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 -BsCe lan Dr. Jana Leong-Skornickova

Singapore Botanic Gardens Singapore Botanic Gardens

(Editor) (Assistant Editor)

Dr PSY, Tan Ms. C. Soh

Singapore NParks Board Singapore Botanic Gardens

(Assistant Editor) (Journal Business Manager)

EDITORIAL BOARD

DrS:C. Chin Dr. M.C. Roos

Singapore Botanic Gardens National Herbarium Netherlands

Leiden University, The Netherlands

Dr. M.J.E. Coode

Royal Botanic Gardens Dr. E. Soepadmo

Kew, U.K. Forest Research Institute Malaysia

Kepong, Malaysia

Prof. Sir P. Crane

University of Chicago Prof. T. Stuessy

Chicago, USA University of Vienna

Austria

Dr. R.T. Corlett

National University of Singapore Dr. W.K. Tan

Singapore National Parks Board, Singapore

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

Department of Botany, NUNH Research Associate

Smithsonian Institute Singapore Botanic Gardens

Washington DC, USA

The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore.

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[Cover photo: Bulbophyllum entobaptum (see p. 145); photo by J.J. Vermeulen]

M =

ie /, JAN 05 2009 ROD ARCS ETM

RES RY

The Gardens’ Bulletin

Singapore

VOL. 60 (1) 2008 ISSN 0374-7859

CONTENTS

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

Studies on Homalomeneae (Araceae) of Borneo I. Four new species and preliminary

thouenis ON IntOnMall SPECIES! CLOUPS Ii) Sala WAK. .-.:.<..<0cc-s<2-0--2<-2<.esacessencessaeeeseceess oe

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

Hapaline celatrix (Araceae: Caladieae) — a new record for Sarawak, Malaysian Borneo

sengecessedesec sae enes aoele mabe oats coker? aide ae eee ES Ao nse ea 3]

Jutta, M. P.T. Ong and S. N. Phoon

New records for the Flora of Peninsular Malaysia, Family Orchidaceae |

Appendicula floribunda, Bulbophyllum elevatopunctatum, Cymbidium sigmoideum

ATR CTIALO C/U CIAGRATACHSE weer tot seec eons eens coe cvaen ane ov vecesueeyssoceeeeseeseertaeess 37)

Kurzweil, H.

Studies in the Peristylus tentaculatus-complex (Orchidaceae) in Thailand ................... 45

Kurzweil, H.

Habenaria mandersii (Orchidaceae) newly recorded from Thailand with notes

UBL E ST OSSCUSTI LOU Wren once ct sea tare ean eat oes Bec es oa ca dacc <5 cb Sad ceens dewaceeeunoevsteceetenese ce: 55

Turner, I. M.

Polyalthia saprosma (Annonaceae), a new species from Borneo................::..0:++6+- = 03

Veldkamp, J. F.

Sarcotheca lunduensis (Oxalidaceae), a new species from W. Sarawak, Malaysia........ 69

Vermeulen, J. J. and P. O’Byrne

Thirty two new Species of Bulbophyllum (Orchidaceae) from Sulawes!1 ...............::::0++ Se,

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

Studies on Schismatoglottideae (Araceae) of Borneo II: Schismatoglottis confinis,

a putative sister taxon to Schismatoglottis bauensis from Sarawak, Malaysian

BOLD EXO ys Rs ec os Be Re a ee

Yeh, C.-L., J.-H. Chen, C.-R. Yeh, S.-Y. Lee, C.-W. Hong, T.-H. Chiu and Y.-Y. Su

Musa yamiensis C. L. Yeh & J. H. Chen (Musaceae), a new species from Lanyu,

GIN GUD se scoe cated eee sodeseescecetccocbodecdeL Rca aaai aoe eee BRS aSoCRe OEE ee 165

Book Reviews

Vascular Flora of Ko Hong Hill, Songkla Province, Thailand by J.F. Maxwell

| 2 Rl) ee SE eee cence accdl Sa scaesoccanosbaccdondoocosse 173

Date of publication: September 2008

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Photoplates Pte Ltd

Gardens’ Bulletin Singapore 60 (1): 1-29. 2008 |

Studies on Homalomeneae (Araceae) of Borneo I.

Four New Species and Preliminary Thoughts on

Informal Species Groups in Sarawak

PETER C. BOYCE ' AND WONG SIN YENG ”

'Malesiana Tropicals, Level 5, Tun Jugah Tower,

No. 18, Jalan Tunku Abdul Rahman,

93100 Kuching, Sarawak, Malaysia

"Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Samarahan, Sarawak, Malaysia

Abstract

Four new species of Homalomena (Araceae: Homalomeneae) are described

from Sarawak. The current supraspecific taxonomy of the genus is reviewed

and reasons for not recognizing at this time formal supraspecific units are

justified although the need for species groupings is restated. In line with

other large, taxonomically intractable groups, informal groups are proposed

and circumscribed. All new species are illustrated.

Introduction

Homalomena Schott is a genus of in excess of 150 species distributed in

the new and old world tropics with the majority of species and greatest

diversity centred on perhumid South East Asia where there are three centres

of diversity: Sumatera, Borneo and New Guinea.

Phylogenetically Homalomena, together with Furtadoa M.Hotta

(two species, west Malesia, differing from Homalomena by male flowers

each associated with a pistillode), comprises the tribe Homalomeneae

M.Hotta, which is sister to tribe Philodendendreae Schott and part of a clade

containing otherwise African genera Cul/casia P.Beauvois and Cercestis

Schott (together comprising tribe Culcasieae Engl.).

Since the now long out-of-date full revision of Engler (1912) there

have been fragmentary floristic accounts by Ridley (1905), Merrill (1922),

Alderwerelt (1922a, b) and Furtado (1939, 1940), an uncritical species

listing for Malesia focusing primarily on Sumatera (Hotta, 1985), a revision

for New Guinea and the Bismark Archipelago (Hay, 1999), and various

ad hoc new taxa published by Hotta (1986, 1993), Boyce (1994), Hay

Gard. Bull. Singapore 60 (1) 2008

& Herscovitch (2002) and Sulaiman & Boyce (2005), but no attempt to

undertake a full revision of Homalomena. The lack of a reliable taxonomy

poses considerable problems for field workers since Homalomena is one of

the most speciose and taxonomically intractable aroid genera in the Asian

tropics.

The problems presented by a lack of reliable a taxonomy are

compounded by the poor state of preservation of many of the historical

types, the cryptic nature of most of the systematically significant

morphologies, notably the presence, absence and disposition of sterile

flowers, the generally large and complex vegetative structures that do not

lend themselves readily to traditional herbarium vouchering methodologies,

and the fleeting anthetic period such that even well prepared herbarium

specimens are frequently taxonomically useless because inflorescences

were prepared post anthesis by which time many significant structures

have deliquesced or been subjected to pre-preservation damage by the most

frequent inflorescence visitors, chrysomelid beetles, and post-preservation

destruction by herbarium beetles.

Homalomenaisataxonomically complex group and notwithstanding

the above difficulties, is in urgent need of a rigorous study aimed at

resolving the taxonomy and phylogeny. This is imperative not only because

Homalomena is one of the most abundant, speciose and least understood of

the mesophytic aroid genera in tropical Asia, but also because the genus is

now becoming the focus of interest for pharmaceutical research due to the

terpenoids and flavonoids occurring in the plant tissues; such studies must

have a basis in sound taxonomic understanding or risk being futile. That a

taxonomic study is required is no better exemplified than by the work of

Hanne Christensen (Christensen, 2000) in which five Homalomena species

are highlighted as having moderate to significant importance as medicinal

plants among indigenous people in Sarawak; four of the five species are

scientifically novel.

