Gardens’ Bulletin

Singapore

~ VOL. 62 (1) 2010 ; ISSN 0374-7859

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 Dr. Jana Leong-Skorni¢kova

Singapore Botanic Gardens Singapore Botanic Gardens

(Editor) (Assistant Editor)

De PY. Tan Ms. C. Soh

National Parks Board Singapore Botanic Gardens

(Assistant Editor) (Journal Business Manager)

EDITORIAL BOARD

Dr. S.C. Chin Dr. M.C. Roos

Singapore Botanic Gardens National Herbarium Netherlands

Singapore Leiden University, The Netherlands

Dr. M.J.E. Coode Dr. E. Soepadmo

Royal Botanic Gardens Forest Research Institute Malaysia

Kew, U.K. Kepong, Malaysia

Prof. Sir P. Crane Prof. T. Stuessy

Yale University University of Vienna

New Haven, USA Austria

Dr. R.T. Corlett Dr. W.K. Tan

DBS, National University of Singapore National Parks Board

Singapore Singapore

Dr. W.J. Kress Dr. I.M. Turner

Department of Botany, NMNH Research Associate

Smithsonian Institution Singapore Botanic Gardens

Washington DC, USA Singapore

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[Cover photo: Inflorescence of Curcuma vitellina (see p. 114); photo by Jana

Leong-Skorni¢kova. ]

VOL. 62 (1) 2010

ot DEC 21 2010

The Gardens' Bulletin

Singapore

CONTENTS

Ahmad Sofiman bin O. and P.C. Boyce

Studies on Monstereae (Araceae) of Peninsular Malaysia I: Rhaphidophora

(MEV ALIN ALAAMO WIPE ECOL OiOR WIESE NI Al aY SIA sores scrapers eace ses u- si ce-wancsessassiesuaserscescsecsestccesetes

Ahmad Sofiman bin O., P.C. Boyce and Chan L.-K.

Studies on Monstereae (Araceae) of Peninsular Malaysia III: Scindapsus lucens,

a New Record for Malaysia, and a Key to Peninsular Malaysian Scindapsus..............:.00+-

Ardi, W.H. and M. Hughes

Begonia droopiae Ardi (Begoniaceae), a New Species of Begonia

RE GUELTR VES ER ULET cl Rites eee ce cc cece eee eee Sosa sco suceo lececawesueny sos vavabeunendocabossascecsaatevees

Arifiani, D.

Newly Recorded Endiandra R. Br. (Lauraceae) from Waigeo Island, Raja Ampat,

AGU AMMUTT CHOTA Scene esas eee ee cee etna evcn ene veseeccnuscccanncsodesu cane sasstesteccacereaessseseee

Atik R.

Species of Marasmius (Agaricales: Tricholomataceae) from Kayan Mentarang

IN AEC IT Ale ya Konya CATT ATI GANS Gl OMESTA oe ce cccsescacsoceovwovsacnvecasacdecscesarcecesesessscosencesscenccteees:

Hadiah, J.T. and B.J. Conn

Lectotypification of Elatostema subscabrum H.Schroet. (Urticaceae) .........:eeeeeeeeee

Haerida, I., S.R. Gradstein and S.S. Tjitrosoedirdjo

Lejeuneaceae subfamily Ptychanthoideae (Hepaticae) in West Java «0.0...

Kurzweil, H., S. Watthana and S. Lwin

Phaius takeoi (Orchidaceae) Newly Recorded from Thailand and Myanmat ...................

Leong-Skornickova, J., Tran H.-D. and M.F. Newman

Curcuma vitellina (Zingiberaceae), a New Species from Vietnam ..........:.:ccceeseeteeeteeeeeees

Lindsay, S.

New Combinations in Haplopteris (Adiantaceae) for the Flora of Peninsular Malaysia...

Nguyen, V.D., J.C. Regalado, Jr. and Vu T.C.

A New Species of Alocasia (Araceae-Colocasineae) from Cambodia ............:seeeee

ARNOLD A2B0RER ae]

UBRARY

ISSN 0374-7859

ore 9

43

119

121

Nor-Ezzawanis, A.T.

New Combinations VMalaysianis tap biylea cea ce sereeneeete teers cane nen veserse te seated seer et enree sans 127

Poulsen, A.D. and C.D. Specht

JN INDY S| VSOES Olt (COMES 1H KOA BOTAN EO) ecccecencaseospeccecnccerconsecontcnos acc -peccoconaaconnSotcenocecpenanancec 135

Rajbhandary, S.,M. Hughes and K.K. Shrestha

Three New Species of Begonia Sect. Platycentrum from Nepal ..............:csscccssesesesseseseneeees 143

Sudarmono and B.J. Conn

Genetic Variation of Populations of Scutellaria slametensis and S. discolor

(Lamiaceae) on Gunung Slamet, Jawa Tengah (Indonesia) ............::::ceeccescsssseeseseeseeecseesenees 155

Turner, I.M.

A new species of Polyalthia (Annonaceae) from Sabal «.............200.ccsccssscsececcesseeseesseossosseesece 173

Wong S.Y.

Studies on Schismatoglottideae (Araceae) of Borneo XIII: A Revision of the

SGI TRY UCT ONRMOPMAKY HA MUON! SI OASIS (COO) OES. cc rooecence-eeceecoccoccoreracecoccececoccocchLce ce pococtaASHocenSDOE SOS 177

Book Review

Flora of Peninsular Malaysia, Series I. Ferns and Lycophytes. Vol. 1. by B.S. Parris, R. Kiew,

R.C.K. Chung, L.G. Saw & E. Soepadmo (eds).

TE 1 Pal 24 (Opa 7 DEON BREET Oto dceoancoosoDODSOOOODNs0NS 2A

Date of publication: August 2010

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 62 (1): 1-8. 2010 l

Studies on Monstereae (Araceae) of Peninsular Malaysia II:

Rhaphidophora latevaginata, Newly Recorded for

West Malaysia

AHMAD SOFIMAN BIN OTHMAN AND PETER C. BOYCE *

Pusat Pengajian Sains Kajihayat

Universiti Sains Malaysia

11800 USM, Pulau Pinang, Malaysia

*Corresponding author: phymatarum@gmail.com

Abstract

Rhaphidophora latevaginata M.Hotta, a neotenic, shingling, climbing aroid,

hitherto considered a Bornean endemic, has recently been found and

collected from the southern part of the east coast of Peninsular Malaysia

(Johor: Kota Tinggi and Mersing), where so far it appears to be restricted

to kerapah and the drier (raised podzol) facies of seasonally inundated

peatswamp forest. This discovery of R. /atevaginata takes to 18 the number

of Rhaphidophora known to occur in Peninsular Malaysia, of which three

are endemic. An updated description of R. latevaginata, a key to the

Rhaphidophora species of Peninsular Malaysia, and a plate illustrating the

diagnostic characters of those with shingle-stage juveniles is presented. A

brief note on the significance of the new record with regard the Riau Pocket

is made.

Introduction

Since the publication of an alpha-taxonomy for Peninsular Malaysia (Boyce,

1999). and the Peninsular-relevant taxonomic alterations made for Borneo

(Boyce, 2001), further study of Rhaphidophora in Peninsular Malaysia

has generated additional data that sheds light on possible biogeographical

patterns not earlier apparent. Most recently the discovery in Perak of R.

megasperma Engl. (Baharuddin & Boyce, in press), previously regarded as

a Bornean endemic and furthermore belonging to a species group until now

considered to be restricted to E Sunda, Papuasia, and the tropical Western

Pacific, and now the discovery of R. latevaginata in Johor, is providing

compelling non-woody plant support to the Riau Pocket phytochore (Corner,

1960; Ashton, 2005). All terminology used here follows Boyce (1999).

— —

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Gard. Bull. Singapore 62 (1) 2010

Key to Rhaphidophora in Peninsular Malaysia (adult plants)

. Leaf lamina variously pinnately divided and/or perforated ...............:esceeseeseeeeeees 2

- Leaf lamina Cmte. is: si.i2..ccsecducascvesantie tessoee oe tae te oansateol cos canes: evs tee eudaegussaceacnde costeemn ere 9

Leaf lamina abaxially pubescent, especially the mid-rib and primary lateral veins.....3

Leaf lamina abaxially pla brotis:c.22:.jcccc. cede cccxecsee cece oe ceceseere ace ee 4

. Plants flowering on adherent stems; mature leaves with numerous perforations

along both) sides of the mtd nib x2. 22. -sssescssncee nen - sesecee a saseeeee ee eee eee R. foraminifera

. Plants flowering on free stems; mature leaves with one to three perforations on

one side of the mii rilb) ia. cs ccscesesoce-neesten as tt cps cece etna ae eee cee R. puberula

Rheophytes; leaves of flowering plants occasionally entire.................... R. beccarii

. Medium to high-climbing lianes; leaves of flowering plants never entire.............. 5)

. Active shoot apices with sparse to copious netted fibre; feeding roots conspicuously

ramentose-scaly; lamina of mature plants pinnatisect, the pinnae often perforated

basally and appearing stilted. Juvenile plants with leaves overlapping in the manner

ofroor shingles (Shingle: ClimDErs) he ccc. csee tte cee eee R. korthalsii

. Active shoot apices glabrous; feeding roots smooth or slightly corky-lenticillate;

lamina of mature plants variously split and perforated but never with the pinnae

perforated basally and appearing stilted. Juvenile plants not shingle climbers)...6

Leal lamina.at leastpartiallly pom ates. -c-ocerecseeecceee aces eee ee 6

. Leaf lamina not pinnate, either entire or with a few, large ovate to rhomboid or

trapezoid perforations, these sometimes extending almost to the margin............. 8

.Leaf lamina up to 53 x 105 cm; sparsely to rarely + entirely pinnatipartite,or

pinnatisect; petiole 40-70 cm long, petiolar sheath extending % - %4 along petiole;

spadix up to 14 x 2 cm, stoutly cylindrical, inserted decurrently from 2 cm on peduncle;

plants:exclusively of montane forest: se ccece tenes eeeee eee R. nicolsonii

Leaf lamina up to 42 x 38 cm but usually much smaller, sparsely to + entirely deeply

pinnatipartite to nearly pinnatisect, occasionally with large rhombic perforations

adjacent to mid-rib; petiole 10-34cm long, petiolar sheath extending to base of apical

pulvinus, inflorescences a few together, spadix up to 3.5 x 1 cm, cylindrical, inserted

slightly decurrently on peduncle; plants of lowland forest.............. R. tetrasperma

. Inflorescence two (sometimes more?) together, each subtended by a prominent,

soon falling, cataphyll, and arising from an elongated reiterative floral sympodium

at the tip of a plagiotropic free lateral shoot; spathe caducous; stigma impressed

irregularly elliptic, longitudinally ontentated in cc... cree ere eee R. corneri

. Inflorescence three or more together, each subtended by a prominent persistent,

later marcescent cataphyll, and arising from an much abbreviated reiterative floral

sympodium arising laterally on a climbing shoots; spathe marcescent; stigma raised,

PUT CHLORINE ones. 50s dcdsancaenscoanere <oseueseeevec hes oone Seem eens oe eee eo R. megasperma

Studies on Monstereae (Araceae) of Peninsular Malaysia II 3

9. Leaves always shingling, even in flowering individuals; leaf laminas stiffly coriaceous,

broadly oblong-ovate-elliptic, 848 x 6.5-20.5 cm, bright green, slightly to markedly

glaucous, base truncate-cordate to broadly cuneate. Flowering on clinging shoots

Pe ee rane eee aac ac ey connie reusing ssundesesdes sdesuoesnsseassssieeshosuseessacakdouews R. latevaginata

9. Leaves spreading in adult and flowering individuals; leaf laminas variously coloured

but never glaucous. Flowering on free or clinging ShHOOTS.............c:ecceseeeteeteeees 10

10. Abaxial surface of lamina and apical pulvinus pubescent.................. R. puberula

Mabamind anGapical PUlvinUs MOL AS-ADOVE...z.......<....c--censecessnesacsnssssesnnerteseonsedsatsacs 11

11. Flowering shoots consisting of scattered fans of large litter-trapping leaves carried

on short stout shoots and held at about 90” to the + leafless main stem................

ae SE tee Ae Sea oe cans cn satvandeqhdsnecsdustaidesnoenpcaestseateaveaueoeadeswedecetens R. angustata

GND OMe GIMPISMOOUS MOLI AS AD OW Cxco2.2520.520:.0.acdsascnseuccessvscosueccveccnsseecsecstetsaseoesbeedaesonced oz 12

12. Plant climbing; feeding roots smooth, or minutely asperate........... eee 13

12. Plant rheophytic; feeding roots conspicuously ramentaceous............... R. beccarii

13. Stems sub-terete to weakly 4-angled, scabrid to asperous, older portions with thin,

brittle pale brown epidermis; spathe exterior minutely puberulent......... R. lobbii

13. Stems conspicuously 4-angled, smooth, older portions remaining green; spathe

EE NAMEN AS EONS Bes eo os ns on Ses nas atnsabavoeisAuve susedibaceeoanseeswaveeteseasvoseacecsesvinveats 14

14. Apices of active stems with netted prophyll, cataphyll and petiolar sheath remains

SAS TASC a severe rece a csc SSST SELL cus SSoasee ve evensoscecesusdvessessisvsonsacavssvessueecieessies R. maingayi

Pes MAIC OMAC (IVC SU CMS TI AKC Glew sias2covnsn sien ieecenseanccoeceanrsnest-cosecsccsncsovsscuecoessssetsesedseceee 15

iSaspadix at anthesis 9-20 cm long, tapering apicallly.....................:cce-ceoresesseeseosonees 16

15. Spadix at anthesis 3.5-6 cm long, cylindrical to weakly clavate...........c:cceeee 17

igropadin 9-iticna lone; Samdstonme Se PTAMILE............2.200csseissseooesesccoseeneeeee R. montana

16s; Spadix 15-20 cm long; limestone obligate......:........:.s:.s-csssessessecessens R. burkilliana

1s SS ORYC TDS (57 SAG Ui Oe eee er Ee a PS Pe 18

RNiemS a AG aI ANY CAKINICIAN AL Coes ake canc eaters cessavececeaneecvtvevesertorsoectsbeenecteuseosessecevnenes R. sylvestris

18. Leaf lamina thickly coriaceous to almost fleshy, falcate-elliptic- lanceolate to

falcate- oblong or falcate-oblanceolate, 4.5-25.5 x 1.5-5 cm; margins slightly reflexed,

this becoming greatly accentuated in dried material.................0004 R. crassifolia

18. Leaf lamina thinly coriaceous, narrowly falcate-elliptic to falcate-lanceolate

or falcate-oblanceolate, 2.5-16 x 1.2-3 cm, margins flat, leaf drying pale straw-

CONIA sees ca ca wpe ence sab os soesacU vas osnstnap ac cosceseedsscdsnssbbeesdéadpesaasatnconaeesas R. minor

4 Gard. Bull. Singapore 62 (1) 2010

a

Plate 1. A-B. Rhaphidophora latevaginata M.Hotta. A. Leaf (abaxial view) showing the

strongly unequal wings of the long-persistent petiolar sheath. Note that the sheath is mainly

adnate to the moderately short petiole, with only a short free-auriculate portion. B. Juvenile

plant. C-D. Rhaphidophora kothalsii Schott. C. Leaf (abaxial view) showing the equally winds

to the swiftly-marcescent petiolar sheath. Note that the sheath is mostly free from the very

short petiole, with the greater part forming a long, narrowly triangular, free-ligulate portion.

D. Feeding root showing the diagnostic ramenta. Images © P.C.Boyce

Studies on Monstereae (Araceae) of Peninsular Malaysia II 5

Rhaphidophora latevaginata M. Hotta, Acta Phytotax. Geobot. 22: 44

(1966); Boyce, Gardens’ Bulletin Singapore 53: 51-54, Fig. 10 (2001). -Type:

Malaysia, Sarawak, Bintulu (‘4th Division’), Bintulu District, about 4 km

east from Minah Camp, Sg. Kakus, 4 Oct 1963, Hirano & Hotta 140 (KYO,

holo!). Plate 1A & B.

Moderate to very large, robust, pachycaul, homeophyllous neotenic liane to

12 m; seedling stage a non-skototropic shingling juvenile shoot; pre-adult

plants forming small terrestrial colonies of shingling closely appressed

leaves; adult shoot architecture comprised of clinging, physiognomically

unbranched, shingling to very densely leafy, sterile stems and almost identical

fertile stems. Stems weakly compressed-terete to weakly rectangular in cross-

section, smooth, pale green, without prophyll and cataphyll fibre, internodes

to 12 x 2 cm, separated by prominent straight scars, but scars obscured by

leaf bases on all but the oldest stems, lower parts of stem later sub-woody

with slightly shiny cracking thin pale brown epidermis; flagellate foraging

stems moderately well developed although often somewhat short and leafy;

clasping roots arising densely from the nodes and internodes, prominently

scaly; feeding roots ca 3 mm diam., brown, minutely pubescent. Leaves

distichous, appressed, ascending and shingling, becoming slightly scattered

and spreading (often litter-trapping) towards fertile tips; cataphylls and

prophylls membranous,soon drying black and persisting briefly before falling;

petiole deeply canaliculate and winged, 3-22 x 0.5-2 cm, smooth, apical and

basal pulvinus obscure in young leaves, later becoming prominent, especially

the basal pulvinus; petiolar sheath very pronounced, up to 2.5 cm wide,

rather thickly membranous, adnate to the petiole for much of their length,

prominently rounded short-auriculate, especially the larger (outer) sheath,

outer sheath (away from climbing surface) greatly expanded and partly to

completely obscuring stem, both sheaths persisting some considerable time,

much later (and then almost exclusively on adult plants) rotting to produce

two large scars extending to the top of the petiole; lamina broadly oblong-

ovate-elliptic, 8-48 x 6.5-20.5 cm stiffly coriaceous, bright green, slightly to

markedly glaucous, base truncate-cordate to broadly cuneate, very briefly

decurrent, apex rounded to acute with a tiny apicule; midrib prominently

raised abaxially, slightly sunken adaxially; primary venation pinnate, slightly

raised abaxially, more so adaxially; interprimaries sub-parallel to primaries,

slightly raised on both leaf surfaces; secondary venation tessellate-reticulate,

slightly raised abaxially, + flush adaxially, all veins much more prominent in

dried material. Inflorescence solitary on a clinging shoot, subtended by a

fully developed foliage leaf and one or more cataphylls; peduncle laterally

compressed-cylindrical, 6.5-11 x 0.5-0.7 cm; spathe not observed; spadix

stoutly cigar-shaped, sessile, inserted + level on stipe, 17.5 x 1.5 cm, pale

6 Gard. Bull. Singapore 62 (1) 2010

green; stylar region rhombohexagonal, ca 2 x 1 mm, truncate; stigma slightly

raised, elongated, longitudinally orientated, ca 0.75 x 0.2 mm; anthers not

exserted at male anthesis; infructescence stoutly cigar-shaped, 15 x 2 cm,

stylar region becoming convex at fruit maturity.

Specimens seen: MALAYSIA. Johor Bahru, Mersing, Kluang - Mersin Road,

km 39, 02° 15’ 78.2”; 103° 43’ 79.2” 56, 18 Apr 2010, PC Boyce, Siti Nurfazila

Abdul Rahman & Ooi Im Hin AR- 3039 (KEP); Johor Bahru, Kota Tinggi,

Hutan Simpan Panti, 01° 51’ 65.6”; 103° 54 10.7” 28, 19 Apr 2010, PC. Boyce,

Siti Nurfazila Abdul Rahman & Ooi Im Hin AR- 3046 (KEP). For Borneo

specimens see Boyce (2001).

Distribution: West Malaysia (Johor), Borneo (widespread throughout the N

and W, but much under-sampled).

Habitat: Primary to secondary moist lowland to hill dipterocarp forest on

clay and sandstone, in West Malaysia in kerapah and the drier (raised podzol)

facies of seasonally inundated peat swamp forest; 20-840 m altitude.

Notes: The juvenile and pre-adult stages of Rhaphidophora latevaginata and

R. korthalsii Schott are superficially similar and to non-specialist difficult to

differentiate. The most readily observable characters concern the petiolar

sheath, which in R. /atevaginata is long-persistent (vs very swiftly marcescent),

mostly adnate (vs mostly free) with the wings strongly unequal (vs weakly or

not at all unequal), and shortly free-auriculate (vs long, narrowly triangular

free-ligulate). Additionally, the petiole of R. latevaginata proportionately

longer (petiole:lamina ca. 1:5 vs ca. 1:12). See Plate 1A & B.

Pre-adult climbing stages of R. latevaginata and R. korthalsii are

also similar but aside from the petiolar sheath characters noted above are

readily distinguished by the feeding roots which are minutely pubescent in

R. latevaginata and conspicuously ramentaceous in R. korthalsii. See Plate

IC & D.

While abundant juvenile plants were located, often intermixed with

those of Rhaphidophora korthalsii, but only a single pre-adult, and no adult,

plants were located during the 4-days fieldwork; by contrast, pre-adult

and adult plants of R. korthalsii were frequent. Nonetheless we have no

doubt that the Bornean and West Malaysian plants are one and the same

species. Adult and fertile details given above are furnished from Bornean

collections.

Studies on Monstereae (Araceae) of Peninsular Malaysia II 7

Phytogeographical Implications

The discovery of another hitherto Bornean endemic in Peninsular Malaysia,

following from the recent finding of Rhaphidophora megasperma in Perak

(Baharuddin & Boyce, in press), gives further weight to the existence of a

‘Riau Pocket’ phytochore (Corner, 1960; Ashton, 2005).

Of particular interest is the growing evidence that the Riau Pocket

phytochore involves plants other than trees, on which distribution and

relationships the Riau Pocket was originally postulated. Of yet further

interest is that while R. /atevaginata occurs in the E coast of Riau Pocket

phytochore fragment, R. megasperma is so far known only from the Perak

phytochore fragment.

Figure 1. The Riau Pocket phytochore of West Malesia (areas enclosed within heavy lines

on northwestern Borneo, Peninsular Malaysia, and Central Sumatra). From: P.S. Ashton.

2005. Lambir’s Forest: The World’s Most Diverse Known Tree Assemblage? p. 199, Fig. 17.6.

Used with permission.

Acknowledgements

This work is funded under the Research University Grant of Universiti

Sains Malaysia (1001/PBIOLOGI/811132).

8 Gard. Bull. Singapore 62 (1) 2010

References

Ashton, P.S. 2005. Lambir’s Forest: The World’s Most Diverse Known Tree

Assemblage? pp. 191-216. In: D.W. Roubik, S.Sakai & A.A. Hamid

Karim (eds.). Pollination Ecology and the Rain Forest: Sarawak Studies.

Springer, New York.

Baharuddin, S. and P.C. Boyce. 2010. Studies on Monstereae (Araceae) of

Peninsular Malaysia I: Rhaphidophora megasperma, a new record for

West Malaysia. Tropical Life Sciences Research: in press.

Boyce,P.C.1999.The genus Rhaphidophora Hassk.(Araceae-Monsteroideae-

Monstereae) in Peninsular Malaysia, and Singapore. Gardens Bulletin

Singapore 51: 183-256.

Boyce,P.C.2001.The genus Rhaphidophora Hassk.(Araceae-Monsteroideae-

-Monstereae) in Borneo. Gardens Bulletin Singapore 53: 19-74.

Corner, E.J.H. 1960. The Malayan Flora, pp. 21-24. In: R.D. Purchon

(ed.). Proceedings of the Centenary and Bicentenary Congress of Biology.

Singapore.

Gardens’ Bulletin Singapore 62 (1): 9-15. 2010 9

Studies on Monstereae (Araceae) of

Peninsular Malaysia ITI:

Scindapsus lucens, a New Record for Malaysia, and a Key

to Peninsular Malaysian Scindapsus

AHMAD SOFIMAN BIN OTHMAN, PETER C. BOYCE*

AND CHAN LAI KENG

Pusat Pengajian Sains Kajihayat

[School of Biological Sciences]

Universiti Sains Malaysia

11800 USM, Pulau Pinang, Malaysia

*Corresponding author: phymatarum@gmail.com

Abstract

Scindapsus lucens Bogner & P.C.Boyce is a species of considerable horti-

cultural potential, arguably rivalling the commercially important S. pictus,

originally describedfromcultivated materialofunknownprovenanceandonly

later found wild in Sumatera. It has recently been discovered and collected

from several localities in southern Peninsular Malaysia, representing both a

new record for Malaysia, and through clonal propagation via various tissue

culture techniques would supply a sustainable potential source of a local

commercial ornamental horticultural product. An enlarged description of

S. lucens, and a key to the Peninsular Malaysian Scindapsus are offered. A

plate illustrating the diagnostic characters of S. /ucens is given.

Introduction

Scindapsus has not been revised in its entirety since 1908 (Engler & Krause,

1908), and not treated for Malaysia since Ridley’s accounts for the flora

(Ridley, 1907, 1925), when he recognized 5 species: Scindapsus beccarii Engl.,

S. hederaceus Migq., S. perakensis Hook.f., S. pictus Hassk., and S. scortechinii

Hook.f. Since Ridley, fieldwork has revealed two further species: S. treubii

Engl.,a widespread species from Sumatera to Borneo, and S. /ucens reported

here.

Scindapsus lucens Bogner & P.C.Boyce (1994) was described based on

material of unknown origin cultivated in the Botanischer Garten Miinchen,

Germany, with a note added late in proof that it had subsequently been

located in KabupatenTapanuli, western Sumatera Utara. Since publication,

10 Gard. Bull. Singapore 62 (1) 2010

consultation of all the major herbaria in Peninsular Malaysia, and other

herbaria with significant Malaysian and Indonesian collections (K, L, SING)

has not revealed a single collection of this distinctive and horticulturally

promising species. It was thus with some considerable surprise that during

a recent series of fieldwork in Peninsular Malaysia S. /ucens proved to be a

widespread species in Johor Bahru, Melaka, Negeri Sembilan, and Pahang.

Plants seemingly favour the drier facies of perhumid podzolic shady lowland

peatforest, although on occasions occurring with no less vigour in lowland

mixed dipterocarp forest.

These new collections have revealed additional morphological details

that were unavailable at the time of original publication, and have also

shown that the leaf colour is more variable than was initially reported, with

the leaf blade ranging from pale grey (as per the typical collection) to an

unusual (for Araceae) shade of jade green.

Key to Peninsular Malaysian Scindapsus

1. Short-climbing to perching epiphyte or scandentsemi-terrestrial............. 2

PANAMA CS oa ces scene iaeecasetss dees Sete bance eRe 2 es eR 3

2. Stout, short-climbing to perching epiphyte; leaves large, up to 75 x 25 cm,

carried in litter-trapping fan, glossy deep-green, oblanceolate. Spathe white.

Teo Watt USix. ssesacavczpicactaceseentiteueeteceanevaneren thet tases ee S. beccarii

2. Scandent semi-terrestrial, leaves small, up to 15 x 9 cm, scattered along

a sprawling stem, medium green, slightly glossy, ovate. Spathe yellow.

Highlands. :scccncontie utero eee ee S. scortechinii

(oS)

. Leaf blade oblong lanceolate, primary lateral veins impressed adaxially

and raised abaxially; petiole broadly conspicuously open-winged to the

| 101) 6p bb Lan Rr eR ear RE gion oe erccecer Acree cisaasustocedocecbocolecdoaconacc-o +

3. Leaf blade mostly cordiforme, if oblong-lanceolate then primary lateral

veins irtually invisible; petiole not or only very obscurely winged to the

yO) 01 kA 01 Uk nee SSE ne nor ne ere en CoC H oon sce accor oceecc nr Sacoestonsrenase 5

A Spadix at anthesis ca S:stemyl om Pees ee eer S. hederaceus

A Spadix at anthesis 1O=1S cml omg seeee cece eee eee S. perakensis

5. Stems smooth; leaf blade sub-succulent, markedly oblique, oblongo-lanceolate,

concolorous deep green, less often with jagged grey markings in juvenile

0) £201 RRP ern RP ee crere noc ta) cccconccdcbsocodsbaueudeoseooaeoc! S. treubii

. Stems scabrid or verruculose; leaf blade coriaceous but not markedly

succulent, not markedly oblique, condifoniness see enna eee 6

Nn

Studies on Monstereae (Araceae) of Peninsular Malaysia III 1]

6. Leaf blade conspicuously bullate, glossy pale grey to pale jade green; stems

Vermuculose, O1GEL POLUONS TEMAIMING PTEEMN.«..<......s2:s.cce0scse0eeseec: S. lucens

6. Leaf blade smooth, adaxially matte to somewhat scintillating, medium

green, usually with silvery jagged spots or with these coalescing into larger

areas; Stems at most scabrid, older portions with conspicuous orange-brown,

EUTUE LS SPDT (SIC ce ae eee eee S. pictus

Scindapsus lucens Bogner & P.C.Boyce, Kew Bull. 49: 789 (1994). — Type:

Origin unknown, cultivated in Botanischer Garten Miinchen, Bogner 2113

(holo,M!; iso,B!, K! (spirit)). Plate 1.

Slender, sparsely branched trunk climber to ca 4 m; primary axis sterile,

physiognomically monopodial, comprising superposed sympodia; lateral

branches fertile, rather few and short in wild plants, producing sub-terminai

continuation shoots after flowering. Stem up to 4mm diam., terete, epidermis

markedly verruculate, glossy medium-green, verrucate similarly coloured,

internodes of primary axis 3-8 cm long with one or two roots at each node;

roots 1-2 mm diam., with grey, spongy epidermis. Leaves of primary axis

rather distant, very regularly arranged, appressed to the substrate, those of

lateral branches, somewhat densely grouped, petiole terete, adaxial surface

slightly canaliculate; 3-5.5 cm x 2-3 mm, smooth, greyish to light green;

pulvinate apically, pulvinus 1-1.5 cm, usually visible only on older leaves

when the blade has moved its position relative to light; petiolar sheath

2-5 cm, almost reaching pulvinus; leaf blade 7-14 x 5-9.5 cm, cordiforme,

conspicuously bullate, coriaceous, base cordate, posterior lobes sometimes

overlapping, apex cuspidate to acuminate, margin entire, narrowly hyaline,

lamina adaxially glossy greyish to jade green with the raised parts of bullae

often paler, or in green forms tinged grey, paler and glossy abaxially; primary

lateral veins 5-7 on each side of the middle vein, ascending towards apex:

interprimary veins barely less prominent; secondary and tertiary venation

reticulate, much less conspicuous. Inflorescence solitary on the tips of lateral

branches: peduncle 5 cm x 4 mm diam., terete, green; subtending cataphyll

with very small lamina, ca 1-2 cm x 5 mm, otherwise resembling petiole;

spathe caducous during male anthesis, 5 x 15 cm, coriaceous, cuspidate,

exceeding spadix, somewhat constricted above spadix apex and thence

rostrate, very pale yellow, apical rostrum ca 7 mm long, pale green prior

to anthesis, all except the very tip becoming same colour as spathe during

anthesis; spadix ca 2.7 x 1 cm diam., oblong-ellipsoid, fusiform, pale yellow,

shortly stipitate, stipe ca 2 mm long. Flowers bisexual, naked, truncate;

gynoecium rhombohexagonal in plan view, 3.5-4.2 x ca 2 mm; stylar region

broader than ovary, ca 1 mm thick, pale yellow; stigma longitudinal-linear,

1- 1.8 mm long, brown; ovary ca 3 mm diam., unilocular, locule globular,

1 Gard. Bull. Singapore 62 (1) 2010

ca 1.2 mm diam.; ovule solitary, ca 1 mm long, funicle short, placentation

basal; stamens shorter than gynoecium, not extending markedly at pistillate

anthesis; filaments flat, ca 1 x 1-1.2 mm; thecae ellipsoid, ca 0.8 x 0.5 mm,

opening by a subapical slit. Infructescence unknown.

Specimens seen: MALAYSIA, Johor Bahru: Mersing, Kluang— Mersing road,

km 39, Lenggor FR.,02° 15.727’, 103 43.767’, 18 April 2010, PC. Boyce, Siti

Nurfazilah bt Abdul Rahman & Ooi Im Hin AR-3056 (KEP); Kota Tinggi,

Panti FR.,01° 52.226’, 103 54.755’, 19 April 2010, PC. Boyce, Siti Nurfazilah

bt Abdul Rahman & Ooi Im Hin AR-3057 (KEP); Kota Tinggi, Hutan Lipur

Panti; 01° 48.077’, 103° 57.202’, 19 April 201, PC. Boyce, Siti Nurfazilah bt

Abdul Rahman & Ooi Im Hin AR-3060 (KEP). Melaka: Machap, Hutan

Simpan Bukit Sedana, 02° 24’ 00.6”; 102° 20° 91.7”, 11 May 2010, PC Boyce

& Ng Kiaw Kiaw AR-3059 (KEP); Negeri Sembilan: Pasoh Forest Reserve,

3° 0’ 0; 102° 19° 60, 13 May 2010, PC. Boyce & Ng Kiaw Kiaw AR-3058(KEP);

Pahang: Kuala Rompin, Taman Negeri Endau Rompin, Waterfall trail, 02°

37° 08.9”; 103°20° 81.3”, 12 May 2010, PC. Boyce & Ng Kiaw Kiaw AR-3063

(KEP); Jerantut, Krau Wildlife Centre, main trail to river, 03° 49° 39.2”;

102°13’ 03.1”, 13 May 2010 PC. Boyce & Ng Kiaw Kiaw AR-3054 (KEP).

Distribution: Malaysia (Johor Bahru, Melaka, Negeri Sembilan & Pahang).

Indonesia, (western Sumatera Utara).

Habitat: Drier facies of perhumid lowland shady peatforest; less often in

lowland mixed dipterocarp forest (e.g., at Pasoh FR., Krau W.R.). 20-125

masl.

Notes: Scindpasus lucens appears to be a widespread species in southern

Peninsular Malaysia, with occurrences so far recorded in four states.

Curiously, given the complete absence of herbarium specimens, it appears

to be more abundant than S. pictus, despite the abundance of material of the

latter species in herbaria.

Potential of Scindapsus lucens as a sustainably-produced

ornamental horticulture subject

Scindapsus lucens is a widespread species in the southern Peninsular, and

although never abundant appears to be more prevalent than S. pictus

Hassk., the species to which S. /ucens is most similar. The very attractive

Studies on Monstereae (Araceae) of Peninsular Malaysia III

Plate 1. Scindapsus lucens Bogner & P.C.Boyce. A. Juvenile primary axis. Note the glossy

quality of the leaf blade (PC. Boyce & Ng Kiaw Kiaw A R-3054). B. Juvenile plant, jade-green

type.(P.C. Boyce & Ng Kiaw Kiaw AR-3063). C. Type clone in cultivation, Jardin Botanique

du Montet, Nancy, France (Bogner 23/1). D. Detail of stem showing the diagnostic verrucate

surface (P.C.Boyce & Ng Kiaw Kiaw AR-3054). Images A & B, D © P.C.Boyce; Image C ©

David Scherberich, used with permission.

14 Gard. Bull. Singapore 62 (1) 2010

Plate 2. A. Scindapsus treubii Engl. Note the subsucculent, strongly oblique leaf blade, and

smooth stem. B-E. Scindapsus pictus Hassk. B. Primary axis showing the diagnostic and

older portions with conspicuous orange, brittle scabrid epidermis. C-E. A selection of leaf

markings typical of S. pictus. Notes the somewhat scintillating quality particulary in C & E.

Images © P.C. Boyce.

Studies on Monstereae (Araceae) of Peninsular Malaysia II] 15

leaves and, for a lianescent aroid, small habit make S. /ucens an ideal subject

for consideration as a commercial ornamental product with via various in

vitro culture technology. The ease with which S. pictus is globally produced

in tens of millions for the ornamental horticulture market augers well that

S. lucens should also prove similarly amenable. In many respects S. /ucens

is a far more attractive plant than S. pictus, not in the least because it has a

much more compact habit, and is not a heteroblastic species and thus the

attractive juvenile stage is retained into adult-hood, unlike that situation

with S. pictus, which requires regular pruning to retain the commercially

desirable leaf form of the juvenile. Since S. /ucens has a high potential as

an attractive ornamental plant, it may be over-collected by irresponsible

individuals to fulfil the public demand for new ornamental plant in the

market. This may lead to the eventual extinction of this plant. To conserve

the germplasm and prevent over-collection from its natural habitat, S. /ucens

can be mass propagated via induction of multiple shoots formation using

shoot culture technique or the production of somatic embryogenesis using

the leaf as explants. Millions of plantlets can then be produced to fulfil the

market demand.

Acknowledgements

This work is funded under the Research University Grant of Universiti

Sains Malaysia (1001/PBIOLOGI/811132)

References

Boyce,P.C.1999.The genus Rhaphidophora Hassk.( Araceae-Monsteroideae-

Monstereae) in Peninsular Malaysia, and Singapore. Gardens’ Bulletin

Singapore 51: 183-256.

Engler, A. & K. Krause. 1908. Araceae — Monsteroideae. In: A. Engler (ed.),

Das Pflanzenreich 37(1V.23B): 4-139.

Ridley, H.N. 1907. Araceae. Materials for a Flora of Malay Peninsula. 3: 1-53.

Ridley, H.N. 1925. The Flora of the Malay Peninsula, 5: 84-131. Reeve & Co.,

London.

Gardens’ Bulletin Singapore 62 (1): 17-22. 2010 17

Begonia droopiae Ardi (Begoniaceae),

a New Species of Begonia

from West Sumatra

W.H.ARDI AND M. HUGHES

Bogor Botanic Garden

Jl. Ir. H. Juanda No. 13, P.O. Box 309, Bogor, Indonesia

- Royal Botanic Garden Edinburgh

20A Inverleith Row, Edinburgh EH3 SLR, U.K.

E-mail: m.hughes@rbge.ac.uk

Author for correspondence: wisn001 @lipi.go.id

Abstract

A new species of Begonia (Begoniaceae), B. droopiae Ardi,is described from

the Indonesian island of Sumatra. It belongs to Begonia sect. Reichenheimia

and is a limestone endemic in the Sawah Lunto District. Its IUCN threatened

category is considered to be ‘Vulnerable’.

Introduction

During an expedition to West Sumatra in August 2009 organised by The

Royal Botanic Garden Edinburgh, Kebun Raya Bogor and Andalas

University Herbarium (ANDA), a new species of Begonia, B. droopiae

Ardi, was collected from a limestone cave in the Batang Pangean I Nature

Reserve, Sawah Lunto District, West Sumatra Province. Begonia droopiae

is placed in Begonia sect. Reichenheimia because it exhibits the typical

characters of the section: rhizomatous habit, protandrous inflorescences and

three locular fruit with entire placentae.

This species-rich genus was previously represented by 52 species in

Sumatra (Hughes, 2008; Hughes er a/. 2009), although it is obvious from

herbarium collections that many more remain to be described from the

island. Begonia sect. Reichenheimia is the second-most species rich section

of the genus in Sumatra, with 11 species. All available herbarium specimens

in ANDA, BO, E and SING have been consulted, and hence it must be

assumed, at least until more intensive collecting in West Sumatra may reveal

otherwise, that this species has a very restricted range (Fig.1).

18 Gard. Bull. Singapore 62 (1) 2010

100°0'0"E 1050'0"E

00'0"

50'0"

95°0'0"E 100°0'0"E

Figure 1. Distribution of Begonia droopiae.

Begonia droopiae Ardi, sp. nov. (Sect. Reichenhemia)

Haec species Begoniae nurii similis est sed foliis magis asymmetricis, floribus

femineis tepalis 3 (nec 2) provisis et stipulis secus costam pilosis recedit.

—Typus: Indonesia, Sumatra, West Sumatra (Sumatra Barat), Batang Pangean

Nature Reserve, Sawah Lunto District, Nagari Solok Ambah, Perkaulan

cave, 00° 43’ 21.7” S, 101° 09’ 01.0” E, 484 m, 21 viii2009, A. J. Droop, W. H.

Ardi, Nurainas & Riki AJD173 (holo, BO; iso, E, BO, ANDA). Plate 1.

Perennial, creeping, monoecious herb, to ca 10 cm tall, rooting at the nodes,

hairy with up to ca 2 mm long, multicellular hairs. Stems rhizomatous,

internodes very short, up to ca 2 mm long, with scattered multicellular

hairs; stipules 3-6 x 3-5 mm, triangular, with an abaxially prominent midrib

forming an up to ca 4 mm long, thin, hairy appendage at the apex, abaxially

densely hairy along the midvein, persistent. Leaves alternate; petioles 5-

19.5 cm long, sparsely to densely hairy; lamina basifixed, 3.5-9.5 x 2-6.8 cm,

very asymmetric, ovate to elliptic, base cordate, lobes sometime slightly

overlapping, apex acuminate, margin broadly crenate and fringed with

hairs, adaxial surface glabrous, purplish-green to dark purple between the

Begonia droopiae Ardi (Begoniaceae) from West Sumatra 19

“>N ra

ee

«%

Plate 1. Begonia droopiae Ardi. A. Habit; B. Stipules (scale bar = 5 mm); C. Male flowers

(scale bar = 5 mm); D. Female flower (scale bar = 5 mm); E. Fruits (scale bar = 5 mm). A-E

based on A. J. Droop, W. H. Ardi, Nurainas & Riki AJD173.

20 Gard. Bull. Singapore 62 (1) 2010

veins, slightly raised, veins green, abaxial surface pale green to dark purple

between the veins, hairy on the veins, venation palmate. Inflorescences

cymose, axillary, protandrous, bisexual; peduncles 5-11 cm long, reddish,

glabrescent to sparsely hairy; bracts sub orbicular, ca 1.5-2.5 x 1-2 mm,

margin fimbriate, deciduous. Male flowers, pedicels 8-20 mm, moderately

hairy; tepals 4, two outer tepals, white or white with a tinge of pink, 11-17

x 6-7 mm, elliptic to sub orbicular, base slightly cordate, apex rounded,

abaxially moderately red hairy; two inner tepals white, 7-9.5 x 3-3.5 mm,

oblong-obovate, glabrous; androecium of ca 40 stamens, yellow, filaments ca

1 mm long, fused at the base, anthers ca 1-1.5 mm long, obovate, dehiscing

through unilaterally positioned slits ca 1/2 as long as the anther, connective

not projecting. Female flowers, pedicels 6-9 mm, glabrous; tepals 3, unequal,

two outer tepals orbicular to sub orbicular, 5.5-6 x 4.5-6 mm, one inner

tepal, obovate, 5-6 x 1.4-1.6 mm; ovary 6-7 x 10-13 mm, orbicular, locules 3,

placentation axile, placentae entire; wings 3, subequal, triangular, rounded at

the and the apex, the widest point at the middle of the ovary, glabrous; style

basally fused, 3-branched, each stylodium bifurcate in the stigmatic region,

stigmatic surface a spirally twisted papillose band, yellow. Fruits, on thin, 6-9

mm long, sparsely hairy pedicels; capsules 8-8.5 x 10-7 mm, (excluding the

wing) deflexed, dehiscent, splitting along the wing attachment, drying pale

brown, glabrous, wings 4-5 mm wide at the widest point (at the middle of the

ovary), wing shape as for ovary; seeds unknown.

Distribution: Indonesia, Sumatra, West Sumatra (Sumatra Barat), Batang

pangean nature reserve, Sawah Lunto District, Nagari Solok Ambah,

Perkaulan cave.

Habitat: This species is growing directly on vertical limestone wall in the

front of cave at 484 m altitude.

IUCN Conservation category: We consider this species to belong to the

VUD2 IUCN category, as it has a very restricted distribution. Although it

resides in a protected area, there is some evidence of small scale agricultural

activities encroaching the habitat nearby.

Notes: Begonia droopiae is morphologically similar to Begonia nurii Irmsch.

Both species exhibit a rhizomatous habit, patterned leaves, and have male

flowers with 4 tepals. However, B. droopiae can easily be distinguished

from B. nurii by its very asymmetric leaves with an acuminate apex, and

stipules which are hairy on the mid-vein (versus slightly asymmetric leaves,

with a rounded apex and stipules which are glabrous on the mid-vein in B.

nurit). The tepal number of the female flowers also differs; B. droopiae has

Begonia droopiae Ardi (Begoniaceae) from West Sumatra 21

three tepals, whilst B. nurii has two. The wing shape of the fruits is a further

difference: in B. droopiae the wings are rounded at the base and the apex,

whilst in B. nurii the wings are rounded at the base and cuneate at the apex.

Begonia rajah Ridlis another allied species from Peninsular Malaysia, similar

in its rhizomatous habit, patterned leaves and male flowers with 4 tepals (R.

Kiew, pers. comm.), but B. droopiae differs in having very oblique leaves

(the midrib is clearly at an acute angle) which are soft and not succulent; B.

rajah has thickly succulent leaves which are more bullate,1.e., raised between

the veins. Further, Begonia rajah has never been found on limestone; most

Peninsular Malaysia Begonia species grow either on limestone and not on

other rock types or vice versa (Kiew, 2005). A comparison of the two species

to Begonia droopiae is presented in Table 1. The epithet is after the first

collector, Alison Jane Droop.

Table 1. Comparison of Begonia droopiae, B. rajah and B. nurii.

Begonia Begonia Begonia

droopiae rajah nurii

Asymmetric Slightly asymmetric

Thickly succulent Thin

3.5-9.5 x 2-6.8 cm 7-15 x 6-15 cm 2-7 x 3-11 cm

: Cordate and Cordate and slightly Cordate and often

Lamina base : ; : ,

slightly overlapping overlapping overlapping

Lamina apex

Surface between

the veins

Character

Acuminate Short and acute Rounded

Slightly prominent Prominently raised Slightly prominent

eee

Hairy on mid vein Midvein glabrous

Stipules Midvein glabrous

Peduncle 5-11 cm long 10-25 cm long 4-19 cm long

8-8.5 x 10-12 mm 6-7 x 5-6 mm 5-9 x 10-16 mm

Fruit size

E th 3

: Rounded at the Romadediat the base Rounded at the base,

Wing shape sub truncate at the

base and the apex cuneate at the apex

apex

i)

i)

Gard. Bull. Singapore 62 (1) 2010

Acknowledgements

We are grateful to the Indonesian Ministry of Research and Technology

(RISTEK), the Indonesian Institute of Sciences (LIPI), Direktorat Jenderal

Perlindungan Hutan dan Konservasi Alam (DITJEN PHKA) for giving

us permission to conduct research in West Sumatra and for their help and

assistance. The curators of herbaria ANDA, BO, E and SING are thanked

for facilitating access to specimens. The fieldwork was supported by the

ASEAN Centre for Biodiversity. Prof. Ruth Kiew is gratefully acknowledged

for her constructive comments, and Dr. Robert Mill is thanked for his Latin

expertise.

References

Doorenbos, J., M.S.M. Sosef and J.J.RE. de Wilde. 1998. The sections

of Begonia including descriptions, keys and species lists (Studies in

Begoniaceae VI). Agricultural University Wageningen Papers 98(2): 1-

266.

Hughes, M. and M. Pullan. 2007. Southeast Asian Begonia Database.

Electronic publication accessible via: www.rbge.org.uk.

Hughes, M. 2008. An annotated checklist of Southeast Asian Begonia. Royal

Botanic Garden Edinburgh, UK.

Hughes, M., D. Girmansyah, W.H. Ardi and Nurainas. 2009. Seven New

Species of Begonia from Sumatra. Gardens’ Bulletin Singapore 61(1): 29-44

Kiew, R.2005. Begonias of Peninsular Malaysia. Natural History Publications,

Borneo.

Gardens Bulletin Singapore 62 (1): 23-30. 2010. 23

Newly Recorded Endiandra R. Br. (Lauraceae) from

Waigeo Island, Raja Ampat, Papua, Indonesia

D. ARIFIANI

Herbarium Bogoriense, Research Center for Biology, Indonesian Institute

of Sciences (Pusat Penelitian Biologi-LIPI), Jl. Raya Jakarta-Bogor

KM 46, Cibinong 16911,

West Java, Indonesia

E-mail: debyarifiani@yahoo.com

Abstract

Endiandra trees are not frequently encountered in the forest in Indonesia.

Recent exploration in Waigeo Island resulted in the collection of three

species of Endiandra. All three species were collected for the first time from

Waigeo Island. The three species are restricted in distribution to eastern

part of Indonesia (Papua Province) and Papua New Guinea. Key to the

species and species description are provided.

Introduction

Waigeo Island is part of the Raja Ampat Islands, an archipelago that

is situated at the northern extremity of Indonesia’s easternmost Papua

province. Waigeo Island is the largest island in the archipelago, 130 km long

and 48 km wide and positioned at the northernmost of Raja Ampat Islands.

Raja Ampat is very rich in both terrestrial and oceanic biodiversity including

plant, coral reefs and fish (Anonymous 2006).

Endiandra is one of the genera in the family Lauraceae, a dominant

plant family in tropical forests and occurs from lowland up to montane

forests. Main distribution area of Lauraceae species is in the tropical forests

of South East Asia and America (Heywood, 1993).

Endiandra was first described by R. Brown (1810) based on the

type species, Endiandra glauca, from Australia. The genus consists of

approximately 100 species distributed from South China, Taiwan, Malesia,

and Australia up to Fiji (Rohwer, 1993). In Malesia, New Guinea is the

main distribution centre of Endiandra with several endemic species. A

floristic survey to Waigeo Island was carried out to study the occurrence of

Endiandra on the island and to make additional collection for the Herbarium

Bogoriense.

24 Gard. Bull. Singapore 62 (1) 2010

Materials and Method

Herbarium specimens studied for this research were collected

from Waigeo Island. Additional specimens of Endiandra available at the

Herbarium Bogoriense were also examined. Main localities explored on

Waigeo Island are forests in the area of Teluk Mayalibit District and riverine

forest along the Werabiyai River.

We collected vegetative, floral and fruiting parts of each plant species

and took their pictures. Information, such as locality, plant habit, uses, color

and scent of specimens, were noted to help make the species identification

easier. Identification was done mainly by consulting references and

comparing the recently collected herbarium with the herbarium available

at the Herbarium Bogoriense. The descriptions for each species were

constructed based on observed morphological characters of the collected

specimens.

Results and Notes

The forests at the Waigeo Island are mostly of moist lowland type. The

areas of Warsamdim and Werabiyai on the island vary from slightly hilly to

steep slope. Easily accessible forests were disturbed, especially by logging

activities in the past. However, good vegetation can still be found on hills

and slopes.

Endiandra species are less known because they are not easily found

in the forests in general, neither in Waigeo forests. The more common

genera of Lauraceae in the area are Actinodaphne, Cryptocarya and Litsea,

of which young trees are easily spotted inside the forests. However, it is

very fortunate that in the recent floristic exploration in Waigeo Island, we

encountered three species of Endiandra.

Three species of Endiandra, i.e., Endiandra beccariana, E. grandifolia

and EF. papuana, were collected from the island.

Endiandra R. Br. (Lauraceae)

Taxonomically, the genus Endiandra is characterized by having alternate,

spiral and pinnately-veined leaves, inflorescences paniculate-determinate

with ultimate flowers not strictly opposite. Flowers trimerous, bisexual,

stamens 3, each with 2 thecae, glands present or absent. In some species

glands are united forming a disc surrounding the stamens and pistil. Fruits

are without cupule, free on receptacle.

Newly Recorded Endiandra R. Br. (Lauraceae) 25

The genus Endiandra is known for its wood. E. palmerstonii produces

high quality wood commonly used for good furniture and for building

construction.

Identification key to species of Endiandra in Waigeo Island

1. Lamina broadly elliptic, lateral veins 11-15 pairs, glands reniform, not united

SRPMS PANICAIS PES. os ee 3 eee ctcs-s.sesersersesesssseees 2. EB. grandifolia

1.Lamina elliptic, lateral veins 7-10 pairs, glands united forming disc-like

2 UL EL LEESS cc this Sok eee eee 2.

2. Leaf surfaces and panicles sparsely pubescent ............. 1. E. beccariana

2. Leaf surfaces and panicles densely pubescent ................ 3. E. papuana

Species description

1. Endiandra beccariana Kosterm., Reinwardtia 7: 5 (1969) 474. —Type:

Morotai Isl., District Tobelo, N. Totodoku, 14 May 1949, Kostermans &

Tangkilisan 146 (A, BO). Plate 1.

Tree up to 32 m, 40 cm dbh. Bark greyish brown, lenticellate. Young twigs

dense erect pubescent, rusty colored. Terminal buds densely pubescent,

rusty colored, 8 mm long and 3 mm wide. Leaves spiral, stiffly chartaceous,

elliptic; 8.5-19 x 6-11 cm; apex acuminate, base cuneate or slightly rounded:

both surfaces finely reticulate; upper surfaces shiny, midrib flat, lateral

veins slightly impressed; lower surfaces slightly shiny, sparsely pilose or

glabrescent, midrib raised; lateral veins 7-10 pairs, prominent below, arcuate

towards margin; petiole terete, densely pubescent to glabrescent, flat, 12-

23 x 10-20 mm. Inflorescences paniculate, axillary, 4-11 cm long, pubescent.

Pedicel slender, ca 3 mm long. Flowers yellow, ca 6 mm in diameter; tepals

fleshy, spreading, narrowly elliptic, slightly acuminate, 2.5-3 mm long; glands

united in a form of a disc, ca 0.75 mm thick, brownish yellow; anthers widely

triangular, emerge from gland-like disc; locules large; ovary ellipsoid,

glabrous; style 0.5 mm long, stigma unconspicous. Fruit ellipsoid, ca 1-2 cm

long, green, free on the receptacle.

Recently collected specimens: INDONESIA. Papua Province: Kabupaten

Raja Ampat, Waigeo Isl., District Teluk Mayalibit, Desa Warsamdim, 120

m, 8 Jun 2007, D. Arifiani & Obaja 596 (BO); ibid., + 50 m, 13 Jun 2007, D.

