Prunny Wa epg ti eae SRN dt ee PY an ree ged
Pav lb geye gy?
ee Mays abe yts 2 yb PAH EPL GH eo oye PSE RIN aya, LS YS Gay Eee Se er arty Sern VIVID EAN EE NGM EN aed Tey Cee Cte ty He T OUT O 7 ee LU UO ee ae PITS NG TL AEE IETS ED LS SEES GENES Doge Bets COR audigy BP) PT AG Ja OANA seeded Vegi gh
Oe eb
a RATE a Maes BOOP hE COR MeO Pye ts Bul: LWT BS aD Y SAT te fee dts eter HE th be
hg hrs une b ete ny acral watae ig hie shadeighen peed parohich sy, Seneca ML the aM all Silk a Fall att ACO SE \attitl eet alba eS Miak lal bas aU a yt at cube cies Lidiabeiedbdetloes what! sha ele & Saar aitb sos ST Mall ARTS bye ae rainerallag th SAE Mt Moth yirsirpersiapl acid sat ah Vick leh
a Vb ae ae pee ng RO Mb an AUN AV Leyak ben Ah ye pyetetehy VARNA VA ah ery FT MA ae og eye el rhe.
MRS UNL pated igs a a TON deers ats Woy CeaP to %
Sey bry WEA EERE ES Abe a NEVA BN ah heme Tt eae
cad abv dire
eee re tent
Be gcwb seen at NTR ALS Cbegh ot rt yi
Maru Hee
outa Shem weba sy aft athe SODA eed ties Pea eee it Vv byte ote eta oat BEN BEEN A TVG Gb LIS TURIN OUR UAE A rd Kiaed Bw a Uae DRM SML SALAM ES pb HL atl caw fh megs ac aned ce die wd ZAMS VUTNPS BST ELEY ERECT STC INARA B SLU LS WN des CURA POE TET obdad eR EN MG dN HAUG inne ta Ey PN BAIT MME AA Se SOT tee SNOB SMG RT Se Lath ik ah yea outta Be 2B SARA WEN oP 9 Sant Astra aA ptbeth BTA 8 LAR os wang Pasta ct, Be Pees te Mabe ERY eh ET dee eee ARAN EMRE ON EE RS Pye he Ae ee FERNS ey hg te tye m, SYM SORT AE ae et et iat ae WO avy Nie Osis, he NH a oe UR IN, cE SW SBR ORM alt Hewah cam has AONE MES ONE VLE NOT SS TR Peas Sree Lene, eee |e es ee eer geyp oa cite ‘ eR Sn ieee ath lbdapas ea ASN ANA AT of ‘ > eres Be dred OMe Tey vere eeu peat A bpoiSih pal er Sirrty AEH Wah at EL don coat amy eT Sart or Ee ‘ Sonu ere atts WP Ul teehee PE EULER ES pe ae ih git q VAUAY ROUTE DES pa be Et neh deka et py tapy Ede dy VAN UREN NT Chey Vy he yb te LAL i ey y EN Let eM PM ya Lad Wee td SE AROS ENTREE ES My wy yet ban a SET RE TERED Hay COMIN EL TIN ere Leah Oe de LS i SEN AEYOMD EGS CPT ge 48 ETH GS gh Lewsey sOvb ed ee bal Ee Dy Svat wah ge Ae Medea ene i Aeyteate Ie Mn at 64 Heng shape Tae yl es eee a Ce ee Oe ee ce ee ee Pn eed RW neck es ae ee cee wl a Aba gs Oly UNO IREE Cope GR TREE UUW Jory geyttytys hae AE RVR EU tad vhs Paitai ot yh al yen dibewe tey RNS Sse IRR N, Auta Tete WENN otis ee wget AUPE AE RANGEL TOTS Phu Dandie Gade Muy a en ae he ho NADINE UV EN Gv Harsh ebets wy whack eb AN nb vis FERN ta te SP Viol atin Ma er Sh ee ee ot rer “pues Neate ye Nt Basie ae ie hoe Le PT oe See Le ees ee ee eT ‘ FP nD ba AUN NSN TC EL ake SUPE INES ob SEE SEM EL ohh ay! wee hatha bet seb abn ae My dts Wy Aue ha We Oy Lae GS wis ro! SUMP PREM PR py tod ah ed ey ey rs Ie.
feves CYS COD Thebes yt kde ase SEALE GY DN Cay ds Pi bE ae oes Sor Te ee eco ere en ey
14 PEO A ie ti IN eaTFS My Se Mdeteg NOPE EIN Sa ire MF thelta Ne lea ate De Zs
Wed lates esac Bar ty OME Ne ELS Hy Wa BS Ge tea ti Wists lap ea
PEE ede Ib ab aa oi Tt a Le Le ee ee ed eT TY AVINASH EYES Eat CUS atestheh pe ee pede ve Dy eS NE bR Buh hoe {haa ck abe ge sty Wud ys, eee ere rah ere ere ere
Eee yey a NAUSEA GN fb UN ERNST EIT EME b EN Pee oo eee Dee eer wayne SUA nT Lik as sod ENE teeta and 1 4 Pleo ee ae a eT eh es Sa DO eee her eaY SR ANE Bayh PMA RM EERILY MET Fee TS Sug ed bet tater pH
SVN GREASE AN od AN US GEN gg AVORP UN EL AMEE ROUBLE Lye SRY SU pare
fe toed yeh AN S50 Sa re ot ae ee mes ee ere ened ee Cre ee Wd RTS ot TELE NWR ALS ON Aes Gyo EUS TaT PL akus ERY Meee ty Hehehe g we Safar ht Sure WA RTE Eee ae oe yee ae, RYO Jere deel: Seer hr eee aec eee OLE Pe ere ene ele ene hy TURE NR AE OR SU Sty MEL wd eet ot VISASID OLIN Sys gegl etal gs
Aas bON a sty bat ye om
Lee eae) Pate?
VIVA ey dy
Ved Rpt Peis Tey mer aes
Bathe Pyey vibe ya Lets Ah Ok eh pa ay eee Wh ENG a th ib NFU a eH AL oe nigeaoes ASM SAD ML AAT AL RLSM Neth SMUT BoP Bt aay M4 ANALY Oiew
vat Oa re nee UR Ruy! ghey a at SPNTV ES Ed Saba ny at ty Spat yet yey ebay aye
De ciheh hod de Weld ee ay beh Sete bkWN EG beg FAME UF Rea NEL sale crt vt oty Ve EWE REET eae ca wh TS ge et wesley AMEE ESS ony PAMtg A PEE TV eR arb EY rk DE UY RENEE Se AEE tee Quek tage FEV ae OORT NEY BN Vd ev EY Ab TUS Abe SD gt att wavy sa eviebt ted Fe he sere AVIV CASTS tie Eyer Rn ee ete eet eer Core trae Fv eA re ey Py ets he em tes evi Eglo y 2) Hatha wth POMS ENE SSL Gi Fahey
mi met nS Anas TYEU Seton
CV vb dae 3 0 Py PEN eh PENI We
ee Bay she Nelatunt Shela! SRSA eh it baba aed wt eRe
Privy a Ped Dade Nebo de & Be PN dete
OFF Bet tat a dis By f
PPAR CEM boa Ny USAT, MEIER EE ot
Pe CNH, oe
Aya Aah Pe ee
Wee wa te te
Te PLO tha ad
AULT Cave wh
et Seshatt Ah aes
Wa ath ct Aaya wits,
Reo Rey Es TNE PRE FR by
(ie tha chai ta Ne ave ie faite efron ety stay Raha rg MMe,
eT eee eT ed Ee] TD hy Mode
deg aeg tena Nyt h ge VELBA See gees eh WES RTP ou bye g ean = eee ee Cee eee eed eae ees SAUDE UGA ORG fo ght
ea bay Oo gm Rf edog sv euds PVN ET dt Ysa WT GRUNT NE de sed roa DURA ge Sh Ode Sad tr ahaeas yates SNe aba geste cst
Sree Ve vothal fhdis cK abn
ay)
een Cee ee eee re ee Rae Re
abate aden tat Tae atl batt
avh yaa, Cv babatt rt ye eT TUN te Sat “Ph ded ye MbET SL erat re eV Bede LSS Ey ety Vet h tee Sve ta She Aye ya WON ae NPL ad gt dN ee Sew Batata / EV e eben tobe h bay ely wha rit
civt os
le HBR Ne Pa ite its eM di
Bede ee et stew Sb al hg wet
Bate ah annie hala
PAN Atty ty Piva Ps
ead Mees Ub VM EE HT gly.
EN Me dread ee ee ed Hee oh
EET UNE LETC cee AEA yy twe
wees
Tety tate ta That Ser artes birt
sthaes Se ylwhylyey ar
4 Med auy eee dey tata PEIN TWIN SEEN ee SONY et et ehy io Seen in gry PEIN RENAE DEES IL OE ohn
Wa tata Phe at
PRTUPE GIT IVE aN ee tg om PAE Yad Ee Ee oN
sh Mout shen at Pyle
SsVitarere
PA Lm Se Meghann, Mavarhial a EWE GPRS LY YUE ew dS Mathes Ee TM pat wove Jee hr Eb ta ey stele oe en a Tay WAN Gd PELE EY VENT TSUN Stabe lerdivistybers Bopeh few rats Ceo Oa eTy LRSENANSE DOP CRN ves IVER wid Sard SINE be tet S ads aby bs say bate oP Nes
a2 i
PMU ST CV aeya ira yh Scar tet gs erg awn yt Weta sta Wa yt
vous age
POE IVAN ES TT eet
vybwern BAVA,
AEGN VIN ILS iegee buns ate SOUT WCE abet gy ree gees
fen Poveeylombe
dates ere vanahs bel ea ee ES
E has Sionte Few ae
PT A Met cee! DIME ed Svein es oe
ae Sa ty
RN, ere et
Weert Ava arate tov ht ewe
Higbee Seda at Say aE
WGI ean ee
rye
ng 84
rer ret teeny sober eit AbEN AGE
etl Oy ie Palate se edwin
YSoy veg! ey
a err:
Weve eon vi
Harvard Botany Librari
HN
1 821
LIBRARY OF
THE
ARNOLD ARBORETUM
HARVARD UNIVERSITY
Digitized by the Internet Archive Sein 2014
http://archive.org/details/gardensbulletins458u nse
rk ee i 1 i
The Gardens' Bulletin
Singapore
VOL. 58 (Part 1) 2006 | oe ISSN 0374-7859 =
NATIONAL PARKSBOARD ———is—t Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 64741165 Telefax: 64754295. is
THE GARDENS’ BULLETIN SINGAPORE
The Gardens’ Bulletin Singapore publishes original papers on plant taxonomy (including revisions), horticulture, phytogeography, Floristics, morphology, anatomy and related fields with emphasis on plants in the West Malesian region.
Dr. B. C. Tan and Dr. R. Kiew (Editors of vol. 58)
Singapore Botanic Gardens
Dr. PY. Tan Ms. C. Soh (Assist. Editor) (Business Manager) National Parks Board Singapore Botanic Gardens EDITORIAL BOARD Dr. S.C. Chin Dr. E. Soepadmo Singapore Botanic Gardens Forest Research Institute Malaysia
Kepong, Malaysia
Dr. M.J.E. Coode Dr. W.K. Tan
Royal Botanic Gardens Kew. U.K. National Parks Board, Singapore Dr.-R.T. Corlett Dr. I. M. Turner
University of Hong Kong Research Associate
Hong Kong Singapore Botanic Gardens
Dr. M.C. Roos
National Herbarium of the Netherlands Leiden University
The Netherlands
The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore. Neither the National Parks Board nor the Editorial Board is responsible for the opinions or conclusions expressed by the contributing authors.
The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including postage. Overseas subscribers are required to make payment in the form of bank drafts or international
money orders in Singapore currency payable to National Parks Board, Singapore.
Instructions for contributing authors are found on the inside back cover.
ae | APR 24 2007 ARNOLD AREA aij:
The Gardens' Bulletin
Singapore
VOL. 58 (Part 1) 2006 ISSN 0374-7859 CONTENTS Boyce, P.C. ihwaphidophora tenuis (Araceae: Monstereac) Resurrected ¢....2:......0.so.sencceenesecesoneseoets if Boyce, P.C. A trio of new Schismatoglottis (Araceae: Schismatoglottideae) from Sarawak, Borneo SSA ee es ee ee ee ee eee eee eee ee i Boyce, P.C.
Rhaphidophora crassifolia Hook.f. (Araceae: Monstereae): a new record for Sarawak
and notes on the Rhaphidophora ‘Hongkongensis” group in Borneo .........eeeeeeeeeeeees i Boyce, P.C. and S. Julia
A review of entire-leaved Tacca (Dioscoreaceae) in Sarawak, Bor ...........ccccccssceeseeeeseees 25 Maknoi, C. and T. Jenjittikul
A new species of Curcuma L. (Zingiberaceae) from Southeast Asia .........ccceeeeeseeeeees 4]
Shafreena, A. and P.C. Boyce | A hitherto overlooked field identification character for Bornean Scaphochlamys Baker Preece cee, hotter a Oa ahaa eee ne Da ees GA Aaah ah eel eddies dbgeah eoesesbeabdacae deed 47
Watthana, S., T. Hidayat, M. Ito and T. Yukawa Phylogeny of the Genus Pomatocaipa Breda (Orehyidaceae))s-......0.sceseee ee ee 55
Zhu, W., H. Wang and B.-G. Li Species composition, physiognomy and biogeography of tropical montane rain forest in
southern: Yuntiam- Of China: 35.5, oscsace asessdensteede.g suze sins cals epee a eee 81 Obituary Humphrey Morrison Burkill O-B:E., FL.S.(19 14-2006) ececenccse-ceeeee- coe ee 133
Date of publication: 18 December 2006
Published by
National Parks Board Singapore Botanic Gardens Cluny Road Singapore 259569
Printed by Oxford Graphic Printers Pte Ltd
Gardens’ Bulletin Singapore 58 (2006) 1—6 1
Rhaphidophora tenuis (Araceae: Monstereae) Resurrected
Pe BOYCE
Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia
Abstract
Rhaphidophora tenuis Engl., a species considered synonymous with the widespread and variable R. korthalsii Hassk. in the most recent revision of Bornean species is resurrected as an endemic to Sarawak and Brunei. A full description of R. tenuis is presented together with a modification to the most recent published key to Rhaphidophora in Borneo and photographs. This reinstatement takes to 15 the number of Rhaphidophora species recognized for Borneo.
Introduction
In the revision of Rhaphidophora for Borneo Boyce (2001) treated Rhaphidophora korthalsii Hassk. as a polymorphic species. Since that publication, the author has been able to undertake extended and on-going fieldwork in Sarawak which has revealed that at least one element of Bornean R. korthalsii sensu Boyce 2001 is a morphologically stable taxon with a suite of characters consistently separating it from R. korthalsii s. s. In particular the smaller stature, slender leaflets and solitary, slender inflorescence, and most strikingly in the form of the leaves in the juvenile shingling plant which are ovate and spreading in R. korthalsii s. s. (Plate 1c & d), but strongly falcate-lanceolate and ascending in the segregate taxon. Such plants match incontrovertibly R. tenuis Engl. based on Beccari collections from Matang, Kuching Division.
Rhaphidophora tenuis Engl.
Rhaphidophora tenuis Engl., Bot. Jahrb. Syst. 1 (1881) 181; Beccari, Malesia 1 (1882) 271-272; Engl. & K. Krause in Engl., Pflanzenr. 37 (I1V.23B) (1908) 53.—Type: Malaysia, Sarawak, Kuching Division, Matang, Beccari PB 1977
2 Gard. Bull. Singapore 58 (2006)
(lecto, FI; isolecto, B; selected by Boyce, 1999).
Rhaphidophora korthalsii var. angustiloba Ridl. ex Engl. & K. Krause in Engler, Pflanzenr. 37 (IV.23B) (1908) 49. — Type: Malaysia, Sarawak, Kuching Division, Matang, Jul 1903, Ridley s.n. (lecto, SING; selected by Boyce, 1999). Plate 1 (a & b).
Moderate slender heterophyllous liana to 5 m; seedling stage a non- skototropic shingling juvenile shoot; pre-adult plants never forming terrestrial colonies; adult shoot architecture comprised of elongated, clinging, physiognomically unbranched, moderately leafy fertile stems; stems smooth, medium green, with sparse prophyll, cataphyll and petiolar sheath fibre, especially at the stem tips, internodes to 11 x 2 cm, separated by prominent oblique leaf scars, older stems subwoody; flagellate foraging stems absent; clasping roots densely arising from the nodes and internodes, prominently pubescent; feeding roots abundant, adherent and free, very robust, densely ramentose-scaly; leaves distichous; cataphylls and prophylls membranous, soon drying and degrading to intricately reticulate fibres, these only very slowly falling; petiole shallowly grooved, upper part + terete, (1—) 9-65 x 0.2-1.5 cm, smooth, apical and basal genicula prominent; petiolar sheath prominent, membranous, strongly to slightly unequal on one side, extending almost to or reaching the apical geniculum, of + short-duration, soon degrading into persistent netted fibres, these eventually falling leaving a prominent, slightly corky scar; shingling lamina entire, ascending falcate- lanceolate, 5—11x 3.5—6 cm, base slightly cordate, pre-adult and adult lamina spreading, entire, pinnatipartite to pinnatisect, 10-30 x 14-45 cm, broadly oblong-elliptic to oblong lanceolate, slightly oblique, subcoriaceous, base truncate and very briefly decurrent, apex acute to acuminate, individual pinnae 1-2 cm wide, perforated basally adjacent to the midrib, thus appearing stilted; midrib very prominently raised abaxially, slightly sunken adaxially; primary venation pinnate, raised abaxially, somewhat impressed adaxially, 2—4 primary veins per pinna; interprimaries sub-parallel to primaries, slightly raised abaxially, slightly impressed adaxially; secondary venation strongly reticulate, slightly raised; tertiary venation invisible; inflorescence solitary subtended by a membranous prophyll and one or more cataphylls, these swiftly degrading to netted fibres; peduncle slightly laterally compressed to terete, 6-26 x 1-1.5 cm; spathe narrowly canoe-shaped, stoutly beaked, 10-15 x 3-3.5 cm, stiffly fleshy, dull yellow, gaping at female anthesis and then caducous leaving a large straight scar at the base of the spadix; spadix cylindrical, sessile, inserted + level on peduncle, 9-13 x 1—1.5 cm, dirty white; stylar region well developed, mostly rhombohexagonal, 1.5—2 x c.2 mm, very slightly conical; stigmas slightly elliptic, mostly longitudinally oriented, c. 0.3-0.5 x 0.2-0.4 mm; anthers barely exserted at male anthesis; infructescence
Rhaphidophora tenuis Resurrected
v
t be ,
Plate 1. Rhaphidophora tenuis Engl.
a. Juvenile shingling stage showing diagnostic ascending falcate-lanceolate leaves. b. Adult plant showing narrow pinnae.
Rhaphidophora korthalsii Hassk. €: Juvenile shingling stage with ovate, spreading and overlapping leaves.
d. Adult plant showing broad pinnae.
4 Gard. Bull. Singapore 58 (2006)
14-18 x 2—3cm, dark green ripening to dull orange, stylar tissue abscissing to reveal pale orange ovary cavity pulp.
Distribution: Sarawak, Brunei. Endemic. Habitat: Primary to disturbed secondary lowland and hill forest, on trees and rocks on a variety of substrates including limestone. 20-650 m altitude.
Other specimens examined: SARAWAK. Kuching Division: Three miles from Kuching, Haviland & Hose 3605 (K); Semengoh E.R., 6 miles west of Kuching, Nicolson 1252 (US); Padawan, Subang, 7 Dec 2004, M. Gibernau AR-837 (SAR); Bau, Kampung Segong, 26 Jun 2005, Jeland ak Kisai & A. Shafreena AR-1262 (SAR); Padawan, Gunung Braang, 2 May 2001, C. Lee AR-68.1 (SAR). Samarahan Division: Tebedu, mile 15, Mohtar et al. S 49245 (K, SAR, US); Serian, G. Penrissen; Paie S 16001 (K, LE, SAR). Bintulu Division: Eastern ridge of Bt. Kans, Bintulu district, Hirano & Hotta 1432 (KYO); Sg. Ma’au, Dataran Tinggi Merurong, Tubau, Othman et al. S 49050 (K, SAR); Bukit Sarang, Ulu Kakus, 02° 39’ 21.08”; 100° 02’ 61.09”, R. Kiew, Julia anak Sang & S.Lee AR-729 (SAR). BRUNEI DARUSSALAM. Belait: Ulu Ingei, Bt. Batu Patam, lower slopes near Sg. Ingei, Boyce 312 (BRUN,K).
Rhaphidophora tenuis can be fitted into the key to Bornean Rhaphidophora (Boyce 2001) as follows:
la. Mature leat lamima:pinnately divided “3-42.4..-2..-e eo eee Z
1b. Mature leaf lamina entire, without or without perforations, but never pinnately divided | svheicnGicsdcss ss ists: weeca eee ee eee ee ee 3
2a. Plants always associated with sandy or rocky forest streams. Flowering plants usually rheophytic, rarely low-climbing on trees beside torrential SEV SANNA is sa Ge eo oo ee leo ole cn - «Re eee R. beccarii
2b. Plants not specifically associated with watercourses.
3a. Juvenile shingling stage falcate-lanceolate non-overlapping ascending leaves; adult plants with leaf pinnae no more than 2 cm wide and frequently much less; inflorescences solitary; spadix slender, not exceeding 13 cm long a ndsinnedidinn a5 nicisina peas sipiph cede os Fedral ie SaeRicayt sae iiad ek ae ere R. tenuis
Rhaphidophora tenuis Resurrected 5)
3b. Juvenile shingling stage with ovate, overlapping spreading leaves; adult plants with leaf pinnae exceeding 3 cm wide; inflorescences several together; SJOEIC ED STC UT EM (0) 720) Gr7 018) (081 ee en eee R. korthalsii
3a = 4a, etc., as in Boyce (2001)
References
Boyce, PC. 1999. The genus Rhaphidophora WHassk. (Araceae Monsteroideae-Monstereae) in Peninsular Malaysia, and Singapore. Gardens’ Bulletin Singapore 51:183-256.
Boyce, PC. 2001. The genus Rhaphidophora Hassk. (Araceae- Monsteroideae-Monstereae) in Borneo. Gardens’ Bulletin Singapore 53: 19-74.
e
Guninad Waves mn pet igi (eroe) ie s gereeval fs Cadre Be leila * 7, ¥ ” - ." ab “oe on we ‘ q rela
pie sini 8 opt eased
fe, ew iA ww nity
4 . | | SARE
beats ean Gangs av ‘awe? 4 a3 eta ad Ris leeds“
} | ats =) oe el?
ae |
"y
Gardens’ Bulletin Singapore 58 (2006) 7—18 7
Studies on Schismatoglottidae (Araceae) of Borneo I: A trio of new Schismatoglottis from Sarawak
PC. BOYCE AND WONG SIN YENG
; Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia
* Faculty of Resource Science and Technolgy, Universiti Malaysia Sarawak, 94300 Samarahan, Sarawah, Malaysia
Abstract Three new species of Schismatoglottis - Schismatoglottis jelandii P.C.Boyce & S.Y. Wong, Schismatoglottis jepomii P.C.Boyce & S.Y. Wong and Schisma- toglottis maelii P.C.Boyce S.Y. Wong - from Sarawak are described and in-
cluded in the amendments to the key to Bornean Schismatoglottis published by Hay & Yuzammi (2000). All are illustrated.
Introduction
Recent revisions and publications on tropical Asian Schismatoglottis (Hay & Yuzammi, 2000; Hay, 2002; Hay & Herscovitch, 2003) have greatly facilitated accurate naming of Bornean Schismatoglottis and provided an excellent platform from which to describe further novelties without the concern that obfuscated species names are being overlooked. Despite a 100% increase in the number of Bornean Schismatoglottis in the past five years resulting from these publications, on-going fieldwork in Sarawak has revealed that there is still a considerable number of Schismatoglottis species awaiting formal description. Three are described here.
I. Schismatoglottis jelandii P.C.Boyce & S.Y. Wong, sp. nov.
Schismatoglottidem asperatam Engl. simulans sed folii lamina stenophylla at inflorescentia gemini differt.-Typus: Malaysia. Sarawak, Bintulu Division, Bintulu, Bukit Satiam, 02° 59’ 33.0”N; 112° 56’ 01.4”E, 12 Aug 2004, PC Boyce & Jeland ak Kisai AR-636 (holo,SAR). Plate 1.
Facultative rheophytic herb to c.30 cm tall, solitary or forming small clumps.
8 Gard. Bull. Singapore 58 (2006)
Stem condensed, erect, to c. 1 cm diam., pleionanthic; internodes obscured by overlapping leaf bases, not conspicuous. Leaves several together (c. 10 per plant); petiole D-shaped in cross-section, 2-4(—7) cm long, c.1/3 the length of the lamina, sheathing in the lower 1/3-2/3, asperulate, narrowly crispulate-alate on each side on the adaxial face; petiolar sheath adnate to petiole for the greater part, then free-auriculate at the top for c. 1 cm, broad and subparallel, the margins inrolled; blade narrowly obovate, thinly subsucculent, 15-21(—26) x 2-4(-5) cm, lustrous deep green adaxially variegated with two longitudinal zones of diffuse greenish white blotches, matt pale brownish yellow-green abaxially, the base slightly obliquely truncate, the tip attenuate and apiculate for c. 2 mm; midrib not prominent adaxially, prominently raised abaxially; primary lateral veins c. 9 on each side, alternating with slightly to considerably lesser interprimaries, diverging at c. 30°; secondary conspicuously tessellate (abaxially); tertiary venation not visible. Inflorescences two together, subtended by one narrow prophyll and one broad cataphyll; peduncle short, not exserted from leaf bases. Spathe 5.5-7.5 cm long, subcylindric, tapering distally, weakly rounded- truncate basally; lower spathe 1.5—2.5 cm long, ovoid, flushed pale pink, hardly differentiated from the limb by a barely perceptible constriction corresponding with the upper part of the fertile male zone; limb 3.5—-6 cm long, narrowly ovate-lanceolate, dirty white flushed and slightly veined pale pink, spreading at onset of female anthesis then crumbling-deliquescent at late male anthesis. Spadix sessile, somewhat shorter than the spathe, to 5.5 cm long; female zone 0.6—1 cm long, with oblique to very shortly adnate (to spathe) insertion, 0.4 cm diam. at base, weakly conoid; pistils crowded, oblong-barrel-shaped, c.1 mm tall, 0.75 mm diam.; stigma sessile, discoid, weakly umbonate, papillate, fractionally wider than ovary and + contiguous with neighbouring stigmas; interpistillar staminodes few, claviform, scattered among the pistils, slightly taller than the pistils, c.0.5mm diam.; sterile interstice 3-8 mm long, slightly wider with top of female and base of male zones, 4—5 whorls of irregularly polygonal staminodes c. 0.5 mm diam., separated from the female zone by a row of more slender staminodes; male zone c. 0.8 mm long, cylindric, fractionally wider at the junction of the appendix; stamens orange (even before anthesis) crowded, truncate with the thick connective slightly elevated above the thecae, more or less rectangular from above; pollen extruded in short strands; appendix pale pink, about one and a half the length of the rest of the spadix, widest c. half way up, then gradually tapering to a rather narrow blunt tip, 2.5-3.5 cm long, 4-6 mm diam.; staminodes of appendix more or less flat-topped, irregularly polygonal, 0.75-1mm diam., densely crowded (fresh). Fruiting spathe not observed.
A trio of new Schismatoglottis from Sarawak, Borneo
Plate 1. Schismatoglottis jelandii a. Holotype plants in habitat; b. Inflorescence at female anthesis. Note orange stamens; Inflorescence at male anthesis, spathe beginning to deliquesce prior to being shed.
10 Gard. Bull. Singapore 58 (2006)
Distribution: Sarawak, Bintulu Division. Known only from the type locality. Habitat: Facultative rheophyte on stream banks on sandstone-derived clays in slightly seasonally dry lowland secondary forest at 18 m altitude.
Notes: Schismatoglottis jelandii falls within the Schismatoglottis asperata group (see Hay & Yuzammi 2000:54) by the pleionanthic shoot, fully attached persistent leaf sheath and the spathe limb opening more or less wide and then crumbling-deliquescent. It is most similar to S. asperata Engl. from which it differs most notably in the stenophyllous habit, the conspicuously tessellate secondary venation (visible on the abaxial surface of the lamina) and in always producing pairs of inflorescences. Additionally the stamens are orange prior to and after anthesis vs. white prior and orange at and after anthesis in S. asperata.
The type locality is subjected to periodic inundation and while the plants of S. jelandii are probably not regularly buffeted by flood waters it does seem likely that during particularly heavy rainfall they are at least temporarily in the flood water flow. It seems reasonable to speculate that the stenophyllous habit is adaptive for this.
Hay & Yuzammi (2000) adopted a broad circumscription for S. asperata with the caveat that, ‘Possibly it will transpire on further field study that more than one taxon has been included here, but at present there is insufficient evidence on which to base a split’. Extensive and focused fieldwork since has revealed that S. asperata is widespread and indeed variable but that there exists a considerable number of localized morphologically distinct taxa, frequently adapted to a specific substrate (shale in particular) and/or ecological niche and that these do, indeed, warrant taxonomic recognition.
Schismato glottis jelandiican be fitted into the key to Bornean Schismatoglottis (Hay & Yuzammi 2000) as follows:
3la-Appeéndrs subeylinGric \i.6::.5cc nk... i eee GW b. Appendix conic-ellipsoid to broadly Conic-OVOIC ........... eee eeseeeeeeeeeeeeees 34 32a. Petiole conspicuously and thickly longitudinally ridged; stigmas minute, punctate, sterile interstice absent (or reduced to a whorl of incompletely
abortive stamens); Southern'Sarawak.s...Af%-- 2.0% 2e eee 27. S. nervosa
b. Petiole finely longitudinally ridged and/or asperous; stigmas button-like to discoid; sterile interstice well-defined: s34.2c.20....2 0 ee ee 35
A trio of new Schismatoglottis from Sarawak, Borneo i
33a. Petiole not ridged; appendix about two thirds of the length of the spadix;
b. Petiole finely longitudinally ridged; appendix about or less than one third of the length of the spadix; Sarawak, ? W. Kalimantan ..... 25. S. latevaginata
34a. Plant with broad leaves and not associated exclusively with riverine habitats; secondary venation obscure; inflorescences solitary; stamens white prior to anthesis, then turning orange. Widespread in NW Borneo ................. eee eee eM oe ER eel sbelradsucuionssa,pbdanuevinssdabdoenaueseaes 14. S. asperata
b. Plant stenophyllous, only associated with riverine habitats (facultative rheophytic); secondary venation conspicuously tessellate; inflorescences paired; stamens orange prior to and after anthesis. Bornean endemic. sig 33.90 Sic Node MBER ES Fico, 8 chee OEE OO neo © S. jelandii
34a in key = 35a, etc.