In conclusion, Homalomena requires a worldwide revision including

investigation of its phylogeny. However, current restraint on available

taxonomic expertise means it is unlikely that Homalomena will be so studied

in the near future. Nonetheless, as noted by Hay and Herscovitch (2002)

and reiterated by Sulaiman and Boyce (2005) there remains a need to be

able to identify at least the more distinctive taxa and, thus, although it is

undesirable to encourage ad hoc description of new species in taxonomically

chaotic genera, at least for the moment description of the more distinctive

taxa is desirable to enable some reliable parameters to be set.

Studies on Homalomeneae (Araceae) of Borneo I.

The genus Homalomena Schott

Homalomena Schott in H.W.Schott & S.L.Endlicher, Melet. Bot.: 20

(1832).

Homalonema Endl., Gen. Pl.: 238 (1837), orth. var.

Spirospatha Raf., Fl. Tellur. 4: 8 (1838).

Cyrtocladon Griff., Not. Pl. Asiat. 3: 147 (1851).

Chamaecladon Migq., Bot. Zeitung (Berlin) 14: 564 (1856).

Adelonema Schott, Prodr. Syst. Aroid.: 316 (1860), syn. prov.

Curmeria Linden & André, Ill. Hort. 20: 45 (1873), syn. prov.

Diandriella Engl., Nova Guinea 8: 20 (1910).

Minute to very large evergreen herbs, erect to decumbent, less often

creeping, usually strongly aromatic (terpenoids — frequently reminiscent of

mango or citrus peel, or ginger in Asian tropics; anise in the Neotropics).

Stems solitary, clustering or creeping and rooting with the terminal portion

erect, physiognomically unbranched or rarely pseudodichotomous. Leaves

spirally arranged, less often spiro-distichous, very rarely distichous,

sometimes together with petioles with conspicuous reddish extrafloral

nectaries; petioles longer to shorter than the lamina, channelled to terete or

D-shaped, the lower part sheathing, petiolar sheath persistent to marginally

marcescent; laminae simple cordate to oblanceolate, glabrous in tropical

Asia (but spiny and/or pubescent in most Neotropical species); primary

veins arising in a cluster at base of the lamina also distributed along the

midrib, veins of the posterior lobes (where present) arcuate, the remainder

running distally to the marginal vein; secondary and tertiary veins poorly

differentiated from one another, striate. Inflorescences several together,

terminal but displaced as to appear to be arising from the axils of the

leaves, each synflorescence with inflorescences developing sequentially,

individual inflorescences erect before and during anthesis, then becoming

decumbent to declinate during fruit development; spathe persistent through

to maturation of fruit, mostly simple boat shape or constricted into a lower

convolute and distal variously opening limb, exterior smooth or externally

ribbed and keeled along the dorsal midline, apex apiculate to strongly

mucronate. Spadix divided into two zones: female proximately, male

distally with the zones occasionally separated by a naked to staminode-

bearing interstice, male part often secreting reddish to brown resin prior to

or during anthesis; female flowers naked, often accompanied by a single

staminode arising from its base on the side nearest to the base of spadix

(hereafter referred to as interpistillar staminodes); ovary incompletely two-

to fully four-locular, style apparently absent or extremely short; stigma

4 Gard. Bull. Singapore 60 (1) 2008

smaller than to exceeding the ovary diam., button like to discoid, sometimes

weakly lobed, sometimes impressed, minutely papillate before and during

anthesis, often sunken afterwards; placentation basal to central, ovules

anatropous, several per locule. Male flowers with two to four, very rarely a

solitary stamen, filaments very short to + absent; anthers opening by short

apical longitudinal slits usually concealed by the expanded connective

forming a flat cap over the top of the stamen. Infructescences pendant

during development and at maturity, fruits contained within the persistent

and somewhat enlarged spathe; fruiting spathe dehiscent from the base

upwards, usually the spathe circumscissile at the insertion of the peduncle,

then splitting into a few irregular strips and these curling upwards to reveal

the ripe fruit. Fruit where known, a small translucent greenish berry,

usually smelling of overripe fruits. Seeds albuminous, very small, ca 1 mm

or less long, longitudinally ridged.

Distribution: Indo-Malesia to southern China eastwards to the Solomon

Islands with centres of diversity in Sumatera, Borneo and New Guinea; ca

8 Neotropical species (section Curmeria but generic status doubtful).

Habitat: Primarily understorey herbs in lowland evermoist tropical forest,

but also reaching mid-montane zone; sometimes rheophytic, very rarely

helophytic, occasionally relictual in regrowth and along road cuttings.

Note: Formerly the generic circumscription of Homalomena_ was

considerably narrower than currently accepted, with seven genera

recognized: Adelonema Schott, Chamaecladon Migq., Curmeria Linden &

André, Cyrtocladon Griff., Diandriella Engl. and Spirospatha Raf. Some

of these former genera are currently employed as subordinal taxa, as

discussed below.

Proposed informal groupings in Asian Homalomena

Current sectional groupings in Homalomena are based upon the work of

Schott (1860) and Engler (1912), with additions by Furtado (1939) and

Hotta (1967). Five sections are presently recognised: Curmeria (Linden

& André) Engl. & K.Krause (including Adelonema Schott) restricted to

the Neotropics, and presently the focus of molecular phylogenetic research

by Barabé and co-workers; Homalomena (‘Euhomalomena’ of Engl. &

K.Krause); Cyrtocladon (Griff.) Furtado; Chamaecladon (Mig.) Engl. &

K.Krause, and Geniculatae M.Hotta. All except Curmeria are restricted

Studies on Homalomeneae (Araceae) of Borneo I. 5

to the Asian tropics, distributed from North East India, eastwards to the

tropical Western Pacific, and, with the exception of Geniculatae, all have

been recognised as genera at some point in their history.

Of the remaining former generic taxa, Spirospatha Raf. (based on

Calla occulta Lour.) is referable to Homalomena occulta (Lour.) Schott

(a species of dubious identity although provisionally placed in section

Homalomena), while monospecific New Guinean Diandriella Engl. was

reduced to a generic and species synonym by Hay (1999) as Homalomena

stollei Engl. & K.Krause.

The present sectional boundaries are based upon overall inflorescence

shapes and sizes, the morphology of the sterile and fertile flowers,

placentation and micropyle direction. Hay (1999) noted that in several of

the New Guinea species these distinctions break down and he declined

to utilize any generic subordinal taxa. However, our observations have

suggested that these simplified inflorescence characters overlay a complex

series of vegetative architectures, including the presence of hapaxanthic

and pleionanthic shoot modules in section Homalomena, presence and

absence of aromatic tissues, in section Geniculatae a complicated shoot

arrangement not paralleled in any other species yet described, and in section

Cyrtocladon complex spathe and spadix movements during anthesis. As

in the case of Schismatoglottis (Schismatoglottideae) another large and

diverse mesophytic genus also distributed in the new and old world tropics

although phylogenetically distant from Homalomena, it seems likely that

vegetative architecture and, particularly, the spathe movement mechanics

during anthesis, are phylogenetically more significant than inflorescence

gross morphology and that the maintenance of the formally recognised

higher taxa is not useful until such time as a comprehensive phylogenetic

study is undertaken.

Nonetheless, in such a large and diverse genus with many novelties

yet to be described, subordinal units are a useful tool to facilitate taxonomic

study. In other taxonomically intractable groups (e.g. A/ocasia G.Don.,

Schismatoglottis Zoll. & Moritzi, the Pothoeae Engl. and Rhaphidophora

Hassk.) the establishment of informal groups has become a standard

approach until such time as phylogenetic testing can be undertaken leading

to the establishment of evolutionarily robust groups (see Boyce 2000a, b, c,

2001a,b; Boyce & Hay 2001; Hay 1998; Hay & Wise 1991; Hay & Yuzammi

2000). We are following this methodology here in proposing the reduction

of the four currently recognised Asian subgeneric groupings into three

informal supergroups: Homalomena, Cyrtocladon and Chamaecladon,

while reducing Geniculatae to Cyrtocladon.

