCCTGAGTAACTA GATGAGTCCTGAGTAACTC GATGAGTCCTGAGTAACTG GATGAGTCCTGAGTAACTT
Musa in New Guinea & Section of Musa 101
Data analysis
For the diversity analysis, bands were scored as present (1) or absent (0) to form a raw data matrix. A square symmetric matrix of similarity was then obtained using Jaccard’s Similarity Coefficient Coefficient [x/(y-z)], where x is the number of fragments in common between two taxa, y is the total number of fragments scored, z is the number of fragments absent in both taxa, from the raw data matrix. Genetic diversity estimates (GDEs) were then calculated as 1 minus Jaccard’s Similarity Coefficient and used for cluster analysis using the Unweighted Pair Group Method with Arithmetic mean (UPGMA) technique of the NEIGHBOR program in PHYLIP version 3.5c (Felsenstein, 1993). The dendrogram was drawn using TREEVIEW version 1.6.1 (Page, 1996).
Results
AFLP Profiles The AFLP profiles generated using eight primer combinations contained highly informative bands, which distinguished all taxa examined and provided valuable information on genetic relationships. Figure | illustrates an AFLP profile obtained. Only unambiguous bands of size 50-500 base pairs were scored. A total of 201 bands were analysed out of which (95%) were polymorphic across all samples examined. On average, 27 bands were scored per primer pair.
Fifty-nine unique bands were observed for the taxa examined (Table 3). All species except for Musa textilis were characterised by unique markers ranging from 1 to 18. M. johnsii is characterised by six unique bands, confirming it is a distinct species. M. banksii is characterised by four unique bands. The genetic markers observed will be useful for the development of probes in Musa breeding programmes. The genetic diversity estimates are shown in Table 4.
Discussion
Position of Musa johnsii
This species has strikingly distinct morphological characters, such as the compact, subglobose head of schizocarpic fruits that are unique in the distal third being filled with pale pinkish orange, mucilaginous pith (Argent, 2001). AFLP analysis confirms it is a distinct species (Fig. 2). Argent (2001) suggested it was closely related to the schizocarpic M. lolodensis as the seeds of the two species are remarkably similar but Table 4 and Fig. 2 show it is genetically more similar to the Australian M. jackeyi than
102 Gard. Bull. Singapore 55 (2003)
it is to the New Guinea M. lolodensis. All three species belong in Sect. Callimusa in the ‘textilis’ group (see below). This is another example where conspicuous morphological differences do not reflect genetic similarity (Jarret & Gawel, 1995).
Status of Musa banksii Simmonds (1956) reduced this species to a subspecies of Musa acuminata based on experiment and field observations. However, Argent (1976) maintained it as a distinct species as it did not hybridise with M. acuminata subsp. malaccensis when they were grown together for many years in the Lae Botanic Garden, Papua New Guinea. This taxon also differs from the other M. acuminata subspecies in its female flowers having some fertile stemens, in the non-imbricating bracts in the male bud, and in producing a very large number of seeds (up to 400) as compared with 40-50 in subspecies from the lowlands of Peninsular Malaysia and Thailand (Simmonds, 1995).
Results of several studies using molecular techniques are now available. Gawel & Jarret (1991) used cpDNA RFLP, which generated a phenogram with M. banksii embedded among the other subspecies of M. acuminata studied, indicating that it is not genetically distinct at the species level. Jarret & Gawel (1995) using total-DNA RFLP again found that M. banksii clustered with the other M. acuminata subspecies in their phenogram. Our study using AFLP showed M. banksii clustering with the other subspecies of M. acuminata (Fig. 2), again indicating that genetically it is not a distinct species and that subspecific rank would be more appropriate. However, the genetic diversity estimates (Table 4) indicate it is the least similar among the four subspecies studied, perhaps a reflection of a combination of its unique characters listed above and its most south-easterly distribution of all M. acuminata subspecies.
Sections and groupings of wild banana species Cheesman (1947) created four sections within the genus Musa - sections Eumusa (now Musa), Rhodochlamys, Callimusa and Australimusa - as a convenient way of grouping the species. He noted that the sections were not of equal rank and that in some characters sect. Australimusa was intermediate between sect. Callimusa and sect. Musa, and that the division between sect. Musa and sect. Rhodochlamys was ‘unessential’. He speculated that sect. Musa and sect. Australimusa were the earliest to diverge and that sect. Rhodochlamys was an offshoot from sect. Musa. He was in two minds as to whether sect. Callimusa had diverged directly from sect. Australimusa or, as he considered the barrel-shaped seeds with a large oil/air space a very significant character, might be an earlier divergence from sect. Musa. Argent (1976a) described a fifth section, /gentimusa, with the single species M. ingens N.W. Simmonds. Subsequent to Cheesman’s work, new species have been described that do not conform to his concepts of the sections so that the distinction between the sections is becoming blurred. Molecular techniques have opened a new avenue of enquiry. Studies carried out so far (Jarret & Gawel, 1995; Wong et al., 2002, and this study)
Musa in New Guinea & Section of Musa 103
Miz3 45