Etymology: Schismatoglottis jelandii is named for Jeland ak Kisai in recog- nition of his considerable field skills and excellent company.
2. Schismatoglottis jepomii P.C.Boyce & S.Y. Wong, sp. nov.
Ab aliis speciebus sect. tecturatae lamina spathae caducosenti differt. — Typus: Malaysia. Sarawak, Kuching Division, Lundu, Kampung Rayu, Sungai Rayu, 01° 36’ 42.6”N; 110° 08’ 59.8”E, 30 m asl, 30 August 2005, PC..Boyce, Jeland ak Kisai, Jepom ak Tisai & Mael ak Late AR-1341 (holo, SAR; iso, SING). Plate 2.
Facultative rheophytic herb to c. 45 cm tall, forming large clumps. Stem condensed, epigeal, pleionanthic, c. 8-10 mm diam., to c. 6 cm long (usually less); foliage leaves many together, each subtended by submembranous, later papery linear dark brown (when dry) cataphylls to 5 cm long; petiole 10-14 cm long, slender, terete, sheathing only at very base; sheath very shortly and truncately ligular; leaf blades spreading to erect, lanceolate,somewhat brittle- coriaceous, glossy mid- to dark green adaxially, concolorous or frequently with irregular jagged or cloudy paler green or creamy grey blotched along the midrib, abaxially paler, the base cuneate, the tip long-acuminate and terminating in a 2-5 mm long tubular mucro, 8-25 x 1-5 cm; venation more or less obscure adaxially; midrib flattened-raised adaxially, usually drying
12 Gard. Bull. Singapore 58 (2006)
with a more or less conspicuous narrow central channel adaxially, abaxially prominent; primary lateral veins, c. 14 on each side of midrib, alternating with scarcely lesser interprimaries and diverging at c. 35°; secondary venation more-or-less obscure forming a weak reticulum, tertiary venation not visible. Inflorescences (solitary to) clustered in groups of up to four, smelling pungently of acetic acid; peduncle moderately robust, much shorter than the petiole c.2 cm long and hardly emerging from the subtending cataphyll. Spathe white, c. 9 cm long; lower spathe ovoid basally becoming slender cylindrical above, c.5 cm long; limb elongate-ovate, gaping at female anthesis then crumbling and falling, distinctly incurved ventrally, c. 4 cm long, apically stiffly mucronate for c.3 mm,soon crumbling-caducous. Spadix sessile, stout, lower part cylindrical, distally weakly elongate-clavate, c.4 (lower part) to 6 mm (upper part) diam., ventrally curved, slightly shorter than the spathe, c. 7 cm long; female zone c. 5 mm long, slightly conical, obliquely inserted but otherwise entirely free from the spathe; pistils crowded; ovary squat- oblong, c. 0.75 mm diam.; style distinct, c. 0.5 mm long; stigma minutely papillate, as wide as the style; interpistillar staminodes scattered throughout the zone, clavate, c. 0.5 mm diam.; male and female zones contiguous; male zone c.2.5 cm long, cylindric; stamens truncate, partly confluent apical pores; appendix sub-cylindric, c.3.7 cm long, weakly elongate-clavate, blunt-tipped, composed of columnar trapezoid to triangular staminodes. Fruiting spathe with the lower part broadly ovoid.
Distribution: Sarawak, Kuching and Sri Aman Divisions. Habitat: Facultative rheophyte in deep sandy-loam along stream banks in lowland secondary forest. 100 m altitude.
Notes: Schismatoglottis jepomiibelongs to the Schismatoglottis tecturata group (see Hay & Yuzammi 2000: 162) based on shoot arrangement (pleionanthic with the leaf sheath very short and fully attached; foliage leaves alternating with cataphylls) but differs from the two species hitherto comprising this group (S. tecturata (Schott) Engl. and S. petri A.Hay) by the spathe limb crumbling and shedding before male anthesis. Schismatoglottis jepomii differs from S. tecturata in the considerably larger size of the inflorescence and the scattered interpistillar staminodes (arranged in a single ring at the base of the female zone in S. tecturata). From S. petri the truncate connective is immediately diagnostic.
Schismatoglottis jepomiican be fitted into the key to Bornean Schismatoglottis (Hay & Yuzammi 2000) as follows:
A trio of new Schismatoglottis from Sarawak, Borneo
Plate 2. Schismatoglottis jepomii. a. Holotype plants in habitat; b. Inflorescence at female anthesis. Note spathe already par- tially shed; ce. Inflorescence emerging from prophyll.
14 Gard. Bull. Singapore 58 (2006)
22a. Connective not or hardly elevated; appendix cylindrical or only weakly elongate-clavate, more or less isodimatric with top of male zone ............. 23
b. Connective much elevated above the thecae; appendix clavate-cylindric, distinctly thicker than male zone; Brine ...........4.-.2 22 87. S. petri
23a. Spathe persistent into anthesis, later marcescent; appendix cylindric, more or less isodiametric with top of male zone; Borneo and Riau Archipelago .s..s.:.cc0seiden-acceeeorsgseee eee: chee 88. S. tecturat
b. Spathe caducous prior to male anthesis; appendix weakly elongate-clavate; Western Sarawak. «....0...0.0030s..s8 eho eeee eventos S. jepomii
23a in key = 25a.jete:
Etymology: Schismatoglottis jepomii is named for Jepom ak Tisai, the most recent member of the Malesiana Tropicals field staff, who has an excellent pair of forest eyes and is a splendid company.
Other specimens examined: SARAWAK: Kuching Division: Bau, Segong, 12 Feb 2004, Jepom ak Tisai AR-222 (SAR); Bau, Rieng Opui, 26 May 2004, Jeland ak Kisai AR-402 (SAR); Bau, Singai, Batu Taring, 19 Jun 2004, Jeland ak Kisai & Jepom ak Tisai AR-465 (SAR); Bau, Rieng Opui, 1 May 2005, Jeland ak Kisai AR-1184 (SAR). Sri Aman Division: Lubok Antu, Batang Ai, Nanga Sumpa, Sungai Delok, 01° 11’ 40.8”N; 112° 04 04.2”E, 28 Jul 2004, P. Boyce, Jeland ak Kisai & Kachong AR-558 (SAR).
3. Schismatoglottis maelii P.C.Boyce & S.Y. Wong, sp. nov.
Ab aliis speciebus sect. multiflorae foliius profunde corrugatis differt.—Typus: Malaysia. Sarawak, Kuching Division, Bau, Segong, Ulu Sungai Adis, Sungai Bronand, 50 m asl, 4 May 2004, Jeland ak Kisai & Jepom ak Tisai AR-47 (holo, SAR). Plate 3.
Lithophytic herb to c. 50 cm tall. Stem condensed, erect to creeping, c. 2—2.5 cm thick, with internodes to c. 0.5 cm long, pleionanthic. Leaves to c. 15 together; petiole 15-25 cm long, terete, mid-green, sheathing only at the extreme base, the wings of the sheath extended into a bicarinate narrowly lanceolate free ligular portion to 13-18 cm long, drying dark brown; lamina oblong, deflexed at insertion on petiole and held subpendent, 14—25 x 5- 15 cm, deep lustrous green adaxially, abaxially almost the same, the base obtuse and slightly decurrent, the tip acute and with a tubular mucro up to 8 mm long; midrib very prominent-rounded abaxially, adaxially impressed
A trio of new Schismatoglottis from Sarawak, Borneo
Plate 3. Schismatoglottis maelii a. Holotype plants in habitat; b. Inflorescence at female anthesis; c. Leaf lamina, adaxial surface to show diagnostic venation.
16 Gard. Bull. Singapore 58 (2006)
to semi-flush with the lamina, with 5—28 primary lateral veins on each side, irregularly alternating with the prominent interprimary veins and diverging at 60-70°; secondary venation very prominently deep green-transparent abaxially, impressed adaxially, arising from the midrib and from the bases of the primary veins; tertiary venation obscure, all primary and secondary veins impressed adaxially and giving the lamina a prominently finely corrugated texture. Inflorescences 1-4 together, nodding, subtended by lanceolate cataphylls resembling the ligular leaf sheaths, strongly fragrant of crushed raspberries at anthesis; peduncle to 7 cm long, terete, pale green, not exceeding the cataphylls, deflexed at the junction of the lower spathe and long-decurrent on the spathe (corresponding to the female zone). Spathe 7.5-8 cm long; lower spathe c. 2 cm long, green, differentiated from the limb by a pronounced constriction level with the top of the interstice; limb 5.5-6 cm long, pale green with darker green veining, apex mucronate for c. 3 mm, interior sticky at female anthesis when conspicuously inflated, then crumbling-caducous at male anthesis. Spadix to 4.5 cm long, subcylindric; female zone c. 1.8 cm long, adnate to the spathe in the lower 2/3, the free part slightly conoid, apically 6.5 mm diam.; pistils numerous and crowded, subcylindric, c. 0.4 mm diam.; stigma sessile, about the diameter of the ovary, button-like, papillate; interpistillar staminodes very few among the pistils, otherwise confined to a single row along the spathe/spadix adnation and a further incomplete row at the junction of the female zone and the interstice, overtopping the pistils, elongate mushroom-shaped, round-topped; sterile interstice 6 mm long, white, somewhat obconoid, distally 6 mm diam.; staminodes of interstice crowded, irregularly polygonal, the lower most resembling the interpistillar staminodes, the upper more flattened, 0.5-1 mm diam.., flat-topped; male zone 2 cm long, cylindrical, basally isodiametric with top of interstice, tapering to a blunt point in the upper half; stamens rather dense, somewhat irregularly rectangular with the connective wide and raised and the pores small on the narrower edges of the connective, 0.7—10 mm across; appendix, c. 1 cm long, blunt-conic, composed of densely packed trapezoid to triangular irregular staminodes, the apex slightly depressed. Fruiting spathe mid-green, narrowly urceolate, up to 3 cm long.
Distribution: Sarawak, known only from the vicinity of Bau, Kuching Division. Habitat: Lithophytic on thin layers of humus and in shallow litter pockets on
sandstone boulders and cliffs, 60 — 100 m altitude.
Notes: Schismatoglottis maelii belongs to the Schismatoglottis multiflora group (see Hay & Yuzammi 2000: 84) by reason of the pleionanthic shoots,
A trio of new Schismatoglottis from Sarawak, Borneo Ey
free-ligular leaf sheath, and the inflorescence held subhorizontal by the bent peduncle and caducous spathe limb. It may immediately be distinguished from all other species in the group by the prominently raised striate veins, and from most species by the curiously brilliant green leaves.
Schismatoglottis maelii is most similar to S. mayoana Bogner & M.Hotta in possessing a sterile appendix with large staminodes and a sterile interstice but is readily differentiated by the markedly corrugated adaxial venation, smaller spadix (up to 4.5 cm in S. maelii vs. 8 cm in S. mayoana) and differ- ent inflorescence odour. The freshly opened inflorescences smell strongly of crushed raspberries in notable contrast to all other Schismatoglottis so far sampled, including species probably most closely related to S. maelii, e.g., S. nicolsonii A.Hay, S. mayoana, etc.,in which the newly opened inflorescence emits a pungent smell reminiscent of acetic acid.
Schismatoglottis maelii can be fitted into the key to Bornean Schismatoglot- tis (Hay & Yuzammi 2000) as follows:
15a. Secondary venation very fine and dense — 2 veins per mm; thecae of SUEAIOUES TSS SE TOCA GH OTR AY (209 010 | else. tee a > ee ne ee ee ee 16
b. Secondary venation density various but 1.5 veins or fewer per mm; thecae Se SUamMe MS CACH WIC OME) DONC ce. Me cecc. .o.-te-secnddesnoeiscaccacnaveresscsendsstoasstapaebents ie
16a. Spadix with a short sterile interstice of sterile stamens between the fertile zones; appendical staminodes large, ca. 1 MM ACTOSS ........ ee eeeeeeeeee i,
b. Spadix with fertile zone contiguous; staminodes small, ca. 0.3 mm across; Bako. Sabawak..0::nle nls Bee (th Bids MS cas, Cand ie Ae beta Sad 42. S. nicolsonii
17a. Spadix usually ca. 8 cm long; leaves smooth adaxially; Matang, Sarawak nésicaoaaces desea eGSGa29 0S eRe REET PRE CED CCRC 5c) Ra AR eee 39. S. mayoana
17a in Key = 18a, etc.
Etymology: Schismatoglottsis maelii is named for Mael ak Late, who is responsible for the construction and maintenance of the Malesiana Tropicals nurseries, and lately an occasional but valued member of our field team.
18 Gard. Bull. Singapore 58 (2006)
Other specimens seen: SARAWAK: Kuching Division: Bau, Kampung Jugan, Sungai Merah, 27 Apr 2004, Jeland ak Kisai AR-32 (SAR) & 6 May 2004, P.Boyce & Jeland ak Kisai AR-55 (SAR); Bau, Segong, Ulu Sungai Adis, 2 Jun 2004, Jepom ak Tisai AR-427 (SAR) & 5 Jun 2004, Jeland ak Kisai & Jepom ak Tisai AR-437 (SAR); Bau, Segong, Gunung Moi, 28 Feb 2005, Jeland ak Kisai AR-1008 (SAR).
Acknowledgements
The collaboration and support of the Sarawak Forestry Department, the Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee, the Forest Research Centre (Kuching), notably L.C.J. Julaihi, are gratefully acknowledged. Thanks are due to Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme Brown and Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd, for their considerable support and funding of fieldwork in Sarawak.
References
Hay, A. 2002. A New Bornean Species of Schismatoglottis (Araceae). Aroideana 25: 67-69.
Hay, A. and C. Herscovitch. 2003. A New Species of Schismatoglottis (Araceae) from Sabah. Gardens’ Bulletin Singapore 55: 27-30.
Hay, A. and Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia I - Schismatoglottis. Telopea 9: 1-177.
Gardens’ Bulletin Singapore 58 (2006) 19—24 19
Rhaphidophora crassifolia Hook.f. (Araceae: Monstereae): a new record for Sarawak and notes on the Rhaphidophora ‘Hongkongensis’ group in Borneo
P.C. BOYCE
Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia
Abstract
Rhaphidophora crassifolia Hook.f., a species hitherto known only from Peninsular Malaysia and southern Thailand is recorded as new for Sarawak and takes to 16 the number of species of Rhaphidophora for Borneo, of which five are endemic. A species description and photographs together with a new key to the Rhaphidophora species in Borneo is given.
Introduction
Revision work on Rhaphidophora (Boyce, 2001, 2005, 2006) in Borneo has established that there are 16 species (excluding that described here), of which five are endemic.
The most speciose group in Borneo with four species is the ‘Hongkongensis’ Group defined by shingling, non-skototropic seedling and shingling juvenile shoots, pre-adult and adult plants with climbing stems square to rectangular in cross-section, simple, often coriaceous leaf laminae, a petiolar sheath extending beyond the leaf base by short to rather long ligules and the sheath soon falling to leave a horseshoe shaped scar around the top of the apical geniculum. All species flower on free lateral shoots that are either angular or more-or-less terete in cross-section. Species in the ‘Hongkongensis’ Group are often lofty climbers and are frequently very inadequately collected.
Key to Adult Flowering Rhaphidophora in Borneo
han Mature leat lanainia pimiiately divide 2...2i...3..ereseasc.-coonnsaonesancssencceenvase 2
20 Gard. Bull. Singapore 58 (2006)
1b. Mature leaf lamina entire, with or without perforations, but never pinnately divided 6.1.4... :c¢..c:eskceech Bosna ee eet ees a eee 3
2a. Plants always associated with sandy or rocky forest streams. Flowering plants usually rheophytic, rarely low-climbing on trees beside torrential SEPSIS ch lets Ae eee cacy. nee ae R. beccarii
2b. Plants not specifically associated with watercourses.
3a. Juvenile shingling stage with falcate-lanceolate non-overlapping ascending leaves; adult plants with leaf pinnae no more than 2 cm wide and frequently much less; inflorescences solitary; spadix slender, not exceeding 13: cnvlon® «:.......ach sae ee ees. ee ee R. tenuis
3b. Juvenile shingling stage with ovate, overlapping spreading leaves; adult plants with leaf pinnae exceeding 3 cm wide; inflorescences several together;
spadix stout, upto: 25 .Cii Ome a ae R. korthalsii 4a. Geniculum and abaxial surface of lamina pubesceNt .............:eeeeeeeeeeees 5 4b. Geniculum and abaxial surface of lamina glabrous ........... ee eeeeeeeeeeees 4
Sa. Plants flowering on clinging stems. Leaves of mature plants extensively perforated, active shoot tips with black mucilage ................. R. foraminifera
5b. Plants flowering on free lateral stems. Leaves of mature plants lacking or with only with scattered perforations; active shoot tips lacking black PICHIA OY oi sas ce osiins neh Seyan a een ea ee 09 fas nosaitdaoey eae ee ee ee R. puberula
6a. Leaves always shingling, even in flowering individuals; leaf laminae stiffly coriaceous, broadly oblong-ovate-elliptic, 8-48 x 6.5—20.5 cm, bright green, slightly to markedly glaucous, base truncate-cordate to broadly cuneate. Ploweriuig’ O17 chinoine SItOOts circ 0s scenic ee a eee R. latevaginata
6b. Leaves spreading in adult and flowering individuals; leaf laminae
variously coloured but never glaucous. Flowering on free or clinging shoots
7a. Stems scabrid to asperous. Spathe exterior minutely puberulent ......... sevevedoassapoosoaatdsonubadtevavsecuqen cespier tired onmEMI- ot sate othe ttyE ds tte ame eee eee R. lobbii
Rhaphidophora in Borneo at
1 OTIN (oh cM S ERS CA RIG 7 G3: Nee ae ee ee ie ee ee ee 2
9a. Clinging stems square or rectangular in cross-section; tips of active stems with netted prophyll, cataphyll and petiolar sheath fibre.......... R. elliptifolia
9b. Clinging stems various shapes in cross-section, including square or heetaneular: tips of achive stems: WithOUt MDE: }...issccicccck.scdsstcdeentecedaccsesceeses 10
10a. Spadix stipitate; leaf lamina very thickly coriaceous ....R. conocephala
10b. Spadix sessile; leaf lamina variously textured but if thickly coriaceous Heme SY CLIK SE SSS ee nee oe tee teeta se a2 ses cs sehay leckagetesascuedccadtensaatadasdetadcsbaieiaccolieiie i]
lla. Inflorescences two, three or more together, each subtended by a prominent chartaceous prophyll and one or more chartaceous cataphyll; leaf lamina oblong-lanceolate or oblong-elliptic, entire to slightly perforated, PM AR OUST SURI e Ce ea eI ee Sat anc ak ids ARR. R. megasperma
11b. Inflorescences almost always solitary, never subtended by prominent chartaceous prophylls and cataphylls; leaf lamina variously shaped but never
pc ote Tia eae oe ae ee er etd BS trees ort 3 ceeierelhlel ok. 12 12a. Clinging stems rectangular 1N CrOSS-SECTION ...........cccceseesseesseeseeeseeeeees 13 12b. Clinging stems more-or-less terete iN CrOSS-SECTION .........ccceeseeeseeeeees 15
13a. Leaf lamina thickly coriaceous to almost fleshy, margins slightly reflexed, this becoming greatly accentuated in dried material. Clinging stems twisting to produce complex ridges ..........cceeseeeseeeeseeeeees R. crassifolia
13b. Leaf lamina thinly coriaceous, margins flat, leaf drying pale straw- coloured. Clinging stenis Mob twistime iil. 14
14a. Leaf lamina lanceolate-elliptic to falcate-lanceolate, 4.5-32 x 1.75-8.5 cm. Spadix weakly clavate-cylindrical, 3-6 cm long ...............6. R. sylvestris
14b. LEAF LAMINA, NARROWLY ELLIPTIC TO NARROWLY ELLIPTIC-OBLONG, 20-29 X 4.5-7 cm. Spadix cylindrical, 8-14 cm LONG ....... eee eeeeesseeeteeeees R. elliptica
22 Gard. Bull. Singapore 58 (2006)
15a. Leaf lamina narrowly falcate-elliptic to falcate-lanceolate or falcate- oblanceolate, 2.5-16 x 1.2-3 cm, drying uniformly pale straw-coloured. Spadix slender cylindrical, 2-5-7 emi ome 2.102550 ee ee ee R. minor
15b. Leaf lamina subfalcate-lanceolate or oblong-elliptic, oblique, 10-19.5 x 2.5-6.5 cm, drying very dark brown. Spadix cylindrical-obtuse, 3-10 x 1 cm (fruiting specimen only) os.20e es 1 R. cylindrosperma
Rhaphidophora crassifolia Hook.f.
Rhaphidophora crassifolia Hook.f., Fl. Brit. Ind. 6 (1893) 543; Ridl., Mat. FI. Malay Penins. 3 (1907) 42; Engl. & K. Krause in Engl., Pflanzenr. 37 (IV.23B) (1908) 22, Fig. 5; Ridl., Fl. Malay Penins. 5 (1925) 122— Type: Malaysia, Perak, Batang Padang District, Batang Padang, Aug 1884, Kunstler 8111 (holo, K).
Large, occasionally very large, moderately robust, semi-pachycaul homeophyllous liane to 10 m; seedling stage a non-skototropic shingling shoot; pre-adult plants rarely forming small terrestrial colonies of appressed shingling shoots; adult shoot architecture comprised of greatly elongated, clinging, physiognomically monopodial, leafy, non-flowering stems and short to moderately elaborated, free, sympodial, densely leafy, flowering stems; stems smooth, climbing stems rectangular in cross section, the angles winged, the surfaces between sulcate, the stem twisting to produce a complex series of ridges and channels, free stems more or less terete to weakly four-angled in cross section, often branching extensively, growing to considerable lengths and then pendent under their own weight, green, later dull brown, without prophyll, cataphyll and petiolar sheath fibre but active apices coated with clear mucilage, internodes to 10x 2cm on adherent shoots, usually shorter and less stout on free shoots, separated by prominent slightly oblique leaf scars, older stems woody; flagellate foraging stems absent; clasping roots densely arising from the nodes and internodes of clinging stems, densely pubescent; feeding roots rather rare, adherent, pubescent; leaves weakly spiralled on adherent and proximal portions of free shoots, densely distichous distally on flowering shoots; cataphylls and prophylls membranous, very quickly drying and falling; petiole shallowly canaliculate to grooved adaxially, 4-7 x 0.2-0.3 cm, smooth, with a slight apical and somewhat prominent basal geniculum; petiolar sheath prominent, extending to and encircling the apical geniculum, very swiftly drying and falling to leave a continuous scar from the petiole base, around the top of the apical geniculum and back to the base; lamina entire, falcate-elliptic-lanceolate to falcate-oblong or falcate-oblanceolate, 2.5-35 x 1-10 cm, thickly coriaceous to almost fleshy, upper surfaces glossy,
Rhaphidophora in Borneo 23
lower surfaces matte, base sub-ovate to acute or briefly decurrent, apex sub-acute with a prominent apiculate tubule, margins slightly reflexed, this becoming greatly accentuated in dried material; mid-rib barely raised abaxially, slightly sunken adaxially; primary venation pinnate, slightly raised abaxially and adaxially; interprimaries parallel to primaries, very slightly raised abaxially and adaxially; secondary and tertiary venation + invisible in fresh material, barely visible in dried specimens; inflorescence solitary, subtended by a fully developed foliage leaf; peduncle compressed-cylindric- clavate, 1.7—2.5 x 0.2—0.3 cm; spathe cigar-shaped, stoutly short-beaked, 4—6.5 x 1-1.7 cm, thickly fleshy, dull green to dull yellow, swiftly falling at female receptivity; spadix cylindrical, sessile, inserted level on peduncle, 3.5-6 x 0.5—1.3 cm, dirty white; stylar region rather well developed, mostly rhombo- hexagonal, 0.8-1 x c. 1 mm, truncate; stigma punctiform, c. 0.3 mm diam.; anthers barely exserted at anthesis, pollen extruded from between ovaries; infructescence not observed.
Distribution: Peninsular Malaysia (Negri Sembilan, Perak, Selangor), southern peninsular Thailand (Narathiwat), Borneo (Sarawak: Miri).
Habitat: Primary lowland rainforest, dry Dryobalanops forest, peatswamp forest, wet evergreen forest, frequently but not exclusively associated with Karst limestone. 10-90 m altitude.
Notes: The remarkably thick leaves and complex patterns of ridges and channels (Plate 1) resulting from the twisting of the stems immediately distinguish R. crassifolia from the two most similar species in Sarawak, R. elliptica Ridl. and R. elliptifolia Merr. Rhaphidophora crassifolia may be further distinguished from R. elliptifolia by the lack of fibrous remains on the tips of the active shoots and from R. elliptica by the smaller (3.5-6) spadix (8-14 cm long in R. elliptica).
Other specimens examined: SARAWAK: Miri, Niah Suai, Niah National Park, trail to Great Cave, 03° 49’ 21.7”; 113° 45°44.1”, PC Boyce, Jeland ak Kisai & Jepom ak Tisai AR-1464 (SAR). PENINSULAR MALAYSIA: Perak: Padang, Batang Padang, Kunstler (‘Dr King’s Collector’) 8111 (K, holo); Wray 2260 (SING); Selangor: Genting Sempah 22 mile, Ulu Gombak ER., Stone 5738 (KLU); Batu Cave ravine, Nicolson 1148 (US); Negeri Sembilan, Pasoh ER., La Frankie 2185 (KEP); Johore, Pengkalan Raja, Pontian, Ngadiman SFN 36774 (B, BO, L, K, P, SING); Sg Kayu Ara, Mawai - Jemaluang road, Corner SFN 29318 (KEP, K, SING). THAILAND: PEN72. Narathiwat, Budho-Phadee N.P, Nam Tok Chatwarin, 6°06’N, 101°50’E, Boyce 1226 (BKF,K).
24 Gard. Bull. Singapore 58 (2006)
Plate 1. Rhaphidophora crassifolia Hook.f. Note the ridges along the twisted stem and coriaceous leaf.
References
Boyce, P.C. 2001. The genus Rhaphidophora Hassk. (Araceae-Monstero1- deae-Monstereae) in Borneo. Gardens’ Bulletin Singapore 53: 19-74.
Boyce, P.C. 2005. A New Species of Rhaphidophora Hassk. (Araceae: Monstereae) from Borneo. Gardens’ Bulletin Singapore 57: 211-216.
Boyce, P.C. 2006. Rhaphidophora tenuis (Araceae: Monstereae) Resur- rected. Gardens’ Bulletin Singapore 58: 1-6.
Gardens’ Bulletin Singapore 58 (2006) 25—40 25
A review of entire-leaved Tacca (Dioscoreaceae) in Sarawak, Borneo
PC. BOYCE AND S.JULIA’
‘Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia
‘Forest Research Centre, Km 10 Jalan Datuk Amar Kalong Ningkan, 93250 Kuching, Sarawak, Malaysia
Abstract
A review of entire-leaved Tacca in Sarawak is presented; four species are recognized. Tacca borneensis Ridl. is resurrected and problems concerning the interpretation of T: integrifolia sensu Drenth are reviewed. The first complete description of 7: bibracteata Drenth is published. One new species, T. reducta P.C.Boyce & S.Julia, is proposed. Significant floral and fruiting morphologies are highlighted and a key to all Tacca species (entire and palmate/dracontioid- leaved) in Sarawak is provided. All entire-leaved species in Sarawak are illustrated.
Introduction The most recent complete revision of Jacca J.R. & G.Forst (including Schizocapsa Hance) was by Drenth (1972), with regional accounts for Flora Malesiana (Drenth, 1976) and Flora of China (Ling, 1985; Ding et al., 2000). All these accounts follow Drenth (1972) in treating T: integrifolia Ker Gawl as a widespread and highly polymorphic species.
Drenth (1972, 1976) recognizes two entire-leaved Tacca species for Sarawak, viz. Tacca bibracteata Drenth and T. integrifolia, in addition to two compound-leaved species: T. leontopetaloides (L.) Kuntze & T. palmata Blume. As is inevitably the situation with wholly herbarium-based monocot, family accounts, subsequent fieldwork has revealed considerable problems interpreting names and a far too broad circumscription of species. In particular there are considerable problems with the circumscription of T: integrifolia, which, far from being a highly variable species, in Sarawak, divides incontrovertibly into three morphologically and, in one instance, an ecologically — limestone -, a distinct species.
26 Gard. Bull. Singapore 58 (2006)
Characters of Taxonomic Significance
During our studies the following characters have proven to be of diagnostic value:
Involucral bracts
Outer and inner pair: heteromorphic or homeomorphic. Outer pair position: lateral or dorso-ventral. Outer pair position: lateral, lateral-ascending or dorso-ventral.
Perianth
Lobes of inner and outer perianth + equal in size or dissimilar. Perianth lobes soon marcescent or persistent into fruiting.
TACCA ON SANDSTONE AND SHALE IN SARAWAK
Tacca integrifolia Ker Gawl & T. borneensis Ridl.
Sarawak has two non-limestone associated Tacca species with large involucral bracts. One has heteromorphic bracts and is discussed in detail later in this paper. The other, with homeomorphic bracts and a truncate, oblique leaf base equates to T- borneensis Ridl., and although treated as a synonym of T. integrifolia by Drenth (1972), is distinct and is here resurrected from 7: integrifolia sensu Drenth.
Tacca borneensis Ridl., J. Straits Branch Roy. Asiat. Soc., 49(1): 45 (1907). Type: Malaysia, Sarawak, Kuching Division, Matang, Aug 1905, Ridley s.n. (holo, SING!). Plate 1a.
Moderate to robust terrestrial herb to 1.2 m. tall. Stem rhizomatous, hypogeal, creeping with the active apex ascending, up to 3.5 cm thick, clothed with persistent leaf bases and frequently rooting through these. Leaves 5 — 15 together; petioles ascending, up to 41 cm long, c. 12 mm diam, sub-terete- in overall cross-section, pronounced deeply canaliculate, sharply 2-keeled on the dorsal side, mid-green, stained deep purple-brown at the base; petiolar sheath c. '« —'/length of petiole, hyaline; lamina ascending to recurved, oblong-lanceolate to oblong, 23 — 65 x 10 — 24 cm, base oblique, broadly
Tacca in Sarawak
Plate la. Tacca borneensis. Note the large homeomorphic involucral bracts. lb. Tacca integrifolia. W Sarawak form. 1b. Tacca integrifolia NE Sarawak form. Note the rather fleshy inner involucral bracts.
