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Harvard Botany Librari

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The Gardens' Bulletin

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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

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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

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Dr. M.J.E. Coode Dr. W.K. Tan

Royal Botanic Gardens Kew. U.K. National Parks Board, Singapore

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conclusions expressed by the contributing authors.

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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

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.

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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)

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.

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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.

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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.

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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.

—

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

10.

th.

£2:

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14.

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).

18.

ry.

20.

ras

De.

23:

24.

20:

28.

22.

30.

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

P. undulatum

P. maculosum

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

64 P. bicolor

P. diffusum

P. macphersonii

P. kunstleri

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

P. floresanum

P. simalurense

P. maculosum

P. fuscum

P. diffusum

P. marsupiale

64 P. kunstleri

P. tonkinense

P. macphersonii

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

2 19 29

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0-1 0-1 0-1

8 9 11 12 20 26

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Ub 0-1 2-30-11-0 1-0 0-1

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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).

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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

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1.37

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0.84

1.68

0.84

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0.84

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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

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.

Tropical Montane Rain Forest in Southern Yunnan of China

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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

Species %

NY iG

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

)

oo)

ioe)

ro)

Rubiaceae

Lauraceae

Euphorbiaceae

Annonaceae

Moraceae

Orchidaceae

Meliaceae

Vitaceae

Apocynaceae

Urticaceae

Rutaceae

Fagaceae

Acanthaceae

Papilionaceae

Piperaceae

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

MM ee

SSS ee

a ee a

———

— ee

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.

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Tropical Montane Rain Forest in Southern Yunnan of China 109

Appendix 3. Species checklist of the montane rain forest in Mengsong,

southern Yunnan.

ACANTHACEAE

ACERACE AE

ACERACEAE

ALANGIACEAE

ALISMATACEAE

AMARANTHACEAE

AMARYLLIDACEAE

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

APOCYNACEAE

APOSTASIACEAE

AQUIFOLIACEAE

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

ARALIACEAE

ARISTOLOCHIACEAE

ASCLEPIADACEAE

BALANOPHORACEAE

BALSAMINACEAE

BEGONIACEAE

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

CONNARACEAE

CORNACEAE

CORYLACEAE

CUCURBITACEAE

CYPERACEAE

DIOSCOREACEAE

EBENACEAE

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

ERICACEAE

ESCALLONIACEAE

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

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

FPAGACEAE

FAGACEAE

FLACOURTIACEAE

FUMARIACEAE

GENTIANACEAE

GESNERIACEAE

GNETACEAE

GUTTIFERAE

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

JUGLANDACEAE

LABIATAE

LARDIZABALACEAE

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

118

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

LAURACEAEB

LAURACEAE

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

MELIACEAE

MENISPERMACEAE

MIMOSACEAE

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

MUSACEAE

MYRICACEAE

MYRISTICACEAE

MYRSINACEAE

MY RSINACEAE

MYRSINACEAE

MY RSINACEAE

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

NYSSACEAE

OLACACEAE

OLEACEAE

OXALIDACEAE

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

PODOCARPACEAE

POLYGALACEAE

POLYGONACEAE

PORTULACACEAE

PROTEACEAE

RANUCULACEAE

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

RHIZOPHORACEAE

ROSACEAE

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

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

SALICACEAE

SAMYDACEAE

SAPINDACEAE

SAPOTACEAE

SAURAUIACEAE

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

THEACEBAE

THEACEAE

THYMELEACEAE

TILIACEAE

ULMACEAE

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

VACCINIACEAE

VERBENACEAE

VIOLACEAE

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

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

prisoner of the Japanese during World War II and served as Director of

Singapore Botanic Gardens up to and after independence.

134 Gard. Bull. Singapore 58 (2006)

Humphrey Burkill’s parents were first cousins and his mother,

Ethel Maud Morrison, was 40 when he was born in the Director’s House,

now Burkill Hall, in Singapore Botanic Gardens. Humphrey was an only

child, and like most children of senior colonial officers he was soon shipped

back to Britain. At four years old he was ‘farmed out’ to the family of

an impoverished vicar in Yorkshire. At eight he was send to preparatory

school where a fellow pupil was one Tommy Disher, nephew to Richmal

Crompton and the model for ‘Just William’. At thirteen Humphrey went

to Repton, followed by Gonville and Caius College, Cambridge, where he

read Natural Sciences like his father did before him. Humphrey, by his own

admission, was more interested in shooting than science when at University

and represented both Cambridge and England at the sport. On graduation

in 1936, after flirting with the idea of becoming a policeman, he got a job

with Dunlop Malayan Estates. Thus he returned to South-East Asia where

he was to spend the bulk of his working life. As a rubber planter, Burkill

had to study plantation management including learning the languages

needed to organise the staff. Fluency in Malay and Tamil was expected of

planters and Burkill, apparently with a facility for languages, also picked

up Telugu as many of the workers on his first estate spoke that language.

Burkill’s adeptness with a rifle and experience with the Officer Training

Corps at university made it natural for him to join the Federated Malay

States Volunteer Force. The Japanese invasion of Malaya saw the immediate

mobilisation of the force and Burkill was part of the retreat to Singapore

where he was seconded to the Royal Engineers. Singapore soon fell and like

most captured servicemen he was imprisoned in Changi. After six months

he was sent, along with many others, to labour camps in Thailand. Burkill

reminisced about the activities of the wood-party he worked on during his

imprisonment in an article written for the newsletter of a prisoners of war

association. Their job was to collect the wood cut in the countryside around

the camp, mostly carrying the cut lengths of wood and stacking it in the

barges that transported it down river. Burkill’s linguistic abilities again came

in useful. The Thai he had learned before the war allowed communication

with the villagers and illicit trading when the guards were not looking. Skin

problems in later life doubtless stemmed from the exposure to the sun

during the forced labour.

After the war Burkill returned to work with Dunlops in Malaya; this

time with Joan Bloomer, his new wife. In 1948 Burkill joined the staff of the

Rubber Research Institute and took on the supervision of the experimental

rubber plantings throughout the Malay Peninsula. In 1954 Burkill was

offered the post of Assistant Director at Singapore Botanic Gardens by

then Director M.R. Henderson. To some extent this was a question of

Humphrey Morrison Burkill 135

returning a favour to the Burkills. I.H. Burkill had helped Henderson in

1923 by offering him a job in Singapore when his post in Kuala Lumpur

was discontinued. H.M. Burkill thought it likely a similar fate awaited

him at RRI due to funding uncertainties. After Henderson’s retirement

Burkill became Acting Director then in 1957 Director of the Gardens.

It had become clear that times were changing and the end of

the colonial era was in sight. Burkill was to oversee the change from a

largely expatriate British staff of administrators and researchers to one of

appropriately qualified locals. People with potential were identified and

sent for training. Chew Wee Lek and Chang Khiaw Lan completed Ph.D.s

under the supervision of former Assistant Director of the Botanic Gardens,

Prof. E.J.H. Corner in Cambridge. Burkill continued in the post of director

after Singapore became part of Malaysia and the subsequent expulsion

of Singapore from the federation and its own independence. He finally

retired in mid-1969. In those final years he tried hard to promote the idea of

continuing Singapore Botanic Gardens as a centre for scientific research as

well as a beautiful and historic garden. Burkill’s administrative burden left

him little time for research, and unlike his predecessors as Director such as

H.N. Ridley, his father or R.E. Holttum he published little original research

while working in Singapore. He had an interest in seaweeds and did go on

many collecting trips, particularly to the small islands around Singapore and

up the east coast of the Peninsula.

Burkill retired to England in late 1969. Looking for something to

do, he was offered work at the Royal Botanic Gardens Kew on a project to

revise Useful Plants of West Tropical Africa. This was written by J.M. Dalziel

and published in 1937. Despite all his tropical experience being in South-

East Asia, Burkill took up the job and for eight years was employed on what

proved to be an enormous task. This included a long trip to Nigeria in 1973.

1.H. Burkill is probably best known in botanical circles for his Dictionary

of the Economic Products of the Malay Peninsula and it must have seen

fitting to his son to also produce a magnum opus on economic botany.

The painstaking work compiling information from published material and

herbarium specimens continued long after Humphrey retired. Working as

an Honorary Research Fellow at Kew, he brought the six volumes covering

5300 species to publication between 1985 and 2004. A masterly work of

great scholarship, it will undoubtedly serve as the pre-eminent source of

information on West African economic botany for decades to come, much

as his father’s dictionary has done for South-East Asia. Humphrey Burkill

certainly followed the model of his predecessors as Director of Singapore

Botanic Gardens in having a long and very productive retirement.

136 Gard. Bull. Singapore 58 (2006)

Humphrey’s commitment to the Useful Plants of West Tropical Africa l\eft

no time for the study of his seaweed collections from his Singapore days.

They were deposited in the algal herbarium at the Natural History Museum

in London.

In relation to South-East Asian botany, H.M. Burkill will be

remembered for his long service (1957—1969) as Director of Singapore

Botanic Gardens and his efforts to maintain the scientific traditions of such

institutions through the transition from colony to independent state. Burkill

made the case for continuing government support for the Gardens and their

research functions at the highest levels. A newly independent nation had

many important items on its agenda for change and the Botanic Gardens

was not among them. Far-sightedly parks and street planting were given

priority in what was to become the ‘Garden City’ policy and in the long-

term the Botanic Gardens was to prosper as a flagship under that policy. But

in the short-term the Gardens became something of a backwater in the civil

service and resources were limited, potentially threatening the continued

existence of the herbarium and other research facilities. Fortunately the

Gardens did come through relatively intact, and Humphrey Burkill will

be remembered for his commitment to the concept of Botanic Gardens as

scientific institutions, particularly to the one in which he was born.

Mrs Joan Burkill died two years before her husband. Humphrey and

Joan are survived by their son and daughter, both of whom have followed in

the Burkill tradition of professional involvement in biology.

Publications

1958

The Botanic Gardens and conservation in Malaya. Gardens’ Bulletin

Singapore 17: 201-205.

1959

Large scale variety trials of Hevea brasiliensis Muell.-Arg. on Malayan

estates 1934-53. Journal of the Rubber Research Institute of Malaya 16: 1-

Dale

Humphrey Morrison Burkill 1S?

1967

Isaac Henry Burkill 1870-1965: a bibliography. Gardens’ Bulletin Singapore

22: 71-105.

H.B. Gilliland, 1911-1965, an appreciation. Gardens’ Bulletin Singapore 22:

107-112.

(with J.P.)

1968

Some studies of Malayan agarophytic and alginophytic seaweeds. Gardens’

Bulletin Singapore 22: 429-441. (with L.H. Greenwood-Barton and P.C.

Crowther)

1969

James Sinclair, 1913-1968: an obituary and an appreciation. Gardens’ Bul-

letin, Singapore 23: 1—xxiil.

1976

R.E. Holttum — in Singapore (1922-1954). Biological Journal of the Linnean

Society 8: 2-4.

LOTT.

Richard Eric Holttum, Croix de Guerre: a note. Gardens’ Bulletin, Singapore

S025: |

1983

Three new combinations in Spermacoce (Rubiaceae). Kew Bulletin 41:

1006.

Murray Ross Henderson, 1899-1983 and some notes on the administration

of botanical research in Malaya. Journal of the Malaysian Branch of the

Royal Asiatic Society 56: 87-104.

1985

The useful plants of West Tropical Africa. Volume 1. Families A—D. Royal

138 Gard. Bull. Singapore 58 (2006)

Botanic Gardens, Kew.

1991

The Singapore Botanic Gardens: a reputation to cherish and preserve. Pp.

11-15 in Proceedings of the Botanic Gardens 130 Anniversary Seminar.

Singapore Botanic Gardens, Singapore.

1994

The useful plants of West Tropical Africa. Volume 2. Families E-I. Royal

Botanic Gardens, Kew.

19%

The useful plants of West Tropical Africa. Volume 3. Families J—L. Royal

Botanic Gardens, Kew.

1997

The useful plants of West Tropical Africa. Volume 4. Families M-R. Royal

Botanic Gardens, Kew.

2000

The useful plants of West Tropical Africa. Volume 5. Families S—Z, addenda,

corrigenda, cryptogamata. Royal Botanic Gardens, Kew.

2004

The useful plants of West Tropical Africa. Volume 6. General index. Royal

Botanic Gardens, Kew.

The Burkills of Burkill Hall. Gardenwise 23: 16-19.

Names authored

Spermacoce intricans (Hepper) H.M. Burkill, Kew Bull. 41 (1986) 1006.

Spermacoce macrantha (Hepper) H.M. Burkill, Kew Bull. 41 (1986)

1006.

Humphrey Morrison Burkill 139

Spermacoce saxicola (K. Schum.) H.M. Burkill, Kew Bull. 41 (1986) 1006.

Eponymy

Scaphium burkillfilii Kosterm., Gard. Bull. Singapore 22 (1968) 44.

Bodo burkillii Skvortzov & Noda, Sci. Rep. Niigata Univ. ser. D 9 (1972) 28.

I.M. Turner

Affiliate Research Associate

Singapore Botanic Gardens

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VOL. 58 (Part 2)2007. | Pe oe cys Sc ee

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THE GARDENS’ BULLETIN SINGAPORE

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The Gardens' Bulletin

Singapore

VOL. 58 (Part 2) 2007 ISSN 0374-7859

CONTENTS

Boyce, P. C.

Studies on the Alocasia Schott (Araceae-Colocasieae) of Borneo: I. Two new species from Sarawak,

AIG WATEIT EONS) ennai th manta erie cet yer Re Nee, ce ea eee ree ee eS Oe ann re ee ne 141

Guo, S.-L., T. Cao, B. C. Tan and G.-Y. Song

Taxonomic notes on Asian species of Orthotrichaceae (Bryopsida): Macromitrium with

SV IMAOS LOM CNIS Ee ATISELIES I oar sensed ee nauenattte tooo ects S642, ae ORs BBs Besa de cba hee’ N veoskUatdCeIBoN, sadecenecbuaele 155

Kidyue, M., T. Boonkerd, O. Thaithong and T. Seelanan

Variations im the Hoya verticillata complexiin Thailand «....:.....:0000..<001-0vaesecsseasaccaspsenaraoesdunceees WS

Kiew, R. and J. Sang

Begonia (Begoniaceae) from limestone hills in the Kuching Division, Sarawak, Borneo, including

AMUN ME NVe SPC CICS 4s. d.dars Sees .eace 8. eee eee ence hoch beer =e Ah Sean a et Eni 199

‘Parris, B. S.

Five new genera and three new species of Grammitidaceae (Filicales) and the re-establishment of

EO 1 CLIATIIIUIS Pa rin coco eete Peccu nae Be sec erase. ono sock sR aN Sess eRe eabab ua sondoaaehoaweab Weed de vendeToaes pe ye,

Turner, I. M.

Two new names and two new combinations in Malesian Annonaceae ................ssccccsecceseceeeees 2I5

Wong, S. Y. and P. C. Boyce

Studies on Schismatoglottideae (Araceae) of Borneo II: Aridarum crassum, a new species from

Sarawak, Malaysian. Borneo. «..0..sceceeeio eae ee ee tee oer PA fe

Book Reviews

Etlingera of Borneo by A.D. Poulsen J. Leong-Skornickova ........................ 287

Spatulata Orchids — Papua New Guinea by J.W. Tkatchenko, BEM and S. Kami

HB. Kuareweil 2.4.44 tee 290

Date of publication: 20 June 2007

Published by

National Parks Board

Singapore Botanic Gardens

Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 58 (2) 2007: 141-154 141

Studies on the Alocasia Schott

(Araceae-Colocasieae) of Borneo: I

Two new species from Sarawak, Malaysian Borneo

PETER C. BOYCE

Malesiana Tropicals, Suite 9-04, Tun Jugah Tower,

No. 18, Jalan Tunku Abdul Rahman

93100 Kuching, Sarawak, Malaysia

Abstract

Two new species of Alocasia, A. chaii P.C.Boyce and A. infernalis P.C.Boyce from Kapit

Division, Sarawak, Malaysian Borneo, are described and included into an updated key to

Bornean Alocasia. Both species are illustrated.

Introduction

Publications on Alocasia for tropical Asia (Hay, 1994, 1998, 1999, 2000; Hay

& Wise, 1991; Hay et al., 1997; Yuzammi & Hay, 1998) have considerably

alleviated the previous problems of accurately naming A/locasia species

and, moreover, provided a stable platform from which to describe further

novelties without the concern that obfuscated species names are being

overlooked.

Prior to the onset of work on Bornean Alocasia by Hay and co-authors

(Hay, 1998, 2000, Hay et al., 1997), the species complement for Borneo stood

(uncritically) at 16 species. Post publication of Hay’s accounts for West

Malesia & Sulawesi (Hay, 1998) and the subsequent naming of an additional

species [A. nebula A.Hay (Hay, 2000) - treated as insufficiently known by

Hay, 1998] the total for Borneo stands at 21 species formally recognized.

Allowing for synonymization, this represents an increase of more than 45%

of the species diversity post 1997.

Since the publication accounts of Alocasiain Malesia and Sulawesi (Hay, 1998,

2000) the author of has been fortunate to be able to spend a considerable

period of time undertaking fieldwork in Sarawak, frequently in areas that

have received little or no botanical investigation in recent years, if ever. One

result of this fieldwork has been the discovery of a significant number of

142 Gard. Bull. Singapore 58 (2) 2007

additional novel Alocasia, such that it is estimated that the total number of

Bornean Alocasia species will eventually exceed 40, all endemic. Although

at the present time there is insufficient fertile material for the preparation

of types for all of the taxa considered to be undescribed; as the preparation

of types becomes possible from plants flowering in cultivation, it is intended

to produce a series of papers dealing with formal recognition of the ca 20

novelties found to date, along with updates on the status of pre-existing

described taxa. This paper is the first in this intended series.

1. Alocasia chaii P.C.Boyce, sp. nov.

Ab Alocasia scabriscula sed stature valde parviore, folii lamina peltato et spathae

fructiferorum extus in toto magenteus vividus differt -TY PUS: Sarawak, Kapit

Division, Pergunungan Hose, foothills of Bukit Batu, Camp E, 01° 51’ 33.6”;

114° 06’ 49.6”, 20 Oct. 2000, C_ Lee AL-41 (holo, SAR; iso, SING). Plates 1

& 2.

Small robust herb to ca 40 cm tall, stem shortly erect to decumbent, eventually

forming a short rhizome; wild plants with rather few leaves (up to 4, usually

less), cultivated plants with several leaves (up to 7); petioles stout, ca 20 cm

long, sheathing in the lower ca 1/4, puberulent, pale green with scattered

deep red spots and speckles in the lower half; sheath persistent, + closed;

lamina broadly ovato-elliptic, up to 40 cm long but frequently much less and

typically reaching c. 23 cm x 10-15, exceptionally up to 25 cm wide, thickly

coriaceous to subsucculent, almost completely peltate save for a shallow

retuse notch between the tips of the connate posterior lobes, margins

reflexed to form a raised smooth rim abaxially, apex acute to obtuse and

mucronate for ca | cm, lamina adaxially pale matt grey, abaxially greenish

white, anterior costa with 1-3 primary lateral veins on each side, diverging

at ca 90° (proximal ones) to 30° (distal ones); primary veins adaxially

somewhat raised proximally to the mid-rib and impressed distally, flush to

very slightly impressed with deep red axillary glands abaxially; secondary

venation impressed adaxially, more or less flush with the lamina abaxially,

forming defined interprimary collective veins when fresh, this decidedly

obscure in dried material; posterior lobes about 1/3 — 1/4 the length of the

anterior, with the posterior costae diverging at ca 30°; inflorescences several

together (up to 4 on vigorous plants), each subtended by a short, broad

prophyll and a single cataphyll; peduncle short, more or less hidden within

cataphyll; spathe ca 7 cm long, externally white with a scattered red flecks on

the lower part, interior uniformly glossy white; lower spathe 2.5—3.5 cm long,

ovoid, separated from limb by a rather weak oblique constriction; limb erect

even after anthesis, narrowly lanceolate-triangular, 3-5 cm long; spadix ca

Studies on the Alocasia Schott of Borneo

Plate. 1. Alocasia chaii P.C.Boyce. A. Adult plant in habitat; note the few leaves, typical of A.

chaii in the wild; B. Cultivated plant produced from tissue culture introduction; note the many

additional leaves and the deep red blands on the visible abaxial leaf surface. Photograph 1A

[copyright Chien Lee, used with permission].

144 Gard. Bull. Singapore 58 (2) 2007

2/3 the length of the spathe, ca 4.5 cm long, briefly stipitate; stipe cylindrical,

ca 2-5 mm tall, glossy white; female zone about 1/4 of the length of the

spadix, pistils moderately densely arranged; ovaries ovoid, ca 1.5 mm diam.,

facing diagonally up, pale greenish white; style + absent; stigma white, single,

bilobed, sometimes trilobed (all variations present in a single inflorescence);

sterile interstice absent or represented by a few (less than 5) synandrodia;

male zone held entirely within the lower spathe, cylindric to barrel-shaped,

about 1/4 the length of the spadix, about 2/5 as wide as long, ivory; synandria

densely arranged, more or less square in plan view, ca 1.5 mm wide, the

thecae very slightly overtopped by synconnective; appendix about 1/3 of

the length of the spadix, narrowly conic; fruiting spathe broadly ovoid, c. 2.5

cm long, erect, glossy brilliant magenta with a few scattered darker spots and

streaks at fruit maturation, then splitting longitudinally into several unequal

strips, these reflexing to reveal the ripe berries; berries bright orange to red,

globose, ca 0.5 cm diam., each with 1-3 seeds; seeds, ca 3mm diam., pace

brown.

Distribution: Sarawak, Kapit Division, to date known only from the foothills

of Gunung Bukit Batu, Hose Mountains and Ulu Kapit.

Ecology: Steep to precipitous leaf litter-covered red clay-loam slopes

beneath open to rather dense canopy of moist upper hill forest in light to

moderate shade, 540—760 m asl.

Notes: Alocasia chaii belongs to the informal Alocasia scabriscula group

(see Hay, 1998), notable for coriaceous, leathery to subsucculent leaves and

the spathe usually constricted at a level above the sterile interstice of the

spadix, thus, including the all or at least the majority of male flower zone

within the lower spathe. Alocasia chaii is most similar to A. scabriscula

N.E.Br. in overall morphology, differing in the considerably smaller, but

hardly less robust habit, the peltate leaves and the persistent lower spathe

that turns bright magenta at fruiting. In general stature and by the grey

leaves A. chaii also vaguely resembles Sabahan A. melo A.Hay, P.C.Boyce

& K.M.Wong, although the latter is readily distinguished by the rugose and

bullate adaxial lamina surface and the fruiting spathe white with slight red

speckling. Alocasia melo is confined to ultramafic substrates.

There appear to be two closely allied species involved here, one in the

lowlands (up to 150 m asl) that has not as yet found fertile, and a higher

elevation element (occurring above 500 m asl), here described as A. chaii.

Aside from the altitudinal differences noted, the lowland element has the

leaves proportionately longer than broad (ovato-triangular in outline), lacks

the deep red abaxial leaf glands (glands concolorous with the abaxial lamina

Studies on the Alocasia Schott of Borneo 145

Plate. 2. Alocasia chaii P.C.Boyce. A. Ripe infructescences; note the bight magenta colour of the

lower spathe and the contrasting orange fruits.

surface in the lowland element), and is overall a less robust plant occurring

in open habitats. The occurrence of related attitudinally differentiated/

morphologically distinct taxa has been noted elsewhere in Alocasia, as for

example, A. beccarii Engl. (lowland) & A. peltata M.Hotta (highland).

Etymology: Alocasia chaii is named for Dr Paul P.K. Chai former Forest

Botanist, now with ITTO, Forest Department, Sarawak.

Other specimens examined: SARAWAK: Kapit Division: Pergunungan

Hose, foothills below Bukit Batu, 02° 14’ 47.2”; 113° 41’ 24.9”, 23 April 2004,

PC.Boyce & Jeland ak Kisai AL-51 (SAR); Ulu Kapit, Sungai Nai, near

Punan Bah, 23 Sept. 1973, P- Chai et al. $.33339 (SAR); Pergunungan Hose,

Ulu Sungai Temiai, 5 July 2003, C. Lee et al., S87433 (SAR).

2. Alocasia infernalis P.C.Boyce, sp. nov.

Ab alli Alocasii borneensibus stature parviore, foliis ascendentis, folii lamina

atropurpureus vel purpureonigris, nitentibus distinguitur —TYPUS: Sarawak,

Kapit Division, Nanga Gaat, Rejang Wood Concession, Batang Baleh, 01° 38’,

113° 09’, 2 April 1998, C.Lee AL-16 (holo, SAR; iso, SING,). Plates 3 & 4.

146 Gard. Bull. Singapore 58 (2) 2007

Small robust herb to ca 55 cm tall, stem slender, erect to ultimately decumbent

with the active shoot tip ascending; leaves several together in nature, in cultiva-

tion up to 12, spreading in juveniles but erect in adult plants; petioles slender,

spreading to ascending, ca 20 cm long, sheathing in the lower ca '2, minutely

puberulent (lens required), bronze-green to purple-green depending on exposure,

stronger light inducing a weak snake-skin marking, particularly on the interior

of the sheath; sheath fleshy-membranous, open and recurving in the lower part;

lamina ovato-triangular, up to 25 cm long but frequently much less and typically

reaching ca 15 cm x 10—12 cm, thinly and somewhat weakly coriaceous, juve-

niles almost completely peltate except for a shallow retuse notch between the tips

of the connate posterior lobes, adult leaves strongly peltate but with a 1—2 cm

deep notch in the sinal tissue, margins smooth, apex acute, acuminate for ca 1 cm,

adaxially glossy, very deep purple, abaxially deep purple, anterior costa with ca 3

primary lateral veins on each side, diverging at ca 60° (proximal ones) to 45° (dis-

tal ones); primary veins impressed distally adaxially, prominently raised abaxi-

ally; secondary venation obscure adaxially, abaxially forming strongly defined

and raised interprimary collective veins; all veins running to a prominently raised

(abaxially) inframarginal collecting vein; posterior lobes about 4% the length

of the anterior, posterior costae diverging at ca. 20°. Inflorescences 2

together, each subtended by a short, broad prophyll and a single cataphyll;

peduncle long, ca 4—6 cm, pale green or purple flushed; spathe 4—9.5 cm

long, lower spathe pale green, spathe limb externally glossy purple with the

margins pale green; lower spathe 1.5—2.5 cm long, ovoid, separated from

limb by a moderate constriction; limb narrowly lanceolate-triangular, at first

erect then soon strongly reflexing and twisting with the margins inrolled,

2—6.5 cm long; spadix ca 1/2 the length of the spathe, ca 4.5 cm long, very

briefly stipitate; stipe umbonate, ca 2 mm tall, glossy white; female zone ca

1/3 of the length of the spadix, pistils moderately densely arranged; ovaries

compressed-globose, ca 2 mm diam., facing diagonally up, pale greenish

white; style absent; stigma white, mostly trilobed, sterile interstice with a few

scattered, compressed white synandrodia; male zone partially held within

the lower spathe, cylindric, ca 1/3 the length of the spadix, ivory; synandria

somewhat laxly arranged, transversely oblong in plan view, ca 2 x 1 mm,

thecae extending slightly from the edge of the synconnective; appendix

about 1/3 of the length of the spadix, narrowly conic, pointed, white; fruiting

spathe broadly ovoid, ca 2.5 cm long, pendent by reflexing of the peduncle, dull

mid-green at fruit maturation, splitting longitudinally into several unequal

strips, these reflexing to reveal the ripe berries; berries bright orange to red,

globose, ca 0.5 cm diam., each with 1—3 seeds; seeds compressed ovoid, ca 2.5

diam., medium brown.

Studies on the Alocasia Schott of Borneo

Plate. 3. Alocasia infernalis P.C.Boyce. A. Juvenile plant in habitat; B. Seedlings, Nanga Gaat; note

the iridescent leaf surface and also the variability in the intensity of the purple colouration.

148 Gard. Bull. Singapore 58 (2) 2007

Plate 4. Alocasia infernalis P.C.Boyce. A. Plant flowering in cultivation; B. Deep purple-

black leaves of a plant in cultivation produced from tissue culture; C. Flowering size plant in

cultivation..

Studies on the Alocasia Schott of Borneo 149

Distribution: Sarawak, Kapit Division. To date known only from the Sungai

Gaat watershed.

Ecology: Valley bottoms in moist to ever-wet lowland forest on deeply leaf

litter-covered red sandstone-derived clay-loams in heavy shade,182—249m

asl.

Notes: Alocasia infernalis belongs to the Alocasia scabriscula group (see

Hay, 1998) by virtue of the pubescent petioles and the positioning of the

spathe constriction above the base of the male zone of the spadix such

that all or at least the basal part of the male zone is held within the lower

spathe chamber. However, it is not at all apparent to which other species

in the group A. infernalis is most closely allied since by the leaf texture and

deflexing infructescences it is unique in the group. In overall appearance

(leaves rather membranous more-or-less completely peltate, spadix distinctly

shorter than the spathe) it is superficially similar to species in the Alocasia

cuprea (C.Koch & Bouché) C.Koch group but is readily distinguished by the

leaves not interspersed with cataphylls.

The metallic-purple leaves of the seedlings and juvenile plants is remarkable

while the lustrous deep purple-black of the ascending leaves of mature

plants, coupled with the dwarf habit, is unmatched by any other species.

Alocasia infernalis is perhaps the most horticulturally significant species of

Bornean Alocasia yet discovered.

Etymology: From the Latin, ‘Hellish’ in fanciful allusion to the remarkable

deep purple-black leaves of mature plants; the epithet is inspired by the no-

less remarkable vampyromorphoid cephalopod, Vampyroteuthis infernalis.

Other specimens examined: SARAWAK: Kapit Division: Nanga Gaat,

Rejang Wood Concession, km 65 road to Camp Gahada, 01° 42’ 01.1”, 113°

31’ 14.8”, 12 May 2004, PC Boyce, Jeland ak Kisai & Jipom ak Tisai AL-

57 (SAR); Nanga Gaat, Rejang Wood Concession, km 55 road to Camp

Gahada, 01° 44’ 44.5”, 113° 28’ 32.3”, 13 May 2004, PC. Boyce, Jeland ak Kisai

& Jipom ak Tisai AL-66 (SAR); Nanga Gaat, Rejang Wood Concession, km

65 road to Camp Gahada, 01° 41’ 59.7”, 113° 31’ 13.7”, oblong leaves, 16 Dec

2004, P.-C. Boyce, Jeland ak Kisai & M.Gibernau AL-123 (SAR).

Conservation

Both species here newly described occur as scattered small populations in

restricted habitats. Fortunately all known populations of both species are

150 Gard. Bull. Singapore 58 (2) 2007

in remote and inaccessible locations and for the moment probably safe

from the deprivations of unscrupulous plant collectors. Alocasia chaii and

A. infernalis were among those species the subject of a joint tissue culture

project between Malesiana Tropicals and UNIMAS funded under MOSTI-

IGS (IGS R&D Proj. No. 16/03), together with a further 14 Sarawakian

Alocasia species and representatives of several other aroid genera that are

now in Sarawak Forestry Department licensed commercial tissue culture

production in Kuching laboratory of Malesiana Tropicals Sdn Bhd.

Key to Bornean Alocasia species

1: Leaf blades not peltate im adult plamts*¢.22.22.2..., 2.232, eee 2

la. Leaf blades distinctly (shallowly to completely) peltate in adult plants

wisbapsneieend8b Bac eninwuld ox sue nadell GeO apes a aetOB oe Mocs URE pee eee 14

2. Secondary venation distinctly prominent abaxially and forming well-

defined interprimary collective veins... A. sarawakensis

2a. Secondary venation not prominent abaxially, or, if prominent, then not

forming well-defined interprimary collective VeINS .........ceeeeeseeseeeeteeeteee es 3

3. Leaf blade membranous, often immense, abaxially waxy-glaucous ..........

dE hbSh EROS AR. Datacard eae A. robusta

3a. Leaf blade of various sizes and textures, not waxy-glaucous (though

sometimes abaxially erey-oneen).. 0222... 20041 eee 4

4.Male zone of spadix completely exerted from lower spathe chamber (always

in association with human disturbance) ................... A. macrorrhizos

4a. Male zone of spadix partly or wholly within lower spathe chamber (plants

of natural forested habitats)i. ac.c.s.0. sce ee 5

5. Leaf blade narrowly to broadly ovato-sagittate, nearly always stiffly

leathery to-subsucculemt (2.33.8 iooe. hatsiseees eee . 6

Sa. Leaf blade hastato-sagittate, triangular in outline, mostly rather fiialy

Fr: 18 1 | ees teeter mee Mier rye AUS eo ascot cncyous 90s 2¢3: . 10

6. Adaxial leaf blade grey-green and distinctly dark green about main veins

racnssitisdeddsions vs sonewscutteaea Geeheedioncete-ecuetiniten site eee ee a ae |

6a. Admaal ieat blade of various colours but not variegated .................. 8

7. Abaxial leaf blade purple; anterior costa with c. 6 primary lateral veins on

each side, with conspicuous subsidiary veins (geology & origin unknown)....

Studies on the Alocasia Schott of Borneo 15h

Pree rr er ee Peete eee toad cen aanidaveal basddcaaseacenroniesncddsends A. nebula

7a. Abaxial leaf blade not purple; anterior costa with 2--3 primary lateral

veins on each side; subsidiary veins absent (limestones: SE Sarawak) ............

_beadyic pages ade He Lace ALE occa dsc ot TERA Ee OO ee A. reversa

8. Inflorescence pairs solitary and secondary venation adaxially impressed

(climates thera SAIVAUIIS) eee cee Oe RE IAD, 31S DRLEAT. aah dha A. reginae

8a. Inflorescence pairs clustered or if solitary then secondary venation not

RTM acs SS IEE et ae Sumer were SIT A SE IIS SSH Jlehad la cadiad a scosauoctees hbeacsbdbe 9

9. Posterior lobes ca 1/2 or more the length of the anterior; blade stiffly

leathery (lithophytic on or terrestrial in close association with limestone

(limestones: SE Sarawak) ........ hots .. A. ridleyi

9a. Posterior lobes less than ioe ca 1/2 ne eae oi ae neta blade thick-

ly coriaceous to subsucculent; terrestrial and not especially associated with

limestone (widespread in Borneo) ......eeeeeeeeeees As Scabriuscula

lO; Plants terrestrial; not lamestone associated .22..26ic.ckccccsslekeleenee vie ecdveeeees 11

Mamrlants lithoplnytic:on limestone. . 20.4. .00.6.05 irae edaste bases AZ

11.Petiolesmottled with wavy oblique zones of dense brown lines, occasionally

scabrid; spathes mostly dusky brownish mauve, the limb darker; lower spathe

narrowly ovoid; limb mostly narrowly lanceolate (Sabah) ....... A. wongii

lla. Petioles variously and more or less haphazardly marked with lines

and/or dots, smooth or occasionally faintly bumpy (glands), but not scabrid;

spathes mostly ivory to yellowish ivory, variously marked or not with

pink to purple, and/or purple-margined; lower spathe broadly ovoid; limb

more or less oblong (widespread in Borneo) ...........eeseeeseeteeees A. princeps

12. Male zone of spadix completely within lower spathe chamber; leaf blades

distinctly grey-green adaxially (limestones: Sabah, E. Kalimantan).................

~useeQSuce dale) abandon haere, mid Aare oie Ot Mick seek gee eee oe A. principiculus

12a. Male zone of spadix partly exserted from lower spathe chamber; leaf

blades dark to bright ereen adaxdatly (Sabal) 2.2. 20.0.....ctsocctecceeeeoeodhessesase 13

13. Leaf blade bright green adaxially; inner side of posterior lobe ovate; male

zone more or less adjunct to female zone or interstice short, not attenuate,

formed of 1-2 whorls of synandrodia, resembling synandria (limestones:

TUNE” hh, [CIT EEA Alene: Se «Lin vate AEA LAG. eee ee ee A, puteri

13a. Leaf blade dark green adaxially; inner side of posterior lobe elliptic

to narrowly ovate; interstice elongate, partly naked, with neuter organs

resembling staminodes below and resembling synandria above (limestones:

152 Gard. Bull. Singapore 58 (2) 2007

Gua Madai):......:.23. 26 Bee eee A. pangeran

14. Leaf blades + membranous and pendent, often solitary or only 2--3

together, often adaxially dark green with whitish major veins (sometimes

adaxially concolorous), often purple-backed, shallowly to deeply peltate;

stigma stellate with pointed lobes; interstice corresponding with spathe

constriction and male zone completely exetied Gu eee

Jin OR See eel ee A. longiloba complex (see Hay, 1998)

14a. Leaf blades variously coriaceous, pendent or not, few to several together,

deeply to almost completely peltate; stigma not lobed or lobes rounded;

interstice and part or all of male zone within lower spathe cham ............ 15

15. Leaf blades metallic greenish brown adaxially, bullate between primary

veins; lower primary veins diverging at first at more than 90° (Sabah & NE

Sama wi) ia v'epciveis.. isdn ad As es ede a

15a. Not be cnn v3 « itidacdinsa inn’ iO MM SIDE ee 16

16. Adaxial leaf surface strongly and minutely rugose with the tertiary venation

raised (ultramafics: Sabah) ................ neler

16a. Adaxial leaf surface sco or Sh eee ae ieee .17

17. Adaxial leaf laminae very dark black-green with white impressed primary

and secondary venation; spadix with isn reduced (limestone: Bukit

Tabin) . ret detdenaattvsei eee, (ALC epmmia

17a. “heal leaf feta an “alegei dh or a wariegated Shes main veins and

neighbouring blade darker than the rest; appendix well developed ............. 18

18. Leaf blades with conspicuous intramarginal vein and marginal vein; laminae

broadly to narrowly elliptic, with the base cuneate; male zone wholly within

the lower spathe: (above S00meBomeo)...ai sie eee A. peltata

18a. Leaf blades with more or less conspicuous marginal vein only; laminae

various; male zone wholly or partly within the lower spathe (mostly below 500

m; Borneo) eget, a eae ee ee eee ee 19

19. Primary lateral veins numerous, 8-10 on each side of midrib; secondary

venation striate; in peat swamp forest (peat swamp forest: Sarawak) ............0

snd Soba tnb beds ABTS AAENVE He TTS Lk AS: RAN a SAT ed nee A. minuscula

19a. Primary lateral veins much fewer;secondary venation clearly colocasioid,

but not forming interprimary collective: veimsi.2.:2.23 ee oi eee 20

20. Leaf blades thickly coriaceous to subsucculent; male zone of spadix

within lower spathe:chamber 2.5:1:):2.291..We a i ee 21

Studies on the Alocasia Schott of Borneo 153

20a. Leaf blades thinly coriaceous or sub-membranous; male zone only

partly mcluded within the lower spathe:chamber ....,..............0csescressscseasess 22

21. Leaf laminae broadly ovato-elliptic, adaxially pale matt grey, abaxially

greenish white with conspicuous deep red axillary glands abaxially; petioles

puberulent; fruiting spathe magenta (Kapit, evergreen upper hill forest on

SAMOS TOMES ADOVE | IW Urmle asl) eee eae ee led.y...Vs2505- oko. G sa leastadasteboeviedcs de A. chaii

21a. Leaf blade narrowly elliptic to ovate to narrowly obovate, mid green

above, slightly paler with inconspicuous pale green axillary glands; petioles

glabrous fruiting spathe pale green (NW Borneo, kerangas below 500 m).........

ee NR te tetra seca cits ip essai ov tule tay + Gctosesvedobubdensinenss A. beccarii

22. Laminae ascending (adult plants) to weakly spreading (juveniles),

thinly and weakly coriaceous, lustrous deep purple-black; petioles minutely

puberulent; infructescences deflexed (Kapit: sandstones) .......... A. infernalis

22a. Laminae pendent to weakly spreading (adult & juvenile plants), thinly,

stiffly coriaceous; never deep purple-black; petioles glabrous; infructescences

SSUES 7070S eee ch Lae a eee ee Re ore oe ents ao? aes ee 23

23. Leaf blades dark green throughout and somewhat darker around mid-

veins; inflorescences to ca 6 together; stigma mostly tri-lobed (limestones: Gua

NEEL) cath telco baw Bl ty, cel Alps nee de inet be ip eto eee Re A. venusta

23a. Leaf blades grey-green and dark blue-green around veins; inflorescences

solitary to paired; stigma mostly bi-lobed (limestones: SE Sarawak) ................

Ne Me ee a he Ae eat RE ear ae ieuate bas acesidustesteuaetteeseorsdses A. reversa

Acknowledgements

I wish to express my 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. Especial

thanks 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.

154 Gard. Bull. Singapore 58 (2) 2007

References

Hay, A. 1994. Alocasia simonsiana - a new species of Araceae from New

Guinea. Blumea 38: 331-333.

Hay, A. 1998. The genus Alocasia (Araceae-Colocasieae) in West Malesia

and Sulawesi. Gardens’ Bull. Singapore 50: 221-334.

Hay, A. 1999. The genus Alocasia (Araceae-Colocasieae) in the Philippines.

Gardens’ Bull. Singapore 51: 1-41.

Hay, A. 2000. Alocasia nebula. Bot. Mag., n.s. 17: 14-18, pl. 381.

Hay, A., P.C. Boyce & K.M. Wong 1997. Alocasia melo. Bot. Mag., n.s. 14(2):

82-86, pl. 315.

Hay, A. & R. Wise. 1991. The genus Alocasia (Araceae) in Australasia.

Blumea 35: 499-545.

Yuzammi & A. Hay 1998. Alocasia suhirmaniana (Araceae-Colocasieae): a

spectacular new aroid from Sulawesi, Indonesia. Telopea 7: 303-306.

Gardens’ Bulletin Singapore 58 (2) 2007: 155—178 Jbes)

Taxonomic notes on Asian species of Orthotrichaceae

(Bryopsida): Macromitrium |

with gymnostomous capsules

SHUI-LIANG GUO , TONG CAO |, BENITO C.TAN :

and GUO-YUAN SONG

College of Life and Environmental Sciences, Shanghai Normal University,

; Shanghai 200234, China

Department of Biological Sciences, National University of Singapore, Science

Drive 4. Singapore 117543

Abstract

Nine taxa of the genus Macromitrium (Bryopsida, Orthotrichaceae)

with gymnostomuous capsules in East Asia, including Macromitrium

benguetense R.S. Williams, M. brevituberculatum Dix., M. formosae Card.,

M. gymnostomum Sull. & Lesq., M. heterodictyon Dix., M. holomitrioides

Nog., M. robinsonii R.S. Williams, M. schmidii C. Muell., and M. taiheizanense

Nog., are taxonomically revised. Macromitrium robinsonii and M.

brevituberculatum are treated as new synonyms of M. gymnostomum,

and M. benguetense as a new synonym of M. schmidii. Neotype was

designated for M. schmidii, and types were selected for M. heterodictyon, M.

gymnostomum, M. brevituberculatum, M. taiheizanense, and M. formosae.

Macromitrium schmidii var. macroperichaetialium S.L. Guo & T. Cao was

described as a new variety. A key to the seven accepted gymnostomous

species of Macromitrium in East Asia 1s also given.

Introduction

Macromitriumis arather large pantropical genus of mesophytic to xerophytic

epiphytes, belonging to Orthotrichaceae, Bryopsida. It is distinguished

by the creeping, prostrate stems giving rise to branches terminated by a

sporophyte, the often contorted leaves twisted around the branches, and

with mostly bulging, conic, papillose, or tuberculate leaf cells. Calyptrae are

large, mitrate, plicate, naked or hairy. The ovate to cylindric capsules mostly

have peristomes, which are at times reduced and fused to a low, single

exostomial membrane.