6 Gard. Bull. Singapore 60 (1) 2008

The supergroups are defined by the following characters -

The Homalomena supergroup comprises medium to large creeping to erect

plants with strongly aromatic tissues, pleionanthic, or rarely hapaxanthic,

shoot modules and spathes greater than 1.5 cm long, with no or only a very

weak constriction between the upper and lower spathe. Spathe movement

during anthesis, where known, comprises simple gaping and then closing

of the spathe limb, no spadix movements have been recorded. Ovary three

to four locular. Male flowers with three to four, rarely five to six, anthers.

The Chamaecladon supergroup comprises small to minute often creeping,

less often erect plants with odourless, or very rarely aromatic, tissues; as far

as is known only pleionanthic shoot modules, and spathes less than 1 cm,

very rarely up to 1.5 cm long, with no constriction between the upper and

lower spathe. Spathe movement during anthesis, where known, comprises

simple gaping and closing of the spathe limb. No spadic movement recorded.

Ovary two to three locular. Male flowers with two to three anthers.

The Cyrtocladon supergroup comprises medium to very large erect to

creeping plants with strongly aromatic tissues, pleionanthic (but very few

studied) shoot modules and spathes greater than 2 cm long, with weak to

moderate to pronounced constriction between the upper and lower spathe.

All of the several species so far studied undergo a complex series of spathe

and spadix movements during anthesis. Ovary three to four locular. Male

flowers with three to four, rarely five to six, anthers.

Previously described Bornean taxa in relation

to the four new species of Homalomena

The four new species described in this paper all belong to the Cyrtocladon

supergroup. Based on our literature review and examination of problematic

types, we are confident that despite the current chaotic state of Homalomena

taxonomy in Borneo, none of our proposed novelties have been featured in

the literature on Bornean Homalomena.

I. The publication of Ridley (1905) -

The first attempt to bring order to Homalomena specifically in

Borneo was that of Ridley (1905). Ridley listed 23 species for Borneo,

of which eight [H. griffithii (Schott) Hook.f, H. intermedia Ridl., H.

Studies on Homalomeneae (Araceae) of Borneo I. il

ovatifolia Ridl., H. paucinervia Ridl., H. pumila Hook. (= H. humilis (Jack)

Hook.f. ), H. saxorum (Schott) Engl. & H. truncata Hook.f | can be dismissed

immediately from this discussion as they belong to the Chamaecladon

supergroup. Of the remaining 15 species, taxonomic and/or nomenclatural

problems exclude three from further discussion.

H. aromatica Schott var. cordata (Schott) Ridl. is based upon a

plant of unknown origin; the superior taxon is endemic to NE India and

Bangladesh and belongs to the Homalomena supergroup.

H. fasiata Ridl. is a synonym of Schismatoglottis tecturata (Schott)

Engl. (Tribe Schismatoglottideae).

H. sagittifolia Jungh. is here regarded as a Javan endemic with

much of the modern interpretation of it as a widespread species in Sunda

based upon a series of misinterpretations of the additional specimens cited

by Schott, and latterly by Engler (1912). The numerous cordato-saggitate

Homalomena species in Sarawak that are commonly referred to as ‘H.

sagittifolia will be the subject of a paper in preparation by the authors.

Of the remaining 12 species treated by Ridley, three (H. beccariana

Engl., H. miqueliana Schott, and H. paludosa Griff.) are helophytes, often

in oligotrophic water systems and not at all related to the species proposed

here. Of the rest, five (H. havilandii Ridl., H. insignis Ridl, H. lancea Ridl.

H. ovata Engl. and H. sarawakensis Ridl.) are all with oblong to lanceolate

leaves, lacking posterior lobes, and while in this morphology approaching

H. pseudogeniculata, all differ in lacking a pulvinus at the insertion of the

lamina on the petiole.

Of the remaining four species, Homalomena borneensis Ridl. has

sagittate, not cordate leaves and a partially naked interstice between the

female and male zones. To date H. borneensis is known with certainty only

from the Kuching type. Homalomena crassinervia Ridl., H. punctulata Engl.

and H. subcordata Engl. are all based upon Matang (Kuching Division)

types, and given their morphological differences coupled with the high levels

of local endemism, which has proven in the work on Schismatoglottis (e.g.,

Hay & Yuzammi, 2000) and in mesophytic terrestrial aroids in general to

be of great importance in delimitating taxa, they can be effectively dismissed

from consideration.

Regarding the status of H. propinqua Schott there exist two problems.

First, Ridley’s (1905) interpretation allies the plants more closely to H.

sagittifolia sensu Ridl. (see above), whereas Schott’s original description

appears to belong to the Chamaecladon supergroup. Secondly, the type of

H. propinqua is from Java.

8 Gard. Bull. Singapore 60 (1) 2008

2. The publications of Alderwerelt (1922) -

Alderwerelt (1922a, b) described and discussed 37 species of Bornean

Homalomena, of which 19 are members of the Chamaecladon supergroup

and 10 are in the Homalomena supergroup. Of the remaining eight species

treated by Alderwerelt, H. raapii Engl. is a Sumateran endemic, while

H. miqueliana is, as noted above, a helophyte, and H. propinqua has the

problems as outlined earlier. Of the remainder, only two (H. /atifrons Eng]

& H. sulcata Engl.) are Bornean (both from Kalimantan) and are narrowly

endemic.

3. The publication of Furtado (1939) -

Furtado’s comprehensive treatment made in 1939 was an attempt to bring

order to the chaotic state of Homalomena primarily in Peninsular Malaysia

and Sumatera. Aside from systematic notes, including complete synonymy,

references and citations, and localities with exsiccatae covering 58 species,

his publication included a review of the sections, together with key and

summary, list of collections seen, and index to all names included, and the

description of a new section, Cyrtocladon. Regrettably the work 1s beset by

a number of problems, no least a tendency to recognize extremely finely

defined subordinal taxa and in several instances heterodox definitions of

taxa in order to ‘mop-up’ problematic plants. Many of the names that can

be applied to Sumateran taxa only are thus ‘stretched’ to fit species from

Borneo and elsewhere. In our opinion, aside from a very few widespread

species (e.g., H. griffithii Hook.f.) Furtado’s work does not apply to

Borneo.

4. The publication of Hotta (1967) -

The only relevant species published by Hotta (1967) is Homalomena

geniculata M.Hotta, which is discussed below under the description of H.

pseudogeniculata P.C.Boyce & S.Y.Wong.

New species of Homalomena

Homalomena ardua P.C.Boyce & S.Y.Wong, sp. nov.

Homalomena ardua differt ab speciebus ceteris Borneensibus laminis

foliorum adaxiali atro-viride, nitentibus, subtus atrorubidus cum nervus

centralis et nervis lateralibus primariis prominentibus, petiolis atrorubidus

vel viride, forma rubidus cum petioli demum viridescens usque ad */, partem

longitudinis petiolo, petioli vagina longa (usque ad 3 partem longitudinis

petiolo). Inforescentia mascula gracilis, 2/3 longitidinis inflorescenta

Studies on Homalomeneae (Araceae) of Borneo I. 9

occupatus. Ab H. josefii (nuper descripto subtus similis) sed stipiteque

longiore (ca 8.5 mm longus versus ca 3.5 mm longus), floribus femineus ad

basin spadice bis magnis et stigmatibus parum impresso et cum alcoholis

atrobrunneus (stigmatii S. josefii elevatus cum alcoholis brunneus pallide),

inflorescentiae feminae staminodiis cum alcoholis brunneus pallides (vs

griseus cum §. josefii), inflorescentia mascula longiore (ad */, versus '>

partem longitudinis spadice) et petioli vagina proportione longa (% partem

longitudinis petiolo H. ardua versus '/, to '/, partem longitudinis petiolo

H. josefii differt.— Typus: Malaysia, Sarawak, Miri Division: Mulu, Long

Lama, Mulu National Park, National Park Headquarters, 04° 02’ 29.4”,

114° 48° 44.3”, 5 Aug 2007, PC. Boyce et al. AR-1938 (Holo, SAR+ spirit).