Arifiani & Obaja 652 (BO).

26 Gard. Bull. Singapore 62 (1) 2010

Additional specimens examined: INDONESIA. Sorong, Kadamah, 14 Aug

1948, Main 592 (BO); Manokwari, Warnapi 15 km N. of Ransiki, 10 m, 25

Sep 1948, Kostermans 448 (BO); Manokwari, Warnapi, 20 m, 30 Sep 1948,

Kostermans 491 (BO); Morotai Isl., along Sambiki R. (S.E. Morotai), 30 m,

22 May 1949, Kostermans 854 (BO); Morotai Isl., along Sambiki R., 100 m,

22 May 1949, Kostermans 890 (BO); Morotai Isl.,23 May 1951, Kostermans

934 (BO); N. slopes of upper Aifat Valley, between Senopi and Aifatfekaan,

W. of Kebar Valley, 10 Dec 1961, Moll BW 12908 (BO).

Note: Endiandra beccariana was first described by Kostermans in 1969 based

on specimens from Morotai Island, Manokwari, Sorong, and Ramoi. In our

exploration, we spotted the species in two locations on Waigeo Island.

This species is similar to the E. papuana Lauterbach, which both can

only be differentiated by checking the amount of indument on the lower leaf

surface and the inflorescences. Kostermans (1969) noted that the panicles

of E. beccariana are glabrous and I would not agree with his statement given

the fact that the panicles bear sparsely erect indument.

The specimens of EF. beccariana collected from this floristic survey were

anew record of the species occurrence on Waigeo Island. Known previously

from Morotai Island and Northwest part of West Papua, its occurrence

in Waigeo Island indicates a possible connection by seed dispersal from

Morotai Island to Papua and/or vice versa (Fig. 2).

2. Endiandra grandifolia Teschn., Engl. Bot. Jahrb. 58 (1923) 417. —Type:

Papua New Guinea, 10 May 1909, Schlechter 17691 (K).

Tree up to 25 m high, 45 cm dbh. Twigs hairy. Leaves simple, coriaceous;

broadly elliptic; 14-32 x 10-17 cm apex acute to rounded; base obtuse;

both surfaces finely reticulate, upper surfaces shiny, midrib slightly raised,

lateral veins slightly impressed, with indument; lower surfaces densely

tomentose; midrib raised below; lateral veins 11-15 pairs, prominent below,

arcuate towards margin; petiole 1.5 cm long, stiff, tomentose. Inflorescences

paniculate, compact, up to 9 cm long, axillary, densely tomentose, reddish

brown, many flowers. Tepals slightly unequal; elliptic; apex obtuse; tomentose.

Stamens elliptic, tip rounded, pubescent; glands reniform, stalked, pilose;

staminodia cordate, pilose, stalked. Ovary ovoid, glabrous. Fruit unknown.

Recently collected specimens: INDONESIA. Papua Province: Kabupaten

Raja Ampat, Waigeo Isl., District Teluk Mayalibit, Desa Warsamdim, low

alt., Jun 2007, Mirmanto & Ruskandi 09305 (BO).

ona

“s

;

'

.

Plate 1. Endiandra beccariana: A. Tree habit (D. Arifiani & Obaja 652); B. Leaf arrangement:

C. Flower buds; D. Mature flower, glands united forming disc-like appendages with stamens

exposed from the middle: E. Young fruit. (Photos C-E: A. Hidayat).

28 Gard. Bull. Singapore 62 (1) 2010

Additional specimens examined: PAPUA NEW GUINEA. Madang District,

Terr. of New Guinea, near the Gogol River by Mawan village (ca 25 km

inland), 60 m, 22 Jun 1955, Hoogland 4919 (BO); Morobe District, Lae

Subdistrict, Oomsis Logging area N.G.I., 180 m, Millar NGF 12036 (BO);

Morobe District, Menyamya Subdistrict, between Aseki & Menyamya,

Spreader Divide, 21 Nov 1970, Streimann & Kairo NGF 42451 (BO).

Note: E. grandifolia may be differentiated from both E. beccariana and

E. papuana by observing the shape or fusion of the glands. Some species

of Endiandra, including E. grandifolia, bear three pairs of glands at the

bases of each stamen. However, these glands sometimes may unite to each

other because of the limited space inside the flower and forming a disc-like

appendage surrounding the stamens. This is the case with both E. beccariana

and E. papuana.

This survey yielded the first collection of Endiandra grandifolia

from western part of New Guinea. Previously, the species was only collected

from Madang and Morobe districts of Papua New Guinea. Occurrence of

E. grandifolia in Waigeo Island extend its distribution toward northwestern

part of New Guinea (Fig. 2).

3. Endiandra papuana Lauterbach, Nova Guinea, Bot. 8: 2 (1912) 819. -

Type: Papua New Guinea, 6 Dec 1907, Branderhorst 263 (BO, K).

Tree up to 30 m. Twigs densely pilose, rusty colored. Terminal buds densely

pubescent, smooth, rusty colored. Leaves spiral, elliptic, stiffly chartaceous,

10-19 x 7-10 cm, apex acute, base obtuse to acute; both surfaces densely

reticulate; upper surface shiny, midrib flat, lateral veins slightly impressed;

lower surface slightly shiny, densely pilose, midrib raised; lateral veins 7-

8 pairs, raised, arcuate towards margin; petiole terete, densely pubescent,

1.5-2.0 x 0.20-0.25 cm. Inflorescences paniculate, densely pubescent, up to

18 cm long, axillary. Tepals spreading; glands large, united to form a disc-

like appendage. Fruits spherical to ellipsoid, 1-1.5 cm long, free on the

receptacles.

Recently collected specimens: INDONESIA. Papua Province: Kabupaten

Raja Ampat, Waigeo Isl., along Werabiyai River, Jun 2007, Mirmanto &

Ruskandi 09304 (BO).

Additional specimens examined: WEST NEW GUINEA, Albatros Bivouac,

7 May 1926, Docters van Leeuwen 9016 (BO). PAPUA NEW GUINEA,

Morobe District, Wareo, 450 m, 5 Feb 1936, Clemens 1782 (BO); Northern

Newly Recorded Endiandra R. Br. (Lauraceae) 29

Division, Terr. of Papua, between Mambare and Arumu Rivers. South of

Botue village (near Kokoda), 350 m, 21 Sep 1953, Hoogland 3955 (BO):

Oomsis Logging area, NW of Lae, Morobe District. T.N.G. 90m. 5 Mar

1959, White NGF 10487, Near Garagos, Lae. Bulolo Road. Morobe District.

450 m, 3 May 1962. Havel & Kairo NGF 11197 (BO): Titapuba, Morobe

District: 10 Jan 1966, Streimann & Kairo NGF 26160 (BO). Tributary of

Busu River, above Sankwep R., Lae Subdistrict, Morobe District. 13 Apr

1972, Wommersley NGF 43919 (BO).

Note: E. papuana

is More commonly encountered in the forests of Papua

New Guinea, around Morobe District than in Indonesian part of New

Guinea. Specimen of EF. papuana collected from Waigeo Island is a good

addition to the Herbarium Bogoriense collection (see Fig. 3).

(far

En? he: Raja Ampat

ESE

a = | . oS,

Si =. ae A ea =

~ ¥ cr oc - "

Paes Ss eee Wis + i

= ej = os : ~~

Py a ae a ee

S x Sa

re aot = — ‘ a.

> . =< fo 4 = %

[r~.* F fr) a

‘ * 5 4 |

| i et eo =o

si ome is ~

i st Es = =t

| —_ , j

sor enn She a

‘sal ee 4 i. TE

Figure 2. Distribution of Endiandra beccariana (m). E. grandifolia (a) and E. papuana (e).

: Waigeo

= £

? aS =<

ae

Fe _ : &

© c —

a —

ee oe

ee ee

Tu";

a ye

2 5S we s

—— rs f

a “Ue? 3

+

fae te

Figure 3. Occurrence of newly recorded species of Endiandra in Waigeo Island: Endiandra

beccariana (m). E. grandifolia (a) and E. papuana (@)

30 Gard. Bull. Singapore 62 (1) 2010

Acknowledgements

The field research was funded by DIPA project of the Research Center for

Biology-Indonesian Institute of Sciences (LIPI) for the fiscal year 2007/2008.

I would like to thank Nurdin, A. Sujadi, A. Ruskandi, people of Warsamdim

village (especially: Obaja, Andi, Yustus, Dani, Marcus, and Mattheus) for

their valuable assistance during the field survey, Suhardjono and Roemantyo

for assisting with distribution map. My respect goes to the late A. Ruskandi

for his enormous knowledge of plant identification who passed away in 2008

after his short battle with cancer. Indonesian Plant Taxonomy Association

(Penggalang Taksonomi Tumbuhan Indonesia) and DIKTI facilitated author

and editor meeting for manuscript preparation.

References

Anonymous. 2006. Atlas Sumberdaya Pesisir Kabupaten Raja Ampat, Provinsi

_ Irian Jaya Barat. Pemerintah Kabupaten Raja Ampat dan Konsorsium

Atlas Sumberdaya Pesisir Kabupaten Raja Ampat, Irian Jaya Barat.

Brown, R. 1810. Laurinae. In: Prodromus Florae Novae Hollandiae et Insulae

Van Diemen. Typis Richardi Taylor et Soci, London.

Heywood, V. 1993. Flowering Plants of the World. B. T. Batsford Ltd.

London.

Kostermans, A.J.G.H. 1969. Material for a revision of Lauraceae II.

Reinwardtia 7(5): 470-496.

Lauterbach, C. 1912. Lauraceae. Nova Guinea 8(2): 819-820.

Rohwer, J.G. 1993. Lauraceae. In: K. Kubitzki, J.G. Rohwer, V. Bittrich (eds.).

The Families and Genera of Vascular Plants IT. Springer Verlag, Berlin.

Teschner, H. 1923, Die Lauraceen Nordost-Neu-Guineas. In: A. Engler

(ed.). Botanische Jahrbiicher fiir Systematik, Pflanzengeschichte und

Pflanzegeographie 58: 413-420.

Gardens Bulletin Singapore 62 (1): 31-42. 2010.

Species of Marasmius (Agaricales: Tricholomataceae)

from Kayan Mentarang National Park, East Kalimantan,

Indonesia

ATIK RETNOWATI

Herbarium Bogoriense

Research Center for Biology, The Indonesian Institute of Sciences

Bogor, Indonesia

Abstract

Five species of Marasmius were encountered from forest surrounding

Pa 'raye village at Kayan Mentarang National Park, East Kalimantan: three

of them are described as new taxa (1 new species and 2 new varieties).

The five species of Marasmius are M. cf. purpureostriatus, M. guyanensis,

M. coklatus var. mentarangensis, var. nov., M. caryote var. pa’rayeensis, var.

nov., and M. gypseus, sp. nov. Comprehensive descriptions, illustrations,

and comparison with similar taxa are presented.

Introduction

Among the Agaricales in the tropics, Marasmius has become a favorite

collection item. Species of Marasmius grow relatively longer period of time

than other agarics. They are very attractive and relatively easy to be spotted.

It is the most common genus in tropic, and grows frequently on forest soil,

fallen leaves, and on dead or living wood and other plant tissues, such as

grass roots and bamboo twigs. Most species of Marasmius are saprophytic,

some are even parasitic, and none goes into mycorrhizal association.

The earliest report of Marasmius from Indonesia with 28 species

was made by Léveillé (1844, 1846) and many of them were reported from

Java. This was followed by several reports by Moritzi (1845-1846), Zollinger

(1854), Hennings (1900), Overeem and Overeem-de Haas (1922), Boedijn

(1940). Desjardin, Retnowati, and Horak (2000) documented 37 species

of the genus from Java and Bali. However, no species of Marasmius was

recorded from Kalimantan, particularly from Kayan Mentarang National

Park, East Kalimantan. This paper presents five species of Maramius

encountered during a three weeks expedition to survey the fungal flora

from the forests surrounding Pa’raye village at Kayan Mentarang National

Park in April of 2003.

ies)

ir)

Gard. Bull. Singapore 62 (1) 2010

Kayan Mentarang National Park, which lies between altitudes

116°15’-115° E and 2°-4°30’ N, embraces a total area of 1.35 million hectares.

It is located in East Kalimantan, bordering with Sabah and Sarawak. The

inventory was done from several collecting sites in the mountainous Pa’raye

village located north of the National Park. Descriptions of the species are

based on fresh collections made by the author. All micro characters were

studied from dried materials rehydrated in distilled water and 3% of KOH

solution, with the use of Melzer’s reagent or Congo Red dye. Color terms

and notations are those of Kornerup and Wanscher (1978). Spore sizes were

based on measurements of 25 basidiospores.

Spore statistics include - X = the arithmetic mean of the spore length

by spore width (+ standart deviation); Q, the quotient of spore length and

spore width in any one spore to indicate the range of variation in n spores

measured; q, the mean of Q-values in a single sample.

The basidiomes in illustration are of natural size, basidiospore (scale

bar = 12 um); basidia, cystidia, pileipellis and stipitipellis (scale bar = 15.7

um). All collections examined are deposited in Herbarium Bogoriense

(BO) and Harry D. Thiers Herbarium (SFSU).

Description of taxa

1. Marasmius cf. purpureostriatus Hongo

J. Jap. Bot. 33: 344. 1958. -Type: Japan, Otsu City, Ishtyama, 7 May 1957,

Hongo 1609 (Isotype, ZT [3221]). Fig. 1.

Pileus 55 mm diam, convex with slightly depressed center, sulcate; margin

crenate, straight; surface pruinose, smooth; greyish magenta (13-E3) overall.

Context thin, pale greyish magenta. Lamellae adnate, distant (12 attached

stipe) with 1 series of lamellulae, narrow, non-marginate; pale greyish

magenta. Stipe 180 x 5 mm, hollow, central, equal, cylindrical; smooth,

pruinose; pale greyish magenta. Odor and taste not distinctive. Basidiospores

unobserved. Basidia 46.4-56 x 12-12.8 um, 4-spored, clavate. Basidioles

clavate. Cheilocystidia common, 16-24 x 7.2-12.8 um, clavate, broadly clavate,

sub-globose to globose, hyaline to weakly yellow, thin-walled. Pleurocystidia

absent. Pileipellis hymeniform, clavate to broadly clavate or subglobose,

16-25.6 x 8.8-9.6 um, non-diverticulate, hyaline, thin-walled. Stipe tissue

monomitic; cortical and medullary hyphae indistinguishable, 6.4-20 um diam,

parallel, cylindrical, hyaline, thin-walled, strongly dextrinoid. Stipe vesture

common, composed of clavate, cylindrical to fusoid or irregular in outline,

18.4-32 x 8-12 um, hyaline, thin-walled. Clamp connections present.

oS)

ie)

Species of Marasmius (Agaricales: Tricholomataceae)

Specimen examined: INDONESIA. East Kalimantan, Kayan Mentarang

National Park, forest surrounding Pa’raye village, 4 Apr 2003, A. Retnowati

391 (BO, SFSU).

Distribution: Japan (type), Malaysia, Thailand, Republic of Korea, Papua

New Guinea, and Indonesia.

Habit and habitat: Solitary on soil.

Figure 1. Marasmius cf. purpureostriatus Hongo (A.Retnowati 39/). A. Basidiomes; B. Basidioles;

C. Cheilocystidia; D. Pileipellis; E. Caulocystidia.

Notes: M. cf. purpureostriatus is characterized by several peculiar features,

which include relatively large, greyish magenta pileus, sulcate, distant

lamellae and absence of pleurocystidia. Basidiospores of the material

examined are unobserved, but other characters match with description of

M. purpureostriatus (fide Desjardin and Horak, 1997). As a result, I identify

the material as M. cf. purpureostriatus.

M. purpureostriatus 1s allied with M. musisporus from Papua New

Guinea (Desjardin and Horak, 1997). M. musisporus differs in forming

smaller basidiome (8-25 mm diam), paller pileus, and the absence of stipe

vesture (Desjardin and Horak, 1997).

34 Gard. Bull. Singapore 62 (1) 2010

2. Marasmius guyanensis Mont.

Mont., Ann. Sci. Nat. Bot. 4(1): 114. 1854. -Type: French Guyana, Leprieur,

Dennis 255. (Holo, K). Fig. 2.

Pileus 2-5 mm diam, convex, with a small dark papilla at the depressed

center; margin sulcate, straight, crenate; surface dull, dry, smooth, glabrous;

orange (6-A6). Context thin, white. Lamellae adnate to a collarium, distant

(7-9 reaching stipe) with no lamellulae, narrow, white to off-white, non-

marginate. Stipe 5-7 x 0.1 mm, central, terete, pliant, smooth, glabrous, shiny,

insititious, cylindrical; dark brown to black; no-nodes, stipe arises from dark

rhizomorphs, black rhizomorphs association. Odor and taste not distinctive.

Basidiospores (11.2)12-13.6 x 4 um [x = 12.83 + 0.67x 4 +0 um, QO = 2.80-

3.40, q= 3.21 + 0.17, n = 25 spores per one specimen], elongate-ellipsoid,

smooth, hyaline, inamyloid, thin-walled. Basidia unobserved. Basidioles

fusoid, clavate. Cheilocystidia common, composed of Siccus-type broom

cells; main body 10.4-23.2 x 6.4-9.6 um, cylindrical to clavate, broadly clavate,

subglobose or irregular in outline; hyaline, thin walled; apical setulae 1.6-3.2

x 0.8 um, cylindrical to conical, subacute to obtuse or irregular in outline,

hyaline, thin-walled. Pleurocystidia absent. Pileipellis hymeniform, mottled,

composed of Siccus-type broom cells; main body 8.8-16 x 5.6-10.4 um,

cylindrical, clavate, broadly clavate, to subglobose or irregular in outline,

hyaline, thin-walled; apical setulae 2.4-3.2 x 0.8 um, cylindrical to conical,

obtuse to acute; hyaline, thin-walled. Stipe tissue monomitic; cortical

hyphae 3.2-5.6 um diam, parallel, cylindrical, smooth, hyaline; dextrinoid,

thin-walled; medullary hyphae 2.4-7.2 um diam, cylindrical, dextrinoid, thin-

walled. Stipe vesture absent. Clamp connections present.

Specimens examined: INDONESIA. Java, Bogor Botanical Garden, 7 Jan

1998, D.E. Desjardin 6713 (BO 98-4, SFSU); same location, 8 Jan 1998, D.E.

Desjardin 6719 (BO 98-19, SFSU);same location, 15 Jan 1998, D. E. Desjardin

6797 (BO 98-198, SFSU); same location, 12 Jan 1999, A. Retnowati 125 (BO

99-245, SFSU). East Kalimantan, Kayan Mentarang National Park, forest

surrounding Pa’raye village, 5 Apr 2003, A. Retnowati 412 (BO).

Distribution: Indonesia, Martinique, Dominica, and French Guiana.

Habit and habitat: Gregarious on leaves or arising directly from rhizomorphs.

Notes: Marasmius guyanensis firstly described by Montagne from French

Guiana (fide Singer, 1976). It is apparently a widespread species throughout

tropical Central and South America (Pegler, 1983) and Asia (Corner, 1996).

This species is easily distingushed by having a small, orange pileus with dark

Species of Marasmius (Agaricales: Tricholomataceae) 35

Figure 2. Marasmius guyanensis Mont. (A. Retnowati 412). A. Basidiomes; B. Basidiopores; C.

Basidioles; D. Cheilocystidia; E. Pileipellis.

papilla at the center, a presence of collarium, and basidiospores 12-13.6 x 4

uum and it is abundant in research sites.

3. Marasmius coklatus var. mentarangensis Retnowati, var. nov.

Pileus latus, convexus, velutinus. Lamellae adnatae, brunneus marginatae.

Stipes teres aequalis, minute pruinose, haud insititious. Odor saporque

nulli. Basidiosporae (5.6) 6.4-7.2 x 3.2-4 um, ellipsoideae, leves, hyalinae,

inamyloideae, tenui-tunicate. Basidiola 21.6-30.4 x 6.4-7.2 pum, anguste

clavata. Cheilocystidia cellulae typi Sicci, 11.2-4 x 4-8 um, setulosae, clavate

vel irregulars, hyalinae; 2-4 setulae ad apicem 6.4-13.6 x 0.8-2.4 um.

Pleurocystidia nulla. Pileipellis hymeniform, cellulae typi Sicci, 12-36 x 4.6-8

pum; 2-6 setulae ad apicem, 4-16 x 1.6-3.2 um. Gregarius ad terra. -Holotypus:

Indonesia. East Kalimantan, Kayan Mentarang National Park, 4 Apr 2003,

A.Retnowati 390 (BO). Fig. 3.

Pileus 26-80 mm diam, conic at first, then convex in age, hygrophanous;

margin incurved, surface strongly wrinkled at the disc, velutinous overall;

dark brown overall (9-F8). Context white, thick up to 6 mm. Lamellae

adnate, 24-26, closely attached to stipe, with 3 series of lamellulae, narrow,

dark brown marginate; pure white. Stipe 50-100 x 9-14 mm, hollow, cylindrical,

equal, center, non-insititious, smooth, pruinose; light brown. Odor and taste

36 Gard. Bull. Singapore 62 (1) 2010

Figure 3. Marasmius coklatus var. mentarangensis Retnowati, var. nov. (A. Retnowati 390,

Holotype). A. Basidiomes; B. Basidiospores; C. Basidia and Basidioles; D. Cheilocystidia. E.

Pileipellis; F. Caulocystidia.

not distinctive. Basidiospores (5.6) 6.4-7.2. x 3.2-4 um (x = 6.46 + 0.51 x

3,39. + 0:35, O = 1.60- 2.2 5, gq =1.92 = 0:17; mn = 25ispores pen lespecimen):

ellipsoid, smooth, hyaline, inamyloid, thin-walled. Basidia 21.6-30.4 x 6.4-7.2

um, clavate, 4-spored. Basidioles clavate. Cheilocystidia common, composed

of Siccus-type broom cells, main body 11.2-4 x 4-8 um, clavate, subcylindrical

to cylindrical or irregular in outline, yellowish brown, thin to thick-walled;

apical setulae 6.4-13.6 x 0.8-2.4 um, 2-4 setulae, narrowly cylindrical to

conical or irregular in outline, subacute to acute, hyaline, thin to thick-

walled. Pleurocystidia absent. Pileipellis hymeniform, mottled, composed of

Siccus-type broom cells; main body 12-36 x 4.6-8 tm, clavate, subclavate,

cylindrical to subcylindrical or irregular in outline, yellowish brown, thin to

thick-walled; apicals setulae 4-16 x 1.6-3.2 um, 2-6 setulae, often branched

at the apex, narrowly cylindrical to irregular in outline, yellowish brown,

thick-walled. Stipe tissue monomitic; cortical hyphae and medullary hyphae

undifferentiated, 6.2-12 um diam, hyaline to weakly yellowish brown,

thin-walled, dextrinoid. Stipe vesture common, composed of two types of

caulocystidia: a) Siccus-type broom cells, main body 20-66.4 x 3.2-4.8 um,

clavate, clavate to broadly clavate, cylindrical or irregular in outline, hyaline,

Species of Marasmius (Agaricales: Tricholomataceae) 337)

thin-walled; apical setulae 3.2-6.4 x 1.6 um, not crowded, narrowly cylindrical

to conical, obtuse to acute, thin to thick-walled up to 0.8 um; b) non-setulae,

main body 35-36 x 5.6 um, clavate, cylindrical, thin-walled, hyaline. Clamp

connections present.

Distribution: Indonesia.

Habit and habitat: Gregarious on soil.

Notes: Marasmius coklatus was described from Cibodas Botanical Garden

by Desjardin, Retnowati and Horak (2000). This javanese materials have a

dark chocolate brown, velutinous pileus, remote to distant, broad, greyish

brown lamellae, a brownish orange to brown, pruinose stipe, moderately

long and broad basidiospores, Siccus-type cheilocystidia and pleurocystidia

with few (2-5), long (-30 um) apical setulae and numerous pileo-, cheilo-,

pleuro-, and caulosetae. The material collected from KMNP differs from

the Javanese material in having shorter basidiospores (6.4-7.2 x 3.2-4) um,

and the absence of pleurocystidia. These two characters separate the KMNP

collection as a new variety of M. coklatus.

4. Marasmius caryote var. pa’rayeensis Retnowati, var. nov.

Differt a pileo sulcatus, lamellis numerosus, basidiosporis longissimus

(28)30.4-33.8 x 4.8-5.6 um longis), stipitis caulocystidiis cylindricis deim

ordinary. Sparsus ad solum. —Holotypus: Indonesia, East Kalimantan,

Kayan Mentarang National Park, forest along Parinibung, 10 Apr 2003,

A. Retnowati 460 (BO). Fig. 4.

Pileus 17-48 mm diam, convex at first, then convex with upturned margin

in age; hygrophanous, sulcate; margin straight at first, upturned in age;

surface glabrous, smooth; light brown. Context thin, light brown. Lamellae

adnate, close (18 reached stipe) with 2-3 series of lamellulae, narrow, non-

marginate; light brown. Stipe 100-145 x 1.5-2 mm, equal, cylindrical, central,

non-insititious; smooth, pruinose; light brown to dark brown; white base

tomentose at the base. Odor and taste not distinctive. Basidiospores (28 )30.4-

B55 645-50) fon | x— 31.85 += 1.42 x 5.12 + 0.40 pm; O = 5.43-7, g = 6.25 +

0.51, n = 25 spores per | specimen], cylindrical, smooth, hyaline, inamyloid,

thin-walled. Basidia 42.4-44 x 10.4 um, 4-spored, clavate. Basidioles clavate.

Cheilocystidia abundant, 10.4-28 x 8-12.8 um, clavate, broadly clavate to

pyriform or globose, hyaline to weakly yellow, thin-walled. Pleurocystidia

absent. Pileipellis hymeniform, not mottled, 12-21.6 x 5.6-9.6 um, clavate

to broadly clavate, subglobose or globose, hyaline, thin-walled. Stipe tissue

monomitic; cortical hyphae 7.2-16.8 tm diam, parallel, cylindrical, hyaline,

38 Gard. Bull. Singapore 62 (1) 2010

thin- walled,smooth, weakly dextrinoid; medullary hyphae 5.6-10.4 um diam,

parallel, cylindrical, hyaline to yellowish brown, thin-walled, dextrinoid.

Stipe vesture common, 16.8-40 x 4.8-7.2 um, cylindrical to clavate or irregular

shape, hyaline, thin-walled. Clamp connections present.

Figure 4. Marasmius caryote var. pa’rayeensis Retnowati, var. nov. (A. Retnowati 460, Holotype).

A. Basiodiomes; B. Basidiospores; C. Basidia and basidioles; D. Cheilocystidia; E. Pileipellis;

F. Caulocystidia.

Distribution: Indonesia.

Habit and habitat: Scattered on soil.

Notes: Some species of Marasmius, e.g., M. bekolacongoli Beeli (fide Singer,

1965) and M. titanosporus Reid & Guillarmod (Reid, 1988), are closely

related to the M. caryote var. pa’rayeensis; however, the Kalimantan material

differs in having longer basidiospores from M. bekolacongoli (16.5-23(-28)

x 3.5-5 um), and shorter basidiospores from M. titanosporus (22-36 x 5-7.2

um). In addition, the presence of stipe vesture in this Marasmius makes it a

new variety different from the related species. M. caryote variety typical had

been reported from Krakatau Island by Boedijn (1940).

Species of Marasmius (Agaricales: Tricholomataceae) 39

5. Marasmius gypseus Retnowati, sp. nov.

Pileus 22-41 mm latus, convexus, hygrophanous, crenatus, glabrus, gypseus.

Lamella adnate, distantes, haud marginatae. Stipes 40-60 x 1.5-3 mm, aequalis,

glabrus, Haud insititious, brunneus. K-Basidiospores 6.4-7.2 x 3.2-4 um,

ellipsoideae, leves, hyalinae, inamyloideae. Basidiola clavate, cheilocystidia

et pleurocystidia nula. Pileipellis hymeniformis, clavate, prelate clavate,

globosus vel subglobosus, leves, 16-26.4 x 8-18.4 um. Stipitis caulocystidis

ordinary, 20-24.8 x 6.4-12 um, clavate, globosus vel subglobosus. Cespitosesus

vel gregarious ad lignum. —Holotypus: Indonesia. East Kalimantan, Kayan

Mentarang National Park, trail to sub-camp 1 in forest surrounding Pa’raye

village, 3 Apr 2003, A. Retnowati 371 (BO). Fig. 5.

Pileus 22-41 mm diam, convex with depressed center, strongly hygrophanous,

crenate; margin straight, wavy; surface glabrous, wrinkle; white overall.

Context thin, white. Lamellae adnate, distant with 2 series of lamellulae,

narrow, non-marginate, white. Stipe 40-60 = 1.5-3 mm, cylindrical, center,

hollow, equal, mooth, glabrous, non-insititious; brown. Odor and taste not

distinctive. Basidiospores 6.4-7.2 x 3.2-4 um [xX = 6.78 + 0.41 x 3.33 + 0.30

um, Q = 1.60-2.25, q = 2.05 + 0.18,n = 25 spores per one specimen], ellipsoid,

smooth, hyaline, inamyloid, thin-walled. Basidia unobserved. Basidioles

clavate, fusoid. Cheilocystidia and pleurocystidia absent. Pileipellis

hymeniform, composed of clavate, broadly clavate, globose to subglobose,

non-diverticulate, 16-26.4 x 8-18.4 um, hyaline, thin-walled. Stipe tissue

monomitic; cortical and medullary hyphae indistinguishable, 5.6-12.2

um diam, parallel, cylindrical, hyaline, smooth, thin-walled, dextrinoid.

Stipe vesture common, composed of clavate, broadly clavate, globose to

subglobose, non-diverticulate, scattered, 20-24.8 x 6.4-12 um, hyaline, thin-

walled. Clamp connections present.

Distribution: Indonesia.

Habit and habitat: Caespitose to gregarious on wood.

Notes: This new Marasmius can be distinguished by having convex with

depressed center pileus, strongly hygrophanous and distant with 2 series of

lamellulae, the lacking of cheilocyctidia and pleurocystidia, and basidiospores

6.4-7.2 x 3.2-4 um. Several members of section G/obulares have a similar

40 Gard. Bull. Singapore 62 (1) 2010

Figure 5. Marasmius gypseus Retnowati, sp. nov. (A. Retnowati 371, Holotype). A. Basidomes; B.

Basidiopores; C. Basidioles; D. Pileipellis; E. Caulocystidia.

feature with this Marasmius. M. albertianus Singer described from Congo

has longer basidiospores 8-10 x (3)-4(-5) um, presence of cheilocystidia,

and is host specific on bamboo (fide Pegler, 1977). M. niveus from French

Guyana (fide Singer, 1976) also has longer basidiospores 6.5-9 x 2.5-3.5 um,

and absence of cheilocystidia and stipe vesture.

Acknowledgment

The author thanks to Simon Taka Nuhamara (BIOTROP), Danang Anggoro

(BKSDA Kalimantan Timur), and Deni Wahyudi (Mulawarman University)

for help during the fieldwork. I am also thankful to Prof. Dr. Mien A. Rifai

for his useful comments on this paper, and to Indonesian Plant Taxonomy

Association (PTTI) which facilitated the editors of scientific journals to

review this paper. This research was funded by World Wide Fund (WWF)

Indonesia programme to do fieldwork in Kayan Mentarang National Park

in East Kalimantan.

Species of Marasmius (Agaricales: Tricholomataceae) 4]

References

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Bulletin du Jardin botanique de Buitenzorg III, 16: 398-399.

Corner, E.J.H. 1996. The agarics genera Marasmius, Chaetocalathus,

Crinipellis, Heimiomyces, Resupinatus, Xerula, and Xerulina in Malesia.

Bethefte zur Nova Hedwigia 111: 1-141.

Desjardin, D.E. and E. Horak. 1997. Marasmius and Gloiocephala in South

Pacific Region: Papua New Guinea, New Caledonia, and New Zealand

Taxa. Bibliotheca Mycologica 168: 1-152 .

Desjardin, D. E.,A. Retnowati and E. Horak. 2000. Agaricales of Indonesia..

2.A preliminary monograph of Marasmius from Java and Bali. Sydowia

52(2): 92-93.

Hennings, P. 1900. Fungi Monsunenses. Monsunia 1: 15-16; 150-151.

Kornerup, A. and J.H. Wanscher. 1978. Methuen Handbook of Colour. 3rd.

Ed. Eyre Methuen, London. 252 pp.

Overeem, C. van and D. van Overeem-de Haas. 1922. Verzeichnis der in

Niederlandisch Ost Indien bis dem Jahre 1920 gefundenen Myxomycetes,

Fungi, und Lichens. Bulletin du Jardin botanique de Buitenzorg III, 4: 88-

89.

Léveillé, J.H. 1844. Champignons exotiques. Annales de la Societe des

Sciences Naturelles (Paris), 3. sér., 2: 167-221.

Léveillé, J.H. 1846. Descriptions des Champignons de |’Herbier du Muséum

de Paris. Annales de la Societe des Sciences Naturelles (Paris), 3. sér., 5:

111-167.

Moritzi, A. 1845-1846. Verzeichnis der von H. Zollinger in den Jahren 1842-

1844 auf Java gesammelten Planzen. Solothurn. 144 pp.

Pegler, D.N. 1977. A preliminary agarics flora of East Africa. Kew Bulletin

Additional Series. 6: 1-615.

Pegler, D.N. 1983. Agarics flora of Lesser Antilles. Kew Bulletin Additional

Series. 9: 195-232.

42 Gard. Bull. Singapore 62 (1) 2010

Petch, T. 1947. A revision of Ceylon Marasmii. Transactions of the British

Mycological Society. 31: 19-44.

Reid. 1988. Marasmius titanosporus, a new species from the eastern Cape,

South Africa. Transactions of the British Mycological Society. 91(4): 707-

909.

Singer, R. 1964. Marasmius congolais recueillis par Mme Goosens-Fontana

et d’autres collecteurs belges. Bulletin du Jardin Botanique de I Etat.

[Bruxelles.] 34: 317-388.

Singer, R. 1965. Monographic Studies on South American Basidiomycetes,

especially those of the east slope of the Andes and Brazil. 2. The genus

Marasmius in South America. Sydowia 18: 106-358.

Singer, R. 1976. Marasmieae (Basidiomycetes-7richolomataceae). Flora

Neotropica Monograph 17: 1-347.

Takahashi, H. 1999. Marasmius brunneospermus,a new species of Marasmius

section Globulares from central Honshu, Japan. Mycoscience 40: 477-

481.

Takahashi, H. 2000. Three new species of Marasmius section Sicci from

eastern Honshu, Japan. Mycoscience 41: 313-321.

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in den Jahren 1842-1848 gesammelten sowie aus Japan empfangenen

Pflanzen. Zurich. 160 pp.

Gardens’ Bulletin Singapore 62 (1): 43-51. 2010 43

Lectotypification of Elatostema subscabrum H.Schroet.

(Urticaceae)

JLT. HADIAH AND BJ.CONN>

‘Kebun Raya Bogor, Jl. Ir. H. Juanda 13, PO Box 309 Bogor 16122,

Indonesia

: Email address: jhadiah@yahoo.com

- National Herbarium of New South Wales, Mrs Macquaries Road, Sydney

NSW 2000, Australia

Email address: bauy.conn@bgsycl.nsw.gov.au

Abstract

Descriptions of Elatostema sessile J.R.Forst. & G.Forst and E. subscabrum

H.Schroet.(Urticaceae) are provided to clarify the morphological differences

between these two species. The lectotype of E. suwbscabrum H.Schroet.

(Urticaceae) is here selected.

Introduction

Elatostema J.R.Forst. & G.Forst. (Urticaceae) is a very speciose genus of at

least 300 species occurring throughout the Old World tropics, subtropics and

subtemperate regions. The general lack of knowledge about the systematics

and circumscriptions of the species has resulted in the frequent incorrect

application of plant names because there are no recent publications on

the systematics of this group. The most recent account of the genus by

Schréter and Winkler (1936) was based on the relatively few collections

that were available. Furthermore, this publication only described species

of subgenera Elatostematoides, Pellionia, and Weddellia (as circumscribed

by them). Species of Elatostema sensu stricto, that Schroter and Winkler

(1935) regarded as consisting of 240 species, were not included in their

publication.

Recently, progress has been made on our understanding of the

taxonomy and phylogeny of the Urticaceae (Friis, 1993; Hadiah er al.,

2003, 2008; Sytsma et al., 2002; Wilmot-Dear, 2009) and tribes, in particular

Elatostemateae (sensu Conn and Hadiah, 2009). Traditionally, the

taxonomic studies of the genus were based on morphological characteristics

(Robinson, 1910, 1911; Schréter and Winkler, 1935, 1936; and Weddell, 1854,

1856, 1857, 1869). A detailed discussion of the morphological characteristics

used to circumscribe taxa within Elatostema and their usefulness in re-

44 Gard. Bull. Singapore 62 (1) 2010

construction of phylogenies are presented in Hadiah and Conn (2009).

There has been a large volume of work on the taxonomy of the Chinese

species of Elatostema sensu lato (Wang, 1980a, 1980b; Yang et al., 1995),

one of the centres of species diversity for the genus. However, little has

been published on the Malesian species, which are part of another major

centre of diversity. With relevance to this paper, the above studies provide

an, albeit incomplete, framework for further systematic study of the genus.

During studies of the Malaysian species of Urticaceae, material of

E. sessile has often been confused with E. subscabrum. Two herbarium

sheets of type material of E. subscabrum were located at the herbarium

of the Singapore Botanic Gardens (SING). These specimens clarify the

circumscription of the latter species.

Elatostema subscabrum H.Schroet.

Repertorium specierum novarum regni vegetabilis 83: 1 (1935) 20 in obs.;

Repertorium specierum novarum regni vegetabilis 83:2 (1936) 85.

Lectotype (here chosen): Malaysia, Pahang, Telom, Nov 1908, H.N. Ridley

13789 (upper left specimen, SING s.n.); isolectotype (lower right specimen,

SING s.n.). Other syntype material: H.N. Ridley 13789 (SING67093) (refer

to discussion of lectotypification, below).

Terrestrial herb, 0.15-0.2 m_ high, self-supporting (erect/suberect) or

spreading; internodes developed (elongate, distinct); branched hairs

lacking; stinging hairs absent; monoecious. Stipules axillary, interpetiolar,

persistent, free, 1-1.3 mm long. Leaves opposite, appearing alternate (by

misinterpretation — nanophylls caducous); petiole absent (or < 2 mm long);

megaphylls with lamina 23-60 mm long, 12-24 mm wide (length to width

ratio 1.92-2.5), unequal-sided, larger side of lamina ovate to elliptic, smaller

side elliptic to obovate; surface flat, not rugose, with 4 or 5 vein-pairs;

venation actinodromous (‘type IVd, semi pinnate nerved’ sensu Schroter

and Winkler, 1935); basal pair of secondary veins arising from above base

of primary vein and arising from different points (more than 2 mm apart),

both directed towards margin (or almost so), joined to next distal secondary

vein; abaxial surface with cystoliths present on interstices (cystoliths linear),

with hairs on primary, secondary and tertiary veins; adaxial surface lacking

cystoliths, glabrous; base oblique, rounded or cuneate; margin toothed, not

lobed, glabrous; apex short-acuminate; nanophylls not known. Inflorescences

unisexual; male inflorescences not known; female inflorescence subsessile

to shortly pedunculate, unbranched, head-like, involucral bracts narrowly

ovate, sparely hairy to almost glabrous; bracteoles very long, narrowly

ovate; flowers condensed/crowded, unisexual; actinomorphic (or slightly

Lh

Lectoiypification of Elatostema subscabrum 45

asymetrical): tepals 4. free. unequal, one tepal c. 1mm long with appendage

1-1.5 mm long: 3 tepals slightly smaller, ca 0.75 mm long without appendage:

staminodes present. 4, inflexed in bud: ovary straight: style absent: stigma

oblong, filiform to linear. Achene not enclosed (or only partly so): surface

ribbed or punctate.

Lectotypification of Elatostema subscabrum H.Schroet.

Hilde Schréter described the new species E. subscabrum from material

collected from Telom, Pahang, Malaysia, with the type cited as Ridley

13789 (SING) (Schroéter and Winkler, 1936). Two sheets of syntype material

are held at SING. One herbarium sheet is annotated: *H.N. Ridley 13789.

Noy, 1908, Telom [all written in Ridley’s hand]. State of Pahang, Malay

Peninsula [State and region part of pre-printed label — lower right of

sheet]? (SING67093), with field identification *Elatostema ?acuminatum™

[in Ridley’s hand] has three separate samples of the plant: all specimens

are female and largely in fruit (Fig. 1). The second sheet. °H.N. Ridley

13789, Nov, 1908, Telom, State of Pahang, Malay Peninsula’ [as annotated

above — upper left of sheet] (SING), except identified as *E. acuminatum.

Brongn. by ‘CXF’ [Caetano Xavier Furtado, Singapore Botanic Gardens,

1923-1952, then 1956-1960] and the label has been stamped “30 MAR,

1909, presumably the date when the material was accessioned into the

herbarium (Fig. 2). This second sheet has two female specimens, upper left

with flowers and fruits, and the lower right specimen largely in fruit. Both

sheets have been examined by Schréter and annotated as “Elatostema

subscabrum Hilde Schroter [in her hand], dated *1935° [in her hand]. and

‘det. Hilde Schréter [pre-printed determinavit slip]. Furthermore, Schréter

notes that she has not examined male inflorescences (refer protologue),

only female inflorescence, female flowers and fruit. The upper left specimen

of the latter sheet (SING) is a good match for the protologue. Therefore, it

is here selected as the lectotype because it has both flowers and fruits (in

accordance with Articles 7.11, 8.1-8.3: McNeil er al., 2006).

Lectotype (chosen by Florence, 1997): J.G.A. Forster [186], Société, Tahiti (P-

Forst): isolectotype: Forster sn. (BM, K): probably isolectotype: *Dorstenia

pubescens’ (MW). For further discussion refer Nicolson and Fosberg (2003,

pp. 677 & 678).

Distribution: Malaysia; Malay Peninsula, Pahang — only known from type.

Notes: Collections of E. sessile from Malaysia have frequently been mis-

identified as E. subscabrum: however, the two species can be distinguished

by the morphological characters listed in Table 1 (below).

46 Gard. Bull. Singapore 62 (1) 2010

: ale ey Atlan fl

Wy bed Y

Rass hla

ee

Herb. Hort. Bot

067093

Singaporensis

Figure 1. Isolectotypes of Elatostema subscabrum H.Schroet. (Urticaceae) (SING67093).

Lectotypification of Elatostema subscabrum

A. neccsrecsc ature, Ssergn.

both —

a P

Gene, AUctrdabntus

: kL ShcAer

‘*

Was. ter Fifide Schroter

Figure 2. Lectotype (upper left) and isolectotype (lower) of Elatostema subscabrum

H.Schroet. (Urticaceae) (SING s.n.).

47

48 Gard. Bull. Singapore 62 (1) 2010

Table 1. Diagnostic morphological features distinguishing Elatostema subscabrum from

E. sessile.

Character E. subscabrum E. sessile

Appearing ‘hairy’ because Not appearing ‘hairy’

Female of very long, narrowly because of shorter

inflorescences ovate, tapering, unequal bracteoles not extending

bracteoles the involucral bracts

4; unequal size, one of

which is longer with a long

appendage

3; unequal, without

appendage

Tepals of female

flower

Female flower tepal

Ree Distinct and readily visible | Minute, difficult to observe

Elatostema sessile J.R.Forst. & G.Forst.

Characteres Generum Plantarum 53, n. 2 (1775); ed. 2 (1776) 106.

Terrestrial herb, 0.3-0.5 m high, self-supporting (erect/suberect); internodes

developed (elongate, distinct); branched hairs lacking; stinging hairs absent;

monoecious. Stipules caducous. Leaves opposite, appearing alternate (by

misinterpretation — nanophylls caducous); sessile (or petiole < 2 mm long);

megaphylls (45-)50-135(-155) mm long, (21-)22-55(-64) mm wide (length

to width ratio (2.1-)2.2-2.5), unequal-sided, larger side of lamina ovate to

elliptic, smaller side as for larger side or to slightly obovate; surface smooth,

not rugose, with 4 or 5 vein-pairs; venation asymmetric, actinodromous;

basal pair of secondary veins arising above base of primary vein and arising

from one point (or less than 2 mm apart), both directed towards apex (or

almost so), joined to next distal secondary vein; abaxial surface with linear

cystoliths on interstices and hairs on primary, secondary and tertiary veins;

adaxial surface with linear cystoliths interstices and on primary, secondary

and tertiary veins, hairs occasionally present on interstices; base oblique,

rounded or cuneate; margin toothed, not lobed, sparsely hairy; apex

acuminate; nanophylls absent. Inflorescences unisexual; male inflorescences

(based on Weddell, 1869; not known by current authors) sessile or shortly

pedunculate, depressed-globose, involucral bracts present, broadly

ovate, hairy on margin; female inflorescence sessile or sometimes shortly

pedunculate, unbranched, discoid; involucral bracts present, with margin

hairy, with long appendage; bracteoles linear-spathulate, ciliate; flowers

Lectotypification of Elatostema subscabrum 49

condensed/crowded, unisexual: actinomorphic (or slightly asymetrical):

tepals 3, very minute (appearing absent), unequal, free, appendage absent:

staminodes 3, inflexed in bud; ovary straight: style absent: stigma oblong.

filiform to linear. Achene 0.4-0.63 x 0.25-0.35 mm not enclosed (or only

partly so); surface smooth or ribbed.

Selected specimens examined: MALAYSIA. Perak: Rotan Segar limestone

Hill near Tambun, 29 Nov 1960, Allen 4652 (SING): Maxwell’s Hill. 7 Dec

1965, Shah & Sidek 1146 (SING): Ah Kee Iron Mine, Ipoh, 21 Oct 1958,

Sinclair 9890 (SING). INDONESIA. Sumatera Utara: Kabupaten Karo, Air

terjun Sikulikap, Desa Mejuah-juah, 21 Jun 2001 Hadiah, Conn & Ariyanti

453 (NSW): Jawa Barat: Gunung Gede-Pangrango National Park, track

to air terjun Cibeureum, 25 Aug 1998, Hadiah 148 (NSW): Bogor, Wana

Wisata Curug Nangka, Curug Sawer, Warung Loa, 29 Sep 1998, Hadiah 253

(NSW).

Distribution: Malaysia. Malay Peninsula: Perak, Perlis, Pahang and Kelantan.

Indonesia. Sumatera: Sumatera Utara, Sumatera Barat and Jambi; Jawa:

Jawa Barat, Jawa Tengah and Jawa Timur: Bali.

Acknowledgements

This research project was supported by two Singapore Botanic Gardens

Research Fellowships and one of us (BJC) was also a recipient of a Royal

Botanic Gardens Sydney Friends’ Science Scholarship. We gratefully

acknowledge the generous support of these two organisations.

References

Conn, B.J. and J.T. Hadiah. 2009. Nomenclature of tribes within the

Urticaceae. Kew Bulletin 64: 349-352.

Florence, J. 1997. Flore de la Polynesiae francaise. Volume 1. ORSTROM,

Paris.

Forster. J.R. and G. Forster. 1775. 53. Elatostema. Characteres generum

plantarum ... B. White, T. Cadell & P. Elmsly, London.

Friis, I. 1993. Urticaceae, pp. 612-63. In: Kubitzki, K., J.G. Rohwer, and V.

Bittrich (eds.). The families and genera of vascular plants II, flowering

plants — dicotyledones. Magnoliid, hamamelid and caryophyllid families.

Springer-Verlag, Berlin.

50 Gard. Bull. Singapore 62 (1) 2010

Hadiah, J.T. and B.J. Conn. 2009. Usefulness of morphological characters for

infrageneric classification of Elatostema (Urticaceae). Blumea 54: 181-

191.

Hadiah, J.T., B.J. Conn and C.J. Quinn. 2008. Infra-familial phylogeny of

Urticaceae, using chloroplast sequence data. Australian Systematic

Botany 21: 375-385.

Hadiah, J.T., C.J. Quinn and BJ. Conn. 2003. Phylogeny of Elatostema

(Urticaceae) using chloroplast DNA data. Telopea 10: 235-246.

McNeill, J., FR. Barrie, H.M. Burdet, V. Demoulin, D.L. Hawksworth, K.

Marhold, D.H. Nicolson, J. Prado, P.C. Silva, J.E. Skog, J.H. Wiersema

and N.J.Turland. 2006. International Code of Botanical Nomenclature

(VIENNA CODE). 2006. Regnum Vegetabile 146. ARG Gantner Verlag,

Ruggell, Liechtenstein. (http://ibot.sav.sk/icbn/main.htm; viewed 25 Oct

2009).

Nicolson, D.H. and E.R. Fosberg. 2003. The Forsters and the botany of the

second Cook expedition. A.R.G. Gantner-Verlag, Ruggell, Leichenstein.

Robinson, C.B. 1910. Philippine Urticaceae. The Philippine Journal of

Science 5: 465-542.

Robinson, C.B. 1911. Philippine Urticaceae II. The Philippine Journal of

Science 6(4): 299-314.

Schroter, H. and H. Winkler. 1935. Monographie der gattung Elatostema

s.l: Allgemeiner teil. Repertorium specierum novarum regni vegetabilis

83(1): 1-71, figs 1-44.

Schroter, H. and H. Winkler. 1936. Monographie der gattung Elatostema s.L:

Spezieller teil. Repertorium specierum novarum regni vegetabilis 83(2):

1-237, figs 9-40.

Sytsma, K. J., J. Morawetz, J. C. Pires, M. Nepokroeff, E. Conti, M. Zjhra, J.

C. Hall and M. W. Chase. 2002. Urticalean rosids: circumscription, rosid

ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences.

American Journal of Botany 89(9): 1531-1546.

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Weddell, H.A. 1856. Monographie de la famille des Urticées. Nouvelle

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=)

Gardens’ Bulletin Singapore 62 (1): 53-103. 2010

Nn

Oo

Lejeuneaceae subfamily Ptychanthoideae (Hepaticae)

in West Java

I HAERIDA ,S.R.GRADSTEIN AND S. S. TJITROSOEDIRDJO~

‘ Herbarium Bogoriense, Botany Division, Research Center for Biology

Indonesian Institute of Sciences, Cibinong Science Center

: Jl. Raya Jakarta Bogor Km. 46, Cibinong, Bogor

Department of Systematic Botany, Institute of Plant Sciences, University of

_ Gottingen, Untere Karspiile 2, 37073 Gottingen, Germany

* Department of Biology, Faculty of Science and Mathematics

Bogor Agriculture University, Bogor, and

South East Asian Regional Center for Tropical Biology (SEAMEO BIOTROP)

P.O. Box 116, Bogor, Indonesia

Abstract

The subfamily Ptychanthoideae of the family Lejeuneaceae (Hepaticae)

in West Java is still poorly known. A study of Ptychanthoideae in this

area reveals the occurrence of 26 species, in 8 genera: Acrolejeunea

(Spruce) Schiffn. (3 species), Archilejeunea (Spruce) Schiffn. (1 species),

Lopholejeunea (Spruce) Schiffn. (10 species), Mastigolejeunea (Spruce)

Schiffn. (3 species), Ptychanthus Nees (1 species), Schiffneriolejeunea Verd.

(2 species), Spruceanthus Verd. (2 species) and Thysananthus Lindenb.

(4 species). Mastigolejeunea indica and Thysananthus minor are newly

reported for Java. The altitudinal and geographical ranges of the species

of Ptychanthoideae from West Java are presented as well as a key to the

species and an assessment of their phytogeography. It appears that the

species are rather widespread; none of the species are endemic to Java or

western Malesia. The widespread distribution of the species is probably due

to their dispersal by spores, which may be easily carried by the wind over

long distances, and by the rather old geological age of the group.

Introduction

Lejeuneaceae is a large, mostly tropical family of leafy liverworts,

containing about 1000 species in 75 genera. According to Gradstein et al.

(2001), Lejeuneaceae 1s subdivided into two subfamilies, Leyjeuneoideae and

Ptychanthoideae; the latter group is treated in this paper.

Ptychantheaeis the only tribe of Ptychanthoideae, while Lejeuneoideae

consists of three tribes, Brachiolejeuneeae, Lejeuneeae and Cololejeuneeae.

54 Gard. Bull. Singapore 62 (1) 2010

The subfamily Ptychanthoideae is characterized by the capsule valves

spreading widely after dehiscence, elaters usually 72 per capsule, underleaves

undivided, ventral merophytes 4 or more cells wide, and ocelli absent. In

Lejeuneoideae the capsule valves are not spreading after dehiscence, elaters

are 34 or less per capsule, underleaves bifid or undivided, ventral merophytes

usually 1-4 cells wide, and ocelli sometimes present.

Ptychanthoideae include fewer genera and species than Lejeuneoideae

and are generally better known (Gradstein, 1991). They contain about 175

species in 19 genera worldwide; 14 genera occur in tropical America. Most

species are xerotolerant epiphytes of forest canopies or rather open, wooded

vegetation, and usually occur at rather low elevation (Thiers and Gradstein,

1989; Gradstein et al., 2001).