156 Gard. Bull. Singapore 58 (2) 2007

Though Asia is rich in the species of Macromitrium, with about 85-90 species

recorded (Crosby et al., 1999; Eddy, 1996; Gangulee, 1976; Noguchi, 1967,

1972, 1989; O’Shea, 2002; Redfearn et al., 1996; Tan and Iwatsuki, 1991, 1993),

little taxonomic revision has been done except for those in Japan (Noguchi,

1989), Malaysia, Philippine and Indonesia (Eddy, 1996), Eastern India and

its adjacent regions (Gangulee, 1976), and Papua New Guinea (Vitt, 1995).

There is a rather special group of Macromitrium with gymnostomous

capsules, including M. benguetense R.S. Williams, M. brevituberculatum Dix.,

M. formosae Card., M. gymnostomum Sull. & Lesq., M. heterodictyon Dix.,

M. holomitrioides Nog., M. robinsonii R.S. Williams, M. schmidii C. Muell.,

and M. taiheizanense Nog. Since these species were published, there has

been little taxonomic information about them, except for M. gymnostomum

and M. holomitrioides (Crosby et al., 1999).

During our taxonomic revision on Asian Macromitrium, we found a

specimen with gymnostomous capsules marked as “China, Kwangtung:

Ngok Shing Shan, Y. M. Taam 402C ” in NY, which is rather similar to

M. schmidii Muell. A further check on this specimen showed that it

differs from the later by its rather long inner perichaetial leaves and its

gymnostomous capsule that are constricted below the mouth into 4-angles

or 4-furrows when dry. This specimen represents a hitherto undescribed

taxon, here we placed it at the variety level and named it as Macromitrium

schmidii var. macroperichaetialium. Our further study of the types showed

that M. brevituberculatum and M. robinsonii are two new synonyms of M.

gymnostomum, and M. benguetense is a new synonym of M. schmidii. In

order to understand these gymnostomous species better and to stabilize the

usage of the species epithet, a neotype was designated for M. schmidii C.

Muell. and types were selected for M. heterodictyon Dix., M. gymnostomum,

M. brevituberculatum, M. taiheizanense and M. formosae, and a key to these

gymnostomous species was also given.

Taxonomic treatment

Macromitrium schmidii C. Muell., Bot. Zeit. 11: 61. 1853. Figs. 1-3.

Type: India, Nilgherri, leg. Perrottet no. 22 (neotype, H-BR 2595002!,

designated here) — Macromitrium benguetense R.S. Williams, Bull. New

York Bot. Gard. 8: 343, 1914. syn. nov. Type: “Philippine, Baguio, on tree,

1570 meters, Oct. 1904, Williams 830 (isotype, H-BR 2572023!).

Taxonomic notes on Asian species of Orthotrichaceae 157

Plants moderate-sized, forming yellowish-brown mats, darkish below. Stems

creeping, up to 10 cm long, with erect branches, branches 10—20 mm high and

1.5-2 mm wide. Branch leaves moderately crisped and somewhat twisted

when dry, widely spreading when moist, entire, lanceolate, (2.5) 3.5-4 mm

long, 0.5—0.7 mm wide, the apex acuminate, somewhat incurved, keeled,

plicate below; margins entire, plane on one side, reflexed-recurved on the

other side, particularly in lower portion; costa single, prominent, ending a

few cells below the apex or in the apex; laminal cells unistratose from top to

base, upper laminal cells subquadrate-rounded to rounded-quadrate, 10-14

yum wide, obscure, densely pluri-papillose; median laminal cells rounded-

quadrate or slightly elongate, 10—15 um wide, 15-20 um long, with a single,

large conical papilla or 3—4 small papillae per cell; basal laminal cells brown-

yellowish, rectangular to sublinear, 30-50 um long, 6—10 ym wide, strongly

thick-walled and sinuous, tuberculate to bulging-papillose.

Sporophytes lateral on branches; inner perichaetial leaves ovate-oblong,

long-acuminate in upper part, 2.8—3.2 mm long, shorter than branch leaves,

the costa ending far below the apex. Setae 4—6 mm, smooth, twisted to right.

Capsules oblong-cylindric or oblong, 1—2 mm long, 0.6-0.8 mm wide, brown,

plicate or ribbed under the mouth when dry, peristome absent. Spores 25-35

uum in diameter. Calyptrae moderate sized, 2—2.5 mm long, campanulate,

with brownish hairs.

Fig. 1. Neotype and habit of M. schmidii: “Nilgherri, Leg. Perrottet 22” at H-BR.

158 Gard. Bull. Singapore 58 (2) 2007

OOOG OG00-

Fig. 2. M. schmidii C. Muell. a-e. Branch leaves; f-g. Perichaetial leaves; h. Transverse section

of the meddle part of leaf; i. Transverse section of the basal part of leaf; }. Transverse section

of the upper part of leaf; k, m. Basal laminal cells; 1. lower laminal cells; n. Middle laminal

cells; o. Upper laminal cells p—q. Capsules; r. Spores. All from the lectotype of M. schmidii at

H-BR. Line scales: A = 0.5 mm (a-g, p-q); B = 50 um (h-o, r).

Taxonomic notes on Asian species of Orthotrichaceae 159

3 : : d e f

Fig. 3. M. schmidii C. Muell. a-d. Branch leaves; e-f. Stem leaves; g. Calyptra; h. Transverse

section of the meddle part of leaf; i. Transverse section of the basal part of leaf; }. Upper

laminal cells of branch leaf; 1. Middle laminal cells of branch leaf; k. Basal laminal cells of

branch leaf; m. Upper laminal cells of stem leaf; n. lower laminal cells of stem leaf; 0. Dry

capsule; p. Wet capsule; q. Spores. All from isotype of M. benguetense at H-BR. Line scales: A

= 0.5 mm (a-d, f, o,p); B = 50 um (g-m, q); C = 1 mm (g).

160 Gard. Bull. Singapore 58 (2) 2007

There has been no more taxonomic information of M. schmidii after it was

published (Crosby et al., 1999), and no citation about the type of the species

could be found in the original publication by Mueller (1853). According to

the title of the publication and context, we surmised that Mueller’s original

description on M. schmidii was based on the collection by Perrottet from

Neigherri, South India. There were two specimens collected by Perrottet

from Neigherri in H-BR, identified as M. schmidii by Mueller himself. We

selected the specimen marked with “Neigherri, Perrottet no. 22” as the

neotype (Fig. 1).

M. benguetense was described by R.S. Williams from Baguio of the Philippine

in 1914. The type of M. benguetense and M. schmidii are rather similar except

that the capsules of M. benguetense are broadly oval with small mouths,

and its calyptrae are sparsely hairy, while the capsules of M. schmidii are

oblong cylindric, and its calyptrae are densely hairy. Here we treated M.

benguetense as a new synonym of M. schmidii.

Based on the type and other specimens of M. schmidii, the diagnostic features

of M. schmidii are: 1) moderate-sized plants with lanceolate branch leaves;

2) upper laminal cells obscure, densely pluri-papillose; median laminal cells

rounded-quadrate or subrectangular, bulging or with a single, large conical

papilla, or 3-4 low papillae per cell; 3) inner perichaetial leaves ovate-oblong,

long-acuminate in upper part, shorter than branch leaves; 4) the laminal

cells of the inner perichaetial leaves elongate to linear-elongate from base

to top, sinuous, thick-walled, with a single conical papilla; 5) capsules oblong-

cylindric or oblong, peristome none, plicate under the mouth.

Distribution: India, Sri Lanka, Philippine, Vietnam (Tan & Iwatsuki, 1993).

Other specimens examined: INDIA: Nilgherri, Perrottet (H-BR 2595001);

Madras, Madura, G.. Foreau 1925 (H); Nilghiri, Suthi B. 7331 (H-BR

2595003). SRI LANKA: Wichura 2744 (H-BR 2595004); Mire Bamforih (H-

BR 2595006); ibid, C. Mueller 41 (H-BR 2595005); Kalak, 77W.N. Beckett c

29/2.( H-BR 2595007).

Macromitrium schmidii var. macroperichaetialium S.L. Guo & Cao T,, var.

nov. (Figs. 4-6)

Macromitrium schmidii Muell. satis similes, sed differt foliis perichaetialibus

ad 5.5 mm longis, excedenibus foliis vegetatiis. Type: China, Kwangtung:

Ngok Shing Shan, Sai-lin-shan Village, Sin-fung district; thicket on steep

slope. 23-31 March 1938, Y. M. Taam 402C (holotype, NY!).

Taxonomic notes on Asian species of Orthotrichaceae 161

Nae

Or~S0

oO O

O a

Te) ese

O O O

Oo Oo O

oy4 OP(O

O ae

_ 0600°

0006 CER”

O,0 ma Olals

Eleat

alc! a

eens

Fig. 4 Macromitrium schmidii var. macroperichaetialium. a. Perichaetial leaf; b, o. upper part

of perichaetial leaf, c-f. branch leaves; g-j. dry capsules; k. calyptra; 1. basal laminal cells of

branch leaf; m. middle laminal cells of branch leaf; n. upper laminal cells of branch leaf; p.

transverse section of the middle part of branch leaf; q. transverse section of the basal part of

branch leaf; r,s. transverse section of the upper part of branch leaf; t, y. transverse section of

the upper part of perichaetial leaf; u. upper laminal cells of perichaetial leaf; v. middle laminal

cells of perichaetial leaf; w. basal laminal cells of perichaetial leaf; x. transverse section of the

middle part of perichaetial leaf. All from holotype at NY. Line scales: A = 0.5 mm (a, c-f); B

= 50 um (I-n, p-y); C = 2 mm (k); D= 1 mm (b, 0); E= 0.5 mm (g-j).

162 Gard. Bull. Singapore 58 (2) 2007

HERBARIUM OF

THE NEW YORK BOTANICAL GARDEN

mOgses or CHINA

COLLECTED ay Y

DETERMINED By

Fig. 5. Holotype and habit of M. schmidii var. macroperichaetialium in NY.

Perichaetial leaves =f

Fig. 6. M. schmidii var. macroperichaetialium showing the perichaetial leaves and the

gymnostomous capsules constricted below the mouth in a 4-angled or 4 -furrowed shape.

Plants moderate-sized, forming yellowish-brown mats, darkish below. Stems

creeping, up to 10 cm long, with erect branches, branches 10-15 mm high

and 1—2 mm wide. Branch leaves moderately crisped and somewhat twisted

when dry, widely spreading when moist, entire, lanceolate, (2.5) 2.7—3.5 mm

long, 0.4—0.6 mm wide, the apex acuminate, somewhat incurved, keeled,

slightly plicate below; margins entire, plane on one side, reflexed-recurved

on the other, particularly in lower portion; costa single, prominent, ending

below the apex or in the apex; laminal cells unistratose from top to base;

upper laminal cells subquadrate-rounded to rounded-quadrate, 9-12 um

wide, obscure, pluri-papillose; median laminal cells rounded-quadrate or

slightly elongate, thick-walled, 10—11 um wide, 13-14 um long, with a single,

large conical papilla or pluripapillae; basal laminal cells rectangular to

sublinear, 22—30 um long, 6-8 um wide, strongly thick-walled and sinuous,

often with a single large papilla per cell.

Taxonomic notes on Asian species of Orthotrichaceae 163

Sporophytes lateral on branches. Inner perichaetial leaves lanceolate,

gradually long-acuminate in upper part, up to 5.5 mm long, much longer

than branch leaves; costa long excurrent, slightly coarse in the upper part;

the laminal cells of the perichaetial leaves elongate to linear elongate from

base to top, sinuous, thick-walled, with a single conical papilla per cell;

Paraphyses numerous. Setae (4) 6—7 mm, smooth, twisted to right. Capsules

oblong-cylindric or oblong, 1.3—1.5 mm long, 0.5—0.65 mm wide, brown,

constricted below the mouth in 4-angled or 4-furrowed shape, peristome

absent. Spores not seen. Calyptrae large, cucullate, 2.0—2.5 mm long, with

many long, flexuose or stiff, brown-yellowish hairs.

The specimen marked as “China, Kwangtung: Ngok Shing Shan, Y. M.

Taam 402C (NY) ” shares the following characteristics with the neotype

of M. schmidii at H-BR: 1) moderate-sized plants, with lanceolate branch

leaves; 2) upper and median laminal cells subquadrate-rounded to

rounded-quadrate, obscure, pluripapillose; basal laminal cells rectangular

to sublinear, strongly thick-walled and sinuous, often with a single

large papilla per cell; 3) the laminal cells of the inner perichaetial leaves

elongate to linear-elongate from base to top, sinuous, thick-walled, with

a single conical papillae; 4) gymnostomous capsules. However, the inner

perichaetial leaves of the specimen (Taam 402C, NY) are long lanceolate,

gradually long-acuminate in upper part, much longer than branch leaves,

its capsules distinctly constricted below the mouth in a 4-angled or 4-

furrowed shape, while the inner perichaetial leaves of M. schmidii are

similar to its branch leaves, and its capsule only plicate or ribbed without

a constriction under the mouth in a 4-angled or 4-furrowed shape. We

consider that the specimen represents a hitherto undescribed taxon, here

we placed it at variety level and named it as Macromitrium schmidii

var. macroperichaetialium based on its longer inner perichaetial leaves.

Distribution: China.

Macromitrium heterodictyon Dix., Hong Kong Naturalist, Suppl. 2: 12. 6.

1933. (Fig. 7). Type: China, Hong Kong, granite rock, 100-200 ft. alt., Amoy

I, 11 Jul 1931, coll. Herklots (B.17 E) (holotype, BM!); Hongkong, coll. Ah

Nin (B. 21E), 12 Jul 1931 (paratype, BM!); Hongkong, granite rock, 1,000-

1,500 ft. alt., Shaukiwan, coll. Ah Nin (H. 4I),5 Oct 1931 (paratype, BM!);

Hongkong, Lan Yau Peak, Lan Tau Is., coll. Herklots 359 (paratype, BM!).

Plants small to moderate-sized, forming yellowish-brown mats. Stems

creeping, up to 4 cm long, with erect branches, branches 5—6 mm high and

1—1.3 mm wide. Branch leaves strongly crisped when dry, widely spreading

when moist, entire, long lingulate or lanceolate, 2.5-3 mm long, 0.4—0.5 mm

164 Gard. Bull. Singapore 58 (2) 2007

wide; the apex acute or acuminate, somewhat incurved, keeled, slightly

plicate below; margins entire, usually plane on one side and recurved on

the other; costa single, prominent, ending below the apex or rarely in the

apex, smooth; laminal cells unistratose, clear and smooth from top to base,

moderate thick-walled; upper and median laminal cells small and clear, in

diagonal rows, subquadrate-rounded to rounded-quadrate, 5-7 um wide,

gradually elongate from lower part to base. Basal laminal cells along costa

rectangular, thin-walled, 7.5—-12.5 um wide, 20-30 um long, looking like

“calymperoid” cells, those near margins elongate, rectangular to sublinear,

17.5—42 um long, 5-10 um wide, thick-walled and slightly sinuous.

Sporophytes lateral on branches. Inner perichaetial leaves similar to branch

leaves, ovate-oblong, acuminate, 2.5—3 mm long, 0.5—0.6 mm wide; costa

smooth, ending below the apex; laminal cells quadrate or subquadrate,

smooth and clear. Paraphyses numerous. Setae 2—3 mm, smooth, twisted to

right. Capsules oblong-cylindric or oblong, 1.6—2 mm long, 0.6—0.9 mm wide,

brown, constricted or slightly constricted below the mouth in 4-angled or

4 -furrowed shape, peristome absent. Calyptrae rather large, campanulate,

2.5—3 mm long, with many long, brown-yellowish hairs. Spores not seen.

M. heterodictyon is somewhat similar to M. schmidii, but differs from the latter

by its smooth and clear laminal cells from leaf top to base, and the capsules

constricted below the mouth in a 4-angled or 4-furrowed shape. It is also

similar to M. schmidii var. macroperichaetialium in their plant and capsule

shape, but the new variety could be distinguished from M. heterodictyon by

its pluripapillose laminal cells and its longer inner perichaetials leaves with

gradually long-acuminate upper part.

In Dixon’s original publication of M. heterodictyon, he considered the

species similar to M. japonicum, only differed from the latter by its

gymnostomous capsules. Our check on the types of M. heterodictyon shows

that M. heterodictyon is distinctly different from M. japonicum by its clear

and smooth laminal cells without any papillae, while the upper and median

laminal cells of M. japonicum are obscure and densely pluripapillose.

M. heterodictyon Dix. is a distinct species, its diagnostic features are: 1) plants

small to moderate-sized, branches short, less than 6 mm high; 2) laminal

cells smooth and clear from top to base, median and upper laminal cells

in diagonal rows, moderately thick-walled, basal laminal cells along costa

rectangular, thin-walled, “calymperoid” in areolation; 3) inner perichaetial

leaves ovate-oblong, acuminate, similar to branch leaves in length; 4)

oblong-cylindrical to oblong gymnostomous capsules, constricted below the

Taxonomic notes on Asian species of Orthotrichaceae 165

Fig. 7. Macromitrium heterodictyon Dix. a-b. Perichaetial leaves; c-g. Branch leaves; h. Basal lamina

cells near costa; i. Upper laminal cells; j. Lower laminal cells; k. Basal laminal cells near margins;

|. Transverse section of the basal part of leaf; m-o. Transverse section of the middle part of leaf; p.

Transverse section of the upper part of leaf; q-r. Dry capsules; s. Wet capsule. All from holotype of

M. heterodictyon at BM. Line scales: A = 0.5 mm (a-g); B = 50 um (h-p); C= 0.8 mm (q-s).

166 Gard. Bull. Singapore 58 (2) 2007

mouth in a 4-angled or 4 —-furrowed shape; and 5) calyptrae rather large,

campanulate, up to 3 mm long, with many long, brown-yellowish hairs.

Distribution: China.

Macromitrium gymnostomum Sull. & Lesq., Proc. Am. Ac. Arts Sc. 4: 278.

1859. (Figs. 8-10)

Type: Japan, Shizuoka Pref., Simoda, Wright (lectotype selected by Noguchi,

1967, FH, not seen; isolectotype, NY00512839!); Japan, Ousima, one of the

northern Loo Choo Islands (syntype, NY00512840!).

Macromitrium brevituberculatum Dix., Hong Kong Naturalist, Suppl. 2:

14. 5. 1933. syn. nov. Type: “Granite rock, 1,000-1,500 ft. alt., Shaukiwan,

Hong Kong Is, 5 Oct 1931, coll. Ah Nin (H.8).” (holotype, BM 000576135!);

“near sea level, Stanley, Hong Kong, 14 May 1931, coll. Ah Nin (H. 43A)”

(Paratype, BM 000576134!).

Macromitrium robinsonii R.S. Williams, Bull., New York Bot. Gard. 8: 344,

1914, syn. nov. Type: “Philippines, Upper Lamao River, on tree, 1000 meters,

Jan 1904, Williams 1760“ (isotype, H-BR 2617006!)

Plants small to medium-sized, in dense, dark or reddish-brown mats, younger

parts dark-greenish. Stems elongate, sparsely leaved, densely covered with

reddish rhizoids; branches erect, up to 5 mm long, simple, or with several short

branchlets. Branch leaves crisped when dry, widely spreading when moist,

linear or linear-lanceolate, 1—2.5 x 0.15—0.2 mm; Keeled, apex obtuse acute,

acute or acuminate, lower half of leaves yellowish and hyaline, upper half

rather obscure, leaf margins sometimes slightly recurved; costa yellowish-

brown, reaching or almost reaching to the apex; laminal cells unistratose

from top to base; median and upper laminal cells rather obscure, rounded or

rounded-hexagonal, 4—5 um, thin-walled, densely pluri-papillose; lower cells

longer, rectangular, with or without papillae, basal cells linear, 8—25 um long,

with thickened walls, smooth.

Sporophytes lateral on branches. Inner perichaetial leaves ovate-lanceolate,

acuminate, costa extending to leaf apex; laminal cells hyaline, smooth.

Paraphyses numerous, slightly exserted beyond leaves. Setae brown, smooth,

usually 5-8 mm long. Capsules obloid-cylindric, brown, deeply plicate,

constricted at mouth when dry, peristome absent; operculum conic-subulate.

Spores finely papillose, 20-25 um diameter. Calyptrae cucullate, 1.7—-2 mm

long, somewhat lobed and plicate, naked.

M. brevituberculatum was described by Dixon in 1933 based on the

Taxonomic notes on Asian species of Orthotrichaceae 167

Fig. 8. M. gymnostomum. a-d, g-n. Branch leaves; 0. Apex of branch leaf; p-q. Stem leaves; r,

t. Basal transverse section of leaf; s. Upper transverse section of leaf; u. Basal laminal cells;

v. Basal laminal cells near margins; w. Upper laminal cells; x. Spores; y-z. Capsules. (a-c, p-r,x

from Hongkong, Merrill 11159, marked as “M. gymnostomoides Broth.”; e-k, s-w, y-z, from

isolectotype of M. gymnostomum; m-o from Japan, Ferrie, marked as “M. subgymnostomum

Ther.). Line scales: A = 0.5 mm (a-n, p-q, y-z); B = 50 um (1-x).

168 Gard. Bull. Singapore 58 (2) 2007

Fig. 9. M. gymnostomum. a-n. Branch leaves; p-r. Upper transverse section of leaf; 0,s. Basal

transverse section of leaf; t, u. Basal laminal cells near margins; v. Low laminal cells; x. Dry capsule;

y. Calyptra. (a-g, o-r, u from holotype of M. brevituberculatum Dix. at BM); (h-n, s-t, v-y from

isotype of M. robinsonii at H-BR). Line scales: A = 0.5 mm (a-n, x-y); B = 50 um (0-w).

Taxonomic notes on Asian species of Orthotrichaceae 169

a b d

Fig. 10. M@. gymnostomum. a-d, h. Wet capsules; e-g, j. Calyptrae in different stages; 1. Perichaetial

leaf. (a from isolectotype of M. gymnostomum at NY; b-c, e from Japan, Ferriei, marked as

“M. subgymnostomum Ther.”; f-i from isotype of M. robinsonii at H-BR; j from holotype of

brevituberculatum Dix. at BM). Line scales: A = 0.50 mm (a-h, j); B = 0.2 mm (j).

170 Gard. Bull. Singapore 58 (2) 2007

collection by Youngsaye from Taimoshan, Hong Kong, and M. robinsonii by

R.S. Williams in 1914 based on the material from the Upper Lamao River in

the Philippine (in memory of C.B. Robinson who lost his life on a collecting

trip). Our check of their types showed that these three species belong to the

same taxon, therefore we treated M. brevituberculatum and M. robinsonii as

two new synonyms of M. gymnostomum.

Macromitrium gymnostomum was described by Noguchi in details in 1967.

The morphological characters of M. gymnostomum are relatively stable.

Most specimens of this species are characterized by their lanceolate, linear-

lanceolate or oblong-lanceolate branch leaves with acute or acuminate

apices, usually 1—2.5 x 0.15—0.2 mm. There is a specimen in H-BR collected

from Hong Kong, China, marked as “Macromitrium gymnostomumoides

Broth. sp. nov.” (HB2541002). This specimen is rather special in having

long linear branch leaves, up to 3.6 mm long, with long acuminate apices.

Another specimen collected from Japan was marked as “Macromitrium

subgymnostomum Ther. sp. nov.” (HB2541001). It is characterized by short

lanceolate branch leaves with obtuse acute apices. We consider that the

branch leaves of M. gymnostomum can vary from lanceolate with obtuse

or acute apices to long linear with long acuminate apices, and so, both

specimens fall into the range of leaf variation of M. gymnostomum. M.

gymnostomum is easy to separate from other gymnostomous taxa in Asia

by its naked calyptrae.

Distribution: China, Japan, Korea, Philippine, Vietnam (Tan & Iwatsuki,

1993).

Other specimens examined: CHINA: Taiwan, collector not clear, no. M3;

Fujian, Mt. Wuyi Guo S.-L. 030001-030008 (SNUH); Amoy, Chung (NY);

Hong Kong, Loh Fau Mountain (Lofaushan), E.D. Merrill 11159, marked

as “Macromitrium gymnostomoides Broth., new sp.’(H-BR 2541002);

Kwangtung (Guangdong), Ngok Shing Shan, Sai-lin-shan village, Y W. Taam

402A (H); Hunan, Badagongshan, Enroth 58960 (H); Changsha, Mt. Yuelu,

Enroth 48962 (H); Daweishan. DAW16, Virtanen 61994 (H); Mangshan,

Koponenetal.51680,51449,51411,50762 (H);Shunhuangshan., Enroth 70674,

70760 (H); Taoyuandong, Koponen et al. 55092 (H); Wulingyuan, Koponen

et al. 53352, 52057, 58525 (H); Yuankou, Enroth 59595 (H); Yunshan, Enroth

70130 (H). Zhejiang, Hangzhou, Tianmu Mountain, W.R. Buck 23916 (BH).

JAPAN: Kagoshima, Sumiyo-mura, Island. Amami-ohshima, K. Saito (H);

Okamura, Mt. Maya Settsu, Shutai 684 (H); Okamura, Mt. Kuishi, Tosa, Shutai

202 (H); Liou, Kine Sushi, 20 Apr 1898, Ferrie, marked as “Macromitrium

subgymnostomum Ther. sp. nov” (H-BR2541001); Wakayama, Syégawa,

Taxonomic notes on Asian species of Orthotrichaceae £7

Shirahama-ch6. Nishimuro-gun, 7! Nakajima 1129 (H); Ohara, Yuya, Oto-mura,

Nishimuro-gun, 7. Nakajima 1229 (H); Higo, Aida, K. Mayebara 3623 (H);

Kumamoto, Koonose, K. Mayebara 430 (H).

Macromitrium taiheizanense Nog., J. Sc. Hiroshima Univ. ser. B(2), Bot. 3:

11.1. 1936. (Fig. 11)

Type: “Formosa: Mt. Taiheizan (ca 2000m), Prov. Taihoku, Aug 1932 (A.

Noguchi 6548-type, holotype, NICH 365243!)

Plants robust, yellow-brownish above, dark-brownish below, in mats.

Stems creeping, branches up to 7-8 mm high, densely covered with reddish

rhizoids. Branch leaves regularly, strongly and spirally twisted when dry,

flexuose-spreading when moist, entire, funiculate, or gradually narrowed to

a slender acuminate acumen or subula from a broadly lanceolate to ovate-

oblong lower portion, 5—~7 mm long, 0.5—0.6 mm wide at base, leaf basal

longitudinal plicate; margins entire plane; costae ending in the apex; laminal

cells unistratose from top to base; upper and median laminal cells smooth

and clear, subquadrate-rounded to rounded-quadrate, 8-10 um wide; basal

laminal cells somewhat sinuous, gradually elongate from lower part to base,

thick-walled, 22-50 um, 3.5-7.5 um wide, bulging and with a papillae per

cell.

Sporophytes lateral on branches. Inner perichaetial leaves ovate-oblong,

long-acuminate in upper part, 6-7 mm long, similar to the branch leaves in

length. Setae rather short, 1—1.2 mm long. Capsules ovate, 1.4—1.5 mm wide,

1.6-1.9 mm long, brown, slightly ribbed under a smooth mouth; peristome

absent. Spores not seen. Calyptra campanulate, with many long, yellowish

hairs.

M. tatheizanense is a very distinct species, and could be separated from its al-

lies by its funiculate or linear to linear-lanceolate branch leaves with smooth

and clear laminal cells from top to middle.

Distribution: China (Taiwan).

Macromitrium formosae Card., Beih. Bot. Centralbl. 19(2): 104. 8. 1905.

(Fig. 12)

Type: “Formosae: Kelung, 1903, Faurie 181” (isotype, H3090260!)

Plants medium-sized, forming yellowish-brown mats. Stems creeping, with

erect branches, branches up to 10 mm high. Branch leaves crisped when

dry, flexuose-spreading when moist, entire, lanceolate to linear-lanceolate,

172 Gard. Bull. Singapore 58 (2) 2007

Fig. 11. M. taiheizanense. a-b. Branch leaves; c-d, k, i-h. Transverse sections of leaf; e. Lower

laminal cells near margin; f. Lower laminal cells;,g. Lower laminal cells near margin; j. Upper

laminal cells; m-n. Capsules. All from holotype of M. taiheizanense at NICH). Line scales: A

= 0.5 mm (a-b, m-n); B = 80 um (c,h, k); C = 50 um (d, 1, f, g, i-j).

Taxonomic notes on Asian species of Orthotrichaceae 173

acuminate, revolute below on one side, and somewhat plicate at base. 2.3—

2.7 mm long, 0.25—0.4 mm wide; costae single, prominent, ending below

the apex; laminal cells unistratose from top to base; upper and median

laminal cells irregularly quadrate, 7-8 um wide, rather obscure, densely

pluripapillose, basal laminal cells 12-20 um long, 3-4 um wide, cell walls

sinuous and thick, without papilliae, those along the basal margins regular

and smooth. Setae 5-6 mm long, smooth. Inner perichaetial leaves similar to

branch leaves. Capsules ovate, 1.2—1.4 mm long, 0.65—0.8 mm wide, brown,

with a small mouth, slightly ribbed below the mouth when dry; peristome

absent. Calyptrae campanulate, with many long, brown-yellowish hairs.

M. formosae Card. was described by J. Cardot based on the collection of

Faurie from Taiwan in 1905. He had compared it with M. tylostomum Mitt.

even the latter was distinctly different from M. formosae by its larger plants,

much wider branch leaves with obtuse to subacute apices with a distinct

apiculus (Eddy, 1996). In fact, M. formosae is somewhat similar to M.

gymnostomum in appearance, branch leaf shape and laminal cells, but could

be separated from the latter by its hairy calyptrae.

Distribution: China (Taiwan), Philippine (Tan & Iwatsuki, 1991)

Macromitrium holomitrioides Nog., J. Sc. Hiroshima Univ. ser. B(2), Bot.

ome epee Bis

Type: “Taiwan, Taihoku, Rahau, Noguchi” (holotype, HIRO, not seen)

M. holomitrioides Nog., a species recorded from Japan (Kyushu) and China

(Taiwan), was described in details by Noguchi (1938, 1967, 1989). Based on

Noguchi’s descriptions and our check on the non-type specimens, we found

that M. holomitrioides is similar to M. schmidii var. macroperichaetialium in

their branch leaf shape, capsules constricted or slightly constricted below

the mouth in a 4-angled or 4 -furrowed shape, but the former could be

separated from the latter by: 1) its broad leaves with large, mammillose or

bulging, collenchymatous upper and middle laminal cells; 2) sublinear or

subrectangular basal laminal cells without papillae.

Specimens examined: CHINA: Hainan island, Jianfengling Mt., RC. Chen et

al. 789b (H); ibid, PC. Wu W1185016 (A).

Distribution: China, Japan.

174

Gard. Bull. Singapore 58 (2) 2007

Fig.12. M. formosae Card. a-c. Branch leaves; d-f. Dry capsules; m. Wet capsule; g. Basal

laminal cells; h. Upper laminal cells; i. Basal laminal cells near margin; j-l. Transverse sections

of leaf. All from isotype of M. formosae at BH. Line scales: A = 0.5 mm (a-f, m); B = 50 um

(g-l).

Taxonomic notes on Asian species of Orthotrichaceae (Ve)

Discussion

Taxonomically, the above seven species belong to Sect. Goniostoma in

the subgenus Macromitrium. Based on our checking on the literature

and specimens, seven taxa with gymnostomuous capsules of the genus

Macromitirum in East Asia are confirmed. The main identification

characteristics of these seven taxa are the calyptra, shape of branch leaves

and inner perichaetial leaves, papillosity of laminal cells and capsule shape.

The following key can be applied to separate the above seven taxa.

1. Calyptrae naked, without hairs....................... VM. gymnostomum

TOM acd PT MN ae a rs chs SPITS ncaa cure ne nll nae sieyainy s-+ais ooF tes wien ees iereni 2

2. Branch leaves funiculate, or linear to linear-lanceolate, up to 5-7 mm long

Ee ea ee ee eee eee M. taiheizanense

2. Branemieaves lanccolate,.shorter than 5. mm.long ....22.........ccds.00. 3

3. Median and upper laminal cells smooth or almost smooth ...... ee.

Renee ern eet ete $e AMG oe bese Ba ces ast Piss Sake debetvay late M. heterodictyon

3. Median and upper laminal cells pluripapillose, mammillose, bulging or

ON Ne Wg TRIO eo at rr By tas Toe hele c nes uae dacs snddededacucedeledennnrsesaceves +

4. Median and upper laminal cells mammillose, ioe collenchymatous,

Basar iamimal Cells SMOOUR: jc... Seeeidetecncsenastede «5 . M. holomitrioides

4. Median and upper eeaiel! cells she Seite eal laminal cells

Peete GUIO OU 41h frrcersa centers bp dcthnatl sdacossenaeusAssyenicoun tenet. ao

>_ Basal laminalcells without papilharc.i.cco-ci docs taseeecekd,cticlad sss M. formosae

5. Basal laminal cells tuberculate or with a high papilla ....... 0 eee 6

6. Inner perichaetial leaves ovate-oblong with acuminate apex, shorter than

LPC) WEE) iS eee Whe oe ee ee eee M. schmidii

6. Inner perichaetial leaves long lanceolate with linear acuminate apex, much

longer than branch leaves ............ M. schmidii var. macroperichaetialium

176 Gard. Bull. Singapore 58 (2) 2007

Acknowledgements

This research was supported by the National Nature Science Foundation

of China (no. 30570121), and the Scientific Research Foundation for the

Returned Overseas Chinese Scholars, State Education Ministry. We also

gratefully acknowledge the financial support given by the State Scholar-

ship Fund of China and the Finnish-Chinese Botanical Foundation for

the visit of the first author to the University of Helsinki in 2003-2004.

Thanks to Gao Chien (Institute of Applied Ecology, Academia Sinica),

J. Hyv6nen, T. Koponen, Xiao-Lan He, J. Heino, S. Sallinen, T. Ahti, and

S. Huttunen at Herbarium of Natural History Museum of the Univer-

sity of Helsinki for their hospitality and help during the present work.

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Garden 8: 332-377.

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Gardens’ Bulletin Singapore 58 (2) 2007: 179-198 179

VARIATIONS IN THE HOYA VERTICILLATA COMPLEX IN

THAILAND

MANIT KIDYUE, THAWEESAKDI BOONKERD, OBCHANT

THAITHONG AND TOSAK SEELANAN

Department of Botany, Faculty of Science, Chulalongkorn University, Bangkok

10330, Thailand

Abstract

Hoya verticillata (Vahl) G.Don s.l. is a climbing epiphyte belonging to

the family Apocynaceae. At present, the taxonomic status of this species

in Thailand is still uncertain due to the great variability of size, shape and

colour of leaf and flower. Morphological variations were explored in 500

fresh plants collected from 50 sites throughout the country. The collected

specimens represent the H. verticillata complex, including two polymor-

phic species, namely H. verticillata s.l. and closely related species, H. rigida

Kerr. Based on qualitative macro- and micro-morphological characters, the

H. verticillata complex can be divided into nine groups. They can be distin-

guished by leaf shape, leaf base, venation pattern, leaf indumentum, and

shapes of sepal, corona and corpusculum. Group I matched with the char-

acteristics of H. rigida Kerr, which has ovate leaves with cordate base, 3-5

prominent nerves running from base to apex; indumentum absent on the

abaxial surface; and lanceolate sepals. In contrast, Group II is an unidenti-

fied taxon that is close to H. verticillata (Vahl) G. Don var. citrina (Ridl.)

Veldkamp. It is charaterized by broad ovate leaves with cordate base, 3-5

prominent nerves running from base to apex; and narrowly oblanceolate-

oblong corpusculum. Groups III and IV have distinctly different vegetative

characters and do not correspond to the previously described varieties of

H. verticillata. Their leaf venation is acrodromous, with 3 prominent nerves

running from leaf base parallel to the midrib and reaching the apex, but

they are different in their shape of leaf and coronal scale. Group III has

ovate leaves with subcordate base and elliptic coronal scale, while Group

IV has elliptic-oblong leaves with cuneate base and ovate-lanceolate co-

rona. Groups V-IX are still of weak validity, having slightly discontinu-

ity in leaf shape, base and venation; and shape of coronal scale. They are

treated in this paper as variable groups within H. verticillata var. verticillata.

180 Gard. Bull. Singapore 58 (2) 2007

Introduction

Hoya verticillata (Vahl) G.Don (Apocynaceae: Asclepioideae) is one of the

most common members of the section Hoya (Hooker f., 1883). It is relatively

widespread, occurring from E. India-Assam through Myanmar, Thailand,

Indo-China, Malay Peninsula, and Sumatra to N. Borneo (Rintz, 1978). The

most recent taxonomic study of the species (Kiew, 1995; Veldkamp et al.,

1995) included three varieties: H. verticillata var. verticillata, H. verticillata

var. citrina and H. verticillata var. hendersonii. These varieties can be

distinguished from each other by leaf characters and habitat. H. verticillata

var. verticillata is distinguished from the other two varieties by its elliptic leaf,

cuneate base, obscure veins with the lowest pair extending about halfway

to apex. The other two varieties are similar in having an ovate leaf with

slightly cordate base, but with conspicuous veins and the lowest vein pair

extending to apex. As for habitat, var. hendersonii grows in lower montane

forest between 1,200 and 1,260 m altitude, while the other two are lowland

varieties (Kiew, 1995).

In Thailand, H. verticillata s.l. is the most common hoya, and is an extremely

variable species. There is variation in texture, shape, size and venation of

leaves, the size of the flower and shape of corolla lobes (Kerr, 1951). These

plants were later named “the H. verticillata complex” (Thaithong, 1995).

However, variations within this complex did not match properly with the

three fomerly recognized infraspecific taxa. Additional intensive research

into relationships within this difficult group of plants has not been carried

out for many years.

Moreover, previously distinct species, H. ridleyi and H. rigida have been

taxonomically confused with H. verticillata s.l. H. ridleyiis found in peninsular

Thailand (Ridley, 1923), but has also been recognized as H. parasitica var.

parasitica, the synonym of H. verticillata var. verticillata (Rintz, 1978). Then,

Veldkamp et al. (1995) suggested that H. rigida, a species endemic to Thailand,

may be included in H. verticillatas.l.. Thus, the taxonomic status of H. verticillata

complex in Thailand is still uncertain and needs to be reinvestigated.

We suspect that H. verticillata complex in Thailand is composed of several

undescribed taxa. We have explored all the available variables in the

complex and have determined macro- and micro-morphological variations

of each group. In this paper, we have (1) described the overall qualitative

morphological variations of characters in the complex, (2) classified

the variations into groups, and (3) described morphological characters,

geographical distribution and ecological features of each recognized group.

Variations in the Hoya verticillata complex in Thailand 181

We then (4) discuss the taxonomic treatment of these recognized groups as

compared with the previous classification.

Materials and methods

Plant Morphology

Plant materials used in this study were collected from 50 sites throughout

Thailand during 2002-2003 (Table 1, Fig. 1) and all were raised in the same

conditions in a greenhouse at the Department of Botany, Faculty of Science,

Chulalongkorn University. Flower and leaf materials were then gathered

from the living collection. These were preserved in 70% ethanol for further

morphological studies. In general, the morphological characters of leaves

and flowers were examined using light microscopy (LM). Leaf indumentum

on both surfaces was observed under LM and scanning electron microscopy

(SEM).

Habitat and Distribution

Habitats of H. verticillata complex were studied and their location recorded

using GPS. Collection sites were plotted on an outline map, thus showing

their distribution, in relation to forest types and elevation, and characterizing

the habitat of particular member of the complex.

Results

General description of Hoya verticillata complex (Fig. 2—4)

Glabrous twiner, epiphyte, with long internodes. Leaves opposite or

whorled; /amina ovate or elliptic to oblong, glabrous or papillate on abaxial

surface; texture coriaceous to succulently coriaceous, depending somewhat

on habitat; venation with three or five principal veins (nerves) arising near

the junction of lamina and petiole; nectary glands present at the junction

of petiole and lamina; petioles usually thick and stout, occasionally slender

in mature leaves. Inflorescences axillary, negative geotropic umbels with

almost equal pedicel; flower actinomorphic, 5-merous; pedicels slender;

calyx glabrous or covered with hairs on abaxial surface and margin, with a

small gland alternately arranged between calyx lobes; corolla rotate, abaxial

surface glabrous, covered with minute hairs on adaxial surface, reflexed

when in full bloom, becoming closed at the end of flowering period; corolla

lobe ovate-triangular with acute or acuminate apex, inflexed (bent inwards)

between corona scales; corona longer than corolla tube, coronal scale ovate-

elliptic or ovate-lanceolate; the first inner angle abruptly apiculate, slightly

raised, the outer angle broadly or narrowly acute, concave on the upper

182 Gard. Bull. Singapore 58 (2) 2007

GULF OF

THAILAND

ANDAMAN SEA ©

Lg Te

100

MALAYSIA 194

Fig. 1. Sampling sites of Hoya verticillata complex in Thailand.

Variations in the Hoya verticillata complex in Thailand

183

Table 1. Locality and collected groups of the Hoya verticillata complex in

Thailand.

Siteno. Locality Group’

i Mueang, Chiang Rai Province V

2 Wang Bua Ban Waterfall, Chiang Mai Province V, IX

. Haew Keaw, Mueang, Chiang Mai Province VI

4. Doi Khun Tan National Park, Lampang Province IX

S. Silaphet Waterfall, Pua, Nan Province III, VI

6. Tat Laung Waterfall, Pua, Nan Province Vil

7. Ban Muang Wang Nhua, Phu Phiang, Nan Province IX

8. Phasing, Mueang, Nan Province V

9. Lhinan, Na Noi, Nan Province VII, 1X

10. Pla Ba Waterfall, Phu Ruea, Loei Province IX

it. Than Thong Waterfall, Sri Chiang Mai, Nong Khai Province IX

12. Than Ngam Waterfall, Nong Wua So, Udon Thani Province IX

is Tat Kham Waterfall, Ban Phaeng, Nakhon Phanom Province IX

1a. Phu Phan National Park, Sakon Nakhon Province IX

15. Tat Ton Waterfall, Mukdahan Province IX

16. Soi Sawan Waterfall, Ubon Ratchathani Province IX

ee Huai Sai Yai waterfall, Sirinthon, Ubon Ratchathani Province IX

18. Phu Chongna Yoi National Park, Ubon Ratchathani Province IX

19. Tat Hai Waterfall, Nam Yuen, Ubon Ratchathani Province IX

20. Sam Long Kiat Waterfall, Khun Han, Si Sa Ket Province IX

PAW Phu Wa Kiew Waterfall, Nakhon Ratchasima Province IX

JPR Khao Yai National Park, Nakhon Ratchasima Province | bal ®.€

23, Pu Kae, Saraburi Province IX

24. Pang Sida National Park, Sa Kaeo Province IX

Uh Prachantakham, Prachin Buri Province VIII

184 Gard. Bull. Singapore 58 (2) 2007

Site no. — Locality Group’

26. Ban Pe, Rayong Province Vill

21: Nam Tok Phriu National Park, Chanthaburi Province VIII

28. Mueang, Trat Province Vill

29: Ko Chang, Trat Province Vill

30. Thi Lo Su waterfall, Tak Province IX

3h Sangkhla Buri, Kanchanaburi Province IX

SZ Koeng Kra Wia Waterfall, Kanchanaburi Province IX

33) Pong Ron, Thong Pha Phum, Kanchanaburi Province IX

34. Pha Suk Pass,Thong Pha Phum, Kanchanaburi Province I

35: Ban Thamadua, Thong Pha Phum, Kanchanaburi Province IX

36. Sai Yok National Park, Kanchanaburi Province IX

3% Bang Saphan, Prachuap Khiri Khan Province Vill

38. Namtok Ngao National Park, Ranong Province IX

39. Ko Wua Ta Lub, Suratthani Province VIII

40. Mueang, Phangnga Province Vil

41. Noppharat Thara Beach, Krabi Province VIII

42. Khao Luang National Park, Nakhon Si Thammarat Province I

43. Pakmeng, Trang Province Vill

44. Hat Chao Mai National Park, Trang Province VI

45. Thung Kai, Trang Province Vill

46. Sathing Phra, Songkhla Province VII

47. Singhanakhon, Songkhla Province Vill

48. Pak Bang Sakom Beach, Songkhla Province Vill

49. Tarutao National Park, Satun Province IV, VIII

50. Sirinthon Waterfall, Waeng, Narathiwat Province II

a = Site numbers correspond to those in Fig.1; b = Nine groups distinguished in this study.