Plates 1 & 2.

Medium to robust herbs, strongly aromatic (mango peel), evergreen,

glabrous, toca 80cm tall. Stem pleionanthic, erect to ascending, ca3 cm thick,

dark red to green, internodes to ca | cm long. Leaves up to ca 20 together;

petiole terete, erect to decumbent with the terminal portion ascending, the

laminae held flat, up to 50 cm long, petiole bases clasping, petioles dark

reddish to green, reddish forms with the */, lower part of petiole ageing to

dark green, matte, drying dark brown, petiolar sheath to ca 31 cm long, near

to '/, of petiole length, unequal, broader side rounded at apex, narrower side,

weakly decurrent at apex, sheath initially long-persistent with the marginal

1.5 mm soon drying paler, eventually the whole sheath marcescent; lamina

broadly ovato-sagittate, 40-50 cm long x 28-34 cm wide, thinly leathery,

glossy dark green adaxially (fresh), drying pale olive green, abaxially dark

red (fresh), drying pale brown, base cordate, posterior lobes spreading,

subtriangular 6-11 cm long, lamina tip obtuse, short-acuminate for ca | cm,

thence, apiculate for ca 3.5 mm, sometimes red; midrib raised abaxially

(fresh and dry), dark red when fresh, drying reddish brown, adaxially flush

with lamina (fresh and dry), ca 3 mm wide, with ca 10 primary lateral veins

on each side, diverging at 50°-90° from the midrib, adaxially impressed

(fresh), flush with lamina when dry, abaxially raised (fresh and dry), curved

sharply towards the apex when near the margin, interprimary veins ca

width of the primary lateral veins, alternating irregularly with primaries,

posterior lobes each with 3-4 primary lateral veins; secondary venation

rather obscure, striate; tertiary venation not visible, all veins running into

a thickened intermarginal vein, this particularly conspicuous at the leaf

tip, drying paler than the lamina. Inflorescences 1-5 together, erect at

anthesis, later declinate, peduncle to ca 10 cm long x ca 3 mm diam, white,

matte. Spathe 9-15 cm long, tightly furled prior to anthesis, lower spathe

inflating at female anthesis, spathe limb loosening at female anthesis,

10 Gard. Bull. Singapore 60 (1) 2008

Plate 1. Homalomena ardua P.C.Boyce & S.Y.Wong. A. Overall plant; B. Abaxial leaf

lamina dark red with equally coloured midrib; C. Red petiole bases; D. Inflorescence

shedding pollen at male anthesis; E. Inflorescences with emerging inflorescence bud; note

the sequential emergence. The inflorescence at the back is at female anthesis; that next is

at male anthesis.

Studies on Homalomeneae (Araceae) of Borneo I. 11

Plate 2. Homalomena ardua P.C.Boyce & S.Y.Wong. A. Whole spadix, spathe removed,

from alcohol collection; B. Close up of female zone; C. Detail of female zone, note

especially the enlarged lowermost female flowers; D. Male zone.

thence, inflating and then opening wide, lower spathe pale green, stained

red at insertion of peduncle, white above, spathe limb white at anthesis,

with apex and mucro shading to dark pink during anthesis; lower spathe

ovoid-ellipsoid, ca 5.3-6.5 cm long, moderately constricted at the junction

of the spathe limb, the constriction coinciding with the lower-most fertile

male flowers, spathe limb narrowly elliptic, ca 8.5 cm long x 2.3 cm wide

(at male anthesis), prominently keeled along the exterior midline, spathe

limb margins with the middle ca 7/, reflexing slightly at male anthesis,

apex mucronate to ca 3.5 mm long. Spadix equalling the spathe, ca 13

12 Gard. Bull. Singapore 60 (1) 2008

cm long, elongate cylindrical-fusiform, narrowing in the lower part male

zone coinciding with the constriction of the spathe and there intergrading

with staminodes, stipitate; stipe ca 8.5 mm long x 4.5 mm diam., strongly

dorso-ventrally flattened, female zone ca 3.6 cm long x ca | cm wide, ca %

length of spadix, weakly fusiform, the surface adjacent to the spathe limb

flattened, female flowers ca 1.5 mm x 0.75 mm, laxly arranged, squat-

cylindrical, stigma as broad or slightly smaller than ovary, impressed and

weakly trisulcate, stained light brown, interpistillar staminodes truncate on

a very slender stipe, ca 0.3 mm diameter, equalling or slightly overtopping

the associated female flower, stained greyish in alcohol, lower-most female

flowers ca twice the size of fertile females, mostly associated with two

or more interpistillar staminodes and seemingly sterile, suprapistillar

staminodes each comprising a single sterile anther; male zone ca 8 cm

long, ca ’/; length of spadix, lower part weakly constricted, clothed with

fertile male flowers intergrading into a single row of staminodes., distal-

and proximal-most flowers apparently sterile; male flowers, ca 3 mm x

.2 mm trapezoid, comprising 3-7 truncate stamens, each overtopped by a

large, slightly raised connective. Infructescence declinate, spathe entirely

persistent, pale green stained reddish pink, peduncle dark red, matte. Fruits

and seeds not observed.

Distribution: Borneo: Sarawak - endemic, Miri Division. Known only from

the type collection.

Habitat: Not known in the wild, planted at the headquarter of Mulu National

Park, 65 m asl, but plants originating from the surrounding areas.

Notes: Homalomena ardua is a distinctive plant with the leaf laminae

glossy dark green adaxially and dark red abaxially with the midrib red and

prominently raised abaxially. Petioles are dark reddish to green, with the

reddish forms having petioles ageing to dark green at the lower ’/, of petiole.

Petiolar sheathes are long, extending to nearly 2 of the petiole length. The

male zone is long and slender which occupies ”/, of the inflorescence.

Homalomena ardua differs from H. josefii but the spadix being

longer stipitate (ca 8.5 mm in H. ardua) vs. ca 3.5 mm (H. josefii), the

female flowers at the lower part of the zone are twice the size those in

the upper zone vs. uniform size of flowers throughout the female zone in

H josefii. \t is speculated that the enlarged female flowers of H. ardua,

which appear to be functionally sterile, may be adapted to attracting or

maintaining pollinators. There are also differences in the stigma, which

is slightly impressed and stains dark brown in alcohol in H. ardua, but is

ws)

Studies on Homalomeneae (Araceae) of Borneo I |

raised and remains pale brown in alcohol in H. josefii. Further, interpistillar

staminodes stain greyish in alcohol in H. ardua but light brown in H. josefii.

Lastly, the male flower zone is much longer in H. ardua (occupying ”/,

length of spadix), as compared with that of H. josefii ('/, length of spadix).

Vegetatively H. ardua is notable by virtue of its leaves dark red abaxially

(green adaxially in H. josefii) and the proportionately longer petiolar sheath

(% of petiole length in H. ardua vs. '/, to '/, of petiole length in H. josefii.

One of the inflorescences (see 4R-/900) was collected at late male

anthesis (based on observation, male anthesis lasts for two to three days). It

was observed that the staminodes at the base of insertion and interpistillar

staminodes were eaten by visiting chrysomelid beetles.

Etymology: The specific epithet is from the Latin arduous — hard work — in

fanciful allusion to the fact that there is much hard work to be done in this

genus!

Other specimens seen: Sarawak, Miri Division: Mulu, Long Lama, Mulu

National Park, National Park Headquarter, 04° 02° 29.4”, 114° 48° 44.3”,

11 Aug 2007, P.-C. Boyce et al. AR-2002 (SAR).

Homalomena josefii P-C.Boyce & S.Y.Wong, sp. nov.