28 Gard. Bull. Singapore 58 (2006)
ovate to unequally weakly cordate, apex acuminate, margins smooth, lamina glossy mid-to deep green, paler and less glossy abaxially; mid-rib strongly raised abaxially, sunken adaxially, primary lateral veins 7 - 9 per side, prominent abaxially, sunken adaxially, interprimary lateral veins less prominent than primaries, secondary veins forming a tessellate network with a variably pronounced interprimary collecting vein running through the middle of each trans-interprimary area. Inflorescence solitary to up to 5 at different developmental stages per plant; peduncle sub-erect to decumbent and apically ascending at anthesis, 3 — 5 angled or weakly 3 — 5 winged, up to 45 cm long, mid-green to more-or-less wholly stained purple-brown; involucral bracts homeomorphic; outer pair dorso-ventrally positioned, ovate-triangular, basally briefly clawed, 5.5 — 7.5 x 6 — 9 cm, velvety very deep purple-black, rarely pale purple with deep purple veining, claw paler, occasionally almost white; inner pair laterally positioned, ovate to broadly ovate, clawed basally, 5 — 10 x 5.5 — 12 cm, deep lustrous purple-black, claw paler to almost white; filiform bracts 10 — 25 or more per inflorescence, 12 — 15cm long, deep purple. Flowers 5 — 25 (— 30) per inflorescence; pedicel triangular in cross-section, 2.5 —4 cm long, dark purple, initially erect, later in anthesis pendent; gynoecium obpyramidal, c. 1.5 cm long x c. 1.3 cm wide at apex, 6-ribbed, purple with the ribs darker, perianth inserted annularly onto top of gynoecium; outer perianth lobes rounded, c. 10 x 9 mm, reflexing at anthesis, velvety deep purple; inner perianth lobes ovate, c. 10 x 10 mm, deep velvety purple. Infructescence prostrate by twisting of the peduncle base, many-fruited, involucral bracts marcescent well prior to fruit maturation; fruits obpyramidal, c. 3.5 x 1.5 cm, semi-glossy deep purple. Seeds weakly laterally compressed-reniform, c. 3.5 x 1.5 —2 mm, pale brown.
Distribution: Sarawak. Endemic, based on known herbarium collections, but most likely occurring also in Kalimantan Barat.
Habitat: Old or disturbed secondary lowland forest on sandstones, very rarely on limestone, but then never on exposed rocks. 40 — 450 m altitude.
Other specimens examined: SARAWAK. Kuching Division: Padawan, Kampung Belimbing, 28 Nov 2003, PC.Boyce & Jeland ak Kisai TA-5 (SAR); Bau, Kampung Jugan, 26 Mar 2004, PC. Boyce & Jeland ak Kisai, TA- 7 (SAR); Padawan, Kampung Sadir, 2 Feb 2006, PC. Boyce & Simon Kutuh ak Paru TA-34 (SAR); Bau, Plaman Kaman, 26 Sep 2000, K.G. Pearce et al. SBC 21 (SBC); Bau, Bukit Kho Z San, Km 1 % Bau — Kuching road, 20 Dec 1994, Rantai Jawa et al. S70124 (SAR); Padawan, Gunung Merubong, Ulu Sungai Sluba, 18 Sep 1987, Yii Puan Ching S.51396 (K, L, SAR). Samarahan
Tacca in Sarawak 29
Division: Serian, Gunung Ampungan, 21 Nov 2003, PC. Boyce & Jeland ak Kisai TA-4 (SAR); Serian, Tebedu, Kampung Saan, 4 Oct 2004, PC. Boyce & Simon Kutuh ak Paru TA-13 (SAR); Serian, Pichin, Bung Biringan, 28 Oct 2004, PC.Boyce & Simon Kutuh ak Paru TA-15 (SAR); Serian, Pichin, Utak Ogong (Ogong Amang Ramping), 5 Jan 2005, PC. Boyce & Simon Kutuh ak Paru TA-18 (SAR). Sri Aman Division: near Sungai Kuap, Ulu Sungai Engkari, 21 Mar 1974, P Chai S34090 (L, K, MO, SAR, USA). Kapit Division: Rejang, G.D.Haviland 936 (SAR). Miri Division: Baram, J. Hewitt 593 (SAR). Bintulu Division: tau Range, Sungai Mayeng, 2 Jun 1956, J. W. Purseglove P.5300 (SAR).
Notes: The oblique, broadly ovate to unequally weakly cordate leaf bases immediately distinguish 7? borneensis from all other entire-leaved Sarawakian Tacca in which the leaf bases are acute and decurrent. The large ovate homeomorphic involucral bracts of 7: borneensis are diagnostic.
Resurrection of 7: borneensis in Sarawak delimits a non-limestone associated Jacca with large heteromorphic involucral bracts. For the present, with the caveat that there appears to be at least two taxa involved (one in W Sarawak and the other in NE Sarawak), and that these require further field study to ascertain their appropriate status, we are maintaining this heteromorphic bracted plant as a single taxon for which the earliest name applicable name 1s 7? integrifolia.
Tacca integrifolia Ker Gawl, Bot. Mag., 35, t.1488 (1812). Type: ‘East Indies’, (holo, K!; iso, L!). Plate 1b, ec.
Slender to moderately robust terrestrial herb to 75 cm tall. Stem rhizomatous, hypogeal, creeping with the active apex ascending (W Sarawak) or epigeal (NE Sarawak), up to 2 cm thick, clothed with persistent leaf bases and fre- quently rooting through these. Leaves 5 — 15 together, petioles ascending, up to41 cm long,c. 12mm diam, D-shaped in overall cross-section, canaliculate,2- keeled on the dorsal side, pale to mid-green sometimes stained purple at the base; petiolar sheath c. / length of petiole, hyaline; lamina thinly to rather thickly coriaceous, ascénding, lanceolate, 11 — 45 x 5 — 12 cm, base acute, decurrent, c. 5 cm along petiole, apex acute to acuminate, margins smooth, lamina pale to mid-green, weakly glossy adaxially, paler and less glossy abaxially; mid-rib prominently raised abaxially, sunken adaxially, pri- mary lateral veins 3 —5 per side, slightly sunken adaxially, interprimary lat- eral veins slightly less prominent than primaries, secondary veins forming a very obscure network with an obscure interprimary collecting vein running through the middle of each trans-interprimary tessellate area. Inflorescence
o>) _
Gard. Bull. Singapore 58 (2006)
solitary; peduncle erect, weakly 3 angled, up to 75 cm tall but usually less, especially in plants from NE Sarawak, pale green stained deep purple espe- cially near the base; involucral bracts heteromorphic: outer pair dorso-ven- trally positioned, narrowly ovate, 2.5—3.5 cm x 10-22 mm, deep purple-black to pale lavender purple, rarely white with lilac veining; inner pair laterally positioned but ascending and ultimately sub-erect at anthesis, spathulate, 3.5 —9 x 2-6 cm, deep purple-black to pale lavender purple, rarely white with lilac veining; filiform bracts 8-15 per inflorescence, 12—19cm long, pale to mid- or deep purple basally, fading to at the tip. Flowers 7-14 per inflo- rescence; pedicel triangular in cross-section, 24cm long, pale to mid-purple, initially erect, later in anthesis reflexing, thence pendent; gynoecium widely obpyramidal,c. 1 cm long x c. 1 cm wide at apex, 6-ribbed, greenish-purple to dark purple, the ribs darker purple, perianth inserted annularly onto top of gynoecium. Outer perianth lobes oblong, rounded 10-17 x 5-7 mm, reflex- ing at anthesis, dark purple; inner perianth lobes oblong, 12-17 x 6-9 mm, deep velvety purple on both surfaces. Infructescence declinate by twisting of the peduncle base, few to many-fruited, involucral bracts marcescent well- prior to fruit maturation; fruit obpyramidal, deep glossy brown-purple c. 3 x 1.5 cm, dull purple. Seeds weakly laterally compressed-reniform, c. 3.5 x 1.5 —2 mm, pale brown.
Distribution: Following the taxonomic interpretation used here, the distribution is southern Peninsular Thailand, Peninsular Malaysia, Singapore and N Borneo.
Habitat: Old or disturbed secondary lowland forest on sandstones. Very occasionally associated with limestone formation but never on exposed limestone rock, 40-420 m altitude.
Other specimens examined: SARAWAK. Kuching Division: Bau, Tanjung Durian, 14 Nov 2003, PC Boyce TA-3 (SAR); Bau, Serikin, PC Boyce TA- 37 (SAR); Kuching, Dec 1906, J. Hewitt 592 (SAR); Matang, Sungai Rayu, 27 Mar 1987, Bernard Lee Meng Hock 8.53351 (K, SAR); 10 Mile Landeh Road, Engkabang Plantation, 26 Feb 1974 S. Laijanai S.33648 (SAR, USA). Samarahan Division: Serian, Pichin, Kampung Kakang, Sungai Sisang, 14 Jan 2005, PC. Boyce & Simon Kutuh ak Paru TA-19 (SAR); Serian, Pichin, Ampan Pichin, 25 May 2005, PC. Boyce & Simon Kutuh ak Paru TA-36 (SAR). Sarikei Division: Sungai Lepong, 01° 57’ 12.9”; 111° 30’ 34.9”, 8 Dec 2005, P.-C. Boyce, Jeland ak Kisai, Jepom ak Tisai, Mael ak Late & Wong Sin Yeng TA-33 (SAR).
Tacca in Sarawak Sil
Bintulu Division: Bukit Satiam, 02° 59’ 26.1”; 112° 55’ 54.4”, 11 Aug 2004, P.C.Boyce & Jeland ak Kisai TA-10 (SAR). Miri Division: Baram, Sept. 1891, C._Hose 1357 (G.D.Haviland 949) (SAR); Marudi, Sungai Slat basin, Sungai Palutan, 2° 50’ 32”; 114° 59° 12”, SPLim & Banyeng L. S.90414 (SAR, KEP).
Notes: Drenth (1972) took a very broad view of the circumscription of 7: integrifolia, synonymizing 10 taxa described from as far apart as NE India to NE Borneo. The present paper is not intended as a critical revision of all the names so treated, but observation of Tacca in Sarawak coupled with knowledge of the level of endemism in ever-wet Sunda of other herbaceous monocots (notably Araceae and Zingiberaceae) are pertinent in suggesting that 7: integrifolia sensu Drenth is a grossly heteromorphic assemblage.
A brief discussion of the taxa treated as synonymous with 7: integrifolia by Drenth (1972) is insightful. Six names attributed to the synonymy of T. integrifolia (Tacca aspera Roxb., T: choudhuriana Deb., T. integrifolia var. pseudolevis Limpr, and T. laevis Roxb. [including var. angustibracteata Limpr. and var. latibracteata Limpr.]) are based on types originating from NE India and Bangladesh, and one (T: integrifolia var. pseudolevis Limpr.) on a type from NW Myanmar. Given the fact that in families, e.g., Araceae and Zingiberaceae — with pronounced bipolar diversity, and endemism in ever-wet Sunda and Indo-China, and tropical and subtropical trans-Himalaya — there are no shared indigenous species between Borneo and the Indo-Himalaya, it is unlikely that Tacca from these areas are synonymous with species present in Borneo.
There are three distinct entire-leaved Tacca species in Peninsular Malaysia and Singapore, one with heteromorphic and two with homeomorphic involucral bracts. The earliest available name for the heteromorphic bracted species is T: integrifolia, and for the homeomorphic bracted, T: cristata Jack and T: chantrieri André. More fieldwork is required to ascertain whether additional taxa require recognition, to clarify the presence, or otherwise, of 7? chantrieri, and to further its status vis a vis other published names for Thailand and Indo-China. In particular 7. minor Ridl. (treated as one of nine synonyms of T: chantrieri by Drenth in 1972) requires field investigation to clarify its status.
Four names (7? sumatrana {incl. var. ovalifolia Limpr.]) and (T: lancifolia {including var. laeviformis Limpr.]) originate from Sumatera and Java respectively. None of the types of these names is in a sufficient state of preservation to place them without question in any one of the known taxa
32 Gard. Bull. Singapore 58 (2006)
for these islands. More fieldwork is required.
The spathulate and ascending inner involucral bracts are diagnostic for T: integrifolia as interpreted here. Inflorescence colour is variable with the involucral bracts and perianth ranging from deep glossy purple through to pale lavender to lilac-flushed white. The ‘white’ form is in cultivation un- der the illegitimate name ‘7! nivea’.
Plants from NE Sarawak differ from those in NW Sarawak by an epigeal stem, distinctly more leathery leaf, shorter peduncle, and somewhat fleshy, glossy, deep purple inner involucral bracts (Plate 1c). More fieldwork is required to investigate the taxonomic significance of these characters.
Tacca bibracteata Drenth
The type description of T: bibracteata (Drenth, 1972) was based on only three herbarium specimens and lacking details of the stem and ripe fruits. The first author has had the opportunity to collect and bring 7? bibracteata into cultivation and we are now able to furnish a more detailed description.
Tacca bibracteata Drenth, Blumea, 20(2): 395 (1972 publ. 1973) & in FI. Mal. Series 1, 7(4): 814, Fig.7 a — c (1976). Type: Malaysia, Sarawak, Kapit Division, Belaga, Long Kapa, Bukit Dulit, Synge 1335 (holo, K!; iso, L!). Plate 2 a,b.
Moderate terrestrial herb to 40 cm tall. Stem rhizomatous, ascending and ultimately epigeal, c. 2 cm thick, clothed with persistent leaf bases. Leaves c. 6-8 together, petioles ascending, 5-19 cm long, c. 2-5 mm diam, D-shaped in overall cross-section, sharply sulcate-canaliculate, bluntly keeled on the dorsal side, pale to mid-green; petiolar sheath c.'4—% length of petiole (3.5— 9.5 cm long), hyaline and shortly (c. 2mm) ligulate; ligule dolabriform-acute, margins minutely erose; lamina ascending to somewhat spreading, oblong- lanceolate,16—27 x 7-10.5 cm, base acute, apex shortly acuminate, margins smooth, lamina deep glossy green adaxially, paler and less glossy abaxially; mid-rib strongly pronounced abaxially and sunken adaxially; primary lateral veins c. 4 per side, very pronounced abaxially, all arising from the basal half of the mid-rib, interprimary lateral veins absent, secondary veins forming a rather obscure untidy reti-tesellate network and with a weak interprimary collecting vein running through. Inflorescence solitary; peduncle + terete, 20 —31 cm tall, pale green lightly to heavily dark purple-mottled, intensifying
Tacca in Sarawak
Plate 2a. Tacca bibracteata. Note the inner involucral bracts nearly indistinguishable from the filiform bracts, the inflorescence giving the impression of only two involucral bracts. 2b. Close up of flowers.
34 Gard. Bull. Singapore 58 (2006)
towards the base; involucral bracts strongly heteromorphic; outer pair dorso- ventrally positioned, ovate, c. 3 x 2 cm, briefly decurrent basally and fused to form a short pocket, mid-green with the primary veins and apical portion stained purple; inner pair laterally positioned, filiform, up to 10.5-14 cm x c. 2 mm, basally mid-purple fading to pale yellow-green at the tips; filiform bracts 8 - 13 per inflorescence, very variable in length with c. 2.5-14 cm lengths in one inflorescence, pale green. Flowers up to 10 but usually fewer (5 or less) per inflorescence; pedicel triangular in cross-section, 1—3.5 cm long, pale greenish purple; gynoecium obpyramidal, c. 1.5 cm long, strongly 6-ribbed, pale greenish-purple with the ribs dull purple, perianth inserted annularly onto top of gynoecium; outer perianth lobes broadly ovate, 7-12 x 5-14 mm, reflexing at anthesis green purple-stained abaxially, deep velvety purple adaxially; inner perianth lobes ovate, tip notched, 10 x 5 mm, bright green abaxially, deep velvety purple adaxially. Infructescence declinate by twisting of the peduncle base, almost always solitary fruited, fruit subtended by the persistent outer involucral bracts, ovary obpyramidal, deep c. 2.5 x 1.5 cm, dull purple, topped by the persistent outer and basal half of the inner perianth lobes. Seeds not observed.
Distribution: NE Sarawak. Endemic. Habitat: Old or disturbed secondary lowland forest on shales, 30-100 m altitude.
Other specimens examined: SARAWAK. Kapit Division: Pelagus, Jeram Pelagus, 02° 11’ 59”; 113° 04 01”, 1 Dec 2004, Jeland ak Kisai TA-16 (SAR) & 02° 11’ 35.7”; 113° 03’ 30.08”, 15 Mar 2005 P.-C. Boyce, Jeland ak Kisai & Jepom ak Tisai TA-21 (SAR); Belaga, Long Kapa, Bukit Dulit, Richards 1569 (K). Bintulu Division, Sebauh, Kampung Tubau, Sungai Tubau, P Ashton S.18369 (K, L, SAR, SING); Bintulu, Bukit Satiam, 02° 59’ 26.1”; 112° 55’ 54.4”, 11 Aug 2004, PC. Boyce & Jeland ak Kisai TA-11 (SAR). Limbang Division, Tg. Long Amok, Sungai Ensungei, Ulu Medamit, 19 Sep 1980, R. George et al. $.42879 (L, K, KEP, SAN, SAR).
Notes: The epithet bibracteata is misleading. The inflorescence of T. bibracteata has four involucral bracts, as indeed realized and noted by Drenth (1972, 1976). The outer pair is dorso-ventrally positioned and conspicuous, while the inner pair is laterally positioned, filamentous and in the absence of close examination of fresh material easily mistaken for filiform bracts.
The outer perianth and basal parts of the inner perianth are persistent into fruit maturity. This character is shared with T. reducta from which T:
Tacca in Sarawak 35
bibracteata is readily separable by the strongly heteromorphic involu- cral bracts and a preference for shales; 7: reducta is limestone associated.
TACCA ON LIMESTONE IN SARAWAK
Fieldwork on the limestone formations of W Sarawak (Bau, Padawan and Serian) has revealed that the common limestone-associated Tacca does not have a published name. It is here formally described.
Tacca reducta P.C. Boyce & S. Julia, sp. nov.
Ab omnibus speciebus Tacca_ borneensibus combinatio bractiorum involucralis idem et valde minoribus et in habitu calcicola differt. -TYPUS: Malaysia, Sarawak, Kuching Division, Bau, Kuching — Bau road, Gunung Serambu, 1 Sep 1976, PJ. Martin S.37798 (holo, SAR). Plate 3 a -c.
Slender to moderately robust terrestrial herb to 50 cm tall. Stem rhizomatous, hypogeal, creeping with the active apex ascending, c. 1.5 cm thick, clothed with persistent leaf bases and frequently rooting through these. Leaves 7-10 together; petioles ascending, up to 11 cm long, c. 5 mm diam, D-shaped in overall cross-section, shallowly to rather pronounced canaliculate, sharply 2-keeled on the dorsal side, mid-green variously speckled and stained deep purple-brown notably at the base and along the dorsal Keels; petiolar sheath c. '/s length of petiole, hyaline; lamina ascending to recurved, lanceolate, 23-30 x 5-9 cm, base acute, decurrent c. 3 cm along petiole, apex acute to acuminate, margins smooth to very slightly crenulate-dentate, lamina mid-green, slightly glossy adaxially, paler and less glossy abaxially; mid- rib strongly subterete to raised abaxially, sunken adaxially, primary lateral veins 3-5 per side, slightly sunken adaxially, interprimary lateral veins much less prominent than primaries, secondary veins forming a rather prominent tessellate network with a obscure interprimary collecting vein running through the middle of each trans-interprimary tessellate area. Inflorescence solitary to 3 at different developmental stages per plant; peduncle terete to weakly 3-5 angled, up to 20 cm tall, pale green more-or-less wholly stained and speckled deep purple-brown; involucral bract homeomorphic: outer pair dorso-ventrally positioned, linear-triangular to linear ovate, 2.5—3.5 cm x 5-7 mm, slightly lustrous very deep purple-black; inner pair laterally positioned, linear-triangular to linear ovate, 3.5—4 cm x 0.5-10 mm, deep lustrous purple-black; filiform bracts c. 8 per inflorescence, very variable in length with 12-15 cm long, deep purple basally, fading to pale green at the tip. Flowers 2-4 (-5) per inflorescence; pedicel triangular in cross-section,
36 Gard. Bull. Singapore 58 (2006)
1.5-2 cm long, mid-purple, initially erect, later in anthesis reflexing, thence pendent; gynoecium broadly obpyramidal, c. 1 cm long x c. 1 cm wide at apex, strongly 6-ribbed, dull greenish-purple with the ribs darker purple, perianth inserted annularly onto top of gynoecium; outer perianth lobes oblong, rounded with a brief acumen, c. 12 x 7 mm, reflexing at anthesis, purple with a darker purple reticulations, deep velvety purple adaxially; inner perianth lobes ovate, 15 x 10 mm, deep velvety purple on both surfaces. Infructescence declinate by twisting of the peduncle base, few- fruited, involucral bracts marcescent well-prior to fruit maturation; fruit obpyramidal, deep slightly glossy purple c.2.5 x 1.5 cm, dull purple, topped by the persistent whole outer perianth and basal half of the inner perianth lobes. Seeds weakly laterally compressed-reniform, c. 3.5 x 1.5—2 mm, pale brown.
Distribution: W Sarawak. Based on known herbarium collections, it is an endemic, but probably (based on plants observed for sale at border markets) also occurring on limestone in Kalimantan Barat.
Habitat: Primary to disturbed secondary lowland forest on limestone, 35- 220 m altitude.
Other specimens examined: SARAWAK. Kuching Division: Bau, Seburan, 6 Aug 1961, J.A.R.Anderson S.2411 (SAR); Bau, Seburan, 31 Oct 1964, J.A.R. Anderson §.26854 (SAR); Bau, south of Bukit Seburan, 30 Apr 1966, J.A.R. Anderson S.25140 (SAR); Bau, Seburan, gully between Seburan and Bukit Krian, 15 Jul 1964, .A.R. Anderson & N.G. Bisset S.20261 (SAR); Bau, Gunung Juita, 01° 23’ 48.7”; 110° 08’ 07.2”, 28 Oct 2005, P.-C. Boyce, Jeland ak Kisai, Angeline anak Simon & Wong Sin Yeng (SAR); Padawan, Kampung Danu, Gunung Temuang, Sungai Abang, 01° 15’ 38.6”; 110° 15’ 31.4”, 16 Feb 2006, P.C. Boyce, Jeland ak Kisai & Wong Sin Yeng TA-38 (SAR); Bau, Kampung Bogag, Gunung Tibugai, 01° 21’ 31.1”; 110° 03’ 48.7”, 31 Mar 2005, PC. Boyce, R.Kneer & Jeland ak KisaiTA-22 (SAR); Serian, Kampung Selabi, Sungai Mawang, 2 Feb 2006, PC. Boyce & Simon Kutuh ak Paru TA- 35 (SAR); Bau, Gunung Poing, 13 May 2002, Malcom D. et al. SBC 1548 (SAR, SBC); Padawan, Bukit Manok, mile 38 old Padawan road, 2 Sep 1979, J.P.Mamit 841074 (L,SAR, USA); Bau, Jambusan, Gunung Batu, 20 Feb 2002, K. Meekiong et al. SBC 1681 (SAR, SBC); ); Bau, Jambusan, Gunung Jebong, 6 Mar 2002, K.Meekiong et al. SBC 1965 (SBC); Bau, Krokong, Gunung Tabai, 12 March 2002, /.S.Steven et al. SBC 1995 (SAR, SBC). Samarahan Division: Serian, Pichin, Umon Murut, Tiab Belanting, 01° 08’ 03.7”; 110° 27° 00.3”, 28 Jun 2005, P.-C. Boyce, Jeland ak Kisai & A.Shafreena TA-24 (SAR); Samarahan, Kuap, Pangkalan Kuap, 01° 26’ 16.7”; 110° 22’ 18.9”, 25 Oct 2004, PC. Boyce, Jeland ak Kisai, Angeline anak Simon & Wong Sin
Tacca in Sarawak
Plate 3a. Tacca reducta. Note the stellate arrangement of the small homeomorphic involucral bracts. 3b. close up of flowers. 3c. Nearly ripe fruit. Note the persistent perianth.
1S) CO
Gard. Bull. Singapore 58 (2006)
Yeng TA-28 (SAR); Serian, Pichin, Bung Biringan, 28 Oct 2005, PC. Boyce & Simon Kutuh ak Paru TA-14 (SAR); Serian, Pichin, Ampan Pichin, 25 May 2005, PC. Boyce & Simon Kutuh ak Paru TA-23 (SAR).
Notes: The small homeomorphic black stellate involucral bracts are immediately diagnostic. Fruiting plants are recognizable by the persistent perianth, a character otherwise found only in T. bibracteata from NE Sarawak.
Tacca reducta bears some resemblance to W Malaysian T. minor Rid. (treated by Drenth (1972) as a synonym of T: chantrieri) in the narrowly triangular homeomorphic involucral bracts, but is readily separable by the pronounced tessellate secondary veins.
The epithet comes from the Latin, reductus, reduced, in allusion to the small involucral bracts and generally few-flowered inflorescence.
Key to Tacca in Sarawak
la. Mature leaf lamina palmate or dracontioid (elaborated forms of sagittate, hastate or trisect leaves in which the anterior and posterior divisions are highly dissected and:subdivided))2..2.. ...icsnccn ee Z
1b. Mature leat lamina entire ieee ccc eee 3
2a. Mature leaves dracontioid. Ripe fruits green, ribbed. Plants of coastal forest on-almost pure Sami ec. -:.-<c-sicnne eer ee T. leontopetaloides
2b. Mature leaves palmate. Ripe fruits red, smooth. Plants of a variety of
habitats, but never in Coastal forest on Sand ....2.....02.-5208 T. palmata 3a. Involucral bracts stronsely heteromorpihic .....25..22:.0 ee ee 4 3b. Involucral bracts + -homieoniorphic .....2.0:..5.252. 220 5
4a. Outer involucral bracts ovate; inner filiform and not readily distinguished from filiform floral bracts (i.e., inflorescence with the appearance of only one pair of involucral bracts). Perianth persistent until fruit maturity. Plants Of SHANG wisi ccc isidte. i ssscndle te tadeane ae ee T. bibracteata
4b. Outer involucral bracts ovate; inner spathulate, ascending. Perianth marcescent early in fruit development. Plants of sandstone and (rarely)
Tacca in Sarawak 39
iii SUI aren bi edna A. etree cunts. 4. si. k se sdevaelavaessaviees T. integrifolia
5a. Involucral bracts broadly ovate. Leaf base rounded, oblique, truncate. Perianth marcescent early in fruit development. Plants of sandstone ........ eee Re tec tcc td Aaee ee cha Tool ouAvns bch teticSebbuedactosteesnendoey T. borneensis
5b. Involucral bracts linear-triangular to narrowly ovate-triangular. Leaf base acute, decurrent. Perianth persistent until fruit maturity. Plants of MOAT OMS ce ee eats eee aot eR ee us Fea es eaghemosdbawbisiudereduusdacvvosncbseccese T. reducta
Acknowledgements
We wish to express our thanks for the collaboration and support of the Sarawak Forestry Department, notably Datu Cheong Ek Choon, the Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee, and the Forest Research Centre (Kuching), especially L.C.J. Julaihi Abdullah. The first author wishes to thank Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme Brown and Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their considerable support and funding of fieldwork in Sarawak.
References
Ding, Z., Ting, C.T. & Larsen, K. 2000. Taccaceae. Flora of China 24: 274- DAD:
Drenth, E. 1972. A Revision of the family Taccaceae. Blumea 20: 367-406. Drenth, E. 1976. Taccaceae. Flora Malesiana 7: 806-819.
Ling, P-P. 1985. Taccaceae. Flora Reipublicae Popularis Sinicae 16(1): 42- 54.
hie sata AE as aad Ma at : nc ARE. oath etalon baa. iat tolte o , TAOLe wit e an ¥ TIT MI : : ‘ ee a ase ene ae | ia oe | oo ire of arti te 1 he wy Yate Ns hime on + tp i rt rt iy? Ieee ris an At a pe ees
s~paULIRn Ge ' af
t ipa adsl lait Tue aS eS ae | I uted agi vee tolwaeic gel epee tort
ini sol su A)dsgliet rei Se aay 1 ponteen bream coo bet ut: itt enates cond) oo on “
; WGA Baise! Anand) a
‘cba Jaehiwh Af thn Twos a
pa
, RS 9
ares on
(0c: sik; ARI ae ‘pry is baler iw piers ¢ (tee) bi a. ait a ‘ ) ne Tek pee hier epee nd: 3 Inhiean oO . | a Pin ~~ LH ry 4 nes
Svs Age iti a a
> op oe
r - \ f ‘sg } i Lif Va i) is bs : ‘i 1 3 . ae
Gardens’ Bulletin Singapore 58 (2006) 41—46 41
A New Species of Curcuma L. (Zingiberaceae) from Southeast Asia
CHARUN MAKNOI_ AND TAYA JENJITTIKUL
: Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112 Thailand
: Department of Biology, Faculty of Science, Mahidol University, Rama VI road, Rajdhevee, Bangkok 10400 Thailand
Abstract
Curcuma larsenii C. Maknoi & T. Jenjittikul, sp. nov. from Southeast Asia (Thailand, Laos and Vietnam) is described and illustrated.
Introduction
The genus Curcuma, established by Linnaeus’ in 1753, comprises approximately 100 species (Skorni¢kova et al., 2003a). In recent years, several new species have been described, for example by Sirirugsa and Newman (2000), Mood and Larsen (2002), Skorni¢kova et al. (2003a, 2003b and 2004), as a result of extensive exploration in South and Southeast Asia.
While revising this genus for Flora of Thailand between 2002 and 2003, the first author found some unidentified specimens that did not match any known species. One of them was collected in Ubon Ratchathani Province, vegetatively similar to C. gracillima. However, it was reported by the second author in 2001 that this species was being sold in Chatuchak Market in Bangkok which was collected from the same province. This species seems to be becoming known as ornamental plants in the market. Therefore, it is described here as Curcuma larsenti C.Maknoi & T.Jenittkul.
Curcuma larsenii C. Maknoi & T. Jenjittikul, sp. nov.