Variations in the Hoya verticillata complex in Thailand 185

\ if \ hy i ; / \ /

/ 4

sig Y NY ot 5 ¥ TY

Fig. 2. Leaves of Hoya verticillata complex. A: Group I, site 42 (166). B: Group I, site 50

(303). C: Group III, site 5 (380). D: Group IV, site 49 (320). E-F: Group V, E. site 1 (404), F. site

8 (378). G-H: Group VII, G. site 6 (385), H. site 5 (384). I: Group VI, site 3 (289). J-M: Group

IX, J. site 11 (559), K. site 14 (436), L. site 24 (3), M. site 33 (258). N-T: Group VIII, N. site 27

(120), O. site 40 (670), P. site 45 (66), Q. site 39 (585), R. site 46(243), S. site 47 (213), T. site 49

(202). Numbers in parenthesis denote the collector number. Bar = 2 cm.

186 Gard. Bull. Singapore 58 (2) 2007

Si ef

Rati

; heal a wan, by ia

Fig. 3. Scanning electron micrographs of the abaxial surface of leaves of Hoya verticillata

complex. A: Group I (absent indumentum); B: Group II (minute and scattered indumentum);

C: groups III-IX (dense indumentum). Bar = 50 um.

Fig. 4. Flower and flower parts of Hoya verticillata complex. A-B (top view of flower), A:

Groups II, IV, VI,VHI and IX (coronal scales ovate-lanceolate); B: Group I, II, V, VII

(coronal scales broad ovate or ovate-elliptic); C-D (bottom view of flower), C: Group I

(sepal lanceolate); D: Group II-IX (sepal ovate); E-F (polinaria), E: Group II (corpuscles

oblanceolate-oblong); F: Groups I, III-[X (corpuscles triangular-oblong). A-D: Bar = 2 mm;

E-F: Bar ='0.2 mm:

Variations in the Hoya verticillata complex in Thailand 187

surface usually with a median ridge; anthers 5, lying just below the inner

coronal angle, each producing two pollen masses or pollinia, each pollen

mass from adjacent anther lobes connected by short translator arms with a

corpusculum, forming a pollinarium; pollinium flattened, oblong, truncate at

the top, with narrow translucent longitudinal wing; corpusculum dark brown

or black. Fruits a follicle, pod-like, straight and glabrous.

Comparative morphology

Despite the great observed variations (i.e., habitat, leaf shape, forms of leaf

base, venation pattern of leaf, forms of sepal and corona), we can tentatively

classify the variations of the complex into nine groups by a comparison of

qualitative morphological characters. These groups are then named as Group

I-[X (IX being the residual group). A key to the Groups and descriptions of

the Groups are presented below.

Key to Groups of wild H.. verticillata complex in Thailand

1. Sepals lanceolate, longer than the corolla tubes, leaves glabrous (Fig. 3A,

BM epeareers ch are iN cyt. Heth nn Lcdehle.d. Yop bicks= Ab adja enthdenadede\ovosisaeess Group I

1. Sepals ovate, equal in length or shorter than corolla tubes, leaves papillate

on abaxial surface (Fig. 3B-C, 4D)

2. Leaves with prominent basal nerves, extending from base to apex,

veins conspicuous (Fig. 1A-D)

3. Leaves ovate, base rounded to cordate

4. Coronal scales ovate-lanceolate (Fig. 4A), corpusculum narrowly

oblanceolate-oblong (Fig ,AF), 2f.p.oi.s3adscdats<ssctoastecarersdeced. Group II

4. Coronal scales ovate-elliptic (Fig. 4B), corpusculum broadly

Plame lar ODlOMO: (HIE LAP)) sccccsscccecessascancdavdeaseaenesesnecatsne Group III

py Leaves OVate-OblOme, DASCICMINGALE s... snc. scecccceoncsaesocsesedeses Group IV

2. Leaves with unclear basal nerves, mostly extending about half way to

apex, veins obscure (Fig. 1E-T)

5. Leaf base rounded to subcordate

6. Coronal scales ovate-elliptic, petioles stout ................ Group V

6. Coronal scales ovate-lanceolate, petioles slender ....... Group VI

188 Gard. Bull. Singapore 58 (2) 2007

5. Leaf base cuneate to obtuse

7.. Coronal scales ovate-clipnc. 2322) 2 Group VII

7. Coronal scales ovate-lanceolate

8. Leaves relatively narrow, 1.8-(3.8)-5.8 cm wide, base cuneate.

Littoral plant: csisclic:duscocteieeaaseee ee eee Group VIII

8. Leaves broad, 4-(5.8)-8.6 cm wide, base variable. Inland plant

found throughout Thailand tee eee Group IX

Group I

Leaves rigidly coriaceous, broad ovate to oblong with rounded to cordate

base, 13.8-(17.8)-19.1 cm long, 6.8-(8.7)-10.1 wide; nerves 3-5, prominent,

extending from base to apex, the other veins conspicuous; glabrous on both

surface; petiole slender, 2.7-(3.4)-3.9 cm long. Umbel 7-(30)-65 flowered,

peduncle 1.5-(2.9)-4.7 cm long; pedicel glabrous, 2-(2.3)-3 cm long. Sepal

lanceolate, longer than corolla tube, 4.9-(5.4)-5.6 mm long, 2-(2.1)-2.2 mm

wide. Corolla creamy white, 1.6-(1.9)-2.1 cm diam. Coronal scale broad

ovate, 4.2-(4.5)-4.8 mm long, 2.2-(2.4)-2.6 mm wide, the outer angle slightly

erected and white, apiculus of the inner angle raised and pink. Pollinia

oblong, 0.62-(0.63)-0.64 mm long, 0.21-(0.23)-0.24 mm wide; translator short;

corpusculum triangular-oblong, 0.16-0.18 mm long, 0.12-0.13 mm wide.

Ecology and distribution: A long climbing epiphyte, usually on trees along

stream banks, in moist evergreen forest, in the eastern, western and southern

parts of the country at 700-900 m elevation.

Specimens examined: Kanchanaburi - Thong Pha Phum, Manit Kidyue 464,

465, 466, 467, 468, 469. Nakhon Ratchasima - Khao Yai, Manit Kidyue 360,

361, 997. Nakhon Si Thammarat - Khao Luang, Manit Kidyue 166.

Group II

Leaves coriaceous, broadly ovate or rhombic with cordate base,

14.7—(15.9)-16.7 cm long, 11.3—(12.3)—13.2 cm wide; nerves 3—5, prominent,

extending from base to apex, veins conspicuous; hairs minute, scattering

on the abaxial surface; petiole slender, 1.8—(2.1)—2.2cm long. Umbel

10-—(26)—49 flowered, peduncle 2.5—(3.3)—4.3 cm long; pedicel glabrous,

1.7-(1.9)-2.0 cm long. Sepal ovate, 1.8—(1.9)—2.0 mm long, 1.5—(1.7)-1.8

mm wide. Corolla creamy white, 1.4-1.5 cm diam. Coronal scale ovate-

Variations in the Hoya verticillata complex in Thailand 189

lanceolate, 3.5—(3.8)—4.0 mm long, 1.5—(1.6)—1.7 mm wide, the outer angle

flattened and pinkish white, apiculus of the inner angle raised and pink.

Pollinium oblong, 0.45—0.46 mm long, 0.15—0.16 mm wide; translator short;

corpusculum narrowly oblanceolate-oblong, 0.2—0.22 mm long, 0.07—0.08

mm wide.

Ecology and distribution: A long climbing epiphyte in moist evergreen forest,

usually on trees along stream banks, restricted to Bala forest, peninsular

Thailand at about 200 m elevation.

Specimens examined: Narathiwat - Sirinthon Waterfall, Manit Kidyue 128,

303, 304, 305, 306, 688.

Group Il

Leaves rigidly coriaceous, broadly ovate with subcordate base, 14—(14.6)—15.3

cm long, 5.9-(7.4)—8 wide; nerves 3, prominent, extending from base to

apex, veins conspicuous; covered with dense hairs on abaxial surface;

petiole stout, 1.1-(1.9)—2.1 cm long. Umbel 22—(43)—59 flowered, peduncle

3.1-(4.5)—7.4 cm long; pedicel sparsely pubescent, 1.9-(2.2)-2.4 cm long.

Sepal ovate, 1.8—(1.9)—2 mm long, 1.4—(1.6)—1.8 mm wide. Corolla yellowish

white, 1.4—1.5 cm diam. Coronal scale ovate-elliptic, 3.5—(3.6)—3.8 mm long,

1.7—(2.0)—2.2 mm wide, the outer angle slightly erect and white, apiculus of

inner angle raised and pink. Pollinium oblong, 0.5—0.6 mm long, 0.19—0.22

mm wide; translator short; corpusculum triangular-oblong, 0.17—0.19 mm

long, 0.11—0.12 mm wide.

Ecology and distribution: A long climbing epiphyte in mixed deciduous

forest, or on rocks along stream banks, at about 400 m elevations, confined

to northern Thailand.

Specimens examined: Nan - Silaphet Waterfall, Manit Kidyue 380, 381, 382.

Group IV

Leaves large coriaceous, elliptic-oblong, with cuneate base, 18—(19.3)—20.4

cm long, 7.3—(7.7)-8 cm wide; nerves 3, prominent, extending from base

to apex, veins conspicuous; covered with dense hairs on abaxial surface;

petiole slender, 2—(2.5)—2.9 cm long. Umbel 65—(68)—71 flowered, peduncles

5.6—(7.3)—9.1 cm long; pedicel glabrous, 2—(2.2)—2.3 cm long. Sepal small

ovate, 1.7—1.8 mm long, 1.5-1.6 mm wide. Corolla creamy white, 1.2—1.3 cm

diam. Coronal scale ovate-lanceolate,3.4—(3.5)-3.6 mm long, 1.82-(1.86)—1.91

mm wide, the outer angle slightly erect and white, apiculus of the inner angle

190 Gard. Bull. Singapore 58 (2) 2007

short raised and pink. Pollinium oblong, 0.45—0.46 mm long, 0.17—0.18 mm

wide; translator short; corpusculum triangular-oblong, 0.16—0.17 mm long,

0.11—0.12 mm wide.

Ecology and distribution: A long climbing epiphyte on tree branches along

stream banks in evergreen forest, restricted to Tarutao Island, peninsular

Thailand.

Specimens examined: Satun - Tarutao National Park, Manit Kidyue 320,

32k:

Group V

Leaves coriaceous to succulently coriaceous, ovate with round to subcordate

base, 8.6—(12.5)—16.2 cm long, 5—(7.1)—10 cm wide; nerves 3, extending about

half way to apex or slightly longer, veins obscure; covered with dense hairs

on abaxial surface; petiole stout, 0.5—(1.2)-2.9 cm long. Umbel 21—(43)-87

flowered, peduncle 0.6-(3.0)-6.5 cm long; pedicel glabrous to pubescent,

1.9-(2.3)—2.8 cm long. Sepal ovate, 1.45—(1.8)—2.1 mm long, 1.4—(1.6)-1.9

mm wide. Corolla creamy white, 1.3—(1.5)-1.6 cm diam. Coronal scale

ovate-elliptic, 3.1-(3.7)-4.4 mm long, 1.9-(2.2)—2.4 mm wide, the outer

angle slightly erect to erect and white, apiculus of inner angle short raised

and pink. Pollinium oblong, 0.51—(0.55)—0.64 mm long, 0.18—(0.2)—0.21 mm

wide; translator short; corpusculum triangular-oblong, 0.15—(0.19)—0.23 mm

long, 0.11-—(0.12)—0.14 mm wide.

Ecology and distribution: On tree branches by stream banks in mixed

deciduous to dry dipterocarp forests, northern Thailand at about 300-600 m

elevation.

Specimens examined: Chiang Rai — Mueang, Manit Kidyue 404, 405, 406,

407, 407/2, 407/3, 408, 409, 409/2, 409/3, 409/4; Chiang Mai - Wang Bua Ban

Waterfall, Manit Kidyue 281, 282, 283, 284, 285, 292, 293, 294, 295, 296, 312,

313, 411, 413, 414, 414/2, 415, 416, 417, 417/2, 4177/3, 418, 419, 425; Nan — Meang,

Manit Kidyue 331, 332, 333, 378, 378/2, 378/3, 378/4, 378/5, 378/6, 378/7, 379,

37/2, 379/3, 379/4, 379/95, 379/6, 379/7, 379/8, 3799, 379/10.

Group VI

Leaves succulently coriaceous, broadly ovate, subcordate at base,

11-(11.2)-11.6 cm long, 7.4—(8.0)—8.3 cm wide, indistinct acrodromous

venation, the lowest pair extending about half way to apex; veins obscure;

covered with dense hairs on abaxial surface; petiole slender, 1.5—(2.2)—3 cm

Variations in the Hoya verticillata complex in Thailand 191

long. Umbel 26—(29)-30 flowered, peduncle 3.5—(5.8)—7.1 cm long; pedi-

cel pubescent, 1.7—(1.8)—1.9 cm long. Sepal ovate, 2—(2.2)—2.3 mm long,

1.6—(1.7)-1.8 mm wide. Corolla creamy white, 1.3—(1.4)-1.5 cm diam. Co-

rona scale ovate-lanceolate, 3.6—(3.7)—3.8 mm long, 1.8-1.9 mm wide, the

outer angle slightly erect and white, apiculus of inner angle short raised and

pink. Pollinium oblong, 0.49-—(0.5)—0.54 mm long, 0.17—0.18 mm wide; trans-

lator short; corpusculums triangular-oblong, 0.17—0.19 mm long, 0.11-—0.12

mm wide.

Ecology and distribution: On tree branches in dry dipterocarp forests at 350

m elevation, northern Thailand.

Specimens examined: Chiang Mai - Haew Keaw, Manit Kidyue 288, 289, 290,

291.

Group VII

Leaves coriaceous to succulently coriaceous, ovate-oblong with cuneate

to obtuse base, 10.4—(13.9)—17.9 cm long, 4.3—(5.6)—7.8 cm wide; 3 nerves,

extending about half way to apex or slightly longer, veins obscure; covered

with dense hairs on abaxial surface; petiole stout, 0.8—(1.4)—2.1 cm long.

Umbel 9-(31)-52 flowered, peduncle 0.6—(3.4)—8.1 cm long; pedicel

glabrous to pubescent, 1.8—(2.3)—2.6 cm long. Sepal ovate, 1.5—(1.9)—2.4 mm

long, 1.4—(1.6)—2.0 mm wide. Corolla creamy white, 1.5—(1.6)—1.8 cm diam.

Coronal scale ovate-elliptic, 3.4—(3.7)—4.1 mm long, 2.1—(2.2)—2.5 mm wide,

the outer angle erect and white, apiculus of inner angle short raised and

pink. Pollinium oblong, 0.54—(0.58)—0.62 mm long, 0.19—(0.2)—0.24 mm

wide; translator short; corpusculum triangular-oblong, 0.16—(0.18)—0.21 mm

long, 0.11-—(0.12)-0.15.

Ecology and distribution: On tree branches by stream banks in mixed

deciduous to dry dipterocarp forests at about 300-600 m elevation, northern

Thailand.

Specimens examined: Nan - Silaphet Waterfall, Manit Kidyue 325, 326, 327,

328, 329, 383, 384/1, 384/2, 384/3, 384/4, 384/5, 384/6, Tat Laung Waterfall,

Manit Kidyue 342, 343, 344, 345, 385, 385/2, 386, 387, Lhinan, Manit Kidyue

391, 392, 393, 394, 395, 391, 392, 393, 394, 396, 396, 402, 402, 403.

Group VIII

Leaves succulently coriaceous, narrowly ovate or elliptic to oblong, cuneate

or rarely obtuse at base, 5—(9.6)—15 cm long, 1.8—(3.8)—5.8 cm wide, in-

192 Gard. Bull. Singapore 58 (2) 2007

distinct acrodromous venation, the lowest pair extending about half way

to apex; veins obscure; covered with dense hairs on abaxial surface; peti-

ole stout, 0.3-(1.2)—2.8 cm long. Umbel 10-(28)—77 flowered, peduncle

0.8—(3.8)—11.4 cm long; pedicel glabrous to pubescent, 1.2—(1.7)—2.6 cm

long. Sepal ovate, 1.4—(2.0)-2.9 mm long, 1.1-(1.7)-2.6 mm wide. Corol-

la creamy white, 1.1—-(1.3)—1.6 cm diam. Coronal scale ovate-lanceolate,

2.9—(3.5)—4.2 mm long, 1.5—(1.7)—2.1 mm wide, the outer angle variable

from flattened to erect and white, apiculus of inner angle short raised and

white to pink. Pollinium oblong, 0.40—(0.48)—0.56 mm long, 0.15—(0.18)—0.2

mm wide; translator short; corpusculum triangular-oblong, 0.13-(0.16)-0.2

mm long, 0.10—(0.12)—0.14 mm wide.

Ecology and distribution: A littoral plant, on tree branches or on rocks, com-

mon in beach forest, coastal areas of islands, or mainland in eastern and

peninsular Thailand.

Specimens examined: Rayong - Ban Pe, Manit Kidyue 80, 81, 82, 83, 84, 85, 94,

95, 96, 99, 100; Chanthaburi - Nam Tok Phriu National Park, Manit Kidyue

110, 411, 113, 114, 115, 116, 117, 198, 119, 120. 121, F220 71235, 174 eee

135, 136; Trat — Mueang, Manit Kidyue 163, 164, 250, Ko Chang Manit Ki-

dyue 655, 656, 657, 658, 659, 660, 661, 662, 663, 664, 665, 666, 667, 668; Prach-

uap Khiri Khan - Bang Saphan, Manit Kidyue 622, 623, 624; Suratthani - Ko

Wua Ta Lub, Manit Kidyue 584, 585, 585, 586, 587, 588, 589, 590, 591, 592, 593,

594, 595; Phangnga — Mueang, Manit Kidyue 669, 670, 671, 672, 673, 674, 675,

676, 677, 678, 671, 672, 673, 674, 675, 676, 677, 678; Krabi [Noppharat Thara

Beach, Manit Kidyue 579, 580, 581, 582, 583, 634, 635, 636, 637, 638, 639, 640,

641, 642, 643]: Trang [Pakmeng, Manit Kidyue 693, 694, 695, 696, 697], [Hat

Chao Mai National Park, Manit Kidyue 628, 629, 630, 631, 632, 633], |Thung

Kai, Manit Kidyue 66, 67, 68, 69, 70, 71, 72, 73, 74]; Songkhla [Sathing Phra,

Manit Kidyue 224, 225, 226, 227, 228, 229, 230, 231, 232, 233, 234, 235, 236,

237, 238, 240, 241, 242, 243, 244, 245, 246, 247, 248], [Singhanakorn, Manit

Kidyue 212, 213, 216, 217, 218, 219, 220, 221, 222|, [Pak Bang Sakom Beach,

Manit Kidyue 607, 608, 609, 610, 611, 612, 613, 614, 615, 616, 617, 618]; Satun

[Tarutao National Park, Manit Kidyue 167, 168, 169, 170, 171, 172, 173, 174,

175, 176, 177, 178, 179, 180, 181, 182, 184, 185, 186, 187, 188, 189, 190, 191, 192,

193, 194, 195, 196, 197, 198, 200, 201, 202, 203, 204, 205, 206, 207, 208, 209, 210,

211, 596, 597, 598, 599, 600, 602, 603, 604, 605, 606].

Group IX (residual group)

Leaves coriaceous to succulently coriaceous, ovate or elliptic to oblong,

cuneate or obtuse at base, 7.7—-(12.9)-18.6 cm long, 4—(5.8)—8.6 cm wide,

Variations in the Hoya verticillata complex in Thailand 193

indistinctly acrodromous venation, the lowest pair extending about half

way to apex; veins obscure; covered with dense hairs on abaxial surface;

petiole stout, 0.5—(1.5)—3.2 cm long. Umbel 5—(27)—75 flowered, peduncle

0.3—(3.8)—14.3 cm long; pedicel glabrous to pubescent, 1.3—(2.0)—2.9 cm long.

Sepal ovate, 1.3-(1.9)—-3 mm long, 1.1—(1.6)—2.1 mm wide. Corolla creamy

white, 1.1-(1.4)-1.8 cm diam. Coronal scale ovate to ovate-lanceolate,

2.8—(3.5)—4.4 mm long, 1.5—(1.9)—2.4 mm wide, the outer angle slightly erect

to erect and white, apiculus of inner angle short raised and white to pink.

Pollinium oblong, 0.41—(0.51)-6.43 mm long, 0.16—(0.19)-—0.22 mm wide;

translator short; corpusculums triangular-oblong, 0.14—(0.17)—0.21 mm long,

0.10—(0.12)—0.15 mm wide.

Ecology and distribution: Along stream banks in mixed deciduous to dry

dipterocarp forests at about 100-600 m elevation. Occurs sporadically

throughout northern, north-eastern, eastern, and western to peninsular

Thailand.

Specimens examined: Chiang Mai - Wang Bua Ban Waterfall, Manit Kidyue

299, 300, 301, 412; Lampang - Doi Khun Tan National Park, Manit Kidyue

206, 209, 270, 271,272, 273,274, 275, 270, 277, 278, 279, 380, 410; Nan [Phu

Pieng, Manit Kidyue 388, 389, 390], [Rhinan, Manit Kidyue 398, 399], Loei

[Pla Ba Waterfall, Manit Kidyue 689, 690, 691, 692]; Nong Khai [Than Thong

Waterfall, Manit Kidyue 551, 552, 553, 554, 555, 556, 557, 558, 559, 560, 561,

562, 563, 564, 565, 566, 567, 568]; Udon Thani [Than Ngam Waterfall, Manit

Kidyue 420, 421, 422, 423, 424]; Nakhon Phanom [Tat Kham Waterfall, Manit

Kidyue 493, 494, 495, 496, 497, 498, 499, 500, 501, 502, 503, 504, 505]; Sakon

Nakhon [Phu Phan National Park, Manit Kidyue 427, 228, 429, 430, 431,

432, 433, 434, 435, 436, 437, 438, 439, 440, 441]; Mukdahan [Tat Ton Waterfall,

Manit Kidyue 484, 485, 486, 487, 458, 489, 490, 491, 492|; Ubon Ratchathani

[Soi Sawan Waterfall, Manit Kidyue 540, 541, 542, 543, 544], [Huai Sai Yai

Waterfall, Manit Kidyue 479, 480, 481, 482, 483], |Phu Chongna Yoi National

Barks Mann Nidyve 312,515; 514, 515,510, 517, 518, 519, 520, 521, 522, 523,

524, 525, 526, 527, 528], [Tat Hai Waterfall, Manit Kidyue 529, 530, 531, 532,

533, 534, 535, 536, 537, 538, 539]; Sti Sa Ket [Sam Long Kiat Waterfall, Manit

Kidyue 506, 507, 508, 509, 510, 511); Nakhon Rat Chasima [Phu Wa Kiew

Waterfall, Manit Kidyue 545, 546, 547, 548, 549, 550], [Khao Yai National

Park, Manit Kidyue 362, 363, 364, 365, 366, 367, 368, 369, 370, 371, 372, 373,

374, 375, 376, 377|; Saraburi [Pu Kae, Manit Kidyue 307, 308, 309]; Prachin

Buri [Prachantakham, Manit Kidyue 315,316,317,318]; Sa Kaeo [Pang Sida

Nauomal Park, Mani Kidyues..2, 3, 4, 5,6, 7, 8, 9, 10, 11, 12, 13, 16, 17, 18,

120, 2122, 23, 24, 2), 20, 27, 26, 29, 30); Yak [Thi Lo Su Watertfall, Manit

Kidyue 570|; Kanchanaburi [Sangkhla Buri, Manit Kidyue 447, 448, 449,

450, 451, 452, 453, 454, 455, 456, 457, 458, 458, 460, 461, 462, 463], [Koeng

194 Gard. Bull. Singapore 58 (2) 2007

Kra Wia Waterfall, Manit Kidyue 475, 476], [Pong Ron, Thong Pha Phum,

Manit Kidyue 258, 259, 442, 443, 444, 445, 446], [Ban Thamadua, Thong Pha

Phum, Manit Kidyue 140, 141, 142, 158, ,165, 473, 474], [Sai Yok National

Park, Manit Kidyue 150, 151, 152, 153, 155, 156, 157, 159, 160, 162|; Ranong

[Namtok Ngao National Park, Manit Kidyue 644, 645, 646, 647, 648].

Discussion

Hoya verticillata has long been recognized as a taxonomically confusing

group in the genus. Many authors have discerned differently the taxa within

the species complex. The confusion was partly due to the great variations

exhibited by the plants in their natural habitats. The results from this

present study indicate that nine morphological groups can be recognized

based on leaf shape, form of leaf base, leaf venation, leaf indumentum,

shape of sepal, and shape of corona scale. However, the Hoya verticillata

specimens could not be consistently allocated into a discrete group based

on lamina texture, color of flower and hair. Lamina texture was found to be

slightly discontinuous in its range of variability. This variation appeared to

be related to growing conditions in different habitats (Hill, 1988). Colour

of flowers has been reported to be highly variable in this species (Kiew,

1995). In our study, we found that the colour of corolla varied from white,

creamy white, yellowish white, greenish white, pinkish white to white with

brown, pink and violet at apices. Hair is frequently found on the stem,

corolla (adaxial surface), calyx (abaxial surface) and pedicel. Young stems

and new branches varied from glabrous to pubescent, or earlier pubescent

then becoming glabrous when getting old, but some plants are consistently

glabrous. Hair on pedicel also varied from glabrous to densely pubescent.

In contrast, in some populations hairiness varied from glabrous to densely

pubescent, while the others were only glabrous or pubescent. Hair on the

corolla varied from minutely pubescent (almost glabrous to the naked eye)

to densely pubescent (obvious to the naked eye). All the above characters

(lamina texture, color of flower and hair) possess sufficient variability for us

to conclude that they are not good discriminant characters for distinguishing

taxa of this complex.

Regarding their habitats and distributions, the nine recognized groups of

the complex occur in different floristic regions and habitats throughout the

country. It was found that Group I, II and IV occur in rather humid forests

with medium to low light conditions, while the other Groups usually grow

in dry habitat to some extent. These differences are possibly related in part

to the degree of succulence and size of leaves that allow plants to withstand

Variations in the Hoya verticillata complex in Thailand 195

different light conditions (Forster and Liddle, 1991). Naturally, the members

of the complex occur in a wide range of habitats and geographic features in

Thailand. Anyway, the habitat and distribution of the nine Groups overlap

to such a degree that they are not good characters to use to distinguish taxa

in the complex.

The previous treatments of the genus Hoya and the Hoya verticillata

complex, were based primarily on discrete characters of floral structures

(Rintz, 1978; Hooker, 1983; Forster and Liddle, 1991 and Kiew, 1995). In

this study, we used floral characters in defining the species, while vegetative

characters and their geographical distribution are used to characterize the

varieties. Hence, we considered that the nine groups in the H. verticillata

complex can be treated as either species or varieties. Group I and Group II

of the H. verticillata complex were already treated at species level, due to

their discontinuity in floral and vegetative characters.

Group I corresponds to the previous described species, Hoya rigida Kerr

(Kerr, 1939). The result from this study shows good diagnostic characters to

recognize this species based on leaf shape, venation, sepal shape and ratio of

corolla tube length to sepal length. We agree with Kerr (1939) in separating

this species from H. verticillata; Veldkamp et al. (1995) had already a

suspicion about including this species in H. verticillata s.l.

Group II is somewhat similar to H. verticillata (Vahl) G. Don var. citrina

(Ridl.) Veldkamp, except for its retention of minute hairs, scattered over the

abaxial surface. Furthermore the corpusculum of anthers are oblanceolate-

oblong. Rintz (1978) describes leaves and flowers of H. verticillata var.

citrina and his illustrations show that lower surfaces of leaves have distinctly

dense hairs and triangular-oblong corpusculum. The result from this study

indicated that Group II is clearly distinguishable from the H. verticillata

s.l. Importantly, since it has some diagnostic characters (leaf broad ovate

with cordate base; prominent, 3-5 nerves, extending from base to apex, hairs

minute, scattered on the abaxial surface; corpusculum oblanceolate-oblong),

which did not match any previous described taxa, we consider it should be

treated as a new species. In addition, the fragrance of flowers in Group II

tends to differ from the other groups. We consider this preliminary evidence

suggests a chemical difference that may be used to recognize Group II. At

present, we tentatively treat Group II as a cryptic and undescribed species

within the Hoya verticillata species complex.

Group III and Group IV have conspicuously discrete vegetative characters.

Thus, we would define them at the varietal level. Group III is similar to H.

196 Gard. Bull. Singapore 58 (2) 2007

verticillata var. citrina. They share some common characters: possession of

ovate leaves, the basal main veins extending to apex and the other veins

being conspicuous, but there are still some differences. Group III has

elliptic coronal scales and occurs in lowlands of northern Thailand, while

H. verticillata var. citrina has ovate-lanceolate coronal scales and occurs in

Malaysia, being common on limestone hills.

Group IV is an intermediate form between H. verticillata var. citrina and H.

verticillata var. verticillata. It shares its venation pattern with the var. citrina,

while leaf shape is elliptic-oblong with cuneate base; this character is similar

to the var. verticillata.

Itis evident that Group II] and Group IV do not correspond to the previously

described varieties. So we think that these two groups can be treated as

undescribed varieties of H. verticillata complex. However, these two taxa

are rather rare in occurrence, each being found only at one site.

Group V-IX have some discontinuity in flower and leaf characters. They

cannot be clearly distinguished as distinct species, neither as varieties. Plants

of Group VIII and IX correspond to the previous infraspecific taxon, H.

verticillata var. verticillata. This is characterized by possession of elliptic leaf

shape, cuneate base, obscure veins and the lowest basal pair of main veins

extending about halfway to apex, while the other groups do not best fit to

this taxon. They have broad ovate or elliptic to oblong leaf shape, obtuse

to rounded at base and varied in corona lobe shape. However, they are

more comparable to var. verticillata than the other two previously described

varieties, i.e. H. verticillata var. citrina and var. hendersonii which have ovate

leaves, cordate base, 3 prominent nerves extending from base to apex and

the other veins are also conspicuous (Table 2). So we have treated Groups

V-IX as being variable forms of H. verticillata var. verticillata.

In conclusion, the above reported possibly new cryptic species and maybe

undescribed varieties of the Hoya verticillata complex in Thailand need to

be researched further to reach more definitive taxonomic conclusions. Such

study, however, is not part of this paper.

Acknowledgements

We would like to express our thanks to Brian Swale for his criticism of

the manuscript. In additional, we would like to thank Dr. Somran Suddee

and Sahut Chantanaorrapint for their generous help in plant collection.

Variations in the Hoya verticillata complex in Thailand 197

This work was supported by the TRF/BIOTEC Special Program for

Biodiversity Research and Training grant T_146004 and the Graduate

School, Chulalongkorn University.

References

Forster, P.I. and Liddle, D.J. 1991. Variation in Hoya australis R. Br. Ex Triall

(Asclepiadaceae). Austrobaileya 3: 502-521.

Hooker, J.D. 1883. Flora of British India 4: 52-63.

Kerr, A. F G. 1939. Contributions to the Flora of Siam. Additamentum II.

Kew Bulletin 463.

Kerr, A.EG. 1951. Flora Siamensis Enumerratio 3: 35-42.

Kiew, R.1995.A new Begonia (Begoniaceae), Hoya (Asclepiadaceae) and

Sonerlia (Melastoataceae) from Fraser’s Hill, Peninsular Malaysia.

Sandakania 6: 63-71.

Hill, K.D.1988. A revision of Hoya (Asclepiadaceae) in Australia. Telopea

3(2): 241-255.

Ridley, H.N. 1923. A Flora of Malay Peninsula 2: 393-402.

Rintz,R.E.1978.The Peninsular Malaysian Species of Hoya(Asclepiadaceae).

Malay. Nat. J. 30: 467-522.

Thaithong, O. 1995. The Genus Hoya in Thailand. pp.83—94 . In: R. Kiew (ed.),

The Taxonomy and Phytochemistry of the Asclepiadaceae in Tropical

Asia. Universiti Pertanian, Malaysia.

Veldkamp, J.F, Vandonkelaar, R. and Kloppenburg, R.D. 1995. The Identity

of Sperlingia Vahl (Asclepiadaceae). Blumea 40: 425-428.

Gard. Bull. Singapore 58 (2) 2007

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Gardens’ Bulletin Singapore 58 (2) 2007: 199-232 199

Begonia (Begoniaceae) from Limestone Hills

in the Kuching Division, Sarawak, Borneo,

including nine new species

RUTH KIEW' AND JULIA SANG’

‘Forest Research Institute Malaysia,

Kepong, Malaysia

“Sarawak Herbarium, Kuching, Malaysia

Abstract

Fifteen Begonia species are described from limestone hills in the Kuching

Division, Sarawak, Borneo, of which nine are new: B. andersonii Kiew &

S.Julia, B. burttii Kiew & S.Julia, B. chaiana Kiew & S.Julia, B. corrugata

Kiew & S.Julia, B. kiamfeeii Kiew & S.Julia, B. paoana Kiew & S.Julia, B.

penrissenensis Kiew & S.Julia, B. punchak Kiew & S.Julia and B. serapatensis

Kiew & S.Julia. The distribution of these begonias confirms that the Bau

limestone flora is phytogeographically distinct and shows that the Padawan-

Serian and Penrissen limestone areas also form two distinct phytogeographic

areas and that there are few species shared between the three areas.

Introduction

In common with limestone elsewhere in Sabah and Sarawak, the limestone

hills in the Kuching Division are tower karst formations with sheer cliffs.

They occur as isolated hills and in the Kuching Division are now surrounded

by cultivation. Geologically, limestone in the Kuching Division has been

subdivided into three areas: the Bau, Padawan-Serian and Penrissen

limestone areas (Banda et al., 2004).

The Bau limestone is best known botanically (Kiew et al.,2004) and includes

both the well-known tourist caves, Wind Cave (formerly called Gunung

Lubang Angin) and Fairy Cave (Gunung Kapur). Being closest to Kuching

town, it was explored botanically as early as 1845 when Hugh Low collected

there (Kiew et al., 2004). Other 19" century collectors include G.D. Haviland

(1891-1895) and H.N. Ridley (1893, 1903), who wrote an account of Bornean

begonias (Ridley, 1906).

200 Gard. Bull. Singapore 58 (2)2007

In the 20th century, major collectors include J.A.R. Anderson (1950-1970),

one of whose special interests was the Sarawak limestone flora (Anderson,

1965) and B.L. Burtt (1978), who specialized in the Gesneriaceae. Many

collections were added to the Sarawak Forest Department Herbarium

(SAR) by local staff.

The most recent survey was carried out in 2001-2003 when 20 hills in the

Bau area were inventoried (Kiew et al., 2004), as part of Kuching Limestone

Biodiversity Project by the Sarawak Biodiversity Centre and the ASEAN

Regional Center for Biodiversity Conservation, with the result that six

begonias, including one new species, Begonia lailana Kiew & Geri, were

documented from the Bau limestone (Kiew & Geri, 2003).

In contrast, the botany, including begonias, of the Padawan-Serian and

Penrissen limestone remained unstudied. The Padawan-Serian area was

better collected before our survey with begonias collected from the Tebedu

area (Teng Bekap, Gua Baju and Gunung Mentawa), Bukit Pait, Bukit

Angob, Gunung Bra’ang (the highest limestone hill in the Kuching Division

reaching 729 m and first climbed by Haviland), Gunung Manok, Gunung

Mas, Gunung Sebakap, Gunung Sebengkam, Gunung Selabor, Gunung

Seburan and 21° Mile Serian Road.

The Penrissen area was even more poorly known previous to our study

with only Bukit Serapat (also known as 13th Mile Kuching-Simanggan

Road or 16th Mile Penrissen Road), Gunung Bah (G. Bar on the herbarium

specimens) and G. Berloban being represented by herbarium specimens.

Our exploration of these hills, as well as of Gunung Burau and Gunung

Rimo, show that the Penrissen area is particularly high in endemics.

This study was undertaken to document and describe the begonia species

found on limestone in the Kuching Division (nine new species are described

below) and in particular to map their distribution. It is likely that as more

hills are explored in the Padawan-Serian and Penrissen areas further new

species will come to light.

Distribution of Begonias in the Kuching Division

Among species of the limestone flora in Borneo, begonias exhibit one

of the highest levels of endemism with many being restricted to a single

Begonia from limestone hill in the Kuching Division 201

110°00°E _ _110°1S’E

1°30°N

Beratok

| 1°00'N

\ Tapah

Legend

co Limestone area

\ — Road

1°00°N

| 1°15'N

INDONESIA

0°85’N

| 0°85'N

INDONESIA

110°00°E ; 110°15E 110°19°E

Figure 1. Limestone hills in the Kuching Division, Sarawak.

202 Gard. Bull. Singapore 58 (2)2007

hill (Kiew, 2001; Pearce, 2003). The distribution of these narrow endemics

is therefore useful in delineating phytogeographic areas (Kiew, 2001).

Indeed, Burtt (1978) did just this for Monophyllaea (Gesneriaceae) in the

Kuching Division and was the first to recognise that there were in fact two

distinct phytogeographic areas: the Bau hills versus the rest based on the

distribution of Monophyllaea species and varieties. This distribution pattern

is also exhibited by other limestone endemics, such as species of Begonia

and Schismatoglottis (Araceae), and led Kiew et al. (2004) to distinguish the

Bau limestone as separate from the Padawan-Serian limestone. Our study

not only confirms that the Bau limestone is a distinct phytogeographic area

but also shows for the first time that the Penrissen limestone is distinct from

the Padawan-Serian limestone in which it had previously been included

(Burtt, 1978, and Kiew et al., 2004) so that in fact there are three distinct

limestone areas in the Kuching Division.

All three areas are equally diverse with six, seven and six begonia species

occurring in the Bau, Padawan-Serian and Penrissen areas, respectively.

Each area is home to its own endemic species: two in the Bau area (Begonia

congesta Ridl. and B. lailana Kiew & Geri); three in the Padawan-Serian

area (B. andersonii Kiew & S.Julia, B. chaiana Kiew & S.Julia and B. paoana

Kiew & S.Julia); and four in the Penrissen area (B. burttii Kiew & S.Julia, B.

kiamfeei Kiew & S.Julia, B. penrissenensis Kiew & S.Julia, B. punchak Kiew

& S.Julia and B. serapatensis Kiew & S.Julia).

In addition, although the Penrissen area geographically lies between those

of the Bau and Padawan-Serian areas, it shares fewer species with the two

other areas: only B. speluncae Ridl. occurs in all three areas; the Padawan-

Serian and Bau areas share three species (B. calcarea Ridl., B. pendula Rid.

and B. rubida Ridl.), while only B. corrugata Kiew & S.Julia is found in both

the Penrissen and Padawan-Serian areas.

Key to Limestone Begonia Species in the Kuching Division

la... Stems, erect. and..cane-like 2... eee eee 2

lb. Stems rhizomatous or cecepine ce... eee ee

2a. Leaves with veins prominently forked and narrowly diverging. Fruits

ODI ONG ao ss cs tice dbeien snapbican wansileae dhs one cde eee cee

2b. Leaves with veins widely diverging where they fork. Fruits wider than

long or much wider distally o0:..:2 vei. anus tees eee ee ee

Begonia from limestone hill in the Kuching Division 203

3a.

3b.

4a.

Ab.

Sak

Sb.

6a.

6b.

Va.

7b.

8a.

Sb.

9a.

9b.

Leaves glabrous. Female flowers and fruits in a compact cluster; pedicels

4-5 mm long ............ ea 0k .. 5. B. congesta

Leaves hairy. Beale ewer ae fruits oe spaced on aan inflorescence;

pedicels 7-13 mm long... sua eS iaaceon Riis O2id: COMMNBALA

Leaves lanceolate or oblanceolate; petioles straight or at a slight angle

iN ee are PN eee ery ernie prccapmrcana'-nacodtacuns els ala 5

Leaves ovate; petioles at a pronounced angle with the midrib .......... 7

Laminas lanceolate, 8-12 cm long. Inflorescences 6-9.5 cm long. Female

RON TSIWITNCS Me past: Teel Cases, AUS allie nc. 4. B. chaiana

Laminas oblanceolate, 12—23 cm long. Inflorescences 10-25 cm long.

emiale Mowers LeMAlsy. 6 A Sen weaken keel Geaned ease devia ce cpanesdeceaees 6

Laminas 12-17 cm long. Inflorescences 10-16 cm long. Female flowers

aig RV Ree AWM Ps Peo GPT Raiedtvicr se eal JAE AL, 8. B. lailana

Laminas 15-24 cm long. Inflorescences 12—25 cm long. Female flowers

in 2 pairs (one pair in the leaf axil and one above) .........9. B. paoana

Basal lobes up to a third of the lamina length. Inflorescences shorter than

Bale aics AV co SeUt O70 ia Com eying et i d5 ie he A alae 8

Basal lobes about half the lamina length. Inflorescences longer than

ETE SORE nn) a) 0) oa: See 0. ee oe ee ee ee 9

Leaves with petioles to 4.5 cm on the lower leaves and blades 9.5—12

by 5-6 cm. Inflorescences with female flowers at the base. Male flowers

with 2 tepals .. eer. Boe oes . 11. B. penrissenensis

Leaves with peice - 2. 5 cm m long and Aedes to 7 by 4 cm. Female

flowers in separate axils from the male inflorescences. Male flowers

WW A) Pe RU ae GAS re Eee ae OL a ee eae 12. B. punchak

Laminas more than 9 x 5 cm. Flowers white. Fruit wings 6-9 mm wide

re ry ae te a Re etm Ss 6 De BE y ia ucaees 14. B. serapatensis

Laminas up to 8 x 5.5 cm. Flowers ruby red. Fruit wings 4-5 mm wide

Sie ee owt holes Voit eh 58 ast ls SRL PS Pela « 22h 13. B. rubida

10a. Stems thin and creeping. Leaves widely spaced, longer than wide .... 11

10b. Stems thick and rhizomatous. Leaves tufted, as wide as long ........... 12

iia,

Leaves hairy, plain green, ovate, basal lobes 24.5 cm long. Inflorescences

with several female flowers and fruits. Flowers white. Male flowers with

204 Gard. Bull. Singapore 58 (2)2007

4 tepals .. ree 13s . 7. B. kiamfeeii

11b. Leaves secs an i pease ote lobes 1.2-2 cm long.