Planta magna usque ad 100cm alta quam foliorum laminii ad vena

submarginali abaxialiter valde prominentibus. Ad H. ardua (nuper descripto

leviter similis) sed stipiteque breviore (ca 3.5 mm longus versus ca 8.5

mm longus), floribus femineus in toto consimilis, stigmatibus elevatus

cum alcoholis brunneus pallide (stigmatii H. ardua parum impresso et

cum alcoholis atrobrunneus), inflorescentiae feminae staminodiis cum

alcoholis griseus (versus brunneus pallides cum H. ardua), inflorescentia

mascula breviore (ad 2 versus */; partem longitudinis spadice) et petioli

vagina proportione breviore ('/, to '/, partem longitudinis petiolo H. josefi

versus 4 partem longitudinis petiolo H. ardua differt. — Typus: Malaysia,

Sarawak, Bintulu Division: Bukit Satiam, 02° 59° 10.0”, 112° 55° 42.8", 14

July 2006, P.C. Boyce et al. AR-1894 (Holo, SAR + spirit). Plates 3 & 4.

Medium to robust herbs, strongly aromatic (ginger/resin), evergreen,

glabrous, to ca 100 cm tall. Stem pleionanthic, erect to ascending, ca 5 cm

thick, dark red to green, internodes to ca 1 cm long. Leaves up to ca 15

together, ca 5-7 per module; petiole terete, erect to decumbent, 50-70 cm

long, petiole bases clasping, eventually falling to leave a conspicuous lunate

scar, petioles dark reddish to green, dark reddish forms with longitudinal

ridges, green forms always with pinkish red bases, matte, drying dark

14 Gard. Bull. Singapore 60 (1) 2008

Plate 3. Homalomena josefii P.C.Boyce & S.Y.Wong. A. Overall plant; B. Abaxial leaf

lamina pale green, with red petiole; C. Red petiole bases; D. Petioles in green form

with inflorescences and infructescences; E. Emerging inflorescence bud with declinate

infructescences.

Studies on Homalomeneae (Araceae) of Borneo I. 15

Plate 4. Homalomena josefii P.C.Boyce & S.Y.Wong. A. Whole spadix, spathe removed,

from alcohol collection; B. Close up of female/male zone transition; C. Detail of female

zone, note evenly sized female flowers; D. Male zone and spadix tip; note the vestigial

naked appendix at the tip.

brown, petiolar sheath ca 16-21 cm long, ca '/, to '/, of petiole length,

equal, sometimes unequal, broader side rounded at apex, narrower side,

weakly decurrent at apex, margin always convolute when fresh, sometimes

wide open with broader petiolar sheath, sheath initially long-persistent

with the marginal 1.5 mm, soon drying paler, eventually the whole sheath

marcescent; lamina broadly ovato-sagittate, 25-45 cm long x 18-32 cm

wide, thinly leathery, glossy dark to pale green adaxially (fresh), drying pale

olive green, abaxially matte green (fresh), drying pale brown, base cordate,

posterior lobes spreading, subtriangular 7-9 cm long, lamina tip obtuse,

short-acuminate for ca | cm, thence, stiffly apiculate for ca 2-7 mm; midrib

raised abaxially (fresh and dry), green when fresh, drying reddish brown,

adaxially flush with lamina, ca 1.5 mm wide, with 6-9 primary lateral veins

on each side, diverging at 50°-90° from the midrib, adaxially impressed

(fresh), flush with lamina when dry, abaxially slightly raised (fresh and

16 Gard. Bull. Singapore 60 (1) 2008

dry), curved sharply towards the apex when near the margin, interprimary

veins ca 2 width of the primary lateral veins, alternating irregularly with

primaries, posterior lobes each with 3-4 primary lateral veins; secondary

venation rather obscure, striate; tertiary venation not visible, all veins

running into a thickened intermarginal vein, often red when fresh, this

particularly conspicuous at the leaf tip and there drying paler than the

lamina. Inflorescences 1-7 together, erect at anthesis, later declinate, each

subtended by prophyll to ca 9 cm long, followed by cataphyll, ca 2-8 cm

long, peduncle to ca 15 cm long x ca 5 mm diam, deep red, matte. Spathe

6.5-15.3 cm long, tightly furled prior to anthesis, lower spathe inflating at

female anthesis, spathe limb loosening at female anthesis, thence inflating

and then opening wide, lower spathe pale green, stained deep red at

insertion of peduncle, flushed pink above, spathe limb white at anthesis,

with apex and mucro shading to dark pink during anthesis; lower spathe

ovoid-ellipsoid, 2.5-6.5 cm long, moderately constricted at the junction

of the spathe limb, the constriction coinciding with the lower-most fertile

male flowers, spathe limb narrowly to broadly elliptic, ca 3.3-8.5 cm long

x 2.3 cm wide (at male anthesis), prominently keeled along the exterior

midline, spathe limb margins with the middle ca ’/; reflexing slightly at

male anthesis, apex mucronate to ca 3.5 mm long. Spadix equalling the

spathe, ca 6-15.3 cm long, elongate cylindrical-fusiform, narrowing in the

lower male zone coinciding with the constriction of the spathe and there

intergrading with staminodes, stipitate; stipe ca 3.5 mm long x 5 mm diam.,

short fusiform, few staminodes present at the insertion of peduncle, similar

to interpistillar staminodes, female zone ca 2 cm long x ca | cm wide, ca '/,

length of spadix, weakly fusiform, female flowers ca 1.3 mm x 0.75 mm,

densely arranged, round, stigma as broad or slightly exceeding the ovary,

raised and weakly trisulcate, staining pale brown in alcohol, extending

beyond the ovary as the translucent collar, mostly associated with two or

more interpistillar staminodes and seemingly sterile, staminodes truncate on

a very slender stipe, ca 0.3 mm diameter, equalling or slightly overtopping

the associated female flower, few pistillodes at the base of interstice, similar

size to female flowers, suprapistillar staminodes zone, to ca 1cm long x 5

mm wide, sometimes wider than female zone, staminodes each comprising

a single anther; male zone to ca 4 cm long, ca '/, length of spadix, separated

from interstice by weakly constricted lower part of male zone, clothed with

fertile male flowers intergrading into a single row of staminodes., distal-

and proximal-most flowers apparently sterile; male flowers, ca 3 mm x

2 mm, trapezoid, comprising 3-5 truncate stamens, each overtopped by a

large, flat connective, terminal-most flowers sterile and spadix often topped

with a vestigial naked appendix. Infructescence declinate, spathe entirely

Studies on Homalomeneae (Araceae) of Borneo I. 17

persistent, pale green stained reddish pink, sometimes wholey reddish pink,

peduncle dark red, matte. Fruits and seeds not observed.

Distribution: Borneo: Sarawak, Bintulu Division — endemic.

Habitat: Terrestrial on shales and seasonally inundated alluvium, 7-120 m

asl.

Notes: Homalomena josefii differs from H. ardua by the much shorter

spadix stipe to ca 3.5 mm in H. josefii vs. 8.5 mm in H. ardua, the uniform

size of the female flowers, the stigma raised and staining pale brown in

alcohol in H. josefii (stigma impressed and staining dark brown in H. ardua).

Additionally, the male zone is much shorter, occupying 2 of spadix length

in H. josefii, but longer (7/, length of spadix) and more slender in H. ardua.

Vegetatively H. josefii is distinctive by its large, robust form up to ca 100

cm, shorter petiolar sheath ('/, to '/, of petiole length) as compared to 2 of

petiole length in H. ardua, and leaf laminae always with a red, markedly

thickened intermargininal vein running to the leaf tip.

Leaf colour is variable in H. josefii, with petioles ranging from green

with red bases to more rarely, petioles red with concolorous longitudinal

ridges present, and leaf laminae green adaxially as compared to red adaxial

leaf laminae in H. ardua.

Etymology: The specific epithet is named for Dr Josef Bogner (Botanischer

Garten Miinchen), one of the foremost experts on the aroids and perhaps

the only person to have seen all currently recognized aroid genera in the

field.