Based on several references (Verdoorn, 1933, 1934; Mizutani, 1961;

Gradstein, 1975, 1991, 1994; Gradstein et al. 2002; Zhu and Gradstein, 2005;

Gradstein et al., 2005; Hasan and Ariyanti, 2004; Ariyanti and Gradstein,

2007), there are about 30 species of Ptychanthoideae in Java, in 10

_genera: Acrolejeunea (Spruce) Schiffn, Archilejeunea (Spruce) Schiffn.,

Caudalejeunea _(Steph.) Schiffn., Dendrolejeunea (Spruce) Lacout.,

Lopholejeunea (Spruce) Schiffn., Mastigolejeunea (Spruce) Schiffn.,

Ptychanthus Nees, Schiffneriolejeunea Verd., Spruceanthus Verd., and

Thysananthus Lindenb. In addition, there is an old, unconfirmed report of

the Pacific genus, Phaeolejeunea Mizut., from Java (Zhu and Gradstein, 2005,

p. 88, originally described as Lopholejeunea latistipula (Schiffn.) Schiffn. var.

minor Schiffn.).

Information about the distribution of the species of Ptychanthoideae

within Java is very scarce. The publications on this subject are mostly from

the 19° or early 20° century and outdated (e.g., Reinwardt, Blume and

Nees von Esenbeck, 1824; Nees von Esenbeck, 1830; Sande Lacoste, 1856;

Schiffner, 1898; Stephani, 1912; Verdoorn, 1933, 1934).

The aim of this study was to improve our understanding of the diversity

of Ptychanthoideae in West Java (including West Java Province and Banten

Province).

Material and Methods

The study was based on herbarium specimens of Ptychanthoideae collected

from West Java and Banten province. A total number of 362 specimens

deposited in the Herbarium Bogoriense (BO) and Herbarium Biotrop

(BIOT), and some in the Herbarium of the University of Géttingen,

Germany (GOET) and Herbarium Bangi, Malaysia (UKMB) kept in BIOT

as duplicates, were examined. Additional specimens were collected by the

in

Nn

Lejeuneaceae subfamily Prychanthoideae

first author at several locations in West Java, i.e. Telaga Warna, Ciater, Bogor

Botanical Garden, Bodogol Education Center and Conservation — Mt. Gede

Pangrango National Park and Mt. Halimun Salak National Park.

Morphological characters were studied with a 10 x 4,10 x 10,10 x 20 and

10 x 40 magnification. Specimens with similar characteristic were grouped

and identified using recent literature on the species of Ptychanthoideae,

such as Thiers and Gradstein (1989), Gradstein er al. (2001), Gradstein et

al. (2002) and Zhu and Gradstein (2005). Descriptions and illustrations of

each species were prepared and an identification key to the species was

developed.

Taxonomic treatment

Key to the species of West Javan Ptychanthoideae

1. Branching always Frullania type.......................18. Ptychanthus striatus

Betseancnins Mejeuned OF Friullania ty Pe......:......0.....0aessscssecssssnecsecesese «iene oe: 2

2. Branching Frullania and Lejeunea type (Schiffneriolejeunea)................3

PEM AME MEM ILCICUIMCE-LYPC ONY. 2. 55.2.5. 25 058. tects coe owe ee cceseecteesstoccetens -

3. Lobule forming a distinct sac at the base and flattened above with 2 large

teeth. Teeth erect, not pointing outwards towards leaf apex. Leaves

squarrose when moist.. 20. Schiffneriolejeunea tumida var. haskarliana

3. Lobule not forming a distinct sac at the base, free margin of the lobule

plane and clearly visible, with 2 teeth. Teeth almost equal in size and

pointing outwards towards leaf apex. Leaves not squarrose when moist

Pe BSS s sheds Sn toca avaiees 19. Schiffneriolejeunea pulopenangensis

4. Lobes with a distinct vitta ...........................25. Thysananthus retusus

2 DE EC ee ae eee 5

Pee cls Orthe LODE With COLMAtE ITIDONES.... ......... ..eerecseseerodeececeeeoeeroveeO

Pecusorte lobe not with cordate trigONes. ...... 15.20.00. sce0e soe eeeeseeeees 14

6. Leaves and underleaves entire. Dorsal epidermis cells larger than inner

DEE FELD 2 2S See ee nee een erry aera eee |

6. Leaves and underleaves usually toothed. Dorsal epidermis cells not larger

pam eECe Stem CEUS( PH YSAVIGIINUS) - .-< <cc-ncetwennnsenconanracedvewsieenscsens 12

Lee

7

Gard. Bull. Singapore 62 (1) 2010

. Perianth 3 keels, with 1-2 innovations (Mastigolejeuned).......ccccccescceneen 8

. Perianth 8-10 keels, innovation lacking (Acrolejeunea)...................... 10

; Leaf lobule with several large; triangular tecthis2eee eee

Fe er Pore eet eet eet Mee A Pete 5A 16. Mastigolejeunea indica

Leaf lobule with one tooth). #c.cge sec: aneks ce eee eee eee eee eee 9

. Lobule tooth 3-5 cells long, sharp, curved....... 17. Mastigolejeunea virens

» Lobule tooth Icell long; blumtynot cukved irc eee eee ee

cabal steve oth Se Sah sd Rupees eetes Meta. cte eee meee 15. Mastigolejeunea auriculata

. Lobule semicircular in outline, with 4-10 teeth, all or at least some teeth

23 Cells 1ONGwee eee ae cde eee ee 2. Acrolejeunea fertilis

-Lobule not semicircular, withi2-siteethi 4.2: ea

. Lobule narrow rectangular, with 2 teeth ......... 1. Acrolejeunea arcuata

. Lobule ovate-rectangular, with 3 teeth ......3. Acrolejeunea pycnoclada

. Plant very small, less than 1 mm wide .............. 24. Thysananthus minor

/ Plant more than fmm wide .2.4.<2.52..-04 eee eee ee 13

. Upper part of leaf asymmetrical, turned to the ventral side (“ventrad”’),

leat apex recive secre. eee eee 23. Thysananthus convolutus

. Upper part of leaf symmetrical, not turned to the ventral side, leaf apex

planeionmecuny edie a. eee 26. Thysananthus spathulistipus

. Stems robust, ventral merophyte 8-12 cells wide (Spruceanthus)........15

. stems' thinner, ventral metrophyte 4-6 cells widen ce. -ee eee ee 16

» Keatapexishanply;actitew:--.-ee.s ee 22. Spruceanthus semirepandus

; beat apexsrounded (oo btusem=e = 21. Spruceanthus polymorphus

. Stem epidermis cells not larger than the inner cells. Plants light green.

Perianth keels smooth or weakly toothed. Innovations present..............

sett osieudeuberubedaenies eeeanee eee ete ce eee eee 4. Archilejeunea planiuscula

. Stem epidermis cells larger than the inner cells. Plants blackish-green,

rarely brown. Perianth keels distinctly toothed (Lopholejeunea).

Innovations lackim pst: -sec.eces-s eeneree ete eee 17

Leaf lobule connected to the leaf lobe by only one single cell............18

Leaf lobule connected to the leaf lobe across several cells.................21

Lejeuneaceae subfamily Ptychanthoideae 57

22:

24.

24.

25.

. Leaf lobe acute .....00.0. 0.0). Lopholejeunea applanata

BEcaAnOuE LOUnGCEd OF ODIUSE.....25:22.2-..edeeten se 0b edoncse ce cWecee te dedeeee 19

. Lobule deeply constricted in the middle, free margin strongly involute

ee ec asin oe pds ene cove sue ncvtendcassaves sade 7. Lopholejeunea herzogiana

. Lobule not deeply constricted in the middle, free margin not involute.....20

. Underleaves distant to contiguous, orbicular. Female bracteole entire

Seno ehecnp scan Sancnstbesbnaitcsuhonssonion 8. Lopholejeunea nigricans

. Underleaves imbricate, wider than long. Female bracteole toothed........

-c sserseceeec oe eee re 6. Lopholejeunea eulopha

. Underleaves very large, 6-8 x as wide as the stem ..... ey

meacericaves smaller 4-6 x as wide as the Stem ............2......:.s000000.- 23

. Plants pale brown in the dried condition. Leaf lobules small, 1/5-1/4 of

feat leapth Female bracteole toothed. Dioicous ..........2.........<.ccessessecessssees

Se cles Soha dnc vas onedncvnsnasneve’ 13. Lopholejeunea wiltensii

Plants blackish or dark brown to blackish in the dried condition. Leaf

lobules larger, 1/3-2/5 of leaf length. Female bracteole usually entire.

8 ICDS TE ae Be ee 14. Lopholejeunea zollingeri

3. Plants minute, less than 1 mm wide............. 10. Lopholejeunea horticola

miaaes medium sized, more than 1 mm wide... ......-..-...2..0.0..se0ssseee8 24

Margins of underleaves and leaves recurved. Underleaf large, more

eee ee ROD EMCH PLN: «ci. 25. c.cccveasesessceeeseco = 28 11. Lopholejeunea recurvata

Margins of underleaves and leaves plane. Underleaf less than % x

C2 DE EOD ae scence sp ere ee oe Ay 2

. Female bracteole entire. Female bract lobule very small or lacking

ses ne neecececiese Coane Oe 12. Lopholejeunea subfusca

Female bracteole crenate. Female bract lobule large... ee

se saghiss nda se ec Ocae 9. Lopholejeunea ceylanica

Species descriptions

1. Acrolejeunea arcuata (Nees) Grolle & Gradst., J. Hattori Bot. Lab. 38: 332

(1974).

Jungermannia arcuata Nees, Enum. Pl. Crypt. Javae 1:38 (1830). Ptychocoleus

arcuatus (Nees) Trevis., Mem. Reale Ist. Lomb. Sci. Mat. Nat., Ser. 3, 4: 405

58 Gard. Bull. Singapore 62 (1) 2010

(1877). -Type: Indonesia. Java: Lebak Mts., Reinwardt s.n. (holotype, STR;

isotypes, G, NY, W) — cf. Gradstein et al. (2002).

Figure 1. Acrolejeunea arcuata (Nees) Grolle & Gradst. Shoot (a); leaf-lobe (b); lobule (c); cells of

midleaf (d); underleaf (e). Drawn from Kornochalert 1408, BIOT.

Plants up to 2 cm long, about 0.9 mm wide, dark brown to reddish brown in

dried condition. Branching Lejeunea-type. Stem diameter 95-108 um. Leaves

closely imbricate, obliquely spreading. Lobe ovate-orbicular to obovate, 520-

650 um long, 410-520 um wide, margin entire, apex subobtuse to sometimes

rounded; cells of lobe thin-walled, hyaline, somewhat elongate, trigones +

cordate, intermediate thickening scarce; marginal cells 5-11 x 6-11 um, mid-

leaf cells 19-29 x 11-26 um, basal cells 24-30 x 10-13 um; oil bodies not seen.

Lobule narrow rectangular, 300-400 um long, 145-160 um wide, apex truncate

with 2 teeth, each tooth consisting of 2-3 cells, 2-3 cells long. Underleaves

closely imbricate, subtriangular to suborbicular, 260-315 tm long, 295-435

um wide, margins entire, apex truncate, base slightly auriculate, insertion

line shallowly curved. Generative structures not seen.

Distribution: Java, Sumatra, Borneo, Philippines, Papua, Peninsular Malaysia.

Specimen examined: INDONESIA. West Java, Mt. Patuha, on bark of tree

along the road to crater, 2000 m, Kornochalert 1408 (BIOT!).

Notes: Diagnostic characters of Acrolejeunea arcuata are: 1) lobules 2(-3)

times longer than wide, with 2 teeth, 2) lobe ovate-orbicular with entire

margins, 3) cordate trigones. This species was found at ca 2000 m and occurs

only at higher elevations in the mountains (Gradstein, 1975). It is readily

in

\O

Lejeuneaceae subfamily Prychanthoideae

distinguished from other species of Acrolejeunea by the very long and

narrow lobule with 2 teeth and the often reddish-brown color.

2. Acrolejeunea fertilis (Reinw. et al.) Schiffn., in Engler and Prantl, Nat.

Pflanzenfam. 1, 3: 128 (1893); Jungermannia fertilis Reinw. et al., Nova Acta

Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 211 (1824): Ptychocoleus

fertilis (Reinw. et al.) Trevis., Mem. Reale Ist. Lomb.Sci. Mat. Nat., Ser. 3, 4:

405 (1877). —Type: Indonesia. Java, Lebak Mts., Reinwardt s.n. (holotype,

STR; isotypes, G, NY, W) — cf. Gradstein et al. (2002).

Figure 2. Acrolejeunea fertilis (Reinw. et al.) Schiffn. Shoot (a): leaf-lobe (b); underleaf (c): cells of

midleaf (d); cells of basal part of the leaf (e); Drawn from van Borssum Waalkes 549, BO.

Plants up to | cm long, about 0.8 mm wide, dark green to dark brown in

dried condition. Branching Lejeunea-type. Stem diameter 100-124 um.

Leaves closely imbricate, obliquely spreading, squarrose. Lobe ovate-

orbicular to subrectangular, 600-825 um long, 500-660 um wide, margin

entire, apex truncate to sometimes rounded; cells of lobe thin-walled, light-

yellow, somewhat elongate, trigones cordate, intermediate thickening scarce;

marginal cells 7-8 x 7-10 um, mid-leaf cells 18-23 x 12-20 um, basal cells 17-30

x 13-17 um; oil bodies not seen. Lobule semicircular, 200-290 um long, 220-

250 um wide, apex obliquely truncate with 4-9 teeth, each tooth consisting of

2-3 cells, 1-2 cells long. Underleaves closely imbricate, broadly orbicular, 300-

350 um long, 310-570 um wide, margins entire, apex truncate, base cuneate,

insertion line shallowly curved. Generative structures not seen.

Specimens examined: INDONESIA. Banten, Pulau Panaitan, van Borssum

Waalkes 549 (BO!). West Java, Depok, Kampus Universitas Indonesia, 60-

80 m, Afiatri Putika 80 (BO!).

60 Gard. Bull. Singapore 62 (1) 2010

Distribution: Java,Sumatra, Borneo, Philippines, Moluccas, Papua, Peninsular

Malaysia, Indochina, India, Sri Lanka.

Notes: Diagnostic characters of Acrolejeunea fertilis are: 1) lobule

semicircular with 4-9 teeth, 2) lobe ovate-orbicular with entire margins, 3)

trigones cordate. The species occurs in the lowlands, at ca 60-80 m, and is

distinguished from other Javanese members of the genus Acrolejeunea by

the squarrose leaves and the semicircular lobule with 4-9 teeth.

3. Acrolejeunea pycnoclada (Taylor) Schiffn., in Engler and Prantl, Nat.

Pflanzenfam. 1, 3: 128 (1893); Ptychanthus pycnocladus Taylor, London J.

Bot. 5: 385 (1846); Ptychocoleus pycnocladus (Taylor) Steph., Sp. Hepat. 5:

52 (1912). -Type: Peninsular Malaysia, without loc., Cantor s.n. (holotype,

FH; isotypes, G, MANCH, NY, S, U, W) — cf. Gradstein et al. (2002).

Figure 3. Acrolejeunea pycnoclada; (Taylor) Schiffn. Shoot (a); leaf lobe (b); leaf lobule (c):

underleaf (d); leaf cells (e); female bract (f); female bracteole (g); perianth (h). Drawn from Haerida

1484, BO.

Autoicous. Plants up to 2 cm long, 0.5-0.7 mm wide, 1-1.5 mm wide in

wet condition; dull green when fresh, greenish-brown in dried condition.

Branching Lejeunea-type. Stem diameter 100-145 um; cross-section of the

stem with epidermis cells larger than medulla cells Leaves imbricate, widely

spreading, convolute when dry. Lobe ovate, 680- 1050 um long, 370-850 um

wide, margin entire, apex rounded; cells of lobe thin-walled, hyaline, in mid-

Lejeuneaceae subfamily Ptychanthoideae 61

leaf elongate, trigones cordate, intermediate thickening frequent; marginal

cells 7-16 x 6-13 um, mid-leaf cells 17-31 x 11-25 um, basal cells 24-52 x

11- 31 um; oil bodies (7-)8-20 per cell, homogeneous, hyaline, globose to

ellipsoidal. Lobule ovate to rectangular, large, 365-680 um long, 170-310 um

wide, inflated, apex obliquely truncate with 2 conspicuous teeth, first tooth

consisting of 3-4 cells, second tooth consisting of 2-3 cells, an additional

small tooth present at the distal end of the free margin beyond the first

tooth. Underleaves distant to contiguous, obovate-orbicular to wider than

long, 270-480 um long, 350-700 um wide, margin entire, apex rounded to

almost truncate, base cuneate, insertion line shallowly curved. Androecia

intercalary on branches, bracts in 3-11 pairs, very similar to leaves but

lobules slightly larger, 400-420 um long, 180-250 um wide, bract lobule 1/2-

2/3 of the bract lobe, epistatic, bracteoles similar to underleaves in size and

shape. Gynoecia on short or long branches, innovations lacking, bracts and

bracteoles in 2 pairs, bract oblong to broadly oblong 720-1070 um long,

270-500 um wide, conspicuously incurved in the upper part, margin entire,

apex obtuse to orbicular, sinus up to 1/3 of lobe length; bracteole oblong to

broadly oblong 890-1010 um long, 520-660 um wide, margin entire to slightly

undulate, apex truncate. Perianth immersed, obovate, 975 um long, 430 um

wide, inflated, with + 10 inflated keels, margins entire. Sporophytes and

asexual reproduction not seen.

Distribution: Java, Sumatra, Borneo, Moluccas, Philippines, Papua New

Guinea, Pacific, Peninsular Malaysia, Thailand, India, Sri Lanka, tropical

Africa.

Specimens examined: INDONESIA. West Java, G. Gede, above Sukabumi,

1500-1900 m, Verdoorn 46b (BO!); Ciater, Subang, Sari Ater, ca 1500 m,

Haerida 1480, 1481, 1484, 1486, 1487 (BO!).

Notes: Diagnostic characters of Acrolejeunea pycnoclada are: 1) lobule with

2 conspicuous teeth and an additional, third small tooth present at the distal

end of the free margin, beyond the first tooth, 2) trigones cordate, 3) female

bracts conspicuously incurved in the upper part, covering the immersed

perianth, 4) perianth with 10 keels. This species was found at ca 1500-1900

m but it has also been recorded at lower elevations, from sealevel upwards.

It differs from other members of the genus Acrolejeunea by the lobule

with 2 conspicuous teeth and a 3° small tooth at the very end of the free

margin, near the junction with the ventral margin of the leaf lobe, and by the

conspicuously incurved female bracts.

4. Archilejeunea planiuscula (Mitt.) Steph., Sp. Hepat. 4: 731 (1911);

Lejeunea planiuscula Mitt., J. Proc. Lin.. Soc. 5: 111 (1861). -Type: Burma.

62 Gard. Bull. Singapore 62 (1) 2010

Rangoon, McClelland s.n. (holotype, NY) — cf. Thiers and Gradstein (1989).

- Archilejeunea caramuensis Steph., Hedwigia 34: 59 (1895). —Type:

Philippines.Caramuan, 1884-85, Micholitz 70 (G) — cf. Verdoorn (1934).

- Archilejeunea falcata Steph., Hedwigia 34: 60 (1895). —-Type: Papua New

Guinea. Madang Prov.: Stephansort, 1888, Kdrnbach s.n. (G) — cf. Verdoorn

(1934).

- Mastigolejeunea paradoxaVerd.,Nova Guinea 18:5 (1935).—Type: Indonesia.

West Irian, Jayawijaya Prov., Prauwenbivak, Lam 1182a (holotype, FH) — cf.

Gradstein et al. (2002).

Figure 4. Archilejeunea planiuscula (Mitt.) Steph. Shoot (a); leaf lobule (b); underleaf (c); cells of

leaf margin (d); cells of midleaf (e). Drawn from Haerida 1489, BO.

Plants up to 3 cm long, 0.8-2 mm wide; light green when fresh, darker green

in dried condition. Branching Lejewnea-type. Stem diameter 90-135 um;

epidermis cells not large than inner cells; ventral merophyte 4-6 cells wide.

Leaves contiguous, widely spreading. Lobe orbicular to oblong, 500-900

um long, 370-700 um wide, margin entire, apex rounded; cells of lobe thin-

walled, hyaline, isodiametrical hexagonal, trigones triangular, never cordate,

intermediate thickening scarce; marginal cells 3.5-7.5 x 5-10 tm, mid-leaf

cells 17-24 x 14-18 um, basal cells 22-37 x 13-17 um; oil bodies not seen.

Lobule ovate, 240-340 um long, 130-200 um wide, apex obliquely truncate

with 1-2 teeth, first tooth with 2 cells long, second tooth small, 1 cell long,

sometimes lacking. Underleaves distant to contiguous, obdeltoid, 280-420

um long, 260-460 um wide, margin entire, apex truncate, insertion line almost

straight. Generative structures not seen [innovations present; perianth with

5 smooth or weakly toothed keels].

Lejeuneaceae subfamily Prychanthoideae 63

Specimens examined: INDONESIA. West Java, Bogor Bot. Garden,

Verdoorn 12a, 12b, 12c, 12e, 131, 134, 232, Meijer B55a, Bola, B49, B76,

B3664, B27a, B56, BS4a (BO!); Cibodas Bot. Garden, 1450 m, Schiffner

233, Haerida 813 (BO!); Ujung Kulon Nat. Park, Dewi Dw922 (BO!).

Distribution: Java, Peninsular Malaysia, Papua New Guinea, Philippines,

Solomon Is., Pacific, Australia, Indochina, India, Sri Lanka.

Notes: Diagnostic characters of Archilejeunea planiuscula are: 1) underleaves

obdeltoid, 2) trigones triangular, never cordate, 3) plant light green in color

when fresh. This species was found at ca 200-1450 m. It is readily distinguished

from other members of the subfamily by the rather light green color and

flat appearance of the plants, lobules with 1-2 teeth, isodiametrical leaf cells

with small, simple trigones, obdeltoid underleaves and thin stems (ventral

merophyte 4-6 cells wide) with epidermis cells not larger than inner cells.

5. Lopholejeunea applanata (Reinw. et al.) Schiffn., in Engler and Prantl,

Nat. Pflanzenfam. 1, 3: 129 (1893); Jungermannia applanata Reinw. et al.

Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 210 (1824):

Phragmicoma applanata (Reinw. et al.) Nees, Naturgesch. Eur. Leberm.

3: 248 (1838); Lejeunea applanata (Reinw. et al.) Nees, in Gottsche et al.,

Syn. Hepat.: 314 (1845): Symbiezidium applanatum (Reinw. et al.) Trevis.,

Mem. Reale Ist. Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type:

Indonesia. Java, collector unknown (holotype, STR: isotypes, G, S, W) — cf.

Zhu and Gradstein (2005).

- Lopholejeunea fleischeri Steph., Sp. Hepat. 5: 79. 1912. —Type: Indonesia.

Java, Tjibodas, Apr 1900, M. Fleischer s.n. (holotype, G) — cf. Zhu and

Gradstein (2005).

Plants up to 5 cm long, 1-1.5 mm wide; dark brown in the dried condition.

Branching Lejeunea-type. Stem diameter 120-200 um Leaves imbricate,

widely spreading. Lobe ovate, 800-1200 um long, 500-800 um wide, margin

entire sometimes crenulate towards the apex, apex acute to acuminate: cells

of lobe thick-walled, yellow, rectangular to isodiametric, trigones triangular,

intermediate thickening frequent; marginal cells 15-21 x 12-15 um, mid-leaf

cells 26-30 x 20-24 um, basal cells 30-40 x 20-31 um: oil bodies not seen.

Lobule ovate, 230-320 um long, 170-220 um wide, inflated, apex truncate

with 1 small tooth, connected to the leaf lobe across only one single cell.

Underleaves large, imbricate, broadly orbicular, 300-400 um long, 400-

900 um wide, margin entire, apex truncate, insertion line deeply curved.

Androecia intercalary on branches, bracts in 3-12 pairs, ovate with obtuse

apex, 200-300 um long, 150-220 um wide, bract lobule about % of the lobe

64 Gard. Bull. Singapore 62 (1) 2010

length; bracteoles similar to underleaves in size and shape. Gynoecia not

seen.

20 Um

Figure 5. Lopholejeunea applanata (Reinw. et al.) Schiffn. Shoot (a); underleaf (b); leaf lobule

(c); leaf lobe (d); cells of midleaf (e). Drawn from Meijer 3696, BO.

Distribution: Java, Sumatra, Borneo, Philippines, Peninsular Malaysia, Papua

New Guinea, Pacific, Indochina, Sri Lanka, India.

Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden,

Meijer B3696 (BO!); above Cibodas, Cibeureum, Hasskarl s.n. (BO!); G.

Pangrango, “bij Tjisaroea” ca 1200 m, Verdoorn 44b (BO!); G. Patuha, Lake

Situ Patengan, ca 1500 m, Kornochalert 1413 (BIOT!).

Notes: Diagnostic characters of Lopholejeunea applanata are: 1) leaf lobes

acute to acuminate at apex, 2) lobule apex connected to the lobe across only

one single cell. This species grows at ca 1200-1500 m. It differs from other

Javanese members of the genus Lopholejeunea by the acute to acuminate

leaf apex and large, imbricate underleaves.

6. Lopholejeunea eulopha (Taylor) Schiffn., in Engler and Prantl, Nat.

Pflanzenfam. 1,3: 129 (1893); Lejeunea eulopha Taylor, London J. Bot. 5: 391

(1846); Phragmicoma eulopha (Taylor) Mitt.,in Seeman, FI. Vit. 413 (1873);

Symbiezidium eulophum (Taylor) Trevis., Mem. Reale Ist. Lombardo Sci.,

Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type: Pacific islands: locality unknown,

in

Lejeuneaceae subfamily Ptychanthoideae 6

Nightingale s.n. (holotype, FH; isotypes, FH. NY) — cf. Zhu and Gradstein

(2005).

- Lopholejeunea nymannii Steph., Sp. Hepat. 5: 84. 1912. -Type: Indonesia.

Java, G. Salak, E. Nyman s.n. (holotype, G;isotype, W) —cf. Zhu and Gradstein

2005.

a

fi

a Se

Figure 6. Lopholejeunea eulopha (Taylor) Schiffn. Shoot (a): leaf lobule (b); leaf lobe (c): cells of

midleaf (d). Drawn from Haerida 710, BO.

Dioicous, sometimes autoicous. Plants up to 3 cm long, 1-1.5 mm wide,

brownish green to dark brown in dried condition. Branching Lejeuwnea-type.

Stem diameter 130-190 um. Leaves imbricate, widely spreading. Lobe oblong-

ovate, 430-1090 um long, 340-900 um wide, margin entire, apex obtuse; cells

of lobe thin-walled, pale brown, hexagonal to nearly isodiametric, trigones

triangular, intermediate thickening frequent; marginal cells 7-15 x 10-17 um,

mid-leaf cells 14-31 x 10-21 um, basal cells 22-40 x 18-24 um; oil bodies

(11-)18-20 per cell, homogeneous, hyaline, ellipsoidal. Lobule ovate, 120-270

um long, 110-200 um wide, inflated, apex with | triangular tooth consisting

of 3-5 cells, connected to the leaf lobe by only one single cell. Underleaves

imbricate, large, reniform, 300-500 um long, 410-1070 um wide, margin entire,

apex rounded, insertion line deeply curved. Generative structures not seen.

Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer

94b, Schiffner 245, van Steenis s.n. (BO!); Bogor, without locality, Verdoorn

s.n. (BO!); Cibodas Bot. Garden, 1400 m. Ariyanti 47] (BIOT!); G. Halimun

Nat. Park, Haerida 710 (BO!); G. Guntur, “bij Kawah Kamoedjan” 1500-

1700 m.alt., Verdoorn 50h, 50d (BO!); G. Megamendung, Verdoorn 68 (BO!):

66 Gard. Bull. Singapore 62 (1) 2010

Kota Batu, Schiffner s.n. (BO!): Kampung Jember, Geger Bentang ca 1350

m, Neervoort 3087 (BO!): Ujung Kulon Nat. Park, Dewi Dw922d (BO!); G.

Patuha, Lake Situ Patengan ca 1500 m, Kornochalert 1418, Dian Apriana &

Afiatri Putrika 25 (BIOT!).

Distribution: Java, Sumatra, Borneo, Bali, Sulawesi, Moluccas, Philippines,

West Irian, Papua New Guinea, Solomon Is., Pacific, Australia, Indochina,

India, Sri Lanka, China, Japan, tropical Africa, tropical America.

Notes: Diagnostic characters of Lopholejeunea eulopha are: 1) lobule ovate,

with | triangular tooth consisting of 3-5 cells,2) lobule apex connected to the

leaf lobe across only one cell, 3) large, reniform underleaves. This common

species grows at ca 200-1700 m. It resembles L. applanata by the very large,

imbricate underleaves but differs by the rounded leaf apex.

7. Lopholejeunea herzogiana Verd., Rec. Trav. Bot. Neerl. 30: 217 (1933). -

- Type: Indonesia. Java, Cibodas, ‘ad arborum truncos,” 1420 m, 1894, Schiffner

s.n. (FH, lectotype designated by Zhu and Gradstein, 2005).

— Lopholejeunea pullei Verd., Nova Guinea 18: 4 (1935). —Type: Indonesia.

Irian Jaya, Hellwig Mts., “an Gymnospermen”, 2600 m, 10 Jan 1913, Pulle

s.n. (holotype, FH: isotype, U) — cf. Zhu and Gradstein 2005.

Figure 7. Lopholejeunea herzogiana Verd. Shoot (a): leaf lobule (b); female bract (c): female

bracteole (d): cells of midleaf (e). Drawn from Verdoorn 64a, BO.

Lejeuneaceae subfamily Prychanthoideae 67

Plants robust, up to 4 cm long, 1-2.5 mm wide: blackish brown in the dried

condition. Branching Lejeuwnea-type. Stem diameter 130-220 um Leaves

imbricate, widely spreading. Lobe obovate-triangular to broadly orbicular,

650-1050 um long, 660-1420 um wide, margin entire, apex obtuse to rounded,

sometimes recurved; cells of lobe thick-walled, brownish-orange, rectangular

to hexagonal, trigones triangular, intermediate thickening frequent; marginal

cells 12-18 x 12-17 um, mid-leaf cells 21-26 x 20-30 um, basal cells 24-50 x

20-30 um; oil bodies not seen. Lobule ovate, large, 300-540 um long, 220-310

um wide, inflated, free margin strongly involuted and with a conspicuous

constriction in the middle, connected to the leaf lobe across only one single

cell. Underleaves closely imbricate, reniform to transversely rectangular,

400-520 um long, 600-1310 um wide, margin entire, apex truncate, insertion

line deeply curved. Androecia not seen. Gynoecia on short or long branches,

innovations lacking, bracts and bracteoles in 1-2 pairs, bract ovate to nearly

rectangular, 2-3 mm long, 1.5-2 mm wide, margin toothed, apex truncate,

sinus up to 1/2 of lobe length, bract lobule ovate, margin ciliate-dentate,

apex truncate to nearly acute; bracteole oblong 1.3-2.2 mm long, 0.5-1.4

mm wide, margin toothed, apex rounded. Perianth immersed, obovate, 550

um long, 530 um wide, inflated, with 5-7 keels, margins widely winged and

toothed. Sporophytes and asexual reproduction not seen.

Specimens examined: INDONESIA. West Java, Telaga Warna, Puncak Pass,

ca 1500 m, Verdoorn 64a (BO!): Cibodas Bot. Garden, Meijer B3800b (BO!):

G. Pangerango, “bij Tjisaroea”, ca 1200 m, Verdoorn 44c (BO!).

Distribution: Java, Peninsular Malaysia, West Irian, Papua New Guinea,

New Caledonia.

Notes: Diagnostic characters of Lopholejeunea herzogiana are: 1) strongly

winged and toothed obovate perianth, 2) lobule conspicuously constricted

in the middle and with strongly involuted free margin, 3) lobule apex

connected to the leaf lobe across only one single cell, 4) large, + reniform

underleaf with deeply curved insertion line. This rare species was found at

ca 1200-1500 m. and usually grows in the mountains at higher elevation. It

is readily separated from other species of the genus by the peculiar lobule

with deep constriction in the middle.

8. Lopholejeunea nigricans (Lindenb.) Schiffn., Consp. Hepat. Arch. Ind.

293 (1898); Lejeunea nigricans Lindenb., in Gottsche et al., Syn. Hepat.

316 (1845): Symbiezidium nigricans (Lindenberg) Trevis., Mem. Reale Ist.

Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877). —Type: Indonesia. Java,

without locality, collector unknown (holotype, W: isotypes. G, S) — cf. Zhu

and Gradstein (2005).

68 Gard. Bull. Singapore 62 (1) 2010

— Lejeunea intermedia Lindenb., in Gottsche et al., Syn. Hepat. 316 (1845);

Symbiezidium intermedium (Lindenberg) Trevis. Mem. Reale Ist.

Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea intermedia

(Lindenberg) Steph., Sp. Hepat. 5:77 (1912). Type: Indonesia. Java, without

locality, collector unkown (holotype, W) — cf. Zhu and Gradstein 2005.

— Lejeunea javanica Nees, in Gottsche et al., Syn. Hepat. 320 (1845);

Symbiezidium javanicum (Nees) Trevis., Mem. Reale Ist. Lombardo Sci., Cl.

Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea javanica (Nees) Schiffn., in

Engler and Prantl, Nat. Pflanzenfam. 1,3: 129 (1893). —Type: Indonesia. Java,

without locality, collector unknown (holotype, W; isotypes, G, S) — cf. Zhu

and Gradstein (2005).

Figure 8. Lopholejeunea nigricans (Lindenb.) Schiffn. Shoot (a): leaf lobule (b); leaf lobe (c);

underleaf (d); female bract (e); cells of midleaf (f); female bracteole (g). Drawn from Meijer 387c, BO.

Plants up to | cm long, 0.5-0.7 mm wide, blackish brown in dried condition.

Branching Lejeunea-type. Stem diameter 40-110 um Leaves imbricate,

widely spreading. Lobe oblong-orbicular, 250-500 um long, 200-400 um

wide, margin entire, apex rounded; cells of lobe thick-walled, dark brown,

isodiametric to hexagonal, trigones triangular, intermediate thickening

scarce; marginal cells 6-10 x 5-10 um, mid-leaf cells 16-22 x 7-20 um, basal

cells 20-28 x 16-20 um; oil bodies not seen. Lobule ovate, 120-230 um long,

80-150 um wide, inflated, apex obliquely truncate, plane, connected to the

leaf lobe across only one single cell. Underleaves distant to contiguous,

obdeltoid to orbicular, 120-200 um long, 150-190 um wide, margin entire,

Lejeuneaceae subfamily Prychanthoideae 69

apex rounded to almost truncate, insertion line shallowly curved. Androecia

not seen. Gynoecia on short or long branches, innovations lacking, bract

oblong. 510-700 pm long, 270-330 um wide, margin entire, apex obtuse to

orbicular, sinus up to 2/3 of lobe length, bract lobule about the same length

as the bract lobe: bracteole obdeltoid to orbicular 330-700 um long, 350-

500 pm wide, margin entire. sometimes recurved, apex broadly rounded.

Perianth exserted, obovate, 300-900 um long, 250-500 um wide. inflated, with

4 keels, margins toothed. Sporophytes and asexual reproduction not seen.

Distribution: Java, Sumatra. Borneo, Sulawesi, Moluccas, West Irian, Papua New

Guinea, Australia, Peninsular Malaysia, Philippines, Indochina, India, Bhutan.

Nepal, China, Japan, tropical Africa, tropical America.

Specimens examined: INDONESIA. West Java, without locality. collector

unknown (BO!): Bogor Bot. Garden, van Borssum Waalkes 122, Meijer

B995, B368, 55d4, Verdoorn 242 (BO!): Telaga Warna, Puncak Pass. 1500 m,

Verdoorn 64d, 64b (BO!): G. Pangrango, “Tjisaroea™, ca 1200 m, Verdoorn

44a (BO!); G. Guntur, “Kawah Kamoedjan”, 1500-1700 m, Verdoorn 50c

(BO!).

Notes: Diagnostic characters of Lopholejeunea nigricans are: 1) ovate leaf

lobules, apex of lobule connected to the leaf lobe across only one single cell,

2) entire margin of female bract and bracteole, 3) female bract lobule of

almost the same length as the bract lobe. 4) perianth long exserted beyond

the bracts, with 4 toothed keels (rarely entire). This species was found

at ca 200-1700 m. It is separated from other species of the genus by the

combination of characters mentioned above.

9. Lopholejeunea ceylanica Steph., Sp. Hepat.5:86 (1912).—Type: Sri Lanka.

Horton Plain, Giesenhagen s.n. (G, lectotype designated by Verdoorn,

1934).

— Lopholejeunea levieri Schiffn., Ann. Bryol. 6: 134 (1933).—Type: Indonesia.

Sumatra, Mt. Singalang. “inter Riccardiam hymenophylloideum™, 1878,

Beccari s.n. (holotype, FH) — cf. Zhu and Gradstein (2005).

—Lopholejeunea schiffneri Verd., Ann. Bryol. 6: 134 (1933).—Type: Indonesia.

Java, Cibodas, “ad cortices,” Jan 1895, Massart 1546 p.p. (holotype, FH) — cf.

Zhu and Gradstein (2005).

Dioicous. Plants up to 1.5 cm long, about 0.8 mm wide: blackish brown in the

dried condition. Branching Lejeuwnea-type. Stem diameter 90-120 um Leaves

imbricate, widely spreading. Lobe ovate-orbicular, 450-870 um long, 350-750

um wide, margin entire, apex rounded to obtuse: cells of lobe thick-walled.

70 Gard. Bull. Singapore 62 (1) 2010

200 um

4 2 q y,

Soe ie 3 _ ee ie

ae =<

9

- Figure 9. Lopholejeunea ceylanica Steph. Shoot (a); underleaf (b); leaf lobule (c); leaf lobe (d);

female bract (e); female bracteole (f); cells of midleaf (g). Drawn from Haerida 1479, BO.

brown, hexagonal to isodiametric, trigones triangular, intermediate thickening

scarce; marginal cells 8-11 x 4-8 um, mid-leaf cells 14-30 x 15-23 um, basal

cells 27-32 x 22-25 um; oil bodies not seen. Lobule oblong 250-450 um long,

200-300 um wide, inflated, apex truncate with | small, unicellular tooth

pointing towards the leaf apex, connected to the leaf lobe across 2-3 cells.

Underleaves distant, orbicular, 150-300 um long, 190-350 um wide, margin

entire, apex rounded, insertion line nearly straight. Androecia intercalary

on branches, bracts in 5-9 pairs, similar to leaves but lobules larger, 400-

430 um long, 250-350 um wide, bract lobule about 2/3 of the bract lobe,

bracteoles similar to underleaves in size and shape. Gynoecia on short

or long branches, innovations lacking, bract oblong to broadly obovate

1000-1370 um long, 1050-1200 um wide, margin crenate, apex truncate to

orbicular, bract lobule large, about the same length as the bract lobe, oblong,

margin entire, apex acute; bracteole obovate 900-1200 um long, 1050-1100

um wide, margin entire, crenulate towards the apex, apex rounded. Perianth

immersed, obovate, 850 um long, 630 um wide, inflated, with 4 keels, margins

toothed. Sporophytes and asexual reproduction not seen.

Specimens examined: INDONESIA. West Java, Telaga Warna, Haerida 1479

(BO!); Cibodas Bot. Garden, ca 1450 m, Meijer B3801, 3810, Neervoort 811,

3353 (BO!);G. Pangrango, tea estate Mandalawangi, ca 1600 m, Meijer B387g

(BO!); G. Gede, Kandang Badak 2400 m, subalpine forest (forest canopy

ca 10 m high), common on tree trunks, Gradstein10241 (BIOT!, GOET!);

Lejeuneaceae subfamily Ptychanthoideae 7h

G. Cikurai, W slope, ca 1700 m, Verdoorn 59] (BO!); Cigombong (Pondok

Gedelanden), ca 500 m, Verdoorn 43c (BO!): G. Patuha, Kornochalert 1405

(BIOT!).

Distribution: Java, Sumatra, Borneo, Peninsular Malaysia, Sri Lanka,

Indochina.

Notes: Diagnostic characters of Lopholejeunea ceylanica are: 1) lobule large,

ca ¥2 x leaf length, with one small, unicellular tooth pointing towards the

leaf apex, 2) apex of leaf- lobule attached to the lobe across 2-3 cells, 3)

crenate margin of female bract, 4) female bract lobule large, acute. This

species grows at ca 500-2400 m. It resembles the very common L. subfusca

but differs by the more brownish color, the larger lobule (ca % x leaf length;

in L. subfusca ca 1/3 x leaf length) and the large lobule of the female bract

(very small in L. subfusca).

10. Lopholejeunea horticola Schiffn., Ann. Bryol. 6: 133 (1933). —Type:

Indonesia. Java, Bogor Bot. Garden, on trees (“In horto Buitenzorgensi ad

arbores”), Dec 1894, Massart 941 (holotype, FH; isotypes, FH, W) — cf. Zhu

and Gradstein (2005).

— Lopholejeunea glomerata Herzog, Ann. Nat. Mus. Wien 53: 370 (1942).

—Type: Indonesia. Sumatra, Padang, Schild 119 (holotype, W) — cf. Zhu and

Gradstein (2005).

( ie ; a 100 Hm) \

Figure 10. Lopholejeunea horticola Schiffn. Shoot (a); underleaf (b); leaf lobule (c); female bract

(d), female bracteole (e); cells of midleaf (f); cells of margin of leaf (g); perianth (h). Drawn from

Haerida 809, BO.

Te Gard. Bull. Singapore 62 (1) 2010

Dioicous. Plants up to 2 cm long, 0.6-0.9 mm wide, dark brown in dried

condition. Branching Lejeunea-type. Stem diameter 90-190 um Leaves

imbricate, obliquely spreading. Lobe orbicular, oblong to quadrate, 450-800

um long, 270-600 um wide, margin entire, apex rounded to obtuse; cells of

lobe thick-walled, dark brown, hexagonal, trigones triangular, intermediate

thickening scarce; marginal cells 7-10 x 6-8 um, mid-leaf cells 17-27 x 11-23

um, basal cells 28-36 x 18-30 um; oil bodies not seen. Lobule ovate, 190-

360 um long, 120-200 um wide, inflated, apex obliquely truncate, plane,

sometimes with 1 small tooth, connected to the leaf lobe across 2-3 cells.

Underleaves distant to contiguous, broadly-orbicular, 200-350 um long, 330-

610 um wide, margin entire, bases cuneate, apex rounded to almost truncate,

insertion line nearly straight. Androecia intercalary on branches, bracts in

3-11 pairs, similar to leaves but lobules larger, 190-230 um long, 140-180

um wide, bract lobule 2/3 of the bract lobe, strongly inflated, bracteoles

similar to underleaves in size and shape, 100-180 um long, 120-200 um wide.

Gynoecia on short or long branches, innovations lacking, bract obovate or

_ oblong + 1200 um long, 900 um wide, margin entire, apex obtuse to truncate,

bract lobule large, about the same length as the bract lobe, oblong, margin

entire, apex obliquely truncate; bracteole obovate, about 800 um long, 900

um wide, margin entire, apex truncate. Perianth immersed, obovate, about

800 um long, 570 um wide, inflated, with 4 keels, margins strongly toothed.

Sporophytes and asexual reproduction not seen.

Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden,

Neervoort 2098 (BO!):G. Gede, Cibeureum, above “Soekaboemi”, 1200-1400

m, Verdoorn 47b, 47a, 46 c (BO!); G. Gede Pangrango Nat. Park, Bodogol,

Haerida 809 (BO!); G. Malabar, SW slope, ca 1650 m, Verdoorn 62a (BO!):

G. Guntur, “Kawah Kamoedjan”, ca 1500-1700 m, Verdoorn 50a (BO!);:

G. Gegerbentang, E slopes, ca 1500-2000 m, Verdoorn 67h (BO!); without

locality, Pondok, Kurz s.n. (BO!); G. Patuha, ca 2000-2400 m, Verdoorn 60a

(BO!).

Distribution: Java, Sumatra, Bali, Moluccas, Peninsular Malaysia, Thailand.

Notes: Diagnostic characters of Lopholejeunea horticola are: 1) minute plant,

less than | mm wide, 2) lobule apex connected to the leaf lobe across 2-3

cells, 3) female bract lobule almost as large as the lobe, 4) margins of female

bract entire, 5) immersed perianth. This species grows at ca 800-2400 m. It

closely resembles the very common L. subfusca but differs by the much

smaller plant size and the large female bract lobule (very small in L. swbfusca).

[e)

Lejeuneaceae subfamily Prychanthoideae 7

11. Lopholejeunea recurvata Mizut., J. Hattori Bot. Lab. 46: 369 (1979).

—Type: Indonesia. Java, Gn. Gede, Cibodas, 1420 m, on bark of trees, Jun

1930, F Verdoorn s.n. (holotype, NICH; isotypes, FH, JE, L. S. U) — cf. Zhu

and Gradstein (2005).

20um

Figure 11. Lopholejeunea recurvata Mizut. Shoot (a): leaf lobe (b); leaf lobule (c); cells of midleaf

(d). Drawn from Verdoorn 20f. BO.

Autoicous. Plants up to 1.5 cm long, 1-1.2 mm wide; dark brown in dried

condition. Branching Lejeunea-type. Stem diameter 60-130 um Leaves

imbricate, widely spreading. Lobe oblong-rectangular, 430-760 um long,

390-530 um wide, margin entire, apex truncate: cells of lobe thick-walled,

brownish-orange, hexagonal toisodiametric,trigones triangular,intermediate

thickening scarce; marginal cells 6-11 x 6-9 um, mid-leaf cells 13-20 x 10-17

um, basal cells 26-33 x 11-19 um; oil bodies not seen. Lobule ovate, 160-220

um long, 130-180 um wide, inflated, apex obliquely truncate, plane, connected

to the leaf lobe across 2-3 cells. Underleaves contiguous to imbricate, wider

than long, 150-250 um long, 220-400 um wide, margin entire, apex recurved,

rounded to almost truncate, insertion line curved. Androecia intercalary on

branches, bracts in 4-8 pairs, similar to leaves but lobules larger, 200-250

um long, 100-150 um wide, bract lobule 1/2-2/3 of the bract lobe, bracteoles

similar to underleaves in size and shape. Gynoecia not seen.

Specimen examined: INDONESIA. West Java, Cibodas Bot. Garden,

Verdoorn 20f (BO!).

Distribution: Java, Sumatra, Bali, Peninsular Malaysia, Papua New Guinea.

74 Gard. Bull. Singapore 62 (1) 2010

Notes: Diagnostic characters of Lopholejeunea recurvata are: 1) recurved

margins of leaf lobes and underleaves, 2) lobule apex connected to the leaf

lobe across 2-3 cells. This rare species grows at ca 500-1900 m. It differs from

other Javanese species of the genus by the recurved leaf margins.

12. Lopholejeunea subfusca (Nees) Schiffn., Bot. Jahrb. Syst. 23: 593 (1897);

Jungermannia subfusca Nees, Enum. Pl. Crypt. Jav. 1: 36 (1830); Lejeunea

subfusca (Nees) Nees & Mont., Ann. Sci. Nat., Bot., Sér. 2, 5: 61 (1836);

Phragmicoma subfusca (Nees) Nees, Naturgesch. Eur. Leberm. 3:248 (1838);

Symbiezidium subfuscum (Nees) Trevis., Mem. Reale Ist. Lombardo Sci.,

Cl. Sci. Mat., Ser. 3, 4: 403 (1877); Lopholejeunea sagraeana var. } subfusca

(Nees) Schiffn., Consp. Hepat. Archip. Ind.: 294 (1898). —Type: Indonesia.

Java, without locality, Blume s.n. (holotype, STR; isotypes, W, S) — cf. Zhu

and Gradstein (2005).

200 ym

(es

on /

iS | bherita, 2008 : és rs

i See

* |200 Hm at

b>

:

Figure 12. Lopholejeunea subfusca (Nees) Schiffn. Shoot (a); leaf lobule (b); female bract (c);

perianth (d); cells of midleaf (e). Drawn from Haerida 850, BO.

Autoicous. Plants up to 2 cm long, 1-1.3 mm wide, dark brown to black

in dried condition. Branching Lejeuwnea-type. Stem diameter 90-120 um.

Leaves imbricate, widely spreading. Lobe oblong-orbicular, 410-700 tm

long, 420-620 um wide, margin entire, apex rounded; cells of lobe thick-

walled, pale brown to orange, hexagonal to isodiametric, trigones triangular,

intermediate thickening scarce; marginal cells 6-9 x 5-7 um, mid-leaf cells 17-

24 x 15-20 um, basal cells 22-30 x 16-23 um; oil bodies not seen. Lobule ovate,

small, 180-220 um long, 120-150 um wide, inflated, apex obliquely truncate,

plane, connected to the leaf lobe across 2-4 cells. Underleaves distant to

Lejeuneaceae subfamily Ptychanthoideae iS

contiguous, broadly-orbicular, 220-400 um long, 340-610 um wide, margin

entire, apex truncate, insertion line shallowly curved. Androecia intercalary

on branches, bracts in 3-16 pairs, similar to leaves but lobules larger, 230-430

um long, 170-320 um wide, bract lobule almost the same size of the bract

lobe, bracteoles similar to underleaves in shape, 170-270 um long, 180-450

um wide. Gynoecia on short or long branches, innovations lacking, bracts and

bracteoles in 2 pairs, bract ovate to broadly oblong 700-850 um long, 700-860

um wide, margin irregularly toothed, apex orbicular to nearly truncate, sinus

up to 1/4 of lobe length, bract lobule very small; bracteole broadly orbicular

170-280 um long, 180-450 um wide, margin entire, apex rounded. Perianth

immersed, obovate, about 870 um long, 600 um wide, inflated, with 2 keels,

margins toothed. Sporophytes and asexual reproduction not seen.

Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer

92b, 26b, Verdoorn 124, 141, 249 (BO!): Cibodas Bot. Garden, Neervoort 952,

Verdoorn 20¢, 20d, Schiffner 250 (BO!), ibid., Lee & Nova Indri 49 (BIOT!,

UKMB)!), ibid., ca 1425 m, Meijer B4017, B3800 (BO!); G. Gede Pangrango

Nat. Park, Bodogol, ca 800 m, Haerida 850, 844 (BO!); G. Pangrango, ravine

above Kampong G. Mas, Meijer B616i (BO!); Geger Bentang, Neervoort

1344 (BO!); G. Halimun Nat. Park, Citalahab, 1065 m, Radhiah Zakaria

254c (BIOT!); G. Papandayan, Tji Paroegpoeg, ca 2300-2500 m, Verdoorn

54b (BO!); G. Cikurai, W slope, ca 1700 m, Verdoorn 59i (BO!); G. Patuha,

Kornochalert 1416, 1400 (BIOT!).

Distribution: Java, Sumatra, Borneo, Bali, Sulawesi, Moluccas, Philippines,

Papua New Guinea, Pacific, Peninsular Malaysia, Indochina, China, Japan,

India, Sri Lanka, Nepal, tropical Africa, tropical America.

Notes: Diagnostic characters of Lopholejeunea subfusca are: 1) lobule apex

truncate, connected to the leaf lobe across 2-4 cells, 2) very small size of

female bract lobule, 3) entire margin of female bracteole, 4) immersed

perianth. This very common species is very grows at ca 200-2500 m. Like

in several other Javanese species of Lopholejeunea, the lobule apex is

connected to the leaf lobe surface across 2-4 cells; the flat leaves with rounded

apex, the orbicular, distant underleaves, the rather small leaf lobule, and the

characters of the gynoecium mentioned above, readily separate this species

from its relatives.

13. Lopholejeunea wiltensii Steph., Hedwigia 35: 112 (1896).—Type: Indonesia.

Sumatra, Padang, A. Wiltens (holotype, G) — cf. Zhu and Gradstein (2005).

76 Gard. Bull. Singapore 62 (1) 2010

— Lopholejeunea serrifolia Steph., Sp. Hepat. 5: 84 (1912). —Type: Indonesia.

Java, without locality, collector unknown (G, lectotype designated by

Mizutani, 1979).

i "

200"um } Hara 2004 ih

a :

Figure 13. Lopholejeunea wiltensii Steph. Shoot (a); leaf lobule (b); cells of midleaf (c); female

bracteole (d): female bract (e). Drawn from Verdoorn 30d, BO.

Dioicous. Plants up to 4 cm long, 0.9-1.3 mm wide; pale brown in dried

condition. Branching Lejeuwnea-type. Stem diameter 50-100 um Leaves

imbricate, widely spreading. Lobe broadly ovate to broadly orbicular, 500-

650 um long, 580-610 um wide, margin entire, apex rounded; cells of lobe

thick-walled, pale yellow, quadrate to isodiametric, trigones triangular,

intermediate thickening scarce; marginal cells 5-7 x 2-6 um, mid-leaf cells

11-13 x 7-10 um, basal cells 20-25 x 13-23 um; oil bodies not seen. Lobule

triangular to quadrate, small, 190-220 um long, 110-130 um wide, inflated

at the basal part forming somewhat a longitudinally elliptical sac, apex

obliquely truncate with 1 tooth consisting of 1-2 cells, connected to the leaf

lobe across 2-3 cells. Underleaves distant to contiguous, reniform, 210-360

um long, 310-610 um wide, margin entire, apex rounded to almost truncate,

insertion line strongly curved. Androecia not seen. Gynoecia on short or

long branches, innovations lacking, bract oblong-ovate to broadly ovate,

about 470 um long, 460 um wide, margin irregularly toothed, apex obtuse to

orbicular, sinus up to 1/3 of lobe length, bract lobule oblong about 1/3 of the

bract lobe length, margin entire; bracteole broadly ovate about 350 um long,

410 um wide, margin irregularly toothed, apex truncate. Perianth immersed,

obovate, about 970 um long, 670 um wide, inflated, with 4 keels, margins

toothed. Sporophytes and asexual reproduction not seen.

Lejeuneaceae subfamily Ptychanthoideae al

Distribution: Java, Sumatra, Peninsular Malaysia, Philippines, Sulawesi.