Inflorescences with one female flower or fruit. Flowers deep pink or

red: Maleflowers with 2tepals z.c..c.52R eee see ee 10. B. pendula

12a: Leaves: peltate qu.i.un.cencewep ear ae ee See 13

12b;, Leal. Dasesicordate..; .. scasvasie are a ae ee 14

13a. Leaf strongly oblique, laminas broader than long, 1.2—3.5(-12) cm long,

apex nounded..<c hia eee Suir . 15. B. speluncae

13b. Leaf scarcely aan e. (ates ieaeem pe ey 4. 5-7 cm long, apex

ACUMTIINALC suns ae Ba eo eee 2. B. burttii

14a. Laminas round; petioles shorter than the laminas, 2-10 cm long.

Inflorescences among the leaves, erect and longer than the petioles.

Blowers pink onwwhite vonssact. ude, aaa eee ee eee 1. B. andersonii

14b. Laminas ovate; petioles longer than the laminas, 14-35 cm long.

Inflorescences developing on the bare prostrate rhizome, shorter than

the petioles. Flowers peachy orange’... 2a2..0. 32.) sa eee

Begonia specimens from the Bau limestone are listed in Kiew & Geri (2003).

Limestone Begonias in the Kuching Division

1. Begonia andersonii Kiew & S. Julia, sp. nov.

Sect: Reichenheimia

A Begonia speluncae Ridl. laminis latioribus 8-15 latis (nec usque 5(-12) cm

latis), petiolis lanatis (nec glabris) et foliis basi cordatis (nec peltatis) differt.

Typus: Borneo, Sarawak, Kuching Division, Teng Bekap, Gua Baju, 8 August

2004, Kiew, Julia & Tan S 93269 (holo, SAR; iso, E, KEP, SING). Figure 2

Rosette begonia; stems rhizomatous, creeping and rooting at nodes, up to 7

cm long, woody, hairy, not branched, stout, 6-7 mm thick; without a tuber.

Stipules densely hairy, narrowly lanceolate, ca 2 x 5 mm, pale green to pale

brown, margin entire, apex entire, persistent. Leaves tufted and splayed

out against the rock face; petiole woolly with pale brown hairs, (2—)5(—10)

cm long, slightly succulent, reddish in young leaves, reddish brown in older

leaves; lamina oblique, young leaves bronzy, adult leaves plain pale green

above with slightly paler green veins, beneath paler and scintillating with

brown veins, thinly succulent, drying papery, slightly asymmetric, orbicular,

Begonia from limestone hill in the Kuching Division 205

4mm 2mm 2mm 4mm

Figure 2. Begonia andersonii Kiew & S.Julia

A. habit; B. lower leaf surface near the midrib; C. leaf margin; D. male bud: E. open male

flower; F. stamen cluster; G. stamens; H. female flower; I. open male flower; J. styles and stig-

mas; K. TS ovary; L. seed; M. fruit. (A from CWL 1308; B-M from S 93269).

206 Gard. Bull. Singapore 58 (2)2007

6.2—11.5 x 8.3-15.75 cm, broad side 4.5—6.5(—9.5) cm wide, base deeply cordate

sometimes overlapping, basal lobes 0.75—2(—3.5) cm, margin hairy, crenate,

apex rounded or shortly acute; venation palmate with 3--4 pairs of veins

branching less than half way to the margin, midrib often branching more or

less dichotomously, veins slightly impressed above, beneath prominent and

densely hairy, hairs brown. Inflorescences terminal, pale red, glabrous, erect,

dichasial cyme, longer than the leaves, 9-31 cm long, peduncle 2.5—-9 cm

long, branches 2 or more, 2.5—9 cm long, male flowers many, female flowers

2-3, protogynous. Bracts obovate, pale green, with long dense hairs, 1-2 mm

long, margin entire; bracteoles similar but smaller. Male flowers with a pale

pink pedicel 15 mm long; tepals 4, pale pink, margin entire, apex rounded,

glabrous, outer two broadly rounded, 6 x 4-6 mm, inner two narrowly oval, 4

x 1.5-2 mm; stamens many, cluster globose, ca 2 mm diam., sessile; filaments

1-2 mm long; anthers yellow, narrowly oblong, 1 mm long, apex slightly

notched, opening by lateral slits. Female flowers with a pedicel ca 20 mm

long; ovary ca 3 x 4 mm, reddish, wings 3, more or less equal, ca 3 mm wide,

locules 3, placenta one per locule; tepals 4, pale pink, glabrous, margin entire,

apex rounded, outer two rounded, ca 3 x 3 mm, inner two smaller 2 x 1 mm;

styles 3, styles and stigma yellow, ca 1 mm long. Fruits pendent on a fine stalk

ca 9-11 mm long; capsule 4-5 x 11-13 mm, glabrous, wings 3, equal, thinly

fibrous, 3-4 mm wide, splitting between the locules and wings. Seeds barrel-

shaped, ca 0.5 mm long, collar cells less than half the seed length.

Distribution: Endemic in Kuching Division, Sarawak; known only from the

Padawan-Serian limestone.

Habitat: Locally common, it is restricted to limestone and grows on dry ver-

tical limestone cliffs in light shade.

Specimens examined: Padawan-Serian limestone - Teng Bekap [Tiang Bekap],

Anderson § 12345 (SAR); Gunung Mentawa, Burtt B 8112 (SAR), Chew

CWL 656 (SAR), CWL 1308 (SAR); Kampung Gayu, Mohizah & Jegong S

66838 (SAR); 21st Mile Serian Road, Anderson S$ 20999 (SAR), Murthy &

Chai S 24699 (SAR); Tebedu, Bukit Kajing, Abang Mohtar S 48287 (SAR).

Notes: Begonia andersonii, like B. speluncae, belongs to sect. Reichenheimia

in its ovary having three locules, each locule with one placenta. They are also

similar in their circular leaves and female flowers that are very like the male

flowers in having four tepals, the outer pair large and round and the inner

pair narrowly oblong. Otherwise, they are different in size - B. speluncae is a

small plant with leaves 1.4-5.5 cm across (exceptionally up to 12 cm wide),

which are almost glabrous (the leaf margin is minutely ciliate) and its leaves

are peltate (not cordate as described by Ridley, 1906).

Begonia from limestone hill in the Kuching Division 207

Begonia andersonii has a very local distribution centred on Teng

Bekap. Interestingly, although B. speluncae Ridl. has a wide distribution in

the Kuching Division, and occupies the same habitat as B. andersonii, these

two species are not found growing on the same hill.

It is named in honour of J.A.R. Anderson, who pioneered the

exploration of the limestone flora in Sarawak (Anderson, 1965). His

collections include this species.

2. Begonia burttii Kiew & S. Julia, sp. nov.

Sect: Reichenheimia

A Begonia speluncae Ridl. laminis foliorum majoribus longitudine latitudinem

aequanti 4.5-6.8 x 5.6-6.4 cm (nec latioribus quam longioribus, 1.2—5(-10)

x 1.4-5.7(-12) cm) apice acuminatis (nec rotundatis) differt. Typus: Borneo,

Sarawak, Kuching Division, Gunung Serapat, 8 August 2004, Kiew, Julia &

Tan §S 93266 (holo, SAR; iso, E, KEP, SING). Figure 3

Rosette begonia; stems rhizomatous, creeping and rooting at nodes, up to

4 cm long, succulent, hairy, not branched, stout, ca 4 mm thick; without a

tuber. Stipules reddish, hairy, narrowly lanceolate, 3-4 x 1-1.5 mm, margin

entire, apex setose, caducous. Leaves tufted, oblique; petiole greenish,

glabrous, 4.7—9 cm long, succulent, slightly glaucous; lamina of young leaves

rosy red above and beneath, mature leaves plain matt, dark green above

with paler green veins, beneath reddish green or rosy red, thickly succulent,

drying papery and with the tertiary veins slightly raised, slightly asymmetric,

peltate, broadly oval, sometimes angular, 4.5—-6.8 x 5.6—6.4 cm, broad side c.

3.7 cm wide, base rounded, 1.5—1.8 cm from junction with the petiole to the

base of the lamina, margin fringed by short hairs, entire to minutely dentate,

apex shortly acute; venation palmate with ca 4 pairs, veins branching half

to one third of the way to the margin, plane above, beneath prominent and

glabrous. Inflorescences terminal, red, glabrous, erect, dichasial cyme, longer

than the leaves, ca 9.5-15 cm long, peduncle 9-13 cm long, branches two

0.5—2 cm long, male flowers many, female flowers 2 or more, protandrous.

Bracts caducous. Male flowers with a white pedicel 15-17 mm long; tepals

4, red in bud, pale pink when open, margin entire, apex rounded, glabrous,

outer two broadly oval to rotund, 4-7 x 5-6 mm, inner two narrowly oval, 3-

4x 1-1.5 mm; stamens many, cluster fan-shaped, stalked; filaments c. 2 mm

long; anthers pale yellow, oblanceolate, 2 mm long, apex slightly notched,

opening by lateral slits. Female flowers with a pedicel ca 5 mm long; ovary

ca 9 x 12 mm, locules light red and wings pale green, wings 3, more or less

equal, ca 3 mm wide, locules 3, placenta one per locule; tepals 5, rosy pink,

glabrous, margin entire, apex rounded, outer two obovate, ca 6 x 3 mm, inner

208 Gard. Bull. Singapore 58 (2)2007

Imm

2mm 5mm

Figure 3. Begonia burttii Kiew & S.Julia

A. habit; B. lower leaf surface; C. male bud: D. open male flower; E. stamen cluster; F. stamens;

G. female flower; H. open male flower; I & J. styles and stigmas; K. fruit; L. TS ovary; M. seed.

(All from S 93266).

Begonia from limestone hill in the Kuching Division 209

three smaller ca 3 x 1 mm; styles 3, styles and stigmas pale yellow, ca 2 mm

long. Fruits pendent on a fine stalk 4-8 mm long; capsule 6-8(-—10) x 7-10(-

13) mm, glabrous, wings 3, equal, rounded, thinly fibrous, 2—3(—5) mm wide,

splitting between the locules and wings. Seeds barrel-shaped, ca 0.3--0.4 mm

long, collar cells ca half the seed length.

Distribution: Endemic in the Kuching Division, Sarawak; known only from

a single hill in the Penrissen limestone.

Habitat: Restricted to limestone, it grows on deeply shaded vertical limestone

rock faces near the base of the hill.

Specimen examined: Penrissen limestone - Bukit Serapat, 13th Mile Kuching-

Simanggan Road, Burtt & Martin B 4746 (SAR).

Notes: This is the second peltate begonia to be recorded for the Penrissen

limestone. The other is the widespread Begonia speluncae collected from

Gunung Burau. Compared with B. speluncae, B. burttii is a larger plant,

has a slightly angular lamina with a distinct midrib and an acuminate apex,

which is longer or as broad as long, whereas laminas of B. speluncae are

broader than long and rounded at the apex and the veins appear to branch

dichotomously so that there is no distinct midrib. In addition, B. burttii has

female flowers with five tepals, while those of B. speluncae have four.

This species is as yet known from a single hill. It is named in honour of

B.L. Burtt, who was one of the early explorers of the Kuching limestone and

who first collected this species, in recognition of his work on Monophyllaea

(Gesneriaceae) that first drew attention to the distinction between the

limestone floras of the Bau and Padawan-Serian/Penrissen areas.

3. Begonia calcarea Rid.

J. Str. Br. Roy. As. Soc. 46 (1906) 260; Kiew & Geri, Gard. Bull. Singapore 55

(2003) 115.

Distribution: Endemic in the Kuching Division, Sarawak; known only

from the Bau (Gunung Kawa, G. Lanyang, G. Tabai) and Padawan-Serian

limestone.

Specimens examined: Padawan-Serian Limestone — Gunung Angob,

Anderson S 27513 (SAR); Gunung Bra’ang, Haviland s.n. (n.v.); Gunung

Manok, Burtt B8134 (SAR).

Notes: Although quite widespread on the Kuching limestone, it is a rare

begonia that on the Bau limestone grows in deeply shaded forest on the

210 Gard. Bull. Singapore 58 (2)2007

base of the hills (Kiew & Geri, 2003). Ridley (1906) first described this

species from Gunung Bra’ang, from where Haviland had collected it from

the summit.

It is an outstandingly decorative begonia - its petioles are densely

clothed in long magenta hairs and its bunches of flowers produced from the

prostrate rhizome are peach-coloured.

4. Begonia chaiana Kiew & S. Julia, sp. nov.

Sect. Petermannia

A Begonia subisensis K.G. Pearce tepalis femineis 3 (nec 5), fructibus pedicellis

brevibus usque 7 mm longis (nec usque 30 mm) et alis angustioribus c.6 mm

latis (nec 10-11 mm) differt. Typus: Borneo, Sarawak, Kuching Division,

Gunung Mentawa, Kiew, Julia & Pearce S93270, 8 August 2004 (holo, SAR;

iso, E, KEP, L, SING). Figure 4

Cane-like begonia, whole plant glabrous; stems erect, 30-50 cm long, young

stem reddish and succulent, mature stem green and woody, slender, 3-4

mm thick, nodes swollen, branching to form a spreading crown; without

a tuber. Stipules narrowly lanceolate, ca 5 x 2 mm, reddish, margin entire,

apex glabrous, persistent. Leaves distant, 24.5 cm apart, slightly oblique;

petiole 1.5—2.2 cm long, succulent, pale red; lamina plain glossy dark green

above, beneath deep red with green or deep red veins, succulent, drying

thinly papery, asymmetric, lanceolate, 8-12 x 2.2-3.5 cm, broad side 1.22.5

cm wide, base unequal, basal lobes rounded 5-10 mm long, margin dentate,

scalloped between the teeth, apex caudate; venation palmate-pinnate with 1

pair at the base and 2 pairs along the margin branching towards the margin,

and with 1 vein in the basal lobe, veins slightly impressed above in life,

raised when dry, beneath prominent. Inflorescences axillary, pale red, erect,

cymose panicle, shorter than the leaves, 6—9.5 cm long, peduncle 2—3 cm

long, branches two 2.5—4 cm long, male flowers many, female flowers | at

the base, protogynous. Bracts obovate 1-2 mm long, margin entire, caducous;

bracteoles similar but smaller. Male flowers with a pale red pedicel 5—7 mm

long; tepals 2, in bud green or tinged red outside, greenish white or pale

reddish green when open, broadly oval, 5 x 4-6 mm, margin entire, apex

rounded, glabrous; stamens many, cluster flattened laterally, ca 2.5 mm

across, stalked; filaments c. 1 mm long; anthers pale yellow, almost rounded,

1 mm long, apex slightly notched, opening by lateral slits. Female flowers

with a pedicel ca 6 mm long; ovary deep red, 5—13 x 6-15 mm, wings 3, more

or less equal, ca 7 mm wide, locules 3, placentas 2 per locule; tepals 3, deep

red, isomorphic, ca 9 x 6 mm, obovate, glabrous, margin entire, apex rounded;

styles 3, styles and stigma pale yellow, ca 4 mm long. Fruits pendent on a stiff

Begonia from limestone hill in the Kuching Division 2A

lcm

Figure 4. Begonia chaiana Kiew & S.Julia

A. habit; B. male bud: C. open male flower; D. stamen cluster; E. stamen; F. female flower; G

& H. styles and stigmas; I.TS ovary; J. fruit. (A-E from S 27424; F-J from S 93270).

212 Gard. Bull. Singapore 58 (2)2007

stalk ca 7 mm long; capsule ca 15 x 20 mm, hairless, locules 2 (one locule not

developing), wings 3, subequal, the longer ca 8 mm wide and the shorter two

ca 6 mm wide, thinly fibrous, splitting between the locules and wings.

Distribution: Endemic in Kuching Division, Sarawak; known from two

limestone hills in the Padawan-Serian area.

Habitat: It grows on limestone rocks or jagged cliffs deep in shade below the

tree canopy.

Specimens examined: Padawan-Serian limestone - Bukit Pait, Erwin & Paul

Chai S 27424 (SAR), Julia & Kiew S 95686 (KEP, SAR); Gunung Mentawa,

Anderson 12346 (SAR), Burtt B 8111(SAR).

Notes: This is a dainty cane-like begonia with slender stems and rather

narrow, glossy dark green leaves that are strikingly deep red underneath.

In habit, it is similar to Begonia subisensis but differs from this species not

only in its female flowers having three tepals (those of B. subisensis have 5)

but the fruit is also quite different in having a short stiff stalk (whereas that

of B. subisensis is flexuose, dangling and up to 30 mm long) and the capsule

is both smaller and has relatively narrower wings (the fruits of B. swbisensis

are about 14 x 23 mm with wings 10-11.5 mm wide).

It shares the inflorescence type (cymose panicle), two tepals in the

male flower and the capsule shape with slightly unequal wings with Begonia

lailana but differs in its smaller size (Begonia lailana grows to 1 m tall, has

larger laminas 12.5-17 x 8-10 cm and longer panicles 10-16 cm long) and

tepal number in the female flower (5 in B. lailana).

In its cane-like habit, its protogynous inflorescences and female

flowers with three locules each with two placentas, it conforms to sect.

Petermannia. However, its fruit wings are slightly unequal and one locule

does not develop in the fruit and is empty of placentas and seeds. In addition,

the slightly longer wing is not thickened (as it is in sect. Platycentrum) and

apart from size 1s identical to the other two wings.

It is an extremely local species known from just two localities.

Itisnamed in honour of Dr Paul Chai Piang Kong, the first Sarawakian

appointed as botanist (1971-1986) in the Sarawak Forest Department.

5. Begonia congesta Ridl.

J. Str. Br. Roy. As. Soc. 46 (1906) 253; Kiew & Geri, Gard. Bull. Singapore 55

(2003) 116.

Distribution: Endemic in the Kuching Division, Sarawak; restricted to the

Bau limestone (Bidi, Fairy Cave (Bukit Kapur), Bukit Krian, G. Batu, G.

Begonia from limestone hill in the Kuching Division 213

Doya, G. Jebong, G. Kawa, G. Krian, G. lanyang, G. Pambur, G. Ropih, G.

Seburan, G. Tai Ton and G. Tongga).

Habitat: In shade on limestone boulders or on damp rock-faces near the cliff

base.

Notes: Begonia congesta belongs to a group of limestone begonias that are

characteristic in their relatively thick stems, often swollen at the nodes, and

their leaves that have narrowly bifurcating veins that are impressed above

giving the lamina a corrugated surface. It is confined to the Bau limestone

where it is common and widespread.

6. Begonia corrugata Kiew & S. Julia, sp. nov.

Section Petermannia.

A Begonia congesta Ridl. foliis pubescentibus (nec glabris) et floribus femineis

remotis pedicellis longioribus 7-13 mm longis (nec 4—5 mm) differt. Typus:

Borneo, Sarawak, Gunung Manok, 6 Sept 2005, Julia & Kiew S 95689 (holo,

SAR; iso, E, KEP, L, SING).

Cane-like begonia, stems greenish or reddish brown, woody, matted

ferrugineous, sometimes bristly with translucent/white hairs 3-4 mm long,

particularly dense on the uppermost internodes, erect, little branched, to 60

cm tall but flowering at ca 10 cm tall, 6-7 mm thick, nodes swollen with a

conspicuous stipular scar, internodes 2-6.5 cm long; without a tuber. Stipules

narrowly triangular, 7-20 x 4-15 mm, pale green, glabrous, margin entire,

apex setose, caducous. Leaves oblique, distant, pendent and held flat against

the rock surface, petioles (2.5-)4.5-6-5(-7.5) cm long, glabrous and glossy,

sometimes densely bristly and bristles 2-3 mm long, narrowly grooved

above; laminas plain deep green above, sometimes with a bluish tinge,

paler beneath with dense translucent erect bristly hairs 1-2 mm long on

the upper surface; thin in life, papery when dry, asymmetric, broadly ovate,

14—22(-25) x 9-16 cm, broad side 7-11 cm wide, base unequally cordate,

basal lobes rounded, not overlapping, the larger 1.5—-4 cm long, margin

ciliate usually almost entire with minute teeth, sometimes serrate, apex

shortly acuminate, acumen to 2.25 cm long; venation palmate-pinnate with

(1-)2 basal pairs of veins, with 3-5 pairs along the midrib and 3 veins in the

basal lobes, veins narrowly parallel and deeply impressed above giving a

corrugate appearance, branching once or twice before the margin, beneath

prominent, in young leaves red, in mature leaves red and minutely pubescent,

sometimes brown and ferrugineous with hairs 1-2 mm long, reticulation of

tertiary veins prominent on both surface in dried leaves. Inflorescences light

green, densely bristly, bristles ca 3 mm long, racemose, erect, shorter than

214 Gard. Bull. Singapore 58 (2)2007

the leaves, 13.5—-23 cm long, protogynous, female flowers produced singly

in the lower part of the inflorescence with up to 5 female flowers ca 3.5—5

cm apart, above branches distant and bearing 2—3 cymules of male flowers

with one male flower opening at a time. Bracts green, 15-23 x 10-16 mm,

minutely hairy especially along the raised midrib, margin entire; bracteoles

green, ca 8 x 6 mm, decreasing in size towards the apex. Male flowers with

a light green pedicel ca 4-5 mm long with dense glandular hairs, tepals 2,

greenish yellow, broadly oval, margin entire and recurved, apex rounded,

on the outer surface densely hairy with short glandular hairs, stamen cluster

globose, ca 2-3 mm across, sessile, stamens 40-50, filament ca 0.75 mm long,

anther light yellow, broadly oblanceolate, ca 0.75 mm long, apex slightly

emarginate, opening by lateral slits. Female flowers with reddish or pale

green pedicel and ovary densely covered in short glandular hairs, pedicel

4-10 mm long, ovary oblong 15-19 x 10-12 mm, locules 3, placentas two per

locule, wings subequal 2.5-4.5 mm wide, tepals (4 or) 5, ovate, 10-12 x 5-7

mm, toothed towards the apex, teeth tipped by a glandular hair, apex acute,

outside glabrous or with dense glandular hairs, greenish yellow or whitish

pink, style and stigma greenish yellow ca 4 mm long, styles 3, bifid, stigma a

papillose spiral band. Fruit pendent on a stiff, minutely hairy pedicel (7-)10-

13 mm long, capsules 15—20 x 12-15 mm, locules 3, wings 3, subequal, thin,

longer wing 4-5 mm wide, narrower wing ca 3 mm wide, splitting between

the ovules and wing, stigma caducous. Seeds barrel-shaped, brown, ca 0. 25

mm long, collar cells ca half the seed length.

Distribution: Endemic in the Kuching Division, Sarawak; known only from

the Padawan-Serian and Penrissen limestone.

Habitat: On steep rock-strewn slopes and on limestone screes in shade under

the tree canopy.

Specimens examined: Padawan-Serian limestone - Bukit Angob, Anderson

S 27493 (SAR, SING); Bukit Paya Payung, Tebedu, Jamree et al. S 73270

(SAR); Gunung Manok, Burtt B 8133 (SAR); Gunung Mentawa, Burtt B

8109 (SAR); Gunung Sabu, Jamree et al. S 75866 (SAR); Gunung Sebakab,

Jamree et al. S75804 (SAR); Gunung Sebengkam, Jamree et al. S75896

(SAR). Penrissen limestone - Gunung Bah, Julia et al. S 95699 (E, KEP, L,

SAR, SING); 16th Mile Penrissen Road, Anderson S 15274 (SAR).

Notes: Similar to Begonia congesta in habit and in its corrugated leaves and in

its small male flowers with two tepals that are hairy outside, like B. congesta

it belongs to sect. Petermannia on account of its three-locular ovary, each

locule of which has a bilamellate placenta. It differs from B. congesta in its

densely hairy leaf surface and inflorescences that bear well-spaced single

Begonia from limestone hill in the Kuching Division DAS

female flowers with long pedicels in the lower part of the inflorescences. (B.

congesta has two pairs of female flowers with short 4-7 mm-long pedicels

that are congested (2-3 mm apart) at the base of the inflorescence).

It is named for its characteristic corrugated leaf surface.

7. Begonia kiamfeei Kiew & S. Julia, sp. nov.

Section: Petermannia

A Begonia pendula Ridl. foliis dense hirsutis (nec glabris) et tepalis masculis

4 (nec 2) differt. Typus: Borneo, Sarawak, Kuching Division, Gunung Bah,

Julia, Kiew & Leong S 95698, 17 Nov 2005 (holo, SAR; iso, E, KEP, L, SING).

Figure 5

Creeping begonia rooting at the nodes, climbing vertically up rocks and

cliffs. Stems slender, reddish, unbranched, 23-50 cm long, internodes 2.5—4.5

cm long, succulent and in life ca 3 mm thick; without a tuber. Indumentum

of uniseriate, translucent or white hairs, scattered and 3 mm long on the

stems and petioles; 3-4 mm long on upper and lower surfaces of veins

and lamina margin. Stipules broadly ovate, 8-13 x 4-6 mm, membranous,

reddish, margin entire, apex setose, rapidly decaying on the plant. Leaves

alternate, distant, pendent; petioles reddish, succulent, 2.5—-6 cm long, terete;

laminas oblique, young leaves pale green, adult leaves mid-green with pale

green veins, paler beneath, succulent, drying papery, asymmetric, ovate, 6.5—

8.5 x 5-8.5 cm, broad side 3.5—5.5 cm wide, base cordate, basal lobes 2—4.5

cm long, margin minutely dentate, teeth tipped by a hair, apex acuminate,

acumen 1-—1.7 cm long; venation palmate with 2 pairs of veins at the base,

1 pair along the midrib and 2 veins in the basal lobe, branching towards

the margin, midrib and veins impressed above, slightly prominent beneath.

Inflorescences axillary, erect, cymose panicle, pale green, sparsely hairy,

13-23 cm long, longer than the leaves, peduncle 6.5-10 cm long, branches

10-15 mm long, male flowers many, female flowers | per branch with up to

6 branches bearing female flowers, protogynous. Bracteole pairs subtending

the male flowers white, lanceolate, 4-6 x 3-4 mm, glabrous, margin entire.

Male flowers with pedicels 6-10 mm long; tepals 4, pure white, margin

entire, apex rounded, glabrous, outer two obovate, 6.5-8 x 6-7 mm, inner

two narrowly oblong, 5—7 x 2-3 mm; stamens 26-30, cluster globose, 2-3 mm

diam., sessile; filaments ca 0.5 mm long; anthers yellow, obovate, 0.75—1 mm

long, apex emarginate, opening by lateral slits. Female flowers with white

pedicels ca 8 mm long; ovary white, ca 7 x 14 mm, wings 3, more or less

equal, ca 5 mm wide, locules 3, placentas two per locule; tepals 4 or 5, white,

glabrous margin entire, apex acute, outer ovate, ca 4 x 2 mm, inner smaller 3

x 2 mm; styles 3, yellow, ca 2 mm long, joined for ca half its length, ultimate

216 Gard. Bull. Singapore 58 (2)2007

SS WN

Figure 5. Begonia kiamfeei Kiew & S.Julia

A. habit; B. leaf surface (a) lower side, (b) upper side; C. male bud: D. male flower; E. stamen

cluster; F. stamens; G. female flower; H & I. styles and stigmas; J. TS ovary; K. seed; L. fruit.

(All from S 95698).

Begonia from limestone hill in the Kuching Division PAG

branches broadly U-shaped. Fruits pendent on fine stalks ca 6-13 mm long;

capsules 6-8 x 13-15 mm, glabrous, wings 3, equal, thinly fibrous, (3—)5-6

mm wide, splitting between the locules and wings. Seeds barrel-shaped, ca

0.5 mm long, collar cells ca half the seed length.

Distribution: Endemic in the Kuching Division, Sarawak; on Penrissen

limestone, known from a single hill.

Habitat: Growing directly on limestone rocks where there is a deep moss

layer in light shade.

Notes: The very pale green leaves and pure white flowers make it a striking

begonia. (Most Bornean begonias have pink flowers). It grows on shaded

damp mossy vertical cliff faces, the same micro habitat as Begonia pendula

with which it shares the creeping habit with slender stems that root at the

node. However, it is very different from B. pendula that is glabrous, has

variegated leaves, deep pink or red flowers and male flowers with two tepals.

It has been collected from a single hill.

It is named for Leong Kiam Fee, botanist with the Singapore Botanic

Gardens, and one of the discoverers of this new species

8. Begonia lailana Kiew & Geri

Gard. Bull. Singapore 55 (2003) 117.

Distribution: Endemic in the Kuching Division, Sarawak; known only from

the Bau limestone (Gunung Aup, G. Batu, G. Doya, G. Kawa, G. Lanyang, G.

Podam, G. Poing, G. Tabai).

Habitat: On soil at the base of limestone hills, in light shade.

Notes: It is a cane-like begonia with large, glossy green leaves sometimes with

scattered dark red bristles on the upper surface. Its attractive red, toothed

tepals in the female flowers are unusual among Bornean begonias.

9. Begonia paoana Kiew & S. Julia, sp. nov.

Sect. Petermannia.

A Begonia lailana Kiew & Geri laminis longioribus 17-23 cm longis (nec

12-17 cm) inflorescentis longioribus 12-25 cm longis (nec 10-16 cm longis)

et paribus florum femineorum 2 (nec 1) differt. Typus: Borneo, Sarawak,

Gunung Rimo, Julia, Kiew & Geri $91390, 29 April 2005 (holo, SAR; iso,

KEE E. L.SING). Figure 6

218 Gard. Bull. Singapore 58 (2)2007

4mm 4mm 4mm Smm

Figure 6. Begonia paoana Kiew & S.Julia

A. habit; B. lower leaf surface; C. male bud: D. male flower; E. stamens; F. female flower; G &

H. styles and stigmas; I. TS ovary; J. seed; K. fruit. (All from § 97390).

Begonia from limestone hill in the Kuching Division 219

Cane-like begonia, stems up to 1.2 m tall, 3-4 mm diam., reddish brown,

woody, minutely pubescent on the uppermost nodes, internodes 3-13.5 cm

long, branching; without a tuber. Stipules lanceolate, 16-35 x 6-10 mm, pale

green, glabrous, margin entire, midrib ridged, apex attenuate, caducous.

Leaves alternate, distant, not oblique; petioles 5—10(-18) mm in the lower

leaves, 6-10(-20) mm in the upper leaves, slightly thickened, minutely

pubescent, brownish red, terete; laminas plain mid-green above, paler below,

in most plants leaves glabrous above, a few plants with minute red hairs on

a raised hair base, oblanceolate, asymmetric, 15—24 x 4.5—11 cm, broad side

4-8 cm wide, succulent in life, papery when dry, base rounded on the broad

side, basal lobe 0.5-1.3 cm long, cuneate or rounded on the narrow side,

margin minutely dentate, apex acuminate; venation palmate-pinnate with 1

pair of veins at the base, 4-6 pairs along the midrib and 2-3 veins in the basal

lobe, branching twice, impressed above, beneath veins slightly prominent,

concolorous and glabrous or minutely pubescent. Inflorescences axillary,

erect, dull red, (3-)11-—25 cm long, longer than the leaves, sessile or with a

peduncle ca 3 cm long, racemose with cymose branches of male flowers,

branches 2-4 cm long, glabrous, protogynous with (rarely 1 or) 2 pairs of

female flowers at the base and many male flowers on side branches above.

Bracts pale green, ca 7 x 3 mm, margin entire, glabrous. Male flowers with

glabrous pedicels 6-7 mm long, tepals 2, pale green flushed crimson, round

with a recurved margin, 5—7 x 5—7 mm, glabrous, stamen cluster conical, ca

2.5 mm across, stamens ca 40; filament c. 0.75 mm long, anther pale yellow,

oblanceolate, ca 1 mm long, apex rounded, dehiscing by longitudinal

slits. Female flowers with pale red pedicels 7-8 mm long, ovary pale red,

glabrous, 15 x 12-14 mm, locules 3, each locule with 2 placentas, wings 3,

equal; tepals 5, greenish yellow, isomorphic, oval, 10-11 x 4-6 mm, slightly

toothed towards the acute apex; styles 3, 3-4 mm long, bifurcating, stigma a

spirally twisted papillose band. Fruits on a stiff, decurved stalk, 10-23 mm

long, capsules glabrous, oblong, widening towards the apex, 20-28 x 21-25

mm, wings 3, subequal, larger wing 6-10 mm wide and the smaller two 6-8

mm wide, thinly fibrous, dehiscing between the locules and the wings, stigma

caducous. Seeds barrel-shaped, brown, ca 0.25 mm long, collar cells more

than half the seed length.

Distribution: Endemic in the Kuching Division; known only from the

Padawan-Serian and Penrissen limestone.

Habitat: Slope of limestone hill or on limestone screes, rubble or rocks at

130-280 m altitude, locally common.

Specimens examined: Padawan-Serian limestone - Bukit Peyang, Yi &

Othman S 46218 (SAR); Gunung Manok, Mamit S 33476 (SAR); Gunung

220 Gard. Bull. Singapore 58 (2)2007

Mas, Julia et al. S 95693 (E, KEP, SAR, SING); Gunung Selabor, Anderson

$20827 (SAR), $28053 (SAR); Selabor, 3 Sept 2005, Julia & Kiew S 95676

(E, KEP, L, SAR, SING); Seburan, Anderson S 12921 (SAR); Teng Bekap,

Mamit S 32641 (SAR). Penrissen limestone — Gunung Rimo (type).

Vernacular names: Kura (Bidayuh), riang (Iban).

Notes: This cane-like begonia is similar to Begonia lailana in habit, size, and

the leaves that are not oblique. However, it differs in the petioles not being

densely hairy, the juvenile leaves are never spotted, nor are the adult leaves

reddish beneath and the leaves are considerably larger, the inflorescences

longer, and there are two pairs of female flowers in the inflorescence.

It is more common on the Padawan-Serian limestone; as yet it is

known from only one hill (Gunung Rimo) in the Penrissen limestone area.

There are two other specimens collected from Padawan limestone

(S 27425 from Bukit Pait and CWL 1306 from Gunung Mentawa) that are

similar in habit to B. paoana but they differ in having narrowly oblong fruits

(15 x 9-25 x 15 mm) that are narrowed towards the apex. These populations

have not been refound and the specimens without inflorescences and flowers

are insufficient to describe.

This species is named in honour of Joseph Pao, botanical artist in the

Sarawak Herbarium.

10. Begonia pendula Rid.

J. Str. Br. Roy. As. Soc. 46 (1906) 257; Kiew & Geri, Gard. Bull. Singapore 55

(2003) 120.

Distribution: Endemic in the Kuching Division, Sarawak: known only from

the Bau and Padawan-Serian limestone. Bau limestone - Bidi, Bukit Boring,

Gunung Apin, G. Aup, G. Batu, G. Berloban, G. Doya, G. Jebong, G. Juita,

G. Lobang Angin, G. Pambur, G. Podam, G. Poing, G. Ropih, G. Setiak, G.

Stulang, G. Tabai, G.Tai Ton, G. Tongga, G. Umbut; Padawan-Serian limestone

- Gunung Selabor, Sinclair SFN 38471 (SING).

Habitat: On damp, mossy vertical limestone cliff faces, in shade beneath the

tree canopy.

Notes: It is a decorative begonia with bright green leaves variegated with

silver green spots between the veins. Its flowers too are attractive being

deep pink or red. On the Bau limestone, it is common. Ridley (1906) named

it ‘pendula’ from its stems that trail down the cliff faces.

Begonia from limestone hill in the Kuching Division 221

11. Begonia penrissenensis Kiew & S. Julia, sp. nov.

Section: Petermannia

A Begonia rubida Ridl. laminis longioribus 9-12 cm longis (nec usque 8 cm

longis), lamina folii quam lobo basali 5-plo longiore (nec lamina lobo basali

duplo longiore) et tepalis masculis 2 (nec 4) differt. Typus: Borneo, Sarawak,

Kuching Division, Gunung Bah, Kiew, Julia & Leong S95700, 17 Nov 2005

(holo, SAR; iso, KEP, E, K, L, SAN, SING). Figure 7

Cane-like begonia, whole plant glabrous; stems erect, up to 50 cm long

and 5 mm thick, reddish brown, several stems arising from branches from

the base, nodes swollen, internodes 2—7 cm long; without a tuber. Stipules

lanceolate, pale green, 7-10 x 3.5-6 mm, margin entire, keeled toward the

apex, apex apiculate, persistent. Leaves distant; petioles deep red, ca 2 cm

long in the upper leaves, elongating to 4.5 cm in lower leaves, succulent,

terete; laminas oblique, deep green above and pale green beneath or bronze

green above and purple-red beneath, sometimes with silver-grey spots with

a deep green centre in rows between the veins, all colour forms with deep

red veins that on the upper surface appear blackish, in life succulent and

brittle, drying thinly leathery, asymmetric, narrowly ovate, 9.5-12 x 5-6 cm,

broad side 3.2—-4 cm wide, base unequal, basal lobe large and rounded, 1.5-

2.5 cm long, margin minutely toothed at the vein endings, apex acuminate,

acumen ca 1-2 cm long; venation palmate-pinnate with 1(-2) pairs at the

base and 2-3 pairs along the midrib and with 2(-3) veins in the basal lobe,

branching twice, veins slightly prominent above and beneath, red beneath.

Inflorescences axillary, reddish, glabrous, erect, cymose panicles with up to 4

order branching, shorter than the leaves, (1.3-)2—3.6(-5.2) cm long, peduncles

1.3-1.7 cm long, female flowers on short branches at the base, subsessile to

2-7 mm long, upper branches with male flowers (7-)10-15 cm long, male

flowers many, female flowers usually 1 or sometimes 2, protogynous. Bracts

pale green, lanceolate, 7-10 x 2-4 mm, margin entire, persistent; bracteoles

similar but smaller. Male flowers with green pedicels 6-10 mm long; tepals

2, pale greenish yellow, glabrous, broadly ovate, 5-8 x 6-8.5 mm, margin

entire, apex acute to rounded; stamens 30-40, cluster broadly conical, 3-5

mm diam., sessile; filaments ca 1 mm long; anthers pale yellow, obovate,

0.75 mm long, apex emarginate, opening by lateral slits. Female flowers with

reddish pedicels 10-22 mm long; ovary pale green, 5—12 x 10-15 mm, wings

3, equal, 2-6 mm wide, pointed to rounded, locules 3, placentas 2 per locule;

tepals 4, pale green, tinged rosy red, glabrous, isomorphic, lanceolate, 6-8 x

4-5 mm, margin entire, apex acute; styles 3, 2-4 mm long, free to the base

and bifurcating with ultimate branches broadly U-shaped. Fruits pendent

on fine stalks 2.2-3 cm long; capsules 8-13 x 20 mm, locules 3, wings 3,

222 Gard. Bull. Singapore 58 (2)2007

Figure 7. Begonia penrissenensis Kiew & S.Julia

A. habit; B. lower leaf surface; C. male bud: D. male flower; E. stamen cluster; F. stamens; G.

female flower; H. open female flower; I & J. styles and stigmas; K. TS ovary; L. seed; M. fruit.

(A-L from S 95700; M from S 91389).

Begonia from limestone hill in the Kuching Division 223

rounded to pointed, equal, fibrous, wings 6-7 mm wide, splitting between

the locules and wings. Seeds barrel-shaped, c. 0.5 mm long, collar cells about

¥% seed length.

Distribution: Endemic in the Kuching Division, Sarawak, known only from

the Penrissen limestone.

Habitat: Confined to limestone, it grows on rocky shoulders of the limestone

hill c. 250 m high, in light shade.

Specimen examined: Penrissen limestone — Gunung Rimo, Julia, Kiew &

Malcom S 91389 (E, KEP, SAR).

Notes: It is an attractive begonia with the large, pale green pairs of stipules

contrasting with its red stems. In addition, some plants have adult leaves

with silver-grey spots or are bronze-coloured.

12. Begonia punchak Kiew & S.Julia, sp. nov.

Section: Petermannia

A Begonia rubida Ridl. petiolis brevioribus usque 2.5 cm longis (nec 5 cm

longis), lamina folii quam lobo basali 3-plo longiore (nec lamina lobo basali

duplo longiore) et fructibus alis angustioribus 2-3 mm latis (nec 4-5 mm latis)

differt. Typus: Gunung Burau, Penrissen, Julia, Kiew & Malcom S 91392, 30

April 2006 (holo, SAR; iso, E, KEP, L, SING). Figure 8

Cane-like begonia, whole plant glabrous. Stems erect, 50-75 cm tall, glossy,

crimson, old stem becoming brown and woody, 6—7 mm thick, several stems

produced from branching at the base, nodes swollen with a conspicuous

stipular scar, internodes 1.5-2.5 cm, upper branches zigzag; without a

tuber. Stipules foliaceous, pale green tinged red, 2.2 x 0.8 cm, margin entire,

broadly lanceolate, outside keeled, apex acute, soon caducous. Leaves

distant, alternate, held almost horizontally; petioles 1-2.5 cm long, dark

crimson, succulent, terete; lamina oblique, plain mid-green above with a red

patch at the base, yellowish green beneath contrasting with deep crimson

veins, glossy, in life succulent, asymmetric, ovate, up to 7 x 4.5 cm, decreasing

markedly in size towards the apex, broad side to 3 cm wide, base unequal,

basal lobe rounded up to 2.75 cm long, margin undulate with minute distant

teeth, apex acuminate; venation palmate-pinnate, 3 veins at the base, 1-2 pairs

along the midrib and 2 veins in the basal lobe, branching once, impressed

above, prominent beneath. Inflorescences axillary, deep crimson, unisexual,

protogynous; without bracts. Female inflorescences with a peduncle 1.5-3.5

224 Gard. Bull. Singapore 58 (2)2007

Figure 8. Begonia punchak Kiew & S.Julia

A. habit; B. lower leaf surface; C. male bud with bracteoles: D. male flower; E. open male

flower; F. stamen cluster; G. stamens; H. female flower with bracteole;I & J. styles and stigmas;

K. TS ovary; L. seed; M. fruit. (All from S 9/392).

Begonia from limestone hill in the Kuching Division 225

cm long terminating in 2 pendent female flowers. Male inflorescences with

two branches, peduncle 1.5-2.5 cm long, branches up to 4 cm long with

5-10 cymule scars, cymules with 3 male flowers, subtended by foliaceous

bracteoles, pale green tinged red, ovate, 5-7 x 6 mm, cucullate, caducous.

Male flowers many, pedicels crimson, 2—3 mm long; tepals 4, margin entire,

apex rounded, outer two broadly ovate, 5-7 x 7 mm, cream with crimson

centre, inner two cream, narrowly lanceolate ca 4 x 2 mm; stamens 23-24 in

a subsessile, globose cluster, ca 2 mm diam., filaments c. 1 mm long, anthers

yellow, obovate, ca 1 mm long, apex emarginate, opening by lateral slits.

Female flowers pendent, pedicels crimson, 15—20 mm long; ovary glossy,

crimson, ca 8 x 12 mm, wings 3, subequal, ca 3 mm wide, slightly pointed

distally, locules 3, placentas 2 per locule; tepals 5, margin entire, apex acute,

outer four isomorphic, elliptic, ca 8 x 5 mm, inner one narrow ca 4 x 2 mm;

styles 3, crimson, widely U-shaped, ca 2 mm long. Fruits pendent, peduncle

and pedicels thin and hair-like, 16-20 mm long; capsules 10-13 x 9-15 mm,

wings 3, thinly fibrous, subequal, two rounded, sometimes slightly wider

distally, 2-3 mm wide, in some fruits the third wing scarcely developed,

dehiscing between the locules and wings. Seeds barrel-shaped, ca 0.3 mm

long, collar cells about 4 of the seed length.