Other specimens seen: SARAWAK: Bintulu Division, Bukit Satiam, 02°

99° 13:37, 112° 55° 57.5”, 14 Jul. 2006, P.-C. Boyce et al. AR-1900 (SAR);

Bukit Satiam, 02° 59’ 07.4”, 112° 55’ 47.0”, 15 Jul. 2006, P.C. Boyce et al.

AR-1908 (SAR); Bintulu, road to Kampung Jepak, ca 3.3 km after bridge

over Batang Kemena en route to Sibu from Bintulu, 03° 08’ 32.3”, 113° 03°

24.3”, 15 Jul. 2006, P.C. Boyce et al. AR-1911 (SAR).

Homalomena pseudogeniculata P.C.Boyce & S.Y.Wong, sp. nov.

Ab omnibus speciebus generis ceteris combinatio caulibus longis repentibus

valde robusti, petiolo ad apicem pulvinato, lamina foliiorum pro ratione

oblongo-elliptica vel ovato, in stato vivo aliquando abaxiali punctis pellucidus

instructa distinguitur. Ab H. geniculata follis spiro-distichis, coriaciis (non

chartaceis), sine staminodiis ad basin inflorescentia feminis et interstitio

18 Gard. Bull. Singapore 60 (1) 2008

quam inflorescentia mascula et femina plus late prompte distinguibilis est.

—Typus: Malaysia, Sarawak, Sarikei Division, Ulu Sarikei, 01° 55’ 05.4”,

111° 29’ 35.8”, 7 Dec 2005, P.-C. Boyce et al. AR-1583 (Holo, SAR). Plates

5& 6.

Medium to moderately robust herbs, strongly aromatic (pine resin),

evergreen, glabrous, to ca 50 cm tall. Stem pleionanthic, decumbent with

apex erect, frequently creeping for several metres and branching laterally

while still continuing a physionogmically unbranched primary axis, green,

internodes to ca | cm long. Leaves ca 8-12 together, ca 5 per module, each

module subtended by prophyll, up to ca 12 cm long; petiole terete, erect to

decumbent, up to ca 30 cm long, pulvinate, ca 3-13 cm from lamina base,

roots penetrating petiole bases, petioles green when fresh, matte, drying

light brown, petiolar sheath to ca 14 cm long, ca '/, of petiole length, sheath

convolute, initially persistent, eventually the whole sheath marcescent;

lamina oblongo-lanceolate, sometimes oblongo-elliptic to ovate, 18-33 cm

long x 6-15 cm wide, rather thinly coriaceous, matte mid-green adaxially,

sometime the mid-rib paler (fresh), drying pale olive green, abaxially matte

pale green, sometimes with conspicuous pellucid dots when fresh, very

occasionally red (fresh), drying pale brown, base decurrent to truncate, tip

obtuse, acuminate for ca 2 cm, thence, apiculate to ca 9 mm; midrib raised

abaxially (fresh and dry), drying straw-coloured, adaxially flush with lamina,

but slightly channelled towards the leaf base, ca 2.5 mm wide, with 6-9 primary

lateral veins on each side, diverging at 45°-55° from the midrib, adaxially

impressed (fresh), flush with lamina when dry, abaxially slightly raised (fresh

and dry), curved towards the apex when near the margin, interprimary veins

ca 2 width of the primary lateral veins, alternating irregularly with primaries,

secondary venation rather obscure, striate; tertiary venation not visible, all

veins drying in intermittent raised and flush strips especially when near to

leaf margin, all veins running into intermarginal vein. Inflorescences 1-5

together, erect, each subtended by a prophyll up to ca 6.2 cm long, peduncle

to ca 12-15 cm long x 1.5- 1.6 cm diam, yellowish green. Spathe ca 10.6

cm long, tightly furled prior to anthesis, loosening at female anthesis and yet

further at male anthesis, lower spathe yellowish green to white at maturity,

spathe limb white prior to and at anthesis, with apex and mucro pale green at

anthesis; lower spathe narrowly ellipsoid, ca 3.5 cm long, weakly constricted

at the junction of the spathe limb, the constriction coinciding with the lower-

most fertile male flowers, spathe limb narrowly lanceolate, ca 7 cm long,

mucronate to ca 7 mm long. Spadix shorter than the spathe, ca 8.3 cm long,

stipitate, stipe ca 4.5 mm long, weakly dorso-ventrally flattened, obliquely

inserted on peduncle; female zone ca 2.2 cm long x 6.5 mm wide, ca 4 length

Studies on Homalomeneae (Araceae) of Borneo I.

Plate 5. Homalomena pseudogeniculata P.C.Boyce & S.Y.Wong. A. Overall plant;

B. Pulvinate petioles; C. Declinate infructescences; D. Emerging inflorescence with

distinctive mucro; E. Two synflorescences.

20 Gard. Bull. Singapore 60 (1) 2008

Plate 6. Homalomena pseudogeniculata P.C.Boyce & S.Y.Wong. A. Whole spadix, spathe

removed, from alcohol collection; B. Close up of female and stipe; C. Detail of female/

sterile interstice/male zone, note the interstice width exceeds that of the male and female

zones; D. Male zone and spadix tip.

of spadix, weakly fusiform, female flowers densely arranged, ca 1.3 mm

diam. x | mm tall, round-cylindrical, lower-most female flowers ca twice

the size of fertile females, stigma exceeding the ovary, coherent to adjacent

stigma, slightly raised, staminodes absent at the base of insertion, and

female flowers with no associated interpistillar staminode, interstice zone

wider in diameter than the rest zones, staminodes truncate, ca 1.5 mm wide,

slightly overtopping the female flowers, male zone ca 4.8 cm long x 5.2 mm

wide, ca '/, length of spadix, cylindrical and tapering to a sharp end, distal

and proximal most flowers apparently sterile, narrowing in the lower part

coinciding with the constriction of the spathe; male flowers ca 3 mm x 1.6

mm, trapezoid, comprising (3)4-7 truncate stamens, overtopped by a large

connective, seemingly fertile to the tip. Infructescence declinate, spathe

Studies on Homalomeneae (Araceae) of Borneo I. Z

entirely persistent, lower spathe dark red, limb green, peduncle green. Fruits

and seeds not observed.

Distribution: Borneo, endemic: Sarawak, Kuching, Sarikei, Kapit and Miri

Divisions; Brunei.

Habitat: Always terrestrial mostly under full shade in deep soil on various

substrates, frequently on shales, rarely on granite. 62 m — 600 m asl.

Notes: Homalomena pseudogeniclata is distinctive by its pulvinate petioles

and remarkable decumbent-creeping stem giving rise to short leafy side

shoots while maintaining a primary axis. Plants frequently occur growing

down steep forested slopes giving the impression of several individual plants

in arow but on investigation revealing a single creeping stem/rhizome with

numerous short lateral branches.

Homalomena_ pseudogeniculata is the third pulvinate-petioled

Homalomena species to be formally described. The first was Homalomena

geniculata M.Hotta from which H. pseudogeniculata differs by spiro-

distichous leaves, with coriaceous leaf laminae and overall, much more

massive habit. Homalomena pseudogeniculata is further characterised by

having laminae mostly oblongo-lanceolate, sometimes oblongo-elliptic to

ovate, sometimes with conspicuous pellucid dots at abaxial surface when

fresh, all veins drying in intermittent raised and flush strips especially

those near the margins. Inflorescences of H. pseudogeniculata differ

in lacking basal staminodes, the female flowers densely arranged and

without associated interpistillar staminodes, and an interstice wider than

the fertile zones. Interestingly both species have a decumbent rhizome-like

stem and a predilection for growing down slopes. The pulvinus is at the

petiole/lamina insertion in H. geniculata; mid-way along the petiole in H.

pseudogeniculata.