Curcuma larsenii Maknoi & Jenjittikul sp nov., C. gracillimae Gagnep. characteribus_ vegetativis et inflorescentiae simulans, sed characteribus sequentibus differt: foliis latioribus; bracteis majoribus, apice staminodii acuto, non truncato; lobis labelli rectangularibus, apicibus truncatis, irregulariter
42 Gard. Bull. Singapore 58 (2006)
erosis. — Typus: C. Maknoi 496, Ban Sanamchai, Piboon Mangsahan, Ubon Ratchathani, 26 Aug 2003 (holo, PSU; iso, AAU, BKF, OSBG). Figure 1 and Plate 1.
Perennial herb, 25-30 cm tall. Vertical underground structure ovoid, 2-3 cm in diameter, yellowish inside. Bladeless sheaths 2-3, 1-6.5 cm long, pale yellow, green or dull red with red veins, apex acute or with a minute beak. Leaves 2-4; sheath up to 4 cm long, glabrous; petiole furrowed, 5-7 cm long, glabrous; ligule c. 1 mm long, membranous, glabrous, truncate; blade lanceolate, 20-25 x 3-5 cm, green, sometimes with a purple midrib, glabrous on both surfaces, except for few short hairs at the tip, base attenuate, apex acute. Inflorescence terminal; peduncle green, 5—10 cm long, glabrous; spike elliptic to elliptic-oblong, 4-6 x 2.5-3 cm; fertile bracts 7-15, obovate, 15- 20 x 18-25 mm, subtending cincinnus of 5—7 flowers, glabrous, green with a white margin, apex truncate, recurved; coma bracts 3—5, smaller and narrower, white or green with white stripes at apex. Bracteole broadly ovate, concave, 4.5-6.5 x 3.5-5.0 mm, white, membranous, glabrous. Flowers 20-25 mm long, exerted from the bract. Calyx funnel-shaped, 4-6 mm long, white, glabrous, apex unequally 3-lobed. Corolla tube 10-14 mm long, white, glabrous; corolla lobes 4.5—5.2 x 2.5-3 mm; dorsal one concave, white or pale yellow, apex hooded, shortly cucullate; lateral ones shallowly concave, white or pale yellow, apex obtuse. Staminodes oblong, spreading, 4.5-6.5 x 1.8—2.3 mm, creamy white to dark orange with translucent veins, red streak along margin near labellum, sparsely hairy at base, apex acute, margin eroded. Labellum broadly obovate, 5.5-6.5 x 6.0-7.5 mm, deeply divided, 2.5-3.5 mm deep, broad sinus, creamy white to dark orange with translucent veins, longitudinal red-streaked on the lower half, raised-band on either sides of sinus creamy white or yellow; lobes c. 2 mm wide, oblong, apex truncate with irregularly eroded. Filament c. 2.5 x 2 mm, white, glabrous. Anther c. 2.0 x 1.5 mm, white, pilose near the base; spurs absent; crest 1 mm long, apex shallowly emarginate. Ovary tri-loculed, oblong, c. 3 x 2.5 mm, white, glabrous; epigynous glands absent; stigma appressed cup-shaped, 0.6 mm wide, mouth serrulate. Fruits subglobose, 1 cm diam., white; seeds obovate, 3.5 mm long, brown with white aril.
Distribution: Thailand — Ubon Ratchathani; Laos and Vietnam. Flowering period: May to October. Ecology: In open forest in moist sandy areas.
Other specimens examined: Eastern Thailand — Larsen 47387, plant from Ubon Ratchathani, cultivated in Aarhus, 8 Oct 2001 (AAU!); Ngamriabsakul 67, Suan Pa Buntharik, Ubon Ratchathani, 5 Aug 1999
Gard. Bull. Singapore 58 (2006) Curcuma from Southeast Asia
Plate 1. Curcuma larsenii. 1. flower — pale colour form, 2. flower — dark colour form, 3. habit; 4. habitat.
Curcuma from Southeast Asia 43
Figure 1. Curcuma larsenii. a) whole plant; b) flower with bracteole at base; c) bracteole; d) calyx and ovary; e) dorsal corolla lobe; f) lateral corolla lobes; g) staminodes and stamen; h) labellum (scale a = 5 cm; b — h = 5 mm). Drawings made by the first author from the type specimen.
(BKF! E!); Tiptabiankarn 6604, Ubon Ratchathani, 29 May 2001 (PBM!); Laos — Maxwell 98-896, Khong Island, E side, base to the summit of Khong Hill, Mae Khong river, Champasak, 11 Sep 1998 (CMU!); Vietnam — Evrard 2324, 3 Jul 1929 (P!); Madris, km 97 route to Saigon, 21 Oct 1920 (P!).
Notes: This species is similar to Curcuma gracillima Gagnep. in vegetative and inflorescence characteristics but differs by the following characters: leaves broader, bracts larger, staminode apex acute instead of truncate, labellum lobes rectangular, apex truncate with irregularly eroded, staminodes and labellum creamy white to orange with translucent veins, red streaks on the
44 Gard. Bull. Singapore 58 (2006)
lower half and creamy white or yellow raised mid-band (table 1).
Table 1. Comparison of key characters of C. larsenii and C. gracillima.
15x20 mm
Staminodes and lip Creamy white to orange with red | Violet color streaks and yellow mid-band
Labellum lobes shape | Oblong, apex truncate, margin Obliquely irregular eroded obovate, apex acute
This species is named in honor of Professor Kai Larsen who initiated the research of Zingiberaceae in Thailand over 40 years ago.
Acknowledgements
The authors wish to thank Professor Puangpen Sirirugsa and Mrs. Supee Saksuwan Larsen for their kind advices. We are also indebted to Benjamin @Mllgaard for latinizing the diagnosis. Thanks are also given to Anni Sloth for the photographs of plant cultivated in Aarhus University. This work was supported by the Royal Golden Jubilee Scholarship (4BPS45E1) and DANIDA (CBBP-QSBG).
Curcuma from Southeast Asia 45
References
Mood, J. and K. Larsen. 2002. New Curcumas from South-east Asia. The New Plantsman 8: 207-217.
Sirirugsa, P. and M. KF Newman. 2000. A new species of Curcuma L. (Zingiberaceae) from S.E. Asia. The New Plantsman 7: 196-198.
Skorni¢kova, J., M. Sabu and M. G. Prasanthkumar. 2003a. New species of Curcuma L. (Zingiberaceae) from Mizoram, India. Garden’s Bulletin Singapore 55: 89-95.
Skorni¢ékova, J.. M. Sabu and M. G. Prasanthkumar. 2003b. Curcuma codonantha (Zingiberaceae) — a new species from the Andaman Islands, India. Gardens’ Bulletin Singapore 55: 219-228.
Skorni¢kova, J.,M. Sabu and M..G. Prasanthkumar. 2004. Curcuma mutabilis (Zingiberaceae): a new species of from South India. Gardens’ Bulletin Singapore 56: 43—54.
ia
ts
7
a
ernie Y Yet aor ywer NOt ucmeamias 7 (wa mcparatg Ike sit = met as ri Yextia a asad 4 Ries “63 / ith idee ; Sha a
Y (ar evindigeald,. ar hy oe ze A zE ap ==
-
} : PES! tA a! V0 for
ry. Qe 4a Ay lawnctending Sis wn eh eee read nn Lg a f ‘ La
ay uc) O00 gam ite oath pnapanaid: Be Fee : veel prilpial ute. mM eit te Hazes eat “a
+e: | o “TV4are) "ia et pear age =
Gardens’ Bulletin Singapore 58 (2006) 47—54 47
A hitherto overlooked field identification character for Borneo Scaphochlamys Baker (Zingiberaceae: Zingibereae)
A. SHAFREENA
Faculty of Resource Science and Technology Universiti Malaysia Sarawak 94300 Kota Samarahan, Sarawak, Malaysia. E-mail: apashafreena@frst.unimas.my
Abstract
The consistent presence of a pulvinus above the insertion at the base of the petiole of Scaphochlamys species has been observed in Sarawak and is here proposed as a useful field character for recognizing and separating Scaphochlamys Baker from its nearest allied taxa Boesenbergia Kuntze and Haplochorema K.Schum.
Introduction The genus Scaphochlamys Baker was last revised for Borneo by Smith (1987) at which time taxonomically important inflorescence characters were discussed (see Table 1). Poulsen & Searle (2005) added another reproductive character, splitting of the bracteole, to those proposed by Smith (1987). In the key to genera of Hedychieae (at that time Scaphochlamys included in Hedychieae) (Smith, 1987), characters other then those pertaining to inflorescence or flowers received very little attention. Recent work (Kress et al., 2002) has shown that Hedychieae is embedded in a newly expanded Zingibereae. The only vegetative character mentioned is well-developed stem in Hedychium Konig as compared to stem-less and tufted or shoots single-leaved (Scaphochlamys Baker, Boesenbergia Kuntze, Haplochorema K.Schum. and Kaempferia L.) (Smith, 1987).
For the present study, numerous species from three closely related genera (Scaphochlamys, Boesenbergia and Haplochorema) have been observed in Sarawak. The presence of a pulvinus at the base of the petiole (Plate 1) has been shown to be generically diagnostic in all the Scaphochlamys so far studied, which includes the six species currently recognized for Borneo
48 Gard. Bull. Singapore 58 (2006)
and at least a further 25 unidentified and seemingly novel taxa revealed during the first author’s ongoing studies in Borneo. This is a very useful character especially if plants are not in flowering at the time of collection since assigning sterile individuals to one of the three above genera is often problematic.
Despite our confidence in the diagnostic pulvinate petiole, care still needs to be exercised when examining Boesenbergia and Haplochorema since the insertion of the leaves can appear to be pulvinate (Plates 2 & 3). Careful comparison with the true pulvinate petiole of Scaphochlamys reveals that in Scaphochlamys the pulvinus is situated above the insertion of the petiole (Plate 4).
Scaphochlamys and its allied genera look very similar in the vegetative state. Thus, the presence of pulvinus will make determining these closely related taxa easier at the genus level.
Plate 1. Scaphochlamys sp. (Boyce & Shafreena Z\-157): note the pulvinate petioles.
Scaphochochlamys in Borneo
Plate 2. Boesenbergia pulchella (Ridley) Merr. showing leaf insertion (Boyce & Shafreena Z1-860)
50 Gard. Bull. Singapore 58 (2006)
Plate 3. Haplochorema magnum R.M.Sm. showing leaf insertion (Boyce & Shafreena ZI- 1013)
Scaphochochlamys in Borneo
Plate 4. Scaphochlamys polyphylla (K. Schum.) B.L. Burtt & R.M. Sm.: note many leaves each with a pulvinate petiole (Boyce & Shafreena ZI-904)
52 Gard. Bull. Singapore 58 (2006)
A comparison of inflorescence characters among Scaphochlamys and allied genera (based on Smith, 1987; Poulsen & Searle, 2005) and additional characters from this study are set out in Table 1 below.
Table 1. Comparative table of inflorescence and vegetative characters of Scaphochlamys, Boesenbergia and Haplochorema.
Inflorescence bract | Spiral Distichous Distichous arrangement
Flowering mode Acropetalous Basipetalous Basipetalous Flower arrangement | Cincinni Solitary Solitary
First bracteole More or less keeled, | Boat-shaped, Boat-shaped or (shape and arising oppositebract | arising at right split, arising at
position) angle to bract right angle to bract
Bracteole splitting | Split to base Split to base Split to 7/3 Labellum Bilobed or entire, Usually saccate, | Bilobed or lobes overlapping rarely bilobed, emarginated, flat
never flat
Base of thecae With very short free | Spurless Spurless basal spurs
Pulvinus Present Absent Absent
Acknowledgements
We wish to express our thanks for the collaboration and support of the Sarawak Forestry Department, notably Datu Cheong Ek Choon, the Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee and the Forest Research Centre (Kuching), especially L.C.J. Julaihi Abdullah. The second author wishes to thank Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme Brown and Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their considerable support and funding of fieldwork in Sarawak. The first author wishes to thank Universiti Malaysia Sarawak for the grant awarded [01(102)/454/2004(191)] for funding this research and Jeland ak Kisai of Malesiana Tropicals for his invaluable assistance in the field.
35)
Scaphochochlamys in Borneo
References
Kress, W.J., Prince, L.M. & Williams, K.J. (2002). The phylogeny and a new classification of the gingers (Zingiberaceae): Evidence from molecular data. American Journal of Botany 89: 1682-1696.
Poulsen, A. D. & Searle, R.J.2005. Scaphochlamys calcicola (Zingiberaceae): a new and unusual species from Borneo. Gardens’ Bulletin Singapore 57:
29-35.
Smith, R.M. (1987). A review of Bornean Zingiberaceae: 3 (Hedychieae). Notes Royal Botanic Garden Edinburgh 44: 203-232.
ies serie my Syit] Srl ae Hi he
7799 2 a? its _ $6 ale ’ 16 sg | a ~ _ w - = ~~ { ie ‘ as \ rit 7 : he ie, iG rg ~ r hid tke TOR Pe ; i 6 —wig’ Piss >, & wijpey eer }
DJ
grees: ro aa ora lini) e wining
aie, » Mee —_ , Ani Re ® Ale rahe
ise re, Gi eee aoa ph ea
46 TL Wie ial >? ty
other epee: acpportamety ret (il bape Sa be oe tded ee ‘ie 2A Add riot hl eae — ‘04 ee
Gardens’ Bulletin Singapore 58 (2006) 55—80 3 )8)
Phylogeny of the Genus Pomatocalpa Breda (Orchidaceae)
SANTI WATTHANA . TOPIK HIDAYAT , MOTOMI ITO” AND TOMOHISA YUKAWA
‘Botanical Garden & Museum, Natural History Museum of Denmark, University of Copenhagen, Gothersgade 130, DK-1123 Copenhagen K, Denmark.
‘Department of Biological Science, Graduate School of Science, The Uni- versity of Tokyo, Hongo 7-3-1, Bunkyou-Ku, Tokyo, Japan.
“Tsukuba Botanical Garden, National Science Museum, 1-1, Amakubo 4, Tsukuba, Japan
Abstract
The phylogeny of the orchid genus Pomatocalpa has been analyzed using morphological and molecular (matK and ITS) data. First, 9 representative species of Pomatocalpa, viz. P. armigerum (King & Pantl.) T. Tang & F Wang, P. bambusarum (King & Pantl.) Garay, P. bhutanicum N.P.Balakr., P. bicolor (Lindl.) J.J.Sm., P. spicatum Breda, P. kunstleri (Hook.f.) J.J.Sm., P. diffusum Breda, P. macphersonii (F.Muell.) T.E. Hunt and P. tonkinense (Gagnep.) Seidenf. and 11 neighboring genera (according to morphological features), viz. Acampe, Cleisostoma, Micropera, Haraella, Pelathanteria, Robiquetia, Sarcoglyphis, Smitinandia, Staurochilus, Trichoglottis and Ventricularia, were included to test the monophyly and phylogenetic position of the genus Pomatocalpa. This analysis was based on morphological data only, and the genera Seidenfadenia, Vanda and Ascocentrum were chosen as outgroups. Second, the interspecific phylogenetic relationships within Pomatocalpa were analyzed on the basis of matK and ITS sequences and morphological characters. The molecular analyses included seven species of Pomatocalpa, viz. P. undulatum (Lindl.) J.J.Sm., P. bicolor, P. diffusum, P. kunstleri, P. macphersonii, P. maculosum (Lindl.) J.J.Sm. and P. spicatum, and used Acampe ochracea (Lindl.) Hochr., Ventricularia tenuicaulis (Hook.f.) Garay and Smitinandia micrantha (Lindl.) Holtt. as outgroups. The morphological analysis included all 13 accepted species of Pomatocalpa by us and used
56 Gard. Bull. Singapore 58 (2006)
Acampe, Ventricularia, and Smitinandia as outgroups. Based on the results of an unpublished precursory study, the monotypic genus Haraella was included in the ingroup in both the morphological and molecular analyses. In addition to the separate analyses, a total evidence analysis was made, combining information from matK, ITS, and morphological data. Pomatocalpa turned out to be monophyletic, but only after exclusion of P. armigerum (= Cleisostoma armigerum King & Pantl.), PR bambusarum (= Cleisostoma bambusarum (King & Pantl.) King & Pantl.) and P. bhutanica (= Cleisostoma bhutanicum (N.P.Balakr.) S. Watthana, comb. nov.). According to matK, morphology, and total evidence, Pomatocalpa is monophyletic without Haraella. According to ITS data, on the other hand, Haraella is nested in Pomatocalpa. The incongruence of matK and ITS sequences may be because Haraella retrocalla is of ancient hybrid origin. When delimited according to this paper, the genus Pomatocalpa is probably monophyletic and is characterized by a single unique synapomorphy, i.e., the presence of a narrow longitudinal groove at the upper end of the front wall of the spur. The interspecific relationships in Pomatocalpa were poorly resolved.
Introduction
The orchid genus Pomatocalpa Breda belongs to subtribe Aeridinae, subfam. Epidendroideae, tribe Vandeae (Dressler, 1993). It contains 13 species distributed from Sri Lanka to Fiji, south to Northern Australia and north to Hainan and Taiwan (Watthana, in prep.).
A comprehensive hypothesis about the phylogeny of the genus Pomatocalpa is not yet available. A recent global phylogenetic analysis of subtribe Aeridinae based on DNA (matK and ITS sequences) contributes some information (Topik et al., 2005). In that analysis, Pomatocalpa is represented by P. diffusum and P. kunstleri.On this limited basis, Pomatocalpa appears to be monophyletic with the monotypic genus Haraella as a sister-
group.
As a precursory work of the present study, the molecular phylogeny of the Aeridinae has been re-analyzed with additionally seven species of Pomatocalpa included, which were available for DNA extraction only, viz P. undulatum s.1. (including P acuminatum (Rolfe) Schltr.), P bicolor, P. diffusum, P. kunstleri, P. macphersonii, P. maculosum and P. spicatum (Topik and Yukawa, unpublished). However, they seem to represent almost all the variations observed in the genus. In this analysis, Pomatocalpa is still
Phylogeny of Pomatocalpa 57
monophyletic according to matK data. In the re-analysis based on ITS data, on the other hand, Haraella is nested in Pomatocalpa.
The objectives of the present study are: 1) to test the monophyly of Pomatocalpa more thoroughly and to identify its phylogenetic posi- tion among closely related genera according to morphological features; 2) to resolve the interspecific phylogenetic relationships in Pomato- calpa based on molecular and morphological data and on total evidence.
Materials & Methods
I. Delimitation and phylogenetic position of Pomatocalpa (morphological data)
A. Sampling
Acampe, Cleisostoma, Haraella, Micropera, Pelathanteria, Robiquetia, Sarcoglyphis, Smitinandia, Staurochilus, Trichoglottis and Ventricularia were identified as the genera being morphologically most similar to Pomatocalpa. Thus, all of them except Robiquetia are placed in the same main group in Seidenfaden’s (1988) comprehensive account of Thai monopodial orchid genera; additionally, certain species of Robiquetia and Cleisostoma have previously been referred to Pomatocalpa. Consequently, these genera were included in a phylogenetic analysis together with nine representative species of Pomatocalpa, viz. P. armigerum, P. bambusarum, P. bhutanicum, P. bicolor, P. spicatum, P. kunstleri, P. diffusum, P. maphersonii and P. tonkinense. Furthermore, 7richoglottis lasiocarpa was treated separately from the rest of Trichoglottis in the analysis, because this species was only recently transferred to the latter genus from Pomatocalpa (Ormerod, 1997). Unfortunately, no specimen of Robiquetia vaupelii (syn. Pomatocalpa vaupelii Ormerod & J. J. Wood) was available for this study. The genera Seidenfadenia, Vanda and Ascocentrum were chosen as outgroups. According to the molecular phylogenetic study of Topik et al. (2005), these three genera belong to the clade that is sister to the clade accommodating the genera of the ingroup of the present study.
The decision to include genera as terminal taxa in the ingroup was made because this practice was expected to increase the influence of ancestral character states by reducing the noise from later-evolved species- specific features (Wiens, 2000). Naturally, the best solution would have been to include all individual species of all the genera concerned. However, this
58 Gard. Bull. Singapore 58 (2006)
would have become a major project, far beyond the scope of the present study. The only realistic alternative would be to include one or few representative species from each genus. Since none of the genera concerned have ever been thoroughly analyzed phylogenetically, a more or less random selection of “representative” species might well influence the analysis through the differences in later-evolved species-specific features. Naturally, we realize that including the genera as terminal taxa in the ingroup probably reduces the final resolution. Still we consider this practice to be the better option.
B. Characters
Morphological characters are usually regarded as qualitative or quantitative. However, so-called qualitative characters often have a quantitatively phenomenological base (Stevens, 1991) and can be expressed quantitatively by the systematist coding them (Baum, 1988). According to Stevens (1991), character states used in phylogenetic analysis should be discrete and based on carefully analyzed discontinuities in the variation. However, it seems that neither characters with a continuous variation pattern, nor characters with overlapping attribute values of the taxa, should be excluded a priori from phylogenetic analyses. Thiele (1993) and Ryding (1998) found such polymorphic characters to track phylogeny almost as accurately as characters with a discontinuous variation pattern. The results of Wiens (1995) suggest that polymorphic characters may contribute significant phylogenetic information, but also that they are less reliable. On this background, it was decided to include polymorphic characters in the present study.
For all genera included in the analysis, the character states were scored from: (1) the keys and descriptions in Schuiteman & de Vogel (2000), Seidenfaden (1988) and Seidenfaden & Wood (1992); (2) herbarium specimens and spirit samples deposited at C; (3) live specimens cultivated in the Botanical Garden, University of Copenhagen.
Some of the measurements are defined in Figure. 1. For this part of the study, 18 characters were scored (Table 1) and the matrix is shown in Table 2. The inflorescence type was excluded due to occasional variation within individual specimens of certain species (e.g., Pomatocalpa bicolor, P. diffusum, P. kunstleri and P. spicatum). It should also be mentioned that several floral characters were excluded because of too much variations seen in many of the terminal taxa.
The interpretation of the variation encountered in some of the
Phylogeny of Pomatocalpa 59
accepted characters should be briefly explained.
Habit (character 1). Plants with moderately elongate stems carrying many condensed leaves (e.g., Vanda) were coded as “stem short”. Certain species of Pomatocalpa, e.g. P. diffusum, are polymorphic. However, P. kunstleri, P. macphersonii and P. spicatum consistently have a short stem, while P. bicolor consistently has a rambling habit with an elongate stem.
—
i. Figure 1. Definitions of selected characters. A, length of peduncle; B, length of terminal rachis shoot; C, hind edge of labellum side lobe; D, front edge of labellum side lobe.
Table 1. List of morphological characters and states used in the phylogenetic analysis of intergeneric relationships.
1. Stem elongate (0) or short (1).
2. Labellum subglabous to minutely papillose (0) or pubescent to
distinctly hairy (1).
Mid-lobe without (0) or with ornaments (1).
Mid-lobe simple (0) or lobed (1).
Spur absent (0) or present (1).
Spur without (0) or with (1) a median septum.
Front wall of spur with (0) or without (1) a narrow longitudinal groove
at its upper end.
8. Front wall of spur without (0) or with (1) a big fleshy callus at its upper end.
9. Back wall of spur without (0) or with (1) an ornament (callus or tongue).
me a
60
10. th. £2: iS. 14. i. 16. ay.
18.
Gard. Bull. Singapore 58 (2006)
Back wall ornament situated near the spur entrance (0) or down in the spur (1).
Back wall ornament a fleshy callus (0) or a membranous tongue (1). Back wall tongue glabrous (0) or hairy (1).
Stipe broadly rectangular to subspathulate (0), (linear-)oblong (1) or narrowly linear (2).
Viscidium large and broader than long (0); large and ovate to oblong (1) or small and subglobose (2).
Pollinia two (0) or four (1).
Rostellum shorter (0) or longer (1) than the diameter of column. Column not dorsally prolonged and incurved (0) or column dorsally prolonged and incurved (1).
Column without (0) or with (1) two horizontally protruding appendages.
Table 2. Character by taxon matrix of the analysis of intergeneric relationships. Characters as in Table 1. ?=unknown, -=inapplicable. Polymorphic taxa are coded with the following symbols; these are equate macros in PAUP: b = (0&1).
5 10 15 Seidenfadenia 00001 0000- --100 000 Vanda 00101 OObO- --120 000 Ascocentrum 00001 OObO0- --1b0 000 Acampe 11101 O000- --101 000 Smitinandia 10001 0010- --021 000 Venticularia 11101 0100- --001 000 Trichoglottis 11bb1 O001b 1b101 000 T: lasiocarpum Ol11 ~~ 00011 ~~ 11101 000 Staurochilus 1ibib 0001 — 11101" 060 Cleisostoma bbbb1 b001b = O0-bb1 b00 Pelathanteria lbbb1 1001b 0-021 O00 Micropera tbhibiy pn 1OLob) pybahiderisl00 Robiquetia bOb01 O00bb b-1b0 bbb Sarcoglyphis 100b1 100bb~ b-111 #000 Haraella 01000 —----- --100 000 P. bhutanicum 12101 00010 #£O-101 O00 P. armigerum 11101 ‘00011 . U=10i-_* Coe P. bambusarum 00101 OO111 TOT * Ti P. bicolor 10001 01011 10101 000 P. spicatum 00001 01011 10101 000 P. kunstleri 00001 O1011 10101 000 P. diffusum bO001 01011 10101 010 P. maphersonii 00001 01011 10101. 010
P. tonkinense b0001 01011 10101 #010
Phylogeny of Pomatocalpa 61
Labellum surface (character 2). Due to the homogeneity of the epidermis (and due to intermediary states), the condition of a densely and minutely papillose surface (e.g. P. diffusum and some specimens of P. macphersonil) was considered conceptually similar to a subglabrous surface (e.g. P. spicatum and P. bicolor). Consequently, these conditions were treated as one character state and opposed to the condition of a properly hairy surface with heterogenous epidermis (e.g. in Acampe, Haraella, Staurochilus and Trichoglottis).
Ornaments on the labellum mid-lobe (character 3). In this context, the term “ornaments” covers a range of morphologically diverse structures, such as keels, warts, or markedly thickened margins. It is uncertain to which extent these structures are homologous.
Front wall groove (character 7). Several species and genera have two small calli at the boundary between the base of the labellum mid-lobe and the upper end of the front wall of the spur. However, only in some (but not all) species traditionally assigned to Pomatocalpa, these calli are adjoin- ing, elongate and extend down into the spur, thus producing a narrow longi- tudinal groove of varying length between them.
Front wall callus (character 8). The presence of two tiny calli at the boundary between the base of the labellum mid-lobe and the upper end of the front wall of the spur is common among the group of genera examined. However, this feature is markedly different from the presence of one big fleshy callus — a character state only observed in Pomatocalpa bambusarum, Micropera and Smitinandia. In the latter genus, it varies from globular (S. micrantha) to somewhat complanate (S. helferi (Hook.f.) Garay).
C. Phylogenetic analyses
The phylogenetic analysis based on maximum parsimony (MP) was performed using PAUP* version 4.0610 for Microsoft Windows (Swofford, 2001). All characters were equally weighted, unordered (Fitch, 1971). The data set was analysed by the heuristic search method with bisection- reconnection (TBR) branch swapping and the MULTREES option on, saving all most parsimonious trees (MPTs). Evaluation of internal support of clades was conducted by the bootstrap analysis (Felsenstein, 1985) with 10,000 replicates with faststep searching.
62 Gard. Bull. Singapore 58 (2006)
Interspecific Relationships in Pomatocalpa A. Molecular data A.1. Sampling
For this part of the study, matK and ITS sequences could be obtained from seven species of Pomatocalpa, viz. P undulatum, P. bicolor, P. diffusum, P. kunstleri, PR. macphersonii, P. maculosum (Lindl.) J.J.Sm. and P. spicatum. Due to the results of a preliminary analysis encompassing many more taxa of subtribe Aeridinae (cf. the Introduction), Haraella retrocalla (Hayata) Kudo (the only species of Haraella) was included in the ingroup, while Acampe ochracea, Smitinandia micrantha and Ventricularia tenuicaulis were used as outgroups.
Material of Pomatocalpa was partly collected in the field in Thailand by the first author and partly obtained from living collection in the Tsukuba Botanic Garden, Japan and from Queen Sirikit Botanic Garden, Chiang Mai, Thailand. A list of voucher specimens is given in Table 3.
Table 3. List of species analysed in the molecular analyses.
Species Voucher Source Pomatocalpa diffusum Breda Watthana 1767 (QBG) Thailand P. bicolor (Lindl.) J.J. Sm. Watthana 1821 (QBG) Malaysia P. maculosum (Lindl.) J.J.Sm. Watthana 1768 (QBG) Thailand P. spicatum Breda Watthana 1771 (QBG) Thailand P. kunstleri (Hook.f.) J.J.Sm. TBG 145833 (TBG) Borneo P. undulatum (Lindl.) J.J.Sm. Yukawa s.n. (TBG) Taiwan P. macphersonii (F.Muell.) Hunt TBG 130189 (TBG) Unknown Acampe ochracea (Lindl.) Hochr. TBG 130163 (TBG) unknown Haraella retrocalla (Hayata) Kudo TBG 133078 (TBG) Taiwan Smitinandia micrantha (Lind1.) Holttum TBG 118427 (TBG) Vietnam Ventricularia tenuicaulis (Hook.f.) Garay TBG 145846 (TBG) Thailand
A.2. Nucleotides preparation
The total DNA was extracted from fresh material or silica-gel dried plant tissues following the instruction of QIAGEN DNeasy Mini Plant Kit. For matk
Phylogeny of Pomatocalpa 63
sequences, the amplification was performed using a primer pair, OMATIF and trnK-2R (Topik et al. 2005). The 20-ul amplification reaction included 2 ul 10 x of Ex-Taq buffer (Takara Bio Inc.), 1.6 ul 2.5 mM of dNTPs mix, 0.5 ul each primer (10 pmol), 0.1 micro liter 5 units/ul of Ex-Taq DNA-polymerase (Takara Bio Inc.), 2 ul of template DNAs and 13.3 ul of MilliQ water. The polymerase chain reaction (PCR) profile consisted of an initial 5 min premelt at 94°C and 30 cycles of 30 s at 94°C, 30 s at 53°C, and 3 min at 72°C, followed by a final 7 min extension at 72°C.