Distribution: Endemic in the Kuching Division, Sarawak; known from a

single hill in the Penrissen limestone.

Habitat: It grows in soil pockets in an exposed position on the limestone

rocks near the summit. The summit of this particular hill had been burned

leaving bare exposed rocks and it was in this area that it was found.

Notes: This new species is remarkable among Sarawak limestone begonias

for its microhabitat. ‘Punchak’ is the Malay word for summit, hence the

species epithet. (In the modern Malay spelling system, ‘ch’ is spelt ‘c’, so

summit would be written ‘puncak’.) This compares with Sabah, where for

example, Begonia keithii Kiew, grows in similar exposed conditions. B.

rubida from the Bau limestone grows near the summit of hills but always on

a substrate of thick peat where it is lightly shaded by the straggly trees that

grow near the summit.

The new species is distinct from B. rubida in its shorter petioles and

the basal lobe less than half the length of the lamina and in its capsules

that have narrower wings. B. penrissenensis also has leaves with a basal lobe

about a third the lamina length, but B. punchak differs from this species in

its smaller leaves with shorter petioles and the male flowers with 4 (not 2)

tepals.

Begonia punchak is distinctive among limestone begonias in sect.

Petermannia in that the female flowers are produced in separate leaf axils

226 Gard. Bull. Singapore 58 (2)2007

from the male inflorescences. The more usual conditions are for the female

flowers to be produced from the lower branches of the inflorescence or from

the base, but still in the same axil. Unlike typical species in this section, its

capsule wings are sometimes unequal, two wings being 2-3 mm wide and the

third being reduced to almost being wingless. It is an extremely decorative

begonia with glossy bright red stems, ovaries and fruits.

13. Begonia rubida Rid.

J. Str. Br. Roy. As. Soc. 46 (1906) 256; Kiew & Geri, Gard. Bull. Singapore 55

(2003) 124:

Distribution: Endemic in the Kuching Division, Sarawak; known only from

the Bau and Padawan-Serian limestone. Bau limestone - Bidi, Bukit Jebong,

Bukit Manok, Bukit Numpang, Gunung Apin, G. Batu, G. Juita, G. Meraja,

G. Pambur, G. Ropih, G. Setiak, G. Tai Ton, G. Tongga, G. Umbut.

Specimens examined: Padawan-Serian limestone - Bukit Regu, Paul et al. S$

37392 (SAR); Gunung Manok, Erwin & Paul S 27416 (SAR).

Notes: This species is likely to be more widespread on Padawan-Serian

limestone but is difficult to collect as it grows on the summit shoulders of

the tower karst hills. Erwin & Chai collected it from the summit of Gunung

Manok at 330 m altitude.

14. Begonia serapatensis Kiew & S. Julia, sp. nov.

Sect: Petermannia

A Begonia rubida Ridl. laminis majoribus 9-11 x 7-10 cm (nec 4.5-8 x 3-5.5

cm), floribus albis (nec rubris) et alis fructus inaequalibus longissima 6-9

mm lata (nec alis inter se aequalibus et 4-5 mm latis) differt. Typus: Borneo,

Sarawak, Kuching Division, Bukit Serapat, 8 August 2004, Kiew, Julia & Tan

$93267 (holo, SAR; iso, E, KEP, SING). Figure 9

Cane-like begonia, whole plant glabrous; stems erect, 20-50 cm long, young

stem reddish brown flecked with white, becoming brown and woody at the

base, several stems produced from branching at the base, nodes swollen

with a conspicuous stipular scar, 6-8 mm thick; without a tuber. Stipules

narrowly lanceolate, pale green, 2 x 4-7 mm, margin entire, apex caducous.

Leaves distant 2.5—3 cm apart; petiole 3.5-5.5 cm long, succulent, reddish

brown; lamina oblique, young leaves green with dark red veins and pale

green linear spots between the veins, glossy mid-green with a red patch at

junction of lamina and petiole, beneath pale green, succulent and brittle,

drying papery, asymmetric, broadly lanceolate, 9-11 x 7-10 cm, broad side

Begonia from limestone hill in the Kuching Division 227

lem

Figure 9. Begonia serapatensis Kiew & S.Julia

A. habit; B. male bud with bracteoles: C. male flower; D. open male flower; E. stamen cluster;

F. stamens; G. female flower; H. open female flower; I & J. styles and stigmas; K. fruit; L. TS

ovary; M. seed; N. fruit. (All from S 93267).

228 Gard. Bull. Singapore 58 (2)2007

5.5-13 cm wide, base unequal, basal lobe large and rounded, 4.5-6.5 cm

long (about half the length of the lamina), margin entire, apex acuminate;

venation palmate-pinnate with 2 pairs at the base and 2 pairs along the

midrib and with 3 in the basal lobe, branching twice, veins slightly prominent

above and beneath, red beneath. Inflorescences axillary, reddish, glabrous,

erect, cymose panicle, longer than the leaves, 9-15 cm long, peduncle ca 6.5

cm long, female flowers on short branches at the base 2.5-3 cm long, upper

branches with male flowers 6-8 cm long, male flowers many, female flowers

2 to 10 or more, protogynous. Bracts pale green, broadly ovate, ca 5 x 7 mm,

margin entire, caducous; bracteoles similar but smaller. Male flowers with

a rosy red pedicel 10-15 mm long; tepals 4, white, deep red towards the

base, margin entire, apex rounded; outer two oval, 8-10 x 6-8 mm, inner

two narrowly oval, 4-6 x 1.5—3 mm; stamens many, cluster globose, 2-3 mm

diam., sessile; filaments almost absent; anthers yellow, obovate, ca 1 mm

long, apex slightly notched, opening by lateral slits. Female flowers with a

rosy red pedicel ca 5 mm long; ovary pale green, ca 12 x 13 mm, wings 3,

unequal, longer wing ca 8 mm wide, shorter wings ca 3 mm wide, locules 3,

placenta two per locule; tepals 5, greenish white, outermost reddish at base,

margin entire, apex rounded, outer four isomorphic 9-10 x 5—7 mm, inner

tepal narrowly oval, 7-8 x 3-4 mm; styles 3, styles and stigmas pale yellow, ca

3 mm long. Fruits pendent on a thin stiff stalk 11-23 mm long; capsule 9-10 x

14-20 mm, locules 3, wings 3, rounded, unequal, fibrous, longer wing 6—9 mm

wide, shorter wings 2-3 mm wide, splitting between the locules and wings.

Seeds barrel-shaped, ca 0.8 mm long, collar cells ca half seed length.

Distribution: Endemic in the Kuching Division, Sarawak, known only from

a single limestone hill in the Penrissen area.

Habitat: It grows on rocky shoulders of a limestone hill ca 250 m high, in

light shade.

Notes: In its woody, slightly zig-zag stems and its oblique leaves that are

succulent and glabrous and with a large basal lobe almost half the length of

the lamina, it resembles Begonia rubida. It differs, however, in a number of

characters: it does not have the fine ruby coloration of the stems and flowers

of B. rubida instead the flowers of B. serapatensis are white; the leaves are

also much larger than those of B. rubida. The apparent difference in tepal

number in the female flowers between these two species [Ridley (1906)

described B. rubida as having three tepals] is not supported as the flowers of

B. rubida we examined had five, three larger and two smaller inner ones.

15. Begonia speluncae Rid.

J. Str. Br.Roy. As. Soc. 46 (1906) 258; Kiew & Geri, Gard. Bull. Singpaore. 53

Begonia from limestone hill in the Kuching Division 229

(2003) 122.

Distribution: Endemic in the Kuching Division; from the Bau, Padawan-

Serian and Penrissen limestone. Bau - Bidi, Fairy Cave, Gunung Aup, G.

Batu, G. Doya, G. Jebong, G. Kawa, G. Lanyang, G. Meraja, G. Podam, G.

Poing, Quop. Padawan-Serian - Bukit Pait ,Gunung Berloban, G. Manok, G.

Mas, G. Sebengkam, G. Selabor, Sungai Serin, Tebedu. Penrissen - Gunung

Bah, G. Burau.

Habitat: On vertical rock faces at the base of limestone cliffs or on the

vertical sides of large boulders; in the Bau area frequently on dry rock faces,

in the Padawan-Serian and Penrissen areas usually on damp, deeply shaded

rock faces.

Notes: This is the most common begonia on limestone in the Kuching

Division, being found on most hills, although it is interesting to note that

it is not found on those hills where the other begonias with a rosette habit,

Begonia andersonii and B. burttii, occur, both of which occupy the same

habitat on sheer, shaded, vertical rock faces.

Ridley (1906) described the leaves of this species as orbicular

cordate, but they are without exception peltate. Plants of the Bau limestone

uniformly have small laminas (1.2-4.5 x 1.4-5.7 cm), but plants in the

Padawan-Serian populations show a much greater range from 3.2 x 4cm to

5 x 5.5 cm with exceptional plants with laminas up to 10 x 12 cm (Patsipun

et al. S 82156). They are otherwise typical, for example, in the broad laminas

with a rounded apex and the indistinct midrib due to its bifurcating.

Specimens examined: Padawan-Serian limestone - Bukit Pait, Erwin & Paul

S 27427 (SAR), Julia & Kiew S 95687 (KEP, SAR); Gunung Berloban, Yii

S46142 (SAR); Gunung Manok, Burtt B 8132 (SAR), Julia & Kiew S 95677

(E, KEP, SAR); Gunung Mas, Julia et al. S 95691 (KEP,SAR, SING); Gunung

Sebengkam, Jamree et al. S 75900 (SAR); Gunung Selabor, Anderson S 20836

(SAR), Ilias Paie S 28044 (SAR); Sungai Serin, Jacobs 5177 (SAR); Tebedu

15th Mile, Abg. Mohtar S 49238 (SAR); Bukit Payung, Tebedu, Patsipun

et al. S 82156 (KEP, SAR). Penrissen limestone: Gunung Bah, Julia et al. S

95694 (SAR), Rantai et al. S 66034 [Gunung Bar] (SAR); G. Burau, Julia et

al. S 91388 (E, KEP, L, SAN, SAR, SING).

Incompletely Known

The begonia collected by Yii & Othman (S 46275, SAR) from Gunung

Selangan (10 km Tebakang/Tebedu Road), Padawan-Serian limestone, is

230 Gard. Bull. Singapore 58 (2)2007

quite unlike any other limestone begonia in its short petiole, non-oblique,

narrowly obovate leaves 10 by 3.5 cm and its short inflorescences. It belongs

to the same group of begonias as Begonia pubescens Ridl., but is very

different in being a larger plant (about 15 cm tall) and in being much less

pubescent. It is obviously a new species but it has not been recollected and

the material available is too incomplete to describe.

Acknowledgements

We are grateful to Datu Cheong Ek Choon, Director of Forests; to Ms Lucy

Chong, Sarawak Forestry Corporation for her strong support for this project

and to the curators of BM, E, K, KEP, SAR and SING for permission to

examine specimens in their care; to Joseph Pao for the botanical plates; to

the Singapore Botanic Gardens for partly funding the project and to Mark

Coode for correcting the botanical Latin.

References

Anderson, J.A.R. 1965. Limestone habitat in Sarawak, p. 49-57. In Symp.

Ecol. Res. in Humid Tropical Vegetation. Sarawak Government &

UNESCO.

Banda, R.M.,R.Gendang and A.U.Ambun. 2004. Geology and geochemistry

of the limestone in Sarawak. In: H.S. Yong, ES.P. Ng and E.E.L. Yen (eds.)

Sarawak Bau Limestone Biodiversity. Sarawak Museum Journal. 59, no.

80; Special issue 6: 41-61.

Burtt, B.L. 1978. Studies in Gesneriaceae of the- Old World: XLV. A

preliminary revision of Monophyllaea. Notes Royal Botanic Gardens

Edinburgh 37. 1-59.

Kiew, R. 2001. The limestone begonias of Sabah - flagship species for

conservation. Gardens Bulletin Singapore 53: 241-286.

Kiew, R. and C. Geri. 2003. Begonias from the Bau limestone, Borneo,

including a new species. Gardens Bulletin Singapore 53: 113-123.

Kiew, R., C. Geri, J. Sang and P.C. Boyce. 2004. The understorey flora. In HLS.

Yong, F.S.P. Ng and E.E.L. Yen (eds.) Sarawak Bau Limestone Biodiversity.

Begonia from limestone hill in the Kuching Division 23k

Sarawak Museum Journal. 59, no. 80; Special issue 6: 105-146.

Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) from the Niah

National Park, Sarawak, Malaysia. Gardens Bulletin Singapore 55:

73-88.

Ridley, H.N. 1906. Begonias of Borneo. Journal Straits Branch of Royal

Asiatic Society 46: 247-261.

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Gardens’ Bulletin Singapore 58 (2) 2007: 233-274 233

Five new genera and three new species of

Grammitidaceae (Filicales)

and the re-establishment of Oreogrammitis

BARBARA S. PARRIS

Fern Research Foundation

21 James Kemp Place, Kerikeri

Bay of Islands, New Zealand 0230

Abstract

Five new genera of Grammitidaceae (Filicales) are described: Ctenopterella Parris (12 species),

Dasygrammitis Parris (six species), Radiogrammitis Parris (28 species), Tomophyllum

(E.Fourn.) Parris (22 species) and Xiphopterella Parris (six species). Oreogrammitis Copel.

(Grammitidaceae) is re-established (110 species). New combinations are made in all six

genera. Synonymy is provided, types are indicated, including those of synonyms, and

lectotypes are chosen for some names. Three new species are described from Peninsular

Malaysia, Oreogrammitis malayensis Parris, O. kunstleri Parris and Xiphopterella gracilis

Parris.

Introduction

Grammitidaceae contains more than 20 genera and more than 750 species,

mostly occurring in montane forests in the wet tropics of both hemispheres.

Parris (2003) lists the currently recognized genera. Parris (1997a) notes that

in Old World Grammitidaceae, Ctenopteris Blume ex Kunze, Grammitis Sw.

s.. and Xiphopteris Kaulf. have been used for some time as unsatisfactory

artificial genera, based on frond dissection, pending further study on generic

boundaries in the family. Ctenopteris is a synonym of Prosaptia C.Presl,

however, while Xiphopteris isasynonym of Cochlidium Kaulf.and Grammiitis

s.str. is a small genus of c. 23 species with blackish sclerotic lamina margins.

Ranker et al. (2004) demonstrate clearly that Ctenopteris and Grammitis are

not monophyletic. Parris 1997a, 1998a, 2002a and 2004 form part of a series

describing new genera in Old World Grammitidaceae and re-assigning the

species formerly treated as Crenopteris, Grammitis s.l. and Xiphopteris.

Preparation of an account of Grammitidaceae for the Flora of

Peninsular Malaysia necessitates the description of five new genera,

Ctenopterella Parris, Dasygrammitis Parris, Radiogrammitis Parris,

Tomophyllum (E.Fourn.) Parris and Xiphopterella Parris, and the re-

establishment of Oreogrammitis Copel.,as part of a revision of the Old World

234 Gard. Bull. Singapore 58 (2) 2007

genera of Grammitidaceae. Two new species of Oreogrammitis and one

new species of Xiphopterella from Peninsular Malaysia are described here.

New species from areas other than Peninsular Malaysia will be described

later. Monographs of the new genera, together with Oreogrammitis, are

currently being prepared and will contain detailed species descriptions,

keys to species and distribution maps. Synonymy is provided below for the

new combinations. Types of all names are indicated and some lectotypes are

chosen here.

Ctenopterella

-Ctenopterella Parris, gen. nov.

Typus generis: C. blechnoides (Grev.) Parris

Rhizomata dorsiventralia, stipitibus in duobus seriebus, interdum articulatis,

interdum phyllopodiis. Paleae pallidae brunneae ad atrobrunneae, non

clathratae, non iridescentes, glabrae, impolitae, subpolitae vel politae, cellulis

turgidis vel non turgidis, sine septis. Pili integri et ramose, albidi vel pallidi

brunnei ad atrobrunnei. Laminae profunde pinnatifidae, pinnatae vel

bipinnatifidae; venae non anastomosantes, plerumque pinnatim ramosae, raro

dimidiate ramosae, extremis venarum plerumque hydathado in superficie.

Pinnae soris plerumque in duobus serialibus, raro uniseriate, sori in lamina

nonimmersi vel parum immersi, rotundi vel elliptici; sporangia glabra. Species

12, Africae ad Polynesiae incolae.

Rhizomes dorsiventral, stipes in two rows, sometimes articulated to

rhizome, phyllopodia sometimes present. Rhizome scales pale to medium

brown, not clathrate, not iridescent, glabrous, dull to glossy, cells turgid

or not, without cross-walls. Hairs simple and branched, whitish or pale to

dark brown. Laminae deeply pinnately lobed, pinnate or bipinnately lobed;

venation free, veins in pinnae usually pinnately branched, rarely dimidiately

branched, ends of vein branches usually with hydathodes on adaxial surface

of lamina. Sori on pinnae, usually in two rows, rarely in one row, superficial

or slightly sunken, + circular to elliptic; sporangia glabrous. 12 species, Africa

to Polynesia.

The generic name is a diminutive of Ctenopteris,in which most of the species

were formerly placed.

Species of Ctenopterella were not sampled by Ranker et al. (2004), so the

position of the genus in their clade system is unknown. Four Old World

Five new genera and three new species of Grammitidaceae 235

clades are recognised (Ranker et al. 2004); I, I, I] and VIII. All species in

Clades II and VIII have dorsiventral rhizomes, with stipes in two rows, as do

most species in Clade 1, except for Radiogrammitis parva (Brause) Parris

and R. hirtelloides (Copel.) Parris, which have radial rhizomes, with stipes

in whorls. Species of Clade III belong to Tomophyllum, Scleroglossum and

Calymmodon and all have radial rhizomes, with stipes in whorls. On the

basis of rhizome form, type of rhizome scale and hair types Crenopterella is

more likely to be in Clade Ia near Themelium [including T: conjunctisorum

(Baker) Parris], or Clade II, than in Clades III or Clade VIII, which contains

the Hawaiian Islands endemic genus Adenophorus.

1. Ctenopterella blechnoides (Grev.) Parris, comb. nov.

Grammitis blechnoides Grev., Ann. Mag. Nat. Hist. 2, 1 (1848) 328, t. 17.

Type: Sibbald s.n., Raiatea, Society Is. (E, lecto., chosen here; E, isolecto.).

- Polypodium blechnoides (Grev.) Hook., Sp. Fil. 4 (1863) 180. - Cryptosorus

blechnoides (Grev.) J.Sm., Hist. Fil. (1875) 87. - Ctenopteris blechnoides

(Grev.) W.H.Wagner & Grether, Univ. Calif. Publ. Bot. 23 (1948) 61. -

Polypodium decorum Brack., U.S. Expl. Exped. 16 (1854) 7,t.2 f.2. Type: U.S.

Expl. Exped. s n., Tahiti, Society Is. (K, US, n.v.). — Polypodium macgregori

Baker, Ann. Bot. 8 (1894) 130. Type: Macgregor s.n., Rossel I., Milne Bay,

Papua New Guinea (K, holo.). - Polypodium moultonii Copel., Philipp.

J. Sci., C. Bot. 10 (1915) 149. Type: Native collector, Bur. Sci. 2572, Retuh,

Sadong, Sarawak, East Malaysia [MICH, lecto. (Parris, 2002b)]. - Ctenopteris

moultonii (Copel.) C.Chr. & Tardieu, Notul. Syst. (Paris) 8 (1939) 181. -

Polypodium revolvens Alderw., Bull. Jard. Bot. Buitenzorg, sér. 3, 2 (1920)

170, f. G. Type: Teysmann 50, Mt Sanggir, Banka Is., Sumatra (BO, n.v.; L,

n.v.).— Polypodium pseudorevolvens Alderw., Bull. Jard. Bot. Buitenzorg, sér.

3,5 (1922) 219. Type: Bunnemeijer 6902, Mt Tanda, Lingga I., Sumatra (BO

n.v.;L).- Polypodium palaoinsulare Hosokawa, Trans. Nat. Hist. Formosa 31

(1941) 476. Type: Hosokawa 9049, Mt Sul, Babeldaob, Palau (Herb. Taihoku

Imperial Univ., holo., n.v.). — Ctenopteris palaoinsularis (Hosokawa) H.Ito,

Bot. Mag. Tokyo 67 (1954) 215.

2. Ctenopterella cornigera (Baker) Parris, comb. nov.

Polypodium cornigerum Baker, Syn. Fil. ed. 2 (1874) 508. Type: Thwaites

C. P. 4005, Horton Plains, Sri Lanka (PDA, holo.). - Grammitis cornigera

(Baker) Ching, Bull. Fan Mem. Inst. 10 (1941) 240. - Xiphopteris cornigera

(Baker) Copel., Gen. Fil. (1947) 215. - Micropolypodium cornigerum (Baker)

X.C.Zhang, Fl. Reipub. Pop. Sin. 6 (2000) 301.

3. Ctenopterella denticulata (Blume) Parris, comb. nov.

Grammiitis denticulata Blume, Fl. Javae 2 (1830) 121, t. 50, f. 4. Type: Blume

236 Gard. Bull. Singapore 58 (2) 2007

s.n., Java (L, n.v.). — Polypodium denticulatum (Blume) C.Presl, Tentamen

(1836) 178. - Calymmodon denticulatus (Blume) T.Moore, Index Fil. (1857)

Ixii. - Ctenopteris denticulata (Blume) C.Chr. & Tardieu, Notul. Syst. (Paris)

8 (1939) 181. - Polypodium solidum Mett., Abhandl. Senckenberg. Naturf.

Gesell. 2 (1857) 53, t. 1, f. 1-3. Type: Zollinger 165, Java (LZ7). - Ctenopteris

solida (Mett.) J.Sm., Hist. Fil. (1875) 184. - Grammitis solida (Mett.) Ching,

Bull. Fan Mem. Inst. Biol. 10 (1941) 241. - Polypodium undosum Baker, J.

Bot. 28 (1890) 108. Type: Macgregor s.n., Mt Musgrave, Central, Papua New

Guinea (K, holo.). - Ctenopteris undosa (Baker) Copel., Philipp. J. Sci. 81

(1953) 103.- Polypodium sibarongae Christ, Ann. Jard. Bot. Buitenzorg, sér.

2,4 (1904) 36. Type: F & P. Sarasin 2011, Mt Sibaronga, Sulawesi (BAS, n.v).

— Polypodium serratodentatum Alderw., Bull. Dép. Agric. Indes Néerl. 18

(1908) 20. Type: Forbes 244, Java? (BO, holo., n.v.). - Polypodium solidum

var. bolanicum Rosenst., Repert. Spec. Nov. Regni Veg. 12 (1913) 177.

Syntypes: Keysser B 21 & 39, Bolan Mts, Papua New Guinea (both UC).

— Polypodium bolanicum (Rosenst.) Copel., Univ. Calif. Publ. Bot. 12 (1931)

403. - Ctenopteris bolanica (Rosenst.) Copel., Philipp. J. Sci. 81 (1953) 103. -

Polypodium serratodentatum var. major Alderw., Bull. Jard. Bot. Buitenzorg,

sér. 2, 16 (1914) 59. Type: Matthew 230, Kapahiang-barisan, Sumatra (BO,

n. v.; K). — Polypodium scalpturatum C.Chr., Svensk Bot. Tidskr. 16 (1922)

100, f. 6. Type: Kaudern 37, Bolaang to Mongondou, Sulawesi (BM, n.v.).

— Polypodium vernicosum Alderw., Nova Guinea 14 (1924) 46. Syntypes:

Lam 1797, 1850, Mt Doorman, Papua (both BO, n.v.). — Polypodium

pycnophyllum C.Chr., Brittonia 2: 308 (1937). Type: Brass 4421, Mt Albert

Edward, Central, Papua New Guinea (BM, holo., n.v.; BRI, MICH, NY, iso.,

n.v.). — Ctenopteris pycnophylla (C.Chr.) Copel., Philipp. J. Sci. 81 (1953)

102.

4. Ctenopterella gordonii (S.B.Andrews) Parris, comb. nov.

Polypodium gordonii Watts, Proc. Linn. Soc. New South Wales 39 (1915)

792, t. 89, f. 12 non Baker (1879). Type: Watts & Gordon s.n., Tully Falls,

Queensland, Australia [NSW, lecto. (Parris, 1998b); BRI, MEL, isolecto.]. -

Ctenopteris gordonii S.B.Andrews, Austrobaileya | (1977) 12, nomen novum

pro Polypodium gordonii Watts (1915) non Baker (1879).

5. Ctenopterella khaoluangensis (Tagawa & K.Iwats.) Parris, comb. nov.

Xiphopteris khaoluangensis Tagawa & K.Iwats., Acta Phytotax. Geobot.

23 (1969) 177, f. 14. Type: Iwatsuki et al. T:14594, Khao Luang, Nakhon Si

Thammarat, Thailand (KYO, holo., n.v.; K, iso.). — Ctenopteris malayana

Parris, Kew Bull. 41 (1986) 494. Type: Piggott 1285, G. Ulu Kali, Pahang/

Selangor, Peninsular Malaysia (K, holo.).

Five new genera and three new species of Grammitidaceae 257

6. Ctenopterella lepida (Brause) Parris, comb. nov.

Polypodium lepidum Brause, Notizbl. Bot. Gard. Berlin-Dahlem 8 (1922)

139. Type: Vaupel 421, Mangaloa, Savaii, Samoa (B, n.v.).

7. Ctenopterella macrorhyncha (Baker) Parris, comb. nov.

Polypodium macrorhynchum Baker, J. Bot. 18 (1880) 370. Type: Kitching

s.n., Tanala, Madagascar (K).

8. Ctenopterella parvula (Bory ex Willd.) Parris, comb. nov.

Polypodium parvulum Bory ex Willd., Sp. Pl. 5 (1810) 182. Type: Réunion

(B, n.v.; photo. FI). - Ctenopteris parvula (Bory ex Willd.) J.Sm., Hist. Fil.

(1875) 185. - Grammitis parvula (Bory ex Willd.) C.V.Morton, Contr. U. S.

Nat. Herb. 38 (1973) 261.

9. Ctenopterella pediculata (Baker) Parris, comb. nov.

Polypodium lobbianum Hook., Sp. Fil. 4 (1863) 226, t. 278B non Hook.

(1853). Type: Lobb, s.n., Sarawak, East Malaysia (CGE, K). — Polypodium

pediculatum Baker in Hook. & Baker, Syn. Fil. (1868) 455, nomen novum

pro P. lobbianum Hook. (1863) non Hook. (1853). — Ctenopteris lobbiana

J.Sm., Hist. Fil. (1875) 185, nomen novum pro P. lobbianum Hook. (1863)

non Hook. (1853). — Polypodium inarticulatum Copel., Philipp. J. Sci. 1,

Suppl. 2 (1906) 160, nomen superff. illegit. pro P. pediculatum Baker (1868).

- Ctenopteris pediculata (Baker) Copel., Fern Fl. Philipp. 3 (1960) 530.

10. Ctenopterella seemannii (J.Sm.) Parris, comb. nov.

Polypodium contiguum Brack., Wilkes, U. S. Expl. Exped. 16 (1854) 6, t. 2,

f. 1 non (G. Forst.) J.Sm. (1841). Type: U.S. Expl. Exped. s. n., Muthuata Mts,

Fiji (K, MICH, US, n.v.).- Cryptosorus seemannii J.Sm., Bonplandia 9 (1861)

262, nomen novum pro Polypodium contiguum Brack. (1854) non (G.Forst.)

J.Sm. (1841). - P. seemannii (J.Sm.) Copel., B. P. Bishop Mus. Bull. 93 (1932)

69. - Ctenopteris seemannii (J.Sm.) Copel., J. Arnold Arboretum 30 (1949)

442.

11. Ctenopterella vodonaivalui (Brownlie) Parris, comb. nov.

Ctenopteris vodonaivalui Brownlie, Nova Hedwigia, Beih. 55 (1977) 357, t.

40, f. 10. Type: Vodonaivalu 2173, Namosi, Viti Levu, Fiji (CHR, holo.).

12. Ctenopterella zenkeri (Hieron.) Parris, comb. nov.

Polypodium zenkeri Hieron., Bot. Jahrb. Syst. 46 (1911) 385. Type: Zenker

1507, Yaunde, Cameroon (B, lecto., chosen here; BM, E, K, P, isolecto., 7.v.).

— Ctenopteris zenkeri (Hieron.) Tardieu in Aubreville, Fl. Cameroun 3 (1964)

327. — Xiphopteris zenkeri (Hieron.) Schelpe, Contr. Bolus Herb. 1 (1969)

238 Gard. Bull. Singapore 58 (2) 2007

12. - Polypodium decorum var. excaudatum Bonap., Notes Ptérid. 4 (1917)

79. Syntypes: Perrier de la Bathie 7533, Forét d’ Analamazaotra, Madagascar

(BM, P,n.v.); Perrier de la Bathie 7593, Manankazo au Nord-Est d’Ankazohe,

Madagascar (P, n.v.); Perrier de la Bathie 7643, Montagnes du Sambirano,

Madagascar (P, n.v.). - Ctenopteris excaudata (Bonap.) Tardieu, Notul. Syst.

(Paris) 15 (1959) 445. - Polypodium excaudatum C.Chr., Arch. Bot. Caen.

2 (1928) 215. Type: Perrier de la Bathie 7533, Forét d’Analamazaotra,

Madagascar (BM, holo.; P, iso., n. v.).

Dasygrammitis

‘ Dasygrammitis Parris, gen. nov.

Typus generis: D. mollicoma (Nees & Blume) Parris

Rhizomata radialia, stipitibus in verticillis, non articulatis. Paleae mediae

brunneae ad atrobrunneae, non clathratae, non iridescentes, plerumque

politae, raro subpolitae, ad marginem pilis mediis brunneis ad atrobrunneis,

raro pallidis brunneis, cellulis plerumque turgidis, raro subturgidis, sine

septis. Pili integri et ramosi, plerumque atrobrunnei, raro medii brunnet.

Laminae plerumque pinnatae, raro pinnatifidae vel bipinnatifidae; venae non

anastomosantes, pinnatim ramosae, extremis venarum hydathado in superficie

destitutis. Pinnae soris in duobus serialibus, raro uniseriati, sori in lamina

non immersi, rotundi vel elliptici; sporangia glabra vel uno vel duobus pilis

jJuxta annulum ad basim sporangio vel apicem versus. Species 6, Ceylonae ad

Polynesiae incolae.

Rhizomes radial, stipes in whorls, not articulated to rhizome. Rhizome

scales medium to dark red-brown, rarely pale red-brown, concolorous, not

clathrate, not iridescent, usually glossy, rarely subglossy, with medium to

dark red-brown, rarely pale red-brown, simple eglandular hairs on margin,

cells usually turgid, rarely subturgid, without cross-walls. Hairs simple and

branched, usually dark brown, rarely medium brown. Laminae usually

pinnate, rarely deeply pinnately lobed or bipinnatifid; venation free, veins

pinnately branched in pinnae, ends of vein branches without hydathodes on

adaxial surface of lamina. Sori in two rows on pinnae, superficial, + circular

to elliptic; sporangia glabrous or with one or two medium to dark red-brown

hairs adjacent to annulus at base of sporangium or part way towards apex.

Six species, Sri Lanka to Polynesia.

The generic name refers to the often shaggy nature of the frond indumentum

and to Grammiitis, the type genus of the family.

Five new genera and three new species of Grammitidaceae 239

No species of Dasygrammitis were sampled by Ranker et al. (2004), so its

position in their clade system is unknown. On the basis of rhizome form and

hair types it is likely to be in Clade III, characterized by radial rhizomes,

together with Zomophyllum, Scleroglossum and Calymmodon. It most

closely resembles Tomophyllum, but differs from it in darker usually glossy

rhizome scales with usually turgid cells, darker hairs and the absence of

hydathodes.

1. Dasygrammitis brevivenosa (Alderw.) Parris, comb. nov.

Polypodium brevivenosum Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 28

(1918) 40. Type: Brooks 299/S, Lebong Simpang, Bengkoelen, Sumatra (BO,

n.v.).- Ctenopteris brevivenosa (Alderw.) Holttum, Rev. Fl. Malaya 2 (1955)

ans:

2. Dasygrammitis crassifrons (Baker) Parris, comb. nov.

Polypodium crassifrons Baker in Hook. & Baker, Syn. Fil. (1867) 325. Type:

Deplanche s.n., New Caledonia (K). - Grammitis crassifrons (Baker) Ching,

Bull. Fan Mem. Inst. Biol. 10 (1941) 240. - Ctenopteris crassifrons (Baker)

Brownlie, Pacific Sci. 14 (1960) 401. - Prosaptia crassifrons (Baker) Parris,

Kew Bull. 50 (1995) 165. - Polypodium stenopteron Baker, J. Bot. 24 (1886)

183. Type: Thurston s.n., Viti Levu, Fiji (K). - Polypodium hispidosetosum

Rosenst., Repert. Spec. Nov. Regni Veg. 10 (1912) 162. Type: Franc 1318, Mt

Mou, New Caledonia (P, 7.v.).

3. Dasygrammitis fuscata (Blume) Parris, comb. nov.

Polypodium fuscatum Blume, Enum. Pl. Javae (1828) 129. Type: Blume s.n.,

Java (L, n.v.). - Ctenopteris fuscata (Blume) Blume ex Kunze, Bot. Zeit. 4

(1846) 425. - P mollicomum Nees & Blume var. fuscatum (Blume) Alderw.,

Malayan Ferns (1908) 601.

4. Dasygrammitis malaccana (Baker) Parris, comb. nov.

Polypodium malaccanum Baker, Ann. Bot. 8 (1894) 129. Type: Ridley

3345, G. Mering, Johor, Peninsular Malaysia (K, holo.; SING, iso., 7.v.). -

Polypodium pachycaulum Copel., Philipp. J. Sci. 56 (1935) 105, t. 10. Type:

Copeland s.n., Mt Apo, Mindanao, Philippines (MICH, UC). - Ctenopteris

pachycaulis (Copel.) Copel., Fern Fl. Philipp. 3 (1960) 529. - Ctenopteris

matutumensis Copel., Philipp. J. Sci. 81 (1952) 45. Type: Copeland s.n., Mt

Matutum, Mindanao, Philippines (MICH, holo.).

5. Dasygrammitis mollicoma (Nees & Blume) Parris, comb. nov.

Polypodium mollicomum Nees & Blume, Nova Acta 11 (1823) 121, t. 12,

f. 2. Type: Blume s.n., Java (L). - Ctenopteris mollicoma (Nees & Blume)

240 Gard. Bull. Singapore 58 (2) 2007

Kunze, Bot. Zeit. 4 (1846) 425. - Polypodium rufescens Brause, Bot. Jahrb.

Syst. 49 (1912) 43 non Blume (1830). Type: Schultze 322, Augusta River,

West or East Sepik, Papua New Guinea (B, left hand plant, lecto., chosen

here). — Polypodium rufidulum C.Chr., Index Fil. Suppl. 3 (1934) 157, nomen

novum pro P. rufescens Brause (1912) non Blume (1830). - Ctenopteris

rufidula (C.Chr.) Parris, Fern Gaz. 12 (1980) 118. - Ctenopteris micropaleata

Copel., Gen. Fil. (1947) 219, nomen superfi. illegit. pro P. rufidulum C.Chr., P.

rufescens Brause (1912), non Blume (1830), non C. rufescens Kunze.

6. Dasygrammitis purpurascens (Nadeaud) Parris, comb. nov.

Polypodium purpurascens Nadeaud, Enum. Pl. Indig. Tahiti (1873) 25.

Type: Nadeaud s.n., Tahiti, Society Is. (P, n.v.). — Ctenopteris purpurascens

(Nadeaud) Copel., J. Arnold Arboretum 30 (1949) 442.

Radiogrammitis

, Radiogrammitis Parris, gen. nov.

Typus generis: R. setigera (Blume) Parris

Rhizomata radialia, stipitibus in verticillis, non articulatis. Paleae pallidae

brunneae ad atrobrunneae, plerumque non clathratae, raro_ clathratae,

plerumque non iridescentes, raro iridescentes, plerumque subpolitae, raro

impolitae, glabrae, cellulis plerumque non turgidis, raro turgidis, sine septis;

paleae interdum absentes. Pili plerumque integri, raro ramosi, albidi vel

pallidi brunnei ad atrobrunnei. Laminae plerumque simplices, raro pinnatim

lobatae; venae non anastomosantes, plerumque dichotome_ uni-bi-(tri-

)ramosae, raro simplices vel pinnatim ramosae, extremis venarum interdum

hydathado in superficie. Sori plerumque in duobus, raro ad duodecimos,

serialibus, plerumque uno, raro ad sextos, utroque medio-venae, plerumque

in lamina non immersi, interdum parum immersi, raro profunde immersi,

rotundi vel elliptici; sporangia praecocia in soris proprie pilis uno usge ad

novena ad apicem juxta annulum, sporangia serotina in soris glabra. Species

28, Ceylonae ad Polynesiae incolae.

Rhizomes radial, stipes in whorls, not articulated to rhizome. Rhizome

scales pale to dark brown, usually not clathrate, rarely clathrate, usually

not iridescent, rarely iridescent, usually subglossy, rarely dull, glabrous,

cells usually not turgid, rarely turgid, without cross-walls; scales sometimes

absent. Hairs usually simple, rarely branched, whitish or pale to dark brown.

Laminae usually simple, rarely pinnately lobed; venation free, veins usually

dichotomously one-two-(three-) branched, rarely simple or pinnately

branched, ends of vein branches sometimes with hydathodes on adaxial

surface of lamina. Sori usually in two rows, rarely up to 12, usually one,

Five new genera and three new species of Grammitidaceae 241

rarely up to six, on each side of the mid-vein, usually on surface of lamina,

sometimes slightly sunken in broad shallow depressions, rarely deeply

sunken in steep-sided pits, + circular to elliptic; first-developed sporangia of

each sorus characteristically with one to nine hairs at apex of sporangium

adjacent to annulus; later-developed sporangia of each sorus glabrous. 28

species, Sri Lanka to Polynesia.

The generic name refers to the radial rhizome and to Grammitis, in which

the species were formerly placed.

Species of Radiogrammitis sampled by Ranker et al. (2004) are in the lower

part of Clade Ia. Radiogrammitis closely resembles Oreogrammitis, but

differs in its radial, rather than dorsiventral rhizome, and in the absence of

rhizome scales in numerous species.

1. Radiogrammitis ahenobarba (Parris) Parris, comb. nov.

Grammitis ahenobarba Parris, Blumea 29 (1983) 142, f. 22, 3. Type: Gibbs

5551, Anggi Lakes, Vogelkop Peninsula, Papua (BM, holo.).

2. Radiogrammitis alepidota (M.G.Price) Parris, comb. nov.

Grammitis alepidota M.G.Price, Philipp. Agricult. 57 (1974) 34, f. 1. Type:

Price 2149, Mt Banahaw, Luzon, Philippines (PNH, holo., n.v.; K, L, US, iso.,

ny).

3. Radiogrammitis beddomeana (Alderw.) Parris, comb. nov.

Polypodium beddomeanum Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 28

(1918) 39. Type: Wall 30, Bogawantalawa, Sri Lanka (BO, n.v.; photo., BM).

- Grammitis beddomeana (Alderw.) Ching, Bull. Fan Mem. Inst. Biol. 10

(1941) 240.

4. Radiogrammitis cheesemanii (Parris) Parris, comb. nov.

Grammitis cheesemanii Parris, N. Z. J. Bot. 31 (1993) 15. Type: Cheeseman s.

n., Te Kou, Rarotonga, Cook Is. (AK, holo.; AK, iso.).

5. Radiogrammitis clavipila (Parris) Parris, comb. nov.

Grammitis clavipila Parris, Blumea 29 (1983) 156, f. 21. Type: Vink 17020,

Ibiwara, Enga, Papua New Guinea, (L, holo.; LAE, iso.).

6. Radiogrammitis coredrosora (Aldetrw.) Parris, comb. nov.

Polypodium coredrosorum Aldetw., Nova Guinea 14 (1924) 44. Type: Lam

1550A, Mt Doorman, Papua [BO, lecto. (Parris, 1983); BM, L, isolecto.].

Grammitis coredrosora (Alderw.) Copel., Philipp. J. Sci. 80 (1952) 179.

242 Gard. Bull. Singapore 58 (2) 2007

7. Radiogrammitis graminella (C.Chr.) Parris, comb. nov.

Polypodium simplex Baker, J. Bot. 14 (1876) 12 non Burm.f. (1868) nec.

Sw. (1801) nec Lowe (1858). Type: Whitmee 123, Samoa (CGE, GH n.v.;

K). - Polypodium graminellum C.Chr., Index Fil. (1906) 530, nomen novum

pro P. simplex Baker non Burm.f. (1868) nec Sw. (1801) nec Lowe (1858).

- Grammitis graminella Sledge, Brit. Fern Gaz. 9 (1967) 340.

8. Radiogrammitis havilandii (Baker) Parris, comb. nov.

Polypodium havilandii Baker, Trans. Linn. Soc. London, Bot. 2, 4 (1894)

253. Type: Haviland 1488, Mt Kinabalu, Sabah, East Malaysia (K, holo.).

- Polypodium multisorum Copel., Philipp. J. Sci., C. Bot. 12 (1917) 61. Type:

Topping 1665, Mt Kinabalu, Sabah, East Malaysia [MICH, lecto. (Parris

1990); BM, US, isolecto., n.v.). - Grammitis havilandii (Baker) Copel., Gen.

Fil. (1947) 211.

9. Radiogrammitis hirtella (Blume) Parris, comb. nov.

Polypodium hirtellum Blume, Enum. Pl. Javae (1828) 122. Type: Blume 71,

G. Gede, Java (L, lecto., chosen here). - Grammitis hirtella (Blume) Tuyama,

Bot. Mag. Tokyo 51 (1937) 125. - Grammitis pusilla Blume, Fl. Javae 2 (1829)

109, t. 46, f. 4, excl. varieties, nomen illegit. superfl. pro Polypodium hirtellum

Blume. — Polypodium pusillum (Blume) Christ, Ann. Jard. Bot. Buitenzorg

15 (1897) 145.

10. Radiogrammitis hirtelloides (Copel.) Parris, comb. nov.

Polypodium hirtelloides Copel., Bull. B. P. Bishop Mus. 59 (1929) 17. Type:

Gillespie 2387, Nadarivatu, Viti Levu, Fiji (BISH, MICH, UC, n.v.). -

Grammitis hirtelloides (Copel.) Copel., Philipp. J. Sci. 80: 146 (1952).

11. Radiogrammitis hirtiformis (Rosenst.) Parris, comb. nov.

Polypodium hirtiforme Rosenst., Repert. Spec. Nov. Regni Veg. 12 (1913)

176. Type: Keysser B.48, Bolan Mts, Morobe, Papua New Guinea [S, lecto.

(Parris 1983); BM, UC, isolecto.].

12. Radiogrammitis holttumii (Copel.) Parris, comb. nov.

Grammitis holttumii Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Holttum SFN 23384, Cameron Highlands, Pahang, Peninsular Malaysia

(MICH, holo.; K, SING, UC, iso.).