Moluccas, West Irian, Papua New Guinea, New Caledonia.

Specimens examined: West Java, Cibodas Bot. Garden, 1400 m, Ariyanti 458

(BIOT!), ibid., along the road to Huis ten Bosch, Meijer B1078 (BO!): above

Cibodas, Cibeureum, Schiffner 252, 253, Verdoorn 30d, 30c, 30a (BO!); G.

Gede, Soekaboemi, Verdoorn 46a (BO!): G. Pangrango, *Tjisaroea”, ca 1200

m, Verdoorn 44a (BO!):Tugu, tea estate G. Mas, Meijer & van der Wijk BO59b

(BO!); above tea estate G. Mas, Jalan Mandalawangi, Meijer B335a (BO!):

G. Guntur, “Kawah Kamoedjan”, ca 1500-1700 m, Verdoorn 50b (BO!).

Notes: Diagnostic characters of Lopholejeunea wiltensii are: 1) pale color of

the plant in the dried condition, 2) lobule small, in the basal part forming a

somewhat longitudinally elliptical sac, 3) lobule apex connected to the leaf

lobe across 2-3 cells, 4) reniform underleaves. This species grows at ca 1200-

1900 m. Lopholejeunea wiltensii is closely related to L. zollingeri and was

sometimes considered a synonym of the latter. It differs from L. zollingeri

by the characters given in the key.

14. Lopholejeunea zollingeri (Steph.) Schiffn.,Consp. Hepat. Arch. Ind.:296

(1898); Lejeunea zollingeri Steph., Hedwigia 29: 14 (1890). —-Type: Indonesia.

Java, without locality, collector unknown (W, lectotype designated by

Mizutani, 1979).

— Lopholejeunea latialata Mizut., J. Hattori Bot. Lab. 46: 365 (1979). —Type:

Indonesia. Java: without locality, collector unknown, ex Herb. Sande Lacoste

(holotype, L) — cf. Zhu and Gradstein (2005).

7

Ye

hy es

Figure 14. Lopholejeunea zollingeri (Steph.) Schiffn. Shoot (a): leaf lobule (b): leaf lobe (c):

underleaf (d); cells of midleaf (e). Drawn from Meijer 387c, BO.

78 Gard. Bull. Singapore 62 (1) 2010

Autoicous. Plants up to 3 cm long, 0.7-1.2 mm wide, blackish brown in dried

condition. Branching Lejeunea-type. Stem diameter 60-160 um. Leaves

imbricate, widely spreading. Lobe orbicular to broadly ovate, 200-420 um

long, 230-460 um wide, margin entire, apex rounded, usually recurved; cells

of lobe thick-walled, brown, hexagonal to isodiametric, trigones triangular,

intermediate thickening frequent; marginal cells 5-11 x 7-9 um, mid-leaf cells

11-19 x 9-15 um, basal cells 25-30 x 19-22 um; oil bodies not seen. Lobule

ovate, large, almost 1/3 of the leaf lobes length, 190-210 um long, 160-180 um

wide, inflated at the middle part forming a globose sac, apex obliquely truncate

with | small tooth or without tooth but forming a sharp angle, connected to

the leaf lobe across 3-4 cells. Underleaves closely imbricate, reniform, 190-420

um long, 370-710 um wide, margin entire, apex rounded to almost truncate,

insertion line strongly arched. Androecia intercalary on branches, bracts in 4-

9 pairs, similar to leaves but lobules larger, 200-220 um long, 160-190 um wide,

bract lobule 1/3 of the bract lobe, bracteoles similar to underleaves in shape,

110-170 um long, 180-300 um wide. Gynoecia not seen.

Specimens examined: INDONESIA. West Java, G. Pangrango, tea estate

Mandalawangi, ca 1600 m, Meijer B387c (BO!); G. Patuha, ca 1600 m,

Kornchalert 1413, Gradstein 12169 (BIOT!).

Distribution: Java, Sumatra, Borneo, Sulawesi, Peninsular Malaysia,

Philippines, Papua New Guinea, Sri Lanka, China, Japan, Fiji.

Notes: Diagnostic characters of Lopholejeunea zollingeri are: 1) broadly

orbicular leaf lobe with recurved apex, 2) leaf lobule large, apex connected

to the leaf lobe across 3-4 cells, 3) closely imbricate, large, reniform

underleaves with strongly arched insertion line. This species grows at ca

1350-1600 m. Lopholejeunea zollingeri has very large, reniform underleaves

like in L. wiltensti, L. eulopha and L. applanata but differs from the latter

two by the lobule apex which is attached to the lobe across 3-4 cells (across

1 cell in L. eulopha and L. applanata). For differences with L. wiltensii see

the characters given in the key.

15. Mastigolejeunea auriculata (Wils.) Schiffn., in Engler and Prantl, Nat.

Pflanzenfam. 1, 3: 129 (1893); Jungermannia auriculata Wils., in Drummond,

Musci Amer. Exsicc. (Southern States) nr. 170 (1841); Ptychocoleus

auriculatus (Wils.) Trevis., Mem. Reale Ist. Sci. Mat. Nat., Ser. 3, 4: 405

(1877). -Type: USA. Louisiana, New Orleans, Drummond s.n. (holotype,

BM; isotypes, MANCH, PC) —cf. Gradstein et al. (2002).

—Phragmicoma humilis Gottsche, in Gottsche et al., Syn. Hepat. 299 (1845);

Mastigolejeunea humilis (Gottsche) Schiffn., in Engler and Prantl, Nat.

Lejeuneaceae subfamily Ptychanthoideae 79

Pflanzenfam. 1, 3: 129 (1893). —Type: Indonesia. Java, without locality,

Junghuhn s.n. (isotypes, B, W) — cf. Gradstein et al. (2002).

Figure 15. Mastigolejeunea auriculata (Wils.) Schiffn. Shoot (a); underleaf (b):; leaf lobe (c); leaf

lobule (d); cells of basal part of the leaf (e); cells of midleaf (f). Drawn from Meijer 92a, BO.

Plants up to 2 cm long, 0.8-1 mm wide, dull green to dark brown in dried

condition. Branching Lejeunea-type. Stem diameter 70-170 um Leaves

imbricate, obliquely spreading, somewhat squarrose. Lobe ovate, 440-1030

um long, 410-730 um wide, margin entire, apex rounded to obtuse; cells of

lobe thick-walled, light yellow, rhomboidal to hexagonal, trigones cordate,

intermediate thickening scarce; marginal cells 11-15 x 6-10 um, mid-leaf cells

18-21 x 9-12 um, basal cells 25-34 x 16-19 um; oil bodies not seen. Lobule

oblong to subrectangular, 190-440 um long, 130-200 um wide, apex truncate

to obliquely truncate with 1 small tooth. Underleaves imbricate, obdeltoid

to nearly triangular, 200-530 um long, 220-490 um wide, margin entire, apex

truncate, base cuneate, insertion line curved. Androecia not seen. Gynoecia

on short or long branches, with 1-2 lejeuneoid innovations, bract oblong to

broadly oblong, 1000-1070 um long, 600-800 um wide, margin entire, apex

subrounded to nearly truncate, sinus 1/3 to 2/3 of lobe length, bract lobule

oblong-ovate to subrectangular, 500-670 um long, 340-360 um wide, margin

somewhat undulated towards the apex; bracteole obdeltoid 600-770 um

long, 850-960 um wide, margin entire, apex truncate. Perianth obovate, 1010

um long, 440 um wide, with 3 keels, margins entire. Asexual reproduction

not observed.

Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, van

Borssum Waalkes 122a, Meijer 92a, 552d (BO!). Banten, Taman Nasional

Ujung Kulon, Dewi Dw 922b (BO!).

80 Gard. Bull. Singapore 62 (1) 2010

Distribution: Java, Borneo, Sulawesi, Moluccas, West Irian, Papua New

Guinea, Australia, Solomon Is., India, Indochina, China, Japan, tropical

Africa, tropical America.

Notes: Diagnostic characters of Mastigolejeunea auriculata are: 1) leaf lobe

ovate in shape, margins entire, 2) leaf cells elongate, with cordate trigones

3) lobule with truncate to obliquely truncate apex, with 1 small tooth, 4)

underleaves obdeltoid, 5) perianth 3-keeled, with smooth keels and 1-2

innovations. This species grows at ca 200-1700 m. It is very similar to M.

virens but differs by the very short, blunt lobule tooth.

16. Mastigolejeunea indica Steph., Sp. Hepat. 4: 776 (1912). —Type: India.

Nicobar Is., Man s.n. (holotype, G) —cf. Gradstein et al. (2002).

—Thysananthus integrifolius Steph., Sp. Hepat. 4: 788 (1912). Mastigolejeunea

integrifolia (Steph.) Verdoorn, Blumea 1: 231, 239 (1934). —Type: Australia.

Torres Str., Possession I., Micholitz s.n. (holotype, G; isotype, FH) -cf.

Gradstein et al. (2002).

™ -

ee

Figure 16. Mastigolejeunea indica Steph. Shoot (a); leaf lobe (b); underleaf (c); leaf lobule (d); cells

of margin of leaf (e); cells of midleaf (f). Drawn from Meijer 75a, BO.

Dioicous. Plants up to 2.5 cm long, 0.6-1 mm wide, 1-1.5 mm wide in wet

condition, greenish-brown in dried condition, becoming more green towards

the apex of the plant. Branching Lejewnea-type. Stem diameter 100-250 um,

epidermis cells very unequal in size, dorsal one much large than ventral ones,

hyalodermis lacking. Leaves imbricate, obliquely spreading, convolute when

Lejeuneaceae subfamily Ptychanthoideae 8]

dry. Lobe ovate-oblong, 520-1150 um long, 300-890 um wide, margin entire,

apex acute to obtuse; cells of lobe thin-walled, light yellow, rhomboidal,

trigones cordate, intermediate thickening scarce; marginal cells 6-11 x 3-5

um, mid-leaf cells 7-18 x 6-14 um, basal cells 21-35 x 17-20 um; oil bodies not

seen. Lobules ovate to triangular, 180-390 um long, 160-280 um wide, slightly

inflated along the keel, apex obliquely truncate with 3-4 triangular teeth, each

tooth consisting of 3-4 cells, 2-3 cells long, first and second tooth sometimes

blunt, only 1-2 cells long, sometimes with or without fourth, 1-2 cells long

tooth. Underleaves imbricate, obdeltoid, 230-520 um long, 250-540 um wide,

margin entire, apex truncate, usually recurved and seemingly emarginate,

base auriculate, insertion line curved. Generative structures not seen.

Specimens examined: Inodnesia. West Java, Bogor Bot. Garden, ca 250m,

Meijer 75a (BO!).

Distribution: Java, China, India (Nicobar), Philippines, Papua New Guinea,

Australia.

Notes: Diagnostic characters of Mastigolejeunea indica are: 1) plant like

M. auriculata but lobule with 3-4 large teeth, 2) stem cross section with

very unequal epidermis cells, dorsal ones much large than ventral ones

(hyalodermis lacking). Mastigolejeunea indica is new to Java. The species

has only been collected in the Botanical Garden of Bogor, at ca 250 m. It

is readily separated from all other species of the genus Mastigolejeunea by

the lobule with 3-4 large triangular teeth (usually only one tooth in other

species of the genus).

17. Mastigolejeunea virens (Angstr.) Steph., in Sp. vee A/G. VOID:

Thysananthus virens Angstr., Ofv. K.Vetensk Akad. Férh. 5: 131. 1873. -

Type: Society Is., Moorea, Andersson s.n. (holotype, S; ieee FH, G) -cf.

Gradstein et al. (2002).

—Mastigolejeunea humilis sensu Verdoorn 1933 p.p.

Autoicous. Plants up to 3 cm long, 0.9-2 mm wide, brown to dark brown

in dried condition. Branching Lejeunea-type. Stem diameter 60-190 um

Leaves closely imbricate, widely spreading. Lobe ovate to oblong, 340-920

um long, 200-640 um wide, margin entire, apex rounded to obtuse; cells of

lobe thick-walled, light-yellow, rhomboidal, trigones cordate, intermediate

thickening scarce; marginal cells 6-11 x 5-6 um, mid-leaf cells 11-16 x 5-

7 um, basal cells 18-26 x 8-10 um; oil bodies not seen. Lobule ovate, 140-

280 um long, 60-170 um wide, inflated, apex obliquely truncate with 1 long

tooth consisting of 3-5 cells. Underleaves imbricate, quadrangular to nearly

82 Gard. Bull. Singapore 62 (1) 2010

triangular, 190-560 um long, 250-600 um wide, margin entire, apex truncate,

sometimes recurved forming a hearth shaped. Androecia intercalary on

branches, bracts in 5 pairs, similar to leaves but lobules larger, bract lobule

2/3 of the bract lobe, 100-110 um long, 50-60 um wide, bracteoles similar to

underleaves in size and shape, 90-120 um long, 100-120 um wide. Gynoecium

with 1-2 subfloral innovations; innovation type lejeuneoid; bract lobe ovate-

oblong, ventral margin incurved, margin entire, apex rounded, 1160 um

long, 460 um wide; bract lobule about % of bract lobe length, oblong, margin

crenulate; bracteole oblong, margin entire, apex rounded. Perianth 820 um

long, oblong, inflated, with 3 keels, margins entire. Sporophytes and asexual

reproduction not seen.

10 Hm

Figure 17. Mastigolejeunea virens (Angstr.) Steph. Shoot (a); leaf lobe (b); underleaf (c); cells of

midleaf (d); cells of basal part of the leaf (e). Drawn from Meijer 62a, BO.

Specimens examined: INDONESIA. West Java, Bogor, without locality,

Verdoorn 149 (BO!); Bogor Bot. Garden, Schiffner 257, W. Meijer B24,

62a, B55f, B998b, Verdoorn 12z, 12d, 143, 255 (BO!); Cibodas Bot. Garden,

Neervoort 154, 918, Meijer B3720, B3815 (BO!), ibid., Lee & Nova Indri 62

(BIOT!, UKMB!), ibid., Ariyanti 472 (BIOT!); Telaga Bodas, Verdoorn 58a,

5Sb (BO!); G. Megamendung, above “Toegoe”, Schiffner 256 (BO!).

Distribution: Java, Borneo, Moluccas, Philippines, Peninsular Malaysia,

Thailand, Sri Lanka, Papua New Guinea, Australia, Pacific Islands.

Notes: The occurrence of Mastigolejeunea virens in Java has not been

reported before, therefore this species is new for Java. Verdoorn (1933)

Lejeuneaceae subfamily Ptychanthoideae 83

treated the species as a synonym of M. humilis (= M. auriculata), but it is

readily separated from the latter by the longer lobule tooth (see key). The

diagnostic characters of Mastigolejeunea virens are: 1) leaf lobe rounded to

obtuse at apex, 2) lobule with | long tooth consisting of 3-5 cells, 3) perianth

with 3 keels. This species grows at ca 200-1500 m.

18. Ptychanthus striatus (Lehm. & Lindenb.) Nees, Naturgesch. Eur.

Leberm. 3: 212 (1838); Jungermannia striata Lehm. & Lindenb., Nov. Stirp.

Pug. 4: 16 (1832); Bryopteris striata (Lehm. & Lindenb.) Mitt., in Seemann,

Fl. Vit. 411 (1873). -Type: Nepal. Wallich s.n. & s.d. (holotype, S; isotypes, G,

W) -cf. Gradstein er al. (2002).

—Jungermannia retusa Reinw. et al. var. f Nees, Enum. PI. Crypt. Javae 1: 39

(1830); Ptychanthus retusus (Reinw. et al.) Nees var. B Nees, in Gottsche et al.,

Syn. Hepat. 292. 1845: Ptycholejeunea retusa (Reinw. et al.) Steph., Hedwigia

28: 258 (1889); Ptychanthus retusus (Reinw. et al.) Steph., Sp. Hepat. 4: 743

(1912): Ptychanthus striatus var. retusus (Reinw. et al.) Verd., Ann. Bryol.

Suppl. 4: 122 (1934). —Type: Indonesia. Java, without locality, Blume s.n.

(holotype, STR; isotype, W) —cf. Gradstein et al. (2002).

Figure 18. Prychanthus striatus (Lehm. & Lindenb.) Nees. Shoot (a); underleaf (b): leaf lobe (c):

cells of midleaf (d). Drawn from Meijer 840, BO.

Autoicous. Plants robust, up to 10 cm long, 1-1.5 mm wide, standing up away

from the substrate or pendent, pinnate, greenish-brown in dried condition.

Branching Frullania-type. Stem diameter 130-300 um Leaves imbricate,

flat, obliquely spreading. Lobe broadly-ovate, 650-2170 um long, 300-1260

um wide, margin entire or slightly toothed towards the apex, apex acute,

base auriculate at the proximal side; cells of lobe thick-walled, light-yellow,

84 Gard. Bull. Singapore 62 (1) 2010

rhomboidal to hexagonal, trigones cordate, intermediate thickening frequent;

marginal cells 7-16 x 4-11 um, mid-leaf cells 20-27 x 8-14 um, basal cells 26-38

x 11-20 um; oil bodies not seen. Lobule ovate, small, 240-370 um long, 110-

170 um wide, inflated, apex with 1-2 teeth, teeth consisting of 1 small cell.

Underleaves distant to contiguous, broadly-ovate, sometimes subrectangular

in outline, 480-760 um long, 470-1000 um wide, base auriculate, margin entire,

apex irregularly toothed. Androecia intercalary on branches, bracts in 5-10

pairs, bract lobe ovate, 560-1150 um long, 300-520 um wide, margin entire,

apex acute, bract lobule 1/2 of the bract lobe, inflated, 390-500 um long, 200-

240 um wide, bracteoles similar to underleaves in size and shape. Gynoecia

on short or long branches, innovation type lejeuneoid, bract lobe ovate with

small bract lobules, 1220-1320 um long, 550-630 um wide, margin toothed

towards the apex, apex acute, bract lobules oblong, 460-710 um long, 80-

150 um wide; bracteole broadly ovate, 1110-1150 um long, 1220-1330 um

wide, margin toothed, apex rounded. Perianth immersed, elliptical, 1770 um

long, 730 um wide, inflated, with + 10 keels, margins entire. Sporophytes and

- asexual reproduction not seen.

Specimens examined: INDONESIA. West Java, G. Salak, ca 1000 m,

Zollinger 3560, Schiffner 259 (BO!); G. Gede, above Cibodas, 1600-1900 m.

Neervoort 107, 129, 227, 2775, 2401, 282, 2271, Verdoorn 30g, 30f, 21a, 30n,

Iwamasa s.n., Meijer B4137, B4076, B160 (BO!), ibid., above Tanjung Mas

waterfall (Pancuran Mas), Alston 12807 (BO!), ibid., Cibeureum, Schiffner

266, Verdoorn 30h, 301, 30}, 30n (BO!), ibid., trail Cibeureum falls to Kandang

Badak 1700-2200 m, montane forest, epiphytes on trunk bases, common,

Gradstein 10215, 10217 (BIOT!, GOET!); G. Pangrango, “Tjisaroea”, ca 1200

m, Verdoorn 44a, 44b, 44c (BO!); Tugu, slope of Gede-Pangrango, above G.

Mas, Meijer B3394a, BS40, B421 (BO!); G. Geger Bentang, E slopes, 1500-

2000 m, Neervoort 2890, Verdoorn 67a, 67b, 267, Meijer B5584 (BO!); G.

Megamendung, Schiffner 265 (BO!); G. Malabar, SW slopes of Punciak

Besar 1650-2300 m, Verdoorn 61a, 61b, 61c, 62b, 62d (BO!); G. Patuha, 2000-

2400 m, Verdoorn 60b (BO!), Kornochalert 1414 (BIOT!).

Distribution: Java,Sumatra, Peninsular Malaysia, Borneo,Sulawesi, Moluccas,

Philippines, West Irian, Papua New Guinea, Australia, New Zealand, Pacific

Islands, India, Sri Lanka, Taiwan, Indochina, China, Japan, tropical Africa.

Notes: Diagnostic characters of Ptychanthus striatus are: 1) Frullania-type

branching, 2) lobe with acute apex and entire or toothed margins, 3) leaf

cells elongate, with cordate trigones, 4) perianth with 10 smooth keels, and

with innovation, 5) plant robust, pinnate, usually pendent. This species is

common in the mountains where it occurs at ca 1000-2400 m. By its large

(oe)

Nn

Lejeuneaceae subfamily Ptychanthoideae

size, its pinnate, Frullania-type branching and acute leaf lobes the speciers

is unmistakable and cannot be confused with any other Javanese member

of Lejeuneaceae.

19. Schiffneriolejeunea pulopenangensis (Gottsche) Gradst., J. Hattori Bot.

Lab. 38: 335 (1974); Phragmicoma pulopenangensis Gottsche, Syn. Hepat.

299 (1845); Ptychocoleus pulopenangensis (Gottsche) Trevis., Mem. Reale Ist.

Lombardo Sci., Cl. Sci. Mat., Ser. 3, 4: 405 (1877).—Type: Peninsular Malaysia.

Pulo Penang, Delessert s.n. (holotype, PC-Mont; isotypes, BM, S, W).

Plants up to6cm long, 1.5-2 mm wide, brown to dark brown in dried condition.

Vegetative branching Frullania-type, or Lejeunea-type. Stem diameter 150-

260 um. Leaves imbricate, obliquely spreading, convolute when dry. Lobe

broadly ovate to somewhat triangular, 700-1060 um long, 500-850 um wide,

margin entire, apex obtuse to nearly acute; cells of lobe thick-walled, hyaline,

rhomboidal to hexagonal, trigones cordate, intermediate thickening scarce;

marginal cells 9-24 x 7-16 um, mid-leaf cells 17-26 x 8-19 um, basal cells 25-

45 x 13-23 um; oil bodies not seen. Lobule ovate, 300-420 um long, 150-280

100pHm

Figure 19. Schiffneriolejeunea pulopenangensis (Gottsche) Gradst. Shoot (a); leaf lobe (b); apex

of leaf lobule (c); underleaf (d); cells of midleaf (e). Drawn from Meijer & Alston 5329, BO.

um wide, apex with 2 conspicuous teeth that point outwards towards the

leaf apex, first tooth consisting of 3-4 cells, second tooth consisting of 2-

3 cells, lobule free margin plane, with 2 clearly visible teeth. Underleaves

imbricate, obdeltoid-orbicular sometimes with recurved apex seemingly

emarginate, 360-420 um long, 480-610 um wide, margin entire, apex truncate,

86 Gard. Bull. Singapore 62 (1) 2010

insertion line deeply curved. Androecia on lateral branches, inflated, in 3-11

pairs, about 1/3-1/2 of the vegetative lobe, hypostatic, bracteoles oblong to

obdeltoid, apex truncate, smaller than the underleaves. Gynoecia not seen.

Specimen examined: INDONESIA. West Java, Bogor Bot. Garden, Meijer

& Alston 5329 (BO!).

Distribution: Indomalesia, Australia.

Notes: Diagnostic characters of Schiffneriolejeunea pulopenangensis are:

1) branching Lejeunea and Frullania type, 2) leaf lobule ovate, with plane

free margin and with 2 long teeth that point outwards to the leaf apex, 3)

leaves not squarrose when moist, convolute when dry, 4) trigones cordate.

This species is apparently rare in Java and has only been collected in the

Botanical Garden of Bogor at ca 250 m. The species is closely related to S.

tumida but differs by the lobule without saccate base and by the flat leaves

- (squarrose in S. tumida).

20. Schiffneriolejeunea tumida var. haskarliana (Gottsche) Gradst. &

Terken, Occas. Pap. Farlow Herb. 16: 77 (1981); Phragmicoma hasskarliana

Gottsche,in Gottsche eral.,Syn. Hepat.299 (1845): Acrolejeunea hasskarliana

(Gottsche) Schiffn.,in Engler and Prantl, Nat. Pflanzenfam. 1, 3: 129 (1893);

Ptychocoleus hasskarliana (Gottsche) Steph., Sp. Hepat. 5: 44 (1912). —Type:

Indonesia. Java, without locality, Hasskarl 20 (holotype, B; isotypes, G, S, W)

— cf. Gradstein and Terken (1981).

—Mastigolejeunea badia Gottsche ex Steph., Sp. Hepat. 4: 779 (1912). —Type:

Solomon Is., Vanikoro, Lesson s.n. (isotypes, BM, FH) —cf. Gradstein and

Terken (1981).

Plants up to 6 cm long, 2-3 mm wide; brown to dark brown in the dried condition.

Vegetative branching Frullania-type, or Lejeunea-type. Stem diameter 160-270

um. Leaves imbricate, obliquely spreading, somewhat squarrose, convolute

when dry. Lobe broadly ovate to somewhat triangular, 900-1150 um long, 700-

1200 um wide, margin entire, apex obtuse to nearly acute; cells of lobe thick-

walled, hyaline, rhomboidal to hexagonal, trigones cordate, intermediate

thickening scarce; marginal cells 13-24 x 9-22 um, mid-leaf cells 19-29 x 18-27

um, basal cells 25-40 = 18-32 um; oil bodies not seen. Lobule ovate-rectangular,

300-620 tm long, 130-270 um wide, apex with 2 teeth, each tooth consisting

of 3-4 cells, lobule free margin strongly involute, forming a sac at the base of

the lobule. Underleaves imbricate, obdeltoid sometimes with recurved apex

seemingly emarginate, 420-840 um long, 490-720 um wide, margin entire, apex

truncate, insertion line deeply curved. Generative structures not seen.

Lejeuneaceae subfamily Ptychanthoideae 87

I.Meerite.2009 | Fa

Imm

Figure 20. Schiffneriolejeunea tumida var. haskarliana (Gottsche) Gradst. & Terken. Shoot (a): leaf

lobe (b); apex of leaf lobule (c); cells of midleaf (d). Drawn from Haerida 724, BO.

Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden,

Neervoort 2120 (BO!), ibid., on bark of Altingia excelsa,ca 1400 m, Gradstein

10202 (BIOT!, GOET!); above Cibodas, trail to Cibeureum waterfall, on

tree trunk, Lee & Nova Indri 58 (BIOT!, UKMB!); G. Halimun Nat. Park,

Haerida 724 (BO!).

Distribution: Java, Papua New Guinea, Solomon Is., Australia.

Notes: Diagnostic characters of Schiffneriolejeunea tumida var. haskarliana

are: 1) branching Lejeunea- and Frullania-type, 2) leaf lobule with 2

conspicuous, erect teeth and with a sac at the base by the strongly involute

free margin, 3) leaves somewhat squarrose when moist, convolute when dry,

4) cordate trigones. This species grows at ca 1000-1500 m. It is closely related

to S. pulopenangensis but occurs at higher elevations; for morphological

differences see under the latter species and the key.

21. Spruceanthus polymorphus (Sande Lac.) Verd., Ann. Bryol. Suppl. 4: 155

(1934): Phragmicoma polymorpha Sande Lac., Ned. Kruidk. Arch. 34: 420

(1854); Phragmolejeunea polymorpha (Sande Lac.) Schiffn., Forschungsr.

Gazelle 4: 25. (1890); Thysananthus polymorphus (Sande Lac.) Schiffn.,

Consp. Hepat. Arch. Ind. 305 (1898): Archilejeunea polymorpha (Sande Lac.)

B. Thiers & Gradst., Mem. N.Y. Bot. Garden 52: 10 (1989). —Type: Indonesia.

Java, Junghuhn s.n. (holotype, L; isotype, NY) —cf. Gradstein et al. (2002).

88 Gard. Bull. Singapore 62 (1) 2010

Autoicous. Plants up to 4 cm long, 1.5-2.5 mm wide, pale green to greenish

brown in dried condition. Branching Lejeunea-type. Stem diameter 100-250

um: ventral merophyte 8-12 cells wide. Leaves imbricate, widely spreading.

Lobe ovate-oblong, 730-1300 um long, 430-640 um wide, margins entire or

toothed, plane or undulate, apex rounded to obtuse; cells of lobe isodiametric,

with large, nodulose intermediate thickenings, hyaline, trigones not cordate:

marginal cells 7-12 x 6-13 um, mid-leaf cells 20-26 x 19-24 um, basal cells

26-52 x 13-18 um: oil bodies not seen. Lobule ovate, 110-230 um long, 80-130

um wide, apex obliquely truncate with 1-2 small teeth. Underleaves closely

imbricate, obdeltoid to orbicular, 230-400 um long, 400-500 um wide, margin

entire at lateral part, serrate towards the apex, apex rounded to truncate,

insertion line shallowly curved. Androecia intercalary on branches, bracts

up to 14 pairs, similar to leaves but lobules larger, 580-650 um long, 250-300

um wide, hypostatic, bracteoles similar to underleaves in size and shape.

Gynoecia with 1-2 lejeuneoid innovations, bract ovate, about 900 um long,

600 um wide, margin entire, apex rounded, bract lobule ovate about 2/3 of

- the lobe length, toothed; bracteole orbicular to oblong, 530-600 um long,

580-900 um wide, margin toothed. Perianth oblong, about 1300 um long,

430 um wide, with 7 smooth keels, margins entire. Sporophytes and asexual

reproduction not seen.

20 um

Figure 21. Spruceanthus polymorphus (Sande Lac.) Verd. Shoot (a); underleaf (b): cells of basal

cells of the lobe (c); leaf lobe (d); cells of midleaf (e). Drawn from Borssum Waalkes 450, BO.

Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden, Meijer

B3820, B178 (BO!), ibid., on bark of Araucaria, ca 1400 m, very common,

Gradstein 10207 (BIOT! GOET!), ibid., near the guest house, on base of

Lejeuneaceae subfamily Ptychanthoideae 89

tree, Gradstein 10206 (BIOT! GOET!); Gn. Pangrango, Meijer B605 (BO!):

Gn. Gede, 1500-1900 m, Verdoorn 46e, 46d, 47c, Meijer B589a2 (BO!), ibid.,

N slope above “Artja”, ca 1100 m, Schiffner 287, 285 (BO!): Rawa Denok, ca

1900 m, Neervoort 2380 (BO!); Gn. Halimun, ca 930 m, collector unknown

(BIOT!): Gn. Cikurai, ca 1700 m, Verdoorn 59a, 59b,59c, 59d, 59e, 59f (BO!):

Geger Bentang, Neervoort 1366, 1089 (BO!); Gn. Malabar, Verdoorn 62c

(BO!). Banten, Pulau Panaitan, Gn. Putri, ca 75 m, van Borssum Waalkes

450 (BO!); Taman Nasional Ujung Kulon, Dewi Dw928, 922c (BO!).

Distribution: Java, Sumatra, Borneo, Sulawesi, Philippines, New Guinea,

Australia, Pacific, Japan, Taiwan, India.

Notes: Diagnostic characters of Spruceanthus polymorphus are: 1) rather

robust plant, ventral merophyte 8-12 cells wide, 2) underleaves serrate

towards the apex, 3) leaf lobes rounded to obtuse at apex, with entire or

toothed, plane or undulate margin, 4) leaf cells isodiametrical,5) female bract

lobe entire, female bracteole toothed 6) perianth with up to 7 smooth keels,

1-2 innovations present. This species grows at ca 75-1900 m. Spruceanthus

polymorphus may be confused with S. semirepandus and Archilejeunea

planiuscula. S. semirepandus differs by the acute leaf apex (rounded to

obtuse in S. polymorphus) and A. planiuscula by the thinner stems and the

segmented oil bodies (although we did not observe the oil bodies, they are

supposedly homogeneous in S. polymorphus).

22. Spruceanthus semirepandus (Nees) Verd., Ann. Bryol. Suppl. 4: 153

(1934): Jungermannia semirepanda Nees, Enum. PI. Crypt. Javae 1:39 (1830);

Ptychanthus semirepandus (Nees) Nees, Naturg. Eur. Leberm. 3: 212 (1838);

Phragmicoma semirepanda (Nees) Gottsche, in Gottsche ef al., Syn. Hepat.

302 (1845); Lejeunea semirepanda (Nees) Mitt., J. Proc. Linn. Soc. Bot. 5: 111

(1861). —Type: Indonesia. Java. Blume s.n. (holotype, STR?) — cf. Verdoorn

(1934).

Autoicous. Plants robust, up to 8 cm long, 1.5-3 mm wide, pale brown to dark

brownin dried condition. Branching Lejeunea-type.Stem diameter 120-300 um;

ventral merophyte 8-12 cells wide. Leaves imbricate, widely spreading. Lobe

ovate, 1000-2300 um long, 550-1590 um wide, margin entire, crenulate towards

the sharply acute apex: cells of lobe thick-walled, light yellow, hexagonal

and almost isodiametric, trigones not cordate, intermediate thickening

frequent; marginal cells 5-15 x 4-10 um, mid-leaf cells 20-31 x 13-20 um,

basal cells 30-41 x 21-30 um; oil bodies not seen. Lobule ovate, 240-450

um long, 120-220 um wide, inflated, apex truncate with 1-2 small teeth.

Underleaves imbricate, obdeltoid to rectangular, 500-1200 um long, 390-850

um wide, margin entire, apex truncate, base auriculate. Androecia not seen.

90 Gard. Bull. Singapore 62 (1) 2010

Gynoecia with 1-2 lejeuneoid innovations, bracts ovate about 3050 um long,

1530 um wide, margin toothed at the upper part, apex acute, bract lobule

oblong, margin entire about 1/3 of the lobe lenght; bracteole broadly ovate

to suborbicular with emarginate apex, toothed, about 2010 um long, 1500

wide. Perianth, oblong-ovate, about 2700 um long, 1600 um wide, with 7-

9 smooth keels, margins entire. Sporophytes and asexual reproduction not

seen.

Figure 22. Spruceanthus semirepandus (Nees) Verd. Shoot (a); underleaf (b); leaf lobe (c); leaf lobule

(d); female bract (e); female bracteole (f); cells of midleaf (g). Drawn from Verdoorn 20c, BO.

Specimens examined: INDONESIA. West Java, Cibodas Bot. Garden,

Verdoorn 20c, 20b (BO!); Gn. Gede, Jwamasa s.n. (BO!); Gn. Malabar, 1800-

2300 m, Verdoorn 61d (BO!); Gn. Cikurai, ca 1700 m, Verdoorn 59k (BO!);

Gn. Patuha, ca 1500 m, Kornochalert 1417 (BIOT!).

Distribution: Java, Borneo, Moluccas, Philippines, China, Taiwan, India, Sri

Lanka, Japan.

Notes: Diagnostic characters of Spruceanthus semirepandus are: 1) robust

plant, ventral merophyte to 12 cells wide, 2) underleaves with entire

margin, 3) leaf lobe acute at apex, coarsely dentate at margin, 4) leaf cells

isodiametric 4) female bract lobule entire, 5) perianth with 7-9 smooth

keels, innovation present. This species grows at 1200-2300 m. It is closely

related to S. polymorphus but the leaf apex in the latter species is broader,

rounded to obtuse. By its acute leaves S. semirepandus may be confused with

Lejeuneaceae subfamily Ptychanthoideae 9]

Ptychanthus striatus but the very different branching types (Lejeunea-type

in Spruceanthus, Frullania-type in Ptychanthus) readily separates the two.

23. Thysananthus convolutus Lindenb., in Gottsche et al., Syn. Hepat. 288

(1845).—Type: Indonesia. Java, unknown locality, collector unknown.., ex hb.

Lindenberg (isosyntype: G) — cf. Gradstein er al. (2002).

Plants up to 5 cm long, 0.8-1.2 mm wide, pale brown to dark brown in

dried condition. Branching Lejeunea-type. Stem diameter 100-280 um

Leaves closely imbricate, attached to the stem at an angle approx. 45-65°.

Lobe ovate with recurved apex, asymmetric, upper part of leaf distinctly

ventrad, 700-1150 um long, 600-810 um wide, margin toothed or entire, apex

rounded; cells of lobe thick-walled, sometimes nodulose, trigones cordate,

intermediate thickening frequent; marginal cells 7-20 x 6-11 um, mid-leaf

cells 22-40 x 11-13 um, basal cells 36-52 x 11-15 um: oil bodies not seen.

Lobule ovate, 250-320 um long, 110-130 um wide, inflated, apex truncate

with 1-2 teeth, first tooth consisting of 3-4 cells, second tooth small consisting

of 1 cell, sometimes without first tooth. Underleaves imbricate, suboblong,

500-720 um long, 290-430 um wide, margins usually toothed, apex truncate,

recurved, base not auriculate, insertion line shallowly curved. Generative

structures not seen.

400 pm

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Figure 23. Thysananthus convolutus Lindenb. Shoot (a): underleaves (b): leaf lobe (c): cell of

midleaf (d). Drawn from Jwamasa 5435, BO.

92 Gard. Bull. Singapore 62 (1) 2010

Specimens examined: INDONESIA. West Java, without locality, Teysmann

s.n. (BO!); Kampung Dawuan, 1390 m, Neervoort 898 (BO!); Cibodas Bot.

Garden, Pasir Sintek, ca 1410 m, Neervoort 3368 (BO!), ibid., on bark of

trees at garden entrance, abundant, Gradstein 10205, on bark of Araucaria,

Gradstein 10203 (BIOT!, GOET!); above Cibodas, trail to Cibeureum

waterfall, Lee & Nova Indri 45 (BIOT!, UKMB!); Cibeureum, ca 1600-1900

m, Verdoorn 30k, 30q, 20L, 21b, Iwamasa 5435, Neervoort 195, 308, Dadi &

Noerta 134, Meijer B3578, B3705, B3789, B3578, V. Schiffner 283, 282a (BO!),

N.S. Ariyanti 472 (BIOT!); Gn. Gede, boven Soekaboem1, ca 1500-1900 m,

Verdoorn 46c, 46b, 46a (BO!); Gede-Pangrango, Tugu, above G. Mas, along

JI. Mandalawangi, Meijer B3374, B3325, B387f, B607 (BO!); Geger Bentang,

1400-1540 m, Neervoort 1178, 1059, 1024, 1806, 1019, 988, 2987, 3005 (BO!);

Rawa Panjang, Neervoort 1166 (BO!); Cihoerang, ca 1380-1460 m, Neervoort

138, 2241, 2242, Meijer B3681, B3741 (BO!); Telaga Warna, Puncak Pass,

“Kratermuurtje”, Verdoorn 64f, 641, 64h, 64e (BO!); Gn. Halimun-Salak Nat.

Park, Gn. Kendeng, ca 1250 m, Radhiah Zakaria 221c (BIOT!); Gn. Malabar,

- SW slopes of Puncak Besar, 1800-2300 m, Verdoorn 61h (BO!).

Distribution: Java, Sumatra, Peninsular Malaysia, Borneo, Sulawesi,

Moluccas, Papua New Guinea, Philippines, Solomon Is.

Notes: Diagnostic characters of Thysananthus convolutus are: 1) asymmetric

leaf lobe with upper part ventrad and recurved, margin toothed or entire 2)

underleaves suboblong, margins usually toothed or crenulate, 3) lobule with

2 teeth. This species grows at ca 1300-2300 m. Forms with toothed leaves

may be confused with 7: spathulistipus but the leaves in the latter species are

symmetrical while those of 7? convolutus are asymmetric. Forms with entire

leaves may be confused with Mastigolejeunea but the dorsal epidermis cells

in Mastigolejeunea are larger than the inner stem cells, while in Thysananthus

the epidermis cells are not larger than the inner cells.

24. Thysananthus minor Verd.,in Rec. Trav. Bot. Neerl. 30: 231 (1933).—Type:

Indonesia. Sumatra, Brastagi, Petani Falls, 1930, Verdooorn s.n. (holotype,

FH) —cf. Verdoorn (1933).

Dioicous. Plants small, up to 1.5 cm long, 0.5-0.7 mm wide, brown to dark

brown in dried condition. Branching Lejeunea-type. Stem diameter 100-

130 um Leaves imbricate, convolute when dry, widely spreading. Lobe

oblong-ovate, 430-900 um long, 250-390 um wide, lateral margin irregularly

toothed toward the apex, apex acute; cells of lobe thick-walled, light yellow,

rhomboidal, trigones cordate, intermediate thickening frequent; marginal

cells 8-13 x 5-7 um, mid-leaf cells 13-29 x 13-17 um, basal cells 16-38 x 6-

Lejeuneaceae subfamily Ptychanthoideae 93

14 um; oil bodies not seen. Lobule ovate, 150-270 um long, 50-90 um wide,

inflated, apex obliquely truncate with small tooth, Underleaves closely

imbricate, oblong, 200-490 um long, 150-650 um wide, margin entire, apex

truncate to emarginate, toothed, insertion line shallowly curved. Androecia

not seen. Gynoecia with 2 lejeuneoid innovations, bract ovate, 970-1380 um

long, 440-550 um wide, toothed at apex and upper margin, apex acute, sinus

up to 1/2 of lobe length; bracteole oblong to broadly oblong about as long

as bracts or shorter, 500-670 um long, 350-410 um wide, margin serrulate,

coarsely serrate towards the apex, apex emarginate. Perianth, oblong, 1800

um long, 500 um wide, with 3 keels, margins serrulate in the upper part.

Sporophytes and asexual reproduction not seen.

Figure 24. Thysananthus minor Verd. Shoot (a); leaf lobe (b); underleaf (c); lobule (d); cells of

midleaf (e); cells of basal part of the lobe (f). Drawn from Neervoort 1018, BO.

Specimens examined: INDONESIA. West Java, Gn. Gede, above Cibodas,

Geger Bentang, ca 1520-1700 m, Neervoort 1112, 1018, 1088, Noerta &

Soekar 1139 (BO!); G. Halimun Nat. Park, ca 1000 m, Haerida 712 (BO!)

ibid., ca 1200 m, collector unknown (BIOT!); Gn. Cikurai, W slope, ca 1700

m Verdoorn 59g, 59h (BO!).

Distribution: Java, Sumatra, Papua New Guinea.

94 Gard. Bull. Singapore 62 (1) 2010

Notes: Diagnostic characters of Thysananthus minor are: 1) very small plant,

less than | mm wide, 2) leaf lobe irregularly toothed towards the apex, 3)

underleaves toothed towards the apex, 4) lobule with only | small tooth.

Thysananthus minor is new to Java. This species grows at ca 1000-1710 m

and is very similar to 7. spathulistipus but differs by the smaller size. Possibly

it is just a form of the latter species.

25. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst., Mem. N.Y. Bot.

Gard. 52:67 (1989); Jungermannia retusa Reinw et al., Acta Phys.-Med. Acad.

Caes. Leop. Carol. Nat. Cur. 12: 214 (1824); Ptychanthus retusus (Reinw. et

al.) Nees var. a, in Gottsche et al., Syn. Hepat. 292 (1845). —Type: Indonesia.

Java, Blume s.n. (holotype, STR; isotype, W) — cf. Gradstein et al. (2002).

—Thysananthus planus Sande Lac., Ned. Kruidk. Arch. 3,4: 419 (1854).—Type:

Indonesia. Java, Junghuhn s.n. (holotype, L; isotype, G) — cf. Gradstein et al.

(2002).

{ 200Hm

Figure 25. Thysananthus retusus (Reinw. et al.) B. Thiers & Gradst. Shoot (a); underleaves (b); leaf

lobe (c); leaf lobule (d); female bracteole (e); female bract (f); vitta (g); cells of the margin of the

leaf (h); cells of midleaf (i); perianth (j). Drawn from Haerida 511, BO.

Plants up to 2 cm long, 0.7-1 mm wide; pale green to brown in the dried

condition. Branching Lejewnea-type. Stem diameter 100-120 um. Leaves

imbricate, convolute when dry, widely spreading. Lobe ovate to oblong, 740-

840 um long, 350-550 um wide, margin entire, apex obtuse to acuminate;

cells of lobe thick-walled, hyaline to light yellow, quadrangular to hexagonal,

Lejeuneaceae subfamily Ptychanthoideae 95

trigones triangular, intermediate thickening scarce; marginal cells 6-8 x

3-5.5 um, mid-leaf cells 8-15 x 7-10 um, basal cells 13-19 x 5-8 um, vittae

consisting of 2-3 rows of rectangular cells, ending at 2/3 of the lobe length,

21-51 x 10-17 um; oil bodies not seen. Lobule oblong-ovate 290-310 um

long, 140-150 um wide, apex with | tooth consisting of 3-4 cells. Underleaves

imbricate, orbicular, obdeltoid to subrectanglar sometimes recurved

seemingly emarginate, 230-310 um long, 290-390 um wide, margins entire,

apex rounded, sometimes with scattered small teeth towards the apex.

Generative structures not seen.

Specimens examined: INDONESIA. West Java, Bogor Bot.Garden, W. Meijer

55dlI (BO!); Gn. Salak, ca 2000 m, Kurz s.n. (BO!); Gn. Gede-Pangerango

Nat. Park, Bodogol, Haerida 851, 811 (BO!, GOET!); G. Pancar, ca 400 m,

Schiffner 287 (BO!); Geger Bentang, ca 1620 m, Neervoort 1070 (BO!).

Distribution: Java, Philippines, West Irian, Papua New Guinea, Australia,

Pacific Islands.

Notes: Diagnostic characters of Thysananthus retusus are: 1) small plant with

flattened leaves, 2) leaf with a distinct vitta, and with isodiametric non-vitta

cells, 3) apical tooth of leaf lobule 3-4 cells long. This species grows at ca 400-

2500 m. By the vitta, the small, isodiametric leaf cells and the long, curved

lobule tooth T: retusus is a very distinct species that cannot be confused with

any other member of Ptychanthoideae.

26. Thysananthus spathulistipus (Reinw. et al.) Lindenb., in Gottsche et al.,

Syn. Hepat. 287 (1845); Jungermannia spathulistipa Reinw. et al., Acta Phys.-

Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 212 (1824). -Type: Indonesia.

Java, Bantam, Leback Mts., Blume s.n. (holotype, STR; isotypes, G, W) — cf.

Gradstein et al. (2002).

Autoicous. Plants robust, up to 3 cm long, 0.5-1 mm wide, brownish green to

dark brown in dried condition. Branching Lejeunea-type. Stem diameter 80-

180 um. Leaves imbricate, widely spreading. Lobe ovate-oblong, 820-1100

um long, 320-630 um wide, margin coarsely toothed towards the apex, apex

acute; cells of lobe thick-walled, light yellow, rhomboidal, trigones cordate,

intermediate thickening frequent, cell walls at the basal part sometimes with

orange color; marginal cells 10-20 x 7-10 um, mid-leaf cells 26-31 x 10-20

um, basal cells 26-41 x 10-15 um; oil bodies not seen. Lobule ovate, 240-330

um long, 40-90 um wide, inflated, apex truncate with | small tooth consisting

of 1 cell. Underleaves imbricate, spathulate, 310-910 um long, 350-480 um

wide, margin coarsely toothed towards the apex, apex truncate to sometimes

96 Gard. Bull. Singapore 62 (1) 2010

emarginate, insertion line almost straight. Androecia terminal on branches,

bracts in 4-15 pairs, similar to leaves but lobules larger, bract lobule 2/3 of

the bract lobe, bract lobe 270-370 um long, 150-160 um wide, hypostatic,

bracteoles similar to underleaves in size and shape, 170-180 um long, 200-230

um wide. Gynoecia on short or long branches, with 1-2 lejeuneoid innovations,

bract ovate 900-1000 um long, 300-400 tm wide, crenulate towards the apex,

apex acute, sinus up to 1/2 of lobe length; bracteole oblong to sub obovate

700-2200 um long, 250-650 um wide, margin crenate, apex truncate. Perianth

oblong, about 2000 um long, 550 um wide, with 3 keels, margins toothed.

Sporophytes and asexual reproduction not seen.

jb

Figure 26. Thysananthus spathulistipus (Reinw. et al.) Lindenb. Shoot (a); leaf lobe (b); underleaf

(c); female bracteole (d); female bract (e); cells of midleaf (f). Drawn from Haerida 725, BO.

Specimens examined: INDONESIA. West Java, Bogor Bot. Garden, ca 250

m, Schiffner 281 (BO!); Cibodas Bot. Garden, Verdoorn 20a, 20k, 301, 30p,

300, Neervoort 1506, 70 (BO!); Gn. Pangrango, Meijer B895, B775, B286,

B530, B447e, B519, B3339, B818c, B773a, B771a, B805, B81 1c, B81 1e, B385c,

B752b, B753, B533c, B287b, B298b, B801a (BO!); Gn. Gede, 1500-1900

m Verdoorn 46m, 46l, 46], 46h, 46g, 46f, Meijer B4093, Schiffner 291la, 289

(BO!); Gn. Gegerbentang, E slopes, 1500-2000 m, Verdoorn 67f 67e, 67g,

Neervoort 1139, 1144, 1985, 1022, 1014, Noerta & Soekar 1144, 50/1089,

Meijer B637a (BO!); Gn. Halimun Nat. Park, Gradstein s.n., Haerida 725

(BO!); Mt. Kendeng, ca 1060-1350 m, Radhiah Zakaria 173c, 181c, 218c,

226c (BIOT!); G. Malabar Verdoorn 61g, 61f, 6le, 61d (BO!); Gn. Guntur,

“Kawah Kamodjan”, 1500-1700 m, Verdoorn 50f, 50e (BO!); Gn. Cikurai, ca

1700 m, Verdoorn 591 (BO!).

Lejeuneaceae subfamily Ptychanthoideae 97

Distribution: Java, Sumatra, Peninsular Malaysia, Borneo, Bali, Soembawa,

Sulawesi, Moluccas, West Irian, Papua New Guinea, Australia, Solomon Is.,

Thailand, India, Sri Lanka, tropical Africa.

Notes: Diagnostic characters of Thysananthus spathulistipus are: 1)

leaves and underleaves symmetrical, with toothed margins, 2) spathulate

underleaves, 3) coarsely toothed female bracts and bracteoles. This species

grows at ca 200-2000 m and is most common in the mountains. It differs

from other Javanese species of Thysananthus by the rather large plant size,

the symmetrical, toothed leaves without vitta, the lobule with a short, blunt

tooth, and the often narrowly spathulate underleaves.

Discussion and conclusion

This study revealed the occurrence of 26 species of Lejeuneaceae subfam.

Ptychanthoideae in West Java, in 8 genera: Acrolejeunea (Spruce) Schiffn.

(3 species), Archilejeunea (Spruce) Schiffn. (1 species), Lopholejeunea

(Spruce) Schiffn. (10 species), Mastigolejeunea (Spruce) Schiffn. (3 species),

Ptychanthus Nees (1 species), Schiffneriolejeunea Verd. (2 species),

Spruceanthus Verd. (2 species) and Thysananthus Lindenb. (4 species).

Two species found in this study were new records of the Hepaticae in

Java: Mastigolejeunea indica and Thysananthus minor. The record of

Mastigolejeunea indica is based on a specimen collected by the Dutch

Bryologist, W. Meijer,in Bogor Botanical Garden in 1951 and in Meru Betiri

National Park, East Java, in 2005. Five additional species recorded from Java

in the literature (Acrolejeunea tjibodensis Verd., Dendrolejeunea fruticosa

(Lindenb. & Gottsche) Lacout., Phaeolejeunea latistipula (Schiffn.) Mizut.

[doubtful record], Spruceanthus sulcatus (Nees) Gradst. and Thysananthus

comosus Lindenb.) were not found in this study.

Based on the number of known localities in West Java the species

of Ptychanthoideae can divided into several categories. Species with “wide

distribution” are known from more than 5 localities, with “moderately wide

distribution” from about 4, with “rather limited distribution” from about

3, with “limited distribution” from about 2, and “very limited distribution”

from only one locality.

Lopholejeunea eulopha, Lopholejeunea ceylanica, Lopholejeunea

subfusca and Thysananthus spathulistipus are the most common species in

West Java, with a wide distribution. Thysananthus spathulistipus (200-1700

m) was found in Bogor Bot. Garden, Cibodas, Gede Pangrango Nat. Park

and Halimun Salak Nat. Park, and on Mt. Guntur, Mt. Cikurai and Mt.

Malabar; Lopholejeunea eulopha (200-1700 m) in Bogor Bot. Garden, Ujung

98 Gard. Bull. Singapore 62 (1) 2010

Kulon Nat. Park and Halimun Salak Nat. Park, and on Mt. Guntur and Mt.

Megamendung; L. ceylanica (500-1700 m) in Bogor Bot. Garden, Cibodas,

Gede Pangrango National Park, Telaga Warna and on Mt. Cikurai; and L.

subfusca (500-2500 m) in Bogor Bot. Garden, Cibodas, Gede Pangrango

National Park and on Mt. Cikurai and Mt. Papandayan.

Lopholejeunea nigricans, Lopholejeunea horticola and Thysananthus

retusus have a moderately wide distribution. Lopholejeunea nigricans (200-

1700 m) has been recorded from Bogor Bot. Garden, Mt. Pangrango, Mt.

Guntur and Telaga Warna (Puncak Pass); L. horticola (800-2400 m) from

Mt. Gede (including Cibodas), Mt. Guntur, Mt. Patuha and Mt. Malabar;

and Thysananthus retusus (400-2500 m) from Bogor Bot. Garden, Mt. Gede,

Mt. Halimun Salak Nat. Park and Mt. Pancar.

Archilejeunea planiuscula, Lopholejeunea herzogiana, L. wiltensii,

Mastigolejeunea virens, Ptychanthus striatus, Spruceanthus semirepandus,

Thysananthus convolutus and Thysananthus minor have a rather limited

distribution in West Java. Archilejeunea planiuscula (200-1450 m) was found

- in Ujung Kulon Nat. Park and on Mt. Gede; Lopholejeunea herzogiana

(1200-1500 m) in Bogor Bot. Garden, Gede Pangrango Nat. Park and Telaga

Warna (Puncak Pass); L. wiltensii (1200-1900 m) in Gede Pangrango Nat.