The other currently recognized species with geniculate (pulvinate)

petioles is the Sumateran endemic H. elegantula A.Hay, which differs

from H. pseudogeniculata, among other characters, by hapaxanthic shoots,

overall much smaller and less robust habit and smaller (1 cm long) spathes

with only a very weak constriction between the limb and lower part.

A note on the application of the terms pulvinate and geniculate

seems appropriate. Currently these terms are used interchangeably in the

aroids to define a swelling or cushion-like structure, most often at the base

or apex of a petiole; such a presence is frequently used to define several

of the major generic divisions in the aroids, notably the subfamilies

Pothoideae and Monsteroideae. In fact, numerous genera outside of these

DD, Gard. Bull. Singapore 60 (1) 2008

families have some or all species with such structures. The strict definition

of geniculate is bent as in a knee whereas that of pulvinate is a swelling or

cushion. It is the latter definition that more accurately fits the situation in H.

pseudogeniculata. A more detailed paper on these terms is being prepared

by the first author.

Etymology: The specific epithet is coined from the superficial similarity of

the leaves of this species to H. geniculata — hence pseudo — false.

Other specimens seen: SARAWAK: Kuching Division: Lundu, Gunung

Gading, trail to Waterfall, trail above Batu Apek, 01° 41’ 48.2”, 109° 50’

20.5”, 14 Dec 2006, P.-C. Boyce et al. AR-2064 (SAR); Kapit Division: Nanga

Gaat, Rejang Wood Concession, stream below Camp Gahada, 01° 41’ 49.4”,

113° 26’ 16.3”, 15 Oct 2003, PC. Boyce & Jeland ak Kisai AR-141.1 (SAR);

Nanga Gaat, Rejang Wood Concession, km 65 road to Camp Gahada, 01° 42’

01.17, 113° 31? 14.8”, 12 May 2004, P.-C. Boyce et al. AR-363 (SAR); Nanga

_Gaat, Rejang Wood Concession, km 55 road to Camp Gahada, 01° 44’ 44.5”,

113° 28’ 32.3”, 13 May 2004, PC. Boyce et al. AR-385((SAR); Nanga Gaat,

Rejang Wood Concession, trail to water catchment behind main camp, 01°

53’ 00.2”, 113° 26’ 53.9”, 14 Dec 2004, P.-C. Boyce et al. AR-882 (SAR);

Nanga Gaat, Rejang Wood Concession, km 65 road to Camp Gahada, 01° 41’

59.7”, 113° 31’ 13.7”, 16 Dec 2004, P.-C. Boyce et al. AR-907 (SAR); Kapit,

Pelagus, Pelagus Rapids, Woodpecker Trail, 02° 11’ 15.1”, 113° 03’ 29.01”,

14 Mar 2005, P.-C. Boyce et al. AR-1034 (SAR); Kapit, Belaga, Belaga road,

02° 43’ 45.8”, 113° 45’ 37.1”, 12 Oct 2005, P.-C. Boyce et al. AR-1455 (SAR);

Kapit, Belaga, Belaga road, 02° 42’ 55.9”, 113° 45’ 29.3”, 12 Oct 2005, PC.

Boyce et al. AR-1457 (SAR); Kapit, Belaga, Belaga road, 02° 42’ 55.9”, 113°

45° 29.3”, 12 Oct 2005, PC. Boyce et al. AR-1461 (SAR + spirit); Kapit,

Belaga, km 10 Bakun, Bintulu-Miri road junction, 02° 50’ 51.7”, 114° OL

57.6”, 11 Oct 2005, PC. Boyce et al. AR-1481 (SAR, + spirit); Miri Division:

Mulu, Long Lama, Mulu N.P., Trail to Gunung Mulu Summit, 04° 02’ 18.7”,

114° 49° 44.2”, 7 Aug 2006, P.-C. Boyce et al. AR-1955 (SAR); Mulu, Long

Lama, Mulu N.P., Trail to Long Lansat, Sungai Licat, 04° 00’ 03.5”, 114° 48’

49.8”, 9 Aug 2006, P.-C. Boyce et al. AR-1985 (SAR); Miri, Marudi, Sungai

Silat Basin, Sungai Palutan, 02° 49.59’, 115° 00.30’, 25 Mar 2003, Lim S.P.

S.90424 (SAR). BRUNEI: Temburong District: Sungai Temburong at Kuala

Belalong, banks of Sungai Belalong. 4°32’N, 225°9’E, 24 Jun 1989, PC.

Boyce 431 (BRUN, K, L)

Homalomena striatieopetiolata P.C.Boyce & S.Y.Wong, sp. nov.

Ab allis Homalomenae borneensibus petiolis dimidium distalis eborinus cum

bho

Studies on Homalomeneae (Araceae) of Borneo I.

striae longitudinalis atrorubra vel atrochermesinus, petioli vagina brevissimo

(usque ad 3 cm longa; ca 1/10 partem longitudinis petiolo), inflorescentiis

pedunculis brevissimo brevissimo (usque ad 3 cm longa), mucro et marginae

inflorescentiis chermesinus coloratus et connectivo antherae pubescenti

differt.— Typus: Malaysia, Sarawak, Miri Division; Mulu, Long Lama, Mulu

N.P., trail to Long Lansat, Sungai Licat, 04° 00° 03.5”, 114° 48° 49.8”, 9 Aug

2007, P.-C. Boyce et al. AR-1988 (Holo, SAR+ spirit). Plates 7 & 8.

Medium to robust herbs, strongly aromatic (mango peel), evergreen,

glabrous, to ca 100 cm tall. Stem pleionanthic, erect to ascending, ca 3 cm

thick, green, internodes to ca 1 cm long. Leaves few, up to ca 8 together:

petiole terete, erect to decumbent, 50-70 cm long, petiole bases clasping,

petioles with the lower 2 green and upper 2 white with prominently striate-

raised glossy dark to cherry red ridges, longer petioles tending to spread,

and these with a somewhat weakly defined pulvinus-like articulation ca

25-35 cm long usually ca '2 way along petiole, both portions of petioles

drying dark brown, petiolar sheath to ca 3 cm long, ca '/,, of petiole length,

equal at both side, sheath initially long-persistent, eventually the whole

sheath marcescent; lamina broadly ovato-sagittate, 27-45 cm long x 22-

36 cm wide, thinly leathery, glossy green adaxially (fresh), drying dark

brown, abaxially pale green (fresh), glaucous, drying paler brown, base

cordate, posterior lobes spreading, unequal, one side round (ca 5.4-8 cm

long), shorter than subtriangular side (ca 5.7-11 cm long), lamina tip obtuse,

short-acuminate for ca 2 cm, acuminate up to ca 2 cm, apiculate up to ca 1.8

mm; midrib raised abaxially (fresh and dry), drying dark brown, adaxially

slightly channelled when fresh, flush with lamina when dry, ca 2-5 mm

wide, with 8-10 primary lateral veins on each side, diverging at 40°-90°

from the midrib, adaxially impressed (fresh), flush with lamina when dry,

abaxially slightly raised (fresh and dry), drying dark brown, curved sharply

towards the apex when near the margin, interprimary veins ca 2 width of the

primary lateral veins, alternating irregularly with primaries, arising from the

primary lateral veins near petiole insertion, but further up the lamina, arising

from midrib, posterior lobes each with 2-4 primary lateral veins; secondary

venation rather obscure, striate; tertiary venation not visible, all veins

running into a thickened intermarginal vein. Inflorescences 1-4 together,

erect, each subtended by prophyll, ca 3 cm long, marcescent, followed by

cataphyll, marcescent, peduncle to ca 3 cm long x 3 mm diam. Spathe

ca 12.8 cm long, tightly furled prior to anthesis, lower spathe inflating at

female anthesis, spathe limb loosening at female anthesis, thense, inflating

and then opening wide, lower spathe white prior to and at anthesis, spathe

limb white prior to and during anthesis, with spathe tip, mucro and spathe

24 Gard. Bull. Singapore 60 (1) 2008

Plate 7. Homalomena striatieopetiolata P.C.Boyce & S.Y.Wong: A. Overall plant with

decumbent petioles; B. Leaf lamina glossy green; C. Abaxial leaf lamina pale green with

striate-raised ridges on petiole; D. Striking striate-raised ridges on petiole; E. Synflorescence

with spathe mucro and margin stained striking cherry red, note the marcescent prophyll.