13. Radiogrammitis jagoriana (Mett. ex Kuhn) Parris, comb. nov.

Polypodium jagorianum Mett. ex Kuhn, Linnaea 36 (1869) 128. Type: Jagor

835, Luzon, Philippines (B, lecto., chosen here; BM, isolecto.). - Grammitis

Five new genera and three new species of Grammitidaceae 243

jagoriana (Mett. ex Kuhn) Tagawa, Acta Phytotax. Geobot. 10 (1941) 284.

- Polypodium koordersii Christ, Ann. Jard. Bot. Buitenzorg 15 (1897) 144, t.

15, f. 19 A-C. Type: Koorders 17135B, Mt Klabat, Minahassa, Sulawesi (BO,

lecto., chosen here; L, isolecto.). - Polypodium christii Copel., Polypodiaceae

Philipp. (1905) 118. Type: Copeland 1520, Mt Apo, Mindanao, Philippines

(MICH, lecto., chosen here; BM, isolecto.). - Grammitis cuneifolia Copel.,

Philipp. J. Sci. 56 (1935) 479, t. 8, f. 2. Type: Clemens 40962, Mt Kinabalu,

Sabah, East Malaysia [MICH, lecto. (Parris et al., 1992)].

14. Radiogrammitis kinabaluensis (Copel.) Parris, comb. nov.

Polypodium kinabaluense Copel., Philipp. J. Sci. 12, C. Bot. (1917) 60. Type:

Clemens 10649 p.p., Mt Kinabalu, Sabah, Malaysia [MICH, lecto. (Parris et

al. 1992); BM, UC, isolecto.]. - Grammitis kinabaluensis (Copel.) Copel.,

Philipp. J. Sci. 56 (1935) 479.

15. Radiogrammitis ligulata (Baker) Parris, comb. nov.

Polypodium ligulatum Baker in Hook. & Baker, Syn. Fil. (1867) 320. Type:

Wilkes in S. Pacific Expl. Exped. 1838-1842, Fiji (K). - Grammitis ligulata

(Baker) Ching, Bull. Fan Mem. Inst. Biol. 10 (1941) 240. - Polypodium

trachycarpum Mett. ex Kuhn, Linnaea 36 (1869) 127. Type: Vesco s.n., Tahiti,

Society Is. (B, lecto., chosen here). - Grammitis trachycarpa (Mett. ex Kuhn)

Copel., Occ. Papers Bishop Mus. 14 (1938) 72.

16. Radiogrammitis meijer-dreesii (Copel.) Parris, comb. nov.

Grammitis meijer-dreesii Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Brass 9898 & Meijer-Drees, Mt Trikora, Papua (MICH, holo.; A, BO, L,

iSO:)}

17. Radiogrammitis multifolia (Copel.) Parris, comb. nov.

Polypodium alpestre Blume, Enum. Pl. Javae (1828) 123 non (Hoppe)

Spenner (1825). Type: Blume s.n., Java [L, lecto. (Parris, 1986)]. - Grammitis

pusilla var. alpestris Blume, Fl. Javae 2 (1829) 110, t. 46, f 5, pro P. alpestre

Blume (1828) non (Hoppe) Spenner (1825).- Grammitis pusilla var. lasiosora

Blume, Fl. Javae 2 (1829) 110, t. 46, f 6. Type: Blume s.n.,G. Panggrango, Java

[L, lecto. (Parris, 1986)]. - Polypodium lasiosorum (Blume) Hook., Sp. Fil. 4

(1863) 166. - Grammitis lasiosora (Blume) Ching, Bull. Fan Mem. Inst. Biol.

10 (1941) 15. Grammitis multifolia Copel., Philipp. J. Sci. 46 (1931) 219. Type:

Copeland s.n., G. Panggerango, Java [MICH, lecto. (Parris, 1986); BM, BO,

UC, isolecto.]. - Polypodium multifolium (Copel.) C.Chr., Index Fil. Suppl.

3 (1934) 154.

18. Radiogrammitis ornatissima (Rosenst.) Parris, comb. nov.

244 Gard. Bull. Singapore 58 (2) 2007

Polypodium ornatissimum Rosenst., Repert. Spec. Nov. Regni Veg. 5 (1908)

41. Type: Rosenstock’s Filices Novoguineensis Exsiccatae 9, Werner 47, Mt

Gelu, Madang, Papua New Guinea [UC, lecto. (Parris, 1983); B, BM, E,

S, isolecto.]. - Grammitis ornatissima (Rosenst.) Copel., Philipp. J. Sci. 80

(1952) 195. - Polypodium ornatissimum var. dichotomum Brause, Bot. Jahrb.

Syst. 180 (1920) 56. Type: Ledermann 11809, Mt Schrader, Madang, Papua

New Guinea [B, lecto. (Parris, 1983); B, BM, S, isolecto.].

19. Radiogrammitis parva (Brause) Parris, comb. nov.

Polypodium parvum Brause, Bot. Jahrb. Syst. 49 (1912) 36, f. 2C. Type:

Schlechter 17156, Kani Mts, Morobe, Papua New Guinea [B, lecto. (Parris

1983); BISH, BM, K, L, MICH, UC, isolecto.]. - Grammitis parva (Brause)

Copel., Gen. Fil. (1947) 211.

20. Radiogrammitis peninsularis (Copel.) Parris, comb. nov.

Grammitis peninsularis Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Holttum SFN 20678, G. Tahan, Pahang, Peninsular Malaysia (MICH, holo.;

SING, UC, iso.).

21. Radiogrammitis plana (Alderw.) Parris, comb. nov.

Polypodium planum Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 16 (1914)

32. Type: Matthew 501, G. Singgalang, Sumatra (BO, lecto., chosen here).

- Grammitis plana (Alderw.) Parris, Fern Gaz. 12: 180 (1981).

22. Radiogrammitis pseudoxiphopteris (Parris) Parris, comb. nov.

Grammitis pseudoxiphopteris Parris, Hooker’s Icon. Pl. 40 (1990) t. 3975.

Type: Dransfield 837B, G. Belumut, Johor, Peninsular Malaysia (KLU,

holo.).

23. Radiogrammitis reducta (Alderw.) Parris, comb. nov.

Polypodium reductum Alderw., Nova Guinea 14 (1924) 41. Type: Lam

1550, Mt Doorman, Papua [BO, lecto. (Parris, 1983); BM, K, L, MICH, UC,

isolecto.]. - Grammitis reducta (Alderw.) Copel., Philipp. J. Sci. 80 (1952)

167.

24. Radiogrammitis setigera (Blume) Parris, comb. nov.

Polypodium setigerum Blume, Enum. Pl. Javae (1828) 123. Type: Blume s.n.,

G. Gede, Java [L, lecto. (Parris 1983); K, L, isolecto.].- Grammitis fasciculata

Blume, Fl. Javae 2 (1829) 112, nomen illegit. superfl. pro P. setiger'um Blume.

— Polypodium fasciculatum C.Presl, Tentamen (1836) 180, nomen illegit.

superfl. pro P. setigerum Blume. — Polypodium intromissum Christ, Verh.

Natur. Gesell. Basel 11 (1896) 440. Type: Sarasin 1368, Wawokaraeng,

Five new genera and three new species of Grammitidaceae 2S)

Sulawesi [BAS, lecto. (Parris, 1983)].- Grammitis intromissa (Christ) Parris,

Fern Gaz. 12 (1981) 180. - Polypodium heanophyllum Copel., Philipp. J.

Sci. 40 (1929) 310. Type: Copeland's Pteridophyta Philippensis Exsiccata 158

[Sphalm. 185], Mt Matutum, Mindanao, Philippines [MICH, lecto. (Parris

1983); BM, K, SING, UC, isolecto.). - Grammitis setigera (Blume) Ching,

Bull. Fan Mem. Inst. Biol. 10 (1941) 241 non J.Sm. (1875). - Grammitis

cyclosora Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type: Clemens s.n.,

Mt Sarawaket, Morobe, Papua New Guinea (MICH, holo.). - Grammitis

latifolia DeVol, Fl. Taiwan 1 (1975) 223. Type: Yamamoto s.n., Taitungshan,

Taitung, Taiwan (TAI, n.v.).

25. Radiogrammiitis silvicola (Parris) Parris, comb. nov.

Grammitis silvicola Parris, Blumea 29 (1983) 141, f. 22,2. Type: Croft 134, Mt

Piora, Eastern Highlands, Papua New Guinea (K, holo.; K, LAE, iso.).

26. Radiogrammitis subpinnatifida (Blume) Parris, comb. nov.

Polypodium subpinnatifidum Blume, Enum. Pl. Javae (1828) 129. Type:

Blume s. n., Java (K, L,n.v.).- Grammitis subpinnatifida (Blume) Blume, FI.

Javae 2 (1830) 118. - Xiphopteris subpinnatifida (Blume) Copel., Gen. Fil.

(1947) 215.

27. Radiogrammitis tehoruensis (M.Kato & Parris) Parris, comb. nov.

Grammitis tehoruensis M.Kato & Parris, J. Fac. Sci. Univ. Tokyo 3, 15 (1992)

131, f. 4. Type: Kato, Ueda & Fanani C-14155, Muselleinan Pass, Seram,

Moluccas (TI, holo.).

28. Radiogrammitis tuberculata (Parris) Parris, comb. nov.

Grammitis tuberculata Parris, Blumea 29 (1983) 99, f. 15,1. Type: Vink 17495,

Mt Ambua, Enga, Papua New Guinea (L, holo.).

Tomophyllum

‘ Tomophyllum (E.Fourn.) Parris, gen. et stat. nov.

Polypodium Sect. Tomophyllum E.Fourn., Ann. Sci. Nat. 5, 18 (1873) 283.

Type species: 7! subsecundodissectum (Zoll.) Parris.

Rhizomes radial, stipes in whorls, not articulated to rhizome. Rhizome scales

pale to medium brown, not clathrate, not iridescent, usually subglossy, rarely

dull, with pale to medium brown hairs on margin and/or apex, cells usually

not turgid, sometimes turgid, without cross-walls; scales sometimes absent.

Hairs simple and branched, whitish or pale to medium brown. Laminae deeply

246 Gard. Bull. Singapore 58 (2) 2007

pinnately lobed to tripinnatifid; venation free, veins pinnately branched in

pinnae and in pinnules of more dissected species, ends of vein branches

with hydathodes on adaxial surface of lamina. Sori usually in two rows,

rarely in one row, on pinnae, sometimes in one or two rows on pinnules in

more dissected fronds, superficial, more or less circular to elliptic; sporangia

glabrous or rarely with one to three hairs at apex of sporangium adjacent to

annulus (7: congregatifolium).22 species, India to Melanesia.

The generic name refers to the pinna dissection of the type species.

Species of Tomophyllum sampled by Ranker et al. (2004) are in the upper

part of Clade III. Scleroglossum, also in Clade III, differs from Tomophyllum

in its simple lamina, with sori deeply sunken in two grooves, one each side

of the mid-vein. Calymmodon, also in Clade III, has the same type and

colour of hairs as Tomophyllum and most species of Calymmodon also have

hydathodes, but the veins of Calymmodon are simple, rather than pinnately

or bipinnately branched, and each fertile pinna bears a solitary sorus that is

protected by a fold of the basiscopic pinna margin, rather than two or more

sori that are not protected. See under Dasygrammitis, above, for differences

between it and Jomophyllum.

1. Tomophyllum beleense (Copel.) Parris, comb. nov.

Ctenopteris beleensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. Type:

Brass 11041, Bele R., Papua (MICH, holo.; A, n.v.; UC, iso.).

2. Tomophyllum bipinnatifidum (Baker) Parris, comb. nov.

Polypodium bipinnatifidum Baker, J. Bot. 28 (1890) 109. Type: Macgregor

s.n., Owen Stanley Range Central, Papua New Guinea (K, holo.). — Cten-

opteris bipinnatifida (Baker) Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. —

Ctenopteris sesquipinnata Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. Type:

Brass 9510, L. Habbema, Papua (MICH, holo.; A, 2.v., K, UC, iso.).

3. Tomophyllum bipinnatum (Copel.) Parris, comb. nov.

Ctenopteris bipinnata Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. Type:

Brass 9433, L. Habbema, Papua (MICH, holo.; A, 1.v., K, UC, iso.).

4. Tomophyllum brachyphlebium (Baker) Parris, comb. nov.

Polypodium brachyphlebium Baker, Ann. Bot. 8 (1894) 129. Type: Hancock

49, Barisan Ra., Bengkoelen, Sumatra (K, holo.).

5. Tomophyllum brooksiae (Alderw.) Parris, comb. nov.

Polypodium brooksiae Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 23 (1916)

Five new genera and three new species of Grammitidaceae 247

19, t.2.f.2a & b. Type: Brooks 265/S, Lebong Tandai, Bengkoelen, Sumatra

(BO, n.v.; WELT). - Ctenopteris brooksiae (Alderw.) Parris, Fern Gaz. 12

(1981) 180.

6. Tomophyllum capillatum (Brause) Parris, comb. nov.

Polypodium capillatum Brause, Bot. Jahrb. Syst. 49 (1912) 39, f. 2, g. Type:

Schlechter 18032, Finisterre Mts, Madang, Papua New Guinea (B, lecto.,

chosen here).

7. Tomophyllum congregatifolium (Alderw.) Parris, comb. nov.

Polypodium congregatifolium Alderw., Nova Guinea 14 (1924) 47. Type:

Lam 1544, Mt Doorman, Papua (BO, n.v., K, SING, UC). - Ctenopteris

congregatifolia (Alderw.) Copel., Philipp. J. Sci. 81 (1953) 112.

8. Tomophyllum epaleatum (Parris) Parris, comb. nov.

Ctenopteris epaleata Parris, Fern Gaz. 16 (2002) 239. Type: Faden 76/282,

Horton Plains, Nuwara Eliya, Sri Lanka (K, holo.; GH, UC, US, iso.).

9. Tomophyllum hymenophylloides (Parris) Parris, comb. nov.

Ctenopteris hymenophylloides Parris, Kew Bull. 59 (2004) 221, f. 1 A. Type:

Edwards 4061 et al., Mt Jaya, Papua (K, holo.; A, n.v., BISH, n.v., BO, 1.v.,

CANB, n.v., FRE, n.v., L,n.v., MAN, n.v., all iso.).

10. Tomophyllum inconspicuum (Blume) Parris, comb. nov.

Polypodium inconspicuum Blume, Enum. Pl. Javae (1828) 130. Type: Blume

s.n., Java (L). - Ctenopteris inconspicua (Blume) Copel., Philipp. J. Sci. 81

(1953) 103. - Polypodium gedeense Alderw., Bull. Jard. Bot. Buitenzorg, sér.

2,16 (1914) 33. Type: Raciborski s.n.,G. Gede, Java (BO, n.v.).

11. Tomophyllum macrum (Copel.) Parris, comb. nov.

Polypodium macrum Copel., Perkins’ Fragm. (1905) 189. Type: Copeland

1016, Mt Apo, Mindanao, Philippines (MICH, lecto., chosen here). -

Ctenopteris macra (Copel.) Copel., Fern Fl. Philipp. 3 (1960) 529.

12. Tomophyllum millefolium (Blume) Parris, comb. nov.

Polypodium millefolium Blume, Enum. Pl. Javae (1828) 134. Type: Blume

s.n., Java (L, n.v.).- Grammitis millefolia (Blume) Ching, Bull. Fan Mem.

Inst. Biol. 10 (1941) 240. - Ctenopteris millefolia (Blume) Copel., Gen. Fil.

(1947) 219.

13. Tomophyllum minutum (Blume) Parris, comb. nov.

Polypodium minutum Blume, Enum. PI. Javae (1828) 130. Type: Blume s.n.,G.

248 Gard. Bull. Singapore 58 (2) 2007

Gede, Java (Ln.v.). Ctenopteris minuta (Blume) Holttum, Rev. Fl. Malaya 2

(1955) 228. Polypodium callophyllum C.H.Wright, Bull. Misc. Inform., Kew

1909 (1909) 362. Type: Matthew s.n., G. Hijau, Perak, Peninsular Malaysia

(K, holo.; K, iso.).

14. Tomophyllum perplexum (Parris) Parris, comb. nov.

Ctenopteris perplexa Parris, Fern Gaz. 16 (2001) 201. Type: Sledge 1330 p.p..

Kandapola, Nuwara Eliya, Sri Lanka (K, holo.).

15. Tomophyllum polytrichum (Copel.) Parris, comb. nov.

Ctenopteris polytricha Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. Type:

Brass 10518, L. Habbema, Papua (MICH, holo.; A, n.v., UC, iso.).

16. Tomophyllum repandulum (Mett.) Parris, comb. nov.

Polypodium repandulum Mett., Abhandl. Senckenb. Naturforsch. Ges. 2

(1857) 50. Type: Gardner 58, Sri Lanka (BM, OXF). - Ctenopteris repandula

(Mett.) C.Chr. & Tardieu in Lecompte, Fl. Gen. Indo-china 7 (1941) 533. -

Polypodium subfalcatum var. glabrum Beddome, Ferns Brit. India (1866)

t. 189, f. B. Type: Thwaites/Gardner C.P. 1290, Sri Lanka [BM, /ecto. (Parris,

2002b); B, BM, CGE, E, K, L, P, PDA, UC, all isolecto.].

17. Tomophyllum sakaguchianum (Koidzumi) Parris, comb. nov.

Polypodium sakaguchianum Koidzumi, Acta Phytotax. Geobot. 1 (1932)

29. Type: Sakaguchi s.n., Kawakamimura Shiwonoha, Honto, Japan (KYO,

n.v.).- Ctenopteris sakaguchiana (Koidzumi) H.Ito, J. Jap. Bot. 11 (1935) 90.

- Grammitis sakaguchiana (Koidzumi) Sugimoto [ex Kurata], Hokuriku J.

Bot. 6 (1957) 9.

18. Tomophyllum sinicum (Christ) Parris, comb. nov.

Polypodium sinicum Christ, Bull. Boiss. 7 (1899) 3. Type: Henry 10186, Mt

Teng Chen Len, Yunnan, China, (K).- Polypodium convolutum Baker, Bull.

Misc. Inform., Kew 1906 (1906) 12. Type: Henry 10186, Ten Chen Liu Mts,

Yunnan, China (K, holo.). - Polypodium trichophyllum Baker, Bull. Misc.

Inform., Kew 1906 (1906) 13. Type: Hancock 153, Red R., Yunnan, China

(K, holo.). — Polypodium tenuissimum Hayata, Icon. Pl. Formosa 4 (Nov.

1914) 254 non Copel. (Feb. 1914). Type: Hayata & Sasaki sn., Arisan, Tatwan

(TI, n.v.; photo. PE). — Polypodium hayatae Masamune, J. Soc. Trop. Agric.

Formosa 2 (1930) 31, nomen novum pro P. tenuissimum Hayata (Nov. 1914)

non Copel. (Feb. 1914). - Ctenopteris hayatae (Masamune) H.Ito, J. Jap. Bot.

11 (1935) 90.

19. Tomophyllum subfalcatum (Blume) Parris, comb. nov.

Five new genera and three new species of Grammitidaceae 249

Polypodium subfalcatum Blume, Enum. Pl. Javae (1828) 130. Type: Java

(L). - Ctenopteris subfalcata (Blume) Kunze, Bot. Zeit. 6 (1848) 120. -

Grammitis subfalcata (Blume) Ching, Bull. Fan Mem. Inst. Biol. 10 (1940)

16. - Polypodium subfalcatum var. f Blume, Enum. Pl. Javae (1828) 130.

Type: collector unknown, Sulawesi (L).

20. Tomophyllum subminutum (Alderw.) Parris, comb. nov.

Polypodium subminutum Alderw., Malayan Ferns (1908) 598. Type: Forbes

s.n., Java (BO, n.v.). - Ctenopteris subminuta (Alderw.) Holttum, Rev. FI.

Malaya 2 (1955) 228.

21. Tomophyllum subrepandulum (Christ) Parris, comb. nov.

Polypodium subrepandulum Christ, Ann. Jard. Bot. Buitenzorg, sér. 2, 5

(1905) 119. Syntypes: Hallier 3138 (SAR), Hallier 3299 (K), Borneo.

21. Tomophyllum subsecundodissectum (Zoll.) Parris, comb. nov.

Polypodium subsecundodissectum Zoll., Syst. Verz. (1854) 37, 48. Type:

Zollinger 1578,Java (G,n.v.).- Ctenopteris subsecundodissecta (Zoll.) Copel.,

Philipp. J. Sci. 81 (1953) 114. — Polypodium subfalcatum var. semiintegrum

Copel., Philipp. J. Sci., C. Bot. 2 (1907) 138. Type: Merrill 5971, Mt Halcon,

Mindoro, Philippines (MICH, lecto., chosen here). — Polypodium secundum

Ridl., Trans. Linn. Soc. London, Bot. 2, 9 (1916) 262. Type: Kloss s.n., Mt

Carstensz, Papua (BM).

22. Tomophyllum walleri (Maiden & Betche) Parris, comb. nov.

Polypodium walleri Maiden & Betche, J. & Proc. Roy. Soc. New South Wales

35 (1911) 799. Type: Waller s.n., Herberton, Queensland, Australia [NSW,

lecto. (Parris, 1998b); BRI, iso.]. - Ctenopteris walleri (Maiden & Betche)

S.B.Andrews, Austrobaileya | (1977) 12.

Xiphopterella

- Xiphopterella Parris, gen. nov.

Typus generis: X. hieronymusii (C.Chr.) Parris

Rhizomata radialia, stipitibus in verticillis, non articulatis. Paleae pallidae

brunneae ad mediae brunneae, non clathratae, non iridescentes, subpolitae,

glabrae vel ad apicem pili uno vel duobus, cellulis plerumque non turgidis,

raro turgidis, sine septis. Pili integri et ramosi, albidi vel pallidi brunnei ad

medi brunnei. Laminae pinnatifidae vel pinnatae; venae non anastomosantes,

plerumque dichotome uni-ramosae, raro dimidiate bi-ramosae, extremis

venarum interdum hydathado in superficie. Sori in duobus serialibus, uno

utroque rhachidis, plerumque uno, raro duo, in pinnam, in lamina non

250 Gard. Bull. Singapore 58 (2) 2007

immersi, rotundi vel elliptici; sporangia glabra. Species 6, Malesiae incolae.

Rhizomes radial, stipes in whorls, not articulated to rhizome. Rhizome scales

pale to medium brown, not clathrate, not iridescent, subglossy, glabrous

or with one or two simple unjointed-hairs or branched hairs at apex, cells

without cross-walls. Hairs simple and branched, whitish or pale to medium

brown. Laminae deeply pinnately lobed to pinnate; venation free, veins

usually dichotomously one-branched, rarely dimidiately two-branched, ends

of vein branches sometimes with hydathodes on adaxial surface of lamina.

Sori in two rows, one each side of rachis, usually one, rarely two, per pinna,

superficial, + circular to elliptic; sporangia glabrous. Six species, in Malesia.

The generic name is a diminutive of Xiphopteris, in which the species were

formerly placed.

No species of Xiphopterella were sampled by Ranker et al. (2004), so its

position in their clade system is unknown. On the basis of rhizome form and

hair types, it is likely to be in Clade III. Xiphopterella resembles Calymmodon

(in Clade ITI) in having similar rhizome scales, similar frond dissection and

one row of sori on each side of the rachis, but has the veins forked rather

than simple and does not have the sori protected by the upward-folded

basiscopic margin of the pinna. Micropolypodium Hayata s.str. (Old World

species only) resembles Xiphopterella in having a radial rhizome and one

sorus per pinna, but has rhizome scales always glabrous, lacks branched

hairs and has dark brown simple hairs.

Xiphopterella gracilis Parris, sp. nov.

Type: Holttum SFN 20673, c. 1520 m (5000 ft) alt., 30 Aug 1928, G. Tahan,

Pahang, Peninsular Malaysia (SING, holo.; BM, n.v., K, iso).

— Xiphopteris alternidens sensu Parris (1986), Parris & Edwards (1988) &

Parris & Latiff (1997) non (Ces.) Copel.

Species X. alternidenti similis aspectu et rheophytica item, sed stipitibus

atris non mediis brunneis, pinnis 18-26 paribus non 8-10, textura coriacea

non tenuiter coriacea, pilis integris non ramosis, rhachidi in superficie plus

minusve immersa non prominenti, hydathodis praesentibus non carentibus

differt.

Stipes winged to base or 1-2 x c.0.2 mm; glabrous. Laminae 35-87 x 2(-3) mm,

linear-oblanceolate, obtuse to bluntly acute at apex, long-attenuate at base,

pinnate or deeply pinnately divided to wing c. 0.1 mm wide along rachis,

pinnae 18-26 pairs, longest pinnae 1 x 1-2 mm, obovate, oblong, triangular

Five new genera and three new species of Grammitidaceae 251

or broadly triangular, obtuse to bluntly acute at apex, entire; glabrous or

with simple jointed hairs (c. 0.1 mm long) occasional to sparse on abaxial

surface of rachis; rachis slightly prominent to prominent and concolorous to

slightly darker on abaxial surface of lamina, slightly sunken and darker on

adaxial surface; lateral veins invisible in transmitted light, not prominent,

simple in sterile fronds and some fertile fronds or 1-forked in fertile fronds,

acroscopic branch + as long as basiscopic branch, not extending beyond

sorus, vein endings with hydathodes. Sori 1.1-1.8 x 1.0-1.3 mm, slightly

sunken in shallow depressions which may appear slightly prominent on

adaxial surface of lamina, discrete when mature, on 9-18 pairs of pinnae, in

apical 1/4 to 2/3 of lamina, one per pinna, more or less throughout pinnae,

nearer to rachis than margin.

Distribution: Peninsular Malaysia, Pahang, (G. Tahan), known only from the

type locality.

Specimens examined in addition to type: Peninsular Malaysia, Pahang, G.

Tahan, 1520-1680 m (5000-5500 ft), 12 July1905, Wray & Robinson 5522

(SING); ibid, July 1911, Ridley 15975 (SING); G. Tahan, 1520 m (5000 ft), 15

June 1922, Haniff & Nur SFN 7989 (SING).

1. Xiphopterella alternidens (Ces.) Parris, comb. nov.

Polypodium alternidens Ces., Atti Ac. Napoli 7 (1876) 25, t. 2, f. 4. Type:

Beccari s.n., G. Gading, Sarawak, Malaysia (K). — Xiphopteris alternidens

(Ces.) Copel., Gen. Fil. (1947) 215. — Polypodium murudense Copel., Philipp.

J. Sci. C, Bot. 12 (1917) 61. Type: Bur. Sci. 2924, G. Murud, Sarawak, Malaysia

(PNH7). - Xiphopteris murudensis (Copel.) Copel., Gen. Fil. (1947) 215.

2. Xiphopterella govidjoaensis (Brause) Parris, comb. nov.

Polypodium govidjoaense Brause, Bot. Jahrb. Syst. 49 (1912) 41, f.2, K. Type:

Schlechter 19747, Govidjoa, Madang or Morobe, Papua New Guinea (B,

lecto., chosen here). — Xiphopteris govidjoaensis (Brause) Copel., Gen. Fil.

(1947) 215.-— Xiphopteris exilis Parris, Kew Bull. 41 (1986) 503. Type: Holttum

SFN 10680, G. Belumut, Johor, Peninsular Malaysia, (K, holo.; SING, iso.).

3. Xiphopterella hecistophylla (Copel.) Parris, comb. nov.

Polypodium hecistophyllum_ Copel., Philipp. J. Sci. 56 (1935) 477, t. 8, f. 1.

Type: Clemens 40837, Mt Kinabalu, Sabah, East Malaysia [MICH, lecto.

(Parris et al., 1992); BM, K, UC, isolecto.]. — Xiphopteris hecistophylla

(Copel.) Copel., Gen. Fil. (1947) 215.

4. Xiphopterella hieronymusii (C.Chr.) Parris, comb. nov.

Polypodium sertularioides J.sSm. ex Hieron., Hedwigia 44 (1905) 98 non

ae Gard. Bull. Singapore 58 (2) 2007

Baker (1876). Syntypes: Cuming 380, Malacca, Peninsular Malaysia (B, n.v.,

K) & Griffith s.n. (B), Wray in herb. Mus. Perak 1551,G. Beremban, Pahang,

Peninsular Malaysia (B,n.v., UC). — Polypodium hieronymusii C.Chr., Index

Fil. (1906) 533, nomen novum pro P. sertularioides J.Sm. ex Hieron. (1905)

non Baker (1876). — Xiphopteris hieronymusii (C.Chr.) Holttum, Rev. FI.

Malaya 2 (1955) 218.

5. Xiphopterella sparsipilosa (Holttum) Parris, comb. nov.

Ctenopteris sparsipilosa Holttum, Gardens’ Bull. 11 (1947) 268. Type: Holttum

SFN 23522, G. Batu Brinchang, Pahang, Peninsular Malaysia (SING, holo.;

K, UC, iso.). — Xiphopteris sparsipilosa (Holttum) Holttum, Rev. Fl. Malaya

2 (1955) 219.

Oreogrammitis

The genus Oreogrammitis was established by Copeland (1917) for one

species, O. clemensiae Copel. [Type: Clemens 10618 p. p., Mt Kinabalu,

Sabah, Malaysia (MICH, lecto. (Parris, 1983); BM, isolecto.)], then known

only from Mount Kinabalu in Sabah, East Malaysia.

The brief generic description reads ‘Frondibus confertis, stipitatis, simplicibus,

integris, parvis; venis (nisi soriferis) liberis; soro costa parallelo et proximo,

lineare, superficiale, nudo; paraphysibus nullis. Genus Eupolypodium

derivatum.

The description of O. clemensiae gives a few more details: “Rhizomate

adscendente, brevissimo, paleis pallido-fuscis lanceolatis vel ovatis acutis 2

ad 3 mm longis dense obtecto; stipitibus filiformibus, castaneis, 1.0 ad 2.5 cm

altis; fronde lineari-oblanceolata, 1.5 ad 3.0 mm lata, 2 ad 3 cm longa, obtusa,

deorsum sensim attenuata, subcucullata, subcoriacea, fusca, pilis sparsissimis

concoloribus mox deciduis vestita; venis inconspicuis, simplicibus vel furcatus;

soris usque 12 mm longis, subapicalibus, lateribus frondis deflexis protectis.’

Copeland regarded his new genus as being related to Scleroglossum, but

differing in its superficial, rather than sunken sori. He did not mention

that the sporangia were setose, nor that the rhizome scales were glabrous.

Christensen (1929) retained Oreogrammitis clemensiae as a dubious taxon

known only from the type collection, commenting on the scanty material

not permitting a detailed examination and being unable to form a definite

idea of its systematic position. He noted that its scales were like those of

Scleroglossum, but it seemed very different in its filiform long stipes, long

brown hairs along the edges and superficial sori. Christensen & Holttum

(1934) also retained the genus after Holttum had collected the species again

on Mount Kinabalu, mentioning that the species was distinct, but the validity

Five new genera and three new species of Grammitidaceae 253

of the genus was very problematical.

Parris made three further collections of the species from Mount Kinabalu

(Parris et al., 1992) and noted (Parris, 1983) that ‘Oreogrammitis clemensiae

was previously placed in a separate monotypic genus because of its fused

sori (Copeland, 1917). Collections from the type locality show all degrees of

soral fusion from complete to all sori (11 per row) discrete when young and

confluent when mature, and the genus is not worthy of recognition. Fused

sori are occasionally also found in the circum-antarctic species Grammitis

poeppigiana (Mett.) Pic. Serm., to which O. clemensiae is not closely related.

The genus Scleroglossum has fused sori, but O. clemensiae cannot be

regarded asa link between Grammitis and Scleroglossum as postulated by

Copeland (1917). Examination of the rhizome and the frond hairs shows

that O. clemensiae has no near relationship to Scleroglossum.’

The acceptance that Grammitis Sw. s./. is an artificial genus based on frond

dissection (Parris, 1997a, 1998a; Ranker et al., 2004) and the restriction of

Grammitis to the black-margined taxa related to and including the type

of the genus, G. marginella (Sw.) Sw., necessitates the removal from it of

numerous Old World species. Oreogrammitis is an available name for those

that are related to O. clemensiae, with the following characters.

Rhizomes dorsiventral, stipes in two rows, sometimes articulated to

rhizome, sometimes with prominent phyllopodia. Rhizome scales pale to

dark brown or blackish, usually not clathrate, rarely clathrate, usually not

iridescent, rarely iridescent, usually subglossy, rarely dull or glossy, glabrous,

cells usually not turgid, rarely subturgid or turgid, without cross-walls

except in O. dolichosora. Hairs usually simple, rarely branched. Laminae

usually simple, rarely pinnatifid or pinnate; venation usually free, rarely

with occasional anastomoses, veins usually dichotomously one-two(-four)-

branched, rarely pinnately branched, ends of vein branches sometimes with

hydathodes on adaxial surface of lamina. Sori in two (to six or more) rows

on + simple laminae, one (to three or more) on each side of the mid-vein, in

two rows, one on each side of pinna mid-vein when pinnate, superficial or

slightly sunken in broad shallow depressions, rarely deeply sunken in steep-

sided pits, on abaxial surface of lamina, + circular to narrowly elliptic; first-

developed sporangia of each sorus characteristically with one to nine simple

eglandular hairs clustered at apex of sporangium adjacent to annulus; later-

developed sporangia of each sorus glabrous. 110 species, India to Polynesia.

The generic name refers to the montane habitat of the type species and

to Grammitis, in which the species were formerly placed. Species of

254 Gard. Bull. Singapore 58 (2) 2007

Oreogrammitis sampled by Ranker et al. (2004) are in the upper part of

Clade Ia. See under Radiogrammitis above for differences between it and

Oreogrammiitis.

Oreogrammitis kunstleri Parris, sp. nov.

Type: Kunstler s.n., April 1880, G. Ledang, Johor, Peninsular Malaysia, (E,

holo.).

Species aspectu O. reinwardtio similis, sed pilis brevioribus densioribus, venis

non manifestis, ramis venarum in longitudine aequalibus non ramo acros-

copico multo breviore quam ramo basiscopico et hydathodis destitutis non

praesentibus differt.

Stipes not articulated to rhizome, phyllopodia absent. Stipes 3-6 x 0.3-0.6

mm; with dense dark red-brown simple unjointed hairs 0.5-0.8 mm long.

Laminae c. 128 x 6 mm, linear-oblanceolate, bluntly acute at apex, long-

attenuate at base, slightly crenulate, teeth to 0.2 mm long; with dark red-

brown simple unjointed hairs 0.4-1.1 mm long frequent to dense on abaxial

surface of lamina, frequent to dense on abaxial surface of mid-vein, frequent

to dense on margin, frequent on adaxial surface of lamina and frequent on

adaxial surface of mid-vein; mid-vein slightly prominent and concolorous

on abaxial surface of lamina, not evident on adaxial surface; lateral veins

invisible in transmitted light, not prominent on either surface, 1-forked,

acroscopic branch extending beyond sorus, more or less as long as basiscopic

branch, vein endings without hydathodes. Sori 1.7-2.5 x 1.3-1.9 mm, on

surface of lamina or slightly sunken in broad shallow depressions, discrete

when mature, in apical 2/3 of lamina, c. 33 in each row, nearer to mid-vein

than margin. Sporangia setose.

Distribution: Peninsular Malaysia, Johor (G. Ledang), known only from the

type collection.

Oreogrammitis malayensis Parris, sp. nov.

Type: Nor Ezzawanis A. T: FRI 52316,ca 1760 m, 12 Aug 2006, Telekom road

to Gunung Ulu Kali, Pahang/Selangor, Peninsular Malaysia (KEP, holo.).

— Grammitis sp. 9 in Parris & Latiff (1997).

Species aspectu. Radiogrammitidi peninsulari simillima, sed rhizomate

dorsiventrale non radiale, pilis in pagina inferiore laminae brevioribus et pilis

sporangiorum longioribus differt.

Stipes sometimes articulated to rhizome, phyllopodia sometimes present.

Five new genera and three new species of Grammitidaceae 255

Stipes 1-7 x 0.2-0.3 mm; with dense dark red-brown simple unjointed hairs

0.3-0.7 mm long. Laminae 26-60 x 2-6 mm, narrowly oblanceolate to linear-

oblanceolate, bluntly acute to obtuse at apex, long-attenuate at base, entire

or slightly crenulate; with dark red-brown simple unjointed hairs 0.2-0.9

mm long scattered to dense on abaxial surface of lamina, sparse to frequent

on abaxial surface of mid-vein, frequent to dense on margin, occasional

to frequent on adaxial surface of lamina and occasional to frequent on

adaxial surface of mid-vein; mid-vein slightly prominent to prominent on

abaxial surface of lamina, not prominent to slightly prominent on adaxial

surface, concolorous on both surfaces; lateral veins invisible in transmitted

light, sometimes slightly prominent and concolorous on adaxial surface

of lamina, not prominent on abaxial surface, 1-2-(3-) forked, acroscopic

branch extending beyond sorus or not, shorter than basiscopic branch, vein

endings without hydathodes. Sori 1.3-3.3 x 1.2-2.5 mm, on surface of lamina

or slightly sunken in broad shallow depressions, discrete to confluent when

mature, in apical 1/3 to 1/2 of lamina, 4-18 in each row, nearer to mid-vein

than margin. Sporangia setose.

Distribution: Peninsular Malaysia, known from Pahang/Selangor and

Johor.

Specimens examined in addition to type: Peninsular Malaysia, Pahang/

Selangor, Telekom road to G. Ulu Kali, ca 1760 m, 12 Aug. 2006, Nor

Ezzawanis A. T. FRI 52318, 52328 (both KEP); Johor, G. Belumut, 910 m

(3000 ft), 25 May1923, Holttum SFN 10860 bis (MICH, SING); G. Ledang,

ca 1070 m (3500 ft), 21 April 1930, Holttum SFN 23701 (MICH, SING, UC);

G. Mering, Ridley 3354 p p. (SING).

1. Oreogrammitis adspersa (Blume) Parris, comb. nov.

Polypodium adspersum Blume, Enum. Pl. Javae (1828) 123 non Schrad.

(1818). Type: Blume s.n., Java [L, lecto. (Parris 1983); L, isolecto.].

— Grammitis adspersa Blume, Flora Javae 2 (1830) 115, t. 48, f. 2, nomen

novum pro Polypodium adspersum Blume (1828) non Schrad. (1818). -

Polypodium sessilifolium Hook., Sp. Fil. 4 (1863) 168 non Liebm. (1849).

Type: Cuming 222, Luzon, Philippines [K, lecto. (Parris 1983); BM, CGE,

isolecto.|. — Polypodium subevenosum Baker in Hook. & Baker, Syn. Fil.

(1867) 320. Type: Mactier s.n., Penang, Peninsular Malaysia [K, holo; E,

iso.|.— Grammitis subevenosa (Baker) C.Chr. & Tardieu, Notul. Syst. (Paris)

8 (1939) 179. — Grammitis sessilifolia J.Sm., Hist. Fil. (1875) 181, nomen

novum pro P. sessilifolium Hook. (1863) non Liebm. (1849). — Polypodium

maxwellii Baker, Bull. Misc. Inform., Kew 1893 (1893) 211. Type: Hose 296,

Mt Gading, Sarawak, East Malaysia (K, holo.; E, SAR, iso.). — Grammitis

maxwellii (Baker) Parris, Fern Gaz. 12 (1980) 118.— Polypodium paucisorum

256 Gard. Bull. Singapore 58 (2) 2007

Copel., Philipp. J. Sci., C. Bot. 2 (1907) 137. Type: Merrill 5964, Mt Halcon,

Mindoro, Philippines [MICH, lecto. (Parris, 1983)].— Polypodium malaicum

Alderw., Malayan Ferns (1909) 577, nomen novum pro P. sessilifolium

Hook. (1863) non Liebm. (1849). — Grammitis malaica (Alderw.) Tagawa,

Acta Phytotax. Geobot. 8 (1939) 173, nomen superfl. illegit. pro G. sessilifolia

J.Sm. —Polypodium pilistipes Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 11

(1913) 20. Type: no collector, no locality (BO, lecto., chosen here).

2. Oreogrammitis albosetosa (F.M.Bailey) Parris, comb. nov.

Polypodium albosetosum F.M.Bailey, Rep. Govt Sci. Exped. Bellenden Ker

(1889) 78. Type: Bailey s.n., Bellenden Ker, Queensland, Australia [BRI, lecto.

(Parris, 1975); BM, BRI, K, MEL, NSW, isolecto.]. — Grammitis albosetosa

(F.M.Bailey) Parris, Bot. J. Linn. Soc. 70: 36 (1975).

3. Oreogrammitis alta (Parris) Parris, comb. nov.

Grammitis stipitata Brownlie, Nova Hedwigia, Beth. 55 (1977) 345, t. 40, f. 4

non Grammitis stipitata Proctor (1966). Type: Brownlie 1785, Mt Tomaniivi,

Viti Levu, Fiji (CHR, holo.). — G. alta Parris, N. Z. J. Bot. 18 (1980), nomen

novum pro Grammitis stipitata Brownlie (1977) non Proctor (1966).

4. Oreogrammitis antipodalis (Copel.) Parris, comb. nov.

Xiphopteris antipodalis Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Brass 10788, L. Habbema, Papua (MICH, holo.; BM, K, UC, iso.).

5. Oreogrammitis archboldii (C.Chr.) Parris, comb. nov.

Polypodium archboldii C.Chr., Brittonia 2 (1937) 305. Type: Brass 4236, Mt

Albert Edward, Central, Papua (BM, holo.; BO, BRI, NY, iso.). - Grammitis

archboldii (C.Chr.) Copel., Philipp. J. Sci. 80 (1952) 186.

6. Oreogrammitis attenuata (Kunze) Parris, comb. nov.

Grammitis attenuata Kunze, Linnaea 24 (1851) 251. Syntypes: Schmidt s.n.,

Koch s.n., Nilgiris, India (both at JE, n.v.).— Polypodium parasiticum Mett.,

Abhandl. Senckenb. Naturf. Gesell. 2 (1857) 36. Type: based on Grammitis

attenuata Kunze non Polypodium attenuatum Willd.

7. Oreogrammitis austroindica (Parris) Parris, comb. nov.

Grammitis austroindica Parris, Fern Gaz. 16 (2001) 202. Type: Herb. Beddome

s.n., Nilgiris, India (BM, holo.).

8. Oreogrammitis baldwinii (Baker) Parris, comb. nov.

Polypodium baldwinii Baker, Ann. Bot. 5 (1891) 463. Type: Baldwin s.n.,

Kauai, Hawaiian Is. (K). — Grammitis baldwinii (Baker) Copel., Philipp. J.

Bot. 80 (1952) 158. - Polypodium knudsenii Hieron., Hedwigia 44 (1905)

Five new genera and three new species of Grammitidaceae 257

79. Syntypes: Knudsen 90, Baldwin 117, Kauai, Hawaiian Is.; Knudsen 7, Mt

Halemanu, Hawaiian Is. (all at B,7.v.).

9. Oreogrammitis brassii (Copel.) Parris, comb. nov.

Grammitis brassii Copel., Philipp. J. Sci. 60 (1936) 112, t. 21. Type: Brass

2926, Hinuahaoro, San Cristobal, Solomon Is. (PNHf#, holo.; A, lecto., chosen

here; BRI, GH, MICH, isolecto.).