Amplification of the ITS region was carried out using a primer pair, 17SE and 26SE (Sun et al., 1994). Total volume of PCR was 30 ul that included 15 ul GC buffer I (Takara Bio Inc.), 4.8 ul 2.5 mM of dNTPs mix, 0.5 ul of each primer (10 pmol), 0.21 ul 5 units/ul of LA Taq DNA-polymerase (Takara Bio Inc.), 2.4 ul of template DNAs and 6.59 ul of MilliQ water. The PCR profile consisted of an initial 2 min premelt at 94°C and 30 cycles of 50 s at 94°C, 1 min at 60°C, and 30 s at 72°C, followed by a final 7 min extension at 72°C. To confirm the number of amplified copies for ITS regions, we performed the single-strand conformation polymorphism (SSCP) analysis based on the method developed by Orita et al. (1989).
The PCR products were cleaned by using Montage PCR Centrifugal Filter Devices (Millipore Co.) and were used for auto-cycle sequencing reaction. The 10-ul auto- cycle sequencing reaction included 3 ul of Master Mix (Beckman Coulter), 1 pl primer (1.6 pmol), and 6 ul of PCR product. The reaction was incubated with 50 cycles of 20-s at 96°C, 20-s at 50°C and 4-min at 60°C.
Auto-cycle sequencing products were cleaned by adding STOP solution (2 ul 3M of NaOAc, 2 ul 100 mM of EDTA, and 1 ul 20 ng/ul of Glycogen) and 60 ul of 100 % ethanol; subsequently, they were centrifuged at 14000 rpm for 15- min at 4°C. The alcohol/salt mix was discarded, and the pellet was subjected to two washes with 200 ul 70% ethanol, each followed by centrifugation at 14000 rpm for 2-min at 4°C. Cleaned auto-cycle products were allowed to dry in the vacuum dry for 15-min. Both forward and reverse sequences were analyzed with CEQ8000 automated sequencer (Beckman Coulter), and electropherograms were edited and assembled with Genetyx-ATGC version 4.1 (Genetyx Corporation).
A.3. Phylogenetic analyses
DNA sequences obtained from matK and ITS were aligned with ClustalX and were then adjusted manually. Phylogenetic analysis and the evaluation of internal support of clades were performed by the same method as described for the morphological intergeneric analysis (see above).
64 Gard. Bull. Singapore 58 (2006)
B. Morphology B.1. Sampling
Nearly all recently accepted species of Pomatocalpa by us were included in the ingroup. However, P. armigerum, P. bambusarum and P. bhutanicum were excluded because of the results from the analysis of the systematic position and delimitation of Pomatocalpa (see below). Due to considerations discussed above, also Haraella retrocalla was included in the ingroup, while the genera Acampe, Smitinandia and Ventricularia were chosen as outgroups. We preferred to use entire genera as outgroups (instead of using representative species of the same genera). This practice was expected to increase the influence from ancestral character states in the outgroup by reducing the noise from late-evolved species-specific features (Wiens, 2000).
Most of the data were obtained by examination of spirit and herbarium material from AAU, AMES, BM, C, K, L, and P. Additionally, the following publications were consulted: Seidenfaden (1988, 1992), Comber (1990), Seidenfaden and Wood (1992).
B.2. Characters
The choice of characters reflects the same general considerations as discussed under “Delimitation and phylogenetic position of Pomatocalpa” above. For this part of the study, 32 characters were scored (Table 4) and the matrix is shown in Table 5. Some of the characters are defined in Figure 1. Several leaf characters (morphology of the sheath, dimensions and shape of the blade, etc.) as well as the outline of the back wall tongue of the spur were excluded, mainly due to high levels of intraspecific variation. The interpretation of the variation encountered in some of the characters should be briefly explained.
Stem length (character 1). Kerr (1985) made a distinction between Pomatocalpa species with a compact fan-shaped habit and those with an elongate rambling habit. However, intermediary states are found in P diffusum, P. fuscum (Lindl.) J.J.Sm. and P. marsupiale (Kraenzl.) J.J.Sm. Therefore, the length of stem was applied instead.
Length of peduncle in relation to rachis (character 3). This character is controversial in species with branched inflorescences. Furthermore, P. spicatum may produce both unbranched and branched inflorescences.
Phylogeny of Pomatocalpa 65
However, if the peduncle is defined to end at the point where the terminal rachis shoot starts, a ratio can be obtained that is comparable to the peduncle: rachis ratio in an unbranched inflorescence (Figure 1).
Hairiness of peduncle (character 4). According to Seidenfaden (1988), P.maculosum (subsyn. P. linearifolium Seidenf.) has a finely pubescent rachis. Judging from the first author’s observation, it varies from minutely papillose to sparsely and finely pubescent on the rachis, but it is not pubescent on the peduncle. Only P. kunstleri has a finely pubescent peduncle.
Table 4. List of morphological characters and states used in the phylogenetic analysis of interspecific relationships in Pomatocalpa.
1. Stem up to 30 cm long (0) or more than 30 cm long (1). 2. Inflorescence erect (0) or horizontal to pendent (1).
Peduncle shorter than or as long as the terminal rachis shoot (0) or peduncle much longer than the terminal rachis shoot (1) — cf. Figure 1.
4. Peduncle glabrous to minutely papillose (0) or distinctly pubescent (1).
Inflorescence with less than 5 flowers (0), 5-30 flowers (1) or more than 30 flowers (2).
Apex of floral bract obtuse to acute (0) or acuminate to caudate (1).
7. Pedicel less than 5.0 mm long (0) 5.0-10.0 mm long (1) or more than 10.0 mm long (2).
8. Dorsal sepal up to 6.0 mm long (0) or more than 6.0 mm long (1).
Dorsal sepal without markings (0), (sub)bordered (1), spotted or patched (2), with two longitudinal stripes (3), with a transverse band (4) or variegated (5).
10. Lateral sepals widely spreading (0) or strongly incurved (1).
il. Petals (oblong-) obovate (0) or linear (1).
12. Petal ground colour pinkish to whitish (0) or yellowish to brownish (1). 13. Labellum glabrous to minutely papillose (0) or distinctly hairy (1).
14. Front edge of each labellum side lobe subequal to the hind edge (0) or much shorter than the hind edge (1) — cf. Figure 1.
15. Side lobes of labellum rounded to obtuse (0) or (sub)acute (1). 16. Mid-lobe of labellum (ob)ovate (0) or oblong to rounded (1).
17. Mid-lobe of labellum not distinctly thickened at base (0) or distinctly thickened at base (i.e. triangular in longitudinal section) (1).
66
18. ry.
20.
ras
De.
23:
24. 2
20: Za
28.
22. 30. ie a2.
Gard. Bull. Singapore 58 (2006)
Mid-lobe of labellum straight (0) or strongly recurved (1).
Abaxial angle between labellum mid-lobe and spur more than 90° (0) or up to 90° (1).
Spur cylindric to conical (0), pyriform to globular (1), hook-shaped (2) or spur absent (3).
Front wall of spur without (0) or with (1) a narrow longitudinal groove at its upper end.
A median callus at the base of the labellum mid-lobe absent (0) or present (1).
Spur not strongly recurved when dry (0) or spur strongly recurved when dry (1).
Back wall tongue in the spur absent (0) or present (1).
Back wall tongue placed near the spur apex (0) or more than 1/3 from the spur apex (1).
Back wall tongue truncate to obscurely emarginate (0) or distinctly bifid (1).
Margins of the back wall tongue only attached to the spur wall at its very base, not forming a distinct pouch (0); or margins of the back wall tongue adnate to the spur wall for a long distance, forming a pouch (1).
Pollinia two, porate (0) or pollinia four, entire, joined in two globular pairs (1).
Stipe spathulate (0) or oblong to linear (1).
Capsule up to 3 cm long (0) or more than 3 cm long (1).
Capsule distinctly stalked (0) or (sub)sessile (1).
Capsule glabrous to minutely papillose (0) or distinctly hairy (1).
Table 5.Character by taxon matrix of the analysis of intrageneric relationships. Characters as in Table 1. ? = unknown, - = inapplicable. Polymorphic taxa are coded with the following symbols; these are equate macros in PAUP: b = (0&1) d=(1&2),e= (0&2), 1 = (a3).
5 10 15 20 25 30 Smitinandia Ob00d bO000 000-0 10001 OO00- --100 10 Acampe bbb0Od Ob150 0111- Obb00 OO00- --11lb 10 Ventricularia bb-00 10001 011-0 00002 QO00- --100 00 Haraella 01b00 01100 01170 100-3 OOO0- --01? # ?? P. kunstleri Ob112. 10bf0' 10001. .01b10» L004 10110: 14 P. spicatum 0100d 100f1 010b0 00111 1b011 Ob110 10
P. macphersonii 01001 00020 01011 01011 10011 01110 10
Phylogeny of Pomatocalpa 67
P. maculosum 1010d Ob020 01010 O0b101 1b011 O111b 10 P. undulatum 01001 60041 010b0 00111 bb011 10110 b0 P. bicolor 1010d 02120 01001 00111 11010 b1111 00 P. diffusum b010d Ob010 01011 00111 1b011 bbilb 10 P. simalurense 10102 01b20 01001 00111 10110 01111 »b0O P. fuscum b0102 01020 0101b 00001 11011 b1110 10 P. marsupiale b0102 OdbeO 01011 Oblib1 1b011 b1111 10 P. angustifolium 01001 00020 01011 00111 1b011 0b110 10 P. tokinense O1b0d 00130 10011 00110 11010 11110 10 P. floresanum 1010d 0b1?0 01001 00111 11010 bi1111 10
Length of dorsal sepal (character 8). To avoid noise from expected allometry, this was the only floral size character to be included. In other words, the length of the dorsal sepal can be seen as a general measure of flower size.
Index of labellum side lobes (character 14). Nearly all Pomatocalpa species have labellum side lobes in which the hind edge produces a right angle to the front edge. The ratio of the lengths of the hind and front edges is informative. However, this character was coded as inapplicable for P undulatum and P. spicatum, because the side lobes are broadly rounded in all specimens of the former and in some specimens of the latter.
Longitudinal section of labellum mid-lobe (character 17). This character was difficult to study from dried material. Therefore, it was only scored from fresh and spirit-preserved material.
A narrow longitudinal groove at upper front of the spur (character 21). In some specimens of P. undulatum the groove is distinct, while it is obscure in others. Consequently, the character was coded as polymorphic for this taxon.
B.3. Phylogenetic analysis
The phylogenetic analysis and the evaluation of internal support of clades were performed by the same methods as described for the intergeneric analysis based on morphological data (see above).
C. Total evidence
It was impossible to perform a congruency test on our data sets because of insufficient memory for simulation — probably due to the large proportion
68 Gard. Bull. Singapore 58 (2006)
of species for which DNA data were missing. However, since all the indi- vidual clades proved to have low bootstrap support, we found it justified to combine the data sets (in case of high bootstrap support for conflicting clades, data sets should not be combined; cf. de Queiroz 1993). Thus, a total evidence analysis of all data sets was performed to get the maximum infor- mation level (Kluge 1989). Acampe ochracea, Smitinandia micrantha and Ventricularia tenuicaulis were used as outgroups (since the DNA sequenc- es were derived from individual species, these species, rather than entire genera, had to be chosen). For those species of Pomatocalpa from which molecular data were not available, the molecular characters were coded as missing. Results
I. Delimitation and phylogenetic position of Pomatocalpa (morphological data)
Of the 18 characters scored, 14 were informative. The MP analysis yielded 3704 most parsimonious trees (length = 33; consistency index (CI) = 0.58; retention index (RI) = 0.73). The strict consensus tree is shown in Figure 2. The consensus tree suggests that Pomatocalpa,as traditionally circumscribed, is non-monophyletic. Most of the Pomatocalpa species, however, make up a monophyletic group. Bootstrap supports for branches in the strict consensus tree were less than 50% — except for the clade of Micropera and Pomatocalpa bambusarum (55%).
II. Interspecific relationships in Pomatocalpa A. Molecular data
The matK alignment had a total of 1835 sites, out of which 110 were variable and 28 were phylogenetically informative. The MP analysis yielded 9 most parsimonious trees (length = 167; CI = 0.89; RI = 0.65). The strict consensus tree and the corresponding branch supports are shown in Figure 3. The resolution of the matK tree is very low. However, the clade made up from all the Pomatocalpa species in the analysis has strongly bootstrap support (91%), while the clade of Haraella and Pomatocalpa is moderately supported (79%).
The ITS alignment had a total of 668 sites, out of which 69 were variable and 24 were phylogenetically informative. The MP analysis yielded 6 most parsimonious tree (length = 116; CI = 0.85; RI = 0.64). The strict consensus tree and the corresponding branch supports are shown in Figure
Phylogeny of Pomatocalpa 69
4.The ITS tree is better resolved than the matK tree, but all clades are weakly bootstrap supported, and Haraella retrocalla is nested in Pomatocalpa.
B. Morphology
Out of the 32 characters scored, 28 were informative. The MP analysis yielded 108 most parsimonious trees (length = 63; CI = 0.61; RI = 0.69). The strict consensus tree with corresponding branch supports is shown in Figure 5. The following relationship has moderate bootstrap support, viz. the monophyly of Pomatocalpa (62%), the clade consisting of P. simalurense, P. bicolor and P. floresanum (68% ), the sister group relationship of P. kunstleri and P. tonkinense (64%) and the sister group relationship of P. spicatum and P. undulatum (51%).
Pomatocalpa bicolor Pomatocalpa tonkinense Pomatocalpa macphersonii Pomatocalpa diffusum Pomatocalpa kunstleri Pomatocalpa spicatum Acampe Ventricularia
BS Micropera Pomatocalpa bambusarum Trichoglottis Trichoglottis lasiocarpa Pomatocalpa armigerum Pomatocalpa bhutanicum Sarcoglyphis Pelathanteria Cleisostoma Staurochilus Smitinandia Vanda Seidenfadenia Haraella Robiquetia
Ascocentrum
Figure 2. Strict concensus of the 3704 most parsimonious trees from the analysis of the phylogenetic position of Pomatocalpa, based on morphological data. Bootstrap support (if more than 50%) is indicated in percent above each branch.
70 Gard. Bull. Singapore 58 (2006)
P. kunstleri P. diffusum P. bicolor 91 P. undulatum P. maculosum 79 P. macphersonii P. spicatum Haraella retrocalla 58 Acampe ochracea Ventricularia tenuicaulis Smitinandia micrantha Figure 3. Strict consensus of the 9 most parsimonious trees from the analysis of the interspecific
phylogenetic relationships in Pomatocalpa, based on matK data. Bootstrap support (if more than 50%) is indicated in percent above each branch.
P. maculosum 61
64 P. bicolor P. diffusum 68 P. macphersonii P. kunstleri 61 Haraella retrocalla P. undulatum P. spicatum Ventricularia tenuicalis Acampe ochracea Smitinandia micrantha Figure 4. Strict consensus of the 6 most parsimonious trees from the analysis of the interspecific
phylogenetic relationships in Pomatocalpa, based on ITS data. Bootstrap support (if more than 50%) is indicated in percent above each branch.
Phylogeny of Pomatocalpa 71
P. bicolor
68 P. floresanum P. simalurense
P. maculosum
P. fuscum
P. diffusum
P. marsupiale 64 P. kunstleri
P. tonkinense
P. macphersonii 62
P. angustifolium 51 P. spicatum
P. undulatum
Smitinandia
Haraella
Ventricularia
Acampe
Figure 5. Strict consensus of the 108 most parsimonious trees from the analysis of the interspecific phylogenetic relationships in Pomatocalpa, based on morphological data. Bootstrap support (if more than 50%) is indicated in percent above each branch.
C. Total evidence
The data matrix of the combined molecular and morphological data sets has 2534 sites, of which 187 were variable and 82 were phylogenetically inform- ative. The MP analysis yielded 45 most parsimonious trees (length = 365; CI = 0.81; RI = 0.60). The strict consensus tree with corresponding branch supports is shown in Figure 6. All analyzed Pomatocalpa species form a monophyletic group with 78% bootstrap support, as opposed to 62% in the morphological analysis. Also the clade consisting of this genus and its sister group, Haraella retrocalla, is fairly well supported (80%).
The interspecific relationships in Pomatocalpa are almost identical to the topology from the exclusively morphological analysis (except for the collapse of the clade containing P. spicatum and P. undulatum).The bootstrap support of the other clades within Pomatocalpa is slightly lower than in the morphological analysis.
a2 Gard. Bull. Singapore 58 (2006)
Discussion I. Phylogenetic position and achievement of monophyly of Pomatocalpa
The strict consensus tree from the phylogenetic analysis of representative Pomatocalpa species and a selection of closely related genera (Figure 2) indicates that Pomatocalpa, as traditionally circumscribed, is non- monophyletic. Thus, while most Pomatocalpa species make up amonophyletic group, three species are found outside this clade. Pomatocalpa bambusarum is sister to Micropera, while P. armigerum and P. bhutanicum form separate branches in the largely unresolved clade, which in addition to Cleisostoma, Pelatantheria, Sarcoglyphis, Staurochilus and Trichoglottis, also contains the monophyletic Pomatocalpa group and the Muicropera/Pomatocalpa bambusarum clade. Unfortunately, we have been unable to obtain DNA data from these “misplaced” Pomatocalpa species. However, we tend to believe in the morphologically based phylogeny and find that P. armigerum, P. bambusarum, and P. bhutanicum should be removed to other genera to
P. bicolor P. floresanum P. simalurense
a P. maculosum
80
2 19 29 2 a
es 0-1 0-1 0-1
57
8 9 11 12 20 26
9 10 15 eae no f
f fj owU i ie fd
Ub 0-1 2-30-11-0 1-0 0-1 0-2 1-0 0-1
78
5 18 2124 one
UU 0-10-10-10-1
P. fuscum
P. diffusum
P. marsupiale
P. kunstleri
P. tonkinense
P. macphersonii P. angustifolium P. spicatum
P. undulatum Haraella Acampe ochraea Smitinandia micrantha
Ventricularia tenuicaulis
Figure 6. Strict consensus of the 45 most parsimonious trees from the analysis of the interspecific phylogenetic relationships in Pomatocalpa, based on total evidence (matK, ITS, morphology). The square boxes indicate the character numbers and character changes according to Table 3 (black boxes: unique synapomorphies; white boxes: homoplacies). Bootstrap support (if more than 50%) is indicated in percent above each branch.
Phylogeny of Pomatocalpa Hs,
achieve monophyly of Pomatocalpa for the following reasons.
Pomatocalpa armigerum (King & Pantl.) T-Tang & FEWang is probably better treated under its basionym, Cleisostoma armigerum King & Pantl., although its place in Cleisostoma is not unequivocal from the phylogenetic analysis in the present study (Figure 2). Just as Pomatocalpa, the very diverse genus Cleisostoma probably needs to be re-circumscribed. The back wall callus in the labellum spur of C. armigerum corresponds well with those of several species traditionally referred to Cleisostoma, e.g. C. krabiense (Seidenf.) Garay.
According to Pearce & Cribb (2002), Ormerod thinks that Pomatocalpa bambusarum (King & Pantl.) Garay might be better treated as Cleisostoma bambusarum (King & Pantl.) King & Pantl. According to the phylogenetic analysis in the present study (Figure 2), this species should rather be referred to Micropera. This, however, would call for a new combination. Therefore, we support the idea of tentatively referring the species to Cleisostoma, awaiting re-circumscription of the latter genus.
According to Pearce & Cribb (2002), Ormerod thinks that Pomatocalpa bhutanicum N.P.Balakr. might be better placed in Cleisostoma. We fully agree, admitting that its systematic position is not unequivocal from the phylogenetic analysis in the present study (Figure 2). The labellum shape and back wall callus in P bhutanicum correspond well with those of several species traditionally referred to Cleisostoma. This calls for anew combination:
Cleisostoma bhutanicum (N.P.Balakr.) S.Watthana, comb. nov.—Pomatocalpa bhutanicum N.P. Balakr., J. Bombay Nat. Hist. Soc. 75: 162. 1978. — Type: Bhutan. Tashiyangtsi, 1870 m alt.; flowered in National Orchidarium, Shillong on 24 June 1965, Balakrishnan 41993 (holotype, CAL; isotype, ASSAM).
Likewise, judging from Figure 2, it seems appropriate that Ormerod (1997) transferred Pomatocalpa lasiocarpa to Trichoglottis — at least, this species clearly does not belong in Pomatocalpa.
When adjusted in this way, a slightly reduced, but monophyletic, Pomatocalpa can be readily recognized from a character state that provides a synapomorphy for the entire genus, i.e., the presence of a narrow longitudinal groove at the upper end of the front wall of the spur.
The phylogenetic relationships between Pomatocalpa and _ its neighbouring genera is very poorly resolved, and it is not unlikely that a
74 Gard. Bull. Singapore 58 (2006)
number of the genera used as Operational Taxonomic Unit (OTUs) in the present analysis would turn out to be polyphyletic themselves, if they were analyzed with species as OTUs. The very high number of equally most parsimonious trees is undoubtedly due to many cases of polymorphism as well as missing data (cf. Wilkinson, 1995). Phylogenetic analysis of the Aeridinae, based on matK and ITS sequences, are somewhat incongruent with macro-morphological data (Topik et al., 2005). This is hardly surprising, since the majority of currently recognized genera under the Aeridinae are mainly, or exclusively, defined by floral traits (e.g. Garay, 1972, 1974; Senghas, 1986-1990; Seidenfaden, 1988). Floral characters are frequently found to show considerably high levels of homoplasy in the Orchidaceae (e.g. Pridgeon, et al. 1997; Bateman et al. 1997, 2003). Micromorphological features, such as the surface structure of velamen, pollen and seed, as well as sequence data from other genomic regions, could be interesting supplements in future analyses.
One of the neighbouring genera is of particular interest, i.e. the monotypic Haraella. As mentioned above, Topik and Yukawa (unpublished) conducted a precursory study which incorporated a large number of genera of the Aeridinae. They found Pomatocalpa (represented by five species) to be monophyletic in a phylogenetic analysis based on matK sequences. However, in the analysis based on ITS sequences, Haraella was nested in Pomatocalpa, rendering the latter paraphyletic. For this reason, Haraella retrocalla was included in the ingroup in each analysis in the present study. While both analyses based on morphology indicated only a distant relationship between Pomatocalpa and Haraella (Figures 2,5), our analyses based on molecular data corroborated the preliminary findings (Figures 3, 4). According to our total evidence analysis, Haraella is not nested in Pomatocalpa (Figure 6).
We tend to assign most weight to the total evidence analysis, because combined data sets can increase the level of information (Kluge, 1989). Consequently, we accept that Pomatocalpa is monophyletic without Haraella. Acceptance on basis of total evidence seems to be reasonable due to relatively low bootstrap values in the separate analyses (cf. de Queiroz, 1993).
Indeed, only the consensus tree based on ITS sequence data suggests that Pomatocalpa and Haraella should be lumped to achieve monophyly of the former. The conflict between the strict consensus trees based on matK and ITS sequences, respectively, may be because Haraella retrocalla is of ancient hybrid origin. This hypothesis finds some support in a forthcoming
Phylogeny of Pomatocalpa aD
paper (Yukawa et al., in prep.) contributing further phylogenetic details on the Aeridinae. In that analysis, the Tatwanese endemic Haraella retrocalla is sister to a Gastrochilus species in the strict consensus tree based on matK data, while it is sister to the Taiwanese endemic Pomatocalpa acuminatum in the strict consensus tree based on ITS sequences. These different positions might well reflect ancient hybridization combined with maternal inheritance of cpDNA and biparental inheritance of ITS. However, the origin and phylogenetic affinities of Haraella retrocalla are in need of much closer scrutiny.
Even if someone finds the analysis based on ITS sequences more convincing than our other analyses, we think that transferring Haraella to Pomatocalpa would be controversial. Such an act would create a morphologically heterogenous genus that would not be recognizable as an entity outside the laboratory. Indeed, there are a number of good reasons to accept paraphyletic genera in such cases (e.g., Sosef, 1997; Brummitt, 2002, 2003; Grant, 2003). Having said this, we still think that there 1s fairly good support for considering Pomatocalpa a monophyletic genus.
II. Interspecific relationships in Pomatocalpa
While the interspecific relationships in Pomatocalpa are completely unresolved in our strict consensus tree based on matK sequences only (Figure 3), the consensus tree based exclusively on ITS data (Figure 4) suggests an infrageneric structure that is somewhat different from the one hypothesized by the consensus trees based on morphological data and total evidence, respectively. However, the clade containing P. bicolor, P. diffusum and P. maculosum is congruent with the morphological data and total evidence data. |
The topology presented by the strict consensus tree based on total evidence (Figure 6) is almost identical to the consensus tree based on morphological data only (Figure 5). The only difference is that the clade containing P. spicatum and P. undulatum in the morphological tree (Figure 5) is found to be collapsed in the total evidence tree. The clade consisting of P kunstleri and P. tonkinense has weak bootstrap support; “petals linear” being its only synapomorphy. The clade accommodating P. bicolor, P. diffusum, P. floresanum, P. maculosum, P. marsupiale and P. simalurense has bootstrap support less than 50%, but “stem more than 30 cm long” constitutes a synapomorphy (despite the fact that this character is variable in P. diffusum, P. fuscum and P. marsupiale). The clade consisting of P. bicolor, P. floresanum and P. simalurense also has weak bootstrap support,
76 Gard. Bull. Singapore 58 (2006)
with no unique synapomorphic character state. P. maculosum is sister to P. fuscum with no unique synapomorphy and the bootstrap support is more than 50%.
The strict consensus trees of the various analyses do not clearly reflect distribution patterns in Pomatocalpa. However, the clade consisting of P. bicolor, P. floresanum and P. simalurense seems confined to the Malesian region (occurring in Peninsular Malaysia, Indonesia and the Philippines) apart from an uncertain collection of P. bicolor from “Cochinchina”.
In conclusion, neither our analyses of morphological data, nor matK or ITS sequence provided detailed resolution of the interspecific relationships in Pomatocalpa (Figures 3, 4,5). Variation of matK and ITS at species level appeared to be very low, 6.0 % and 10.3 %, respectively. The lack of resolution is probably due to a high internal conflict among the sequences collected, as can be deduced from the relatively low RI values (0.65 and 0.64, respectively). To produce a more detailed phylogeny, it would be desirable to obtain more DNA data from additional species of Pomatocalpa as the high amount of missing molecular characters in the total evidence analyses is undoubtedly influencing the results of this study.
Acknowledgements
We are very grateful to Dr. Henrik Aerenlund Pedersen and Dr. Olof Ryding for advice and help and Tetsuya Yamada and Tomoko Fujimoto for technical assistance. Additionally, we wish to thank Dr. Finn N. Rasmussen for participating in discussions concerning the cladistic analyses. This study was financially supported by a grant from the DANIDA project: Capacity Building in the Field of Biodiversity, Queen Sirikit Botanic Garden and Grants-in-Aid to Scientific Research from the Japan Society for Promotion of Science. (no 1737033).
References
Bateman, R. M., Pridgeon, A. M. & Chase, M. W. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 2. Infrageneric relationships and reclassification to achieve monophyly of Orchis sensu stricto. Lindleyana 12: 113-141.
Phylogeny of Pomatocalpa v7
Bateman, R. M., Hollingsworth, P. M., Preston, J., Luo, Y.-B., Pridgeon, A. M. & Chase, M. W.2003. Molecular phylogenetics and evolution of Orchidinae and selected Habenartinae (Orchidaceae). Journal of the Linnean Society, Botany 142: 1-40.
Baum, B. R. 1988. A simple procedure for establishing discrete characters from measurement data, applicable to cladistics. Taxon 37: 63-70.
Brummitt, R. K. 2002. How to chop up a tree. Jaxon 51: 31-41.
Brummitt, R. K. 2003. Further dogged defense of paraphyletic taxa. Taxon 52: 803-804.
Comber, J. 1990. Orchids of Java. Royal Botanic Gardens, Kew.
Felsenstein, J. 1985. Confidence limit on phylogenies: an approach using the bootstrap. Evolution 39: 783-791.
Fitch, W. M. 1971. Toward defining the course of evolution: minimum change for a specific tree topology. Systematic Zoology 20: 406-416.
de Queiroz, A. 1993. For consensus (sometimes). Systematic Biology 42: 368-372.
Dressler, R. L. 1993. Phylogeny and Classification of the Orchid Family. University Press, Cambridge.
Garay, L.A. 1972.On the systematics of the monopodial orchids 1. Botanical Museum Leaflets of the Harvard University 23: 149-212.
Garay, L.A. 1974. On the systematics of the monopodial orchids 2. Botanical Museum Leaflets of the Harvard University 23: 369-375.
Grant, V. 2003. Incongruence between cladistic and taxonomic systems. American Journal of Botany 90: 1263-1270.
Kerr, A. D. 1985.The genus Pomatocalpa. American Orchid Society Bulletin 54: 312-313.
Kluge, A. G. 1989. A concern for evidence and a phylogenetic hypothesis of relationships among Epicrates (Boidae, Serpentes). Systematic Zoology 38: 7-25.
Orita, M., Suzuki, Y., Sekiya, T. and Hayashi, K. 1989. Rapid and sensitive detection of point mutation and DNA polymorphisms using the polymerase chain reaction. Genomics 5: 874-879.
78 Gard. Bull. Singapore 58 (2006)
Ormerod, P. 1997. Notes on the orchids of New Guinea and the Pacific. Part 4. Australian Orchid Review 62: 15.
Pearce, N. R. and Cribb, P. J. 2002. The Orchids of Bhutan. Royal Botanic Garden Edinburgh, Edinburgh.
Pridgeon, A. M., Bateman, R. M., Cox, A. V., Hapeman, J. R. & Chase, M. W. 1997. Phylogenetics of subtribe Orchidinae (Orchidoideae, Orchidaceae) based on nuclear ITS sequences. 1. Intergeneric relationships and polyphyly of Orchis sensu lato. Lindleyana 12: 89-109.
Ryding, O. 1998. Phylogeny of the Leucas Group (Lamiaceae). Systematic Botany 23: 235-247.
Schuiteman, A. and de Vogel, E. 2000. Orchid Genera of Thailand, Laos, Cambodia and Vietnam. Vietnamese-English ed. Nationaal Herbarium Nederland, Leiden.
Seidenfaden, G. 1988. Orchid Genera in Thailand XIV. Fifty-nine Vandoid Genera. Opera Botanica 95: 1-398.
Seidenfaden, G. 1992. The orchids of Indochina. Opera Botanica 114: 1- 502.
Seidenfaden, G. and Wood, J. J. 1992. The Orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Fredensborg.
Senghas, K. 1986-1990. Vandoideae. — In: Schlechter, R., Die Orchideen. Ihre Beschreibung, Kultur und Ziichtung, 3rd ed., pp.959-1422. I/B. E.G. Brieger, R. Maatsch & K. Senghas. Verlag Paul Parey, Berlin & Hamburg.