Park and on Mt. Guntur; Mastigolejeunea virens (200-1500 m) in Bogor

Bot. Garden and on Mt. Gede and Mt. Megamendung; Ptychanthus striatus

(1000-2400 m) in Gede Pangrango Nat. Park and on Mt. Megamendung;

Spruceanthus semirepandus (1200-2300 m) on Mt. Gede, Mt. Cikurai and

Mt. Malabar; Thysananthus convolutus (1300-2300 m) on Mt. Gede, Telaga

Warna (Puncak Pass) and Mt. Malabar; and T. minor (1000-1700 m) on Mt.

Gede, in Mt. Halimun Salak Nat. Park and on Mt. Cikurai.

Acrolejeunea fertilis, A. pycnoclada, Lopholejeunea_ applanata,

Mastigolejeunea auriculata and Schiffneriolejeunea tumida var. haskarliana

were found in an even more limited distribution. Acrolejeunea fertilis was

only found in the lowlands (60-80 m) in Pulau Panaitan and Depok; A.

pycnoclada only at Mt. Gede and Ciater (Subang), in rather high altitude

(1500-1900 m) although the species has also been recorded from lowlands

(e.g. Bogor Bot. Garden) in the literature; Schiffneriolejeunea tumida var.

haskarliana (440-1500 m) only from Mt. Gede and Mt. Halimun Salak Nat.

Park; Lopholejeunea applanata (ca 1200 m) only from Gede Pangrango

Nat. Park; and Mastigolejeunea auriculata (200-500 m) only from Bogor

Bot. Garden and Ujung Nat. Park. This condition may be due to inadequate

collections from the area.

Acrolejeunea arcuata, Lopholejeunea recurvata, L. zollingeri, Mastigolejeunea

indica, and Schiffneriolejeunea pulopenangensis, finally, had the most limited

distribution and were found in only one location, i.e. Acrolejeunea arcuata

only on Mt. Patuha at 2000 m, Lopholejeunea recurvata on Mt. Gede at ca

Lejeuneaceae subfamily Prychanthoideae 99

1450 m, L. zollingeri on Mt. Pangrango at ca 1600 m, and Mastigolejeunea

indica and Schiffneriolejeunea pulopenangensis only in Bogor Bot. Garden

at about 200 m. The records of the latter two species only from Bogor

Botanical Garden emphasize the importance of this garden as a habitat for

Ptychanthoideae in West Java. A searching for additional localities of the

rare species in West Java is needed.

According to Gradstein (1991) the endemic genera of Asiatic

Ptychanthoideae are largely restricted to the subtropical and temperate

areas of Asia and some are also known as fossils in Eocenic amber of Europe.

They are considered to be palaeoendemic, the relictual groups. Endemic

genera of Asiatic Lejeuneoideae, however, occur mainly in the tropical rain

forests of the Malesian archipelago, are often highly specialized, and are

lacking in the fossil record. They probably have co-evolved in the Tertiary

with the rain forest and are to be considered neoendemics.

Ptychanthoideae seem to be older than Lejeuneoideae and may

already have existed in the Mesozoic before the break-up of Laurasia and

Gondwanaland. The Mesozoic age was recently confirmed based on fossil

evidence and DNA sequence analysis by Wilson er al. (2007), who found

that Lejeuneaceae started to diversify in the Late Cretaceous, about 60-90

million years ago.

The geographical distribution and altitudinal ranges of Ptychanthoideae

in West Java are shown in Table 1; the definition of the distribution types

follows Ariyanti and Gradstein (2007). The geographical ranges of the

species were determined based on collected specimens and literature (e.g.,

Mizutani, 1961; Gradstein and Terken, 1981; Menzel 1988; Gradstein er al.,

2002; Zhu and Gradstein, 2005: Aniyanti and Gradstein, 2007).

The data show that the species of Ptychanthoideae of West Java can be

subdivided into 4 groups by their geographical distributions: Malesian species

(8 spp.), tropical Asiatic species (10 spp.), palaeotropical species (tropical

Asia + Africa; 3 spp.) and pantropical species (throughoput the tropics: 4

spp.). It appears that the species are rather widespread; none of the species

are endemic to Java or western Malesia. The widespread distribution of the

species is probably due to their dispersal by spores, which may be easily

carried by the wind over long distances (van Zanten and Gradstein, 1987).

But also the rather old age of Ptychanthoideae (Gradstein, 1991: Wilson er

al., 2007) may play a role.

The majority of the species (about 20) are found at mid-montane

elevations, at 1200-1500 m. Few species found below 100 m and above 2000

m. The data from West Java agree with the general pattern of altitudinal

distribution reported for Lejeuneaceae. According to Gradstein (1995),

the diversity of Lejeuneaceae decreases with elevation and accounts

for about 45% of total hepaticae diversity in the lower montane forest

100 Gard. Bull. Singapore 62 (1) 2010

Table 1. Altitudinal and geographical distributions of the species of Ptychanthoideae

recorded in West Java. Lowland: 0-1200 m. Montane: 1200-3000 m. A: Asiatic. M: Malesian. P:

Pantropical. Pal: Palaeotropical (Asia, Africa). * species new to Java.

Altitudinal — distribution Geographical

Nip Species Lowland Montane distribution

1. | Acrolejeunea arcuata - + M

2. Acrolejeunea fertilis + - M

3 Acrolejeunea pycnoclada + + Pal

4 Archilejeunea planiuscula ~ + A

> Lopholejeunea applanata - . A

6. | _Lopholejeunea eulopha + + P

7 Lopholejeunea herzogiana - Ta M

8 Lopholejeunea nigricans + + P

J) Lopholejeunea ceylanica + + A

10. Lopholejeunea horticola + a A

11. Lopholejeunea recurvata + + M

12. Lopholejeunea subfusca + + 2

13. Lopholejeunea wiltensii - + M

14. Lopholejeunea zollingeri - + A

15. Mastigolejeunea auriculata + + Je

16. | Mastigolejeunea indica* + - M

17. Mastigolejeunea virens + + M

18. Ptychanthus striatus +: + Pal

19. Schiffneriolejeunea

pulopenangnesis oF : M

20. — Schiffneriolejeunea tumida var.

haskarliana - a A

21. Spruceanthus polymorphus + + A

22. Spruceanthus semirepandus - + A

23. Thysananthus convolutus - + A

24. = Thysananthus minor* + + M

25. Thysananthus retusus + + A

u

=

26. Thysananthus spathulistipus + +

Lejeuneaceae subfamily Prichanthoideae 101

(1000/1400-2000/2500 m). 30% in the upper montane forest (2000/2500-

3000/4000 m) and 20% in the subalpine forest (above 3000/4000 m).

The members of the Ptychanthoideae of West Java are epiphytes and

grow on the bark of trees. on treelets and shrubs. As indicated by Thiers

and Gradstein (1989) and Gradstein et al. (2001). many species are rather

xerotolerant epiphytes of trees at forest margins or in rather open vegetation.

None of the species of West Java were found growing on living leaves. which

are generally inhabited by tiny members of the subfamily Lejeuneoideae.

and few species occur on small branches of shrubs. which are the habitat

of the ramicolous bryophytes. Dendroid. feather or bracket-type mosses

and liverworis. belonging to Neckeraceae, Hookeriaceae, Pterobryaceae.

and Plagiochilaceae. as well as various tiny members of Lejeuneaceae are

the specialists of this habitat (Gradstein and Pocs, 1989). A study of the

ecology and optimum habitat conditions of the members of Lejeuneaceae

subfam. Ptychanthoideae, including moisture and temperature rates,

would be desirable. Such data might further improve our understanding

of the distribution of the species of Ptychanthoideae in West Java.

Acknowledgements

The first author is grateful to the ~Program Karyasiswa Dalam Negeri LIPI”

for financial support of her study. She also thanks Dr. Johanis Palar Mogea

for his valuable comments on the manuscript. Furthermore. she gratefully

acknowledges the support of the Directors of Herbarium Bogoriense and

Herbarium Biotrop for providing facilities to conduct her research.

References

Anriyanti, N.S. and S.R. Gradstein. 2007. Wallace’s line and the distribution of

the liverworts of Sulawesi. Cryptogamie. Bryologie 28: 3-14.

Gradstein, S.R. 1975. Monograph of the genus Acrolejeunea. Bryophytorum

Bibliotheca 4- 1-216.

Gradstein, S.R. 1991. Diversity and distribution of Asian Lejeuneaceae

subfamily Ptychanthoideae. Tropical Bryology 4: 1-16.

Gradstein. S.R. 1994. Verdoorm’s Studien iiber Asiatische Jubulae. Hikobia 11:

451-456.

102 Gard. Bull. Singapore 62 (1) 2010

Gradstein, S.R. 1995. Diversity of Hepaticae and Anthocerotae in montane

forests of the tropical Andes, pp. 321-334. In: Churchill, S.P., J. Luteyn, E.

Forero and H. Balslev (eds.), Biodiversity and Conservation of Neotropical

Montane Forests. New York Botanical Garden, New York.

Gradstein, S.R., S.P. Churchill and N. Salazar-Allen. 2001. Guide to the

bryophytes of Tropical America. Memoirs of the New York Botanical

Garden 86: 1-157.

Gradstein, S.R., X.-L. He, S. Piippo and M. Mizutani. 2002. Bryophyte flora

of the Huon Peninsula, Papua New Guinea. LX VIII. Lejeuneaceae

subfamily Ptychanthoideae (Hepaticae). Acta Botanica Fennica 174: 1-

88.

Gradstein, S.R. and T. Pocs. 1989. Bryophytes, pp. 311-325. In: Lieth, H. and

M.J.A. Werger (eds.), Tropical Rain Forest Ecosystems. Elsevier Science

Publishers, Amsterdam.

Gradstein, S.R. and L.Terken. 1981. Studies on Lejeuneaceae subfam.

Ptychanthoideae VI. A revision of Schiffneriolejeunea sect. Saccatae

from Asia. Occasional Papers of the Farlow Herbarium of Cryptogamic

Botany 16: 71-81.

Hasan, M. and N.S.Ariyanti. 2004. Mengenal Bryophyta (Lumut) Taman

Nasional Gunung Gede Pangrango. Vol. 1. BalaiTaman Nasional Gunung

Gede Pangrango, 93 pp.

Menzel, M. 1988. Annotated Catalogue of the Hepaticae and Anthocerotae

of Borneo. Journal of the Hattori Botanical Laboratory 65: 145-206.

Mizutani, M. 1961. A revision of Japanese Lejeuneaceae. Journal of the

Hattori Botanical Laboratory 24: 116-180.

Mizutani, M. 1969. Lejeuneaceae subfamily Ptychanthoideae from Sabah

(North Borneo). Journal of the Hattori Botanical Laboratory 32: 129-

139:

Nees von Esenbeck, C.G. 1830. Enumeratio Plantarum Cryptogamicarum

Javae et Insularem Adjacentium. I. Hepaticas complectens. Breslau.

Piippo, S, T. Koponen and D.H. Norris. 1987. Endemism in the Bryophyte

Flora in New Guinea. Symposia Biologica Hungarica 35: 361-372.

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Reinwardt, C., C. Blume and C.G. Nees von Esenbeck. 1824. Hepaticae

javanicae. Acta Physico-Medica Academiae Caesareae Leopoldino-

Carolinae Naturae Curiosorum 12: 181-238.

Sande Lacoste, C.M. 1856. Synopsis Hepaticarum Javae. Verhandelingen der

Koninklijke Nederlandse Akademie van Wetenschappen 5: 1-112.

Schiffner, V. 1898. Conspectus Hepaticarum Archipelagi Indici. Batavia

Staatsdruckerei. 382 pp.

Schuster, R.M. 1980. Phylogenetic studies on Jungermanniidae IT. Radulineae

(Part I). Nova Hedwigia 32: 637-693.

Thiers, B.M. and S.R. Gradstein. 1989. Lejeuneaceae (Hepaticae) of

Australia. I. Subfamily Ptychanthoideae. Memoirs of the New York

Botanical Garden 52: 1-79.

Verdoorn, F. 1933. Die von V. Schiffner (1893-1894) und von Fr. Verdoorn

(1930) auf den Indomalesischen Inseln gesammelten Lejeuneaceae

Holostipae. De Frullaniaceis XI. Recueil des Travaux Botaniques

Néerlandais 30: 212-233.

Verdoorn, F. 1934. Studien tiber Asiatische Jubuleae. Annales Bryologici

Supplement 4: 1-231.

Wilson, R., J. Heinrichs, J. Hentschel, S.R. Gradstein and H. Schneider. 2007.

Steady diversification of derived liverworts under steady Tertiary climatic

fluctuations. Biology Letters 3: 566-569.

Zanten,B.O.van,andS.R.Gradstein. 1987.Experimental dispersal geography

of tropical liverworts. Nova Hedwigia Beiheft 90: 41-94.

Zhu, R.-L. and S.R. Gradstein. 2005. Monograph of Lopholejeunea

(Lejeuneaceae, Hepaticae) in Asia. Systematic Botany Monographs TA:

1-98.

2 ass -

Gardens’ Bulletin Singapore 62 (1): 105-109. 2010 105

Phaius takeoi (Orchidaceae) Newly Recorded from

Thailand and Myanmar

H. KURZWEIL .S.WATTHANA AND S.LWIN®>

Herbarium, Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569

Queen Sirikit Botanic Garden, The Botanical Garden Organization.

, P.O. Box 7, Mae Rim, Chiang Mai 50180, Thailand

~ Myanmar Floriculturist Association, Alone Road, Yangon, Myanmar

Abstract

The occurrence of Phaius takeoi (Hayata) H.J. Su (Orchidaceae), previously

only known in mainland China, Vietnam and Taiwan, is here reported for

Thailand and Myanmar.

Introduction

In the course of floristic and taxonomic work on the orchids of Thailand and

Myanmar three collections of a yellow and green-flowered Phaius species

were made which did not match any currently known species (Seidenfaden,

1986; Kress et al.,2003; Vaddhanaphuti, 2005, Kurzweil, 2010 ). The specimens

are now positively identified as P. takeoi (Hayata) H.J. Su, currently only

known in mainland China, Vietnam and Taiwan. The new record of this

species in Thailand and Myanmar is reported below.

Phaius takeoi (Hayata) H.J. Su

Quart. J. Exp. Forest Natl. Taiwan Univ. 3 (1989) 77; Chen er al., Fl. China

vol. 25: 290 (2009): Su, in Digital Flora of Taiwan, http://www.efloras.org

florataxon.aspx/flora_id =100&taxon_id =242413999, accessed January 12,

2010). —Basionym: Calanthe takeoi Hayata, Icon. Pl. Formosan. 9 (1920) 111.

—Typus: Taiwan, Toyencho, Urai, Oct 1917, Takeo Ito s.n. (not found). Plate 1.

—Phaius longicruris Z.H.Tsi, Acta Phytotax. Sin. 19 (1981) 505; Averyanov

and Averyanova, Turczaninowia 5 (2002) 93. —Typus: China, Yunnan,

Jinghong, in forest, 1400 m alt., Oct 1936, Wang 79184 (holotype, PE).

Terrestrial herbs, vegetative parts entirely glabrous, basally pseudobulbous, to

1 m tall. Roots to 3 mm in diameter. Pseudobulbs elongate-clavate, 20-40 cm

long and (1.3-)2-2.5 cm in diameter, slightly tapering upwards, with 3-5 nodes,

internodes 2.5-10.7 cm long, green or dark green with green veins, glabrous,

106 Gard. Bull. Singapore 62 (1) 2010

Plate 1. Phaius takeoi (Hayata) H.J. Su. A. Plant; B. Part of inflorescence; C & D. Flowers. All

from Watthana 3287 (Thailand, Tak); D flower photo from Nyan Tun s.n. (Myanmar, Shan

State). Bars: A. 10 cm. B-D.3 cm.

Phaius takeoi (Orchidaceae) 107

covered by pale green sheaths. Leaves 5-8, arising from the apical portion of

the pseudobulbs, elliptic, elliptic-lanceolate or elliptic-oblong. (5.5-)12-40 x

(2.5-)5.5-10(-14) cm, acuminate or long-acuminate, margin entire and slightly

undulate, with 5-7 veins, green above, pale green underneath. Inflorescences

arising from the lower nodes of the pseudobulbs, erect. up to 60 cm long

and 0.4-0.8 cm in diameter, normally racemose but once observed with a

side branch (in the specimen Watthana 3287), glabrous below, glabrous or

sparsely hairy on the uppermost part of the rachis. Peduncles 30-50 cm long,

with up to 9 sterile bracts 16-28 mm long. Rachis 10-13 cm long (rachis in all

three examined specimens still elongating), with 4-15 flowers. Floral bracts

persistent, ovate-lanceolate or narrowly lanceolate, the lower ones to 17 x

3.8 mm, acute, glabrous. Flowers 4-5 cm in diameter, opening widely: sepals

and petals pale green or yellowish green and sometimes darker at the apex.

lip white or cream and sometimes yellow-tinged. Pedicel plus ovary 2-3.2 cm

long. sparsely pubescent or nearly glabrous. Sepals subequal. elliptic-oblong.

obtuse or acute, 27-32(-35) x (7-)8-10.5 mm, 5-7-veined, abaxially sparsely

hairy: laterals slightly oblique, somewhat decurved. Petals oblanceolate or

oblanceolate-oblong, sometimes basally indistinctly clawed, obtuse or acute.

28-33 x 6-8 mm, 3-veined. Lip nearly orbicular, 30-34 x 24-26 mm (excluding

spur), shallowly 3-lobed, glabrous for the most part but basal portion and

inside of spur hairy, basally united with the column for 7-8 mm: midlobe

orbicular-square or suborbicular, apex emarginate and sinus sometimes

mucronate, margins undulate: side lobes ovate or elongate-ovate, strongly

incurved and forming a wide open tube to embrace the column, apex broadly

rounded, margins undulate: disc with three faint yellow and smooth ridges,

the median one extending from the base of the lip to just below the apex of

the midlobe, the lateral ones arising from the middle part of the lip to the

base of the midlobe, spur elongate-conical with broad entrance, clavate, 6-

9 mm long, greenish yellow. Column stout, 15-25 mm long, 6-7 mm broad,

white, hairy on the front face below the stigma and continually into the spur.

apex widened with very prominent lateral appendages.

Specimens examined: THAILAND. Northern Region. Phitsanulok Province.

Phu Soi Dao, Sep 2008, RMK 671/51 (QBG, QBG spirit): Northern Region.

Tak Province, Umphang, Jul 2009, Watthana 3287 (QBG). MYANMAR.

Shan State, Taunggyi, Jul to Aug 2009, Nyan Tun s.n. (SING).

Habitat and flowering time: This species is found in moist, broad-leaved

primary or secondary forest. The Thai specimens were found in a marshy

spot in evergreen forest, while the Myanmar specimen was collected in moist

soil near streams. Populations were quite sizeable in the Myanmar plant

where several dozen individuals were found on a hill slope. but very small

108 Gard. Bull. Singapore 62 (1) 2010

with under 20 individuals in the two Thai specimens’. Altitudes were not

recorded in the Thai and Myanmar specimens reported here, but elsewhere

the species is found at 500-1400 m (Averyanov and Averyanova, 2002; Tsi,

1981: Chen et al.,2009; Su, no date, Digital Flora of Taiwan). Flowering occurs

between July and September, and has elsewhere been reported between

October and December. In Taiwan and Vietnam the species is said to be

uncommon (Averyanovy and Averyanova, 2002; Su, no date, Digital Flora of

Taiwan).

Distribution: China (Yunnan), Taiwan, Vietnam, Thailand, Myanmar.

Notes: Among the Thai and Myanmar Phaius congeners, this species is

unmistakable with its habit and the yellow-green flowers.

Acknowledgements

- We would like to acknowledge the Queen Sirikit Botanic Garden for

assistance during fieldwork and the Rom Klao Botanical Garden for

providing one of the specimens examined. We would also like to thank

the collector of the Myanmar specimen, Mr. Nyan Tun, for providing

information, and the Myanmar Forest Department for issuing a CITES

permit to transport the plant to Singapore for identification. Dr. Jin Xiaohua

is thanked for information on the type specimen of Phaius longicruris.

References

Averyanov,L.andA.Averyanova.2002.Rare species of orchids (Orchidaceae)

in the flora of Vietnam. Turczaninowia 5: 49-108.

Chen S-C, Liu Z-J, Zhu G-H, Lang K-Y,Tsi Z-H, Luo Y-B, Jin X-H, P.J. Cribb,

J.J. Wood, S.W. Gale, P. Ormerod, J.J.Vermeulen, H.P. Wood, D. Clayton

and A. Bell. 2009. Orchidaceae. In: Wu. Z-Y., P-H. Raven and Hong. D.-

Y. (eds.), Flora of China, vol. 25, pp. 1-566. Science Press, Beijing, and

Missouri Botanical Garden Press, St. Louis.

Kress, W.J., R. DeFilipps, E. Farr and Yin-Yin-Kyi. 2003. A checklist of the

trees, shrubs, herbs and climbers of Myanmar. Contributions from the US

National Herbarium 45: 1-590.

Kurzweil, H. 2010. A precursory study of the Calanthe group (Orchidaceae)

in Thailand. Adansonia, sér. 3, 32: 57-107

Phaius takeoi (Orchidaceae) 10°

Su. H-J. Not dated. Phaius. In: Digital Flora of Taiwan. Published on the

internet at http://www.efloras org/florataxon.aspx/flora_id=100&taxon_

id=242413999_ accessed January 12.2010.

Seidenfaden, G. 1986. Orchid genera in Thailand XIII. Thirty-three

epidendroid genera. Opera Botanica 89: 1-216.

Tsi, Z-H. 1981. New species of Orchidaceae from China. Acta

Phytotaxonomica Sinica 19: 505-510.

Vaddhanaphuti, N. 2005. A field guide to the wild orchids of Thailand, ed. 4.

Silkworm Books. Chiang Mai.

5 =z

Gardens’ Bulletin Singapore 62 (1): 111-117. 2010 Wl

Curcuma vitellina (Zingiberaceae), a New Species

from Vietnam

J. LEONG-SKORNICKOVA |, TRAN H.D.” AND M.E NEWMAN’:

‘The Herbarium, Singapore Botanic Gardens

5 1 Cluny Road, 259569 Singapore

University of Science, Vietnam National University

227 Nguyen Van Cu, QO5, Ho Chi Minh City, Vietnam

* Royal Botanic Garden Edinburgh

20A Inverleith Row, Edinburgh EH3 5LR, Scotland.

Author for correspondence: jana_skornickova@seznam.cz

Abstract

Curcuma vitellina,anew species of Zingiberaceae from Tay Nguyén, Vietnam,

is described, illustrated and compared to its closest ally, C. pierreana.

Introduction

Cambodia, Laos and Vietnam reportedly form a diversity hotspot for the

family Zingiberaceae, although detailed data are lacking because the most

recent comprehensive account of the family there is over a century old

(Gagnepain, 1908). The main centres of diversity of the genus Curcuma

are usually said to be India, Burma and Thailand (e.g. Leong-Skorniékova

et al. 2008), but our recent explorations of Zingiberaceae for the Flora of

Cambodia, Laos and Vietnam indicate that these countries are at least as

rich in Curcuma and other Zingiberaceae.

Three Curcumaspecies, C. bicolor Mood & K.Larsen,C. glans K. Larsen

& Mood from Thailand and C. rhomba Mood & K. Larsen from Vietnam

were described in 2001 (Mood & Larsen 2001). As a revision of Curcuma in

Cambodia, Laos and Vietnam progresses,specimens collected at two localities

in Vietnam have been shown to represent a new species which is described

and illustrated here. Like the species described by Mood and Larsen, this

new one lacks a clear distinction between fertile and coma bracts, but differs

in overall shape and coloration of the flower as well as anther morphology.

Curcuma vitellina Skorniék. & H. D. Tran, sp. nov.

Curcumae pierreanae comae absentia, bractearum formae, anthera basi

calcaribus filamentaceis 2 similis, sed rhizomate ramis lanceolatis verticaliter

fasciculatis (contra rhizomate ramis horizontaliter repentibus), corollae lobis

ee Gard. Bull. Singapore 62 (1) 2010

dilute luteis (contra albis), labello staminodtisque lutee aurantiacis (contra

albis vel albis apicibus profunde rosee purpurascentibus), lamina tenui

valide plicata basi rotundata (contra coriacea vents principalibus minus

prominentibus, basi cuneata ad attenuata) differt. -Typus: Vietnam, Lam

Dong Prov., Pongour waterfall; 11° 41’ 07.0” N, 108° 16’ 06.1” E; 787 m;

23 Jun 2008, Tran et al. 70 (holotype, SING incl. spirit; isotype, E, P, VNM,

National University of Laos).

Rhizome ovoid, ca 2.5-5 x 1.5-3 cm, with lanceolate to narrowly lanceolate

branches held upright, 3-8 x 1-1.5 cm, brown externally, light yellow internally,

slightly aromatic, root tubers elliptic, 2-4 cm long, light brown externally,

cream white internally, at 5-15 cm from rhizome. Pseudostem to 15 cm long,

green, composed of leaf sheaths and enclosed by 2 sheathing bracts, ligule to

5 mm long, bilobed, hyaline, greenish white, translucent, turning papery with

age, hairy at the apex, hairs ca 0.3 mm long; leafy shoot to 70 cm tall with up to

5-6 leaves when flowering; petiole 5-20 cm long (petiole of first leaf shortest,

innermost leaves longest), green, glabrous; lamina elliptic to elliptic-ovate,

20-45 x 8-15 cm, glabrous on both surfaces, prominently plicate, adaxially

bright green, shiny, abaxially lighter green, shiny; midrib glabrous, green;

base rounded, margin hyaline, translucent white, ca 0.5 mm wide, glabrous;

apex acute, shortly hairy. Inflorescence central, many flowered. Peduncle 4-

20 cm long, up to 1 cm diam., greenish-white, puberulent, embedded within

pseudostem. Spike 8-15 cm long, ca 4-5 cm diam. at the middle, without

coma. Fertile bracts 15-60, larger at the base of the inflorescence, ca 3.5-4.5

x 2.5-3.5 cm, ovate to trullate, smaller and ovate at the apex, cream white or

pale greenish, sometimes with slight pinkish tinge, both sides shortly densely

pubescent, connate in the lower 1/3 to 1/4. Cincinni with 4-6 flowers at the

base of the inflorescence, 2-3 flowers at the top. Bracteoles one per flower,

ovate, boat-shaped, ca 7 x 4 mm to 15 x 7 mm (outer ones larger, inner

ones are gradually smaller), hyaline, translucent white, glabrous, but for the

apex, upper part and margins sparsely hairy. Flowers 5-5.5 cm, exserted from

bracts. Calyx ca 17 mm long, teeth 3, unilaterally split ca 7 mm, translucent

white, sparsely hairy on the three main veins leading from the tooth to the

base. Floral tube ca 3 cm long, narrowly cylindrical at base for ca 2 cm above

the ovary, funnel-shaped at apex, externally white turning pale yellowish

towards the apex, with glandular hair, internally white with dorsally placed

groove holding the style: dorsal corolla lobe ca 20 x 11 mm, triangularly

ovate, concave, glabrous, cream white with pale yellowish apex, apex

mucronate, mucro less than 1 mm long with a few short hairs; lateral corolla

lobes ca 18 x 9 mm, triangular with a rounded, slightly concave apex, cream

white with pale yellowish apex, glabrous. Lateral staminodes obovate, ca 18

x 11 mm, light yellow at base, yellow-orange towards the apex, glandular

Curcuma vitellina (Zingiberaceae) from Vietnam 113

hairs present on the raised middle portion facing the centre of the flower.

Labellum ca 21 x 20 mm, obscurely trilobed, lateral lobes folding upwards,

middle lobe emarginate with an incision up to 8 mm long, which splits at the

apex of the lobe as flowering progresses, cream white at base, yellow at apex

with deep yellow-orange band running through the centre (golden median

band). Filament 4-6 mm long, pale yellowish, 4.5 mm at base, 2 mm at apex,

with glandular hair at the back. Anther spurred, connective densely covered

with short glandular hairs, anther spurs ca 0.5 mm long, filamentous, cream

white, anther crest present, 1-1.5 x ca 2 mm, deep yellow, anther thecae 5

mm long, white, dehiscing along their whole length. Style white, glabrous,

stigma ca 1 mm wide, white, ostiole facing upwards, ciliate. Epigynous glands

two, cream, 4 x 0.8 mm, with blunt apex. Ovary 3 x 2 mm, trilocular, hairy,

hairs ca 0.2 mm long. Fruits not seen.

Flowering: June to August.

Distribution & habitat: So far known only from two localities (Bao Loc Pass

and Pongour Waterfall) in Lam D’6ng province, Tay Nguyén, Vietnam. It

grows among rocks in open and semi-open shrubby vegetation.

Vernacular names & uses: None so far recorded.

Etymology: Medieval Latin vitellinus, from Latin vitellus, egg yolk, referring

to the deep yellow colour of the flowers.

Other specimens examined: Among the vast amount of Curcuma herbarium

material revised and digitised by the first author, we have found only one

collection from Pongour (24 Aug 1924, Evrard 1193 x 2, P), which can be

identified with certainty as C. vitellina. There are a few other specimens

collected within a radius of 100 km of Pongour, which might represent C.

vitellina (e.g. Poilane 30582, P and Poilane 5031 x 2, P), but the condition

of the specimens and poor accompanying notes do not allow a definite

identification to be made.

IUCN preliminary assessment: Endangered (E): B2 ab (iii). The area of

occupancy is estimated to be less than 500 km. The species is known to

exist at only two locations with continuing decline in extent and quality of

habitat, which is disturbed by erosion and development for tourism.

114 Gard. Bull. Singapore 62 (1) 2010

Figure 1. Curcuma vitellina, sp. nov. A. Habit; B. Inflorescence; C. Flower (front view): D.

Detail of anther (front and side view); E. Liguie; F Rhizome and base of leafy shoot: G.

Dissected flower. Photographs by J. Leong-Skorniékova taken from Tran et al. 70.

Curcuma vitellina (Zingiberaceae) from Vietnam MNS

Figure 2. Curcuma pierreana Gagnep. A. Habit; B. Inflorescence; C. Anther (front and side

view) Photographs by J. Leong-Skorniékova from Tran et al. 26.

Notes: It is expected that Curcuma vitellina will be a seed-setting species as

the presence of young seedlings in natural populations has been observed.

Curcuma vitellina is similar to C. pierreana by its inflorescence composed

of cream or greenish flower bracts (which may be tinged pink), reflexed

at the tips and lacking a distinct coma (the inflorescence of C. vitellina is

more robust and with more bracts than that of C. pierreana). The anthers

of both species have a well-developed crest and two filamentous spurs at

the base, but the overall shape of the anther is different (see Figs. 1-3). The

two species differ in their rhizomes, the branches of which are narrowly

116 Gard. Bull. Singapore 62 (1) 2010

Figure 3. Anthers in side, front, and back view. A. Curcuwma vitellina (Tran et al. 70); B.

Curcuma pierreana (Tran et al. 26). Photographs by H.D. Tran.

lanceolate and vertically clustered in C. vitellina, but creep horizontally in

C. pierreana. The lamina in C. vitellina is thin, prominently plicate, glabrous

on both sides and has a rounded base, whilst it is rather leathery with dense,

short hairs abaxially, with less prominent venation, and with a cuneate to

attenuate base in C. pierreana. The flowers are yellow to yellow-orange in

C. vitellina, while C. pierreana has white flowers with a yellow band on the

labellum and dark maroon tips to the labellum and lateral staminodes.

Acknowledgments

We thank the curators of BK, BKF, E, K, P, SING, VNM herbaria for letting

us examine specimens in their care, the Asian Zingiberaceae Information

Centre at Singapore Botanic Gardens, and the Zingiberaceae Resource

Curcuma vitellina (Zingiberaceae) from Vietnam 117

Centre at the Royal Botanic Garden Edinburgh (http://elmer.rbge.org.uk/

ZRC/) for providing protologues and related references. We are grateful to

Prof. Lé Cong Kiét and Dr. Tran Triét for their support during our fieldwork

in Vietnam, and to Dr. J.F. Veldkamp (L) for translating the diagnosis into

Latin. The second author thanks Singapore Botanic Gardens for granting

an SBG Fellowship to work on the Zingiberaceae of Cambodia, Laos

and Vietnam. The fundings by Sud Expert Plantes, France [SEP project

350], by the National Parks Board (Singapore), and by the Czech Science

Foundation, GACR [grant numbers 521/09/0202 and P506/10/0623] are

gratefully acknowledged.

References

Gagnepain, F. 1908. Zingibéracées, pp. 25-121. In: Lecomte, H. (ed.), Flore

Générale de ’ Indo-Chine, vol. 6. Masson & Co., Paris.

Leong-Skorniékova, J. , Sida, O., Sabu, M. & K. Marhold. 2008. Taxonomic

and nomenclatural puzzles in Indian Curcuma: the identity and

nomenclatural history of C. zedoaria (Christm.) Roscoe and C. zerumbet

Roxb. Taxon 57: 949-962.

Mood, J. & K. Larsen. 2001. New curcumas from South-east Asia. The New

Plantsman 8: 207-217.

NY

4

Gardens’ Bulletin Singapore 62 (I): 119-120. 2010 119

New Combinations in Haplopteris ( Adiantaceae) for the

Flora of Peninsular Malaysia

S. LINDSAY

Royal Botanic Garden Edinburgh,

20A Inverleith Row

Edinburgh EH3 5LR, Scotland, UK.

Abstract

Two new combinations are made here: Haplopteris hirta (Fée) S.Linds. and

Haplopteris angustissima (Holttum) S.Linds.

In preparation for the account of the Adiantaceae for the Flora of Peninsular

Malaysia new combinations are necessary in the genus Haplopteris C.Presl.

The family delimitation to be used in the Flora account is modified from

Smith et al. (2006, 2008) who recognized a broadly delimited Pteridaceae

but acknowledged that there are five monophyletic groups within the family

that could be raised back to family level. The Adiantaceae, comprising

Adiantum L. and all members of the former family Vittariaceae, is one of

these monophyletic groups.

As so defined, the Adiantaceae in Peninsular Malaysia consists of

the five genera Adiantum L., Antrophyum Kaulf.. Haplopteris C.Presl,

Monogramma Comm. ex Schkuhr., and Vaginularia Fée. The generic

delimitation of the Vittarioid genera in the Flora account will largely

follow Crane (1998). In this work it was shown that the Old and New

World Vittaria species did not form a monophyletic group. All Vittaria

species in Peninsular Malaysia (see Holttum, 1955) now belong in the

genus Haplopteris. Crane (1998) and Zhang (2003) made most of the

mecessary name changes but two new combinations are still required.

Haplopteris hirta (Fée) S.Linds. comb. nov.

Basionym: Vittaria hirta Fée, Mém. Foug. 10 (1865) 12. -Type: Wallace s.n.,

Borneo (holotype, BM).

Haplopteris angustissima (Holttum) S.Linds.. comb. nov.

Basionym: Vittaria angustissima Holttum, Gard. Bull. Singapore 11 (1947)

274. -Type: C_E.Carr s.n., Pahang. Fraser’s Hill, alt. 4000 ft. March

1929 (holotype, SING).

120 Gard. Bull. Singapore 62 (1) 2010

Acknowledgements

I thank the Forest Research Institute Malaysia (FRIM) for a visiting

Fellowship to pursue research on Malaysian Adiantaceae. I also thank

Serena Lee (SING) and Alison Paul (BM) for their help in locating type

material.

References

Crane, E.H. 1998 [1997]. A revised circumscription of the genera of the

fern family Vittariaceae. Systematic Botany 22: 509-517.

Holttum, E.H. 1955. A Revised Flora of Malaya, vol. 2., Ferns of Malaya.

Government Printing Office, Singapore.

Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and P.G.

Wolf. 2006. A classification for extant ferns. Taxon 55: 705-731.

Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and

P.G. Wolf. 2008. Fern Classification, pp. 417-467. In: T.A. Ranker and

C.H.Haufler (eds). Biology and Evolution of Ferns and Lycophytes,

Cambridge University Press, Cambridge.

Zhang, X-C. 2003. New combinations in Haplopteris (Pteridophyta: Vitta-

riaceae). Annales Botanici Fennici 40: 459-461.

Gardens’ Bulletin Singapore 62 (1): 121-126. 2010 (Dil

A New Species of Alocasia (Araceae-Colocasieae)

From Cambodia

V.D.NGUYEN ,J.C. REGALADO, JR.’ AND VU TIEN CHINH!

‘Institute of Ecology & Biological Resources, Vietnam Academy of

Science and Technology, 18 Hoang Quoc Viet Road, Hanoi, Vietnam

d Email: nguyenvandu@fpt.vn

- Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299,

USA

Abstract

Alocasia jiewhoei V.D.Nguyen, a new species from Cambodia, is described

and illustrated.

Introduction

The genus Alocasia (Araceae-Colocasieae) includes about 75 species

distributed mainly in tropical Asia, Malesia, Australia and Melanesia

(Govaerts and Frodin, 2002). In Indo-China, eleven species have been

recorded in the Flore Général de |’ Indo-Chine by Gagnepain (1942). In 2007,

during fieldwork in Cambodia, V.D. Nguyen’s group found a new species of

Alocasia growing on dry leaf litter in forest in Kulen National Park of Siem

Riep Province. The plant does not match any known species in the genus

(Boyce, 2007, 2008; Hay, 1998, 1999; Hay and Wise, 1991; Medecilo et al.,

2007; Wang et al., 2005).

A comparison of morphological characters with other species in

the genus showed the species resemble A. odora (Lindl.) K.Koch and A.

navicularis (K.Koch & C.D.Bouché) K.Koch & C.D.Bouché, in having

peltate, cordate-ovate leaf blades, 5-10 pairs of lateral veins, the greenish

spathe and conical appendices. However, it is distinguished from A. odora

and A. navicularis by its smaller size, slender petioles, stolons with bulbils

at the apex, conspicuous styles and prominently lobed stigmas (Table 1). It

closely resembles A. navicularis in having erect, cymbiform spathes.

Alocasia jiewhoei V.D. Nguyen, sp. nov.

Alocasiae odorae similis in folii lamina peltata et spathae lamina dilute

viridi, sed spathae lamina cymbiformi, stylis longis, stigmatibus longe lobatis

differt.-Typus: Cambodia, Siem Riep Province, Phrom Ku Len National

Park, 13 36’58.1” N 104 02°20.4” E, 150 m alt., V.D. Nguyen & Rattana CB-

VN 212 (holotype, HN; isotypes, K, P, MO). Fig. 1 & Plate 1.

122

Table 1. A comparison of Alocasia jiewhoei and closely related species.

Characters

]

A. odora

Plant height

40 to >250 cm

Rhizome internal

Light green and white

colour

Petiole Stout

Bulbils on the

None

stolons

Style Very short or

i inconspicuous

Spathe limb

- shape Hooked cymbiform

- colour Green to dull green

Stigma Shallowly lobed

Habitat Wet, evergreen forest

Gard. Bull. Singapore 62 (1) 2010

A. navicularis A. jiewhoei

50 to150 cm 30 to 70 cm

fen Ce aud Light yellow

white

None At the apex

Very short or

inconspicuous

Erect cymbiform

Dull yellow

Conspicuous, 1-1.5

mm long

Erect cymbiform

Medium green

Shallowly lobed

Wet, evergreen

forest

Prominently lobed

Dry, deciduous

forest

Rhizomatous plants, ca 70 cm tall. Rhizomes decumbent, slender to stout,

aerial parts 10-15 cm long, 1-4 cm in diameter, internodes 1.5-3 mm long,

grey to dark brown externally, light yellow internally, covered by traces of

decayed cataphylls. Roots stout, 10-15 cm long, not branched. Stolons several,

subterranean, apex aerial, bearing bulbils at the apex; bulbils obovate,

winged, six-angular in cross section, 1.5 cm long, ca 1 cm in diameter, oblong

at base, abruptly acuminate, covered by several brown, small triangular

scales. Leaves peltate, 4-7 together; petioles slender, 40-70 cm long, sheathed

up to >1/3 of the length, green with dull dark green veins sunk below the

surface; leaf blades ovate, 22-50 cm long, 17-30 cm at widest point, basal

lobes broadly triangular, 7-16 cm long, 6-13 cm wide at base, apices obtuse,

connection between 2 lobes 5-8 cm wide from petiole apex, dark green

Alocasia jiewhoei from Cambodia 123

Wd Z

Figure 1. Alocasia jiewhoei V.D. Nguyen. 1. Habit; 2. Inflorescence; 3. Detail of female flower

zone; 4. Male flower; 5. Ovary in longitudinal section; 6. Ovule.

and glossy above, dull green beneath; lateral veins 7-8 pairs in anterior

lobes, 2-3 veins in posterior lobes, pronounced above, prominent beneath,

secondary veins originating from lateral veins and midrib, run parallel and

ascend towards leaf margins, connected by reticular veins. Inflorescences

one per axil, several per plant; peduncles slender, 25-30(-45) cm long, 7-10

mm in diameter, expanded at top, green, covered by a thin white chalky

layer; spathes 11-12 cm long, constricted about 1/5 of the way from the base;

lower part ellipsoid or subglobose, green; limb 8.3-10 cm long, ca 4 cm wide,

cymbiform, green when young, medium green at anthesis; spadices 8.5-10 cm

long; female zones cylindrical, 0.8-1 cm long, female flowers congested, light

yellow; ovaries subglobose in general, shallowly concaved into three lobes,

2 mm in diameter, green; styles 1 mm long, white; stigmas three lobed, 1-1.2

mm wide, white; ovules ca 10, placentation basal; sterile interstice conical

at lower part, constricted near apex then slightly expanded at apex, 1.8-2.3

cm long, 9 mm in diameter at base, 3-4 mm at the constriction and 4-5 mm

at apex, bearing 2 whorls of white, acute-ovate, flattened sterile flowers in

124 Gard. Bull. Singapore 62 (1) 2010

P

Plate 1. Alocasia jiewhoei V.D.Nguyen. A. Plant habit; B. Appendix and male portion; C.

Staminodes and female portion in detail. (Photos: V.D. Nguyen).

Alocasia jiewhoei from Cambodia LS)

upper part; base covered by 2 rows of sterile flowers, columnar or ring-like;

male zones cylindrical, 1.8-2.3 cm long, 8-10 mm in diameter, oblong to base,

white; appendices conical, 3.5-4.8 cm long, smaller in diameter than male

zone, acute at apex, very shallowly concaved on surface, without veins, dull

white; male flowers sub-rectangular, 3 mm long, ca 2 mm wide, bearing 4-6

stamens. Berries subglobose, 3-4 mm in diameter; seeds 2-3.

Ecology: Dry seasonal forest, altitude 0-500 m.

Distribution: Endemic in Cambodia.

Vernacular name: None recorded.

Uses: None recorded. However the new species has potential as an

ornamental plant.

Etymology: The plant is named in honour of Tan Jiew Hoe of Singapore.

Conservation status: The plant is found in the Phrom Ku Len National Park

of Siem Riep Province where the forest is protected. Although the new

species is rare, there is little data to show if it is threatened.

Acknowledgements

In our study of the family Araceae in Indochina, the first author received

support from the U.S. National Geographic Society Grant No. 8283-07 for

conducting fieldwork in Laos and Cambodia and additional support from

Mr. Tan Jiew Hoe in Singapore for the collecting expedition to Cambodia

in 2007. The authors also thank Dr. J.F Veldkamp for the preparation of the

Latin diagnosis and Dr. Ruth Kiew and the editor of the Gardens’ Bulletin

Singapore for help with the English text and the constructive comments.

References

Boyce, P.C. 2007. Studies on the Alocasia Schott (Araceae-Colocasieae) of

Borneo: I. Two new species from Sarawak, Malaysian Borneo. Gardens’

Bulletin Singapore 58: 141-154.

Boyce, P.C. 2008. A review of Alocasia (Araceae: Colocasieae) for Thailand

including a novel species and new species records from Southwest

Thailand. Thai Forestry Bulletin (Botany) 36: 1-17.

126 Gard. Bull. Singapore 62 (1) 2010

Engler,A.andK.Krause.1920.“Additamentum adAraceas-Philodendroideas,

Araceae-Colocasioideae”, Das Pflanzenreich T1(1V.23E): 3-132.

Gagnepain F. 1942. Aracée. In: Lecomte, H. Flore Général de Indo-Chine

6(9): 1075-1196.

Govaerts, R. and D.G. Frodin. 2002. World Checklist and Bibliography of

Araceae (and Acoraceae). Kew Royal Botanic Gardens, 560pp.

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

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

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

Gardens’ Bulletin Singapore 51: 1-41.

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

Blumea 35(2): 499-545.

Medecilo MP, G.C. Yao and D.A. Madulid. 2007. A new species of Alocasia

(Araceae: Colocasieae) from Panay Island, Philippines. Journal of the

Botanical Research Institute of Texas 1(2): 815-818.

Wang, Yue-Hua, Jian-Tao Yin and Zai-Fu Xu, 2005. Alocasia hypnosa

(Araceae), a new species from Yunnan, China. Annales Botanici Fennici

42: 395-398.

Gardens’ Bulletin Singapore 62 (1): 127-133. 2010 127

New Combinations in Malaysian Staphyleaceae

A.T. NOR-EZZAWANIS

Forest Research Institute of Malaysia

52109 Kepong, Selangor, Malaysia

Abstract

New combinations are made for the seven species of Dalrympelea

(Staphyleaceae) from Malaysia that were previously included in Turpinia.

Dalrympelea pomifera Roxb. is distinct from D. sphaerocarpa (Hassk.)

A.T.Nor-Ezzawanis and does not occur in Malaysia and Turpinia ovalifolia

‘Elmer from the Philippines is distinct from D. trifoliata (Ridl.) A.T.Nor-

Ezzawanis.

Introduction

Staphyleaceae currently includes two genera, Dalrympelea Roxb. and

Staphylea L. (Simmons, 2007; Stevens, 2010), that can be distinguished by

the following morphological characters: stoloniferous shrubs (2 m) to upper

canopy trees of 25-30 m in Dalrympelea, and small trees reaching 15 m in

Staphylea; the bark ranges from creamy yellow and flaky to smooth grey

in Dalrympelea, and grey to black and somewhat mottled, with or without

lenticels in Staphylea: the stipules of opposed leaves are fused and sometimes

becoming bifid at the apex, often having a colleter in Dalrympelea, and the

stipules are free and multi-veined in most species of Staphylea. The former

occurs in the Old World and the latter in both the Old and New Worlds.

Based on molecular analysis using nuclear and chloroplast markers,

Staphyleaceae divides into two clades that are treated as distinct at the

generic level (Simmons & Panero, 2000; Simmons, 2007). One clade,

Staphylea, includes all species of Staphylea, all New World species of the

Turpinia Raf.,and one Asian species, Turpinia cochinchinensis (Lour.) Merr.

and the monotypic Euscaphis japonica (Thunb.) Kanitz (Simmons. 2007).

The other clade includes the Old World Turpinia species, except

for Turpinia cochinchinensis. Members of this latter clade are generally

characterized by being evergreen trees with fused stipules (at least at the

base), coriaceous leaves and a berry with a thickened pericarp (Simmons.

2007). Because the type species of Turpinia is T. cochinchinensis, which is

now included in Staphylea,the name Turpinia cannot be applied to the Asian

species. The earliest name for this clade is Dalrympelea Roxb. (Roxburgh,

128 Gard. Bull. Singapore 62 (1) 2010

1819). The revision of the family for the Flora of Peninsular Malaysia

necessitates making new combinations for local Turpinia species besides

reassessing the status of some species. New combinations are also provided

for species in the Malaysian states of Sabah and Sarawak in Borneo that

were revised by Pereira (1995).

DALRYMPELEA Roxb.

Hort. Beng. (1814) 17, nomen, Pl. Corom. 3 (1819) 76, t. 279; Simmons in

Kubitzki, Fam. Gen. Vasc. Pl. 9 (2007) 443. Type species: Dalrympelea

pomifera Roxb., Pl. Corom. 3 (1819) 76. ¢. 279.

Synonym: Turpinia Vent., Choix (1803) 31, t.31, p.p. excl. New World species

and T. cochinchinensis; Ridley, Fl. Malay Pen. 1 (1922) 511; Linden, FI.

Malesiana 1, 6 (1960) 51; Whitmore, Tr. Fl. Malaya 1 (1972) 447; Pereira, Tr.

Fl. Sabah & Sarawak (1 (1995) 454.

Evergreen shrubs to large trees to 30 m tall, sometimes with buttresses.

Leaves pinnately (1- or 3-) or 5-15-foliolate, coriaceous; stipules interpetiolar,

basally connate to fused along their length. Inflorescences terminal or

axillary panicles with more than 100 flowers. Flowers with sepals free, shorter

than the pink, cream, yellow or green-white petals; stamens arising between

the lobes of the nectary disc; ovary (2-)3(-4)-locular, sometimes partially

imbedded in the disc, more or less syncarpous, with 2-8 ovules per locule.

Fruit an ellipsoid, nearly globose or trilobed berry, fleshy or leathery, usually

green to purple; exocarp thick and fleshy to woody. Seeds 1-6 per fruit.

Distribution: About 20-25 species from Sri Lanka and S India to China and

Japan and southwards to New Guinea (Simmons, 2007) with 8 species in

Malaysia (2 species in Peninsular Malaysia and 6 species and | variety in

Sabah and Sarawak).

New Combinations for Malaysian Species

1. Dalrympelea borneensis (Merr. & L.M.Perry) A.T.Nor-Ezzawanis, comb.

nov.

Basionym: Turpinia montana var. borneensis Merr. & L.M.Perry, J. Arnold

Arbor. 22 (1941) 553. -Homotypic synonym: Turpinia borneensis (Merr. &

L.M.Perry) B.L.Linden, Fl. Malesiana 1, 6 (1960) 56; Pereira, Tr. Fl. Sabah

& Sarawak 1 (1995) 456. -Type: Borneo, Sabah [British North Borneo],

Gunung Kinabalu, Tenompok, J. & M.S. Clemens 29391 (holotype, UC;

isotypes, BO, K).

Distribution: Borneo (Kalimantan and Sabah) and the Philippines.

New Combinations in Malaysian Staphyleaceae 129

2. Dalrympelea calciphila (J.T.Pereira) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Jurpinia calciphila J.T.Pereira, Sandakania 5 (1994) 18, fig. 1, Tr.

Fl. Sabah & Sarawak 1 (1995) 457. -Type: Borneo, Sarawak, Miri Division,

Gunung Api, Anderson S 4710 (holotype, SAR; isotypes, BO, K, L, SING).

Distribution: Endemic in Sarawak, Borneo — Kuching Division near Bau

(Bukit Buan and Bukit Gebung) and Miri Division (Gunung Api and

Gunung Buda).

3. Dalrympelea grandis (B.L.Linden) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Turpinia grandis B.L.Linden, Fl. Malesiana 1,6 (1960) 55; Pereira,

Tr. Fl. Sabah & Sarawak 1 (1995) 457. -Type: E Borneo, W Koetai, Kiau

River, Endert 4669 (holotype, L; isotypes, BO, K).

Distribution: Endemic in Borneo (Kalimantan, Sabah and Sarawak).

4. Dalrympelea nitida (Merr. & L.M.Perry) A.T.Nor-Ezzawanis, comb. nov.

Basionym: TJurpinia nitida Merr. & L.M.Perry, J. Arnold Arbor. 22 (1941) 549;

Linden, Fl. Malesiana 1, 6 (1960) 58; Pereira, Tr. Fl. Sabah & Sarawak 1

(1995) 459.-Type: Borneo, Sabah [British North Borneo], Gunung Kinabalu,

Penibukan, J. & M.S. Clemens 30840 (holotype, UC; isotypes, BO, K).

Distribution: Endemic in Sabah, Borneo (Ranau, Sandakan, Kudat and

Lahad Datu Districts).

5. Dalrympelea sphaerocarpa (Hassk.) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Turpinia sphaerocarpa Hassk., Flora 25, 2 (1842) Beibl. 1, 42:

Ridley, J. Str. Br. Roy. As. Soc. 82 (1920) 179; Linden, Fl. Malesiana 1, 6

(1960) 49; Whitmore, Tr. Fl. Malaya 1 (1972) 448; Pereira, Tr. Fl. Sabah &

Sarawak 1 (1995) 460-Homotypic synonym: Turpinia pomifera (Roxb.) DC.

var. sphaerocarpa (Hassk.) King, J. As. Soc. Beng. 65 (1896) 453. —Type: Java.

sine coll., s.n., Herb. Reinwardtianum (holotype, L, acc. no. 908272875).

Heterotypic synonyms: Turpinia latifolia Wall. ex Ridl., J. Str. Br. Roy. As.

Soc. 82 (1920) 178, Fl. Mal. Pen. 1 (1922) 512Type: Singapore, Wallich 4939

(lectotype, K, isotype, CAL). -Turpinia laxiflora Ridl., J. Str. Br. Roy. As. Soc.

82 (1920) 179, FI. Malay Pen. 1 (1922) 512; Linden, Fl. Malesiana 1, 6 (1960)

57.—-Type: Peninsular Malaysia, Perak, Larut, Kunstler 2824 (holotype, K).

Taxonomic notes: Wallich (1828) in his Catalogue named Wallich 4939 as

‘Turpinia ?latifolia’. Hiern (1875) included Turpinia latifolia as a synonym

of Turpinia pomifera (Roxb.) DC. with Wallich 4939 as the only specimen

cited for Malaya.

130 Gard. Bull. Singapore 62 (1) 2010

King (1896) described two varieties of Turpinia pomifera from

Peninsular Malaysia. The typical variety was represented by asingle specimen,

Kings’ Collector 4243 from Perak, and Turpinia pomifera var. sphaerocarpa

(Hassk.) King was described as ‘the common form in the Malayan Provinces’.

King cited 7: latifolia based on Wallich 4939 as asynonym of var. sphaerocarpa.