10. Oreogrammitis caespitosa (Blume) Parris, comb. nov.

Grammitis caespitosa Blume, Enum. Pl. Javae (1828) 115. Type: Matthew s.n.,

G. Gede, Java [K, neo. (Parris, 1983)]. — Polypodium caespitosum (Blume)

Mett., Ann. Mus. Bot. Lugd. — Bat. 2 (1866) 219.

11. Oreogrammitis ceratocarpa (Copel.) Parris, comb. nov.

Grammitis ceratocarpa Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Clemens 11349, Ulap, Morobe, Papua New Guinea (MICH, holo.; MICH,

UC, iso.).

12. Oreogrammitis cervicornis (Alderw.) Parris, comb. nov.

Polypodium cervicorne Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2,28 (1918)

39. Type: Brooks 303/S, Lebong Simpang, Bengkoelen, Sumatra (BO, lecto.,

chosen here; BM, isolecto.). — Grammitis cervicornis (Alderw.) Adelbert in

van Borssum Waalkes, Annales Bogor. 4 (1960) 25.

13. Oreogrammitis christophersenii (Copel.) Parris, comb. nov.

Grammitis christopherseniti Copel, Occ. Papers B. P. Bishop Mus. 15 (1939)

89, f. 8. Type: Christophersen & Hume 2277 p. p., Matavanu, Savaii, Samoa

(BISH, holo.; MICH, UC, iso.).

14. Oreogrammitis clavata (Parris) Parris, comb. nov.

Grammitis clavata Parris, Kew Bull. 52 (2004) 219, f. 1 C. Type: Johns 10068,

Mt Jaya, Papua (K, holo.; BO, n.v., FRE, n.v., MAN, 7.v., all iso.).

15. Oreogrammitis collina (Parris) Parris, comb. nov.

Grammitis collina Parris, Blumea 29 (1983) 174, f. 28, 3. Type: Parris 9294

[sphalm. 9224] & Croxall, Baiyer-Jimi Divide, Western Highlands, Papua

New Guinea (K, holo.; K, LAE, iso.).

16. Oreogrammitis conformis (Brack.) Parris, comb. nov.

Polypodium conforme Brack., U.S. Expl. Exped. 16 (1854) 4, t. 1, f. 2. Type:

U.S. Expl. Exped. 3, Ovalau, Fiji (K, US, n.v.).- Grammitis conformis (Brack.)

J.Sm., Hist. Fil. (1875)181.

258 Gard. Bull. Singapore 58 (2) 2007

17. Oreogrammitis congener (Blume) Parris, comb. nov.

Grammitis congener Blume, Enum. Pl. Javae (1828) 115. Type: Blume s.n.,

Java [L, lecto. (Parris, 1986)].— Polypodium congenerum (Blume) C.Presl,

Tentamen (1836) 180. — Polypodium alatellum Christ, Verh. Natur. Gesell.

Basel 11 (1897) 441. Type: F & P Sarasin 1368 p. p., Wawokaraeng, Sulawesi

(BAS, n.v.). — Grammitis brevipila Copel., Brittonia 1 (1931) 69. Type:

Williams 1581, Trinidad, Luzon, Philippines (NY, holo.; MICH, UC, iso.).

— Polypodium brevipilum (Copel.) C.Chr., Index Fil. Suppl. 3 (1934) 145.

— Grammitis duplopilosa Copel., Brittonia 1 (1931) 69. Type: Williams 1582,

Mt Santo Tomas, Luzon, Philippines (NY, holo; UC, iso.). — Polypodium

duplopilosum (Copel.) C.Chr., Index Fil. Suppl. 3 (1934) 148. — Grammitis

petrophila Copel., Philipp. J. Sci. 56 (1935) 478. Type: Clemens 50133, Mt

Kinabalu, Sabah, East Malaysia [MICH, lecto. (Parris, 1986); BM, K, UC,

isolecto.].— Grammitis hirtella var. major Holttum, Rev. Fl. Malaya 2 (1955)

216, nomen invalidum.

18. Oreogrammitis crinifera (Parris) Parris, comb. nov.

Grammitis crinifera Parris, Blumea 29 (1983) 180, f.29, 2. Type: Pulle 857, Mt

Hellwig, Papua (L, holo.).

19. Oreogrammitis crispatula (Holttum) Parris, comb. nov.

Grammitis crispatula Holttum, Gardens’ Bull. 11 (1947) 268. Type: Burkill

SFN 12705, Maxwell’s Hill, Perak, Peninsular Malaysia (SING, holo.; BM,

BO, K, iso.).

20. Oreogrammitis curtipila (Parris) Parris, comb. nov.

Grammitis curtipila Parris, Blumea 29 (1983) 112, f. 16, 4. Type: Brass 25776,

Normanby I., Milne Bay, Papua New Guinea (K, holo.; A, CANB, L, LAE,

1SO.).

21. Oreogrammitis debilifolia (Copel.) Parris, comb. nov.

Grammitis debilifolia Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Brass 10036 & Meijer-Drees, Mt Trikora, Papua (MICH, holo.; A, BM, BO,

K,L, LAR. UC iso.).

22. Oreogrammitis dictymioides (Copel.) Parris, comb. nov.

Grammitis dictymioides Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Brass 11848, L. Habbema, Papua (MICH, holo.; A, BO, L, iso.).

23. Oreogrammitis dolichosora (Copel.) Parris, comb. nov.

Polypodium dolichosorum Copel., Philipp. J. Sci. 1 Suppl. (1906) 159, t. 16.

Type: Copeland 1524, Davao, Mindanao, Philippines [MICH, lecto. (Parris,

Five new genera and three new species of Grammitidaceae 259

1983); K, NSW, UC, isolecto.]. — Grammitis dolichosora (Copel.) Copel.,

Philipp. J. Sci. 80 (1952) 188. - Polypodium ciliolatum Alderw., Nova Guinea

14 (1924) 43. Type: Lam 1518, Mt Doorman, Papua [BO, lecto. (Parris, 1983);

L, SING, frag. of L at MICH, photo. of L at BM, isolecto.). - Grammitis

ciliolata (Alderw.) Copel., Philipp. J. Sci. 80 (1952) 187.

24. Oreogrammitis dorsipila (Christ) Parris, comb. nov.

Polypodium dorsipilum Christ in Warburg, Monsunia 1 (1900) 59. Type:

Warburg s.n., South China (B, n.v.).— Grammitis dorsipila (Christ) C.Chr. &

Tardieu, Notul. Syst. (Paris) 8 (1939) 179.

25. Oreogrammitis excelsa (Parris) Parris, comb. nov.

Grammitis excelsa Parris, Blumea 29 (1983) 171, f. 27. Type: Bergman 416,

Kadubaka, Central Range, Papua (S, holo.).

26. Oreogrammitis fasciata (Blume) Parris, comb. nov.

Grammitis fasciata Blume, Enum. PI. Javae (1828) 116. Type: Blume s.n., G.

Gede, Java [L, lecto. (Parris, 1983); BO, K, L, NSW, isolecto.].— Polypodium

fasciatum (Blume) C.Presl, Tentamen (1836) 180. - Themelium fasciatum

(Blume) Parris, Kew Bull. 59 (2004) 225.— Polypodium integrum Brause, Bot.

Jahrb. Syst. 49 (1912) 37. Type: Schultze Jena (26) 21, Tami R., West Sepik,

Papua New Guinea [B, lecto. (Parris, 1983)]. — Grammitis integra (Brause)

Copel., Philipp. J. Sci. 80 (1952) 171.-— Polypodium fuciforme Rosenst., Nova

Guinea 8 (1912) 726. Type: von Romer 846, Hellwig Mts, Papua [BO, lecto.

(Parris, 1983)]. - Polypodium alcicorne Ridl., Trans. Linn. Soc. London, Bot.

2, 9 (1916) 261 non Baker (1888). Type: Kloss s.n., Mt Carstensz, Papua

[BM, lecto. (Parris, 1983); K, isolecto.]. — Polypodium dichotomum Brause,

Bot. Jahrb. Syst. 56 (1920) 190 non Houttyn (1783) nec Thunb. (1784).

Type: Ledermann 9246, Etappenberg, East Sepik, Papua New Guinea [B,

lecto. (Parris, 1983); BM, S, isolecto.]. — Grammitis angustifolia Gilli, Ann.

Naturhist. Mus. Wien 81 (1978) 24, non (Sw.) Heward (1838). Type: Gilli 165,

Mingende, Simbu, Papua New Guinea (W, holo., 7.v.).

27. Oreogrammitis fenicis (Copel.) Parris, comb. nov.

Grammiitis fenicis Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type: Ramos

Bur. Sci. 80147, Batanes Is., Philippines, (MICH, holo.; K, PNHt, SING, UC,

iSO.).

28. Oreogrammitis flavovirens (Alston) Parris, comb. nov.

Polypodium flavovirens Alston, J. Bot. 78 (1940) 228. Type: Clemens 6764,

Sambanga, Morobe, Papua New Guinea, (B, holo.).— Ctenopteris flavovirens

(Alston) Parris, Kew Bull. 41 (1986) 69. — Ctenopteris clemensiae Copel.,

260 Gard. Bull. Singapore 58 (2) 2007

Univ. Calif. Publ. Bot. 18 (1942) 225. Type: Clemens s.n., Rawlinson Range,

Morobe, Papua New Guinea (MICH, holo.; UC, iso.). — Ctenopteris

schizophlebia Copel., Univ. Calif. Publ. Bot. 18 (1942) 225. Type: Clemens

s.n., Rawlinson Ra., Morobe, Papua New Guinea (MICH, holo.; UC, iso.).

29. Oreogrammitis forbesiana (W.H.Wagner) Parris, comb. nov.

Grammitis forbesiana W.H.Wagner, Contr. Univ. Michigan Herb. 19 (1993)

70. Type: Wagner 65386, Haelaau to Puu Kukui, Maui, Hawaiian Is. (MICH,

holo., n.v.; BISH, iso., 7.v.).

30. Oreogrammitis frigida (Ridl.) Parris, comb. nov.

Polypodium frigidum Ridl., Trans. Linn. Soc. London, Bot. 2, 9 (1916) 259.

Type: Kloss s.n., Mt Carstensz, Papua [BM, lecto. (Parris, 1983)].— Grammitis

frigida (Ridl.) Copel., Univ. Calf. Publ. Bot. 18 (1942) 223. — Grammitis

caricifolia Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type: Clemens s.n.,

Mt Sarawaket, Morobe, Papua New Guinea (MICH, holo.). — Grammitis

plurisetulosa Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type: Brass 10109

& Meyer-Drees, Mt Trikora, Papua (MICH, holo.; A, BO, BRI, L, UC, iso.).

31. Oreogrammitis glabrata (Brownlie) Parris, comb. nov.

Grammitis glabrata Brownlie, Nova Hedwigia, Beih. 55 (1977) 348, t. 40, f. 1.

Type: Brownlie 800, Mt Delaikora, Vanua Levu, Fiji, (CHR, holo.).

32. Oreogrammitis glossophylla (Parris) Parris, comb. nov.

Grammitis glossophylla Parris, Blumea 29 (1983) 47, f. 3, 2. Type: Brass

28495, Rossel I., Milne Bay, Papua New Guinea (L, holo.).

33. Oreogrammitis graminifolia (Copel.) Parris, comb. nov.

Grammitis graminifolia Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Brass 5092, Mt Tafa, Central, Papua New Guinea (NY, holo.; BM, BO, BRI,

GH, MICH, iso.). — Themelium graminifolium (Copel.) Parris, Kew Bull. 59

(2004) 225.

34. Oreogrammitis habbemensis (Copel.) Parris, comb. nov.

Grammitis habbemensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type:

Brass 10520, L. Habbema, Papua (MICH, holo.; A, BO, L, iso.).

35. Oreogrammitis hispida (Copel.) Parris, comb. nov.

Grammitis hispida Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type: Brass

10257, L. Habbema, Papua (MICH, holo.; A, BM, BO, K, L, LAE, iso.).

36. Oreogrammitis hookeri (Brack.) Parris, comb. nov.

Five new genera and three new species of Grammitidaceae 261

Polypodium setigerum Hook. & Arnott., Bot. Beechey’s Voy. (1832) 103, t.

21A non Blume (1828). Type: Bidwill s.n., Hawaiian Is. (K). - Polypodium

hookeri Brack., U.S. Expl. Exped. 16 (1854) 4, nomen novum pro P. setigerum

Hook. & Arnott (1832) non Blume (1828). - Grammitis hookeri (Brack.)

Copel., Philipp. J. Sci. 80 (1952) 157. - Grammitis setigera J.Sm., Hist. Fil.

(1875) 181, nomen novum pro P. setigerum Hook. & Arnott (1832) non

Blume (1828).

37. Oreogrammitis imberbis (Parris) Parris, comb. nov.

Grammitis imberbis Parris, Blumea 29(1) (1983) 128, f. 19,3. Type: Croft &

Lelean LAE 65885, Mt Scorpion, West Sepik, Papua New Guinea (LAE,

holo.).

38. Oreogrammitis inconstans (Alderw.) Parris, comb. nov.

Polypodium inconstans Alderw., Nova Guinea 14 (1924) 43. Type: Lam

1882, Mt Doorman, Papua [BO, lecto. (Parris, 1983) ].— Grammitis inconstans

(Alderw.) Copel., Philipp. J. Sci. 80 (1952) 193.

39. Oreogrammitis insularis (Copel.) Parris, comb. nov.

Grammitis insularis Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type: Vaupel

451,Samoa (US, holo.,.v.;B,n.v., MICH, iso.).- Grammitis impressa Copel.,

Philipp. J. Bot. 80 (1952) 242, f. 95. Type: Bartlett 7964A, Dalok Soeroengam,

Tapianoeli, Sumatra (US, holo.; UC, iso.).

40. Oreogrammitis interrupta (Baker) Parris, comb. nov.

Monogramma interrupta Baker, Ann. Bot. 5 (1891) 482. Type: Macgregor

s.n., Mt Yule, Gulf, Papua New Guinea (K, holo.; BM, iso.).— Pleurogramme

interrupta (Baker) Christ, Farnkr. der Erde (1897) 55. — Nematopteris

interrupta (Baker) C.Chr., Dansk Bot. Ark. 6 (1929) 31. — Grammitis

interrupta (Baker) Copel., Gen. Fil. (1947) 214. — Polypodium pyxidiforme

Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 1 (1911) 28. Type: de Kock 45, Mt

Goliath, Papua [BO, lecto. (Parris, 1983); BM, isolecto].

41. Oreogrammitis jeraiensis (Parris) Parris, comb. nov.

Grammitis jeraiensis Parris, Hooker’s Icon. Pl. 40 (1990), t. 3976. Type:

Abdullah 615, G. Jerai, Kedah, Peninsular Malaysia (KLU, holo.).

42. Oreogrammitis kjellbergii (C.Chr.) Parris, comb. nov.

Polypodium kjellbergii C.Chr. in Kjellberg & C.Chr., Bot. Jahrb. Syst. 66

(1933) 63. Type: Kjellberg 1102, B. Watoewila, Sulawesi, (BM, BO, S, 7.v.).

43. Oreogrammitis knutsfordiana (Baker) Parris, comb. nov.

262 Gard. Bull. Singapore 58 (2) 2007

Polypodium knutsfordianum Baker, J. Bot. 28 (1890) 107. Type: Macgregor

s.n., Mt Knutsford, Central, Papua New Guinea (K, holo.). — Grammitis

knutsfordiana (Baker) Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. —

Polypodium oleandroides Baker, Ann. Bot. 8 (1894) 128. Type: Macgregor

s.n., Mt Suckling, Milne Bay, New Guinea (K, holo.).— Polypodium warburgii

Christ in Warburg, Monsunia 1| (1900) 59. Type: Warburg 17882, G. Sibella,

Bacan, Moluccas [B, lecto. (Parris, 1983)].— Polypodium setosum var. calvum

Copel., Philipp. J. Sci., C. Bot. 2 (1907) 136. Type: Merrill 5965, Mt Halcon,

Mindoro, Philippines (MICH). — Polypodium diplosoroides Rosenst., Nova

Guinea 8 (1912) 724. Type: von Romer 1167, Hellwig Mts, Papua [BO, lecto.

(Parris, 1983)]. — Grammitis nigrosetosa Copel., Univ. Calif. Publ. Bot. 18

(1942) 224. Type: Stresemann 404, G. Fogha, Buru, Moluccas (L 936.248...162,

holo.; photos at CGE, PARRIS). — Grammitis stresemannii Copel., Univ.

Calif. Publ. Bot. 18 (1942) 224. Type: Stresemann 268, Ceram, Moluccas

(L936.248...170, holo.; photos at CGE, PARRIS).

44. Oreogrammitis leonardii (Parris) Parris, comb. nov.

Grammitis leonardii Parris, Bot. J. Linn. Soc. 70 (1975) 38, f.5 A & B. Type:

Brass 20075, Mt Finnegan, Queensland, Australia (BRI, holo.; A, iso.).

45. Oreogrammitis locellata (Baker) Parris, comb. nov.

Polypodium locellatum Baker, J. Bot. 28 (1890) 108. Type: Macgregor 26,

Mt Victoria, Central, Papua New Guinea (K, holo.; BM, iso.). — Grammitis

locellata (Baker) Copel., Occ. Papers Bishop Mus. 15 (1939) 86.— Grammitis

stenocrypta Copel., Philipp. J. Sci. 46 (1931) 220. Type: Copeland s n.,G. Gede,

Java [MICH, lecto. (Parris, 1983); BM, BO, UC, isolecto.]. — Polypodium

stenocryptum (Copel.) C.Chr., Index Fil. Suppl. 3 (1934) 158. — Grammitis

stomatocarpa Copel., Univ. Calif. Publ. Bot. 18 (1942) 223. Type: Brass 9858A

& Meijer-Drees, Mt Trikora, Papua (MICH, holo.; A, BM, BO, BRI, L, iso.).

46. Oreogrammitis loculosa (Alderw.) Parris, comb. nov.

Polypodium loculosum Aldetw., Nova Guinea 14 (1924) 42. Type: Lam 1267,

Doorman R., Papua [L, lecto. (Parris, 1983); BM, isolecto.]. - Grammitis

loculosa (Alderw.) Copel., Univ. Calif. Publ. Bot. 18 (1942) 223.

47. Oreogrammitis longiceps (Rosenst.) Parris, comb. nov.

Polypodium longiceps Rosenst., Repert Spec. Nov. Regni Veg. 12 (1913) 177.

Type: Keysser B.72, Bolan Mts, Morobe, Papua New Guinea (S, 7.v., UC). -

Ctenopteris longiceps (Rosenst.) Copel., Philipp. J. Sci. 81 (1953) 103.

48. Oreogrammitis maireaui (Copel.) Parris, comb. nov.

Grammitis maireaui Copel., B. P. Bishop Mus. Occ. Papers 14 (1938) 71.

Five new genera and three new species of Grammitidaceae 263

Type: St John & Maireau 15553, Mt Perahu, Rapa, Austral Is. (BISH 498689,

lecto., chosen here; BISH 10935, K, MICH, WELT, isolecto.).

49. Oreogrammitis marivelesensis (Copel.) Parris, comb. nov.

Grammitis marivelesensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Copeland 215, Luzon, Mt Mariveles, Philippines (MICH, holo.).

50. Oreogrammitis medialis (Baker) Parris, comb. nov.

Polypodium mediale Baker in Hook. & Baker, Syn. Fil. ed. 2 (1874) 507.

Type: Wall s.n., Sri Lanka [K, lecto. (Parris, 2002b)]. — Grammitis medialis

(Baker) Ching, Bull. Fan Mem. Inst. Biol. 10 (1940) 15.

51. Oreogrammitis merrillii (Copel.) Parris, comb. nov.

Polypodium merrillii Copel., Perkins’ Fragm. (1905) 188. Type: Merrill 754,

Iwahig R., Palawan, Philippines [US, lecto. (Parris, 1983); PNH7#, isolecto.].

- Grammitis merrillii (Copel.) Copel., Philipp. J. Sci. 80 (1952) 224.

52. Oreogrammitis mesocarpa (Alderw.) Parris, comb. nov.

Polypodium mesocarpum Alderw., Nova Guinea 14 (1924) 42. Type: Lam

1878, Mt Doorman, Papua (BO, lecto., chosen here; L, isolecto.).— Grammitis

mesocarpa (Alderw.) Copel., Philipp. J. Sci. 80 (1952) 181.

53. Oreogrammitis miltiblepharis (Copel.) Parris, comb. nov.

Grammitis miltiblepharis Copel., B. P. Bishop Mus. Occ. Papers 14 (1938)

71. Type: Mumford & Adamson 586, Ooumu, Nukuhiva, Marquesas (BISH,

holo.; MICH, iso.).

54. Oreogrammitis mollipila (Baker) Parris, comb. nov.

Polypodium mollipilum Baker, J. Bot. 28 (1890) 107. Type: Macgregor s.n.,

Owen Stanley Range, Central, Papua New Guinea, (K, holo.; BM, iso.). —

Grammitis mollipila (Baker) Copel., Univ. Calif. Publ. Bot. 18 (1942) 224.

55. Oreogrammitis montana (Parris) Parris, comb. nov.

Grammitis montana Parris, Blumea 29 (1983) 167, f.26, 1. Type: Clemens s n.,

Mt Sarawaket, Morobe, Papua New Guinea (BM, holo.; MICH, iso.).

56. Oreogrammitis monticola (Sledge) Parris, comb. nov.

Grammitis monticola Sledge, Brit. Fern Gaz. 9 (1967) 344. Type: Sledge 1835,

Mt Fito, Upolu, Samoa (K, holo.).

57. Oreogrammitis murrayana (C.Chr.) Parris, comb. nov.

Polypodium murrayanum C.Chr., Brittonia 2 (1937) 304. Type: Brass 4726,

264 Gard. Bull. Singapore 58 (2) 2007

Wharton Range, Northern, Papua New Guinea (BM, holo.; BRI, MICH,

NY, iso., 7.v.). - Grammitis murrayana (C.Chr.) Copel., Philipp. J. Sci. 80

(1952) 200.

58. Oreogrammitis neocaledonica (Copel.) Parris, comb. nov.

Grammitis neocaledonica Copel., Univ. Calif. Publ. Bot. 18 (1942) 222. Type:

Schlechter 15346, New Caledonia (L, holo., 1.v.; E, K, MICH, NSW, iso.).

59. Oreogrammitis nigropaleata (Copel.) Parris, comb. nov.

Grammitis nigropaleata Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Clemens 41018, Matap, Morobe, Papua New Guinea (MICH, holo.; MICH,

UC. iso.).

60. Oreogrammitis nipponica (Tagawa & K.Iwats.) Parris, comb. nov.

Grammitis nipponica Tagawa & K.Iwats., Amer. Fern J.51 (1961) 150, f. A-C.

Type: Nakai 5052, San-no-taki Falls, Honshu, Japan (KYO, holo., .v.).

61. Oreogrammitis nuda (Tagawa) Parris, comb. nov.

Grammitis nuda Tagawa, Acta Phytotax. Geobot. 19 (1941) 284. Type:

Tagawa 2092, Tyosyu-gun, Takaio, Taiwan (KYO, holo, n.v.).

62. Oreogrammitis oblanceolata (Baker) Parris, comb. nov.

Polypodium oblanceolatum Baker,Annals Bot.8 (1894) 128. Type: Macgregor

s.n., Mt Suckling, Milne Bay, Papua New Guinea (K, holo.). — Grammitis

oblanceolata (Baker) Copel., Philipp. J. Sci. 80 (1952) 166. — Polypodium

sparsipilum Copel., Philipp. J. Sci., C. Bot. 6 (1911) 139. Type: Brooks 14,

Bengkarum, Sarawak, East Malaysia [MICH, lecto. (Parris, 1983); BM,

isolecto.|. — Grammitis sparsipila (Copel.) Parris, Fern Gaz.12 (1980) 118.

— Polypodium hirtellum var. subcalvum Alderw., Bull. Jard. Bot. Buitenzorg,

sér. 2, 28 (1918) 39. Type: Brooks 302/S, Lebong Simpang, Bengkoelen,

Sumatra (BM, BO). — Grammitis brevisetulosa Copel., Univ. Calif. Publ.

Bot. 18 (1942) 223. Type: Clemens 40969 p.p., Matap, Morobe, Papua New

Guinea (MICH, holo.).

63. Oreogrammitis oncobasis (M.G.Price) Parris, comb. nov.

Grammitis oncobasis M.G.Price, Contr. Univ. Michigan Herb. 17 (1990) 274,

f.1,D & E. Type: Price 3376, Mt Burnay, Luzon, Philippines (MICH, holo.).

64. Oreogrammitis padangensis (Baker) Parris, comb. nov.

Polypodium padangense Baker, J. Bot. 18 (1880) 213. Type: Beccari s.n., G.

Singgalang, Sumatra (K, holo.). — Grammitis padangensis (Baker) Copel.,

Philipp. J. Sci. 80 (1952) 212. — Polypodium stanleyanum Baker, J. Bot. 28

Five new genera and three new species of Grammitidaceae 265

(1890) 107. Type: Macgregor 25, Owen Stanley Range, Central, Papua New

Guinea (K, holo.; BM, iso.).— Grammitis stanleyana (Baker) Copel., Philipp.

J. Sci. 80 (1952) 168. — Polypodium durum Copel., Leafl. Philipp. Bot. 3

(1910) 837. Type: Elmer 11689, Mt Calelan, Mindanao, Philippines [MICH,

lecto. (Parris, 1983)]. — Polypodium ciliiferum Alderw., Bull. Jard. Bot.

Buitenzorg sér. 2, 16 (1914) 32. Type: Matthew 708, G. Sago, Sumatra [BO,

lecto. (Parris, 1983)].— Polypodium trichocarpum Alderw., Nova Guinea 14

(1924) 41. Type: Lam 1693, Mt Doorman, Papua [L, lecto. (Parris, 1983); BM,

L, isolecto.].— Polypodium trichocarpum var. inerme Alderw., Nova Guinea

14 (1924) 42. Type: Lam 1854, Mt Doorman, Papua [BO, lecto. (Parris, 1983);

L, isolecto.].

65. Oreogrammitis palauensis (Hosokawa) Parris, comb. nov.

Grammitis palauensis Hosokawa, Trans. Nat. Hist. Soc. Formosa 31 (1941)

475. Type: Hosokawa 9769, Sin-Gaspan, Babeldaob, Palau (TI, holo., n.v.;

BISH, iso.).

66. Oreogrammitis papuensis (Alderw.) Parris, comb. nov.

Polypodium papuanum Rid1., Trans. Linn. Soc. London, Bot. 2, 9 (1916) 260

non Baker (1886). Type: Kloss s.n., Mt Carstensz, Papua [BM, lecto. (Parris,

1983)]. — Polypodium papuense Alderw., Handbook Malayan Ferns Suppl.

(1917) 521, nomen novum pro P. papuanum Ridl. (1916) non Baker (1886).

- Grammitis papuensis (Alderw.) Parris, Fern Gaz. 12 (1980) 118.

67. Oreogrammitis pilifera (Ravi & J. Joseph) Parris, comb. nov.

Grammiitis pilifera Ravi & J.Joseph, J. Bombay Nat. Hist. Soc. 76 (1979) 348,

f. 1-5. Type: Ravi 5711, Ponmudi Hills, Kerala, India (CAL, holo., n.v.; MH,

n. v., RH, n.v., iso.).

68. Oreogrammitis pilosiuscula (Blume) Parris, comb. nov.

Grammitis pilosiuscula Blume, Enum. Pl. Javae (1828)115. Type: Java, Blume

s.n. (L, lecto., chosen here). — Polypodium pilosiusculum (Blume) C.Presl,

Tentamen (1836) 180. - Grammitis hirta Blume, Enum. Pl. Javae (1828) 115

non Humb., Bonpl. & Kunth (1815). Type: Blume s n., G. Gede, Java (L,

lecto., chosen here). — Polypodium hirtum C.Presl, Tentamen (1836) 180 pro

G. hirta Blume (1828) non Humb., Bonpl. & Kunth (1815), non P. hirtum Sw.

(1788), nec Schkuhr (1806), nec C. Presl (1825), nec Vell. (1827). — Grammitis

hirta var. B Blume, Fl. Javae 2 (1829) 111. Type: Van Hasselt s.n.,G. Karang,

Java (L).

69. Oreogrammitis piruensis (M.Kato & Parris) Parris, comb. nov.

Grammitis piruensis M.Kato & Parris, J. Fac. Sci. Univ. Tokyo 3, 15 (1992) 127,

266 Gard. Bull. Singapore 58 (2) 2007

f.2. Type: Kato, Ueda & Fanani C-12998, Tanahgoyang to G. Tiang Bendera,

Seram, Moluccas (TI, holo.).

70. Oreogrammitis pleurogrammoides (Rosenst.) Parris, comb. nov.

Polypodium pleurogrammoides Rosenst., Repert. Spec. Nov. Regni Veg.

5 (1908) 42. Type: Werner 22, Mt Gelu, Madang, New Guinea [UC, lecto.

(Parris, 1983); S, isolecto.]. — Grammitis pleurogrammoides (Rosenst.)

Copel., Occ. Papers Bishop Mus. 15 (1939) 86.- Grammitis vitiensis Brownlie,

Nova Hedwigia, Beth. 55 (1977) 346, t. 40,2 & 3. Type: Vodonaivalu 2170, Mt

Voma, Viti Levu, Fiji (CHR, holo.).

71. Oreogrammitis polita (Brause) Parris, comb. nov.

Polypodium politum Brause, Bot. Jahrb. Syst. 56 (1920) 185. Type: Ledermann

12937A p.p., Felsspitze, East Sepik, Papua New Guinea (B, lecto., chosen

here).

72. Oreogrammitis pseudolocellata (Parris) Parris, comb. nov.

Grammitis pseudolocellata Parris, Blumea 29 (1983) 189. Type: Brass 22893,

Mt Dayman, Milne Bay, Papua New Guinea (BM, holo.; A, iso.).

73. Oreogrammitis pseudospiralis (Alderw.) Parris, comb. nov.

Polypodium pseudospirale Alderw., Bull. Jard. Bot. Buitenzorg, sér. 2, 1

(1911) 29. Type: de Kock 49, Mt Goliath, Papua (BO, n.v.). — Xiphopteris

pseudospiralis (Alderw.) Parris, Alpine & Subalpine Flora Mt Jaya (2006)

118. - Polypodium roemerianum Rosenst., Nova Guinea 8 (1912) 725. Type:

von Rémer 1227, Hellwig Mts, Papua [L, lecto. (Morton, 1973)].

74. Oreogrammitis queenslandica (Parris) Parris, comb. nov.

Grammitis queenslandica Parris, Bot. J. Linn. Soc. 70 (1975) 39,f.5 E & EF

Type: Brass 18222, Speewah, Queensland, Australia (BRI, holo.; A, iso.).

75. Oreogrammitis raiateensis (J.W.Moore) Parris, comb. nov.

Polypodium raiateense J.W.Moore, B. P. Bishop Mus. Bull. 102 (1933) 10.

Type: Moore 183, Mt Temehani, Raiatea, Society Is. (BISH 498920, lecto.,

chosen here; BISH 79273, BISH 498921, isolecto.). — Grammitis raiateensis

(J.W.Moore) Copel., Occ. Papers B. P. Bishop Mus. 14 (1938) 72.

76. Oreogrammitis reinwardtii (Blume) Parris, comb. nov.

Grammitis reinwardtii Blume, Enum. Addend. (1828) 2. Type: Reinwardt

s.n., Sulawesi [L, lecto. (Parris, 1983)]. — Polypodium reinwardtii (Blume)

C.Presl, Tentamen (1836) 180. - Polypodium sucklingianum Baker, Ann.

Bot. 8 (1894) 128. Type: Macgregor s.n., Mt Suckling, Milne Bay, Papua New

Five new genera and three new species of Grammitidaceae 267

Guinea (K, holo.). — Grammitis sucklingiana (Baker) Parris, Fern Gaz. 12

(1980) 118. -— Polypodium brooksii Copel., Philipp. J. Sci., C. Bot. 12 (1917)

60, non (Copel.) C.Chr. (1913). Type: Brooks s.n., G. Bongo, Sarawak,

East Malaysia (MICH, lecto., chosen here; K, isolecto.). — Polypodium

trichopodum F.Muell. var. serratolobatum Brause, Bot. Jahrb. Syst. 56 (1920)

179. Type: Ledermann 11576, Lehm River, East Sepik, Papua New Guinea

[B, lecto. (Parris, 1983)].— Polypodium bongoense Copel., Philipp. J. Sci. 38

(1929) 153, nomen novum pro P. brooksii Copel. (1917) non (Copel. ) C.Chr.

(1913). — Grammitis bongoensis (Copel.) Copel., Philipp. J. Sci. 80 (1952)

271.— Grammitis reinwardtioides Copel., Philipp. J. Sci. 56 (1935) 478. Type:

Clemens 40792, Mt Kinabalu, Sabah, East Malaysia [MICH, lecto. (Parris

et al, 1992); BM, isolecto.]. - Grammitis mariae Copel., Univ. Calif. Publ.

Bot. 18 (1942) 223. Type: Clemens 10251A, Mt Sarawaket, Morobe, Papua

New Guinea (MICH, holo.). — Grammitis matapensis Copel., Univ. Calif.

Publ. Bot. 18 (1942) 223. Type: Clemens 11176, Matap, Morobe, New Guinea

(MICH, holo.; BM, MICH, UC, all iso.).

77. Oreogrammitis reptans (Parris) Parris, comb. nov.

Grammitis reptans Parris, Blumea 29 (1983) 156, f. 24, 1. Type: Brass 9841 &

Meijer-Drees, Mt Trikora, Papua (L, holo.; A, BO, MICH, iso.).

78. Oreogrammitis rivularis (Parris) Parris, comb. nov.

Grammitis rivularis Parris, Kew Bull. 41 (1986) 514. Type: Matthew s.n., G.

Bubu, Perak, Peninsular Malaysia (K, holo.).

79. Oreogrammitis rupestris (Parris) Parris, comb. nov.

Grammitis rupestris Parris, Blumea 29 (1983) 172, f. 28, 1. Type: Wakefield

1427, Narirogo Creek, Central, Papua New Guinea (BM, holo.).

80. Oreogrammitis salticola (Parris) Parris, comb. nov.

Grammitis salticola Parris, Blumea 29 (1983) 92, f. 13, 1. Type: Parris 5943 &

Croxall, Mt Giluwe, Southern Highlands, Papua New Guinea (K, holo.; K,

LAE, iso.).

81. Oreogrammitis scabristipes (Baker) Parris, comb. nov.

Polypodium scabristipes Baker, J. Bot. 28 (1890) 108. Type: Macgregor

sn., Owen Stanley Range, Central, Papua New Guinea (K, holo.). —

Grammitis scabristipes (Baker) Copel., Univ. Calif. Publ. Bot. 18 (1942)

223. — Polypodium ludens Baker, Ann. Bot. 8 (1894) 129. Type: Hancock

53A, Java (K, holo.). — Grammitis ludens (Baker) Copel., Gen. Fil. (1947)

211.-— Polypodium bulbotrichum Copel., Philipp. J. Sci. 40 (1929) 309. Type:

Copeland’s Pteridophyta Philippensia Exsiccata 136, Mt Pulog, Luzon,

268 Gard. Bull. Singapore 58 (2) 2007

Philippines [MICH, lecto. (Parris, 1983); BM, BO, K, SING, UC, isolecto.].

— Grammitis bulbotricha (Copel.) Copel., Philipp. J. Sci. 46 (1931) 219.

— Grammitis limapes Copel., Philipp. J. Sci. 46 (1931) 218. Type: Copeland

s.n., G. Gede-Panggrango, Java [MICH, lecto. (Parris, 1983); BM, BO, UC,

isolecto.].— Polypodium limapes (Copel.) C.Chr., Index Fil. Suppl. 3 (1934)

152.- Grammitis novoguineensis Copel., Univ. Calif. Publ. Bot. 18 (1942) 223.

Type: Bamler B.41, Bolan Mts, Morobe, Papua New Guinea (UC, holo.).

82. Oreogrammitis scleroglossoides (Copel.) Parris, comb. nov.

Grammitis scleroglossoides Copel., B. P. Bishop Mus. Occ. Papers 15 (1939)

86, f. 6. Type: Takamatsu 1095, Mt Nan-a-lant, Ponape, Caroline Is. (BISH

498214, lecto., chosen here; BISH 103965, MICH, UC, isolecto.).

83. Oreogrammitis setosa (Blume) Parris, comb. nov.

Grammitis setosa Blume, Enum. Pl. Javae (1828) 116. Type: Blume s.n., Java

(L, lecto., chosen here). — Polypodium setosum (Blume) C.Presl, Tentamen

(1836) 180 non Thunb. (1784) nec G.Forst. (1786). — Polypodium sundaicum

C.Chr., Index Fil. Suppl. 3 (1934) 160, nomen novum pro Grammitis setosa

Blume non P. setosum Thunb. (1784), nec G.Forst. (1786). — Grammitis

obscura Blume, Enum. Pl. Javae (1828) 115. Type: Blume s.n., G. Gede-

Panggrango, Java, (L, lecto., chosen here). — Polypodium obscurum (Blume)

Mett., Abhandl. Senckenb. Naturf. Gesell. 2 (1857) 38. - Polypodium

reinwardtii var. obscurum (Blume) C.Chr., Index Fil. (1906) 332.— Grammitis

pubinervis Blume, Enum. Pl. Javae (1828) 116. Type: Blume s.n., Java (L,

lecto., chosen here). — Polypodium pubinerve (Blume) Christ, Ann. Jard.

Bot. Buitenzorg 15 (1897) 146. — Grammitis obscura var. angustata Blume,

Fl. Javae 2 (1830) 113. Type: Kuhl & Van Hasselt s. n., G. Panggrango, Java

(L).— Grammitis longa Fée, Genera (1852) 233. Type: Lobb 271,““Singapore”

(BM, K). — Polypodium diplosorum Christ, Verh. Natur. Gesell. Basel 11

(1896) 440. Type: Sarasin 1358, Wawokaraeng, Sulawesi (BAS, n.v.).

84. Oreogrammitis setulifera (Alderw.) Parris, comb. nov.

Polypodium pumilum Brause. Bot. Jahrb. Syst. 49 (1912) 38, f. 2, F non

Cockayne (1909). Type: Schlechter 19112, Finisterre Mts, Morobe, Papua

New Guinea (B, n.v.). - Polypodium setuliferum Alderw., Bull. Jard. Bot.

Buitenzorg, sér. 2, 16 (1914) 32. Syntypes: Matthew 408, Malacca, Peninsular

Malaysia (BO, n.v.); Matthew 651, Barisan Mts, Sumatra (BO, n.v.; K). -

Xiphopteris setulifera (Alderw.) Parris, Kew Bull. 41 (1986) 69.

85. Oreogrammitis sledgei (Parris) Parris, comb. nov.

Grammitis sledgei Parris, Fern Gaz. 16 (2002) 240. Type: Sledge 1281, Kalu

Ganga, Ratnapura, Sri Lanka, (BM, holo.; US, iso.).

Five new genera and three new species of Grammitidaceae 269

86. Oreogrammitis subdichotoma (Racib. ex Alderw.) Parris, comb. nov.

Polypodium subdichotomum Racib. ex Alderw., Bull. Dept. Agric. Indes

Néerl. 18 (1908) 20. Type: Forbes 2408, Sumatra (BO, lecto., chosen here).

Grammitis subdichotoma (Racib. ex Alderw.) Parris, Fern Gaz. 12 (1981)

180.

87. Oreogrammitis subfasciata (Rosenst.) Parris, comb. nov.

Polypodium subfasciatum Rosenst., Repert. Spec. Nov. Regni Veg. 5 (1908)

41.Type: Werner 25, Mt Gelu, Madang, Papua New Guinea [UC, lecto. (Parris,

1983); S, isolecto.].-— Grammitis subfasciata (Rosenst.) Copel., Philipp. J. Sci.

80 (1952) 170.— Polypodium ceramicum Alderw., Bull. Jard. Bot. Buitenzorg,

sér. 3,2 (1920) 168. Type: Kornassi 1505, G. Murkele, Seram, Moluccas [BO,

lecto. (Parris, 1983); L, isolecto.]. —- Grammitis ceramica (Alderw.) Ching,

Bull. Fan Mem. Inst. Biol. 10 (1941) 240.

88. Oreogrammitis subreticulata (Copel.) Parris, comb. nov.

Polypodium subreticulatum Copel., Philipp. J. Sci. C. Bot. 9 (1914) 6. Type:

Shaw in Copland King 395,Taupota, Milne Bay, Papua New Guinea [MICH,

lecto. (Parris, 1983); BM, NSW, isolecto.].— Grammitis subreticulata (Copel.)

Copel., Philipp. J. Sci. 80 (1952) 185.

89. Oreogrammitis subspathulata (Brack.) Parris, comb. nov.

Polypodium subspathulatum Brack., U.S. Expl. Exped. 16 (1854) 3, t. 1, f.

1. Type: U.S. Expl. Exped. s.n., Tahiti, Society Is. (US, n.v.). — Grammitis

subspathulata (Brack.) Farw., Amer. Midland Nat. 12 (1931) 294.

90. Oreogrammitis sumatrana (Baker) Parris, comb. nov.

Polypodium sumatranum Baker, J. Bot. 18 (1880) 214. Type: Beccari 448,

G. Singgalang, Sumatra (K, lecto., chosen here; BM, BO, FI, n.v., isolecto.).

— Grammitis sumatrana (Baker) Copel., Philipp. J. Sci. 56 (1935) 105.

— Polypodium subpleiosorum Racib., Pterid. Buitenzorg (1898) 89. Type:

Raciborski s.n., Salak West, Java [BO, lecto. (Parris, 1983); KRA, n. v., L,

n. v., isolecto.]. — Polypodium pleiosoroides Copel., Perkins’ Fragm. FI.

Philipp. (1905) 189. Type: Copeland 1011, Mt Apo, Mindanao, Philippines

(PNH?#?, n.v.).— Polypodium serraeforme Brause, Bot. Jahrb. Syst. 49 (1912)

36 non (Wall. ex Hook.) J.Sm. (1841). Type: Schlechter 18142, Finisterre Mts,

Morobe, Papua New Guinea [B, lecto. (Parris, 1983); BM, UC, isolecto.].

— Polypodium subrepandum Brause, Bot. Jahrb. Syst. 49 (1912) 37. Type:

Schultze Jena (26) 16,Tami, West Sepik, Papua New Guinea [B, lecto. (Parris.

1983)]. — Grammitis subrepanda (Brause) Copel., Philipp. J. Sci. 80 (1952)

183. — Polypodium biseriale Ridl., Trans. Linn. Soc. London, Bot. 2, 9 (1916)

260 non Baker (1867). Type: Kloss s. n., Mt Carstensz, Papua [BM, lecto.

270 Gard. Bull. Singapore 58 (2) 2007

(Parris, 1983)]. — Polypodium carstenszense Ridl., Trans. Linn. Soc. London,

Bot. 2,9 (1916) 260. Type: Kloss s.n., Mt Carstensz, Papua [BM, lecto. (Parris,

1983); K, isolecto.]. — Polypodium brauseanum Alderw., Malayan Ferns

Suppl. (1917) 521, nomen novum pro P. serraeforme Brause non (Wall. ex

Hook.) J.Sm.

91. Oreogrammitis taeniophylla (Parris) Parris, comb. nov.

Grammitis taeniophylla Parris, Blumea 29 (1983) 67, f. 7,3. Type: Hoogland

& Craven 11034, Mt Hunstein, East Sepik, Papua New Guinea (LAE, holo.:;

CANB, iso., 7.v.).