Sosef, M. S. M. 1997. Hierarchical models, reticulate evolution and the inevitability of paraphyletic supraspecific taxa. Taxon 46: 75-85.
Stevens, P. F 1991. Character states, morphological variation, and phylogenetic analysis: a review. Systematic Botany 16: 553-583.
Sun Y., Skinner D.Z., Liang G.H., Hulbert S.H. 1994. Phylogenetic analysis of Sorghum and related taxa using internal transcribed spacers of nuclear ribosomal DNA. Theory Apply Genetics 89: 26-32.
Swofford, D. L. 2001. PAUP*4.0b10. Phylogenetic analysis using parsimony (*and other methods). Version 4. Sinauer Associates, Sunderland, Massachussets, USA.
Phylogeny of Pomatocalpa 79
Thiele, K. 1993. The Holy Grail of the perfect character: the cladistic treatment of morphometric data. Cladistics 9: 275-304.
Topik, H., Yukawa, T. and Ito, M. 2005. Molecular phylogentics of subtribe Aeridinae (Orchidaceae): insights from matK and nuclear ribosomal ITS sequences. Journal Plant Research 118: 271-284.
Wiens, J. J. 1995. Polymorphic characters in phylogenetic systematics. Systematic Biology 44: 482-500.
Wiens, J. J. 2000. Coding morphological variation for phylogenetic analysis. In Wiens, J. J. (ed.). Phylogenetic Analysis of Morphological Data. Smithsonian Institution Press, Washington.
Wilkinson, M. 1995. Coping with abundant missing entries in phylogenetic interference using parsimony. Systematic Biology 44: 501-514.
Anni lam (em ‘tee Ps +f} aH
+ ey eid
. , : a Pri
tt i i el ao RG
Ane: RNR
5- - ;
an ititet abn. bowictras: a pone iti
speyy ¥ jel Le
q afi Tre. oo.
ities Cy Gai) RAE wig aah malian: 4 Al
| ss iis i : Thea) : Cipy ra os
LimAwnr f
use ‘4 er ze i PRIOd ' ante |
Gardens’ Bulletin Singapore 58 (2006) 81—132 81
Species Composition and Biogeography of Tropical Montane Rain Forest in Southern Yunnan of China
ZHU H.,WANG H., AND LI B.-G.
Xishuangbanna Tropical Botanical Garden, The Chinese Academy of Sciences, Kunming 650223, P. R. China (mail address: Xue Fu Road 88, Kunming 650223, Yunnan, P. R. China; e-mail: zhuh@xtbg.ac.cn)
Abstract
The species composition, physiognomy and biogeography of tropical montane rain forest in southern Yunnan, SW China, have been studied based on data from 10 sampling plots and a complete floristic inventory. Two separate communities are recognized: a Mastixia euaonymoides-Phoebe megacalyx forest and a Parakmeria yunnanensis-Gymnanthes remota forest based mainly on species composition and forest structures. The tropical montane rain forest is characterized by evergreen meso-phanerophytes and micro-phanerophytes with simple, leathery and entire mesophyllous leaves, more or less frequent woody lianas and epiphytes, abundant herbaceous phanerophytes. However, it has few buttresses or cauliflory in physiognomy. The montane rain forest has similar species diversity to the lowland seasonal rain forest in the region. This indicates that species richness is not necessarily reduced with increasing altitude. We suggest this rain forest is a type of lower montane rain forest based mainly on its physiognomy, structure and floristics, but one that occurs at a higher altitude than those in equatorial SE Asia. The montane rain forest is dominated, in terms of species richness, by Lauraceae, Euphorbiaceae, Fagaceae, Theaceae, Rubiaceae and Papilionaceae, but by Lauraceae, Magnoliaceae, Euphorbiaceae, Fagaceae, Mastixiaceae and Nyssaceae in terms of phytosociological importance. In floristic composition, a total of 623 native species in 327 genera and 115 families of seed plants were recorded from the montane rain forest, of which recognizably ‘tropical’ elements contributed about 78.9% at the generic level and more than 80% at the specific level. Plants of tropical Asian distribution contribute 63.7% of the total sum of species. We conclude that the montane rain forest has strong tropical Asian affinities floristically even though it occurs at the northern margin of mainland SE Asia and at a higher altitude.
82 Gard. Bull. Singapore 58 (2006)
Introduction
Southern Yunnan in southwestern China is exceptionally interesting to botanists because of its diversified biota and unique geological- biogeographical history. The region is the most species rich and has the largest tropical-subtropical forest cover in southern China. Geographically, the region is at a transitional zone between tropical Southeast Asia and subtropical East Asia, and is also to be at a conjunction area between the Shan-Tai fragment of Gondwanaland and the southeastern margin of the Asian continent, geologically (Fortey et al., 1998, Metcalfe, 1998). Accordingly, southern Yunnan is a Key area in biogeography as well as being a global ‘hot spot’ for biodiversity (Myers, 1998).
The vegetation of southern Yunnan was mentioned, albeit briefly, for the first time by C.W. Wang in 1939 (Wang, 1939), but little was known until late 1950s because of poor access to the area. Many studies on the tropical forests in southern Yunnan have been done in the past, although little has been published in English (Zhu, 1992, 1993; Wu, 1987; Jin, 1997; Cao, 1996; Cao & Zhang, 1997; Zhu, 1997; Zhu et al., 1998a, 1998b, 2003, 2004; Zhu & Roos, 2004). Previous work on the forest in southern Yunnan has been restricted, largely, to the tropical rain forests at lowland sites below 900 m in elevation. The tropical montane rain forests are still poorly known (but see the descriptive works of Wu, 1987; Wang et al., 2001; Zhu et al., 2004).
Pristine montane rain forests were discovered recently at Mengsongin southern Yunnan, in the border between Myanmar and Yunnan (Wang et al., 2001).The montane rain forests occur in valleys and on some mountain slopes between 1500-1800m andareatype of‘lower montanerain forest’ according to Ashton’s (2003) categorization ofaltitudinal forest zonation in Southeast Asia.
Montane forests and their altitudinal zonations in tropical southeastern Asia have been studied by a great many authors since Brown (1919). The more important of these include Steenis (1935, 1984), Whitmore & Burnham (1969), Whitmore (1984), Ohsawa et al. (1985), Ohsawa (1991, 1993,1995), Kitayama (1992), Nakashizuka (1992), Pendry & Proctor (1996), Aiba & Kitayama (1999), Buot & Okitsu (1999) and Ashton (2003). It has been accepted, commonly, that there is an ecotone between the tropical lowland forest and lower montane forest with changes in physiognomic, structural and floristic attributes, occurring usually between 900-1200 m altitude. The montane rain forest in southern Yunnan occurs at much higher altitudes on the northern margin of tropical Southeast Asia. Its physiognomy, floristics and biogeography are accordingly of special interest.
Tropical Montane Rain Forest in Southern Yunnan of China 83
Site Description
Southern Yunnan is located in the southernmost part of Mainland China (Figure 1). It borders Myanmar and Laos, and has a mountainous topography with the mountain ridges running in a north-south direction and becoming lower in elevation southward. Altitude ranges from 480 m at the bottom of the lowest valley in the south (Mekong River) to 2429.5 m at the top of the highest mountain in the north. The Mekong River traverses the region from northwest to southeast (Xu & Jiang, 1987).
Mengsong is an administrative district in the west of southern Yunnan occupied by Hani people, an indigenous ethnic group. It is located in the border between Myanmar and Yunnan. Topographically it is a high basin surrounded by mountains, and varies in altitude from 1557 m within the basin to 2100 m at the top of the surrounding mountains. The region has a monsoon climate. From the climatic observation at 1600 m elevation, the mean annual temperature is 16.7 °C; the extreme minimum air temperature is 1.7 °C, the maximum air temperature, 28.5 °C, and the annual temperature accumulation (the sum of daily temperature means of > 10°C), 6083 °C. The mean annual precipitation is between 1800 and 2379 mm. More than 80% of the precipitation falls during the rainy season between May and the end of October, and the annual mean relative humility is 83.4%.
China
Yunnan
Mengsong el ietnam
Myanmar (Burma) *
Thailand
Figure 1. Map showing the location of Mengsong region in Xishuangbanna, southern Yunnan.
84 Gard. Bull. Singapore 58 (2006)
Methodology
The study was conducted in two stages: First, there was a general, landscape- scale, floristic inventory of the tropical montane rain forest in Mengsong, in which all plant species in the forest were recorded and specimens collected whenever possible. When habitat-related floristic variation had been identified, a systematic plot-based study was carried out. Five sampling plots, each 25 <x 20 m in size, were established 1n each assemblage in order to characterize the floristic variation. All trees in these plots were identified and their dbh (minimum 5 cm), height, and crown cover were measured. In each plot, five 5x5 m sub-plots were established to facilitate floristic survey of the understorey. In these sub-plots, saplings, shrubs and herbaceous plants were counted. Lianas in these plots were identified and their abundance estimated. The Importance Value Index (IVI), suggested by Curtis and McIntosh (1951), was calculated. Physiognomy (life forms and leaf sizes) was analyzed using Raunkiaer’s criteria (1934) as revised by Mueller-Dombois and Ellenberg (1974). Webb (1959) split off the lower end of Raunkiaer’s big mesophyll class (2025 —18225 mm’) as notophylls (2025-4500 mm’), which is to be preferred for detailed categorization of leaf size spectrum. Nonetheless, Chinese botanists and their local audience are familiar with Raunkiaer’s big mesophyll class. Accordingly we retain the big mesophyll class of Raunkiaer in this analysis.
Based on intensive floristic inventory of the forest, a more or less complete species list has been compiled, from which the floristics and geographical elements have been analyzed. Physiognomic comparisons between the montane rain forest and lowland rain forests in southern Yunnan and the equatorial tropics, and other montane rain forests in Southeast Asia have been made to demonstrate further the characteristics of the Yunnan montane rain forests. Specimens were identified and voucher material is lodged in the herbarium of Xishuangbanna Tropical Botanical Garden (HITBC). Species authorities follow the recently published and still on- going project of “Flora of China”.
Results
The vegetation
Based mainly on their habitats, species composition and forest profiles, we have divided the vegetation of the montane rain forest into two distinct assemblages which we have named based on their dominant and subdominant species, viz.:
Tropical Montane Rain Forest in Southern Yunnan of China 85
1 Mastixia euonymoides- Phoebe megacalyx forest; 2 Parakmeria yunnanensis- Gymnanthes remota forest.
Mastixia euonymoides- Phoebe megacalyx forest ((ME-PM’)
The ME-PM forest occurs mainly in wetter montane valleys. The forest has usually two tree layers. The upper layer is up to 35 m high with a crown cover of 70-80%, and is dominated by M. euonymoidos, Manglietia hookeri, Michelia cavaleriei and Nyssa wenshanensis var. longipedunculata. In some sites M. euonymoides grew sufficiently tall as to be considered as emergents. The lower tree layer was further divided into two sub-layers in some sites. The upper sub-layer was 10-20 m high with a crown cover of 60- 70%, and was dominated by Phoebe megacalyx, Syzygium brachythyrsum and Dysoxylum binectariferum. The lower sub-layer is 5-10 m high with a cover of 40-50%. The most frequently encountered species are Ardisia thyrsiflora, Cylindrokelupha kerrii, Ostodes kuangii and Brassaiopsis lepidota (see Appended Table 1).
The shrub layer is up to 1-5 m high and is dominated by juvenile trees. The most frequently seen shrub species are Psychotria symplocifolia, Brassaiopsis fatsioides, Mycetia gracilis, Brachytome hirtellata var. glabrescens and Oxyspora vagans.
The herbaceous layer is well developed with a cover of 50-70%. Frequent species are Ophiorrhiziphyllom macrobotryum, Allantodia dilatata, Ctenitopsis sp., Microsorum dilatatum, Porandra scandens, Rhynchotechum obovatum and Strobilanthes sp.
There are a few lianas, but some big woody individuals belonging to species such as Epigynum auritum, Bousigonia angustifolia, Calamus nambariensis, and Gnetum montanum are present.
Epiphytes are abundant. They include Pothos chinensis, Neottopteris nidus, Rhaphidophora hongkongensis, Aeschynanthus bracteatus, Pholidota
imbricata and Asplenium normale.
Parakmeria yunnanensis- Gymnanthes remota forest (PY-GR)
The PY-GR forest occurs on shady slopes and the tops of hills. The forest is 25-30 m high with a very even canopy. It also has two tree layers. The upper layer with a crown cover of 80% is dominated by P. yunnanensis, Nyssa
86 Gard. Bull. Singapore 58 (2006)
wenshanensis, Cinnamomum javanicum and Calophyllum polyanthum. The lower layer is at 5-20 m with a cover of 70-80%, and is dominated by G. remota, Syzygium brachythyrsum, Xanthophyllum yunnanensis and Wendlandia pingpiensis (see Appended Table 2).
Appended Table 1. Importance Values Index (IVI) of tree species in Mastixia euonymoides- Phoebe megacalyx forest.
Altitude: 1650-1780 m Height of canopy: 35 (m) Plot number and size: 5 (25x20) = 2500 m Coverage: > 90% Slope degree: 10-35 No. of sp. (©5 cm dbh): 62
No. of stems: 263 Species name RA RF RD Fvit Mastixia euonymoides 0.76 1.64 23.46 25.86 Phoebe megacalyx 9.13 4.1 6.00 19,22 Syzygium brachythyrsum 951 4.1 3.01 16.62 Dysoxylum binectariferum Ooi 4.1 235 13.95 Manglietia hookeri 0.38 0.82 14.14 15.34 Michelia cavaleriei jhe) 246 © 8.73 13.09 Nyssa wenshanensis var. L352 2.46 7.12 11.10 longipedunculata Linociera insignis 4.94 226, 1500 9.88 Ardisia thyrsiflora 4.56 4.1 0.87 023 Cinnamomum javanicum 2.66 S282 9:15 Helicia pyrrhobotrya 4.18 528-1058 8.05 Calophyllum polyanthum 2.66 a6. bie 7.66 Ostodes kuangii 3.8 2.46 1.38 7.64 Xanthophyllum yunnanensis 3.42 3.28. ) 0.88 7.58 Brassaiopsis lepidota 2.28 26> AOU. 6.64 Cylindrokelupha kerrii 3.8 ZAG = O79 6.55 Cryptocarya rolletii 3.04 B20 9 OL 6.49 Alcimandra cathcartii 1:52 246 2.29 6.27 Litsea vang var. lobata 152 3:28 Oils 4.93 Litsea lancifolia var. pedicellata 2.26 ZAG | “O82 4.86
Randia sp. 2.66 1.64 0.52 4.82
Tropical Montane Rain Forest in Southern Yunnan of China
Michelia hedyosperma 1.14 h6FS "1293 4.71 Drypetes salicifolia 0.76 164 2.30 4.70 Hovenia acerba var. kiukiangensis 0.76 O32. 2:62 4.40 Lithocarpus hancei 0.76 032° ~ 2.56 4.14 Litsea verticillata 2 1.64 0.03 Zal9 Mastixia pentandra var.chinensis 1.14 1.64 ~,0:28 3.06 Reevesia thyrsoidea 1.14 164 0.26 3.04 Randia wallichii 1.14 1.64 0.25 3.03 Dimocarpus yunnanensis 0.76 164 0.45 2ES5 Macaranga henryi 52 O82 O17 Zool Machilus shweliensis 0.38 82.09 2 2.47 Alseodaphne andersonii 0.38 O82 O98 Ze Litsea lancifolia 0.76 O82 039 is) Walsura yunnanensis 0.38 O82. - G73 1.93 Cinnamomum tamala 0.76 OS2-— 027 S85 Elaeocarpus glabripetalus var. alata 0.38 0:32 061 1.81 Rhododendron moulmainensis 0.76 0.32 022 1.80 Alsophila costularis (H76 0.82 0.18 L.76 Beilschmiedia roxburghiana 0.38 0.82 0.45 1.65 Alphonsea tsangyuanensis 0.38 0.82 0.43 1.63 Cyclobalanopsis chrysocalyx 0.38 0.82 0.38 1.58 Meliosma simplicifolia 0.38 O82) 4.0: O33 LSS Tapiscia yunnanensis 0.38 0:32.90 028 1.48 Alseodaphne pectiolaris 0.38 OLS 25. OI2F 1.47 Eriobotrya bengalensis var. angustifolia 0.38 O82 O27 1.47 Gymnanthes remota 0.38 O:s2 "0.26 1.46 Michelia floribunda 0.38 (hS2°) O95 f39 Diospyros kaki var. sylvestris 0.38 Ors2=° O18 38 Laurocerasus jenkinsii 0.38 S27 “OstS | e's, Nyssa wenshanensis 0.38 0.82 0.14 1.34 Beilschmiedia linocieroidea 0.38 Ors2, O12 1.31 Ficus auriculata 0.38 0.82 0.09 1:29 Walsura robusta 0.38 0.82 0.08 1.28
Artocarpus nitidus 0.38 0.82 0.08 128
88 Gard. Bull. Singapore 58 (2006)
Lithocarpus pseudoreinwardtii 0.38 0.82 0.04 1.24 Lindera latifolia 0.38 0.82 0.03 1.23 Oxyspora vagans 0.38 082. , 1.03 123 Litsea garretii 0.38 0.82 0.02 22 Castanopsis argyrophylla 0.38 0.82 0.01 124 Microtropis tetragona 0.38 0:82 «0:00 1.20 Gymnosphaera gigantea 0.38 0.82 0.00 1.20 Total (62 species) 263 stems 100 100 100.00 300.00
* RA: Relative abundance; RD: Relative dominance; RF: Relative frequency: IVI: Importance value index (Curtis & McIntosh, 1951)
Appended Table 2. Importance values Index (IVI) of tree species in Parakmeria yunnanensis- Gymnanthes remota forest.
Altitude: 1650-1700 m Height of canopy: (m) Plot number and size: 5 (25x20) = 2500 m Coverage: > 90% Slope degree: 5-30 No. of sp. (>5 cm dbh): 70
No. of stems: 293 Species name RA RF RD IVI* Gymnanthes remota £5.36 4.20 3.97 23-03 Parakmeria yunnanensis 1,02 257 11.08 14.62 Xanthophyllum yunnanensis 7 AF 3.36 D222 1275 Syzygium brachythyrsum sag 3.36 1.65 12.18 Wendlandia pingpiensis 6.83 336 1.19 11.38 Nyssa wenshanensis 1.02 1.68 8.13 10.84 Cinnamomum javanicum 3.07 3.36 3.83 10.26 Calophyllum polyanthum 3.41 3.36 3.38 10315 Nyssa wenshanensis var. longipedunculata 2.05 1.68 5.93 9.66 Mastixia pentandra subsp. chinensis 4.10 1.68 3.7/2 9.49 Cyclobalanopsis chapensis 3.42 2.52 3.29 9.23 Manglietia insignis 0.68 0.84 6:17 7.70 Acer decandrum 2.05 3.30 2:26 767 Ostodes kuangii 4.44 0.84 0:53 5.80 Cyclobalanopsis chrysocalyx 0.34 0.84 3.97 5:16
Machilus shweliensis 37 Ome y) li 5.00
Tropical Montane Rain Forest in Southern Yunnan of China
Engelhardtia spicata Alcimandra cathcartii Michelia floribunda Podocarpus neriifolius Craibiodendron stellatum Lithocarpus gagnepainianus Dimocarpus yunnanensis Gomphanara tetrandra Cinnamomum bejolghota Lithocarpus pseudoreinwardtii Linociera ramiflora Castanopsis hystrix
Litsea lancifolia
Lindera metcalfiana var. dictyophylla Castanopsis argyrophylla Lithocarpus fohaiensis Reevesia thyrsoidea
Ardisia thyrsiflora
Randia griffithii
Schima wallichii
Symplocos wikstroemiifolia Dysoxylum binectariferum Pygeum henryi
Litsea euosma Cylindrokelupha kerrii Eurya aurea
Linociera insignis Eriobotrya obovata Rhododendron moulmainensis Acer huianum
Alangium chinensis Elaeocarpus howii Machilus rufipes
Ternstroemia gymnanthera
0.34 0.68 37 0.68 02 ia Zi05 ZAS 1.37 2.05 0.68 0.34 0.34 lesv 0.68 0.34 0.68 1.02 0.68 0.68 02 1.02 0.68 0.34 0.68 0.68 0.68 0.68 102 0.34 0.34 0.34 0.34 0.34
0.84 1.68 1.68 0.84 0.84 0.84 1.68 0.84 22 1.68 1.68 0.84 0.84 1.68 0.84 1.68 1.68 1.68 1.68 0.84 1.68 1.68 1.68 0.84 1.68 1.68 1.68 0.84 0.84 0.84 0.84 0.84 0.84 0.84
3.78 2.58 |e a2 L255 1.85 USD 0.69 0.28 0.24 1.16 2.24 D394 0.23 1.47 0.90 0.52 0.15 0.47 eo? 0.07 0.06 0.33 1.26 0.06 0.06 0.04 0.84 0.44 1.03 0.92 O77 0.61 0.54
4.96 4.94 4.83 4.75 4.42 4.40 4.28 4.26 4.17 70 B52 3.43 339 o:27 2099 Pisa iy 2.88 2.86 2.84 2.80 DEG 2516 209 2.44 2.43 2.42 2.40 233] 2.30 Dok 210 1.95 LJ9 ig
90 Gard. Bull. Singapore 58 (2006)
Itea macrophylla 0.68 0.84 0.16 1.68 Beilschmiedia robusta 0.34 0.84 0.39 Sy Lithocarpus truncatus 0.34 0.84 0.38 1.56 Pittosporum kerrii 0.34 0.84 0.26 1.45 Laurocerasus jenkinsii 0.34 0.84 0.23 1.41 Helicia tsati 0.34 0.84 Oh 1.35 Tricalysia fruticosa 0.34 0.84 0.17 1.35 Styrax grandiflora 0.34 0.84 0.16 1.34 Bruinsmia polysperma 0.34 0.84 0.15 1.33 Garcinia cowa 0.34 0.84 0.08 12g Eurya prunifolia 0.34 0.84 0.06 1.24 Casearia velutina 0.34 0.84 0.04 1.22 Carallia lanceaefolia 0.34 0.84 0.03 174 Sarcosperma griffithii 0.34 0.84 0.02 1.20 Oxyspora vagans 0.34 0.84 0.02 1.20 Platea latifolia 0.34 0.84 0.02 120 Cyclobalanopsis myrsinaefolia 0.34 0.84 0.01 1.20 Amoora yunnanensis 0.34 0.84 0.01 [19 Paramichelia baillonii 0.34 0.84 0.01 1.19 Anneslea fragrans 0.34 0.84 0.01 1.19 Total (70 species) 293 stems 100.00 100.00 100.00 300.00
* See Appended Table 1.
The shrub layer is 1-5 m high with a cover of 30%-40%, and is dominated by juvenile trees. Frequent shrub species are Euodia lepta, Fargesia plurisetosa, Lasianthus lucidus, Psychotria symplocifolia, Oxyspo- ra vagans and Lasianthus inodorus.
The herbaceous layer is usually less developed than in the preceding forest type. Frequent species are Davallia mairesii, Pteris insignis, Ophiopogon graminifolia, Colysis pothifolia and Strobilanthes sp.
Lianas are fewer but there are some big woody lianas such as Connarus paniculatus, Celastrus monospermum, Epigeum _ auritum, Bousigonia angustifolia, Gnetum montanum and Alyxia balansae.
Tropical Montane Rain Forest in Southern Yunnan of China 91
Epiphytes are fewer than in the ME-PM forests.
We have analyzed the forest physiognomy based on 261 vascular species from the 10 plots of these two montane rain forest types. Both forests are dominated by phanerophytes, which make up 79.3% of all species (Table 1). In terms of the spectrum of leaf sizes, the plants with mesophyllous leaves contribute up to 68.2% of the total species, and 76.4% of tree species (Table 2). Woody plants with simple leaves contribute up to 90.6% and those with entire leaf margins, up to 76.5% (Table 3).
The flora
623 native seed plant species (including varieties) in 327 genera and 115 families of seed plants were recorded from the montane rain forest (see Appendix 3). The families with highest species richness included Lauraceae (51 species), Euphorbiaceae (36), Rubiaceae (23), Fagaceae (20), Liliaceae (20), Rosaceae (19), Araceae (18), Theaceae (17) and
Table 1. Life form spectrum of the tropical montane rain forest in southern
Yunnan. Lite form” Number of species % Megaphanerophyte 2 4.6 Trees Mesophanerophyte 61 23.4 Microphanerophyte 54 20.7 (All trees) (127) (48.7) Shrubs Nanophanerophyte LE 8.4 Herbaceous phanerophyte 24 2 Herbaceous Geophyte 5 1g plants Chamaephyte is 9.6 (All herbs) (54) (20:7) Liana Liana phanerophyte 34 15:0 Epiphyte Epiphyte 24 oe Total species 261 100
* The Raunkiaer’s criteria (1934) as revised by Mueller-Dombois and Ellenberg (1974): Megaphanerophyte (perennials, over 30 m_ high); Mesophanerophyte (perennials, 8 to 30 m high); Microphanerophyte (perennials, 2 to 8 m high); Nanophanerophyte (perennials, 0.25 to 2 m
92 Gard. Bull. Singapore 58 (2006)
high); Herbaceous phanerophyte (herbaceous perennials, over 0.25 m high); Chamaephytes (perennials, less than 0.25 m high above ground); Geophyte (perennials, dying back above ground).
Table 2. Leaf sizes of the tropical montane rain forest in southern Yunnan.
Macrophyll Mesophyll Microphyll | Total oe 164025 ea 18225 226-2025 mm’
(Number of species [1 [97 [9 MY nS [Number ofspecies |= fois 22 2.0) as faite sdl a reihvalbaed AO Bees le aie al Nunkerafgecis [ax fas 54 [Mosiscse ainsaab a doxif LOT ya tint [SO staal OAS a
Number of species | 1 EAR ERT eee Number of species 2 ee Cn Neca al pane 1 198 Cams ij Eb eee
Table 3. Leaf types, leaf textures and leaf margins of the tropical montane rain forest in southern Yunnan.
127
Shrubs
Herbs
Lianas 34
Epiphytes 74
Total species 261
Leaf type Leaf texture Leaf margin S C P L B N
Trees Number of iB 14 a1 76 i 30 127
species
%o 89.0 110 .402 598. 64. | 23% Shrubs Number of ik iL iN 5 iy 5 pb,
species
% 95.4 4.6 TIS) ay Tat ULE
All woody Number of 134 Ls 68 81 114 35 149 plants species species
% 90.6 9.4 45:6). Sa4ia T6Seues
Tropical Montane Rain Forest in Southern Yunnan of China 93
S: Simple; C: Compound; P: Papery; L: Leathery; E: Entire; N: non-entire. Papilionaceae (16).
The various types of geographic distributions of seed plants from China at the generic level have been documented by Z.Y. Wu (1991). Using Wu’s documentation, we have quantified the distribution types of the flora of montane forest at the generic level and these are summarized in Table 4.
Distribution described as ‘tropical Asian’, such as Mastixia, Pterospermum and Knema, represent up to 27.5% of total genera of the flora. ‘Pantropic’ distribution, such as those of Gnetum, Piper, Lasianthus and Bauhinia, contribute up to 26 %.*Old World Tropical’ distribution, such as those of Thunbergia, Pandanus and Carallia are the next most abundant. These tropical distributions (Types 2-7) compose 78.9% of the total genera. This indicates that the flora of the montane rain forest in southern Yunnan is of tropical nature and has strong tropical Asiatic affinity.
At the specific level, nine geographical elements (distribution types) were recognized from 623 seed plant species of the montane forest (see Table 4). “Tropical Asian’ elements and their subtypes contribute up to 63.7% of the total sum of species, including those of ‘Indo-Malesian’ distribution, such as Garcinia cowa, Knema furfuracea and Gironniera subaequalis. Others belong to ‘Southern Asian’ to ‘Mainland Southeast Asian’ distributions, such as Alcimandra cathcartii and Silvianthus bracteatus; and those of ‘Mainland SE Asia’ to ‘SW and SE China’ distributions, such as Vaccinium exaristatum, Metadina trichotoma and Semecarpus reticulata. The elements of ‘Chinese Endemics’ and subtypes, which were defined on available references, contribute up to 26%, including those of ‘SW to SE China’ distribution, such as Lithocarpus fordianus and Craspedolobium schochii; and the ‘Yunnan Endemics’, such as Lithocarpus fohaiensis and Cryptocarya rolletii.
Comparison with the lowland rain forest in southern Yunnan and _ the equatorial tropical and montane rain forests in SE Asia
Compared with the tropical montane rain forest in Java at similar altitude (Meijer, 1959), the montane rain forest in southern Yunnan has fewer epiphytes (Figure 2), but a higher proportion of woody phanerophytes.
Compared with the tropical seasonal rain forests at lower altitude in southern Yunnan (Zhu et al., 1998a) and equatorial lowland rain forests (Beard, 1946; Paijmans, 1970; Givnish, 1978; Proctor et al., 1998), the
94 Gard. Bull. Singapore 58 (2006)
montane rain forest has fewer mega-and meso-phanerophytes and lianas, fewer plants with compound leaves, fewer plants with macrophyllous leaves, but more abundant herbaceous plants and more plants with non-entire leaf margins (Figures 3 and 4).
The families with highest species richness in the montane rain forest are, to some extent, similar to those in the seasonal rain forests at lower altitudes in the region, but there is greater species richness in Fagaceae, Theaceae, Liliaceae, Rosaceae and Magnoliaceae (Figure 5). In terms of phytosociological importance, most of the dominant families in the montane forest are also dominant families in the lowland seasonal rain forests, but Magnoliaceae, Fagaceae, Mastixiaceae, Nyssaceae and Polygalaceae are of greater importance (Figure 6).
50 i TMRF Yunnan LJ TMRF Java
Tree Shrub Herb Liana Epiphyte Life form
Figure 2. Comparison of life form spectra between the tropical montane rain forest of Mengsong in southern Yunnan and the tropical montane rain forest in Java, Indonesia. TMREF Java: montane rain forest at altitudes 1450-1500 m in Java (Meijer, 1959); TMRF Yunnan: tropical montane rain forest at altitudes 1500-1800 m in Mengsong, southern Yunnan.