Turpinia sphaerocarpa Hassk. was originally described from Java

and has small fruits 0.5-1.8 cm diameter with a thin pericarp 1-4 mm thick

as compared with Turpinia pomifera from Continental Asia that Linden

(1960) described as having larger fruits 2.5-3.7 cm diameter with a pericarp

5 mm thick. Re-constructing the fragments of the fruit on Kings’ Collector

4243 shows that it has a small fruit about 1.5 cm diameter with a pericarp

ca 3 mm thick showing conclusively that it is not a specimen of Turpinia

pomifera, but is typical of Turpinia sphaerocarpa. Ridley (1920) was not

able to examine Kings’ Collector 4243 (there is apparently no specimen at

Kew) but considered it was unlikely to be the same as the Turpinia pomifera

and instead he used Wallich’s original name, T- latifolia Wall. ex Ridl., to

describe this common species in Peninsular Malaysia. (Incidentally, he gave

Penang Hill as the locality for Wallich 4939 but in fact Wallich collected it

from Singapore). Ridley’s description mentioned the fruit diameter as ca

1.3 cm (within the Turpinia sphaerocarpa range), but unfortunately he did

not record pericarp thickness. Linden (1960) too was doubtful that Turpinia

pomifera occurred in Peninsular Malaysia and considered Kings’ Collector

4243 as "too inadequate" for identification. He therefore reinstated Turpinia

sphaerocarpa to specific rank with Turpinia pomifera var. sphaerocarpa and

T. latifolia as synonyms. Whitmore (1972) noted that Turpinia pomifera was

based‘onone doubtfulcollection’, namely Kings’ Collector 4243. Itis therefore

now clear that Turpinia pomifera does not occur in Peninsular Malaysia.

Of the three type specimens at L (acc. nos. 908272875, 908272863 and

908272865), sheet number 908272875 is chosen as the lectotype because it

is the most complete specimen with more fruits and it is from Herbarium

Reinwardtianum.

Ridley (1920) described a new species of Turpinia, T laxiflora Ridl., but

without a description of the fruit. In his Flora (Ridley, 1922), he distinguished

it from 7: latifolia by panicle size: 20 cm [8 in] long in T: latifolia as opposed

to 38 cm [15 in] in 7: laxiflora. Linden (1960) followed Ridley in retaining

T: laxiflora as a distinct species noting that it differed from 7. sphaerocarpa

in its wider leaflets (8-24 x 3.5-12 cm as opposed to 3-18 x 2-10 cm in 7.

sphaerocarpa), larger panicles (to 45 cm vs. to 30 cm long), narrower petals

(2.25-2.5 x 1-1.25 mm vs. 2.5 x 1.5 mm), shorter filaments (1.5-1.75 mm vs.

2-2.5 mm long), smaller fruits (0.5-1 cm vs. 1-1.5 cm diameter) and very thin

pericarp (0.2-0.9 mm thick vs. 1-3 mm thick in 7: sphaerocarpa). However,

Whitmore (1972) synonymised T: laxiflora with T. sphaerocarpa but without

giving a reason for his decision.

New Combinations in Malaysian Staphyleaceae 13]

Examination of specimens from Peninsular Malaysia identified as 7.

latifolia/T. sphaerocarpa and T. laxiflora show that the characters listed by

Linden (1960) are not mutually exclusive. For example, Corner 28690 (KEP)

has a panicle ca 21 cm long, which falls within Dalrympelea sphaerocarpa,

but the petals measure ca 2 x 1.25 mm and filaments ca 1.25-1.5 mm long

and so fall within the range for T: laxiflora. Similarly, Jaamat 10275 (KEP)

has a fruit ca 1.2 cm diameter with a pericarp ca 4 mm thick that fits D.

sphaerocarpa but the petals measure ca 2 x 1 mm and the filaments are ca

1.5 mm long that fall within the range of T: laxiflora. Linden (1960) also

mentioned colour of the dried leaf as a distinguishing characters (brown on

both surfaces in T: laxiflora, but discolorous in D. sphaerocarpa), but again it

was found that this character did not distinguish these two taxa. There being

no constant character or combination of characters to keep 7: laxiflora apart,

it is here treated as a synonym of D. sphaerocarpa.

Burkill (1966) confused the picture by recognizing Turpinia pomifera

and treating both 7. Jatifolia and T. laxiflora as synonyms of it. He described

T. pomifera as a “fairly large tree in the lowlands” which indicates that he

was probably referring to Dalrympelea sphaerocarpa.

Pereira (1994) recognized a second variety, var. microcerotis, from

Borneo, which differs from var. sphaerocarpa in having shorter leaf rachises

(2-8 cm vs. 10-18 cm long), shorter leaflets (5-10 cm vs. 7-19 cm long) and

a slightly trilobed fruit with three apical horns compared with the globose,

hornless fruit of var. sphaerocarpa.

Distribution: Sumatra, Peninsular Malaysia, Singapore, Borneo, Java, Lesser

Sunda Islands (Bali, Flores), Sulawesi, the Philippines and Maluku (Ceram,

Ambon).

5a. Dalrympelea sphaerocarpa (Hassk.) A.T. Nor-Ezzawanis var.

microcerotis (J.T. Pereira) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Turpinia sphaerocarpa Hassk. var. microcerotis J.T. Pereira,

Sandakania 5 (1994) 21, fig. 2; Tr. Fl. Sabah & Sarawak 1 (1995) 461. —

Type: Borneo, Sabah [British North Borneo], Ranau, Gunung Kinabalu,

Tenompok, J. & M.S. Clemens 28707 (holotype, SING; isotypes, BO, K).

Distribution: Endemic in Borneo (Sabah and Kalimantan).

6. Dalrympelea stipulacea (B.L.Linden) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Turpinia stipulacea B.L.Linden, Fl. Malesiana 1, 6 (1960) 55;

Pereira, Tr. Fl. Sabah & Sarawak 1 (1995) 461.—Type: Borneo, Sabah, Gunung

Kinabalu, near Tibabah River, Carr SFN 27516 (holotype, SING).

Distribution: Endemic to Sabah (Ranau district only).

132 Gard. Bull. Singapore 62 (1) 2010

7. Dalrympelea trifoliata (Ridl.) A.T.Nor-Ezzawanis, comb. nov.

Basionym: Turpinia trifoliata Ridl., J. Str. Br. Roy. As. Soc. 82 (1920) 178, FI.

Malay Pen. 1 (1922) 511, fig. 50. -Type: Peninsular Malaysia, Melaka, Nyalas,

Goodenough 1771 (lectotype, SING, here chosen).

Heterotypic synonym: Turpinia ovalifolia Elmer, Leafl. Philip. Bot. 2 (1908)

490, auct. non: Linden, Fl. Malesiana 1,6 (1960) 58; Whitmore, Tr. Fl. Malaya 1

(1972) 448.-Type: Philippines, Luzon, Lucban Tayabas, Elmer 8088 (SING).

Taxonomic notes: Some authors (Linden, 1960; Whitmore, 1972) considered

this species in Peninsular Malaysia to be conspecific with Turpinia ovalifolia.

However, careful comparison between Peninsular Malaysian specimens

and the type specimens of both 7. ovalifolia and T. trifoliata show that the

Malaysian specimens previously identified as 7. ovalifolia in fact belong

to T. trifoliata. Turpinia ovalifolia, known only from the type specimen, is

characterized by its small leaves ca 4 x 2.5 cm with a more-or-less entire

margin. In contrast, T° trifoliata specimens cited by Ridley (1920), namely

Goodenough 1771 and Ridley 15906, are characterized by larger leaves ca 7-

14 x 3.5-7 cm with a crenate margin with fine, dark brown (when dry) callous

points. These characters are shared by more recently collected specimens

from Malaysia and indeed some have even larger leaves up to ca 9.5-20

x 4-9.5 cm. Therefore it is concluded that 7: ovalifolia does not occur in

Peninsular Malaysia.

Ridley (1920) cited two specimens in his original description of the

species. Goodenough 1771 is here chosen as the lectotype because it is a

more complete specimen and the sheet has Ridley’s description of the

flower (in his handwriting) and is named ‘7urpinia trifoliata’.

Distribution: Peninsular Malaysia (Perak, Selangor, Negeri Sembilan,

Pahang and Johor).

Acknowledgements

This study was carried out for the Flora of Peninsular Malaysia Project

funded by the Ministry of Science, Technology and Innovation (MOSTI)

through the National Council for Scientific Research and Development

(MPKSN) under Project No. 01-04-01-0000 Khas 2 entitled ‘Safeguarding

the Forest Plant Diversity of Peninsular Malaysia’. I am indebted to P.F.

Stevens for advice on the current status of Staphylea and Dalrympelea and

to Drs. R. Kiew, Saw L.G., R.C.K. Chung, and E. Soepadmo for help with

preparing the manuscript. I should also like to thank the Curators of the

herbaria at KLU, UKMB and SING for loan of specimens and to BM, K and

L for access to their collections.

New Combinations in Malaysian Staphyleaceae 133

References

Burkill, 1.H., 1966. Turpinia pomifera. In: A Dictionary of the Economic

Products of the Malay Peninsula. p. 2234.

Hiern, W.P. 1875. Turpinia. In Hooker,f Flora of British India 1: 698-699.

King, G. 1896. Turpinia. Journal of the Asiatic Society of Bengal 62: 452-453.

Linden, B.L. van der. 1960. Staphyleaceae. In: Flora Malesiana 1 (6): 49-59.

Pereira,J.T.1994. Anewspecies andanewvariety of Turpinia (Staphyleaceae)

from Borneo. Sandakania 5: 15-23.

Pereira, J.T. 1995. Staphyleaceae. In: Tree Flora of Sabah & Sarawak 1: 453-

462.

Ridley, H.N. 1920. New and rare species of Malayan plants. Journal of the

Straits Branch of the Royal Asiatic Society 82: 177-179.

Ridley, H.N. 1922. Turpinia. In: Flora of the Malay Peninsula 1: 511-512.

Roxburgh, W. 1819. Dalrympelea pomifera. In: Plants of the East Coast of

Coromandel, pp. 76-77, t. 279.

Simmons, S.L. 2007. Staphyleaceae. In: Kubitzki, K.(ed.). The Families and

Genera of Vascular Plants 9: 443-445.

Simmons, S.L. and J.L. Panero. 2000. Phylogeny and biogeography of

Staphyleaceae (DC.) Lindl. American Journal of Botany 87(6, suppl.):

S77.

Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9,

June 2008. http://www.mobot.org/MOBOT/research/A Pweb/.

Latest update January 2010.

Wallich, N. 1828. Turpinia latifolia. In: Numerical List of Dried Specimens of

Plants, p. 173.

Whitmore, T.C. 1972. Staphyleaceae. In: Tree Flora of Malaya 1: 446-448.

dir? @& que 7

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Gardens Bulletin Singapore 62 (1): 135-142. 2010. 135

A New Species of Costaceae from Borneo

A.D.POULSEN’ AND C.D. SPECHT”

Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh

: EH3 5LR, Scotland, U.K.

- Plant and Microbial Biology, University of California, Berkeley,

111 Koshland Hall, MC 3102, Berkeley, CA 94720, USA

Abstract

A new species, Cheilocostus borneensis, is described. Specimens were

collected in Sarawak in 1987 and Kalimantan in 2000, but only intensified

surveys of gingers in Sarawak in 2002-2004 provided sufficient collections

to recognize the new species, which is here described and illustrated. It is

closely related to the widespread C. globosus from which it differs by the

chocolate-brown sheaths, absence of axillary shoots on vegetative stems,

larger leathery leaves, and by its calyx that is not prickly.

Introduction

Members of Bornean Costaceae were previously placed in the genus

Costus L. (Maas, 1979). which is now circumscribed as a clade comprising

only African and neotropical species based on phylogenetic analyses of

morphological and molecular data (Specht and Stevenson, 2006). Following

this evaluation of the generic circumscription of Costaceae, only two genera,

Cheilocostus C. Specht and Paracostus C. Specht, are native to Borneo where

seven species of Costaceae are presently known (Maas. 1979: Meekiong

et al., 2006; Meekiong et al., 2008). The exact generic placement has not

been established for all Bornean species and an updated revision for both

Cheilocostus and Paracostus is pending and will likely include other recently

discovered and described species.

The genus Cheilocostus is easily distinguished from Paracostus

by consisting of larger plants (> 1.5 m high) with erect shoots, and a

condensed inflorescence with conspicuous bracts, each subtending a single

flower. Paracostus, in contrast, is characterized by smaller plants (< 1.5

m), prostrate stems with few leaves, and inflorescences with few flowers

supported by inconspicuous bracts. Cheilocostus is closely related to the

genus Japeinochilus Mig. which is only found east of Sulawesi into New

Guinea, Australia and the Pacific (Gideon, 1996; Poulsen er al. 2010).

136 Gard. Bull. Singapore 62 (1) 2010

During expeditions targeting gingers in Sarawak in 2002-2004, the

first author collected, with Malaysian collaborators, new material of several

species of Costaceae. One of these had already been collected several times,

and specimens were deposited in several herbaria but without pickled

flowers essential for its description. The material now being available, the

species is described below.

Cheilocostus borneensis A.D. Poulsen, sp. nov.

Cheilocostus borneensis in inflorescentia radicaliC. globoso similis est

sed ab eo foliis ad apicem caulis aggregatis plerumque majoribus et calyce

molliter acuto (haud pungenti nec aculeato) differt. -Typus: Malaysia, Borneo,

Sarawak, Batang Ai, Sungai Senkabang, small stream connecting to Sg.

Delok opposite of Ng. Sumpa longhouse, 1°12’S 112°3’E, 130 m, flowering

8 Dec 2002, A.D. Poulsen & Bakir Raymond 1964 (holo, SAR; iso, AAU,

Sarawak Biodiversity Centre Flora Depository). Figs. 1 & 2.

Figure 1. Cheilocostus borneensis photographed by A.D. Poulsen (Poulsen 2696, cultivated).

A New Species of Costaceae from Borneo 137

Terrestrial, perennial herb. Leafy shoots in a dense clump, 1.5-2 m tall. Base of

leafy shoot to 3 cm diam., pale yellow-green when fresh, covered by reddish

brown sheaths. Stem leafless in lower ca 1.4 m, sheaths reddish chocolate-

brown (uppermost yellow-green), glabrous, with 6-10 leaves consistently

clustered toward the shoot apex. Ligule 1-2 mm long, + truncate, slightly

longer laterally to the petiole. Petiole 5-12 mm long, swollen, pale yellowish,

slightly canaliculate, glabrous. Lamina to 27-42 x 9-16(-20) cm, narrowly

obovate, mid-green above, pale green beneath, coriaceous, slightly plicate,

glabrous throughout, base narrowly cuneate, apex acuminate, ca 1 cm long.

Inflorescence radical (i.e. at the base of the plant terminating a separate

leafless shoot that emerges directly from the rhizome), 12-21 cm long (bracts

only), lax. Peduncle horizontal to ascending, 6-12 cm long, sheaths + tubular,

brown, glabrous, margin ragged. Spike 4-12 x 5-8 cm (bracts only). Fertile

bracts to 3.2-3.5 x 2-2.5 cm, elliptic, cucullate, margin membranous, apex

+ emarginate, softly mucronate, dark brown to greenish or pale reddish

brown, glabrous. Rachis (distance from base of lowermost ovary to base

of uppermost ovary) 2 cm long extending with age to 7(-9) cm, with 40-80

flowers, 1-2 open at a time. Bracteole 2-2.5 cm long, split to base adaxially,

reddish brown, glabrous, apex rounded with one minute mucro, 1-lobed

developed laterally, cucullate, not closely adhering to calyx, sometimes with

a second shorter lobe without mucro. Flower 8.5-10 cm long, exerted ca 5

cm above the supporting bract. Calyx 3-3.2 cm long, tubular, dark reddish

brown, glabrous, apex 3-lobed, lobes 0.7-1 cm long, slightly involute, apex

acute to mucronate, soft (not pungent). Corolla tube (from apex of ovary

to base of divergence of labellum and stamen) 2.5-3.2 cm long, fused solid

with style in lowest 1-1.5 cm, white at base; lobes 3.5 x 1.5 cm, narrowly

obovate, translucent white, glabrous, apex rounded, finely apiculate to 1

mm long. Labellum tube (from insertion of dorsal corolla tube to base of

divergence of labellum and stamen) 1.1-1.3 cm long with coarse yellow hairs

inside and outside. Labellum 5-5.3 x 5.2-6 cm, broadly obovate, thickened

in centre with course, short hairs, white with yellow center, margin finely

undulating, glabrous. Stamen ca 2.5 cm long (ca 3.2 cm when crest flattened),

ca 1 cm wide, petaloid, white. Anther crest slightly or irregularly 3-lobed,

recurved, 1.1-1.2 cm long, with yellow spot in lower centre, coarse hairs at

margin. Thecae 0.7-0.8 cm long, 0.3-0.4 cm across both, dehiscing for their

entire length. Ovary 1 x 0.6 cm, flattened ellipsoid, glabrous. Style 3.5-3.7

cm long (free part), glabrous. Stigma ca 0.3 cm wide, fan-shaped, flattened

2-lobed, lobes overlapping, one larger than the other, hairy, pale yellow.

Infructescence head to 12 x 8 cm, often still flowering at apex, with persistent

bracts, bracteoles and, calyces. Fruit 0.9-1 x 0.8-0.9 cm, obovoid, flattened-

triangular, with an apical column (base of calyx and corolla tube), 3-locular,

glabrous, cream to pale green. Seeds 1.5-1.7 x 1-1.2 mm, irregularly barrel-

shaped, black, aril white, basal and not enclosing the seed.

138 Gard. Bull. Singapore 62 (1) 2010

£

&

fb fl br ca

Plate 1. Cheilocostus borneensis. A. Habit; B. Leaves; C. Infructescence with flowers at apex;

D. Close up of labellum and stamen; E. fb = fertile bract, fl = fertile bract with single flower, br

= bracteole, ca = calyx, co = flower with bracteole and calyx removed, tr = longitudinal section

of flower, sd = stamen, dorsal view, sv = stamen, ventral view; F. Fruit with semipersistent

calyx. Photographs by A.D. Poulsen, A, C—D, F of Poulsen & Bakir Raymond 1964 (the type):

B, E of Poulsen 2696 (cultivated).

A New Species of Costaceae from Borneo 139

Additional materials examined: INDONESIA. Borneo. West Kalimantan.

Camp Betung Kerihun NP, Putussibau, 0°56’N 113°19E, 150 m, fruiting 28

Feb 2000, Ambriansyah, Kade Sidiyasa & Albertus AA 2238 (BO, L, WAN).

MALAYSIA. Borneo, Sarawak: Batang Ai, Sungai Senkabang, small stream

connecting to Sg. Delok opposite of Ng. Sumpa longhouse, 1°12’S 112°3’E,

130 m, flowering 3 Jun 1993, Christensen & Poulsen 1997 (AAU): Gunung

Mulu National Park, R. Ubong, between Mulu N.P. and logging concession,

near Base Camp, 200 m, flowering, 7 Nov 1990, Warwick MW177 (E bar code

E00128356): Kapit, Balleh, Ulu Sungai Mengiong, Wong Kijang, flowering 26

Oct 1988, Othman et al. S.56077 (SAR): Kapit, Balleh, Ulu Sungai Mengiong,

Nanga Sebaning, 1°25’N 113°25’E (indicated approximately on map on

rear side of label), flowering 1 Nov 1988, Othman et al. 8.56464 (AAU, E,

K n.v.); Kapit, Batang Baleh, Sungai Mengiong, Sungai Entulu, 18 Jul 1987.

Bernard Lee. S. 54624 (AAU, E (bar code E00320502: bar code E00304605),

K n.v.); Cultivated at Royal Botanic Garden Edinburgh, Accession number

and qualifier: 20040728*A, collected 16 Aug 2007, Poulsen 2596 (E, SING),

origin Poulsen & Bakir Raymond 1964 (the type).

Distribution: Endemic to Borneo where it is known from three main areas

in Sarawak (one being three collections from the Kapit area) and one in

Kalimantan. The furthest localities are about 450 km apart.

Local names and uses: Pa’bu (Iban language; Poulsen & Bakir Raymond

1964). The plant was used in the past by Iban people but it is not certain for

what purpose; pasat baju (Iban; Othman et al. 8.56077).

Etymology: The epithet refers to the species being endemic to Borneo.

Ecology and habitat. Lowland primary or secondary (logged) mixed

dipterocarp forest, along riverbanks, at 130-200 m.

Conservation status: C. borneensis is found at the foot hills of the central

mountain range of Borneo in an area covering at least 2200 km” but with

fewer than 10 localities. Also its sexual reproduction seems dependent on

natural pollinators that may not persist in degraded habitats (see Notes

below). Currently we propose the category of Vulnerable (Vu Blab(iii);

IUCN, 2001), but because of the logging activities and land use is changing

rapidly in Borneo, this category could soon change to “Endangered”.

Notes: Cheilocostus borneensis deviates from the generic description (Specht

and Stevenson, 2006) by not having axillary branching of the vegetative

140 Gard. Bull. Singapore 62 (1) 2010

fi02 115°

Figure 2. Distribution of Cheilocostus borneensis presently known from Sarawak (Malaysia) and

West Kalimantan (Indonesia).

shoots. Vegetative branching is shared by Cheilocostus and Tapeinochilos

but is lost in some Tapeinochilos species and, similarly, may have been lost

in C. borneensis. Molecular evidence using four gene regions (ITS, ETS,

rpb2 and trnL-F) and samples from a single accession (the type) places this

species in Cheilocostus, sister to C. globosus (Blume) C. Specht.

Cheilocostus borneensis is similar to C. globosus in having a radical

inflorescence, but is easily distinguished by having distinctly chocolate-

brown sheaths on the stem (also obvious in herbarium specimens), having

few larger leaves congested at the top of the stem, the leathery almost

plastic-like texture of the leaves, and by not having axillary branching. As

the flowers of C. globosus vary in colour of the labellum from white to

yellow, dark orange to red, this character by itself is not always conclusive

in separating the two. A more useful floral character to separate the two

species is the apex of the calyx: C. borneensis is soft, whereas C. globosus has

rigid points that, if touched, will easily make you bleed (i.e. pungent).

Recently, Costus mulus Meekiong, Ipor & Tawan was described from

Sarawak (Meekiong et al., 2008). This species also has a radical inflorescence

containing flowers with a white labellum but the yellow patch is confined to

a dot at the terminal margin of the labellum. Even though the description

leaves many questions open and we have not been able to examine the type

A New Species of Costaceae from Borneo 14]

material, this species has smaller leaves (13-21 x 6.5-9.5 cm vs. 27-42 x 9-

15 cm) with a rounded to cordate base (not narrowly cuneate) and thus is

clearly distinct from Cheilocostus borneensis.

The northernmost locality presently known of C. borneensis is Mulu

National Park in Sarawak. Despite Smith’s inventory (1984) on the richness

of gingers of this area, no material was collected during her studies. Thus. she

only reported Paracostus paradoxus (K.Schum) C. Specht and Cheilocostus

speciosus (J. KOnig) C. Specht from this area. The collection from Mulu.

Warwick MW177, is a convincing match to the other collections. but the

mucro on the bracts are exceptionally prominent.

In nature, the spike of Cheilocostus borneensis may have mature fruits

in the lower bracts while simultaneously flowering towards the apex (Plate

1C). In cultivation mature fruits were never observed to develop and it is

likely that manual pollination is needed to produce seeds. This may indicate

that C. borneensis is only able to set fruits in habitats where the natural

pollinator is present.

On 11 Dec 2002. at Sungai Rirang near the type locality, a plant of

Cheilocostus borneensis with normal leafy shoots was seen bearing an erect,

54 cm long peduncle that was similar in appearance to the chocolate-brown

sheathed stem of a leafy shoot, but instead of bearing leaves it terminated

in a spike. This aberrant inflorescence has only been seen this once, never

occurred in cultivation, and is apparently a rare phenomenon.

Acknowledgements

The first author thanks his adopted family at Nanga Sumpa for help

with the fieldwork during which this species was discovered, Sarawak

Biodiversity Centre and the National Parks and Nature Reserves of

Sarawak for processing permits, HRH Crown Prince Frederik’s Foundation

for sponsorship of fieldwork, and Paul and Hiltje Maas for confirming the

species as new during a visit to Utrecht in 2003. We thank Robert Mill and

Philip Oswald for help with the Latin diagnosis and John Mood, as well as

two anonymous reviewers, for critically reading an earlier version of the

manuscript.

References

Gideon, O.G. 1996. Systematics and evolution of the genus Tapeinochilos

Mig. (Costaceae — Zingiberales). Thesis submitted for the degree of

Doctor of Philosophy. Department of Botany and Tropical Agriculture,

James Cook University. Queensland, Australia. 385 pp.

142 Gard. Bull. Singapore 62 (1) 2010

IUCN 2001. IUCN Red List Categories and Criteria. Version 3.1. IUCN

Species Survival Commission. Gland, Switzerland and Cambridge, UK.

Maas, PJ.M. 1979. Notes on Asiatic and Australian Costoideae

(Zingiberaceae). Blumea 25: 543-549.

Meekiong, K., Ipor, I.B. and C.S. Tawan. 2006. Three new species from

Sarawak, and notes on the cauliferous Costus Group (Costaceae,

subgenus Paracostus). Folia Malaysiana 7: 55-72.

Meekiong, K., Ipor, I.B., Tawan, C.S. and M. Muliati. 2008. A new Costus

(Costaceae) from Sarawak, Borneo, Malaysia. Rheedea 18: 87-89.

Poulsen, A.D., Gideon, O.G. and M. Ardiyani. 2010. Names of Tapeinochilos

(Costaceae) in Wallaceae. Blumea 55: 61-64.

Smith, R.M. 1984. Zingiberaceae of Mulu National Park, pp. 85-96. In: A.C.

Jermy (ed.), Studies on the flora of Gunung Mulu National Park, Sarawak.

Kuching, Sarawak.

Specht, C.D. and D.W. Stevenson. 2006. A new phylogeny-based generic

classification of Costaceae (Zingiberales). Taxon 55: 153-163.

Gardens’ Bulletin Singapore 62 (1): 143-153. 2010 143

Three New Species of Begonia Sect. Platycentrum

from Nepal

S.RAJBHANDARY .M. HUGHES’ AND K.K.SHRESTHA |

Central Department of Botany

Tribhuvan University. Kirtipur, Kathmandu, Nepal

Royal Botanic Garden Edinburgh

20A Inverleith Row, Edinburgh, U.K.EH3 5LR

Author for correspondence: S. Rajbhandary:imogine3@gmail.com

Abstract

Three new species of Begonia (Begoniaceae) are described from Nepal. All

three species (Begonia nuwakotensis S. Rajbhandary. Begonia panchtharensis

S. Rajbhandary and Begonia taligera S. Rajbhandary) belong to Begonia

section Platycentrum, and they increase the number of Begonia species

known from Nepal to 22. All are considered to belong to the IUCN threat

category VUD2.

Introduction

Begonia inhabits moist, shady locations in humid lowland to upland forests.

The greatest number of species is found in mid-elevations ranging between

1200-2500 m, growing in cloud forest habitats in rock crevices and on moist

boulders and moss-covered tree trunks. Begonia picta Sm. and Begonia

dioica Buch.-Ham. ex D. Don are the most common species in Nepal, found

growing from 150-2700 m altitude. The genus was previously represented

by 18 species in Nepal (Hara er al., 1978; Doorenbos er al., 1998: Press et

al., 2000) and one new record (Rajbhandary and Shrestha, 2009) bringing

this total to 19. These are placed within five different sections, Diploclinium

(Lindl.) A.DC.. Monopteron (A.DC.) Warb., Platycentrum (Klotzsch)

A.DC, Putzeysia (Klotzsch) A.DC and Sphenanthera (Hassk.) Warb. (Smith

et. al. 1986; Doorenbos et al. 1998). There are four endemic species, Begonia

tribenensis C.R. Rao, Begonia minicarpa H. Hara, Begonia fiagellaris H.

Hara and Begonia leptoptera H. Hara.

Section Platycentrum (Klotzsch) A.DC. includes about 110 species

from India through central China to Southeast Asia. and is one of the largest

sections of the genus Begonia (Shui et al., 2002: Ye et al., 2004). This section

is comprised of relatively large and robust species. characterised by a large

144 Gard. Bull. Singapore 62 (1) 2010

androecium on a distinct column, anthers with an extended connective,

two highly convolute or spiralled styles and two-locular, drooping fruits

with three unequal wings, one of which is considerably lengthened.

During botanical expeditions from 2006-2008 to different parts of Nepal

three new species of Begonia belonging to section Platycentrum were

discovered, which are described and illustrated here, bringing the total

number of Begonia species known from the country to 22.

Species descriptions

Begonia nuwakotensis S. Rajbhandary, sp. nov. [§ Platycentrum|

Begoniae palmatae similis sed foliis basalibus longe petiolatis, laminae base

cum lobis superpositis et in margine sine lobis vel dentibus longis curvatis, flore

femineo tepalis exterioribus 4 similibus et tepalo interiore uno parvo, placentis

trilamellatis et fructu cum ala laterali lata oblonga differt. —Typus: Central

Nepal, Nuwakot, Kakani, Doman, 1700 m, 9 Aug 2007, S. Rajbhandary, S.

Ranyitkar, K. P. Thapa and S. R. Bista S31 (holotype, E; isotype, KATH). Plate 1.

Perennial, monoecious, rhizomatous, creeping herb, 14-30 cm tall. Stems

rhizomatous at base, to 2 cm in diameter; erect portions semi-woody, reddish

brown, becoming procumbent, tomentose, trichomes soft red. Stipules

persistent, membranous, ovate, 16-21 x 8-10 mm, green to scariet, with an

abaxially prominent midrib forming a thin, long appendage at the apex,

margin entire, outer surface sparsely hairy on the midrib, trichomes red,

inner glabrous and glossy. Leaves cauline, petiole 6-24 cm long, green, villose,

trichomes, red, multicellular up to 2 mm long; lamina asymmetric, broadly

ovate or oblate-orbicular, 5-22 x 5-19 cm, base slightly oblique, cordate, with

overlapping lobes 1.5-3 cm long, margin remotely and shallowly denticulate,

distinctly divided into short 0.5-2 cm long lobes, apex acute to acuminate;

upper surface green strigulose/sericeous, trichomes small and white, lower

surface pale green, densely pilose, trichomes red and long; palmately veined,

7-8 veins. Inflorescences bisexual, axillary, cymose, few flowered, protandrous;

peduncle 10-25 cm long, green, villose, trichomes red, multicellular. Bracts

caducous, membranous, cymbiform, ovate, 12-15 x 8-11 mm, pale green to

scarlet, glossy and pubescent, trichomes red, margin ciliate, apex acute. Male

flowers: pedicel 1-2.3 cm, green, hairy, trichomes multicellular, red; tepals 4-5,

white, outer 2 tepal orbicular, 13-15 x 13-20 mm, abaxially pilose, trichomes

red, base cordate, margin ciliate, apex rounded to acute; inner 2-3 tepals

obovate, 12-17 x 8-12 mm, smallest innermost 10-12 x 5-6 mm, white,

glabrous, base cuneate, apex acute or sometimes rounded; androecium

receptacle flat, free filaments almost equal, but slightly longer in the middle

of the androecium, stamens ca 100, filaments 1-2 mm; anthers narrowly

Three New Species of Begonia from Nepal 145

Plate 1. Begonia nuwakotensis S. Rajbhandary. A. habit; B. Male flower; C. Stamens; D.

Androecium; E. Upper leaf surface; F. Lower leaf surface; G. Bracts; H. Female flower; I.

Stigma; J. Ovary; K. Prostrate stem; L. Fruits; M. Transverse section of fruit; N. Pollen; O.

seed.

146 Gard. Bull. Singapore 62 (1) 2010

globose, on a small column, 6-10 mm diameter, yellow, symmetrical, obovoid,

1.2-1.5 mm long, anthers towards the outside are longer than the filaments

and those at the centre are shorter than the filaments, dehiscing through

unilateral slits as long as the anther, connective slightly extended. Female

flowers: pedicel 1-2 cm, hairy, trichomes red, multicellular; tepals 5, white,

unequal, four outer tepals are orbicular, 20-22 x 18-22 mm, innermost tepal

smallest, ovate, 13-15 x 7-8 mm, except for the innermost all the tepals are

abaxially pubescent, margin entire, base rounded, apex rounded, acute in

the innermost tepal; styles 2, deciduous, 3-6 mm long, fused at the base and

ends into a broad wide apex, yellowish green; stigma green, a papillose

band, strongly spiralled over the broad apex of the style and twisted at two

ends; ovary green, pubescent, trichomes red, 2 locular; placentation axile,

trilamellate. Fruits with pedicel 15-25 mm long, glabrous; 17-25 x 10-12 mm,

nodding or pendulous, unequally 3-winged, with one larger wing and two

narrow wings, larger wing oblong, 12-20 mm; lateral wings much smaller,

5-6 mm, small wings recurved; capsule obovoid, ca 10-12 x 6-10 mm, green

and hairy when young and dark brown when mature, glabrous; drying dark

brown, dehiscing along the sutures between the two smaller wings.

SEM studies: Pollen: large, perprolate with slightly concave sides and slightly

pointed poles, irregular striate ornamentation with few scattered pores, P

22-22.5 um, E 7.6-8.4 um, P/E 2.6-2.9, the endoporus elliptical with costae,

margo distinct, narrow. Seeds: ellipsoidal to obovoid, mean size 325 x 240

um; collar cells elongated, testa cells polygonal with straight or slightly

curved anticlinal walls; anticlinal walls of testa cells thin and slightly raised;

operculum short, flat and nipple-shaped; cuticular pattern consisting mainly

of short linear zigzag striate ornamentation.

Distribution: Known only from the type locality (Fig. 1).

Ecology: Growing on wet shady slopes by the side of waterfalls, at ca 1700 m.

Notes: This species is similar to Begonia palmata but differs in lacking long

and curved lobes or teeth in the leaf margin. It also differs in having female

flowers with 4 similar outer tepals and one small inner tepal, trilamellate

placentae, and a fruit with a broad oblong wing. It is known only from

the type locality and so it is named after the place of its collection. The

population at the moment is healthy and reproducing well. However, as the

single locality is not under protection and the capability of the species to

cope with less shade and moisture is not likely to be good, we consider it to

belong to the IUCN threat category VUD2.

Three New Species of Begonia from Nepal 147

Figure 1. Distribution of Begonia taligera, Begonia nuwakotensis, Begonia panchtharensis in

Kaski, Nuwakot and Panchthar Districts, Nepal.

Begonia panchtharensis S. Rajbhandary, sp. nov. [§ Platycentrum|]

Begoniae sikkimensi similis sed foliis solum basalibus praesentis eis magnis

suborbicularibus profunde lobatis longe petiolatis et fere glabris, floribus

femineis 6-tepalis et fructu cum ala laterali lata falcata differt. -Typus:

Cultivated plant grown from Eastern Nepal, Panchthar, Tinubote, Sisire,

Prangbung, VDC, 2240-2300 m, 2 x 2007, U. Thamsuhang s.n., vouchered as

S. Rajbhandary S74 (holotype, E, isotype, KATH). Plate 2.

Perennial, monoecious, rhizomatous herb, 50-60 cm tall. Stem rhizomatous,

8-12 cm long, 20-30 mm in diameter, covered with long roots. Stipules

caducous, membranous, cymbiform, broadly ovate, 18-25 x 8-10 mm, pink

with a light green tinge, glabrous with red spots on the abaxial surface,

margin entire, apex acuminate. Leaves several, arising from the rhizome,

petiole 33-46 cm long, succulent, glabrous, grooved, yellowish green with

red linear dots; lamina slightly asymmetric, suborbicular and deeply lobed,

24- 40 x 36-48 cm, base deeply cordate; margin irregularly serrulate or

deeply dentate, distinctly divided to 2/3 of leaf length; lobes 6-8, apex

acuminate, dark green above and pale green below, adaxially sparsely

hirsute more so on the periphery, trichomes small, multi-cellular, crimson or

whitish transparent, abaxially glabrous except for sparse hairs on the veins

near the periphery, palmately veined, 6-8 veins. Inflorescences bisexual,

axillary, cymose, dichotomously branched, protandrous; peduncle 42-50cm

long, glabrous, grooved, sub-woody, yellowish green with red linear dots.

Bracts caducous, in unequal pairs, membranous, cymbiform, elliptic to

broadly elliptic, 25-30 x 15-18 mm, green, narrowly ridged abaxially and

with linear or circular red granules, papery, margin entire, apex acuminate.

148 Gard. Bull. Singapore 62 (1) 2010

Plate 2. Begonia panchtharensis S. Rajbhandary. A. Habit; B. Male flower; C. Androecium; D.

Stamens; E. Bracts; F. Rhizome; G. Petioles; H. Leaf venation; I. Leaf surface; J. & K. Inflorescence;

L. Style and stigma; M. Stipule; N. Fruits; O. Transverse section of fruit; P. Seed; Q. Pollen.

Three New Species of Begonia from Nepal 149

Male flowers: pedicel 1-8 cm, light pink with linear red spots. glabrous:

tepals 4, white to pink: outer 2 tepals broadly ovate. 1.2-1.8 x 0.9-1.2 cm,

abaxially glabrous. with a pink tinge near the tip and the margin, base

truncate, margin entire. apex acute: inner 2 tepals oval to narrowly oval.

1.5-1.8 x 0.8-1.3 cm: white. glabrous, base cuneate, apex retuse: androecium

globose, on a small column. 9-12 mm long. yellow. symmetrical. receptacle

slightly raised: stamens 125-141. filaments 0.5-2.5 mm long: anthers oblong.

1-1.5 mm, anthers towards the outside are longer than the filaments and

those at the centre are shorter than the filaments, dehiscence through long

straight slits which are slightly curved at the tip. connectives extended. acute.

Female flowers: pedicel 1.5-2 cm. glabrous: tepals 6-8, outermost tepals pink.

innermost white. symmetrical. oval 9-11 x 8-11 mm, glabrous. base rounded.

Margin entire, apex acute: styles 2. persistent, 2.5-6 mm long, fused at the

base. greenish yellow. stigmatic surface spirally convolute, papillose: ovary

oblong and slightly curved, locular part 10-13 x 3-4 mm. glabrous with red

- circular or linear granules. with three unequal wings: largest wing oblong

with a rounded tip and two smaller more or less equal rim-like blunt wings:

2-locular, placentae axile, bilamellate. Fruits: pedice] 2.5-3 cm, glabrous: 10-

25 x 10-15 mm, nodding or pendant, falcate. unequally 3-winged: longer

wing obovoid—oblong, 9-25 x 10-15 mm. lateral wings smaller, 1 mm long.

wings red, capsule oblong, 10-15 x 2-4 mm. light greenish yellow when young

becoming brownish green when mature, glabrous: drying dark brown with

green tinge, dehiscing along the sutures between the two smaller wings.

SEM studies: Pollen: large. perprolate with smooth convex sides and rounded

poles, faint striate ornamentation with pores, P 16.8-21 um. E 8-10 um. P.

E 2.1, the endoporus elliptical with costae. margo present. the grains not

syncopate. Seeds: ellipsoidal, 325-345 x 215-240 um: collar cells elongated

with straight and slightly curved anticlinal walls: testa cells polygonal with

straight anticlinal walls; operculum long nipple-shaped with layers of cells:

anticlinal boundaries broad and flat: the cuticle on the testa consisting

mostly of patches without orientation while in the collar and operculum the

cuticule has a long linear loose striate ornamentation.

Distribution: Known only from the type locality (Fig. 1).

Ecology: Growing on shady river banks and edge of the forest near rivers at

ca. 2240-2300 m.

Notes: This distinct species is somewhat similar to Begonia sikkimensis A.

DC. in leaf shape but differs in having only basal. large and almost glabrous

leaves, female flowers with six tepals, and broad falcate winged fruit. As it is

150 Gard. Bull. Singapore 62 (1) 2010

known only from the type locality, the species is named after the place of its

collection. As the single locality is not under protection and the capability

of the species to cope with less shade and moisture is not likely to be good,

we consider it to belong to the IUCN threat category VUD2.

Begonia taligera S. Rajbhandary, sp. nov. [§ Platycentrum|]

Begoniae hatacoa affinis sed folii margine subintegro usque remote triangulari-

serrulato dentibus interdum attenuatis, floribus haud lineatis, stigmate spiraliter

torto, fructu cum ala laterali longa lataque apice rotundato et in petiolis

radicibus ubi e planta separatis effractisve crescentibus recedit. —Typus:

Central Nepal, Kaski, Bharat Pokhari, near Pokhara 700-740 m, 5 Sep 2007,

S. Rajbhandary and § Adhikari S52 (holotype, E; isotype, KATH). Plate 3.

Perennial, monoecious, caulescent herb, 30-70 cm tall. Stem rhizomatous

at base, subwoody, 3.5 cm in diameter, covered with long roots; erect

portions terete, 17-32 cm long, dark brownish red, branched, hairy. Stipules

persistent, membranous, lanceolate, 6-11 x 7-13 mm, cymbiform, greenish

white, transparent, abaxially glabrous, glossy, apex acuminate. Leaves

cauline, petiole 6-10 (-32) cm long, succulent, densely covered with brown

scales or multicellular two-armed short stalked trichomes, dark reddish

brown at the base becoming ruby red adaxially near the top, readily forming

adventitious roots when detached or broken; lamina asymmetric, cordate,

ovate-lanceolate to elliptic-lanceolate, 10.5-23 x 5.5-18 cm, base slightly

asymmetric, rounded or shallowly cordate, margin entire to attenuate-

dentate, apex long-acuminate to attenuate, deep green adaxially, glossy,

glabrous, yellowish-green abaxially, hairy with veins being covered densely

with brown multicellular two-armed stalked trichomes, palmately veined,

5-7 veins. Inflorescences bisexual, axillary, branched cymose, protandrous,

peduncle glabrous 6-20 cm long, red, bearing few terminal flowers.

Bracts: caducous, in unequal pairs, membranous, cymbiform, long elliptic

to lanceolate, 5-8 x 1.5-3.5 mm, light green, transparent, apex acuminate,

margin entire, glabrous, glossy. Male flowers: pedicel 1.5-3.5 cm, white to

brownish green, glabrous; tepals 4, outer 2 tepals broadly oval, 9-17 x 7-

12 mm, abaxially glabrous and glossy with a green or pinkish tinge more

on the upper tepal, base rounded, margin entire, apex acute; inner 2 tepals

oval, 9-18 x 7-10 mm, white, glabrous, base rounded, apex obtuse or retuse;

androecium a loose sessile globose cluster, 7-9 mm long, golden yellow,

symmetrical, receptacle slightly raised; stamens 90-95, filaments 1-2 mm long

and free; anthers narrowly obovate, 2-2.5 mm, longer than the filaments,

dehiscence through long straight slits, connectives extended, round or acute.

Female flowers: pedicel 1-2 cm, glabrous; tepals 5, white, outer 2 tepals

broadly ovate, 1-13 x 1-12 mm, glabrous, glossy, base rounded, margin entire,

Three New Species of Begonia from Nepal 15]

=

Plate 3. Begonia taligera S. Rajbhandary. A. habit; B. Male flower; C. & D. Androecium: E.

Stamens; F. Rhizome; G. Upper leaf surface; H. Lower leaf surface; I. Bracts: J. Broken petiole

with adventitious roots; K. Female flower; L. Style and stigma; M. Two-armed trichome; N.

Ovary; O. Transverse section of fruit; P. Pollen; Q. Fruit; R. Recurved small wings of the fruit;

S. Seed; T. Androecium.

152 Gard. Bull. Singapore 62 (1) 2010

apex obtuse or rounded, the inner 3 oblong or lanceolate, 1.5-2 x 6-7 mm,

apex obtuse to acute, but sometimes all the tepals are either broadly ovate

or all oblong; styles 2, sometimes 3-4, persistent, 0.4-0.5 mm long, free, each

style ending into a broad apex and connate at the base, green when young

and becoming more yellow when mature; stigmas bifid, spirally tortuous

and papillose all over; ovary oblong, 9-12 mm long, glabrous and glossy,

unequally 3-winged, 2-locular, placentae axile, bilamellate. Fruits: pedicels

1.2-2 cm long, glabrous; 19-25 x 9-15 mm, pendulous, with 3 unequal wings,

the largest wing broadly falcate, 15-25 x 9-14 mm, apex rounded, the other 2

wings smaller and narrow, ca 4-5 mm long with rounded tips, capsule oblong,

10-15 x 3-4 mm, reddish brown when young and pale brown when mature,

glabrous; drying pale brown, dehiscing along the sutures between the two

smaller wings.

SEM studies: Pollen: large, perprolate, pollen grain with smooth convex

sides, rounded poles, compact irregular striate ornamentation with pores in-

between, P 17.4-20 um, E 8 um, P/E 2.1-2.5, elliptical endoporus with costae,

margo present around the endoporus, the grains are not syncolpate. Seeds:

ellipsoidal, mean size 392 x 228 um; collar cells elongated with straight

anticlinal walls; testa cells polygonal with straight and slightly curved

anticlinal walls; operculum very short, flat nipple-shaped with a layer of flat

elongated cells; anticlinal boundaries thin and slightly elevated; the cuticule

with long linear striate ornamentation.

Distribution: Known only from the type locality (Fig. 1).

Ecology: Growing on moist shady rock ledges near streams at ca. 700-740 m.

Notes: Allied to Begonia hatacoa Buch.-Ham. ex D. Don but differs in

having an entire to attenuate-dentate leaf margin, while in B. hatacoa the

leaf margin is usually always entire or only slightly dentate. Begonia taligera

also lacks striped flowers, the stigma is spirally tortuous, the lateral long

wing of the fruit is more broad with a rounded tip, and in the pollen the

ornamentation of the testa cells is compact with irregular striate with pores

in-between and margo present. One of the characteristic features of this

species is the development of roots when a petiole is broken or detached

from the main stem. The epithet taligera is derived from the Latin talea,

and means producing cuttings for propagation. The population is currently

healthy and reproducing well. However, as the species is restricted to a

single locality, which is not under formal protection, and the capability of

the species to cope with less shade and moisture is not likely to be good, we

consider it to belong to the IUCN threat category VUD2.

Three New Species of Begonia from Nepal 153

Acknowledgement

The authors thank the curators of BM, E, K, KATH, MICH and TUCH

for allowing access to herbarium material; Dr. Mark Watson and Dr. Colin

Pendry of the Royal Botanic Garden Edinburgh for their support and

encouragement; the team of the Lower Kanchenjunga-Singalila Ridge

CEPF Project, Eastern Nepal, for their extensive help in the collection of

specimens, and to Dr. Robert Mill for translating the diagnoses into Latin.

The Critical Ecosystem Partnership Fund (CEPF) USA, WWF (Nepal),

University Grants Commission (UGC), and the M.L. MacIntyre Trust are

thanked for providing financial support for this work. Finally, special thanks

go to S. Ranjitkar, K. P. Thapa, S. Adhikari and the S. R. Bista for their help

during field trips.

References

Doorenbos,J.,M.S.M.Sosefand J.J.F.E de Wilde. 1998. The sections of Begonia,

including descriptions, key and species lists (Studies in Begoniaceae V1).

Wageningen Agricultural University Papers 98(2): 1-266.

Hara, H., W.T. Stearn and L.H.J. Williams. 1978. An Enumeration of the

Flowering Plants of Nepal. Vol. 1. Natural History Museum, London.

Press, J.R., K.K. Shrestha and D.A. Sutton. 2000. Annotated Checklist of the

Flowering Plants of Nepal. Natural History Museum, London.

Rajbhandary, S. and K.K. Shrestha. 2009. Begonia flaviflora H. Hara

(Begoniaceae), New Record for Flora of Nepal. Journal of Japanese

Botany 84(1): 16-18.

Shui, Y.-M., C-I. Peng and C.-Y. Wu. 2002. Synopsis of the Chinese species

of Begonia (Begoniaceae), with a reappraisal of sectional delimitation.

Botanical Bulletin of Academia Sinica 43: 313-327.

Smith, L.B., D.C. Wasshausen, J. Golding and C.E. Karegeannes. 1986.

Begoniaceae. Part I: Illustrated Key, Part II: Illustrated Species List.

Smithsonian Contributions to Botany 60. Smithsonian Institution Press,

Washington.

Ye, H.-G., E-G. Wang, Y.-S. Ye and C.-I Peng. 2004. Begonia coptidifolia

(Begoniaceae), a new species from China. Botanical Bulletin of Academia

Sinica 45: 259-266.

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Gardens’ Bulletin Singapore 62 (1): 155-171. 2010 15

in

Genetic Variation of Populations of Scutellaria slametensis

and S. discolor (Lamiaceae) on Gunung Slamet.

Jawa Tengah (Indonesia)

SUDARMONO AND B.J. CONN>

’ Centre for Plant Conservation — Kebun Raya Bogor. Indonesian Institute of

Sciences (LIPI). Jl. Ir. H. Juanda No. 13. Bogor, Indonesia.

; Email: s_darmono@ yahoo.com

National Herbarium of New South Wales. Mrs Macquaries Road. Sydney NSW

2000, Australia. Email: barry.conn@rbgsyd.nsw.gov.au

Abstract

_ Genetic variation within and between populations of Scutellaria slametensis

Sudarmono & B.J.Conn and S. discolor Colebr. on Gunung Slamet (Jawa

Tengah, Indonesia) are evaluated by allozyme electrophoresis. Gels stained

by 4 enzyme systems, namely, Aspartate aminotransferase (Aat). Esterase

(Est). Malate dehydrogenase (Mdh) and Peroxidase (Per). were used to

evaluate the number of polymorphic loci. The mean of total number of

observed alleles per locus (A), mean of total number of effective alleles

per locus (A-), percentage of polymorphic loci (Pp%). and expected genetic

heterozygosity (H-) have been generated as parameters of genetic variation.

The interpopulation genetic differentiation (Fsr) and estimated geographic

distance between populations were used to evaluate the correlation

between genetic differentiation and geographic effect. It was found that S.

slametensis is genetically distinct from S. discolor (D = 1.4572). The mean

genetic variation of S. slametensis (Pp = 75 %,A =2.00, Hz =0.450) is greater

than that of S. discolor (Pp = 25 %, A = 1.25, He = 0.125). Almost all loci of

the latter species are monomorphic and homozygotic. especially population

9 near Baturaden (Pp = 0%: He= 0; Allele frequencies all = 1). There is

a moderately high degree of variation between populations of these two

species (Fst = 0.585, SE + 0.092). whereas within-population variation is

low (1—Fsr= 0.415). Both species are out-breeding (at subpopulation level:

Fis = -0.973, SE + 0.015: and population level: Fir = 0.180, SE + 0.183), with

low levels of gene flow within and between populations (N= = 0.249. SE =

0.065). The chromosome number of S. slametensis and S. discolor is 2n = 24.

156 Gard. Bull. Singapore 62 (1) 2010

Introduction

Scutellaria (Lamiaceae) is the largest genus of the family with about

360 species (Huang, 1994; Paton, 1990, 2004). The genus is widespread,

subcosmopolitan, but poorly represented in moist tropical lowlands. There

are currently four known species in Indonesia, namely, S. discolor Colebr.,

S. indica L., S. javanica Jungh. and S. slametensis Sudarmono & B.J.Conn

(Backer and Backhuizen van den Brink Jr, 1965; Keng, 1978; Steenis van,

1972: Sudarmono and Conn, 2010). The Indonesian species are all members

of subgenus Scutellaria sect. Scutellaria (Paton, 1990) and are informally

classified by Paton into several ‘species-groups:’ ‘S. discolor species-group’

(including S. discolor and probably S. slametensis);*S. humilis species-group’

(including S. javanica); *S. violacea species-group’ (including S. indica).

Allozymes have provided the most abundant source of information

regarding genetic variation in natural populations (Hamrick and Godt, 1990).

Genetic variation parameters such as amount of interbreeding between

populations, allele heterozygosity, genetic diversity, genetic differentiation

and the amount of gene flow can indicate the level of gene mutation, genetic

drift, genetic ‘bottle necks’, and even the possible level of vulnerability of

endangered populations. These parameters are equally useful for assessing

genetic variation within and between populations of a species as they are

between species (Hamrick and Godt, 1990). Maki (1972) and Luzuko et al.

(2000) found that there was a significant correlation between low gene flow

and geographic distance or isolation by distance. Wright (1943) was the first

to describe the genetic process of isolation by distance that may operate

when populations conform to a ‘stepping stone’ model, where gene flow

occurs only between adjacent populations. He demonstrated that random

genetic drift within localized populations, combined with limited migration

among populations, can result in increased genetic differentiation with

increasing spatial distance between populations.

Grant (1981) concluded that geographic isolation was an important

factor that may result in allopatric speciation with reproductive isolation

playing an important role in sympatric speciation. Both of these speciation

phenomena might occur within populations of Scutellaria on Gunung Slamet.

Scutellaria slametensis occurs at elevations of more than 1,000 metres,

whereas S. discolor occurs at elevations of less than 800 m. Chromosome

numbers were recorded and within-population allozyme variation was

analyzed for both species.

Genetic Variation of Populations of Scutellaria slametensis lay)

Materials and methods

Sample collections

Samples of Scutellaria slametensis were collected from six populations on

Gunung Slamet (Jawa Tengah, Indonesia, Fig. 1). These populations occur

at different elevations. Populations | (elevation 1,980 m) and 3 (1,802 m)

occurring on the southwestern slopes, along the Kaligua and Brebes route

in the Perseroan Terbatas Perkebunan Nusantara (PTPN) IX National

Tea Plantation (in Paron and Sokarata Blocks of the Protected Forest,

respectively), Forest Holding Unity Division — Balai Kesatuan Pemangku

Hutan (BKPH), with population 2 occurring closer to the summit of Gunung

Slamet (at 2,002 m) (Table 1). Populations 4-6, from the southern part of

Gunung Slamet, on the Baturaden climbing track occur between elevations

of 1,390 and 2,215 m (Table 1), on the eastern and southeastern slopes.