92. Oreogrammitis temehaniensis (J.W.Moore) Parris, comb. nov.

Polypodium raiateense var. temehaniense J.W.Moore, B. P. Bishop Mus. Bull.

102 (1933) 11. Type: Moore 104, Temehani Plain, Raiatea, Society Is. (BISH

498924, lecto.,chosen here; BISH 79247, BISH 498925, isolecto.).—Grammitis

temehaniensis (J.W.Moore) Parris, Holttum Mem. Vol. (1997) 82.

93. Oreogrammitis tenuis (Parris) Parris, comb. nov.

Grammitis tenuis Parris, Blumea 29 (1983) 130, f. 19, 2. Type: Parris 6040

& Croxall, Watut-Aseki Divide, Morobe, Papua New Guinea (K, holo.; K,

LAE, iso.).

94. Oreogrammitis tomaculosa (Parris) Parris, comb. nov.

Grammitis tomaculosa Parris, Blumea 29 (1983) 152, f. 23, 3. Type: Brass

10042A & Meijer-Drees, Mt Trikora, Papua (BM, holo.; A, BO, L, MICH,

1SO.).

95. Oreogrammitis torricelliana (Brause) Parris, comb. nov.

Polypodium torricellianum Brause, Bot. Jahrb. Syst. 49 (1912) 45, f.3A. Type:

Schlechter 14347, Torricelli Mts, East or West Sepik, Papua New Guinea [B,

lecto. (Parris, 1983)]. — Grammitis torricelliana (Brause) Parris, Fern Gaz.

12 (1981) 180. - Grammitis microtricha Copel., Philipp. J. Sci. 56 (1935) 105,

t. 11. Type: Copeland s.n., Mt Apo, Mindanao, Philippines [MICH, lecto.

(Parris, 1983)].

96. Oreogrammitis trichopoda (F.Muell.) Parris, comb. nov.

Polypodium trichopodum F.Muell., Trans. Roy. Soc. Victoria 1 (1889) 41.

Type: Macgregor 28, Mt Victoria, Central, Papua New Guinea [MEL, lecto.

(Parris, 1983); BM, K, isolecto.]. — Grammitis trichopoda (F.Muell.) Copel,

Univ. Calif. Publ. Bot. 18 (1942) 224.

97. Oreogrammitis trogophylla (Copel.) Parris, comb. nov.

Five new genera and three new species of Grammitidaceae 271

Grammitis trogophylla Copel., Univ. Calif. Publ. Bot. 18 (1942) 224. Type:

Clemens s.n., Mt Sarawaket, Morobe, Papua New Guinea (MICH, holo.).

98. Oreogrammitis tuyamae (H.Ohba) Parris, comb. nov.

Grammitis tuyamae H.Ohba, Sci. Rep. Tohuku Univ. 4, 36 (1971) 121, f. 8.

Type: Tuyama s.n., Minamiiwojima, Volcano Is., Japan (TI, holo., n. v.).

99. Oreogrammitis uapensis (E.D.Br.) Parris, comb. nov.

Polypodium hookeri var. uapense E.D.Br., B. P. Bishop Mus. Bull. 89 (1931)

86. Type: Jones 155A (1210), Ua Pou, Marquesas [BISH, lecto. (Parris

1997b)].— Grammitis uapensis (E.D.Br.) Parris, Allertonia 7 (1997) 302.

100. Oreogrammitis universa (Baker) Parris, comb. nov.

Polypodium universa Baker, Annals Bot. 5 (1891) 464. Type: Curtis 633,

Penang, Peninsular Malaysia (K, lecto., chosen here). — Grammitis universa

(Baker) Copel., Philipp. J. Sci. 80 (1952) 218.

101. Oreogrammitis velutina (Parris) Parris, comb. nov.

Grammitis velutina Parris, Kew Bull. 52 (2004) 220, f. 1 B. Type: Johns 10678,

Mt Jaya, Papua (K, holo.; BO, n.v., FRE, n.v., MAN, 7.v., iso.).

102. Oreogrammitis viridula (Alderw.) Parris, comb. nov.

Polypodium viridulum Alderw., Nova Guinea 14 (1924) 41. Type: Lam 1853,

Mt Doorman, Papua [L, lecto. (Parris, 1983); BO, isolecto., n.v.].— Grammitis

viridula (Alderw.) Parris, Fern Gaz. 12 (1980) 118.

103. Oreogrammitis vittariifolia (C.Chr.) Parris, comb. nov.

Polypodium minimum Baker, J. Bot. 17 (1879) 41 non Aublet (1775). Type:

Burbidge s.n., Labuan or Kinabalu, Sabah, Malaysia [K, lecto. (Parris, 1986);

BM, E, K, isolecto.]. — Polypodium vittariifolium C.Chr., Index Fil. (1906)

574, nomen novum pro P. minimum Baker (1879) non Aublet (1775). —

Grammitis vittariifolia (C.Chr.) Parris, Fern Gaz. 12 (1980) 118.

104. Oreogrammitis wallii (Beddome) Parris, comb. nov.

Polypodium wallii Beddome, Suppl. Ferns S India & Brit. India (1876) 20,

t. 380. Type: Thwaites C. P. 3921, Adam’s Peak, Nuwara Eliya, Sri Lanka

[K, lecto. (Parris, 2002b); GH, isolecto., n.v.].— Grammitis wallii (Beddome)

Copel., Philipp. J. Sci. 80 (1952) 240.

105. Oreogrammitis whitmeei (Baker) Parris, comb. nov.

Polypodium whitmeei Baker, J. Bot. 14 (1876) 12. Type: Whitmee 124, Samoa

(CGE, GH, n.v., K). - Grammitis whitmeei (Baker) Copel., Philipp. J. Sci. 80

272 Gard. Bull. Singapore 58 (2) 2007

(1952) 150.

106. Oreogrammitis wurunuran (Parris) Parris, comb. nov.

Grammitis wurunuran Parris, Bot. J. Linn. Soc. 70 (1975) 40,£.5 G & H. Type:

Parris 3703 & Croxall, Mt Bellenden Ker, Queensland, Australia (BRI, holo.:

K, iso.).

107. Oreogrammitis zeylanica (Fée) Parris, comb. nov.

Grammitis zeylanica Fée, Genera (1852) 234. Type: Gardner 56, Sri Lanka

(BM,K,OXF,P).—Polypodium zeylanicum (Fée) Mett.,Abhandl. Senckenb.

Naturf. Gesell. 2 (1857) 38.

Acknowledgements

I am most grateful to the Flora of Peninsular Malaysia project, Forest

Research Institute of Malaysia, for a Visiting Research Fellowship to

complete the account of Grammitidaceae for the Flora, which was the

catalyst for this paper. I should like to thank the curators of A, AK, B, BISH,

BM, BO, BRI, CANB, CGE, CHR, E, FI, GH, K, KEP, KLU, LAE, MEL,

MICH, NSW, NY, P, PDA, PE, S, SAR, SING, TI, UC, US and WELT for the

use of facilities to examine specimens and/or the loan of material. Special

thanks are due to Ewen Cameron at AK for facilitating numerous loans.

References

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70: 21-43.

Parris, B. S. 1983. A taxonomic revision of the genus Grammitis Swartz

(Grammitidaceae: Filicales) in New Guinea. Blumea 29: 13-222.

Parris, B. S. 1986. Grammitidaceae of Peninsular Malaysia and Singapore.

Kew Bulletin 41: 491-517.

Parris, B. S. 1990. Noteworthy species of Grammitidaceae from Southeast

Asia. Hooker's Icones Plantarum 40(4): pp. 1-iv, 1-128.

Parris, B. S. 1997a. Themelium, a new genus of Grammitidaceae (Filicales).

Kew Bulletin 52: 737-741.

Parris, B. S. 1997b. Two new species of Grammitis from the Marquesas

Islands. Allertonia 7: 296-303.

Parris, B. S. 1998a. Chrysogrammitis, a new genus of Grammitidaceae

(Filicales). Kew Bulletin 53: 909-918.

Parris, B. S. 1998b. Grammitidaceae. In: Flora of Australia 48: 450-468,

713-714.

Parris, B. S. 2002a. New species and new combinations in African

Grammitidaceae (Filicales). Kew Bulletin 57: 423-434.

Parris, B. S. 2002b. Notes on Grammitidaceae (Filicales) of Sri Lanka and

the description of three new species. Fern Gazette 16: 239-244.

Parris, B. S. 2003. The distribution of Grammitidaceae (Filicales) inside and

outside Malesia. Jelopea 10: 451-466.

Parris, B. S.2004. New combinations in Acrosorus, Lellingeria, Prosaptia and

Themelium. Kew Bulletin 59: 223-225.

Parris, B. S., R.S. Beaman, and J.H. Beaman. 1992. The Plants of Mount

Kinabalu. 1. Ferns and Fern Allies. Royal Botanic Gardens, Kew. 165

Pp.

274 Gard. Bull. Singapore 58 (2) 2007

Parris, B. S. and P.J. Edwards. 1988. A provisional checklist of ferns and fern

allies in Taman Negara, Peninsular Malaysia. Malayan Naturalist 41:

5-10.

Parris, B. S. and A. Latiff. 1997. Towards a pteridophyte flora of Malaysia: a

provisional checklist of taxa. Malayan Nature Journal 50: 235-280.

Ranker, T.A, A.R. Smith, B.S. Parris, JM.O. Geiger, C.H. Haufler, S.C.K.

Straub and H. Schneider. 2004. Phylogeny and evolution of grammitid

ferns (Grammitidaceae): a case of rampant morphological homoplasy.

Taxon 53: 415-428.

Gardens’ Bulletin Singapore 58 (2) 2007: 275-278 eS

Two New Names and Two New Combinations in Malesian

Annonaceae

IAN M. TURNER

Research Associate, Singapore Botanic Gardens, Singapore

and

P.O. Box 20, Winchelsea, East Sussex TN36 4WA, UK

Abstract

Two later homonyms in Polyalthia are provided with new names: Polyalthia sinclairiana

I.M.Turner replaces P. macropoda King and Polyalthia dolichopoda 1.M.Turner replaces P.

tenuipes Merr. (1922, non Merr. 1912). The combination Dasymaschalon dasymaschalum

(Blume) I.M.Turner, which is not a tautonym under the current code, is made. It replaces the

superfluous Dasymaschalon blumei Finet & Gagnep. A new combination for Dasymaschalon

dasymaschalum var. wallichii is also made. Lectotypes are proposed for Polyalthia macropoda

King, Polyalthia tenuipes Merr. (1922), and Unona dasymaschala Blume var. wallichii Hook.

f. & Thomson.

A new name for Polyalthia macropoda King

Polyalthia macropoda King 1s a cauliflorous tree species widely distributed

in the Malay Peninsula. Unfortunately the name is a later homonym, having

already been used for a species from New Guinea. A new name is therefore

proposed and it is fitting to honour James Sinclair, whose revision of the

Annonaceae of the Malay Peninsula (Sinclair 1955) has proved seminal.

Polyalthia sinclairiana [.M. Turner, nom. nov.

Pro Polyalthia macropoda King (non (Miq.) F. Muell.), J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 61 (1892) 60. Type: King’s Collector, Perak (King’s Collector

4279, lectotype nov., designated here, K!; isolectotype, BM!).

A new name for Polyalthia tenuipes Merr.

Polyalthia tenuipes Mertr. is a distinctive and quite widely collected Bornean

species. Unfortunately Merrill coined the name for this species having

already used the same binomial for a different species from the Philippines.

It is therefore necessary to propose a new name for the Bornean species.

Polyalthia dolichopoda I.M. Turner, nom. nov.

Pro Polyalthia tenuipes Merr. (non Metr., Philipp. J. Sci., C. 7 (1912) 269), J.

Straits Branch Roy. Asiat. Soc. 85 (1922) 184. Type: Sabah, near Sandakan, M.

276 Gard. Bull. Singapore 58 (2) 2007

Ramos 1501 (holotype, PNH destroyed; isotype, SAN destroyed; lectotype

nov., designated here, K!; isolectotype, US).

New combinations in Dasymaschalon

Presumably Finet and Ganepain, when they published the name

Dasymaschalon blumei for Unona dasymaschala Blume, did so in the belief

that the combination Dasymaschalon dasymaschalum would be a tautonym

and thus illegitimate. However, as recent edition of the International Code of

Botanical Nomenclature (Art. 23.4) states: ‘The specific epithet ... may not

exactly repeat the generic name (such repetition would result in a tautonym).’

Therefore Dasymaschalon dasymaschalum is not a tautonym because the

repeat between generic name and specific epithet is not exact. An analogous

case (Art. 14, Ex. 1) is Lycopersicon lycopersicum (L.) H. Karsten, which is

not a tautonym. This combination, however, had to be formally rejected to

avoid replacement of the widely used name for the tomato, Lycopersicon

esculentum Mill. The legitimacy of Dasymaschalon dasymaschalum makes

Dasymaschalon blumei Finet & Gagnep. a superfluous renaming of Unona

dasymaschala Blume. Below I formally make the new combination for

Unona dasymaschala in Dasymaschalon. Dasymaschalon blumei could be

revived either through a change of the ICBN code by widening the definition

of tautonym, or a formal rejection of the new combination proposed below

based on other justifications.

Dasymaschalon dasymaschalum (Blume) I.M. Turner, comb. nov.

Basionym: Unona dasymaschala Blume, Fl. Javae Anonaceae 23-24 (1830)

55,t.27. Type: Java, Blume s.n. [lectotype designated by Nurmawati (2003),

L (barcode: L0186230); isolectotype, BO].

Homotypic synonyms: Dasymaschalon blumei Finet & Gagnep., Bull. Soc.

Bot. France 53, Mémoire 4 (1906) 143, nom. superfi.

-Desmos dasymaschalus (Blume) Safford, Bull. Torrey Bot. Club 39 (1912)

Su,

-Unona dasymaschala var. blumei Hook.f. & Thomson, FI. Ind. 1 (1855) 135,

nomi. inval.

Heterotypic synonyms: Unona coelophloea Scheff., Flora 52 (1869) 300; -

Dasymaschalon coelophloeum (Scheff.) Merr., Philipp. J. Sci., C. 10(4) (1915)

pines

-UnonacleistogamaBurck ex Boerl.,Icon.Bogor.1 (1899) 127;Dasymaschalon

cleistogamum (Burck ex Boerl.) Merr., Philipp. J. Sci., C. 10(4) (1915) 237.

Two New Names and Two New Combinations in Malesian Annonaceae DVT

Dasymaschalon dasymaschalum (Blume) I.M.Turner var. wallichii (Hook.f.

& Thomson) I.M.Turner, comb. nov.

Basionym: Unona dasymaschala Blume var. wallichii Hook.f. & Thomson,

Fl. Ind. 1 (1855) 135. Type: Singapore, Wallich catalogue 6421A (lectotype

nov., designated here, K-WALL!).

Homotypic synonyms: Desmos dasymaschalus (Blume) Safford var. wallichii

(Hook.f. & Thomson) Ridl., Fl. Malay. Penins. 1 (1922) 47; Dasymaschalon

blumei Finet & Gagnep. var. wallichii (Hook.f. & Thoms.) Ban, Bot. Zhurn.

60 (1975) 228.

Acknowledgements

Prof. Simon Owens is thanked for permitting acess to the facilities of the

herbarium and library at the Royal Botanic Gardens Kew. Dr J.F. Veldkamp

(L) provided helpful comments on a draft of this paper.

References

Nurmawati, S. 2003. Malesian species of Dasymaschalon (Annonaceae).

Floribunda 2(3): 67—81

Sinclair, J. 1955. A revision of the Malayan Annonaceae. Gardens’ Bulletin,

Singapore 14: 149—516.

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Gardens’ Bulletin Singapore 58 (2) 2007: 279-286 279

Studies on Schismatoglottideae (Araceae) of Borneo II:

Aridarum crassum, a New Species from Sarawak,

Malaysian Borneo

WONG SIN YENG' AND PETER C. BOYCE

‘Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Samarahan, Sarawak, Malaysia

% Malesiana Tropicals, Suite 9-04, Tun Jugah Tower, No. 18, Jalan Tunku Abdul Rah-

man,

93100 Kuching, Sarawak, Malaysia

Abstract

Aridarum crassum S.Y.Wong & P.C.Boyce is described as a new species from Sarawak,

Malaysian Borneo. The existing key to the genus Aridarum Ridl. is amended to include the

new species.

Introduction

Aridarum Ridl.is a genus consisting of nine species of rheophytic, sometimes

facultatively lithophytic, forest herbs, are all endemic to Borneo. Previous

work by Bogner & Hay (2000) recognised nine species of which one

species was at that time inadequately known and thus not fully described.

The species, treated as sp. A in Bogner & Hay (2000), was known from a

single collection from the summit (850 m asl) of Gunung Gaharu (Sri Aman

Division, Sarawak). Fieldwork by the second author at the same location,

although at lower altitude (the summit is no longer accessible) resulted in

three collections of what we believe to be the same species. This species is

clearly distinguished from the rest of the genus Aridarum by the leaf laminae

very stiffly coriaceous, glossy deep green adaxially and conspicuously raised

punctate abaxially when fresh. We are hereby describing this novelty as

Aridarum crassum S.Y.Wong & P.C.Boyce.

Key to species of Aridarum

ira; Bhecac on cach end of the amther (Sect. ATidarus) ....-.....0ccesen0s see 2

1b. Thecae together on one side of the anther (Sect. Caulescentia) .......... 6

280 Gard. Bull. Singapore 58 (2) 2007

2a. Leaf laminae almost linear; horns of anther thecae long and thin; Sarawak

(Santubong.... ee Leteeeeeeeeeeeee-A. MOontanum

20: Leat pee a7 oie sists a ilps foe om anther thecae

short and StubDy \..ccccccssctneterececcnete cee tote torent ee ee a 3

3a. Leaf arrangement distichous; Sarawak (vicinity of Matang) ...............

ssionssnsweidi boubgianeeuacauttea ge tede Saadere eeeen tector hcestee eee ecard ee ee A. borneense

4a. Spadix fertile to apex (possibly except a few apical sterile stamens); an-

thers not excavated; horns of thecae short but robust, their bases occupying

the whole upper surface of the anther; West Kalimantan ...... A. incavatum

4b. Spadix with an appendix of staminodes; anthers excavated; horns of the

thecae small, on each end of the anther; Sarawak and West Kalimantan ...5

5a. Leaf laminae very stiffly coriaceous, glossy deep green adaxially when

fresh, conspicuously raised-punctate abaxially when fresh, weakly and

minutely punctate in dry material, margins revolute in dry material, primary

lateral venation weakly raised adaxially when fresh, 5-8 primary lateral veins

at each side but obscure in dry material, obscure abaxially when fresh and

in dry material; sweet smelling (esterases) at female anthesis, ovary ovoid to

elongate-ovoid, stigma c of ovary diam. strongly papillate, umbonate, stain-

ing brownish red in alcohol; thecae horns very short, rounded at the end;

Sarawak (Gunung Gaharu) .. 2 ee ‘ooepuxvesansytsdlas An Rte

5b. Leaf laminae rubbery-coriaceous, ee aeditries asea rear when

fresh, inconspicuously raised punctate abaxially when fresh, conspicuously

raised punctate in dry material, margins planate in dry material, primary

lateral venation obscure adaxially when fresh and in dry material, weakly

raised abaxially when fresh and in dry material, to ca 3 primary lateral veins

on each side; inflorescence with no smell at female anthesis, ovary ovoid,

stigma as wide as ovary, weakly papillate, scutiform, staining light brown

in alcohol; thecae horns long, pointed at the end; Sarawak (Bako National

Park) and. West, Kalama mia voi siccec:suactete ae ce stoes cee ee A. nicolsonii

6a. Stamens arranged in pairs; thecae on the inner face of each member of

the Stamen Paid oii. .:scs.d.chcene2-.descts veces ee ee ee |

6b. Stamens single (but crowded); thecae on the proximal (with respect to the

spadix axis) side of the anther, 6.2.5 cscs eet ee ee 8

7a. Horns of thecae shorter than width of stamen; Sarawak and Brunei......

sande ovdnenls vag ebeecavndedsv dda chaaiuene ice Be ee A. caulescens

Tb. Hors of thecae longer than width of stamen; Sarawak .... A. purseglovei

Studies on Schismatoglottideae of Borneo 281

8a. Stamens and staminodes coarsely verruculate; appendix well-

differentiated; spathe beaked for more than half its length; West Kalimantan

EL Wnt err 05210) ReeReese gee ek caer ees en A. rostratum

8b. Stamens and staminodes not verruculate; appendix reduced to a few

terminal sterile stamens; spathe not long-beaked; Sarawak and West &

CMe IMENT) 2 oT a9 5: ea ee eee ee A. burttii

Aridarum crassum S.Y.Wong & P.C.Boyce sp. nov.

Aridarum crassum differt ab speciebus ceteris laminis foliorum valde rigide

coriaceis, adaxiali atro-viride, nitentibus, abaxiali punctis elevatis instructa

(in stato vivo). Aridarum crassum A. nicolsonii similis videtur sed laminis

foliorum abaxiali punctis elevatis instructa (in stato vivo) sed folia siccitatae

abaxiali perpunctulatus, nervis lateralibus primariis parum prominentibus,

5-8 in quoque latere in stato vivo, valde obcuriis in stato sicco, inflorescentia

odorus per anthesin feminae, quam stigmatibus quam ovaria ca / diametro,

cum alcoholis rubiginosus, cornu thecis antherae brevissimus, obtusus differt.

Typus: Malaysia, Sarawak, Sri Aman Division, Pantu, Gn. Gaharu, 01° 02’

39.5”; 110° 53’ 18.3”, 100 m asl, 23 Sept. 2004, PC. Boyce & Jeland ak Kisai

AR-692 (holo,SAR). Plates 1 & 2

Small, moderate to robust rheophytic herbs to ca. 20 cm tall. Stem epigeal,

pleionanthic, erect to semi-decumbent, condensed, to 1.6 cm long x 1.3 cm

diam., roots green, photosynthetic, robust to ca 2 mm diam. Leaves few

together, spirally arranged, to ca 4 per module; petioles stout, glabrous, 4—8

cm long x 3.5—5 mm diam., dark green, similar colour to adaxial side of leaf

lamina, D-shaped adaxially in cross section; sheathing only at the extreme

base, the sheaths fused and extending into a very narrowly triangular

marcescent ligular portion to ca 2.8 cm long, ‘| to % of petiole length; lamina

broadly oblanceolate to broadly oblong-lanceolate to elliptic or narrowly

obovate, very stiffly coriaceous, glabrous, 11—14 cm long x 3—5 cm wide, base

cuneate to decurrent, oblique, apex acuminate, sometimes acute, acuminate

for ca 1.3 cm, tubular apiculate to ca 1.5 mm; margins entire, markedly

thickened when fresh, revolute (in dry material), adaxially shining dark

green, abaxially matte paler green, minutely rugulose, marked conspicuously

with dense, raised punctate when fresh but weakly and minutely punctate

in dry material; midrib adaxially flush with lamina, abaxially prominent,

raised canaliculate in cross section; primary lateral venation weakly raised

adaxially when fresh, 5-8 primary lateral veins at each side but obscure in dry

material, very obscure abaxially when fresh and in dry material, secondary

venation obscure, primary and secondary venation diverging at 30°-45° from

mid-vein, running into obscure marginal vein; tertiary venation invisible.

282 Gard. Bull. Singapore 58 (2) 2007

Inflorescence solitary, sub-erect, sweet smelling (esterases) at female

anthesis, peduncle to ca 8 cm long, always exceeding petioles, subtended by

a pair of fused ligules, to ca 4.5 cm long these spliting into two at the end,

to ca 2.3 cm long, sometimes two rudimentary leaves present, these to ca

2 mm followed by a prophyll and a cataphyll; spathe more or less broadly

ovate, not constricted, softly coriaceous, interior glossy, exterior semi-glossy,

to 6-8 cm long; lower spathe cup-shaped, obconic, 2.2 cm long x 1.9 cm diam.

(across), spathe abcission layer coinciding with base of male zone, green

when fresh, persistent into fruiting, margins convolute; limb broadly ovate,

convolute, white, to ca 3.8 cm long, tip mucronate to ca 3 mm, caducous

after male anthesis; spadix sessile, adnate isodiametrically to the spathe in

the lower 4% of female zone on the dorsal side, cylindric, to ca 4.5—5.5 cm

long, subequalling spathe; female zone cylindric, often constricted at the

adnation, green when fresh, ca 1 cm long x 8.6 mm-1 cm diam., '/; of spadix

length; pistils many, somewhat crowded, ovoid to elongate-ovoid, ca 1.2 mm

long x 0.85 mm diam., turning white in alcohol, stigma sessile, raised, notably

papillate, ca 0.60-1.1 mm diam., */; of ovary diam., staining brownish red in

alcohol; interpistillar staminodes, staminodes confined to a single row at the

base of spadix, laxly arranged, more-or-less square to round, impressed in the

middle, shorter than pistils, about same size as ovary, shortly stipitate; sterile

interstice cylindric, always wider in diameter than female zone, 4—6 mm long

x 8 mm-1.1 cm diam., '/10 of spadix length, without constriction, clothed with

staminodes, more-or-less circular, sometimes rectangular, truncate, ca to 1

mm across, excavation circular deep, sometimes wide impressed; male zone

tapering-cylindric, basally slightly thickened above interstice, 2.2 cm long x

8 mm long, % of spadix length; stamens arranged in longitudinally aligned

pairs, crowded, rectangular, ca 1 mm across; connective deeply excavated,

with the cavity not septate; thecae opposite on the distal and proximal (with

respect to the spadix axis) sides of the anther; horns very short, round to

slightly point at end, erect to suberect; appendix tapering-cylindric to a blunt

tip, ca 8.5 mm long, */s of spadix length, staminodes more or less circular,

sometimes irregularly polygonal, 0.6 mm to 1 mm wide, excavations circular

deep, sometimes wide impressed. Infructescence solitary; peduncle to ca 8

cm long, not exserted, erect; fruiting spathe obconic, ca 2 cm.

Distribution: Endemic to Sarawak: Sri Aman Division (Gn. Gaharu).

Habitat: Rheophytic on large sandstone boulders at low altitudes, mainly on

wet sandstone cliff surfaces at higher altitudes, 100 to 850 m asl.

Notes: Aridarum crassum is clearly distinctive from the rest of the genus

by the leaf laminae very stiffly coriaceous, glossy deep green adaxially and

conspicuously raised punctate abaxially when fresh. Aridarum crassum is

Studies on Schismatoglottideae of Borneo

Plate 1: Aridarum crassum S.Y.Wong & P.C.Boyce (Figs. A, B & D) and Aridarum nicolso-

nii Bogner (Fig. C): A. Note very stiffly coriaceous and glossy deep green leaf laminae and

stout petioles of Aridarum crassum. B. Adaxially weakly raised primary lateral venation seen

in Aridarum crassum. C. Note in Aridarum nicolsonii the matte medium green leaf laminae

and primary lateral venation obscure adaxially, note slender petioles. D. Inflorescence of A.

crassum in bud.

Gard. Bull. Singapore 58 (2) 2007

Plate 2: Aridarum crassum S.Y.Wong & P.C.Boyce (Figs. A, C & E) and Aridarum nicolsonii

Bogner (Figs. B, D & F): A. Abaxial surface of leaf lamina of Aridarum crassum showing

obscure primary lateral venation and conspicuous raised punctae. B. Abaxial surface of leaf

lamina of Aridarum nicolsonii showing showing weakly raised primary lateral venation. C.

Spadix of A. crassum (spathe artifically removed) to display stigma smaller than ovary and

tapering male zone. D. spathe of Aridarum nicolsonii at gaping stage. E. Inflorescence of A.

crassum at male anthesis. F. Spadix (spathe artifically removed) of Aridarum nicolsonii to

display stigma smaller than ovary and cylindrical male zone.

Studies on Schismatoglottideae of Borneo 285

most similar to A. nicolsonii Bogner but is readily separable by the above

noted characters plus in having stigmas smaller than ovary diameter, by

inflorescences fragrant of esterases at female anthesis, and a tapering-

cylindric (vs cylindric) male zone. Additionally, A. crassum is more robust

with stouter petioles than A. nicolsonii, which tends to be gracile with

slender petioles.

Leaf laminae of A. crassum are abaxially conspicuously marked with raised

punctae when fresh, but weakly and minutely punctate in dry material

at which time the margins are conspicuously revolute as compared to A.

nicolsonii with inconspicuous raised punctae abaxially when fresh but

conspicuous raised punctate and a planate leaf margins in dry material.

Leaf laminae of A. crassum are broadly oblanceolate to broadly oblong-

lanceolate to elliptic or narrowly obovate with the apex shortly acuminate

(to ca 1.3 cm), as compared to elliptic to broadly oblanceolate with the leaf

acuminate to ca 2 cm in A. nicolsonii. Primary lateral venation of A. crassum

consists of 5-8 primary lateral veins per side are weakly raised adaxially

when fresh but obscure in dry material, abaxially these veins are obscure

both in fresh and dry material. In A. nicolsonii primary lateral venation (to

ca 3 primary lateral veins per each side) is obscure adaxially when fresh and

in dry material, but weakly raised abaxially both in fresh and dry material.

Aridarum crassum emits a sweet smell (esterases) at female anthesis whereas

A. nicolsonii does not emit any smell at any time during anthesis. Ovaries in

A. crassum are ovoid to elongate-ovoid (compared to ovoid in A. nicolsonit)

with a strongly papillate, umbonate, smaller stigma (to */; of ovary diam.)

staining brownish red in alcohol (vs a less papillate, scutiform stigma as wide

as ovary and staining light brown in alcohol in A. nicolsonii). The thecae

horns are very short and rounded at the tips in A. crassum, compared to

longer thecae horns that are pointed at the end in A. nicolsonii.

Aridarum crassum is endemic to Gunung Gaharu, Sri Aman Division,

Sarawak whereas, A. nicolsonii is widespread in Bako National Park,

Kuching Division, Sarawak and in West Kalimantan.

Etymology: The species epithet is derived from the Latin, crassus, ‘thick’

reflecting the markedly stiffly coriaceous leaf laminae that immediately

distinguishes this species.

Other specimens examined: SARAWAK. Sri Aman Division: Pantu, Gn.

Gaharu, 01° 02’ 39.5”; 110° 53’ 18.3”, 22 March 2004, PC. Boyce & Jeland

ak Kisai AR-251 (SAR); Ibid, 01° 01’ 19.5”; 110° 52’ 52.8”, 22 March 2004,

286 Gard. Bull. Singapore 58 (2) 2007

P.C. Boyce & Jeland ak Kisai AR-263 (SAR + spirit).

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee and

the Forest Research Centre (Kuching), notably L.C.J. Julaihi & Lucy Chong.

Thanks are due to Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme

Brown & Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their

continued support and funding of fieldwork in Sarawak. The primary author

is grateful for the support provided by Faculty of Resource Science and

Technology, UNIMAS.

Reference

Bogner, J. and Hay, A. 2000. Schismatoglottideae in Malesia II — Aridarum,

Bucephalandra, Phymatarum and Piptospatha. Telopea 9(A): 183-194.

Additional notes on Aridarum crassum

Aridarum crassum also occurs on Gn. Lingga (Sri Aman Division) which is

150 km further eastwards from Gn. Gaharu. Aridarum nicolsonii also occurs

on Gn. Lesung, Lingga (based on a collection of Carlo Hansen No. 1044);

the occurrence of two Aridarum species together is unusual. Especially

interesting is the occurrence of a species at high elevation (A. crassum)

and another allopatrically at low elevation (A. nicolsonii); this is also the

instance at Gn. Bako (Kuching Division) where the enigmatic Aridarum

montanum Ridl. occurs towards the summit, while the lower slopes harbours

A. nicolsonii.

Additional specimens examined: SARAWAK. Sri Aman Division: Pantu,

Gn. Gaharu, 8 Oct. 1974, Illias & Azahari S. 35676 (SAR); Gn. Gaharu, 7

Dec. 2004, Bogner 2835 (M); Gn. Lingga, June 1906, Hewitt series 826 (SAR);

ibid, June 1906, Hewitt series 446 (SAR); ibid, June 1906, Hewitt series 1095

(SAR).

Gardens’ Bulletin Singapore 58 (2) 2007: 287-292 ek

Book Review: A.D. Poulsen. 2006. Etlingera of Borneo. Natural History

Publications (Borneo) Sdn. Bhd. in association with Royal Botanic Garden

Edinburgh. x + 263 pp. ISBN: 983-812-117-7. Price: US $55.00

The ginger family (Zingiberaceae) has more than 1500 species worldwide.

The genus Eftlingera comprises more than 100 species, all confined to

Indo-Pacific region. The best-known member of this genus is the widely

appreciated ornamental torch ginger, Etlingera elatior.

The striking plant diversity of Borneo is well known, and, likewise, is the

importance of gingers as spice, vegetable, dye, medicine and other uses. There

are few vegetation types in humid or monsoonal tropics where gingers are

not commonly found. Borneo is home to at least 300 ginger species of which

the genus Etlingera is perhaps the most prominent, most diverse, and also

most attractive.

Gingers are, however, one of the more difficult plant families to collect

and study. Meaningful work can only be done on live and fresh flowering

plant materials and requires fieldwork with round-a-year observation

to catch the appearance of flowers and fruits, both bearing important

diagnostic characters. Fresh flowers of gingers are difficult to preserve, and

unfortunately, many studies on gingers, especially early ones, are based on

herbarium specimens only. Thus, the common outcome of people trying to

key out the ginger species using a published Key is - giving up in frustration.

Fortunately, this is not likely to happen when using the present book.

The new book on the Etlingera of Borneo is the result of the author’s

15 years of extensive fieldworks and his thorough study of herbarium

Specimens in more than 20 herbaria. What more, it is a delightful blend of

strictly scientific information with beautiful color photographs, illustrations,

and color distribution maps, facilitating the dissemination of information to

botanists, foresters, eco-tourists, and even, laymen.

The book starts off with a short introduction and is followed by a chapter

introducing the typical Etlingera plant. Details of the morphological

structures shown in this chapter are especially clarified by images made

under the scanning electron microscope. The third chapter, Etlingera in

Borneo, includes a general topical history, distribution, ecology, species

richness, reproductive biology, and ends with ethnobotany and notes on

conservation. The last chapter deals with the materials and methods and,

in particular, highlights the necessity of good photographic documentation

and field sketches as two indispensable supplements to herbarium and spirit

288 Gard. Bull. Singapore 58 (2) 2007

materials. The last few pages, just before the revision itself, are occupied by

a key to the Bornean species of Etlingera.

The main part of the book features 42 taxa all together. Sixteen species, one

subspecies, one variety, and one combination are described. Each species

is accompanied by relevant information about the source of publication,

synonymy, nomenclatural details, and the type(s). This is followed by a

thorough species description, local names and uses, etymology, ecology and

habitat, distribution (supported by colour map clearly showing the locality

of type specimen), conservation status, list of materials examined, and

ends by elaborate notes helping to pinpoint the differences of the species

under discussion from its closest taxa, or discussing the tricky issues of

nomenclatural history and taxonomic confusions.

Just before the book concludes, there is a big bonus for all botanists printed

between the sections of references and indices to scientific and vernacular

names. It is a list of identification of all Etlingera specimens examined by

the author during the course of his preparation work, alphabetically sorted

according to a collector’s surname, followed by the collection number of

species. In case you get stuck in research, for example, with a sheet of

specimen collected by Gobilik J. et al. 738 kept at SAN Herbarium, you will

know within seconds of checking this book that the species of the specimen

on this sheet is E. inundata.

I tried to find negative points in this new book, however, it turned out to be

a tough job. As a true ginger fan, I would welcome the more than 25 pages of

introduction part of the book. Nonetheless, information about ethnobotany

of various Etlingera species is rather little, despite the fact that more than

20 species are used for various purposes. The Table 1 (p. 18) reported uses

and number of species used for a particular purpose, e.g., shoot is eaten in 20

species, but the reader is not informed which species are these without going

through all the taxa listed. An appendix with an overview of all species and

their uses would have been useful.

To facilitate the use of the identification key, more illustrations explaining

the various morphological terms would have been welcome because the

perception of various structures seen in gingers traditionally deviates from

the definition of botanical terms used in other monocot groups. To note,

the flower structure is explained well, however, the terms regarding leaf

terminology would be particularly useful, if illustrated.

In my opinion, it remains as a matter of fact, that it is faster and much more

Gard. Bull. Singapore 58 (2) 2007 289

enjoyable to identify a species while leisurely flipping through the pages

full of stunning and beautiful photographs, including close-up details of

each species, rather than using the dichotomous key to the species. I have a

feeling that most ginger specialists will use the species key published in the

new book only for the first few times to satisfy their curiosity rather than

using the key as a regular tool for species identification.

Last but not least, it is always necessary to remind ourselves that in our

quickly disappearing plant world today, it is difficult to protect something

that we do not know properly, nor recognize, and neither can we identify.

This published revision study of the genus Eftlingera is therefore a much

welcome step forward, enabling us to protect the endemic and endangered

species of a group of prominent gingers in Borneo.

Etlingera of Borneo is definitely a benchmark publication. It shall serve in

many years to come as a great example and guide, showing how a modern

revision of a complicated genus should be done.

Jana Leong-Skornickova

The SING Herbarium

Singapore Botanic Gardens

1 Cluny Road

Singapore

290 Gard. Bull. Singapore 58 (2) 2007

Book review: Justin W. Tkatchenko, BEM and Steven Kami. 2006. Spatulata

Orchids - Papua New Guinea. Orchid Society of Papua New Guinea. 96 pp.

ISBN 9980-86-070-7.

This publication deals with Dendrobium sect. Spatulata,a fascinating group of

attractive orchids. The horticulturally very important group is characterised

by mostly cane-like stems, erect and frequently twisted petals (giving this

group the common name ‘antelope dendrobiums’) and a usually three-

lobed, concave lip with a 3-5-ridged callus. The area of distribution of this

group is in tropical Asia from Java and the Philippines eastwards as far as the

Pacific Islands Fiji, Tonga and Samoa, and southwards as far as Queensland

(Australia). However, by far the greatest diversity is found in New Guinea

where over half of the altogether ca 50 species of this Dendrobium section

are found.

The brief preface of this semi-scientific book gives relevant background

information on Dendrobium sect. Spatulata, together with notes on its

distribution. In the following main part twenty-eight species plus several

varieties, colour forms and suspected new species are described in detail,each

of them with a short botanical description and comments on its cultivation

requirements. An informative outline map indicates its distribution within

Papua New Guinea and the adjacent islands, and in the accompanying text

the habitat ecology is briefly described. One or several excellent colour

pictures are provided for all taxa, in a high quality of photography and

reproduction which is not always found in publications. A chapter on the

natural habitat in the end of the book explains the growing conditions of

the plants, and also chapters of cultivation, watering and fertilising are given

here. Also these chapters are illustrated by magnificent photographs. A short

list of references as well as an index conclude the book.

There are very few negative points. I personally feel that a detailed map of

the area would have helped, as not all of the readers are familiar with the

geography of Papua New Guinea and its offshore islands (particularly those

not living in this region).

The very minor errors, like author names italicised (p. 13), species and

variety names not italicised (p. 13, 89) or missing authors (p. 13), hardly

affect the value of this publication. Neither do the few distribution maps

which do not exactly reflect what the text actually says (p. 12, 15, 20, 42). I

personally would have liked more detailed notes on the distribution outside

Papua New Guinea (however, occasional comments on the occurrence in

Irian Jaya and Australia are indeed given in the book); but I know that this

Gard. Bull. Singapore 58 (2) 2007 291

is not always done in publication of this nature. Perhaps it would also look

better if the introductory chapters on habitat and cultivation would be in

the front section of the book just behind the preface, but, of course, this too

is mainly a matter of personal preference.

Those who would like to cite this publication will not find it easy to find out

who actually wrote it as only the preface and the biographies on p. 95 says

who the authors are; their names should have been indicated on the title

page.

This book is a very useful and informative contribution on an attractive

plant group in a floristically incompletely know though extremely diverse

country, and can, as such, be recommended to everybody who 1s interested

in tropical Asian orchids. The authors, Mr. Justin W. Tkatchenko, BEM and

Mr. Steven Kami, president and vice-president (respectively) of the Orchid

Society of Papua New Guinea, can be congratulated for their valuable

publication.

H. Kurzweil

The SING Herbarium

Singapore Botanic Gardens

1 Cluny Road

Singapore

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INSTRUCTIONS TO AUTHORS

The Gardens’ Bulletin publishes original research findings and reviews of progress in the fields of plant

taxonomy, horticulture and allied subjects. Contributions must be original and the material must not

have been submitted for publication elsewhere.

Authors should look at the layout of articles recently published in the journal to ensure that submitted

manuscripts conform as closely as possible to the accepted format. Particular care should be taken

with the format of the references. Manuscripts may be submitted in electronic form (PC-compatible

WORD) together with a hardcopy and original drawings and illustrations as appropriate.

Titles and authors: The title should give a concise description of the contents of the article. It should

include the family name, if a taxon name is included in the title. The names(s) and affiliation(s) of the

author(s) must be given below the title. A short running title should also be provided. Lengthy papers

must have contents listed at the beginning of the paper. Avoid footnotes.

Abstract: An abstract of up to about 100 to 200 words should be provided. It should comprehensively

summarise the contents of the article as it is likely to be reproduced without the text.

Scientific names: The complete scientific name - genus, species, authority with family in parenthesis

- must be cited for every organism at the time of first mention.

Abbreviations: Standard abbreviations may be used in the text, but the full term should be given on the

first mention. Dates should be cited as: 1 Jan 2000. Units of measurement should be spelled out except

when preceded by a numeral where they should be abbreviated in standard form: g, ml, km, etc., and

not followed by stops.

Tables: All tables should be numbered and carry a heading with their content. These should be

comprehensive without reference to the text.

Illustrations: For black and white drawings, the original drawings are still preferred. Scale bars should

be used to indicate magnification. Provide a photocopy of the illustrations to indicate the lettering for

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When grouping photographs, the page size of the journal must be taken into account to optimize

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For figures including photographs, type the captions in numerical order on a separate sheet.

Literature citation: Citation in the text should take the form: King and Gamble (1886), or (King and

Gamble, 1886). If several papers by the same author in the same year are cited, they should be lettered in

sequence, 2000a, 2000b, etc. When papers are by three or more authors they should be cited as King et

al. (1886) in the text, but with all the authors’ names given in the reference section. All references must

be placed in alphabetic order according to the family name of the first author. The journal title must be

given in full, as in the following examples:

Stone, B.C. 1994. Additional notes on the genus Glycosmis (Rutaceae). Gardens’ Bulletin

Singapore 46: 113-119.

Kress, W. J., L. M. Prince, and K. J. Williams. 2002. The phylogeny and a new classification

of the gingers (Zingiberaceae): evidence from molecular data. American Journal of Botany 89:1682-

1696.

References to books and monographs should be cited according to the following form:

Ridley. H.N. 1930. The Dispersal of Plants Throughout the World. L. Reeve, Ashford, U.K.

For literature citations in taxonomic papers, the following style is required:

Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. I:1 (1849) 19.

Sterculia acuminatissima Merr., Philipp. J. Sci. 21 (1922) 524.

Offprints: Authors will be given 50 offprints gratis. Additional copies must be ordered and paid for in

advance of publication.

Manuscripts should be sent to: The Editor, Gardens’ Bulletin Singapore. The Singapore Botanic

Gardens, Cluny Road, Singapore 259569.

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