95
Tropical Montane Rain Forest in Southern Yunnan of China
a ne eee 0001 co so1seds [e107
aaa [ye ueUUTLA O} SIWOpUy *7-6 O'O0T ECE BIOUIS [IO],
91 16 POO aS CV ANS 16] 8c Vv BUI) O} SIUTOpuy “CT (97) (z9T) sodAjqns sji pue eulyD Oo} ormepuq’6 = ¢"p val BISWA PT 6C ST PISV UIoIsey °8 9°0 C BISV D 0} BISV ‘AA pure UBOUBLIOIIPO] “TT
2 eC Lvl BUY) AS Puke MS O} BISW AS pueule~‘e-, 90 % oye1od uray, PION PIO “OT 6'0¢ O¢l BISV HS puelurey] OV PISV S°7-L LE @| poyounltsip eolouly ‘N pur eIsy “Yq °6 OL OCI BIso[eIN-Opul TL OL SC oye 1odulay, YON *g (L'€9) (L6¢€) sodAjqns sji pue eisy [eoidoly "2 $1Z 06 ( eIskeyeJ-opuy) eisy [eoidody “7 ST II POLITY [eoidory 0} eisy [esidoip9 —¢'L VC POLY [eoidory, 0} eisy eoidory, “9 9'T OL Blyersny [eoidoly 0} eisy yeoidolp,¢ = 6p On elyensny [eoidoly 0} eisy eotdory,*¢ 9°0 Vv SoIdolL PHOM PIO 'h = 76 O€ SoIdOd], PHIOAA PIO ‘h C4) @ yountstp votoury yeoidoy 2 eisy yeotdoiy,¢ =p €[T younlsip eoroury [eoidoiy, 2 visy [eotdory,¢ Lt L soidonued ‘Z (97 $8 soidonued ‘7 6T cl ueyyodousoy *~ 1'¢ OT ueyyodourso_ "| eed, ae a eee RU % jJO'ON Jada] oytoods ye odd} UOTNGINSIG % JO'ON [PA] D19UNS ye sodA} UOTNNISIG
‘ueuUuN,X UlOYyINOS ‘SUOSSUAJ] Ul JSOIOJ UTR QUR}UOU dy} JO LAO dy} JO s[aAcy] oyoods pure ILI9USS Je SJUOUTITS [eoTydeIsOIH “p oqey,
96 Gard. Bull. Singapore 58 (2006)
Species %
Ep Lph Mega- Micro- Hph+Ch G Mesoph Nanoph
Life form
Figure 3. Comparison of life form spectra between the tropical montane rain forest in Mengsong and seasonal rain forests in southern Yunnan.
LHSR: lower hill seasonal rain forest;
RSR: ravine seasonal rain forest;
TMRF: tropical montane rain forest in Mengsong.
Ep=Epiphyte; Ch=Chamaephyte; G=Geophyte; Lph=Liana-phanerophyte Hph=Herbaceous phanerophyte; Mega-Mesoph=Megaphanerophyte + Mesophanerophyte; Micro-Nanoph=Microphanerophyte + Nanophanerophyte
Tropical Montane Rain Forest in Southern Yunnan of China 97
100
LJ LHSR RSR TMRF LRI
NI SFT
Y
Species %
GY Y GQ TZ GY WZ WZ 7
N \ \ \ \ \ \ \ \ \ SN
NY iG
al Nano-Micro. Meso. Macro. Gigan.
Leaf size
Figure 4. Comparison of leaf size spectra between the tropical montane rain forest in Mengsong and the seasonal rain forests in southern Yunnan, as well as the ones from the equatorial lowland. LHSR: lower hill seasonal rain forest in southern Yunnan; RSR: ravine seasonal rain forest in southern Yunnan; TMRF: tropical montane rain forest in southern Yunnan; LRI: lowland tropical evergreen rain forest in India SFT: evergreen tropical seasonal forest in Trinidad Nano-Micro.: Nanophyll + Microphyll; Meso.: Mesophyll; Macro.: Macrophyll; Gigan.: Gigantophyll : from Beard (1946); from Proctor et al. (1998)
98 Gard. Bull. Singapore 58 (2006)
TMRF Species %
0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0
Lauraceae ee ee ee Euphorbiaceae Rubiaceae Fagaceae Liliaceae Rosaceae Araceae Theaceae Papilionaceae Verbenaceae Moraceae Myrsinaceae Piperaceae
Meliaceae
Commelinaceae
LHSRF Species % 0.0 2.0 4.0 6.0 8.0 10.0 12.0
Rubiaceae ee Euphorbiaceae Moraceae Papilionaceae Lauraceae Rutaceae Apocynaceae Meliaceae Acanthaceae Annonaceae Myrsinaceae Asclep iadaceae Vitaceae
Rhamnaceae
Sap indaceae
Tropical Montane Rain Forest in Southern Yunnan of China
RSRF Species %
3.0 4.0 5.0 6.0
=) o i ) NN o = oo) ioe) ro)
Rubiaceae Lauraceae Euphorbiaceae Annonaceae Moraceae Orchidaceae Meliaceae Vitaceae Apocynaceae Urticaceae Rutaceae Fagaceae Acanthaceae Papilionaceae Piperaceae
99
9.0
Figure 5. Comparison of abundant families with most species richness between the montane
rain forest and seasonal rain forests at lower altitudes in the region. TMREF: tropical montane rain forest in southern Yunnan;
LHSR: lower hill seasonal rain forest in southern Yunnan;
RSR: ravine seasonal rain forest in southern Yunnan.
100
Lauraceae Magnoliaceae Euphorbiaceae Fagaceae Mastixiaceae Nyssaceae Myrtaceae Rubiaceae Meliaceae Polygalaceae Guttifferae Oleaceae Myrsinaceae Aceraceae Proteaceae
Euphorbiaceae Sapindaceae Combretaceae Moraceae Anacardiaceae Annonaceae Lauraceae Myristicaceae Ebenaceae Ulmaceae
Burs eraceae Sterculiaceae Meliaceae Bignoniaceae Verbenaceae
Gard. Bull. Singapore 58 (2006)
TMRF IVI 0 5 10 | 20 D 30 35 40 45 50 a MM ee SSS ee SS a ee a a ——— aS a — ee <r a a) Sa eal RSRF IVI (0) 5 10 15 20 25 30 i 40
Tropical Montane Rain Forest in Southern Yunnan of China 101
LHSRF
0 So eV 20 25° 30° 35 AS _ ME = =
Whites: ee eS Whitt . = eS a ee | A GMaAcCeAc \ Lae ee ee Euphorbiaceae Sa ee eee ee ee eee LW EGS? sae eet a a i papmdaceae (Ge ee Bartingtoniace [= =2> Rubiaceae (=== Guttiiemer = 2 ee] io isc¢1ceie ee hetraineleaced ¢ Less AT Papilionaccad ja Miystacede: ase Rutaccad | ==
Figure 6. Comparison of families with the highest phytosociological importance between the montane rain forest and seasonal rain forests at lower altitudes in the region.
TMRF: tropical montane rain forest in southern Yunnan;
LHSR: lower hill seasonal rain forest in southern Yunnan;
RSR: ravine seasonal rain forest in southern Yunnan.
102 Gard. Bull. Singapore 58 (2006)
Discussion
Altitudinal zonation of tropical forest
Montane vegetation zones in tropical America have been classified by Beard (1944, 1955) into rain forest, lower montane rain forest, montane rain forest, montane thicket and elfin woodland, with increasing altitude. Similarly, Richards (1952) used the terms tropical rain forest, submontane rain forest and montane rain forest for the vegetation zonation in tropical mountains. In contrast, Grubb et al. (1963), Whitmore (1984, 1990) and Ashton (2003) prefer the terms of lowland rain forest, lower montane rain forest and upper montane rain forest. The tropical montane rain forest in southern Yunnan occurs at an altitude comparable with lower montane rain forest zone as defined by Grubb et al. (1963), Whitmore (1984, 1990) and Ashton (2003).
Equatorial lower montane rain forests are 15—33 m tall and have two tree strata, few emergent trees, few trees with buttresses and cauliflory, few big woody lianas, and fewer plants with pinnate leaves. Plants with mesophyll (Grubb et al., 1963; Whitmore, 1984, 1990) or notophyll leaves (Ashton, 2003) are dominant among the woody plants, and there are abundant vascular epiphytes. Floristic zonation of forests in tropical mountains has been discussed by Ashton (2003), who stresses the laurel-oak attributes of the floras of lower montane rain forests in SE Asia.
The montane rain forest in southern Yunnan is similar to equatorial lower montane rain forests in SE Asia in physiognomy, but differs in having fewer epiphytes and more tree species with pinnate leaves (which contribute up to 11% of the sum of tree species).
The montane rain forest is dominated, in terms of species richness, by the families Lauraceae, Euphorbiaceae, Fagaceae, Rubiaceae, Papilionaceae and Theaceae. In terms of phytosociological importance the dominant families are Lauraceae, Magnoliaceae, Euphorbiaceae, Fagaceae, Mastixiaceae and Nyssaceae. The laurel-oak floristic attribute of the montane forest is overshadowed by some dominant families, such as Euphorbiaceae, Rubiaceae and Magnoliaceae, which are more commonly associated with lowland rain forests.
These differences may be due to the monsoonal climate (seasonal dryness) in southern Yunnan and the so-called “Massenerhebung”, or ‘mass elevation effect’ (Whitmore, 1990). This may reflect the fact that these montane forests in Yunnan have characteristics more usually associated
Tropical Montane Rain Forest in Southern Yunnan of China 103
with lowland sites. The montane rain forests in Yunnan may represent a transition between lowland and lower montane forest in physiognomy and floristics, but appears closer to lower montane rain forest.
The physiognomic changes observed with increasing altitudes in southern Yunnan are similar to those in tropical America (Grubb ef al., 1963). Microphyllous leaves increased with increasing altitudes.
Tropical montane rain forests in Yunnan were generally classified into a subtype of tropical rain forest by Wu (1987) based on their floristic composition and physiognomy. They are most similar to the lower montane rain forest in equatorial Asia, which was included under the category of tropical rain forest by Whitmore (1990). We agree with Wu and Whitmore’s classification that the montane rain forest in southern Yunnan is a type of lower montane rain forest within the broader category of tropical rain forest.
Biogeographical affinity
Floristically, the montane rain forest in southern Yunnan has strong tropical Asian affinities even though it occurs at the northern margin of mainland of Southeast Asia and at a high altitude. The tropical elements contribute about 78.9% at the generic level and more than 80 % at the specific level of its total flora. Elements with ‘tropical Asian’ affinities contribute 63.7% of the total sum of species.
Some species of particular biogeographical importance were encountered in these tropical montane rain forests in Yunnan. Mastixia euonymoidos is a dominant and the biggest tree in the montane rain forest. This species occurs only in the limited border area between Myanmar, Yunnan and Thailand, but it was widely distributed in European and America Tertiary flora, which has even been called the Mastixioidean European Flora (Mai, 1993; Eyde et al., 1990; Tiffney et a/., 1996). Its vicarious species, Mastixia octandra, occurs in mountains of central Sumatra in Indonesia (Matthew, 1976) at similar altitude (1700-1800 m alt.).
Gymnanthes remota (Euphorbiaceae), a relic and dominant species in the lower tree layer of the montane rain forest, occurs disjunctively in Mengsong in southern Yunnan and in Sumatra (Zhu et al., 2000). The frequent shrub species, Lasianthus inodorus (Rubiaceae), which is distributed in mainland SE Asia and Sumatra, as well as Java, also occurs vicariously on Mt Kinabalu
104 Gard. Bull. Singapore 58 (2006)
in Borneo (Zhu, 2001). It is interesting that many taxa in the montane rain forest in southern Yunnan have their vicarious species in Malesian montane forests, suggesting a special biogeographical significance for the region. Further floristic and biogeographical studies on the pristine montane rain forest in southern Yunnan are needed.
Acknowledgments
This project was funded by The National Natural Science Foundation of China (30570128). Figure 1 was made by Dr. Hu Huabin. I thank Professor R. Kitching from Griffith University of Australia for his English grammati- cal improvements in my manuscript. I also thank reviewers very much for their important and constructive comments.
References
Aiba, S. & K. Kitayama. 1999. Structure, composition and species diversity in an altitude-substrate matrix of rain forest tree communities on Mount Kinabalu, Borneo. Plant Ecology 140: 139-157.
Ashton, P.S. 2003. Floristic zonation of tree communities on wet tropical mountains revisited. Perspectives in Plant Ecology, Evolution and Systematics 6: 87-104.
Audley-Charles, M.G. 1987. Dispersal of Gondwanaland: Relevance to Evolution of the Angiosperms. In: Whitmore, T.C. (ed.) Biogeographical Evolution of the Malay Archipelago. Clarendon Press, Oxford.
Beard, J.S. 1944. Climax vegetation in tropical America. Ecology 25: 127- 158.
Beard, J.S. 1955. The classification of tropical American vegetation types. Ecology 36: 359-412.
Brown, W.H. 1919. Vegetation of the Philippine Mountains. Bureau of Science, Manila, Publication 13.
Buot, LE.Jr. & S. Okitsu. 1999. Leaf size zonation pattern of woody species along an altitudinal gradient on Mt. Pulog, Philippines. Plant Ecology 145: 197-208.
Tropical Montane Rain Forest in Southern Yunnan of China 105
Curtis, J.T. & R.P. McIntosh. 1951. An upland forest continuum in the prairie- forest border region of Wisconsin. Ecology 32: 467-496.
Eyde, R.H. & Q.Y. Xiang. 1990. Fossil Mastixioid (Cornaceae) alive in east- ern Asia. American Journal of Botany 77: 689-692.
Givnish, T.J. 1978. On the adaptive significance of compound leaves, with particular reference totropical trees. In:Tomlison,P.B.& M.H.Zimmerman. (eds.) Tropical trees as living systems, pp. 351-380. Cambridge Univ. Press, London.
Grubb, PJ., Lloyd, J.R., Pennington, T.D. & T.C. Whitmore. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy and floristics. Journal of Ecology 51: 567-601.
Hall, R. & J.D. Holloway. 1998. Biogeography and Geological Evolution of SE Asia. Backbuys Publishers, Leiden.
Kitayama, K. 1992. An altitudinal transect study of the vegetation on Mount Kinabalu, Borneo. Vegetatio 102: 149-171.
Mai, D.H. 1993. On the extinct Mastixiaeae (Cornales) in Europe. Geophytology 23(1): 53-63.
Matthew, K. M. 1976. A revision of the genus Mastixia (Cornaceae). Blumea 23: 51-93.
Meier, W. 1959. Plant sociological analysis of montane rain forest near Tjibodas, West Java. Acta Botanica Neerlandica 8: 277-291.
Metcalfe, I. 1998. Paleozoic and Mesozoic geological evolution of the SE Asia region: multidisciplinary constraints and implications for biogeography, pp. 25-41. In: Hall R. & Holloway J.D. (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden.
Morley, J.R. 1998. Palynological evidence for Tertiary plant dispersals in the SE Asian region in relation to plate tectonics and climate, pp. 221-234. In: Hall, R. & J.D. Holloway. (eds.) Biogeography and Geological Evolution of SE Asia. Backhuys Publishers, Leiden.
Mueller-Dombois, D. & H. Ellenberg. 1974. Aims and Methods of Vegetation Ecology. John Wiley & Sons.
Myers, N. 1998. Threatened biotas: “Hotspot” in tropical forests. Environmentalist 8: 1-20
106 Gard. Bull. Singapore 58 (2006)
Nakashizuka, T., Zulkifli, Y. & N. Rahim. 1992. Altitudinal zonation of forest communities in Selangor, Peninsular Malaysia. Journal of Tropical Forest Science 4: 233-244.
Ohsawa, M., Nainggolan, P.H.J., Tanaka, N. & C. Anwar. 1985. Altitudinal zonation of forest vegetation on Mount Kerinci,Sumatra: with comparisons to zonation in the temperate region of East Asia. Journal of Tropical Ecology 1: 192-216.
Ohsawa, M. 1991. Structural comparison of tropical montane forests along latitudinal and altitudinal gradients in south and east Asia. Vegetatio 97: 1-10.
Ohsawa, M. 1993. Latitudinal pattern of mountain vegetation zonation in southern and eastern Asia. Journal of Vegetation Science 4: 13-18.
Ohsawa, M. 1995. Latitudinal comparison of altitudinal changes in forest structure, leaf types, and species richness in humid monsoon Asia. Vegetatio 121: 3-10.
Paijmans, K. 1970. An analysis of four tropical rain forest sites in New Guinea. Journal of Ecology 58 (1): 77-101.
Pendry C. & J. Proctor. 1996. Altitudinal zonation of rain forest on Bukit Belalong, Brunei: soils, forest structure and floristics. Journal of Tropical Ecology 13: 221-241.
Proctor, J., Haridasan, K. & G.W. Smith. 1998. How far north does lowland evergreen tropical rain forest go? Global Ecology and Biogeography Letters 7: 141-146.
Qu, Z.-X. 1960. Nature reserves in Yunnan. Journal of Yunnan University (Natural Science ) 1: 1-4 (in Chinese).
Raunkiaer, C. 1934. The Life Forms of Plants and Statistical Plant Geography. Oxford University Press, Oxford.
Richards, P.W. 1952. The Tropical Rain Forest. Cambridge University Press, London.
Richards, P.W. 1996. The Tropical Rain Forest, an Ecological Study. Second edition. Cambridge University Press, London.
Steenis, C.G.G.J. van. 1935. On the origin of the Malaysian mountain flora. 2. Altitudinal zones, general considerations, and renewed statement of the problem. Bulletin du Jardin Botanique de Buitenzorg series 3, 13: 289-
Tropical Montane Rain Forest in Southern Yunnan of China 107
417.
Steenis, C.G.G.J. van. 1984. Floristic altitudinal zones in Malesia. Botanical Journal of the Linnean Society 89: 289-2972.
Tiffney, B.H. & K.K. Haggard. 1996. Fruits of Mastixioideae (Cornaceae) from the Paleogene of western North America. Review of Palaeobotany and Palynology 92: 29-54.
Wang, H., Zhu, H. & B.-G. Li. 2001. A study on the tropical montane rainforest in Mengsong, Xishuangbanna, S. Yunnan. Guihaia 21: 303-314 (in Chinese with English abstract).
Whitmore, T.C. & C.P. Burnham. 1969. The altitudinal sequence of forests and soils on granite near Kuala Lumpur. Malayan Natural Journal 22: 99-118.
Whitmore, T.C. 1984. Tropical Rain Forest of the Far East. Second edition. Clarendon Press, Oxford.
Whitmore, T.C. 1990. An Introduction to Tropical Rain Forests. Clarendon Press, Oxford.
Wu, C-Y. 1987. Vegetation of Yunnan, pp. 143-163. Science Press, Beijing (in Chinese).
Wu, C.-Y. 1991. The areal-types of Chinese genera of seed plants. Acta Botanica Yunnanica, Supplement IV (in Chinese with English abstract).
Zhu, H. 1997. Ecological and biogeographical studies on the tropical rain forest of south Yunnan, SW China with a special reference to its relation with rain forests of tropical Asia. Journal of Biogeography 24: 647-662.
Zhu, H., Wang, H. & B-G. Li. 1998a. Research on the tropical seasonal rainforest of Xishuangbanna, South Yunnan. Guihaia 18: 371-384 (in Chinese with English abstract).
Zhu, H., Wang, H. & B-G. Li. 1998b. The structure, species composition and diversity of the limestone vegetation in Xishuangbanna, SW China. Gardens’ Bulletin Singapore 50: 5-33.
Zhu, H., Wang, H. & B-G. Li. 2000. Gymnanthes Sw. (Euphorbiaceae), new to China and its biogeographical implication. Acta Phytotaxonomica Sinica 38: 462-463.
Zhu, H. 2001. New Plants of Lasianthus Jack (Rubiaceae) from Kinabalu,
108 Gard. Bull. Singapore 58 (2006)
Borneo and its biogeographical implication. Blumea 46: 447-455.
Zhu, H., Wang, H., B-G. Li & P. Sirirugsa. 2003. Biogeography and floristic affinity of the Limestone flora in southern Yunnan, China. Annals of the Missouri Botanical Garden 90: 44446.
Zhu, H., Wang, H. & B-G. Li. 2004. Plant diversity and physiognomy of a tropical montane rain forest in Mengsong, southern Yunnan, China. Acta Phytoecologica Sinica 28: 351-360.
Zhu, H. & M.C Roos. 2004. The tropical flora of S China and its affinity to Indo-Malesian flora. Telopea 10: 639-648.
Tropical Montane Rain Forest in Southern Yunnan of China 109
Appendix 3. Species checklist of the montane rain forest in Mengsong,
southern Yunnan.
ACANTHACEAE ACANTHACEAE ACANTHACEAE ACANTHACEAE ACANTHACEAE ACANTHACEAE ACANTHACEAE ACANTHACEAE ACERACE AE ACERACEAE ACERACEAE ALANGIACEAE ALANGIACEAE ALANGIACEAE ALISMATACEAE AMARANTHACEAE AMARANTHACEAE AMARYLLIDACEAE ANACARDIACEAE ANACARDIACEAE ANACARDIACEAE ANACARDIACEAE ANACARDIACEAE
ANACARDIACEAE
Lepidagathis incurva Buch.-Ham. ex D. Don Mananthes patentiflora ((Hemsl.) Bremek. Phaulopsis imbricata (Forssk.) Sweet
Phlogacanthus curviflorus (Wall.) Nees Pseuderanthemum malaccense (C.B. Clarke) Lindau Pteracanthus alatus (Wall.) Bremek. Rhaphidosperma vagabunda (R.Ben)C.Y.Wu ex Y.C.Tang Rungia pectinata (L.)Nees
Acer decandrum Mert.
Acer huianum W.P. Fang & C.K. Hsieh
Acer jingdongense T.Z. Hsu
Alangium barbatum (R. Br.) Baill.
Alangium chinense (Lour.) Harms
Alangium kurzii Craib
Sagittaria trifolia L.
Achyranthes bidentata Blume
Aerva sanguinolenta (L.) Blume
Allium hookeri Thwaites
Choerospondias axillaris (Roxb.) B.L. Burtt & A.W. Hill Pegia nitida Colobr.
Rhus chinensis Mill.
Semecarpus reticulata Lecomte
Spondias lakonensis var. hirsuta C.Y.Wu & T.L. Ming
Toxicodendron acuminatum (DC.) C. Y. Wu. T. L. Ming
110
ANACARDIACEAE ANNONACEAE ANNONACEAE ANNONACEAE ANNONACEAE ANNONACEAE ANNONACEAE ANNONACEAE ANNONACEAE APOCYNACEAE APOCYNACEAE APOCYNACEAE APOCYNACEAE APOSTASIACEAE AQUIFOLIACEAE AQUIFOLIACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE ARACEAE
ARACEAE
Gard. Bull. Singapore 58 (2006)
Toxicodendron succedaneum (L.) Kuntze Alphonsea boniana Finet & Gagnep. Alphonsea monogyna Merr. & Chun Alphonsea squamosa Finet & Gagnep. Alphonsea tsangyuanensis P.T. Li Fissistigma acuminatissimum Merr.
Fissistigma maclurei Merr.
Fissistigma polyanthum (Hook. f. & Thomson) Merr.
Mitrephora maingayi Hook. f. & Thomson Alstonia rostrata C.E.C. Fisch.
Bousigonia angustifolia Pierre
Epigynum auritum (C.K. Schneid.) Tsiang & P.T. Li Tabernaemontana corymbosa Roxb. ex Wall. Apostasia odorata Blume
Ilex polyneura (Hand.-Mazz.) S.Y. Hu
Ilex tetramera var. glabra (C.Y. Wu) T.R. Dudley Alocasia macrorrhizos (L.) Schott Amorphophallus bannanensis H. Li Amorphophallus rivieri Durieu ex Carriere Amorphophallus ximengensis H. Li
Arisaema austroyunnanense H. Li
Arisaema inkiangense H. Li
Colocasia esculenta (L.) Schott
Colocasia gigantea (Blume) Hook. f.
Gonatanthus pumilus (D. Don) Engl. & K. Krause Pothos chinensis (Raf.) Merr.
Pothos scandens L.
Tropical Montane Rain Forest in Southern Yunnan of China eg
ARACEAE
ARACEAE
ARACEAE
ARACEAE
ARACEAE
ARACEAE
ARACEAE ARALIACEAE ARALIACEAE ARALIACEAE ARALIACEAE ARALIACEAE ARALIACEAE ARALIACEAE ARISTOLOCHIACEAE ARISTOLOCHIACEAE ARISTOLOCHIACEAE ASCLEPIADACEAE BALANOPHORACEAE BALSAMINACEAE BALSAMINACEAE BEGONIACEAE BEGONIACEAE BEGONIACEAE BETULACEAE BETULACEAE
BETULACEAE
Remusatia hookeriana Schott
Remusatia vivipara (Lodd.) Schott Rhaphidophora crassicaulis Engl. & K. Krause Rhaphidophora decursiva (Roxb.) Schott Rhaphidophora hookeri Schott Rhaphidophora lancifolia Schott Rhaphidophora megaphylla H. Li
Aralia armata (Wall.) Seem.
Brassaiopsis producta (Dunn) C.B. Shang Macropanax dispermus (Blume) Kuntze Macropanax undulatus var. simplex H.L. Li Schefflera chapana Harms
Schefflera octophylla (Lour.) Harms Tupidanthus calyptratus Hook. & Thomson Aristolochia cathcartii Hook. f. Aristolochia fangchi Y.C. Wu ex L.D. Chow & S.M. Hwang Aristolochia tagala Cham.
Hoya villosa Costantin
Balanophora harlandii Hook. f.
Impatiens balansae Hook. f.
Impatiens mengtzeana Hook. f.
Begonia augustinei Hemsl.
Begonia crassirostris Irmsch.
Begonia versicolor Irmsch.
Alnus nepalensis D. Don
Betula alnoides Buch.-Ham. ex D. Don
Betula luminifera H.J.P. Winkl.
12 Gard. Bull. Singapore 58 (2006)
BIGNONIACEAE Mayodendron igneum (Kurz) Kurz BURSERACEAE Canarium pimela Leenh. BURSERACEAE Canarium strictum Roxb. BURSERACEAE Canarium tonkinense (Leenh.) Engl.
CAESALPINIACEAE Bauhinia variegata L. CAESALPINIACEAE Caesalpinia cucullata Roxb. CAESALPINIACEAE Cassia agnes (De Wit) Brenen
CAESALPINIACEAE Gleditsia fera (Lour.) Merr.
CAPPARIDACEAE Capparis fohaiensis B.S. Sun CAPRIFOLIACEAE Viburnum cylindricum Buch.-Ham. ex D. Don CAPRIFOLIACEAE Viburnum punctatum Buch.-Ham. ex D. Don
CARLEMANNIACEAE Silvianthus bracteatus Hook. f.
CELASTRACEAE Celastrus angulata Maxim.
CELASTRACEAE Celastrus paniculata subsp. multiflorus (Roxb.) Hou CELASTRACEAE Celastrus paniculatus Willd.
CELASTRACEAE Glyptopetalum sclerocarpum (Kurz) Lawson CELASTRACEAE Microtropis discolor (Wallich) Arn. CELASTRACEAE Microtropis tetragona Merr. & F.L. Freeman
CHLORANTHACEAE Sarcandra glabra subsp. brachystachys (Blume) Verdc.
COMMELINACEAE Amischotolype hispida (Less. & A. Rich.) D.Y. Hong COMMELINACEAE Amischotolype hookeri (Hassk.) H. Hara COMMELINACEAE Commelina paludosa Blume
COMMELINACEAE Cyanotis cristata (L.) D. Don
COMMELINACEAE Cyanotis vaga (Lour.) Roem. & Schult. COMMELINACEAE Dictyospermum conspicuum (Blume) Hassk. COMMELINACEAE Floscopa scandens Lour.
COMMELINACEAE Porandra scandens D.Y. Hong
Tropical Montane Rain Forest in Southern Yunnan of China 113
COMMELINACEAE COMPOSITAE COMPOSITAE COMPOSITAE COMPOSITAE COMPOSITAE CONNARACEAE CORNACEAE CORNACEAE CORYLACEAE CUCURBITACEAE CUCURBITACEAE CUCURBITACEAE CYPERACEAE CYPERACEAE DIOSCOREACEAE DIOSCOREACEAE DIOSCOREACEAE DIOSCOREACEAE EBENACEAE EBENACEAE EBENACEAE EBENACEAE ELAEAGNACEAE ELAEAGNACEAE
ELAEAGNACEAE
ELAEOCARPACEAE
Rhopalephora scaberrima (Blume) Faden Artemisia argyi H. Lév. & Vaniot Dichrocephala benthamii C.B. Clarke
Emilia prenanthoidea DC.
Senecio scandens Buch.-Ham. ex D. Don Vernonia cinerea (L.) Less.
Connarus paniculatus Roxb.
Mastixia euonymoides Prain
Mastixia pentandra subsp. chinensis (Merr.) K.M. Matthew Carpinus londoniana H.J.P. Winkl. Gynostemma laxum (Wall.) Cogn. Gynostemma pentaphyllum (Thunb.) Makino Gynostemma pubescens (Gagnep.) C.Y. Wu
Carex baccans Nees
Mariscus sumatrensis var. subcompositus (C.B. Clarke) S. Karthikeyan
Dioscorea bulbifera L.
Dioscorea chingii Prain & Burkill
Dioscorea esquirolii Prain & Burkill Dioscorea glabra Roxb.
Diospyros kaki var. silvestris Makino Diospyros kerrii Craib
Diospyros nigrocortex C.Y.Wu
Diospyros yunnanensis Rehder & E.H. Wilson Elaeagnus conferta var. menghaiensis W.K. Hu & H.F. Chow Elaeagnus gonyanthes Benth.
Elaeagnus macrantha Rehder
Elaeocarpus apiculatus Masters in Hook. f.
114
ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ELAEOCARPACEAE ERICACEAE ERICACEAE ESCALLONIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE
EUPHORBIACEAE
Gard. Bull. Singapore 58 (2006)
Elaeocarpus austroyunnanensis Hu Elaeocarpus decipiens Hemsl.
Elaeocarpus glabripetalus Merr.
Elaeocarpus glabripetalus var. alatus (Kunth) Hung T. Chang
Elaeocarpus howii Merr. & Chun Elaeocarpus petiolatus (Jack) Wall. ex Kurz Sloanea mollis Gagnep.
Sloanea tomentosa (Benth.) Rehder & E.H. Wilson Craibiodendron stellatum (Pierre) W.W. Sm. Rhododendron moulmainense Hook.
Itea macrophylla Wall.
Antidesma fordii Hemsl.
Antidesma montanum Blume
Aporusa dioica (Roxb.) Mill. Arg.
Aporusa villosa (Lindl.) Baill.
Aporusa yunnanensis (Pax & K. Hoffm.) E-P. Metcalf Baccaurea ramiflora Lour.