Samples of three populations of S. discolor were collected at elevations less

than 800 m, from the Baturaden area of Gunung Slamet (Table 1: populations

7-9). Although Keng (1978) concluded that S. discolor occurred over a wide

range of elevations, from 500-2,400 m, this species was not found above

800 m. on Gunung Slamet. Ten individual plants were sampled from each

population. Collections of all populations of both species were cultivated

at Kebun Raya Bogor (Indonesia) and dried herbarium vouchers lodged

at Herbarium Bogoriense (BO), Herbarium of Kebun Raya Bogor (KRB)

and National Herbarium of New South Wales (NSW).

4 Baturaden

eoreters

Figure 1. Distribution map of populations of Scwtellaria included in this study on Gunung

Slamet, Jawa Tengah, Indonesia. Populations 1-6 = S. slametensis; 7-9 = S. discolor, red square

= summit of Gunung Slamet, elevation 3,428m.

158 Gard. Bull. Singapore 62 (1) 2010

Chromosome observation

Growing root tips were incubated in 0.05 % colchicine aqueous solution

for 2 hours at 18°C. They were fixed with the fixative solution (ethanol:

chloroform:glacial acetic acid = 2:1:1) for more than 45 minutes at 5°C. The

root tips were then macerated with 1N HCl at 60°C for 18 seconds. The

meristematic tissues were stained with 2% aceto-orcein for 5-10 minutes on

a glass slide, one drop of 45% acetic acid was added and the tissue covered

and squashed gently.

Allozyme analysis

Within-population allozyme variation was analyzed by enzyme systems

for six populations of Scutellaria slametensis and compared to the within-

population variation found within three populations of S. discolor, both

occurring on Gunung Slamet (Jawa Tengah, Indonesia, Figure 1).

Leaf samples were prepared for allozyme electrophoresis by extracting

0.5 cm’ samples of fresh leaf material. Young leaves were ground with 0.1 M

Tris-HCl pH7.5,0.1% 2-mercaptoethanol,0.001 M EDTA (tetrasodium salt).

Extract was absorbed by filter paper (Whatmann No.3) andrun by 12% starch

gel (4.5 hours, 300 volt; 45 mA). Four horizontal enzyme systems analyzed

were Aspartate aminotransferase (Aart; EC 2.6.1.1), Esterase (Est; EC 3.1.1),

Malate dehydrogenase (Mdh; EC 1.1.1.37) and Peroxidase (Per; EC 1.11.1.7).

Staining procedure followed Soltis et al. (1983), with some modification in

buffer pH and concentration (Mdh buffer with 1.5 M TRIS-HCI pH 8.0).

The locus specifying the most anodally migrating isozyme was designated

as 1, the next 2, and so on. Similarly, the most anodally of a gene was labeled

‘a’, the next ‘b’ and so on, as done by Shield er al. (1983) and Kephart (1990).

The genetic variation of populations is expressed as a percentage

of polymorphic loci (Pp %), mean number of observed alleles per locus

(A), and Nei’s unbiased genetic diversity or expected heterozygocity (/7.),

using POPGENE version 1.32 (Yeh et al., 1999). Gene flow (Nm — the

number of migrations per generation) for all loci was also estimated. The

analysis of genetic identities (7) and genetic distance (D) for each pair-wise

combination of populations were also estimated following Nei (1978). In

this study, unbiased genetic identity was used to accommodate for the bias

caused by small sample size (<50 individuals).

There are several indices used to evaluate genetic diversity, including

total genetic diversity (Hr), genetic diversity within populations (Hs),

genetic diversity among populations (Ds7), and the among populations

genetic differentiation coefficient (Gsr). These statistics were generated

using FSTAT program (Goudet, 2002). Total genetic diversity (7/7) was

partitioned into within-population (Hs) and between-population (Dsr)

components. Hr was estimated for each polymorphic locus from following

Genetic Variation of Populations of Scutellaria slametensis 159

equations:

Hr =Hs+ Dsr

oP)

v= 5

Where, ji is gene identity in sub-population and S is number of sub-

populations. ji was estimated by:

ji = Sa

Where, xis the frequency of the Ath allele in ith sub-population

Genetic diversity between sub-populations (Ds) was estimated by:

Dsr= ( Ly ID¥), Si

1

Dy= ) 5 Pa-Pa¥

Where, Pix is the frequency of the kth allele in ith sub-population, while Pi

is frequency of kth allele in jth sub-population.

Gene differentiation that occurs among populations (Gsr) was examined

with Nei’s genetic diversity indices (Nei, 1977, 1986). Gsr was expressed

relative to total genetic variations among populations as:

Dsr

Gsr =

Mean values of Hr, Hs, Dsr and Gsrwere the average of all polymorphic

loci within each group. All calculations were generated for each pair-wise

comparison of populations and species using FSTAT program (Goudet,

2002).

The genetic structure of the studied populations was also analyzed in term of

the following F-statistics: Fis - fixation index related to non-random mating

within populations; Fir - mean inbreeding coefficient, Fs7 - interpopulation

genetic differentiation (following Weir and Cockerham, 1984).

Gene flow (N,,) for all loci was estimated:

_G~6y)

N,

m 4Gsr

160 Gard. Bull. Singapore 62 (1) 2010

Assuming that populations have reached equilibrium between the effects

of migration and random genetic drift, the degree of population subdivision

was quantified using:

1

Fy =>

st” (4N,, + 1)

Fsr is the proportion of total genetic variance contained in subpopulation

S relative to total genetic variance 7. Unbiased data matrices (Nei, 1986)

were generated using POPGENE (Yeh et al. 1999). It was also used to

calculate mantel test (Sokal and Rohlf, 1995) for testing the null hypothesis

of independence between genetic differentiation and geographic distances

separating populations (Yeh et al., 1999). Allozyme data were analyzed

using UPGMA (Unweighted Pair-Group Method using Arithmetic

Average) clustering techniques to construct a dendrogram to assist in the

interpretation of these genetic data between species and populations. The

dendrograms were generated using NTS YS (Rohlf, 2000).

Results

Chromosome analysis

Chromosome number of Scutellaria slametensis (Figs.2A & B) and S. discolor

(Figs. 2C & D) are both diploid 2n = 24. The length of mitotic metaphase

chromosome of S. slametensis varies from 2-2.5 um, whereas those of S.

discolor are smaller (1.5-2 um).

Figure 2. Mitotic metaphase chromosomes of Scutellaria slametensis (A, B) and S. discolor

(C, D), both 2n=24. A & C, microphotographs; B & D, line-drawings of microphotographs.

Scale bar = 5 um.

Genetic Variation of Populations of Scutellaria slametensis 161

Genetic Variation

Four loci were detected in all populations of both species of Scutellaria

(Table 1). The four enzyme systems examined (namely, Aat, Est, Mdh and

Per) were consistent for all nine populations studied. Populations 1, 4-6

(Table 1) are genetically highly variable with 75% of loci polymorphic (Pp).

Population 4 has the highest mean number of alleles per locus (A = 2.00),

with a high mean number of A=1.75 for populations 1,5, 6 (Table 1). The

expected heterozygosity is also regarded as high for these four populations

(population 1: He= 0.375; 4: He = 0.401; 5: He= 0.375; 6: He = 0.375) (Table

1). In contrast, the lowest genetic variation occurs in population 9 (Pp= 0

%,A = 1.00, He = 0.000) (Table 1). Others populations that are regarded as

having low genetic variation include: population 2 (Pp = 25 %, A=1.25, He=

0.139) and population 8 (Pp = 25 %, A = 1.25, He = 0.125).

Table 1. Mean genetic variation of populations of Scutellaria slametensis and S. discolor

on Gunung Slamet (Jawa Tengah, Indonesia). Ten samples of each population (N) were

examined. Populations 1-6 = Scutellaria slametensis; 7-9 = S. discolor. Percentage polymorphic

loci (Pp), mean number of observed alleles per locus (A), mean of effective alleles per locus

(Ae), mean of expected heterozygosity (He) (Nei, 1978). *Comparison of genetic variation

categorized at regional geographic distributed means Pp=36.4 %, A=1.55, Ae =1.16 and He

= 0.118 (Hamrick and Godt, 1990).

Eep Location eititnde Geocode Pp% A Ae He Category *

no. (m)

1 Protected forest block 7.2683° S z 2 2 ;

: : : fo Ths) 1S: 75 0.375 high

Paron, Kaliwadas oe 109.1584° E Te : 's

2 __ Protected forest block, A pe SUIS ae - z

, 2,002 ae 25 1255 125) 05139) Sbow

Sakub, Kaligua a 109.1456° E

3 Protected forest block 7. 2829° S 2 P E e

: , 1,802 z 50 =1.50 1.50 0.250 Low

Sokarata, Kaligua 109.1298° E

4 Protected forest, Petak II 1,390 eS 75 2.00 1.88 0.401 high

2 s 109.2004° E a = i

5 Protected forest, Post III, 2215 7.2668° S 75 1.75. 1.75 0.375 high

Baturaden 109.2011° E

6 Protected forest, Post I 7.2668° S : = ae =e

z 3 5778 Lee 75 1.75 1.75 0.375 high

Baturaden 109.2023° E

7 Camping ground, 7.3119°S = Bs Z =

rie 771 ore 50 1.50 1.50 0.250 Low

Baturaden 109.2351° E

| 30522 S egy er ee

8 Baturaden Botanic Garden 771 109.2333°E 1.25 1.25 0.125 Low

7.3069°S

9 Telaga Sunyi, Baturaden at 0 1.00 1.00 0.000 low

109.2427° E

162 Gard. Bull. Singapore 62 (1) 2010

Population structure and gene flow

Genetic diversity indices (Table 2) demonstrate that Aat-] and Mdh

contain the highest total diversity (Hr) of those loci surveyed with 62.1%

and 64.8%, respectively. The enzyme Est-1 contained the least genetic

diversity (Hr= 44.4%). The mean of total genetic diversity (H7) was 58%

(SE +0.030), whereas genetic diversity within populations (Hs) was 26.7%

(SE +0.058) and among populations (Dsr) 31.3% (SE +0.034). There was no

genetic diversity within-populations in the Est-/ locus (Hs= 0%). Likewise,

the genetic diversity within and among populations of Aat-/ was low (Hs=

29.1% and Dsr = 35.1%, respectively). Although the genetic diversity within-

populations of Mdh and Per were similar (both Hs = 38.9%), the among

population genetic diversity was Dsr= 25.9% and Dsr= 19.8%, respectively.

The mean genetic differentiation between populations (Gs7) is 57.1% (SE

+ 0.095).

Table 2. Genetic diversity indices, F-statistic and estimation of gene flow between populations

of Scutellaria. Hs, the genetic diversity within populations; Dsr, the genetic diversity among

populations; Hr, the total genetic diversity; Gsr, the among populations gene differentiation

coefficient; Fis, the fixation index related to non random mating within populations; Fir,

the mean inbreeding coefficient of a set of a populations; Fs7, the interpopulation genetic

differentiation; Nm, gene flow estimated from Fsr = 0.25(1 - Fsr)/Fsr. SE, standard error.

Genetic diversity indices F-statistic Gene flow

LOGTS Hs Dsr Hr Gsr Fis Fir Fsr Nm

Aat-1 0.291 0.351 0.641 0.547 | -0.919 0.135 0.549 | 0.205.

Est-1 0.000 0.444 0.444 = 1.000 ht! 1.000 1.000 0.000

Mdh 0.389 0.259 0.648 0.400 | -1.000 -0.091 0.455 0.300

Per 0.389 0.198 0.587 01337 | -1.000 -0.326 0.337 0.492

Mean 0.267 0.313 0.580 0.571 | -0.973 0.180 0.585 0.249

SE 0.058 0.034 0.030 0.095 | 0.015 0.183 0.092 0.065

The estimation of the genetic variation among and within populations

indicates a moderately high degree of differentiation among populations

(Fst = 0.585; SE +0.092), whereas variation within-populations (1-Fs7)

represented 41.5% of the total variance. These species exhibit a considerable

degree of out-breeding at both the subpopulation (Fis = -0.973; SE +0.015)

and population levels (Fir = 0.180; SE +0.183), even though the estimate

of mean gene flow was low (Nm = 0.249; SE +0.065). However, the null

hypothesis that these populations are at equilibrium is rejected because

there is no correlation between pair-wise genetic differentiation values (Fs7)

and geographic distance (r = 0.365; Mantel t-test= 2.259; p < 0.01) (Fig. 4).

Genetic Variation of Populations of Scutellaria slametensis 163

Based on genetic distance (Nei 1978), two population-groups are

differentiated (Fig. 3), representing Scutellaria discolor (populations 7-9)

which is genetically distinct from all populations of S. slametensis (Genetic

distance, D=1.4572). Within the populations of the latter species, populations

1 (Paron block, Brebes) and 5 (population in Post III, along walking track

from base camp Baturaden to summit of G. Slamet) are genetically similar

(Fig. 3). Likewise, populations 4 (Post II) and 6 (Post I) are also genetically

similar and this latter population pair is genetically close to the previous

pair. Populations 8 and 9 (S. discolor) are also genetically similar (Fig. 3).

Allele frequency of 9 populations and allele shared among species

Most alleles are relatively common within the sampled populations

(occurring in at least 50% of loci) (Table 3). However, two alleles with low

frequencies in population 4 are Aat-1 with only ahs of chromosomes carry

this allele and Aar-I" (35%). Alleles Aat-1 , Mdh-I' and Per’ were shared

between S. slametensis and S. discolor, whereas, Aat-1° ’, Est-I° and Per" were

specific to S. slametensis. Alleles Est-1 and Per are specific to S. discolor.

Table 3. Allele frequency of 4 polymorphic loci in 9 populations of Scutellaria. Populations

1-6 = S. slametensis; 7-9 = S.discolor

Population Aat-I’ Aat-I’ Aat-I' Est-I° Est-I’ Mdh-I' Mdh-I’ Mdh-I' Per" — Per’ Per’

1 0.50 0.50 1.00 0.50 0.50 0.50 0.50

2 1.00 1.00 1.00 0.50 0.50

3 0.50 0.50 1.00 0.50 0.50 1.00

4 0.50 035 0.15 0.50 0.50 0.50 0.50

5 0.50 0.50 1.00 0.50 0.50 0.50 0.50

6 0.50 0.50 1.00 0.50 0.50 0.50 0.50

7 1.00 1.00 0.50 0.50 0.50 0.50

8 1.00 1.00 1.00 0.50 0.50

9 1.00 1.00 1.00 1.00

Discussion

Paton (2004) noted that the chromosome numbers for Scutellaria are 2n=12-

88, with most frequent numbers being 2n=20, 22, 24, 32 and 34. Chromosome

numbers of Scutellaria subgenus Scutellaria, which includes all Indonesian

species, are 2n=24-34 (Paton, 1990). Since the chromosome numbers of S.

discolor and S. slametensis are both 2n=24, these two species are possibly

closely related. Both of these species have small chromosomes, less than 5

um long.

Allozymes have been successfully used to compare mating system,

migration and local differentiation within and between populations (Brown,

164 Gard. Bull. Singapore 62 (1) 2010

1990). Factors such as, regional distribution of a taxon, geographic range,

breeding system, seed dispersal mechanisms and successional status

have been associated with differences in the percentages of polymorphic

loci (Pp%), mean number of observed alleles per locus (A), and genetic

diversity within-populations (He) (Hamrick and Godt, 1990). Values

of Pp%, A and He obtained for S. discolor and S. slametensis are very

similar to those obtained for out-breeding plants, wide-spread species,

and for those with seeds that are dispersed by gravitational forces.

Scutellaria slametensis is a species with a restricted distribution and

has mean genetic variation (Pp= 75 %, A = 2.00, He = 0.450) equivalent to

that of S. montana (Pp = 75.42 %, A = 2.21, He = 0.19; see Cruzan, 2001),

a species restricted to parts of Georgia and Tennessee (U.S.A.) (Cruzan,

2001). Scutellaria slametensis exhibits greater genetic variation than plants

of S. discolor sampled from G. Slamet. Almost all loci of the latter species

are monomorphic and homozygotic, especially population 9 (Pp = 0 %; He

= (); Allele frequencies all=1).

The value of allozyme electrophoresis in delimiting taxa has been

found to be useful by several researchers (such as, Coates and Hnatiuk,

1990; Crawford, 1985; Gottleib, 1984). Although all populations of S. discolor

and S. slametensis are relatively close geographically, they are genetically

distant based on allozyme divergence. This is possibly a result of the steep

mountainous terrain restricting the potential pollinators tosmall geographical

areas within the region (Tyler, 2003). Within S. s/ametensis, populations 4

and 6 are genetically and geographically close. Population 5 is unexpectedly

genetically close to population | even though the two are not geographically

close. Together, populations | and 5, and 4 and 6 form a genetically distinct

grouping (Fig. 3). Although the two high altitude populations, 5 (elevation

at 2,215 m) and 2 (2,002 m) are geographically relatively close, these two

populations are genetically distant (Fig. 3). Although population 1 is located

in between populations 2 and 3, the genetic variation, hence gene flow,

within the latter two populations is low suggesting that they do not share

a common population of pollinators. Contrary to this, within-population

variation in the population group | and 4-6 is high, representing high

gene flow between these populations (Table 1). The three populations of

S. discolor (populations 7-9) are both geographically close and genetically

similar. Species characterized by a low level of gene-flow, such as S. discolor

(Table 1), or a low level within or between populations, such as populations

2 and 3 (S. slametensis), may indicate high levels of selfing. Cruzan (2001)

hypothesized that the smaller fragmentation threshold may reflect higher

levels of selfing in isolated populations because of the absence of pollinators.

Unpublished results from observations of floral behaviour and crossing

tests of S. slametensis clearly demonstrate that this species is mostly self-

—

Genetic Variation of Populations of Scutellaria slametensis 165

compatible as has been found in S. indica (Sun, 1999). However, within the

current study, the low overall proportion of total genetic variance within a

subpopulation relative to total genetic variance (Fsr= -0.9769) implies a low

level of differentiation among populations, suggesting that S. discolor and S.

slametensis are out-breeding. The low level of gene flow (mean Nm = 0.1988)

observed suggests that although these populations are geographically close,

they are relatively isolated genetically. Slatkin (1987) concluded that values

of N» < 1 mean that genetic drift will result in substantial differentiation

between populations. This is possibly a consequence of pollinators only

visiting flowers within a restricted area, hence mostly visiting plants in

closely adjacent areas (such as, between populations 4 and 6, and between 8

and 9). However, this does not explain the lack of genetic distance between

populations 1 and 5 (D = 0.000).

]

5

4

6 S. slametensis

3

2

: S. discolor

8

9

0.02 0.35 0.71 1.08 1.45

Figure 3. Dendrogram of Nei’s genetic distance (Nei, 1978) between six populations of

Scutellaria slametensis and three populations of S. discolor on Gunung Slamet, Jawa Tengah,

Indonesia.

Mean total genetic diversity (Hr) showed majority of partitioning

occurred among populations (Dsr= 31.5%) rather than within populations

(Hs = 0.267). The large amount of genetic differentiation among populations

166 Gard. Bull. Singapore 62 (1) 2010

(Gsr = 0.571; Fsr = 0.585) indicate strong genetic differentiation between

these populations (Table 3). F-statistics have similar values and indicate that

populations of S. discolor and S. slametensis are genetically structured (Table

2). High levels of genetic structure within-populations are supported by the

Est-I locus but with low levels of gene flow (Nm) (Table 2). High levels

of pollen transfer indicate movement of pollinators that will naturally lead

to increased genetic differentiation but decreased gene flow (Ellstrand and

Elam, 1993). If genetic variation within species is predominantly affected

by shared alleles, then there may be a significant correlation between

shared alleles and the local geographical distribution pattern of the species.

An understanding of nature of shared allele may be used to understand

conservation implication for future.

Throughout this study we have assumed a ‘stepping-stone’ model

(Kimura, 1953) of population structure among these plants whose dispersal

ability is constrained by distance such that gene flow is most likely to occur

between neighbouring populations (Hutchinson and Templeton, 1999).

Consequently, it would be expected that adjacent populations tend to be

more genetically similar than more distant populations. Therefore, assuming

a‘stepping-stone’ model of regional population structure, the null hypothesis

that these regional populations are at equilibrium is rejected because there

is no association between pair-wise genetic differentiation values (Fs7)

and geographic distance (r = 0.365; Mantel t-test= 2.259; p < 0.01) (Fig. 4).

In this study, the populations of Scutellaria on Gunung Slamet consist of

mostly small, more or less isolated populations, such that allele frequency

drifts independently of geographic distance between populations and is

much more influential in determining the population structure than gene

flow. These results are consistent with limited pollination events occurring

between populations and/or the inhibition of dispersal in this mountainous

terrain.

The dendrogram of the sampled populations (UPGMA - based on Ne1’s

genetic distances; Nei, 1978) indicates a correlation between shared alleles and

genetic distance (D) (Fig. 3). The topology of this dendrogram is congruent

with morphological differences between S. discolor and S. slametensis. Based

on allozyme data, the greatest genetic distance (D =1.45) was found between

S. discolor and S. slametensis. There are several morphological features that

distinguish S. discolor and S. slametensis (Sudarmono and Conn, 2010). The

shape of the leaf, height of stem and number of flowers at inflorescence

nodes of S. discolor are very different from those of S. slametensis. Scutellaria

discolor has flowers with corolla purple, whereas, S. slametensis has corollas

that are white basally, pink distally (populations 1 & 2) or purple distally

(populations 3-6). Scutellaria slametensis has obovate leaves, whereas

those of S. discolor are ovate. The former species has two flowers at each

node of the inflorescence, whereas S. discolor has four flowers per node.

Genetic Variation of Populations of Scutellana slametensis 167

0.63

3

S 0.47

=

:

ps 0.32

=

2 0.16

E .

0.00 AE 5.45 8.18 10.90

Geographic distance (km)

Figure 4. Correlation between geographic distance (km) and genetic differentiation (Fs7)

of nine populations of Scutellaria discolor and S. slametensis on Gunung Slamet. r=0.365:

Mantel f test = 2.259: p < 0.01.

Conclusion

Scutellaria discolor and S. slametensis are morphologically and genetically

distinct. Genetic variation within populations 1, and 4-6 are high (S.

slametensis), whereas within populations 2, 3 (both S. slametensis), and 7-9

(S. discolor) are low. Although the genetic diversity within S. slametensis

is partitioned, with one group with high genetic variation (populations 1,

4-6) and one with low values (populations 2 and 3). overall this species has

a high level of genetic heterozygosity that makes it a species with a high

frequency of heterogamy. Consequently, from a conservation point of view,

this species is probably a low-risk species (Fracaro and Echeverrigaray.

2006). Even though the two species at the subpopulation level (Fis = -0.973:

SE +0.015) and population level (Fir = 0.180; SE +0.183) are out-breeding,

the mean gene flow was low (Nm = 0.249; SE +0.065). Historically, genetic

drift has affected the overall structure of the populations of this species,

rather than gene flow. However, within the distribution of S. slametensis, the

mean gene flow was high for the local populations 1. 4-6. Since this species

is only known by a limited number of individuals in a restricted locality, it

may prove to be vulnerable (IUCN, 2001) should the populations become

more isolated, resulting in the level of gene flow and hence, heterozygosity

becoming further reduced.

168 Gard. Bull. Singapore 62 (1) 2010

Although S. discolor is a widespread species, all loci of the plants

sampled on Gunung Slamet are monomorphic and homozygotic. An

assessment of the genetic variability of this species throughout its range

is required. Even though a species may be widespread, an increased

homozygosity may result in a reduction of the vigor of individuals, expression

of deleterious characters, increased seed abortion, reduced fertilization and

germination rates. These are factors that may lead to the disappearance of

the populations (Dubash and Fenster, 2000).

Acknowledgements

The authors are much indebted to P. Daryono, P. Budiyanto, and P. Kartam

who provided field assistance in the Protected Forest of Gunung Slamet from

Kaligua, Kaliwadas and Baturaden. We thank Ir. Sinung Nugroho (BKPH

Gunung Slamet Barat), Dr. Irawati (KRB) for permission to undertake

this study, and Prof. Mien A. Rifai (BO) for comment on this manuscript.

Prof. Dr. Syamsuardi, (Andalas University, Sumatera Barat, Indonesia) for

critical comment on technical analysis of allozymes. The first author received

financial assistance from DIPA-LIPI Pelaksanaan Kegiatan Penelitian Hasil

Seleksi Workshop Kiat-Kiat Memenangkan Proyek TA 2008.

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species from Central Java, Indonesia. Telopea 12(4): 463-468.

Sun, M. 1999. Cleistogamy in Scutellaria indica (Labiatae): effective mating

system and population genetic structure. Molecular Ecology 8: 1285-

1295:

Tyler, T. 2003. Allozyme variation in Carex sect. Digitatae — Evidence

of introgression, genetic distinctiveness and evolution of taxa. Plant

Systematic and Evolution 237: 219-231.

Weir, B.S. and C.C. Cockerham. 1984. Estimating F-statistics for the analysis

of population structure. Evolution 38: 1358-1370.

Wright, S. 1943. Isolation by distance. Genetics 28: 114-138.

Yeh, F.C., R. Yang and T. Boyle. 1999. POPGENE version 1.32 — Population

Genetics Analysis, University of Alberta and Center for International

Forestry Research, Alberta.

Gardens’ Bulletin Singapore 62 (1): 173-176. 2010 173

A New Species of Polyalthia (Annonaceae) from Sabah

I.M. TURNER

Research Associate, Singapore Botanic Gardens, and

Royal Botanic Gardens, Kew

Abstract

Polyalthia lasioclada 1.M. Turner, sp. nov. is described. It is a small tree

known from the Mount Kinabalu area of Sabah.

When looking through some Annonaceae specimens recently collected from

the Kinabalu Park area of Sabah, I realised several were the same thing as

an odd specimen I had tentatively placed in Polyalthia microtus Miq. The

leaves drying greyish with auriculate bases and orange-red corollas clearly

pointed to membership of the informal ‘Polyalthia insignis species-group’

of Johnson and Murray (1999). The unusual feature of the specimens was

the villose twigs reminiscent of Polyalthia borneensis Merr. and P. bullata

King rather than typical P. microtus. The availability of more collections

confirmed that there were some consistent differences in flower and fruit

form that separated the shaggy-twigged plant and therefore I here describe

it as a new species.

Polyalthia lasioclada \.M. Turner, sp. nov.

Polyalthiae microtus ramulis_ petiolisque_ villosis, sepalis minoribus,

monocarpiis globosis vix cylindricis, in sicco aureobrunneis laevibus vix

brunneis verruculosisque differt. -Typus: Malaysia. Borneo, Sabah, Ranau

District, west of Kg Takutan, 2500 ft, 25 May 1973, G. Shea & Aban, SAN

77174 (holotype, SING; isotypes, K, L). Plate 1.

Small tree. Twigs drying dark grey or grey-brown with shallow longitudinal

grooves and some cracking, youngest parts brown or reddish brown densely

covered with long (3-4 mm) more or less straight golden brown hairs. Leaves

chartaceous to subcoriaceous, drying grey or grey-brown above, brown

beneath, glabrous except for long straight hairs on midrib below, densest near

base, midrib and lateral nerves immersed above, prominent beneath, lamina

elliptic to elliptic-obovate, 4.5-20 x 1.5-7.7 cm, apex acute to shortly acuminate,

base auriculate, lateral veins 15-19 pairs, angled or arching forward, looping

distinctly via a distinct intramarginal, tertiary venation reticulate. Petioles 1-

174

Plate 1. Photograph of the holotype of Polyalthia lasioclada 1.M. Turner, sp. nov.

Gard. Bull. Singapore 62 (1) 2010

FLORA OF SABAH

HERBARIUM OF TILE FORPSS VEYARTMENT SANDARAN

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A new species of Polyalthia 175

5 mm long, 1-3 mm thick, densely villose. Inflorescences single-flowered,

intranodal. Flowers with pedicel 12-24 mm long, less than 1 mm wide,

drying red brown with scattered pale hairs, faintly wrinkled longitudinally,

sepals ovate or triangular 1-2 x 2-3 mm, drying brown with pale hairs

externally, glabrous within, petals lanceolate 15-30 mm long, orange-red,

outer slightly wider than inner, 5 vs 3-4 mm, drying dark brown or almost

black, outside slightly wrinkled longitudinally, minutely pimpled near the

base with sparsely scattered short white hairs, inside smoother with hairs

confined to the apex. stamens many ca 2 mm long, connective apex truncate,

carpels ca 12, ca 1.5 mm long, densely pale hairs. Fruits with pedicel ca 18

mm long, 1 mm wide, drying reddish brown, monocarps ca 12, globose, 8-

10 mm diameter, apex apiculate, drying golden brown, sparsely adpressed

pale hairy, drying relatively smooth, stipe 5-8 mm long, | mm thick. Seeds 1-2,

drying pale brown.

Specimens seen: MALAYSIA. Sabah, Kota Marudi District, Kampung

Monggis, 4 batu dari pusat Kampung Monggis Utara, 9 Apr 1996, Matamin

Rumutom 186 (K); Kampung Kawasan Taman 100 m dari Sungai Mokodon,

8 Mar 1996, Daim Andau 344 (K); Kampung Kawasan Taman, Jalan ke Palu

Agayo, 16 May 1995, Kinsum Bakia 44] (K); Ranau District, Kampung

Nalumad 2 batu from Kg Nalumad, 3 Oct 1998, Daim Andau 1020 (K);

Ranau District, Kampung Nalumad 5 batu dari Kg Nalumad, 9 Oct 1996,

Daim Andau 855 (K).

Notes: The chosen specific epithet is derived from Greek (/asios = shaggy,

woolly, clados = branch, shoot) and reflects the characteristic villose twigs

of the species.

Polyalthia lasioclada is similar to the widespread and rather

variable Polyalthia microtus, particularly in flower form. The sepals of P.

lasioclada are generally smaller than those of P. microtus (1-2 x 2-3 mm

versus 4-6 x 4-6 mm). The monocarps of P. lasioclada are dry globose,

relatively smooth and a golden brown, whereas those of P microtus are

more cylindrical, dry with the surface minutely warty and dark brown.

The villose twigs of Polyalthia lasioclada are similar to those of P.

borneensis and P. bullata. However these species both have cream to yellow

flowers with short villose pedicels rather than the orange-red corollas and

relatively long, sparsely hairy pedicels of P. lasioclada. The many-nerved

leaves of P. bullata (25-40 pairs) are unlikely to be confused with P. lasioclada

(15-19 pairs).

176 Gard. Bull. Singapore 62 (1) 2010

Acknowledgements

Many thanks to Dr J.R Veldkamp (L) for translating the Latin diagnosis.

Support for this reasearch from the Arnold Arboretum, Forest Research

Institute Malaysia, Singapore Botanic Gardens and Royal Botanic Gardens

Kew is gratefully acknowledged.

References

Johnson, D. M.and N.A. Murray. 1999. Four new species of Polyalthia from

Borneo and their relationship to Polyalthia insignis. Contributions from

the University of Michigan Herbarium 22: 95-104.

Gardens’ Bulletin Singapore 62(1): 177-209. 2010 aT

Studies on Schismatoglottideae (Araceae) of Borneo XIII:

A Revision of the Schismatoglottis nervosa

Species Complex

WONG SIN YENG

Department of Plant Science and Environmental Ecology, Faculty of

Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Samarahan, Sarawak, Malaysia

Abstract

A revision of the Schismatoglottis nervosa Ridl. species complex is presented.

Ten species are recognized, of which three species are pre-existing (S.

nervosa, S. elegans A.Hay, and S. brevicuspis Hook.f.) and seven are novel

and described here (Schismatoglottis adoceta S.Y.Wong, S. linae S.Y.Wong, S.

matangensis S.Y.Wong, S. simonii S.Y.Wong, S. tessellata S.Y.Wong, S. turbata

S.Y.Wong, and S. ulusarikeiensis S.Y.Wong.) The S. nervosa species complex

is readily delimitated by the pungent terpenoid smell when the vegetative

tissues are crushed, and by the presence of longitudinally ridged petioles.

A key to the S. nervosa species complex is presented and all species are

illustrated.

Introduction

Schismatoglottis is a genus of in excess of 150 species of terrestrial, lithophytic,

and rheophytic herbs occurs within Malesia, but mainly in Borneo. Hay &

Yuzammi (2000) presented an alpha-taxonomic revision of the Malesian

species, one of the results of which was the recognition of six informal

species groupings based primarily on shoot architecture and the senescence

mechanics of the upper spathe: Asperata Group, Calyptrata Group, Corneri

Group, Multiflora Group, Rupestris Group and Tecturata Group. However,

Schismatoglottis has been proven as a polyphyletic assemblage (Wong er

al., 2010), and this has resulted in the removal of the Rupestris Group into

a resurrected genus, Apoballis (Wong & Boyce, 2010). Further molecular

analysis is being undertaken and will form the basis of further paper (Ting

et al., in prep.).

Hay and Yuzammi (2000) placed S. nervosa and S. elegans (both from

Sarawak) in the Asperata Group based on the combination of pleionanthic

shoots, the leaf sheath open and nearly always fully attached and persistent,

and by the spathe limb opening more-or-less wide and then crumbling-

178 Gard. Bull. Singapore 62 (1) 2010

deliquescent. One species, which occurs in West Malaysia, Schismatoglottis

brevicuspis Hook.f also belongs to this complex. In addition to these

species, a further 7 species, all novel, are recognized here from Sarawak:

Schismatoglottis adoceta S.Y.Wong, S. linae S.Y.Wong, S. matangensis

S.Y.Wong, S. simonii S.Y.Wong, S. tessellata S.Y.Wong, S. turbata S.Y.Wong,

and S. ulusarikeiensis S.Y.Wong. The S. nervosa species complex is clearly

distinguished within the Asperata Group by a pungent terpenoid smell

when crushed (tissues odourless when crushed in Asperata Group s.s¢r.,

i.e., sensu Wong), coriaceous to thinly coriaceous + elliptic leaves with veins

prominent abaxially (leaves thinly sub-succulent and fragile with veins

obscure to invisible abaxially in Asperata Group s. str), and longitudinally

ridged petioles (petioles asperous to puberulent or hispid in Asperata Group

Ss. Str.), Supporting the recognition as a distinct species complex within the

group.

The S. nervosa species complex comprises terrestrial forest-dwelling

herbs commonly on steep soil banks or occurring as obligate or facultative

lithophytes on limestones, shales, sandstones and granite. Schismatoglottis

nervosa and S. elegans are restricted to Karst limestones; Schismatoglottis

matangensis and S. turbata to sandstones, and S. adoceta, S. tessellata and S.

ulusarikeiensis to shales, S. simonii to both limestones and sandstones, and S.

linae and S. brevicuspis to granite.

Allied taxa

There are additional species in the S. asperata group that possess

longitudinally and/or puberulent ridged petioles and/or are weakly (not

pungently) terpenoid smelling when crushed, of which one, Schismatoglottis

latevaginata Alderw., described and numerous other species are awaiting

formal description. Based on initial observations of the spathe senescence

mechanics (spathe at least partially circumscissile before shedding in large

pieces) and leaf shape (lamina oblongo-ovate on a disproportionately long

petiole), these species are morphologically more closely related to one

another than any of them are to species of the S. nervosa species complex,

and will be the subject of a subsequent paper.

Schismatoglottis nervosa species complex

Small to robust herbs, 40-70 cm tall, occasionally up to ca 1 m tall, with

vegetative tissues emitting a pungent aromatic (terpenoids) smell when

crushed. Stem epigeal, pleionanthic, erect to decumbent, sometimes

ascending and then rooting (‘terrestrial climber’ sensu Boyce), 30-50 cm

Studies on Schismatoglottideae 179

long x 2-4 cm thick, adventitiously branched from older portions, rooting

along entire length on contact with ground, roots often penetrating petiole

bases; leaf scars prominent. Leaves few to many (5-15) together; innovations

yellowish green; senescent lamina sometimes rotting and falling together

with distal portion of petiole to leave the lower portion of the petiole

attached to plant, this rotting and falling at a later stage; petioles terete, ca

30-40 cm long, sometimes, up to ca 70 cm long, adaxially channelled or D-

shaped in cross section, weakly to strongly longitudinally ridged (resembling

celery - Apium graveolens - Apiaceae) especially abaxially, minutely (strong

lens required) and densely verruculate or glabrous; petiole sheathing

in the lower '4- %, sheaths fully attached, thinly coriaceous, sometimes

leathery, marcescent or persistent, tapering, closed or less often wide open,

sometimes with a short rounded free ligular portion; lamina broadly ovate

to oblongo-ovate to elliptic, coriaceous or thinly coriaceous, base broadly

rounded to sub-truncate, slightly retuse or cuneate, apex acute to strongly

acuminate, adaxial surface semi-matte, bright deep to medium green, always

slightly bullate, abaxial surface paler green, often glaucous, drying strongly

discolorous; midrib adaxially flush with lamina, abaxially very prominent;

primary venation impressed adaxially, sometimes flush with lamina, strongly

raised abaxially, alternating with lesser interprimaries, interprimaries

occasionally arising from the bases of the primary veins, both diverging

at 45°- 90° and gradually curving towards the apex before reaching the

intermarginal collecting vein; secondary venation mostly arising from the

midrib, occasionally from near the bases of the primary veins, prominent

abaxially; tertiary venation forming an obscure to prominent tessellate

pattern, variously prominent adaxially and abaxially. Inflorescences up to

three together (rarely 4), erect, white, moderately fragrant (esterase) at

female anthesis; peduncle to ca 2 cm long, concealed by leaf bases, prophylls,

and cataphylls at flowering, slightly exserted in fruit; spathe interior glossy,

exterior semi-glossy; lower spathe differentiated from the limb by a weak

constriction correlating with spadix interstice; limb oblongo-lanceolate, white

to partially green, weakly coriaceous to somewhat spongy, semi-truncate to

shortly to strongly acuminate (to ca 5 mm long), apex mucronate (to ca 2

mm), limb either caducous by crumbling at or just after male anthesis, with

remaining fragments deliquescing, or briefly persistent until end of anthesis

and thence deliquescent; spadix sessile, isodiametrically adnate on the lower

abaxial side relative to the spathe opening; female zone conic-cylindric;

pistils numerous, close-packed or laxly arranged; stigma sessile, punctiform,

minute to large but always smaller in diameter than ovary; interpistillar

staminodes confined to a more-or-less single ring at the base of the female

zone (rarely among the pistils); sterile interstice present, supra-pistillate

pistillode zone often constricted; pistillodes half to twice the diameter of

180 Gard. Bull. Singapore 62 (1) 2010

ovary, close-packed or laxly arranged and followed by a whorl of staminodes,

this zone often constricted; male zone contiguous with interstice staminodes,

cylindric; stamens close-packed, the whole butterfly or dumbbell-shaped

from above; pores oblong to C-shaped, deep to shallow; appendix sub-

cylindric, white or yellow when fresh, distally tapering to a blunt or sharp

point, basally merging with the male zone or weakly or distinctly wider and

slightly truncate; appendix staminodes small to rather large, densely packed,

flat-topped to slightly impressed, irregularly polygonal.

Key to Schismatoglottis nervosa species complex

1. Petiole longitudinally ridged; vegetative tissues pungently aromatic

(terpenoids) when crushed; leaves + elliptic, thickly to thinly coriaceous

with vems prominent abaxialll yi ro scscee ss ccsc ae ee eee 2

1. Not the above combination ......... remainder of genus (see Hay and

Yuzammi, 2000; Hay, 2002)

2. Spathe hardly opening; upper part of spathe limb remaining green during

anthesis; stigma green whemftesins ce:cccse eters eee 5

2. Spathe opening wide; upper part of spathe turns to white during anthesis;

stigma white when fresh (except for S. tessellata)) i. secccccess-s-0ee stress -

3. Petiolar sheath persistent, opening wide; lamina longer and narrower

(ratio >3:1); primary venation adaxially flush with surface and with less

than 10-15 on each side. Central Sarawak: Sarikei.......8. ulusarikeiensis

3. Petiolar sheath marcescent, opening narrow; lamina shorter and wider

(ratio < 3:1); primary venation adaxially sunken with surface and with 15-

20 on each side. West Malaysia: widespread.................::00+ S. brevicuspis

4. Appendix white when fresh; petioles puberulent, densely (sometimes

minutely) verruculate, longitudinal ridges prominent; leaf apex shortly

acuminate to Ca’ 1 CIM... <2. ciecccecsectencecepeteenst a eer ee 5

4. Appendix yellow when fresh; petioles glabrous, not verruculate,

longitudinal ridges not prominent; leaf apex strongly acuminate to ca 3

5. Tertiary venation obscure-tessellate; female zone exceeding male

zone, pistils laxly arranged, stigma small, 1/5 of ovary diam., turning

yellow in alcohol; spadix interstice sharply constricted. West Sarawak:

Matai gi siié cc. scssesenseenseosnzenc ene ctess tenet es tetas cestee aes Sess S. matangensis

5. Tertiary venation prominent tessellate; male zone exceeding female zone,

pistils crowded, stigma large, 1/3 to % of ovary diam., remaining white

in alcohol; spadix interstice weakly constricted. Plants of limestone,

sandstone and granite. West Sarawak except Matang ............c.ceeeeeeees 6

Studies on Schismatoglottideae 18]

10.

Male zone irregularly arranged with uneven surface, male flowers round:

appendix markedly wider than male zone and slightly truncate basally

fete MC FET el ATITS OF SATISTOMEs:.-......s005.cccsa+escdescsasscaceecstcstoesenss S. turbata

Male zone evenly arranged with smooth surface, male flowers butterfly

shape or dumbbell shape; appendix contiguous with male zone. Plants on

2) VETTE OLE SUDO IS ICIS Aa a

Spathe limb splitting into longitudinal strips before deliquescing. Plants

CL SETILG SIDING soso ee ee S. linae

Spathe limb deliquescent in a whole piece. Plants of limestone.............. 8

Stamen pores narrowly C-shaped, deep, connective narrow, rims not

alate; stigma 2 ovary diam.; secondary and tertiary venation not strongly

pronounced. Plants weak to robust, sometimes colony forming. Always

lithophytic in soil collected in crevices and sinkholes in limestone. Full

shade, less than 200 m asl. West Sarawak: Bau........... eee S. nervosa

Stamen pores oblong, shallow, connective broad, rims widely alate; stigma

13 ovary diam.; secondary and tertiary venation pronounced. Plant robust,

always colony forming. Always terrestrial in deep soil on limestones and

sandstones. Sometimes in full sun, less than 600 m asl. West Sarawak:

PVE AC RIR SYST 210 ease so hoe ee S. simonii

Leaf lamina thinly coriaceous, occasionally with grey stripe, adaxially

glossy; tertiary venation more tessellate adaxially than abaxially;

female flowers green; appendix !/ of spadix length. Central Sarawak:

NGM 222 e doc vont ccc seu sede actbosssvesiSieesonesdcotisnnetenss S. tessellata

Leaf lamina coriaceous, never variegated, adaxially semi-matte; tertiary

venation more tessellate abaxially than adaxially; female flowers white;

ASM ie VARO a Ol SPACES LOMO UN cc cescstseinsdevesstwaceceseesavsosesestacarseecsonentsad 10

Petiolar sheath leathery; leaf lamina glaucous abaxially, leaf apex shorter

(to ca 2 cm); stigma large (2 of ovary diameter), staining yellow in alcohol;

connective narrow, pore oblong. Plants of shales in central and northeast

SS ll Ad Kept ECS hc Seance de seeen os Eases cose soneoteaceatenseeeshs S. adoceta

10. Petiolar sheath coriaceous; leaf lamina not glaucous abaxially, leaf apex

longer (to ca 3 cm); stigma small (1/5 of ovary diameter), remaining white

in alcohol; connective broad, pore C-shaped. Plants of limestones in

STINE SE SRE ease Rie eRe ee S. elegans

1. Schismatoglottis adoceta S.Y.Wong, sp. nov.

Ab Schismatoglottis elegans foliis subtus glaucis, laminae foliae apice

acuminata breviore (ad 2 cm longa), stigmatibus maioribus (quam ovaria

ca 2 diametro vs. 1/5 in S. elegans) et cum alcoholis flavescente (stigmatil S.

elegansii sum alcoholis alba remenans). Connectivo antherae anguste, poris

182 Gard. Bull. Singapore 62 (1) 2010

oblongo (vs. connectivo antherae crasso, poris hippocrepiformis: S. elegans)

distinguitur. Schismatoglottis adocetae in habitu saxa shaleiaca restrictus est

(S.elegans in habitu calcicola unica).—Typus: Malaysia, Sarawak, Kapit Division,

Belaga, km 10 Bakun-Bintulu-Miri road junction, 02° 50° 51.7”; 114° OV 57.6”,

182 m asl, 11 Oct 2005, PC Boyce et al. AR-1408 (holo, SAR). Plate 1.

Leaf petioles glabrous, slightly channelled and carinate adaxially in cross

section; sheathing in the lower \s to 4%, sheath fully attached, closed, tapering,

leathery, persistent; lamina broadly ovate to oblongo-ovate, more-or-less

elliptic, thinly coriaceous, slightly bullate, 7-9 cm wide x 13-23 cm long, base

broadly rounded to sub-truncate, slightly retuse or cuneate, apex acute to

strongly acuminate to ca 2 cm, adaxial surface semi-matte mid-green, abaxial

surface paler green, glaucous, drying strongly discolorous, midrib adaxially

flush with lamina, abaxially prominent, raised canaliculate in cross section;

primary venation impressed adaxially, strongly raised abaxially with 17-22

primary lateral veins on each side, alternating with lesser interprimaries,

frequently with a few branches from near the base, both diverging at 45°- 60°

and gradually curving towards the apex before reaching the intermarginal

collecting vein; secondary venation, arising from the midrib and the

primary veins; tertiary venation obscure tessellate adaxially and abaxially

in both living plants and dry specimens. Inflorescences up to three in quick

succession alternating with foliage leaves ca 5 cm long, (1.e. not forming a true

synflorescence), fragrant (esterase) at female anthesis; peduncle to ca 1 cm,

not exserted from leaf bases, slightly exserted in fruit; spathe interior glossy,

exterior semi-glossy, coriaceous, ca 7.5 cm long, lower spathe narrowly ovoid,

green when fresh, ca 2.5 cm long x | cm diam., differentiated from the limb

by a weak constriction; limb sub-truncate, white, ca 5 cm long, coriaceous,

opening wide except for convolute base, at first over-arching, then reflexed,

finally more or less twisted and then caducous-crumbling, apex mucronate

ca 4mm long, green when fresh; spadix sessile, adnate isodiametrically to the

spathe in the lower !/2 of female zone, whitish, to ca 5.5 cm long, less than the

length of spathe; female zone ca 1.7 cm long, 13 of spadix length; pistils many,

crowded, round to slightly rhomboid, with weakly bisulcate longitudinal

ridges, ca 0.5 mm diam. x 0.45 mm long; stigma large, bun-shaped, to ca 0.3

mm diam., ! of ovary, punctiform, papillate, yellow in alcohol; interpistillar

staminodes very occasional among the pistils, crowded into a row along

the spathe/spadix adnation, stalked, clavate and flat-topped, ca 0.5 mm

diam., similar size to pistils; sterile interstice strongly narrowed, ca 1.4 mm

diameter in the middle, (2.5-) ca 7 mm long which is 'gof spadix length,

basally more or less covered with inflated abortive pistils (reduced stigmas),

twice the diameter of ovary, transversely sulcate, apically covered with two

rows of sterile stamens, irregular shape, slightly wider than stamens; male

Studies on Schismatoglottideae 183

Plate 1. Schismatoglottis adoceta S.Y.Wong. A. Whole plant: B. Leaf lamina adaxially: C. Leaf

lamina abaxially; D. Female zone; E. Male zone and yellow appendix; F. Inflorescence during

female anthesis; G. Inflorescence during male anthesis; H. Spadix with the spathe artificially

removed; I. Spathe at male anthesis; J. Spathe prior to female anthesis. Images © P.C. Boyce

184 Gard. Bull. Singapore 62 (1) 2010

zone cylindric, ca 3.6 mm diam. x 1.2 cm long, 13 of spadix length, marked

increased in diameter from interstice; stamens crowded, truncate, ca 0.4 mm

across, dumbbell-shaped with the connective not elevated above the thecae,

connective narrow, pore oblong and deep; appendix sub-cylindric, ca 1.8

cm long, 1/3 of spadix length, yellow when fresh, turning white in alcohol,

frequently chewed by beetles, tapering to a sharp point, basally more or less

not isodiametric with top of male zone; staminodes of appendix more or less

columnar, irregularly polygonal, flat-topped, ca 0.5-1 mm diameter.

Other specimens seen: SARAWAK. Bintulu Division: Bintulu, Bk. Satiam,

02° 59° 33.0”; 112° 56’ 01.4”, 18 m asl, 12 Aug 2004, PC. Boyce & Jeland ak

Kisai AR-632.1 (SAR); Bk. Merairi, 02° 46’ 26.9”; 113° 39° 19.8”, JS/LC-24

AR-1284 (SAR).

Distribution: Central and northeast Sarawak, Kapit and Bintulu Divisions.

Habitat: Lithophytic on shale (Bintulu and Kapit Divisions), 18-180 m asl.

Notes: Schismatoglottis adoceta is clearly distinguished from the rest of the

S. nervosa complex by having leathery petiolar sheath (coriaceous petiolar

sheath in the rest of the complex). S. adoceta is most similar to S. elegans

although readily distinguished by the leaf laminae abaxially glaucous, the

leaf apex much shorter-acuminate (to ca 2 cm), larger stigma (S. adoceta

12 of ovary diameter vs. S. elegans 1/5 of ovary diameter) staining yellow in

alcohol (remaining white in S. elegans), anthers with a narrow connective

and oblong pore (S. adoceta) whereas anthers in S. elegans have a broad

connective and a C-shaped pore. Schismatoglottis adoceta is restricted to

shales as compared to S. elegans endemic on limestones. Schismatoglottis

adoceta differs from S. tessellata (also of shales) by coriaceous leaf laminae

with the adaxially much less pronounced tertiary venation and the spadix

appendix in S. adoceta much longer and narrower.

Etymology: The specific epithet is derived from the Greek ‘adocetus’,

unexpected, in allusion to the realization late in the preparation of this

manuscript that several elements of S. elegans sensu Hay and Yuzammi

required a specific recognition.

2. Schismatoglottis brevicuspis Hook.f.

FI. Brit. Ind. 6 (1894) 537; Ridl. Materials Fl. Mal. Pen. 3 (1907) 33 & J. Straits

Branch Roy. Asiat. Soc. 57 (1910) 113 & Fl. Mal. Pen. 5 (1925) 113; Engl. &

Krause, Pflanzenr. 55 (I[V.23Da) (1912) 98, pro parte excl. specim. cit. Curtis

(Penang, Waterfall), Ridley (Selangor, Petaling; Pahang, Tahan River) [i.e. S.

Studies on Schismatoglottideae 185

brevipes Hook.f., q.v.]. —Type: Malaysia, Perak, Scortechini 612 (holo, K; iso,

CAL, SING).

Other specimens seen: PENINSULAR MALAYSIA. Johor: Muar, Gunung

Ledang F.R., Gunung Ledang, (Mt Ophir), 22 Jan 1994, (orig. coll. Hay, A.,

Samy, A. & Ban Ka 9172) sub. C. Herscovitch NSW407380 (KEP 41490, L

0832670); Bukit Tunjok Laut, Ngadiman 37088 (SING). Melaka: Base of G.

Mering, Ridley s.n. (SING). Pahang: Pulau Tioman, Jason Bay, Burkill 1042 (K,

SING); Kemaman, Ulu Ayam, Bukit Kajang, Corner 30249e (SING); Raub-

Bentong boundary, Furtado 33097a (SING). Negeri Sembilan: Beremban

Forest Reserve, foot of Gunung Angsi, Furtado s.n. (SING). Selangor: Ulu

Gombak, Croat 53276 (K, MO); Gombak valley, 13 Jan 1994, C. Herscovitch

NSW407381 (KEP 41493); near Klang Gates Reservoir, Nicolson 1140 (K):

Genting Peras, Ridley s.n. (SING). Perak: Hulu Perak, Tasik Banding, n.d.,

Baharuddin bin Sulaiman s.n. (sub. AR-2599, living collection in Semenggoh

Botanical Research, Kuching, Sarawak) (SAR, USM); Sungei Batang Padang,

Tapah, Furtado 33096 (SING); Larut, 14 Jan 1994, Hay, A., Samy, A. & Ban

Ka 9075 (L 0832668); Kuala Kangsar, Bubu F.R., foothills of Gunong Bubu,

18 Jan 1994, (orig. coll. Hay, A., Samy, A. & Ban Ka 9130; Cult. RBG Sydney,

Acc. No. 940126 (KEP 41475, L 0832671). Kelantan: Gua Musang, Relai FR.

5°02’ N, 102°23’ E, 1992, Kiew, B.H. KBH10 (KEP 38211). Kedah: Gunung

Bongsu, nr Terap, Bogner 1692 (K). INDONESIA: Sumatera: Sibolangit,

Alston 14481 (BM); Sibolangit, Bukit Semiak, Md Nur 7368 (K); Aceh,

Middle Alas River (Lae Sauraya) area, ca 15 km N of Gelombang, S of

Bengkong R., 21 Jul 1985, de Wilde & de Wilde-Duyfjes 20188 (L 0239831).

Distribution: Malay Peninsula and Sumatra (North Sumatera and Aceh

provinces)

Habitat: In wet gullies and among rocks by streams in lowland rain forest

and lowland hill forest, to ca 900 m alt.