Baliospermum effusum Pax & Hoffm. in Engl. Baliospermum montanum (Willd.) Mill. Arg. Bischofia javanica Blume
Breynia fruticosa (L.) Hook. f.
Bridelia tomentosa Blume
Croton caudatus Geiseler
Croton damayeshu Y.T. Chang
Drypetes cumingii (Baill.) Pax & K. Hoffm. Drypetes salicifolia Gagnep.
Glochidion assamicum (Miill. Arg.) Hook. f.
Tropical Montane Rain Forest in Southern Yunnan of China
EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE EUPHORBIACEAE FAGACEAE
FAGACEAE
FAGACEAE
FAGACEAE
FAGACEAE
FAGACEAE
FAGACEAE
Glochidion hirsutum (Roxb.) Voigt Glochidion khasicum (Mill. Arg.) Hook. f. Glochidion lanceolarium (Roxb.) Voigt Glochidion puberum (L.) Hutch. Gymnanthes remota (Steenis) Esser Macaranga denticulata (Blume) Mill. Arg. Macaranga henryi (Pax & K. Hoffm.) Rehder
Macaranga indica Wight
Macaranga kurzii (Kuntze) Pax & Hoffm. in Engl.
Mallotus barbatus (Wall.) Mill. Arg. Mallotus macrostachyus (Miq.) Mill. Arg. Mallotus paniculatus (Lam.) Mill. Arg. Mallotus philippinensis (Lam.) Mill. Arg. Mallotus tetracoccus (Roxb.) Kurz Ostodes katharinae Pax
Ostodes kuangii Y.T. Chang
Ostodes paniculata Blume
Phyllanthus emblica |i,
Sapium baccatum Roxb.
Sapium discolor (Champ. ex Benth.) Miill. Arg.
Castanopsis argyrophylla King ex Hook. f.
Castanopsis calathiformis (Skan) Rehder & E.H. Wilson
b15
Castanopsis carlesii var. spinulosa W.C. Cheng & C.S. Chao
Castanopsis ceratacantha Rehder & E.H. Wilson
Castanopsis echidnocarpa Hook. f. & Thomson ex Miq.
Castanopsis hystrix Miq.
Castanopsis indica (Roxburgh ex Lindl.) A. DC.
116
FAGACEAE FAGACEAE FAGACEAE FAGACEAE FAGACEAE FPAGACEAE FAGACEAE FAGACEAE FAGACEAE FAGACEAE FAGACEAE FAGACEAE FAGACEAE FLACOURTIACEAE FLACOURTIACEAE
FUMARIACEAE
GENTIANACEAE GESNERIACEAE GNETACEAE GNETACEAE GNETACEAE GUTTIFERAE GUTTIFERAE HAMAMELIDACEAE HAMAMELIDACEAE HYDRANGIACEAE
HYPERICACEAE
Gard. Bull. Singapore 58 (2006)
Castanopsis mekongensis A. Camus
Castanopsis tcheponensis Hickel & A. Camus Cyclobalanopsis kerrii (Craib) Hu
Cyclobalanopsis myrsinifolia (Blume) Oerst. Lithocarpus fohaiensis (Hu) A. Camus
Lithocarpus fordianus (Hemsl.) Chun
Lithocarpus grandifolius (D. Don) S.N. Biswas Lithocarpus hancei (Benth.) Rehder
Lithocarpus hypoglaucus (Hu) C.C. Huang
Lithocarpus microspermus A.Camus
Lithocarpus pseudoreinwardti A. Camus
Lithocarpus rhabdostachyus subsp. dakhaensis A. Camus Lithocarpus truncatus (King ex Hook. f.) Rehder & E.H. Wilson Xylosma congesta (Lour.) Merr.
Xylosma longifolia Clos
Corydalis balansae Prain
Tripterospermum membranaceum (C. Marquand) Harry Sm. Rhynchotechum ellipticum (Wall. ex D. Dietr.) A. DC. Gnetum montanum fo. megalocarpum Markgr.
Gnetum montanum Markgr.
Gnetum pendulum C.Y. Cheng
Calophyllum polyanthum Wall. ex Choisy
Garcinia cowa Roxb.
Altingia excelsa Noronha
Distyliopsis yunnanensis (Hung T. Chang) C.Y. Wu
Dichroa febrifuga Lour.
Cratoxylum cochinchinense (Lour.) Blume
Tropical Montane Rain Forest in Southern Yunnan of China
HYPPOCRATEACEAE ICACINACEAE ICACINACEAE ICACINACEAE ICACINACEAE ICACINACEAE ICACINACEAE ICACINACEAE JUGLANDACEAE JUGLANDACEAE JUGLANDACEAE JUGLANDACEAE LABIATAE LABIATAE LABIATAE LABIATAE LABIATAE LABIATAE LARDIZABALACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE
LAURACEAE
Pristimera arborea (Roxb.) A.C. Sm. Apodytes dimidiata E. Mey. ex Arn. Gomphandra tetrandra (Wall.) Sleumer Todes ovalis Blume
Mappianthus itodoides Hand.-Mazz. Natsiatopsis thunbergiaefolia Kurz Nothapodytes collina C.Y. Wu
Platea latifolia Blume
Engelhardia roxburghiana Wall. Engelhardia serrata Blume
Engelhardia spicata Lesch. ex Blume Juglans sigillata Dode
Gomphostemma arbusculum C.Y. Wu Gomphostemma crinitum Wall. ex Benth. Gomphostemma stellatohirsutum C.Y. Wu Leucosceptrum canum Sm.
Paraphlomis javanica (Blume) Prain Pogostemon glaber Benth.
Stauntonia Denies Wall. ex Hemsl. Actinodaphne henryi Gamble Actinodaphne obovata (Nees) Blume Alseodaphne andersonii (King ex Hook. f.) Kosterm. Alseodaphne petiolaris (Meisn.) Hook. f. Beilschmiedia linocieroides H.W. Li Beilschmiedia percoriacea C.K. Allen Beilschmiedia purpurascens H.W. Li
Beilschmiedia robusta C.K. Allen
iy
118
LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE LAURACEAE
LAURACEAE
Gard. Bull. Singapore 58 (2006)
Beilschmiedia roxburghiana Nees Beilschmiedia yunnanensis Hu
Cassytha filiformis L.
Cinnamomum austroyunnanense H.W. Li Cinnamomum bejolghota (Buch.-Ham.) Sweet Cinnamomum glanduliferum (Wall.) Nees Cinnamomum iners Reinw. ex Blume Cinnamomum mollifolium H.W. Li Cinnamomum tamala (Buch.-Ham.) T. Nees & Eberm. Cinnamomum tenuipilis Kosterm.
Cryptocarya brachythyrsa H.W. Li Cryptocarya calcicola H.W. Li
Cryptocarya densiflora Blume
Cryptocarya rolletii H. Wang & H. Zhu Iteadaphne caudata (Nees) H.W. Li
Lindera latifolia Hook. f.
Lindera menghaiensis H.W. Li
Lindera metcalfiana var. dictyophylla (C.K. Allen) H.B. Cui
Litsea atrata S.K. Lee
Litsea balansae Lecomte
Litsea baviensis Lecomte
Litsea chinpingensis Yen C. Yang & P.H. Huang Litsea cubeba (Lour.) Pers.
Litsea elongata (Nees) Benth. & Hook. f.
Litsea euosma W.W. Sm.
Litsea garrettii Gamble
Litsea glutinosa (Lour.) C.B. Rob.
Tropical Montane Rain Forest in Southern Yunnan of China 119
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAE
LAURACEAEB
LAURACEAE
LAURACEAE
LAURACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
LILIACEAE
Litsea lancifolia (Roxb. ex Nees in Wall.) Benth. & Hook. f. ex Villar Litsea lancifolia var. ellipsoidea Yen C. Yang & P.H. Huang Litsea lancifolia var. pedicellata Hook. f.
Litsea liyuyingi H. Liu
Litsea longistaminata (H. Liu) Kosterm.
Litsea magnoliifolia Yen C. Yang & P.H. Huang
Litsea vang Lecomte var. lobata Lecomte
Litsea verticillata Hance
Machilus salicina Hance
Persea robusta (W.W. Sm.) Kosterm.
Persea rufipes (H.W. Li) Kosterm.
Persea shweliensis (W.W. Sm.) Kosterm.
Phoebe lanceolata (Nees) Nees
Phoebe macrocarpa C.Y. Wu
Phoebe puwenensis Cheng
Phoebe rufescens H.W. Li
Asparagus subscandens F.T. Wang & S.C. Chen
Aspidistra typica Baill.
Campylandra Holeues (Baker) M.N. Tamura, S.Yun Liang & Turland Chlorophytum malayense Rid.
Dianella ensifolia (L.) DC.
Disporopsis longifolia Craib
Disporum calcaratum D. Don
Disporum cantoniense (Lour.) Merr.
Liriope graminifolia (L.) Baker
Ophiopogon tsaii FT. Wang & Ts. Tang
Peliosanthes sinica ..T. Wang & Ts. Tang
120 Gard. Bull. Singapore 58 (2006)
LILIACEAE Reineckea carnea (Andrews) Kunth LILIACEAE Smilax hemsleyana Craib
LILIACEAE Smilax hypoglauca Benth.
LILIACEAE Smilax megacarpa A. DC.
LILIACEAE Smilax myrtillus A. DC.
LILIACEAE Smilax ocreata A. DC.
LILIACEAE Smilax perfoliata Lour.
LILIACEAE Smilax quadrata A. DC.
LILIACEAE Tupistra grandistigma F.T. Wang & S.Yun Liang LOGANIACEAE Buddleja officinalis Maxim.
LYTHRACEAE Rotala rotundifolia (Buch.-Ham. ex Roxb.) Koehne MAGNOLIACEAE Alcimanara cathcartii (Hook. f. & Thomson) Dandy MAGNOLIACEAE Manglietia forrestii W.W. Sm.ex Dandy MAGNOLIACEAE Manglietia garrettii Craib
MAGNOLIACEAE Manglietia insignis (Wall.) Blume MAGNOLIACEAE Michelia cavaleriei Finet & Gagnep. MAGNOLIACEAE Michelia floribunda Finet & Gagnep. MAGNOLIACEAE Michelia hedyosperma Y.W. Law MAGNOLIACEAE Parakmeria yunnanensis Hu MAGNOLIACEAE Paramichelia baillonii (Pierre) Hu MALVACEAE Hibiscus indicus (Burm. f.) Hochr. MALVACEAE Kydia calycina Roxb.
MALVACEAE Kydia glabrescens var. intermedia S.Y. Hu MALVACEAE Sida szechuensis Matsuda
MALVACEAE Urena lobata L.
MARANTACEAE Phrynium placentarium (Lour.) Merr.
MARANTACEAE Stachyphrynium sinense H. Li
Tropical Montane Rain Forest in Southern Yunnan of China
MELASTOMACEAE MELASTOMACEAE MELASTOMACEAE MELASTOMACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MELIACEAE MENISPERMACEAE MENISPERMACEAE MIMOSACEAE MIMOSACEAE MIMOSACEAE MIMOSACEAE MIMOSACEAE MIMOSACEAE MIMOSACEAE MORACEAE MORACEAE MORACEAE
MORACEAE
Medinilla septentrionalis (W.W. Sm.) H.L. Li Melastoma affine D. Don
Melastoma normale D. Don
Oxyspora vagans (Roxb.) Wall.
Aglaia abbreviata C.Y. Wu
Aglaia perviridis Hiern
Amoora yunnanensis (H.L. Li) C.Y. Wu
Dysoxylum binectariferum (Roxb.) Hook. f. ex Bedd.
Dysoxylum lukti Merr.
Melia toosendan Siebold & Zucc.
Toona ciliata M. Roem.
Toona sinensis (Juss.) Roem.
Trichilia connaroides (Wight & Arn.) Bentv. Walsura yunnanensis C.Y. Wu
Cocculus laurifolius DC.
Stephania forsteri (DC.) A. Gray
Albizia bracteata Dunn
Albizia chinensis (Osbeck) Merr.
Albizia crassiramea Lace
Albizia lucidior (Steud.) I.C. Nielsen Albizia odoratissima (L. f.) Benth. Cylindrokelupha kerrii (Gagnep.) T.L. Wu Pithecolobium clypearia Benth.
Artocarpus lakoocha Wall. ex Roxb.
Artocarpus nitidus subsp. griffithii (King) F.M. Jarrett
Artocarpus tonkinensis A. Chev.
Broussonetia papyrifera (L.) L’Hér. ex Vent.
120
122
MORACEAE MORACEAE MORACEAE MORACEAE MORACEAE MORACEAE MORACEAE MUSACEAE MYRICACEAE MYRISTICACEAE MYRISTICACEAE MYRISTICACEAE MYRISTICACEAE MYRISTICACEAE MYRISTICACEAE MYRSINACEAE MY RSINACEAE MYRSINACEAE MY RSINACEAE MYRSINACEAE MYRSINACEAE MYRSINACEAE MYRSINACEAE MYRSINACEAE MYRSINACEAE MYRSINACEAE
MYRTACEAE
Gard. Bull. Singapore 58 (2006)
Ficus auriculata Lour.
Ficus cyrtophylla Wall. ex Miq.
Ficus esquiroliana H. Lév.
Ficus fistulosa Reinw. ex Blume
Ficus hookeriana Corner
Ficus semicordata Buch.-Ham. ex Sm. Morus macroura Miq.
Musa acuminata Colla
Myrica esculenta Buch.-Ham. ex D. Don Horsfieldia glabra (Reinw. ex Blume) Warb. Horsfieldia tetratepala C.Y. Wu
Knema cinerea vat. glauca (Blume) Y.H. Li Knema erratica (Hook. f. & Thomson) J. Sincl. Knema furfuracea (Hook. f. & Thomson) Warb. Knema globularia (Lam.) Warb.
Ardisia corymbifera Mez
Ardisia depressa C.B.Clarke
Ardisia thyrsiflora D. Don
Ardisia villosa Roxb.
Ardisia virens Kurz
Embelia laeta (L.) Mez
Maesa indica (Roxb.) A. DC.
Maesa macilentoides C. Chen
Maesa perlaria (Lour.) Merr.
Maesa permollis Kurz
Myrsine seguinii H. Lév.
Decaspermum fruticosum J.R. Forst. & G. Forst.
Tropical Montane Rain Forest in Southern Yunnan of China 123
MYRTACEAE MYRTACEAE MYRTACEAE MYRTACEAE MYRTACEAE MYRTACEAE MYRTACEAE NYSSACEAE NYSSACEAE NYSSACEAE OLACACEAE OLEACEAE OLEACEAE OLEACEAE OLEACEAE OLEACEAE OLEACEAE OLEACEAE OXALIDACEAE PAPILIONACEAE PAPILIONACEAE PAPILIONACEAE PAPILIONACEAE PAPILIONACEAE PAPILIONACEAE PAPILIONACEAE
PAPILIONACEAE
Syzygium brachythyrsum Merr. & L.M. Perry Syzygium cathayense Merr. & L.M. Perry Syzygium polypetaloideum Merr. & L.M. Perry Syzygium rockii Merr. & L.M. Perry Syzygium tetragonum (Wight) Wall. ex Walp. Syzygium thumra (Roxb.) Merr. & L.M. Perry Syzygium yunnanense Merr. & L.M. Perry Nyssa wenshanensis Fang & Soong
Nyssa wenshanensis var. longipedunculata W.P. Fang & Soong Nyssa yunnanensis W. C. Yin
Schoepfia fragrans Wall.
Chionanthus ramiflorus Roxb.
Fraxinus floribunda Wall.
Jasminum attenuatum Roxb. ex G. Don Jasminum lanceolarium Roxb.
Ligustrum sinense Lour.
Linociera insignis C.B. Clarke
Olea rosea Craib
Oxalis corniculata L.
Craspedolobium schochii Harms
Dalbergia assamica Benth.
Dalbergia pinnata (Lour.) Prain
Dalbergia stipulacea Roxb.
Erythrina subumbrans (Hassk.) Merr.
Fordia cauliflora Hemsl.
Fordia microphylla Dunn ex Z. Wei
Millettia leptobotrya Dunn
124 Gard. Bull. Singapore 58 (2006)
PAPILIONACEAE Millettia pachycarpa Benth. PAPILIONACEAE Millettia tetraptera Kurz PAPILIONACEAE Millettia unijuga Gagnep. PAPILIONACEAE Mucuna pruriens (L.) DC. PAPILIONACEAE Ormosia fordiana Oliv. PAPILIONACEAE Ormosia olivacea L. Chen PAPILIONACEAE Pycnospora lutescens (Poir.) Schindl. PAPILIONACEAE Spatholobus pulcher Dunn PASSIFLORACEAE Passiflora siamica Craib PASSIFLORACEAE Passiflora wilsonii Hemsl. PINACEAE Pinus kesiya Royle ex Gordon PIPERACEAE Peperomia blanda (Jacq.) Kunth PIPERACEAE Peperomia heyneana Mia. PIPERACEAE Peperomia pellucida (L.) Kunth PIPERACEAE Peperomia tetraphylla (G. Forst.) Hook. & Arn. PIPERACEAE Piper chaudocanum C. DC. PIPERACEAE Piper flaviflorum C. DC. PIPERACEAE Piper longum L.
PIPERACEAE Piper macropodum C. DC. PIPERACEAE Piper thomsonii (C. DC.) Hook. f. PIPERACEAE Piper yunnanense Y.Q. Tseng PITTOSPORACEAE Pittosporum kerrii Craib PLANTAGINACEAE Plantago erosa Wall. ex Roxb. PLANTAGINACEAE Plantago major L.
POACEAE Fargesia plurisetosa T.H. Wen POACEAE Imperata cylindrica (L.) P. Beauv.
POACEAE Microstegium ciliatum (Trin.) A. Camus
Tropical Montane Rain Forest in Southern Yunnan of China ‘75
POACEAE POACEAE PODOCARPACEAE POLYGALACEAE POLYGALACEAE POLYGALACEAE POLYGONACEAE POLYGONACEAE POLYGONACEAE POLYGONACEAE POLYGONACEAE POLYGONACEAE POLYGONACEAE PORTULACACEAE PROTEACEAE PROTEACEAE PROTEACEAE PROTEACEAE PROTEACEAE PROTEACEAE PROTEACEAE PROTEACEAE RANUCULACEAE RANUCULACEAE RANUCULACEAE RHAMNACEAE
RHAMNACEAE
Setaria palmifolia (J. Konig) Stapf Thysanolaena maxima (Roxb.) Kuntze Podocarpus neriifolius D. Don
Polygala arillata Buch.-Ham. ex D. Don Polygala glomerata Lout.
Securidaca inappendiculata Hassk. Polygonum chinense L.
Polygonum chinense vat. hispidum Hook. f. Polygonum chinense vat. ovalifolium Meisn. Polygonum hydropiper L.
Polygonum lapathifolium L.
Polygonum orientale L.
Polygonum perfoliatum L.
Portulaca oleracea L.
Helicia cochinchinensis Lour.
Helicia nilagirica Bedd.
Helicia pyrrhobotrya Kurz
Helicia reticulata W.T. Wang
Felicia iciaiets W.W. Sm.
Helicia silvicola W.W. Sm.
Helicia tsati W.T. Wang
Heliciopsis terminalis (Kurz) Sleumer Clematis fulvicoma Rehder & E.H. Wilson Clematis peterae Hand.-Mazz.
Clematis subumbellata Kurz
Gouania leptostachya DC.
Hovenia acerba var. kiukiangensis (Hu & Cheng) C. Y. Wu ex Y. L. Chen
126
RHAMNACEAE RHAMNACEAE RHIZOPHORACEAE RHIZOPHORACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE ROSACEAE RUBIACEAE RUBIACEAE RUBIACEAE
RUBIACEAE
Gard. Bull. Singapore 58 (2006)
Rhamunus leptophylla C.K. Schneid.
Ventilago calyculata Tul.
Carallia brachiata (Lour.) Merr.
Carallia diplopetala Hand.-Mazz.
Cerasus cerasoides (Buch.-Ham. ex D. Don) S.Y. Sokolov Docynia delavayi (Franch.) C.K. Schneid. Duchesnea chrysantha (Zoll. & Moritzi) Mig. Eriobotrya bengalensis var. angustifolia Cardot Eriobotrya obovata W.W. Sm.
Laurocerasus jenkinsii (Hook. f.) Browicz Laurocerasus menghaiensis T.T. Yu & L.T. Lu Laurocerasus zippeliana (Mig.) Yu et Lu Photinia glabra (Thunb.) Maxim.
Potentilla kleiniana Wight & Arn.
Pygeum arboretum (Bl.) C. Kalkman
Pygeum topengii Merr.
Pyrus pashia Buch.-Ham. ex D. Don
Rubus pirifolius Sm.
Rubus poliophyllus Kuntze
Rubus rufus var. palmatifidus Cardot
Sorbus corymbifera (Miq.) Khep & Yakovlev Sorbus globosa T.T. Yu & Tsai
Stranvaesia oblanceolata (Rehder & E.H. Wilson) Stapf Aidia cochinchinensis Lour.
Brachytome hirtellata var. glabrescens W.C. Chen Canthium parvifolium Roxb.
Discospermum fruticosum (Hemsl.) Kuntze
Tropical Montane Rain Forest in Southern Yunnan of China 27
RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUBIACEAE RUTACEAE
RUTACEAE
RUTACEAE
RUTACEAE
RUTACEAE
RUTACEAE
RUTACEAE
RUTACEAE
Geophila herbacea (Jacq.) K. Schum.
Hedyotis capitellata var. mollissima (Pit.) W.C. Ko Hedyotis diffusa Willd.
Hedyotis scandens Roxb.
Lasianthus inodorus Bl.
Lasianthus lucidus B1.
Lasianthus sikkimensis Hook.f.
Metadina trichotoma (Zoll. & Moritzi) Bakh. f. Mussaenda hossei Craib
Mycetia gracilis Craib
Ophiorrhiza mungos L.
Oxyceros sinensis Lour.
Psychotria symplocifolia Kurz
Tarennoidea wallichii (Hook. f.) Tirveng. & Sastre Uncaria laevigata Wall. ex G. Don
Uncaria sessilifructus Roxb.
Wendlandia pingpienensis F.C. How
Wendlandia scabra Kurz
Wendlandia tinctoria (Roxb.) DC.
Acronychia pedunculata (L.) Miq.
Evodia austrosinensis Hand.-Mazz.
Evodia glabrifolia (Champ. ex Benth.) C.C. Huang Evodia lepta (Spreng.) Merr.
Evodia lepta var. cambodiana (Pierre) C.C. Huang Evodia simplicifolia Ridl.
Evodia trichotoma (Lour.) Pierre
Paramignya rectispina Craib
128
RUTACEAE SABIACEAE SABIACEAE SALICACEAE SAMYDACEAE SAMYDACEAE SAPINDACEAE SAPINDACEAE SAPINDACEAE SAPOTACEAE SAPOTACEAE SAPOTACEAE SAPOTACEAE SAPOTACEAE SAPOTACEAE SAPOTACEAE SAURAUIACEAE SAURAUIACEAE SAURAUIACEAE SAURAUIACEAE SAURAUIACEAE SAURAUIACEAE SCHIZANDRACEAE SCHIZANDRACEAE SCHIZANDRACEAE SCHIZANDRACEAE
SCHIZANDRACEAE
Gard. Bull. Singapore 58 (2006)
Toddalia asiatica (L.) Lam.
Meliosma simplicifolia (Roxb.) Walp.
Meliosma velutina Rehder & E.H. Wilson
Salix tetrasperma Roxb.
Casearia balansae Gagnep.
Casearia velutina Blume
Dimocarpus yunnanensis (W.T. Wang) C.Y. Wu & T.L. Ming Nephelium chryseum Blume
Sapindus rarak DC.
Pouteria grandifolia (Wall.) Baehni
Sarcosperma arboreum Buch.-Ham. ex C.B. Clarke Sarcosperma griffithii Hook. f. ex C.B. Clarke Sarcosperma kachinense var. simondii (Gagnep.) H.J. Lam & P. Royen Xantolis boniana var. rostrata (Mertr.) P. Royen Xantolis stenosepala (Hu) P. Royen
Xantolis stenosepala var. brevistylis C.Y. Wu Saurauia cerea Griff. ex Dyer
Saurauia macrotricha Kurz ex Dyer
Saurauia miniata C.F, Liang & Y.S. Wang
Saurauia napaulensis DC.
Saurauia punduana Wall.
Saurauia yunnanensis C.F. Liang & Y.S. Wang Kadsura ananosma Kerr
Kadsura angustifolia A.C.Smith
Schisandra henryi var. yunnanensis A.C. Sm. Schisandra neglecta A.C. Sm.
Schisandra plena A.C. Sm.
Tropical Montane Rain Forest in Southern Yunnan of China 129
SCROPHULARIACEAE _ Lindenbergia indica (L.) Vatke
SLADENIACEAE Sladenia celastrifolia Kurz
SOLANACEAE Lycianthes biflora (Lour.) Bitter SOLANACEAE Lycianthes biflora var. subtusochracea Bitter SOLANACEAE Solanum aculeatissimum Jacq. SOLANACEAE Solanum anguivi Lam.
SOLANACEAE Solanum erianthum D.Don SOLANACEAE Solanum merrillianum Liou SOLANACEAE Solanum spirale Roxb.
SOLANACEAE Solanum torvum Sw.
STAPH YLACEAE Tapiscia yunnanensis W.C. Cheng & C.D. Chu STAPHYLACEAE Turpinia cochinchinensis (Lour.) Merr. STAPHYLACEAE Turpinia pomifera (Roxb.) DC. STEMONACEAE Stemona tuberosa Lour. STERCULIACEAE Pterospermum acerifolium Willd. STERCULIACEAE Reevesia pubescens Mast. STERCULIACEAE Reevesia thrsoidea Lindl. STERCULIACEAE Sterculia lanceifolia Roxb. STERCULIACEAE Sterculia lanceolata Cav.
STYRACACEAE Bruinsmia polysperma (Clarke) Steenis STYRACACEAE Styrax grandiflorus Griff.
STYRACACEAE Styrax rugosus Kurz
STYRACACEAE Styrax tonkinensis (Pierre) Craib ex Hartwich SYMPLOCACEAE Symplocos sulcata Kurz SYMPLOCACEAE Symplocos wikstroemiifolia Hayata TACCACEAE Tacca chantrieri André
THEACEAE Adinandra megaphylla Hu
130
THEACEAE THEACEAE THEACEBAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THEACEAE THYMELEACEAE TILIACEAE TILIACEAE TILIACEAE ULMACEAE ULMACEAE ULMACEAE ULMACEAE URTIACEAE URTIACEAE
URTIACEAE
Gard. Bull. Singapore 58 (2006)
Camellia sinensis var. assamica (J.W. Mast.) Kitam. Camellia pachyandra Hu
Camellia sinensis (L.) Kuntze
Eurya aurea H.T. Chang
Eurya austroyunnanensis T.L. Ming & H. Chu Eurya groffti Merr.
Eurya jintungensis Hu & L.K. Ling
Eurya persicaefolia Gagnep.
Eurya pseudocerasifera Kobuski
Gordonia chrysandra Cowan
Pyrenaria yunnanensis Hu
Schima argentea E. Pritz.
Schima khasiana Dyer
Schima wallichii Choisy
Ternstroemia gymnanthera (Wight & Arn.) Bedd. Tutcheria pingpienensis Hung T. Chang Eriosolena composita (L. f.) Tiegh.
Colona floribunda (Wall. ex Voigt) Craib Microcos chungii (Merr.) Chun
Microcos paniculata L.
Celtis sinensis Pers.
Celtis timorensis Span.
Gironniera subaequalis Planch.
Trema orientalis (L.) Blume
Boehmeria macrophylla Hornem. Debregeasia libera Chien et C.J. Chen
Debregeasia longifolia (Burm. f.) Wedd.
Tropical Montane Rain Forest in Southern Yunnan of China 151
URTIACEAE URTIACEAE URTIACEAE VACCINIACEAE VACCINIACEAE VACCINIACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VERBENACEAE VIOLACEAE VIOLACEAE VITACEAE VITACEAE
VITACEAE
XANTHOPHYLLACEAE
ZINGIBERACEAE
Debregeasia squamata King ex Hook. f.
Dendrocnide sinuata (Blume) Chew
Oreocnide rubescens (Blume) Miq.
Agapetes lobbii C.B. Clarke
Agapetes mannii Hemsl.
Vaccinium exaristatum Kurz
Callicarpa arborea Roxb.
Callicarpa bodinieri H. Lév.
Callicarpa cathayana H.T. Chang
Callicarpa giraldii Hesse ex Rehder
Callicarpa longifolia Lam.
Clerodendrum bungei Steud.
Clerodendrum colebrookianum Walp.
Clerodendrum japonicum (Thunb.) Sweet Clerodendrum serratum (L.) Moon
Clerodendrum serratum var. amplexifolium Moldenke Clerodendrum serratum var. herbaceum (Roxb. ex Schauer) C.Y. Wu Clerodendrum villosum Blume
Premna er pos Roxb.
Vitex quinata var. puberula (H.J. Lam) Moldenke Viola diffusoides Ching J. Wang
Viola hossei W. Becker
Ampelopsis cantoniensis (Hook. & Arn.) Planch. Cayratia timoriensis var. mekongensis (C.Y. Wu) C.L. Li Tetrastigma obovatum (Lawson) Gagnep. Xanthophyllum yunnanense C.Y.Wu
Amomum koenigii J.F. Gmel.
132
ZINGIBERACEAE ZINGIBERACEAE ZINGIBERACEAE ZINGIBERACEAE
ZINGIBERACEAE
Boesenbergia rotunda (L.) Mansf. Globba barthei Gagnep. Globba racemosa Sm.
Globba schomburgkii Hook. f.
Rhynchanthus beesianus W.W. Sm.
Gard. Bull. Singapore 58 (2006)
Gardens’ Bulletin Singapore 58 (2006) 133—139 133
OBITUARY
Humphrey Morrison Burkill O.B.E., EL.S. (1914—2006)
The career and achievements of Humphrey Burkill, who died recently aged 92, parallel those of his father, Isaac Henry Burkill, to a remarkable degree. Both went to Repton School and Gonville and Caius College, Cambridge, both served as Director of Singapore Botanic Gardens and both authored monumental works on the economic botany of large tropical regions. While the direct or indirect influence of his father may have set Humphrey off in his footsteps fate seems to have conspired to keep him returning to the same track. Presumably even his longevity owed much to his parents’ staying power.
However, father and son were of very different generations. Burkill senior was very much of the Colonial Age. He served in British India and Malaya at a time when empire was unquestioned. Burkill junior was a