Notes: P.C. Boyce and the author saw a living collection of this species at

Nancy Botanical Garden, France, and realised that this species belongs to

the S. nervosa complex, although somewhat unusual in the spathe hardly

opening and the spathe limb green, a character set otherwise shared only

with S. uwlusarikiensis. A later trip to the aroid garden in Universiti Sains

Malaysia, Penang, which also has the living material of S. brevicuspis further

confirmed the placement of this species.

3. Schismatoglottis elegans A.Hay

Telopea 9(1) (2000): 67. Type: Cultivated in RBG Sydney Acc. No. 940510,

186 Gard. Bull. Singapore 62 (1) 2010

ex Malaysia, Sarawak, Niah National Park, Niah Caves area, along path

from Niah town (orig. coll. Hay et al. 9359), Feb 1996, C. Herscovitch s.n.

(holo, SAR; iso, K, KEP, NSW, all + spirit). Plate 2.

Plate 2. Schismatoglottis elegans A.Hay. A. Population in the wild, spreading across the floor,

the base of a limestone hill; B. Whole plant occurs in deep litter on limestone; C. Leaf lamina

apex (to c.3 cm long); D. Inflorescence at male anthesis; E. Creeping rhizome; F. Inflorescence

post anthesis showing the deliquescing spathe limb. Images © P.C. Boyce

Studies on Schismatoglottideae 187

Other specimens seen: MALAYSIA. Sarawak. Miri Division: Niah Suai, Niah

National Park, trail to Great Cave, 03° 49’ 09.9”: 113° 46’ 52.3”, 46 m asl, 13

Oct 2005, P.C. Boyce et al. AR-1428 (SAR); Niah Suai, Niah National Park.

Madu Trail, 03°48 57.9»; 113° 4618.3», 34 m asl, 13 Jul 2006, PC. Boyce et

al. AR-1877 (SAR + spirit); P.-C Boyce et al. AR-1878 (SAR + spirit): Subis,

Gua Sibau, Niah National Park, Trail from Rh. Chang, 24 Aug 2002, Julaihi

A. et al. S. 89319 (SAR).

Distribution: Sarawak. Miri Division, endemic to Niah area (Northeast

Sarawak).

Habitat: Humus-filled crevices in limestone in wet to swampy lowland

rainforest, 34-200 m asl.

Notes: Schismatoglottis elegans is distinguished from S. nervosa (in West

Sarawak limestone) by leaf laminae not glaucous abaxially and the spadix

appendix yellow when fresh. The tip of the leaf lamina acuminate up to ca

3 cm. Stamens of S. elegans are much rounder, with a broad connective and

are the more dumbbell-shaped as compared to other species in the complex.

Stamen pores are deepest as compared to other species in the complex. Hay

and Yuzammi (2000) mentioned that the appendix staminodes dried with

the stalk collapsed but the tops remaining expanded and tending to cohere

in groups. However, inflorescence of AR-/877 does not behave in this

manner while some inflorescences of S. nervosa behave in the same manner

suggesting that this is not a strong differentiating character. The spadix of

AR-1877 differs from other collections in that it has ovaries tetrasulcate with

the stigma staining yellow and interstice staminodes and stamens turning

darker yellow in alcohol. It is not clear if these are artefacts of the period

(post anthesis) of preservation.

Schismatoglottis elegans resembles S. tessellata although aside from

ecological differences (S. tessellata is restricted to shales) the latter also

differs by having thinly coriaceous leaf laminae with the tesselate tertiary

venation adaxially markedly more pronounced, a larger stigma and much

shorter and broader spadix appendix.

Hay and Yuzammi (2000) noted that S. elegans is found in a variety

of other localities in rainforest between 800 and 1300 m alt. However,

there are considerable doubts about the four specimens cited and pending

further investigation, the author suspect these observations represent

morphologically similar but different species.

4. Schismatoglottis linae S.Y.Wong, sp. nov.

Schismatoglottis linae simillima speciebus S. nervosa et S. simoni quae

spathe senescens differt. Spathae laminorum findens longitidinalis ante

188 Gard. Bull. Singapore 62 (1) 2010

deliquescens, staminis connectivo producto latus, poris antherae profundis (vs.

staminis connectivo producto latus, poris non-profundis: S. simonii, et staminis

connectivo anguste et poris profundis: S. nervosa) distinguitur. Inter specibis

Schismatoglottodorum grex nervosae habitu graniticola terrestri uniqus. —

Typus: Malaysia, Sarawak, Kuching Division, Lundu,Gunung Gading, below

plank walk to swimming area, 01° 41’ 31.0”; 109° 50° 44.5”, 700 m asl, 14 Dec

2006, P.C. Boyce et al. AR-2062 (holo, SAR). Plate 3.

Leaf petioles terete, obscurely longitudinally ridged (clearer under

magnification), glabrous, up to ca 40 cm long, slightly channelled and

carinate adaxially in cross section; sheathing in the lower 1/3 to %, sheath fully

attached, closed, tapering, coriouceous, marcescent; lamina broadly ovate to

oblongo-ovate, more-or-less elliptic, thinly coriaceous, slightly bullate, 8-10

cm wide x 15-25 cm long, base broadly rounded to sub-truncate, slightly

retuse or cuneate, apex acute to strongly acuminate to ca 3 cm, adaxial

surface semi-matte mid-green, abaxial surface paler green, glaucous, drying

strongly discolorous, midrib adaxially flush with lamina, abaxially prominent,

raised canaliculate in cross section; primary venation impressed adaxially,

strongly raised abaxially with 18-22 primary lateral veins on each side,

alternating with lesser interprimaries, frequently with a few branches from

near the base, both diverging at 45°- 60° and gradually curving towards the

apex before reaching the intermarginal collecting vein; secondary venation,

arising from the midrib and the primary veins; tertiary venation obscure

tessellate adaxially and abaxially in both living plants and dry specimens.

Inflorescences up to three in quick succession alternating with foliage leaves

ca 5 cm long, (i.e. not forming a true synflorescence), fragrant (esterases) at

female anthesis; peduncle to ca 1 cm, not exserted from leaf bases, slightly

exserted in fruit; spathe interior glossy, exterior semi-glossy, coriaceous, ca

13 cm long, lower spathe narrowly ovoid, green when fresh, ca 3.5 cm long

x 1 cm diam., differentiated from the limb by a constriction; limb truncate,

white, ca 8 cm long, coriaceous, opening wide except for convolute base,

at first over-arching, then reflexed, finally more or less twisted and then

caducous-crumbling in longitudinal stripes, delisquescent post male anthesis,

apex mucronate ca 4 mm long; spadix sessile, adnate isodiametrically to the

spathe in the lower 2 of female zone, whitish, to ca 10 cm long, less than the

length of spathe; female zone ca 2.5 cm long, 1/3 of spadix length; pistils many,

crowded, round to slightly rhomboid, with weakly bisulcate longitudinal

ridges, ca 0.5 mm diam. x 0.5 mm long; stigma smaller than ovary, bun-

shaped, to ca 0.3 mm diam., punctiform, papillate; interpistillar staminodes

confined into a row along the spathe/spadix adnation, stalked, clavate and

flat-topped, ca 0.5 mm diam., similar size to pistils; sterile interstice strongly

narrowed, ca 0.4 cm diameter in the middle, ca 5 mm long, basally more

Studies on Schismatoglottideae 189

Plate 3. Schismatoglottis linae S.Y.Wong. A. Whole plant; B. Inflorescence at male anthesis

with an infructescence on the right side; C. Leaf lamina abaxially; D. Emerging inflorescence

with two infructescences; E. Inflorescence post anthesis showing the spathe limbs caducous

in longitudinal stripes. Images © P.C. Boyce

190 Gard. Bull. Singapore 62 (1) 2010

or less covered with inflated abortive pistils (reduced stigmas), twice the

diameter of ovary, transversely sulcate, apically covered with sterile stamens,

irregular shape, slightly wider than stamens; male zone cylindric, ca 5 mm

diam. x 3.5 cm long, '4 of spadix length, slightly increased in diameter from

interstice; stamens crowded, truncate, ca 0.4 mm across, dumbbell-shaped

with the connective not elevated above the thecae, connective broad, pore

oblong and deep; appendix subcylindric, ca 3.5 cm long, '4 of spadix length,

white when fresh and in alcohol, tapering to a sharp point, basally more or

less not isodiametric with top of male zone; staminodes of appendix more or

less columnar, irregularly polygonal, flat-topped, ca 0.5-1 mm diameter.

Distribution: Sarawak, Kuching Division, endemic to Gunung Gading,

Lundu (West Sarawak).

Habitat: Terrestrial on pockets of loose soil in deep litter along the stream

running through at the base of the granites (Kuching Division), 70 m asl.

Notes: Schismatoglottis linae resembles S. nervosa and S. simonii but is

distinguished from these two species by the spathe limb senescence.

The spathe limb splits into longitudinal stripes before deliquescing. The

connective is broad but stamen pores are deep (broad connective and

shallow pore in S. simonii, narrow connective and deep pore in S. nervosa).

Plants are terrestrial along the stream in deep litter on granite surface (the

only species in the complex on this geology).

Etymology: This species is named for Lin Jenkins who is an avid lover of

tropical aroids.

5. Schismatoglottis matangensis S.Y.Wong, sp. nov.

Schismatoglottis matangensis cum S. nervosa confunditur sed inflorescentia

femina quam inforescentia mascula excedens, pistillis sublaxis ordinatis,

stigmate parva (quam ovaria ca 's diametro) cum alcoholis flavescente,

spadice cum interstitio sterile abrupte constricta et habitu lapis arenaricola

restrictus differt. -Typus: Malaysia, Sarawak, Kuching Division, Matang,

Kubah National Park, Waterfall Trail, 01°35’ 40.2”; 110° 10°45.9”, 190 m asl,

7 Jun 2006, P.-C. Boyce & S.Y.Wong AR-1830 (holo, SAR). Plate 4.

Leaf petioles terete, channelled canaliculate adaxially in cross section, 13-

20 cm long, strongly longitudinally ridged (like celery) especially abaxially,

very minutely (strong lens required) and densely verruculate, sheathing

in the lower '4 - % with the sheaths tapering, closed, and fully attached,

marcescent, sometimes with a short rounded ligular portion; lamina

Studies on Schismatoglottideae 19]

Plate 4. Schismatoglottis matangensis S.Y.Wong. A. Whole plant; B. Leaf lamina abaxially; C.

Inflorescence at male anthesis with an infructescence on the left side; D. Inflorescense at male

anthesis; E. Female zone, note the stigma in orange; F. Spathe limb deliquescent post male

anthesis; G. Male zone; H. Spadix with the spathe artificially removed. Images © PC. Boyce

192 Gard. Bull. Singapore 62 (1) 2010

broadly ovate to oblongo-ovate, not elliptic, coriaceous, 13-22 cm long x

7-11 cm wide, the base broadly rounded to slightly retuse, the apex acute

and shortly acuminate to ca | cm, undulate along margin, adaxial surface

matte deep green, always slightly bullate, abaxial surface glaucous, paler

green, drying strongly discolorous; midrib adaxially flush with lamina but

centrally channelled toward leaf base, abaxially very prominent, raised

canaliculate in cross section; primary venation impressed adaxially, strongly

raised abaxially, numerous, 15-22 on each side of the midrib, alternating

with lesser interprimaries or these occasionally arising from the bases of

the primary veins, both diverging at 45°-90° and gradually curving towards

the apex before reaching the intermarginal collecting vein; secondary

veins mostly arising from the midrib, occasionally from near the bases of

the primary veins; tertiary venation forming faint tessellate adaxially and

abaxially. Inflorescences up to 3 together, erect, white, fragrant (esterases)

smell at female anthesis, peduncle to ca 1.5 cm long, concealed by cataphylls

deliquesing and adhering to spathe limb exterior, slightly exserted in fruit;

spathe, interior glossy, exterior, semi-glossy, softly coriaceous, ca 9.5 cm long,

lower spathe very pale green, ca 4 cm long x 1.3 cm width, differentiated

from the limb by a faint constriction at interstice, narrowly ovoid; limb

obovate, white, ca 5.5 cm long x 2.2 cm width, upper margin reflex, abruptly

acuminate, ca 7 mm long and ultimately mucronate apex, ca 2 mm long,

caducous by crumbling at or just after male anthesis; spadix adnate,

isodiametrically attached to 4 of female zone, subcylindric, 5.2 cm long, %

of spadix length; female zone, conic-cylindric, ivory in fresh and alcohol, 1.6

cm long x 7 mm diam., '4 of spadix length; pistils many but laxly arranged,

irregularly round to slight rhomboid from above, ca 0.46 mm diam. x 0.72

mm long, stigma, sessile, minute, ca 0.15 mm diam., '5 of ovary, punctiform,

papillate, orange staining in alcohol, ovaries of on lower part zone tending

to be bisulcate, ovaries of on upper part zone tending to be trisulcate or

tetrasulcate; interpistillar staminodes confined to less than a single ring at

the base of the female zone, similar or twice the size of female flower, flat-

topped: sterile insterstice presents, '/s of spadix length, supra pistilate pistilode

zone (ca 2.5 mm long), pistilode larger than pistil, irregularly round, 0.6 mm

diam. transitioning into naked zone (1.5 mm long x 2.3 mm diam.) in AR-

1830 or contiguous with infra staminate staminodes zone (ca 2.5 mm long),

irregularly polygonal, flat-topped, white when fresh but stained orange

in alcohol (AR-1830 & AR-1865), strongly narrow at the staminode zone

corresponding with the spathe constriction; male zone cylindric, 10 mm long

x 4mm diam., 4 of spadix length; stamens, butterfly-shaped from above,

white when fresh, turning dirty yellow in alcohol (AR-1830 & AR-1865),

remaining white in alcohol (A R-/864), close-packed, 0.3 mm diam., anthers

truncate (flat-topped), connective narrow and thecae large, pores small and

Studies on Schismatoglottideae 193

deep, C-shaped with the convex side innermost; appendix sub-cylindric, ca

1.9 cm long, *4 of spadix length, white when fresh and in alcohol, strongly

tapering to a sharp end, middle slightly thicker than the base: staminodes

of appendix densely packed, flat-topped, irregularly polygonal, ca 0.3-1 mm

diam.

Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Matang,

trail to Indian Temple, 2 Mar 2004, PC. Boyce & Jeland ak Kisai AR-145.3

(SAR); Matang, Kubah National Park, Waterfall Trail, 01°35’ 40.2”; 110°

10°45.9”, 190 m asl, 7 Jun 2006, PC. Boyce & S.Y.Wong AR-1831 (SAR); ibid.,

P.C.Boyce & S.Y.Wong AR-1832 (SAR); ibid., PC. Boyce & S.Y.Wong AR-

1833 (SAR); ibid., PC. Boyce & S.Y¥.Wong AR-1834 (SAR); ibid., 11 Jul 2006,

P.C. Boyce & S.¥.Wong AR-1864 (SAR + spirit); ibid., PC. Boyce & S.Y.Wong

AR-1865 (SAR + spirit); ibid., PC. Boyce & S.Y.Wong, AR-1866 (SAR).

Distribution: Sarawak, Kuching Division, endemic to Matang (West

Sarawak).

Habitat: Evergreen moist forest on Matang series sandstones where terrestrial

in deep litter over exposed black soil, sometimes between sandstone rocks,

190-450 m asl.

Notes: Schismatoglottis matangensis is morphologically different from S.

nervosa by characters of the inflorescence: female zone exceeding male

zone, pistils laxly arranged [in marked contrast to most other species of the

S. nervosa complex (except for S. tessellata) in which pistils are crowded]

and minute (‘4 of ovary diam.) two-three lobed stigma staining yellow

in alcohol. Other distinguishing characters for S. matangensis include a

sharp constriction at the interstice, interstice staminodes staining orange

in alcohol (AR-1830 & AR-1865) and stamens staining orange in alcohol

(AR-1865) although the remainder of the observed inflorescences have

stamens remaining white in alcohol and these colour differences may be an

artefact of inflorescence anthesis phase at preservation although stamens of

S. nervosa always remain white in alcohol irrespective of the phase of the

spadix. In robust inflorescences, the appendix displays a marked increase in

diameter at the junction of the male zone, contiguous with the male but not

isodiametrical.

The petiolar sheath is marcescent in S. matangensis. Primary venation

of S. matangensis is more prominent than in S. nervosa, however, on average

secondary and tertiary venation are less pronounced. Schismatoglottis

matangensis is restricted to sandstones whereas S. nervosa are restricted to

Karst limestones.

194 Gard. Bull. Singapore 62 (1) 2010

Etymology: The specific epithet from the Matang Massif, the type locality

and so far the only known site for this species.

6. Schismatoglottis nervosa Ridl.

Schismatoglottis nervosa Ridl., Journ. As. Soc. Straits 49 (1907) 50; Hay

& Yuzammi, Telopea 9(1) (2000) 73. —Typus: Cult. in Singapore Botanic

Gardens, ex Malaysia, Sarawak, Bau, Jan 1907, H. N. Ridley s.n. (holo, SING,

2 sheets). Plate 5.

Other specimens seen: MALAYSIA. Sarawak. Kuching Division: Bau, Bk.

Krian, 28 May 1972, J.A.R.Anderson S. 31966 (SAR); Bau, Krokong, Gua

Peri-peri (Fairy Cave),01° 22° 51.9”; 110° 07? 09:37, 30 measl, 29 Oct 2003;

PC.Boyce & Jeland ak Kisai AR-145.1 (SAR); Bau, Krokong, Kampung

Tringgus, Sg. Bong, 01° 157 32.272 110° 05/°37-2-2. 8 aisasle 2 ule O04:

PC.Boyce & Jeland ak Kisai AR-525 (SAR); Bau, Gn. Bidi, 01° 23’ 27.0”;

110° 07° 07.6”, 50 m asl, 6 Jan 2005, P.C.Boyce & Jeland ak Kisai AR-944

(SAR + spirit); Bau, Kampung Bogag, Gn. Tibugai, 01° 21° 31.1”; 110° 03°

48.7”, 80 m asl, 6 Jan 2005, PC. Boyce & Jeland ak Kisai AR-950 (SAR); Bau,

Krokong, Gua Peri-Peri (Fairy Cave), 01° 22’ 51.9”; 110° 07’ 09.3”, 30 m asl,

25 May 2006, P.C.Boyce & S.Y.Wong AR-1823 (SAR); ibid., P-C.Boyce &

S.Y¥. Wong AR-1824 (SAR); Ibid., RC. Boyce & S.Y.Wong AR-1825 (SAR);

ibid., PC.Boyce & S.Y.Wong AR-1826 (SAR + spirit); Ibid., PC. Boyce &

S.Y¥.Wong AR-1827 (SAR + spirit); Bau, Kampung Jugan, 01° 28° 46.4”; 110°

0S’ 08.5”, 72 m asl, 25 Jun 2004, P.-C. Boyce et al. AR-491 (SAR); Bau, Gn.

Singai, 17 Nov 2004, PC. Boyce etal. AR-752 (SAR); Bau, Krokong, Kampung

Tringgus, 01° 15’ 40.2”; 110° 05’ 35.9”, 80 m asl, 19 Feb 2005, PC Boyce et al.

AR-992 (SAR); Bau, Gn. Juita, 01° 23’ 48.7”; 110° 08’ 07.2”, 35 m asl, 28

Oct 2005, P.-C. Boyce et al. AR-1499 (SAR); Bau, Segong, Gn. Opar, 01° 27°

07.3”; 110° 04’ 00.5”, 79 m asl, 9 Nov 2005, P.-C. Boyce et al. AR-1502 (SAR);

Bau, Krokong, Kampung Tringgus, 01° 15’ 40.2”; 110° 05° 35.9”, 81 m asl, 27

Jun 2006, P.C.Boyce et al. AR-1845 (SAR + spirit); Bau, Gn. Lanyang, 10

Apr 2002, Connie, G et al. SBC 2807 (SAR); Bau, Kampung Jugan, 19 Jun

2004, Jeland ak Kisai & Jipom ak Tisai AR-474 (SAR); Bau, Gn. Poing, 23

Sep 2001, Julia, S. et al. SBC 345 (SAR); locality not recorded, 26 Feb 2002,

Julia, S. et al. SBC 2155 (SAR); Bau, Bk. Jebong, 6 Jul 1970, RE Lehmann

S. 30136 (SAR); Bau, Jambusan, Gn. Batu, 19 Feb 2002, K. Meekiong et al.

SBC 1662 (SAR); Bau, Jambusan, Gn. Batu, 19 Feb 2002, K. Meekiong et al.

SBC 1929 (SAR); Bau, Jambusan, Gn. Jebong, 5 Mar 2002, K. Meekiong et

al. SBC 1953 (SAR); Bau, Bengoh range, Pangkalan Tebang, logging road,

6 Jul 1996, M. Mohizah S 73890 (SAR); 2 miles east of Bau, 6 Aug 1961, D.

H. Nicolson (SAR, US); Bau, Krokong, Gua Peri-Peri (Fairy Cave), 22 Mar

1999, Patsipun et al. S. 79985 (SAR); Bau, Gn. Tabai, 13 Mar 2002, Shaevy W.

et al. SBC 2298 (SAR).

Studies on Schismatoglottideae 195

Plate 5. Schismatoglottis nervosa Ridl. A. Whole plant; B. Inflorescence prior to anthesis:

C. Inflorescence at male anthesis: D. Whole plant with an inflorescence at male anthesis: E.

Spadix with the spathe artificially removed; F. Female zone: G. Spathe limb deliquescent post

anthesis. Images © P.C. Boyce

196 Gard. Bull. Singapore 62 (1) 2010

Distribution: Sarawak, Kuching Division, endemic to the Bau area (West

Sarawak).

Habitat: Evergreen moist forest on karst limestone in the Bau area. Terrestrial

in deep litter or lithophytic in soil collected in crevices and sinkholes in

limestone; rarely along stream banks, 30-160 m asl.

Notes: Hay and Yuzammi (2000) were unclear whether the interstice

was present or the fertile zones were contiguous. Based on the author’s

observation, the interstice is always present with pistillodes at the lower

part and staminodes at the upper part and without the presence of naked

zone. In the type specimen, the presence of the interstice is not clear due to

damage to the spadix at the crucial area of the interstice. In all collections

the female zone is shorter than the male zone, whereas S. matangensis has

a longer female zone than male zone. Stigmas of S. nervosa are larger than

those of S. matangensis and always remain white in alcohol.

In the type specimen, there are two types of staminodes on the

appendix of the inflorescence. The lower staminodes are very irregularly

rhomboid, smaller (ca 0.5 mm diam.) while the upper staminodes are

regularly rhomboid and larger (0.5-1 mm diam.). A naked zone, which has

dried black, is present at the tip of appendix.

7. Schismatoglottis simonii S.Y.Wong, sp. nov.

A Schismatoglottis nervosa habitu robustis coloniis, veneris laminorum

seconadariis et tertiariis plus prominentis, stigmate plus late, connectivo

antherae crasso, poris oblongo, non profundus quam marginem late alatis

differt. -Typus: Malaysia, Sarawak, Samarahan, Serian, Kidadum, Sugun

Karang, 01°06’ 17.6”; 110° 29°04.5”, 100 m asl, 29 Jun 2006, P.C Boyce et al.

AR-1859 (holo, SAR). Plate 6.

Leaf petioles terete, channelled slightly canaliculate adaxially in cross

section, 13-40 cm long, strongly longitudinally ridged (like celery) especially

abaxially at the base, very minutely (strong lens required) and densely

verruculate, sheathing in the lower '4- % with the wings tapering, and fully

attached, marcescent,sometimes with a short rounded ligular portion; lamina

broadly ovate to oblongo-ovate, sometimes elliptic, 12-30 cm long x 6-20

cm wide, coriaceous, adaxial surface semi-glossy deep green, always slightly

bullate, abaxial surface paler green, glaucous, drying strongly discolourous,

the base broadly rounded to sub-truncate, slightly retuse to cuneate, the

apex acute and shortly acuminate to ca | cm, undulate along margin; midrib

= 4 : a

Studies on Schismatoglottideae 197

Plate 6. Schismatoglottis simonii S.Y.Wong. A. Whole plant; B. Inflorescence during male

anthesis; C. Leaf lamina adaxially; D. Leaf lamina abaxially: E. Emerging inflorescence with

three infructescense flanking on each side; F. Spadix with spathe artificially removed: G.

Female zone; H. Spathe limb starting to deliquescent post male anthesis; I. Infructescence

ripen to reveal the seeds.

198 Gard. Bull. Singapore 62 (1) 2010

ip

adaxially flush with the lamina but centrally channelled toward the leaf base,

abaxially very prominent raised canaliculate; primary venation impressed

adaxially, strongly raised abaxially, numerous, 16-25 on each side of the

midrib, alternating with lesser interprimaries or these occasionally arising

from the bases of the primary veins, diverging at 60°-90° and gradually

curving towards the tip before reaching the intermarginal colllecting vein;

secondary veins mostly arising from the midrib, occasionally from near the

bases of the primary veins; tertiary venation forming tessellate reticulum

adaxially and abaxially, more notable abaxially. Inflorescences up to

three together, erect, white, fragrant (esterases) smell at female anthesis;

subtended by lanceolate cataphylls to ca 6 cm long; peduncle to ca 2.5 cm

long, concealed by leaf bases and cataphylls at flowering, slightly exserted in

fruit; spathe, interior glossy, exterior semi-glossy, softly coriaceous, ca 9 cm

long; lower spathe, ovoid, pale green when fresh and white in alcohol, ca 2.5

cm long x 1.5 cm diam., differentiated from the limb by a faint constriction

coinciding with interstice; limb oblongo-lanceolate, white when fresh and

in alcohol, ca 5.5 cm long, apex, green when fresh, mucronate for ca 5.8

mm, caducous by disintegrating or crumbling at or just after male anthesis;

spadix sessile, adnate isodiametrically to the spathe in the lower '/ of female

zone, subcylindric, ca 7 cm long subequalling the spathe; female zone ovoid,

ivory when fresh and in alcohol, 1.8 cm long x 0.8 cm diam., '4 of spadix

length; pistils numerous and close-packed, rhomboid to round from above,

ca 1.19 mm long x 0.56 mm diam.; stigma sessile, punctiform, large, 0.20 mm

diam., '4 of ovary, raised; interpistillar staminodes confined to more or less a

single ring at the base of the female zone, irregularly polygonal from above,

about the same size as the ovaries shorter than pistils; sterile insterstice

present, supra pistillate pistillodes zone, white, ca 1.8 mm long, '/4 of spadix

length, strongly narrow in the upper half of pistillodes zone corresponding

with the spathe constriction, pistillodes irregularly polygonal, large, ca 1 mm

diam., twice the size of ovary, centrally impressed; sometimes naked zone

present; followed by a whorl of incompletely abortive stamens, ca 2.6 mm

long, staminodes, irregularly squat-columnar, flat-topped, 0.62 mm diam.;

male zone contiguous with interstice staminodes, cylindric, 1.3 cm long x

4.2 mm diam., slightly less than '4 of spadix length; stamens close-packed,

irregular dumbbell-shaped from above and neighbouring anthers with their

lobes interdigitating, white when fresh and remaining white in alcohol, ca

0.6 mm diam., anthers truncate, connective broad and elevated, thecae large;

pores oblong and shallow, rims widely alate; appendix cylindric, 2.6 cm long

x 4.8 mm diam., slightly less than '4 of spadix length, white when fresh and in

alcohol, abruptly widen at base, distally tapering to sharp point; staminodes

of appendix densely packed, flat-topped to slightly centrally impressed,

irregularly polygonal, ca 0.5-1 mm diam.

Studies on Schismatoglottideae 199

Other specimens seen: MALAYSIA. Sarawak, Kuching Division: Padawan,

Kampung Bengoh, Danu road, Gn. Temuang, Sg. Abang, 01° 15’ 38.6”; 110°

15° 31.4”, 50 m asl, 16 Feb 2006, PC. Boyce et al. AR-1707 (SAR); Padawan,

10 mi s.w. of main Kuching-Serian Highway, 01° 10°; 110° 20°, 30 Sep 1981,

T: B.Croat 53179 (SAR); Siburan, Kampung Giam, Air Terjun Giam, 01° 19”

11.2”; 110° 16° 11.4”, 37 m asl, 7 Feb 2006, P.-C. Boyce et al. AR-1693 (SAR);

Samarahan Division: Serian, Mongkos, Kampung Batuh Mawang, Labak

Ebang, Utak Samat, 5 Jan 2006, Simon Kutuh ak Paru AR-1666 (SAR);

Serian, Gn. Ampungan, 01° 09’ 08.2”; 110° 37° 21.2”, 450 m asl, 21 Nov 2003,

P.C.Boyce & Jeland ak Kisai AR-92.4 (SAR); Serian, Pichin, Umon Murut,

Tiab Belanting, 01° 08’ 03.7”; 110° 27’ 00.3”, 90 m asl, 15 Jun 2005, P.C. Boyce

et al. AR-1215 (SAR); ibid., 15 Jun 2005, P.-C. Boyce et al. AR-1220 (SAR +

spirit); Serian, Mongkos, Kampung Batuh, Gn. Selabur, 00°57° 26.2”; 110°

30’ 15.8”, 100 m asl, 15 Mar 2006, PC. Boyce et al. AR-1724 (SAR); Serian,

Kidadum, Sugun Karang, 01°06’ 17.6”; 110° 29°04.5”, 100 m asl, 7 Apr 2006,

P.C.Boyee et al. AR-1764 (SAR); Serian, Gn. Ampungan, 01° 09’ 10.1”; 110°

37 26.2”, 568 m asl, 28 Aug 2006, P.-C. Boyce et al. AR-2003 (SAR + spirit);

Serian, Pichin, Gunung Kedadum, Sugun Kerang, 13 Nov 2004, Simon Kutuh

ak Paru AR-750 (SAR); Serian, Pichin, Labu, Sg. Tiyab, 26 Jul 2005, Simon

Kutuh ak Paru AR-1299 (SAR); Serian, Kampung Selabi, Sg. Mawang, 2

Feb 2006, Simon Kutuh ak Paru AR-1703 (SAR): Serian, Taman Rekreasi

Rachan, 01° 08’ 34.9”; 110° 35’ 02.4”, 57 m asl, 18 Oct 2006, PC. Boyce &

S.Y.Wong AR-2038 (SAR).

Distribution: Sarawak, Kuching & Samarahan Divisions, endemic to the

Padawan/Serian areas (West Sarawak).

Habitat: Always terrestrial in deep soil on limestones and sandstones,

sometimes lithophytic on permanent wet areas of limestone but not in

direct water flow, sometimes not in full shade. Large colony forming, 50-600

m asl.

Notes: Schismatoglottis simonii is clearly closely allied to S. nervosa but leat

laminae are wider in S. simonii than S. nervosa [{length: width ratio (to 3:2

in S. simonii, to 2:1 in S. nervosa)]. Secondary and tertiary venations in S.

simonii are more pronounced than S. nervosa. The zonation in the spadix

of S. simonii is equally distributed among the female, male and appendix

(‘seach) as compared to S. nervosa (female zone, 4; male zone *s; appendix

',). The stigma narrower in S. simonii as compared to S. nervosa while the

stamens have a broad connective, with shallow oblong pores. S. simonii tends

to form large colony and much more robust than S. nervosa.

200 Gard. Bull. Singapore 62 (1) 2010

Etymology: This species is named for Mr Simon Kutuh ak Paru, occasional

member of our field team, who has so ably organise field trips to Padawan

limestones.

8. Schismatoglottis tessellata S.Y.Wong, sp. nov.

Folit lamina tenuiter coriaceis, nervis tertiariis tessellatis abaxialiter et

adaxialiter valde prominentibus quamquam veniis adaxialiter valde

prominentibus, stigmate amplis globuliformis (quam ovaria ca 42 diametro);

appendice crasso brevioribus, usque ad tertiam partem longitudinis spadice

toto in complexus nervosae unica est. In habitu saxa shaleiaca restrictus. —

Typus: Malaysia, Sarawak, Kapit, Taman Rekreasi Sebabai, 01° 56° 45.6”;

112° 54 16.8”, 50 m asl, 16 Mar 2005, P.-C. Boyce et al. AR-1087 (holo, SAR).

Plate 7.

Leaf petioles, slightly D-shaped but slightly carinate channelled to the

base of lamina adaxially in cross section, glabrous, longitudinal ridges only

visible with magnification, long, 20-30 cm long; sheathing in the lower ‘4 to

¥Y2 with the sheaths tapering, fully attached, open, persistent to marcescent;

lamina broadly ovate to oblong ovate, thinly coriaceous, 13-21 cm long x 5-

10 cm wide, sometimes variegated with grey stripes on each side adaxially,

the base cuneate to slightly retuse, the apex acute and strongly acuminate

for ca 0.7-2 cm, adaxial surface semi-glossy deep green, abaxial surface

paler green, always slightly bullate, glaucous, drying strongly discolourous;

midrib adaxially flush with the lamina but centrally channelled at the leaf

base, abaxially very prominent, raised canaliculate in cross section; primary

venation impressed adaxially, strongly raised abaxially, numerous, 11-20

on each side of the midrib, alternating with lesser interprimaries or these

occasionally arising from the bases of the primary veins, diverging at 45°-

60° and gradually curving towards the tip before reaching the intermarginal

collecting vein; secondary veins mostly arising from the midrib, occasionally

from near the bases of the primary veins; tertiary venation forming distinctive

tessellate adaxially and abaxially, more notable adaxially. Inflorescences up

to three together, erect, white, fragrant (esterases) smell at female anthesis;

concealed by oblongo-lanceolate cataphyll,ca 7 cm long, peduncle to ca 2 cm,

slightly exserted in fruit; spathe, interior glossy, exterior semi-glossy, softly

coriaceous, ca 9 cm long; lower spathe broadly ovoid, ca 1.7 cm diam. x 3.8

cm long, differentiated from the limb by a faint constriction coinciding with

interstice; limb broadly lanceolate, rather coriaceous, caducous by crumbling

at or just after male anthesis; spadix, sessile, isodiametrically attached to the

spathe in the lower '4of female zone, conic-cylindric, ca 5.9 cm long; female

zone, conic-cylindric, white when fresh but very slightly yellow in alcohol, 1.4

Studies on Schismatoglottideae 201

Plate 7. Schismatoglottis tessellata S.Y.Wong. A. Whole plant on shales; B. Leaf lamina

adaxially revealing the tessellate venation; C. Leaf lamina variation; D. Inflorescence at male

anthesis; E. Spathe at male anthesis; F. Spadix with spathe artificially removed; G. Female

zone, interstice and part of male zone. Images © P.C. Boyce

202 Gard. Bull. Singapore 62 (1) 2010

cm long x 7 mm diam., 4% of spadix length; pistils numerous, laxly-arranged,

elongate-ovoid to subcylindric, ca 1.70 mm long x 0.53 mm diam.; stigma

sessile, large bun-shaped, ca 0.26 mm diam., 2 of ovary, raised, papillate;

interpistillar staminodes confined to more or less a single ring at the base of

the female zone, irregularly polygonal from above, twice the size of ovaries,

flat-topped, shorter than pistils; sterile interstice presence, 1.2 cm long, '/

of spadix length, pistils transitioning gradually into laxly spirally arranged

pistilode that are ca half to two times larger than pistil, irregularly round,

white when fresh and in alcohol, ca 7 mm long, sharp constriction in between

pistillode and staminode zones; staminodes, larger than stamens, irregular

polygonal, flat-topped, white when fresh and turning yellow in alcohol, ca

5 mm long; male zone, cylindric, ca 2.2 cm long x 4.8 mm diam., '4 of spadix

length; stamens close-packed, irregularly butterfly-shaped from above and

neighbouring anthers with their lobes interdigitating, 0.5 mm diam., anthers

truncate, connective broad and thecae large, pores large accounting of the

entire top of the thecae and shallow, C-shaped, the rims narrowly alate;

appendix conic-cylindric, 1.2 cm long x 5.3 mm diam., 4 of spadix length,

yellow when fresh, turning white in alcohol, distally tapering to a very blunt

point, basally slightly thicker than the male zone, middle thicker than basal;

staminodes of appendix loosely arranged, large staminode, flat-topped, 0.7-

1.2 mm across, squat-columnar.

Other specimens seen: MALAYSIA. Sarawak, Kapit Division: Nanga Gaat,

Rejang Wood Concession, Sg. Piat, 01° 38’ 09.1”; 113° 24’ 09.9”, 200 m asl,

14 Oct 2003, P.C.Boyce & Jeland ak Kisai AR-103.1; P.-C. Boyce & Jeland ak

Kisai AR 103.2 (SAR); ibid., 14 Oct 2003, P.C. Boyce & Jeland ak Kisai AR-

105 (SAR); Nanga Gaat, km 3.5 after heli-logging camp on road to Camp

Gahada, Sg. Bereng, 01° 45’ 36.0”; 113° 27’ 54.7”, 228 m asl, 15 Dec 2004,

P.C. Boyce et al. AR-888 (SAR); ibid., 19 Apr 2006, P.-C. Boyce et al. AR-1792

(SAR); ibid., 19 Apr 2006, P.-C. Boyce et al. AR-1794 (SAR).

Distribution: Central Sarawak (Kapit Division).

Habitat: Evergreen moist forest on shale where either on steep earth banks

or lithophytic on muddy shale, 50-228 m asl.

Notes: This species is distinguishable by its thinly coriaceous leaves texture

and tertiary venation strongly raised-tessellate in both surfaces of leaf,

but more prominently adaxially. The pistils are laxly arranged with large

(compared to ovary diameter) bun-shaped stigmas, while the appendix is

shorter and broader ( ' of spadix length) than other species of the S. nervosa

species complex. Schismatoglottis tessellata bears some resemblance to S.

Studies on Schismatoglottideae 203

elegans but is readily separable by its thinly coriaceous leaf lamina, adaxially

strongly prominent tessellate tertiary venation (hence the epithet) and

shorter leaf apex. The stigma is large bun-shaped in S. tessellata as compared

to small stigma in S. elegans. Schismatoglottis tessellata is restricted to shales,

as observed so far. Occasionally, leaf laminae of S. tessellata are variegated

with grey stripes on each side adaxially.

Etymology: The epithet reflects the prominently raised-tessellate venation

that immediately distinguishes this species.

9. Schismatoglottis turbata S.Y.Wong, sp. nov.

Schismatoglottis turbata ad alii specibis Schismatoglottodorum grex floribus

masculinus parvus, rotundis, densiter irregularis dispositae, poris profunde

differt. Appendice quam inflorescentiae masculae apicem sigillatim crassiore

distinguitur. —Typus: Malaysia, Sarawak, Kuching Division, Sempadi, Sg.

Limau, Bukit Kankar, 01°39" 44.2”; 109°59°56.5”, 41 m asl, 25 Aug 2007,

P.-C. Bovyee et al. AR-2143 (holo, SAR). Plate 8.

Leaf petioles terete, obscurely longitudinally ridged (clearer under

magnification), glabrous, up to ca 30 cm long, slightly channelled and

carinate adaxially in cross section; sheathing in the lower ‘4 to 4, sheath

fully attached, closed, tapering, leathery, persistent; lamina broadly ovate

to oblongo-ovate, more-or-less elliptic, thinly coriaceous, slightly bullate, 7-

9 cm wide x 13-23 cm long, base broadly rounded to sub-truncate, slightly

retuse or cuneate, apex acute to strongly acuminate to ca 2 cm, adaxial

surface semi-matte mid-green, abaxial surface paler green, glaucous, drying

strongly discolorous, midrib adaxially flush with lamina, abaxially prominent,

raised canaliculate in cross section; primary venation impressed adaxially,

strongly raised abaxially with up to 20 primary lateral veins on each side,

alternating with lesser interprimaries, frequently with a few branches from

near the base, both diverging at 45°- 60° and gradually curving towards the

apex before reaching the inter-marginal collecting vein; secondary venation,

arising from the midrib and the primary veins; tertiary venation obscure

tessellate adaxially and abaxially in both living plants and dry specimens.

Inflorescences up to four in quick succession alternating with foliage leaves

ca 5 cm long, (i.e. not forming a true synflorescence), fragrant (esterases) at

female anthesis; peduncle to ca 1 cm, not exserted from leaf bases, slightly

exserted in fruit; spathe interior glossy, exterior semi-glossy, coriaceous, ca

7.5 cm long, lower spathe narrowly ovoid, green when fresh, ca 2.5 cm long

x 1 cm diam., differentiated from the limb by a weak constriction: limb sub-

truncate, white, ca 5 cm long, coriaceous, opening wide except for convolute

base, at first over-arching, then reflexed, finally more or less twisted and

204 Gard. Bull. Singapore 62 (1) 2010

Plate 8. Schismatoglottis turbata S.Y.Wong. A. Whole plant; B. Two emerging inflorescences

with two infructescences; C. Inflorescence at male anthesis; D. Spadix with spathe artificially

removed; E. Female flowers, interstice and part of male zone; F. Male zone and appendix; G.

Spathe limb deliquescent post anthesis. Images © P.C. Boyce

Studies on Schismatoglottideae 205

then caducous-crumbling, apex mucronate ca 4 mm long, green when

fresh; spadix sessile, adnate isodiametrically to the spathe in the lower ‘4

of female zone, whitish, to ca 5.5 cm long, less than the length of spathe:

female zone ca 1.7 cm long, 4 of spadix length; pistils many, crowded, round

to slightly rhomboid, with weakly bisulcate longitudinal ridges, ca 0.5 mm

diam. x 0.45 mm long; stigma large, bun-shaped, to ca 0.3 mm diam., ‘sof

ovary, punctiform, papillate, yellow in alcohol; interpistillar staminodes very

occasional among the pistils, crowded into a row along the spathe/spadix

adnation, stalked, clavate and flat-topped, ca 0.5 mm diam., similar size to

pistils; sterile interstice strongly narrowed, ca 1.4 mm diameter in the middle,

(2.5-) ca 7 mm long which is ‘4 of spadix length, basally more or less covered

with inflated abortive pistils (reduced stigmas), twice the diameter of ovary,

transversely sulcate, apically covered with two rows of sterile stamens,

irregular shape, slightly wider than stamens; male zone cylindric, ca 3.6 mm

diam. x 1.2 cm long, '4 of spadix length, marked increased in diameter from

interstice; stamens crowded, truncate, ca 0.4 mm across, dumbbell-shaped

with the connective not elevated above the thecae, connective narrow, pore

oblong and deep; appendix subcylindric, ca 1.8 cm long, '4 of spadix length,

yellow when fresh, turning white in alcohol, frequently chewed by beetles,

tapering to a sharp point, basally more or less not isodiametric with top

of male zone; staminodes of appendix more or less columnar, irregularly

polygonal, flat-topped, ca 0.5-1 mm diameter.

Other specimens seen: MALAYSIA. Sarawak, Kuching Division, Sempadi,

Sg. Limau, 250 m asl, 26 Mar 2004, RC. Boyce & Jeland ak Kisai AR-271

(SAR).

Distribution: Southern Sarawak, Kuching Division.

Habitat: Lithophytic on sandstone (Kuching Division), 41-250 m asl.

Notes: Schismatoglottis turbata is clearly distinguished from the rest of the S.

nervosa complex by having small and round male flowers with deep pores.

The male zone is dense but irregularly arranged. The appendix is markedly

increased in diameter than the male zone.

Etymology: The specific epithet is derived from the Latin ‘twrbatus’,

exasperating, in allusion to the feeling on discovering yet another novel

species in the S. nervosa complex.

10. Schismatoglottis ulusarikeiensis S.Y.Wong, sp. nov.

Ab alii speciebus grex nervosae Borneensibus spathae laminorum pro parte

206 Gard. Bull. Singapore 62 (1) 2010

majore viridis (non in toto albis) et per anthesin haud aperiens distinguitur.

~Typus: Malaysia, Sarawak, Sarikei Division, Ulu Sarikei, 01° 55’ 05.4”; 111°

29’ 35.8”, 59 m asl, 7 Dec 2005, P.-C. Boyce et al. AR-1588 (holo, SAR). Plate 9.

Leaves few together (to ca 5), innovations yellowish green, the lamina

sometimes rotting and leaving behind portion of petiole, with this rotting

away at a later stage; petioles terete, channelled slightly canaliculate adaxially

in cross section, sometimes D-shaped in cross section, strongly longitudinally

ridged, very minutely (strong lens required) and densely verruculate, 13-20 cm

long, sheathing in the lower '/3 to % with the sheaths tapering, fully attached,

wide open persistent; lamina always oblongo-ovate, sometimes ovate, softly

coriaceous, 13-21 cm long x 5-9 cm wide (ratio up to ca 3:1), the base broadly

rounded to sub-truncate, slightly retuse to cuneate, the apex acute to ca 1

cm, adaxial surface semi-glossy mid green, abaxial surface paler green, always

slightly bullate, glaucous, drying strongly discolourous; midrib adaxially

flush with the lamina but centrally channelled at the leaf base, abaxially very

prominent, raised canaliculate in cross section; primary venation prominent

but flushed on adaxial surface, sometimes slightly impressed, strongly raised

abaxially, numerous, 10-15 on each side of the midrib, alternating with lesser

interprimaries or these occasionally arising from the bases of the primary

veins but only occur at the leaf bases, diverging at 60°- 80° and gradually

curving towards the tip before reaching the intermarginal collecting vein;

secondary veins mostly arising from the midrib, occasionally from near the

bases of the primary veins, obscure adaxially and abaxially; tessellate tertiary

venation, obscure adaxially and prominent abaxially. Inflorescences up to

three together, erect, white, 6.5 to 7 cm long, peduncle 1.5 to 2.5 cm, slightly

exserted during fruiting; spathe, interior glossy, exterior semi-glossy, thickly

coriaceous, ca 9 cm long; lower spathe broadly ovoid, ca 1.5cm diam. x 3.5 cm

long, white when fresh, differentiated from the limb by a faint constriction

coinciding with the upper part of interstice; limb lanceolate, thickly coriaceous,

caducous by melting at or just after male anthesis, upper part of the limb

green prior to anthesis; spadix, sessile, isodiametrically attached to the spathe

in the lower '/s of female zone, cylindric, 5.5-6 cm long; female zone, cylindric,

green when fresh but yellow in alcohol, 1.5-2.2 cm long x 7-10 mm diam.,

’ of spadix length; pistils numerous, tightly arranged, elongate-ovoid to

subcylindric, ca 0.50 mm diam.; stigma sessile, large bun-shaped, ca 0.40 mm

diam., overtopping ovary, raised, papillate; interpistillar staminodes confined

to more or less a single ring at the base of the female zone, but sometimes in

robust inflorescence, a few scattering towards the distal part of the female

tzone, irregularly polygonal from above, twice the size of ovaries, flat-topped,

shorter than pistils; sterile interstice presence, 0.6-1 cm long x 0.7-1 cm wide,

Studies on Schismatoglottideae 207

Plate 9. Schismatoglottis ulusarikeiensis S.Y.Wong. A. Whole plant: B. Petiolar sheath

persistent and open; C. Inflorescence at just before female anthesis; D. Female zone, interstice

and part of male zone: E. Spathe prior to anthesis; F. Spadix with spathe artificially removed:

G. Inflorescence post male anthesis with the appendix and spathe limb deliquescent.

208 Gard. Bull. Singapore 62 (1) 2010

pistillodes that are c. half to two times larger than pistil, irregularly round,

white when fresh and in alcohol, ca 7 mm long, sharp constriction in between

pistillode and staminode zones; staminodes, larger than stamens, irregular

polygonal, flat-topped, white when fresh and turning yellow in alcohol, ca 5

mm long; male zone, cylindric, ca 1.8-2.2 cm long x 7 mm diam., '4 of spadix

length; stamens close-packed, irregularly butterfly-shaped from above and

neighbouring anthers with their lobes interdigitating, 0.5 mm diam., anthers

truncate, connective narrow and thecae large, pores large accounting of

the entire top of the thecae and shallow, C-shaped, the rims narrowly alate;

appendix cylindric, 2.1-2.3 cm long x 7 mm diam., '4 of spadix length, white

when fresh and in alcohol, blunt point at distall, basally contiguous with the

male zone; staminodes of appendix tightly arranged, large staminode, flat-

topped, 0.7-1.2 mm across, squat-columnar, staminodes at the distal portion,

sometimes forms slit. Fruits immature during observation.

Other specimens seen: MALAYSIA. Sarawak, Sarikei Division: Ulu Sarike1;01°

55’ 05.4”; 111° 29” 35.8”; 59 m asl, 7 Dec 2005, P.-C. Boyce et al. AR-1577 (SAR);

ibid., 7 Dec 2005, R-C. Boyce et al. AR-1579 (SAR); ibid., 7 Dec 2005, PC. Boyce

et al. AR-1588 (SAR) ; ibid., 7 Dec 2005, PC. Boyce et al. AR-1635 (SAR).

Distribution: Central Sarawak, Sarikei Division. Known only from the type

locality.

Habitat: Evergreen moist lowland forest on shales, 59 m asl.

Notes: Schismatoglottis ulusarikeiensis is immediately distinguishable from

the rest of the S. nervosa complex with primary venation adaxially flush with

surface and with less than 10-15 on each side. Petioles are terete, strongly

longitudinally ridged and without hyaline margin in S. ulusarikeiensis. The

lamina is longer but narrower as compared to the rest of the species in the

complex. The upper part of the spathe limb remains in green form prior to

anthesis and this is unobserved in any other species in the complex. The

spathe limb is barely open during anthesis and this is only observed in S.

brevicuspis. The spadix is robust and forms a uniform width throughout the

different zones. The female flowers are green when fresh and this is only

observed in S. tessellata.

Etymology: The specific epithet is derived from the name of the type and

only known locality.

Studies on Schismatoglottideae 209

Acknowledgements

This study is funded by the Ministry of Higher Education, Malaysia by

fundamental research grant scheme No. FRGS/01(04)/609/2006(42) under

Sarawak Forestry Department Research Permit No. NPW.907.4.2(I)-101 &

Park Permit No. 58/20076. The collaboration and support of the Sarawak

Forestry Corporation, the Forest Research Centre (Kuching), and the

Sarawak Biodiversity Centre, are gratefully acknowledged. Special thanks

are due to P. C. Boyce for his comments on the manuscript.

References

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Soepadmo (eds). 2010. Flora of Peninsular Malaysia, Series I. Ferns and

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Institute Malaysia, Ministry of Natural Resources and Environment,

Malaysia, 249 pp. Price: RM80/USD60

A new flora useful to the study of Singapore pteridophytes is now available.

The volume | of the new Flora of Peninsular Malaysia, Series 1, dealing with

fern and lycophyte groups, has just been published.

The flora, according to the new publication, was initiated to document

the indigenous plant diversity by providing reliable and accurate accounts

of plant families, genera and species found in Peninsular Malaysia, with

updated nomenclatures. It will consist of two series, with Series IT dealing

with gymnosperms and angiosperms.

The Vol. 1 of Series 1 of the new flora described 100 species in 21

genera and 9 families of ferns and lycophytes, representing about a sixth of

the spore producing vascular flora of the peninsula. The families treated

by various authors and included in vol. | are Selaginellaceae (K.M. Wong),

Psilotaceae (R. Kiew), Equisetaceae (R. Kiew), Osmundaceaee (R. Jaman),

Matoniaceae (R. Jaman & Y. Umi Kalsom), Schizaeaceae (R. Jaman & Y.

Umi Kalsom), Cibotiaceae (A.T. Nor Ezzawanis), Loxogrammaceae (R.

Jaman) and Grammitidaceae (B.S. Parris). All species are shown with a

distribution map and provided with an assessment of their conservation

status. But only selected species of each family are nicely and accurately

illustrated with line drawings. However, each family is further illustrated

with beautiful coloured photos of a few more representative species.

There is one nomenclatural novelty proposed in vol. 1 of the new

flora, namely, Tomophyllum callophyllum (C.H. Wright) Parris, and no

new species record is reported. The biggest taxonomical change is seen in

the Family Grammitidaceae where several new generic concepts recently

proposed based on molecular evidence are accepted.

Aside from the useful information on the taxonomy and nomenclature,

the book includes also a conspectus of the orders, families and genera of

ferns and lycophytes of Peninsular Malaysia, keys to the families, genera

and species of the plant groups treated, a historical account of botanical

collecting of ferns and lycophytes in Peninsular Malaysia, and interestingly,

a chapter on the assessment and conservation of these plant groups in

Peninsular Malaysia. A glossary and a separate index to the scientific names

and vernacular/common names of the taxa conclude the volume.

I have one question in mind after browsing through the hundred pages

of the flora. Figures A & B in Plate 2 represent two different species?

PIMP Gard. Bull. Singapore 61 (2) 2010

As a student of Malesian pteridophyte flora, I definitely look forward

to the completion of the publication of Series | of this new flora. The authors

of vol. 1 of Series 1 of the new flora are to be congratulated for this timely

and well-done accomplishment.

BC Van

The Herbarium

Singapore Botanic Gardens

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 name(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 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. The standard for authority citations is Brummitt & Powell,

Authors of Plant Names, RBG Kew.

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: | 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.

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 the final

reproduction.

When grouping photographs, the page size of the journal must be taken into account to optimize the

space. Colour photographs should only be included where colour adds significantly to the information content

of the article. High resolution digital images may be submitted. 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 er 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 main standards are Stafleu & Cowan, Taxonomic Literature,

ed. 2, Regnum Vegetabile, Utrecht, for abbreviated names of books, and Botanico-Periodicum-Huntianum (B-

P-H), Pittsburgh for abbreviated names of periodicals. The following style is required:

Medinilla alternifolia Blume, Mus. Bot. Ludg.-Bat. 1:1 (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 or sumitted electronically to gbs@nparks.gov.sg

NATIONAL

PARKS

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© BOTANY

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