DS)

Studies on Homalomeneae (Araceae) of Borneo I.

Plate 8. Homalomena striatieopetiolata P.C.Boyce & S.Y.Wong. A. Whole spadix, spathe

removed, from alcohol collection; B. Close up of female/male zone transition; C. Detail

of female zone; D. Male zone.

margin cherry red; lower spathe narrowly ellipsoid, ca 4.6 cm long, weakly

constricted at the junction of the spathe limb, the constriction coinciding

with the lower-most fertile male flowers, spathe limb narrowly lanceolate,

ca 8.2 cm long, bluntly mucronate to ca 5 mm long. Spadix equalling the

spathe, ca 12 cm long, shape, stipitate, stipe to ca 4.4 mm long, obliquely

inserted on peduncle, obpyramidal, female zone ca 2.7 cm long x 9.5 mm

wide, ca “% length of spadix, weakly fusiform, female flowers densely

arranged, ca 1.5 mm diam. x 1.3 mm tall, squat-cylindrical, stigma exceeding

the ovary, coherent to adjacent stigma, umbonate and weakly tetrasulcate,

interpistillar staminodes clavate, on a very slender stipe, remaining pale

brown in alcohol, ca 0.5 mm wide x 1.1 mm long, slightly overtopping

the female flowers, lower most female flowers ca twice the size of fertile

females mostly associated with two or more interpistillar staminodes and

seemingly sterile, suprapistillar pistillodes in three rows and merging with

the lower most male flowers these seemingly sterile, male zone ca 8 cm

long x 7.7 mm wide, ca 7/,; length of spadix, very weakly fusiform and

26 Gard. Bull. Singapore 60 (1) 2008

minutely pubescent, distal and proximal most flowers apparently sterile,

narrowing in the lower part coinciding with the constriction of the spathe

and there intergrading with staminodes, male flowers, ca 3 mm x 1.6 mm

trapezoid comprising (3)-4 truncate stamens each overtopped by a large,

minutely pubescent, connective. Infructescence unknown.

Distribution: Sarawak, Miri Division. Known only from the type locality.

Habitat: Terrestrial under full shade on seasonally inundated alluvium and

shale mud banks mostly in deep soil of riverine forest. 32-60 m asl.

Notes: Homalomena._striatieopetiolata 1s immediately distinctive by

the distal half of the petioles white with dark to cherry red striate-raised

glossy longitudinal ridges; in this feature it is one of the most attractive

Homalomena species yet described. Plants individually carry few leaves,

normally ca 8 together, with these tending to spread. Other notable features

include the very short petiolar sheath (ca 3 cm, ca '/,, of petiole length),

leaf laminae with distinctly oblique base, one round posterior lobe and one

subtriangular, the very short, peduncle (ca 3 cm) and the spathe mucro

and margins are stained cherry red when fresh. The minutely pubescent

connective of the male flowers is noteworthy.

Etymology: The specific epithet alludes to the strikingly striate petioles.

Other specimens seen: Miri Division: Mulu, Long Lama, Mulu N.P., Trail

to Deer Cave, 04° 02’ 23.8”, 114° 48’ 54.6”, 5 Aug 2007, PC. Boyce et al.

AR-1936 (SAR); Mulu, Long Lama, Mulu N.P., Trail to Deer Cave, 04° 02’

02.0”, 114° 49’ 00.0”, 6 Aug 2007, P.-C. Boyce et al. AR-1945 (SAR).

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee and the

Forest Research Centre (Kuching), notably L.C.J. Julaihi & Lucy Chong.

Thanks are due to Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme

Brown and Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their

support and funding of fieldwork in Sarawak. The first author is grateful

for the support provided by Faculty of Resource Science and Technology,

UNIMAS. This study is funded by the Ministry of Science, Technology and

Innovation, Vot: E-Science 05-01-09-SF0006 and permitted under Sarawak

Studies on Homalomeneae (Araceae) of Borneo I. 2/

Forestry Department Research Permit No. NPW.907.4.2(II)-80 and Permit

to enter Park No. 66/2007 valid until 10" October 2008, on yearly renewal

basis.

References

Alderwerelt van Rosenburgh, C.R.W.K. van. 1922a. New or noteworthy

Malayan Araceae II. Bulletin du jardin botanique de Buitenzorg 3: 163-

229.

Alderwerelt van Rosenburgh, C.R.W.K. van. 1922b. New or noteworthy

Malayan Araceae III. Bulletin du jardin botanique de Buitenzorg 3: 320-

347.

Boyce, P.C. 1994 New species of Araceae from Brunei. Kew Bulletin 49:

793-801.

Boyce, P.C. 2000a. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the southern and western Indonesian

archipelago. Gardens’ Bulletin Singapore 52: 101-183.

Boyce, P.C. 2000b. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in the Philippines. Gardens’ Bulletin

Singapore 52: 213-256.

Boyce, P.C. 2000c. The genus Pothos (Araceae: Pothoideae: Potheae) of

Thailand and Indochina. Blumea 45: 147-204.

Boyce, P.C. 200la. The genus Rhaphidophora Hassk. (Araceae-

Monsteroideae-Monstereae) in Borneo. Gardens’ Bulletin Singapore

53: 19-74.

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

Monsteroideae-Monstereae) in New Guinea, Australia and the tropical

western Pacific. Gardens’ Bulletin Singapore 53: 75-183.

Boyce, P.C. and A. Hay. 2001. A taxonomic revision of Araceae tribe

Potheae (Pothos, Pothoidium and Pedicellarum) for Malesia, Australia

and the tropical Western Pacific. Telopea 9: 449-571.

28 Gard. Bull. Singapore 60 (1) 2008

Christensen, H. 2002. Ethnobotany of the Iban & the Kelabit. Forest

Department Sarawak, Malaysia, NEPCon Denmark & University of

Aarhus, Denmark. 384 pp.

Engler, A. and K. Krause. 1912. Homalomena, pp. 25—81. In: A. Engler

(ed.), Das Pflanzenreich, IV (Heft 55.) (Araceae-Philodendroideae-

Philodendreae, [I]: Allgemeiner Teil, Homalomeninae und

Schismatoglottidinae). Leipzig.

Furtado, C.X. 1939. Notes on some Indo-Malaysian Homalomena species.

Gardens’ Bulletin Straits Settlement 10: 183-238.

Furtado, C.X. 1940 (published 1941). The variability and distribution of

the Indo-Malaysian species of Homalomena. Proceedings of 6th Pacific

Science Congress (California, 1939) 4: 577-578.

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

Blumea 35: 499-545

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

and Sulawesi. Gardens s Bulletin Singapore 50: 221-334.

Hay, A. 1999. A revision of Homalomena (Araceae-Homalomeneae) in

New Guinea, the Bismarck Archipelago and Solomon Islands. B/umea

44: 41-71.

Hay, A. and C. Herscovitch. 2002. Two Remarkable New West Malesian

Homalomena (Araceae) Species. Gardens’ Bulletin Singapore 54: 171-

178.

Hay, A. and Yuzammi. 2000. Schismatoglottideae in Malesia I —

Schismatoglottis. Telopea 9: 1-178.

Hotta, M. 1967. Notes on Bornean plants, Il. Acta Phytotaxonomica

Geobotanica 22: 153-162.

Hotta, M. 1985. New species of the genus Homalomena (Araceae) from

Sumatra with a short note on the genus Furtadoa. Gardens’ Bulletin

Singapore 38: 43-54.

Studies on Homalomeneae (Araceae) of Borneo I. 29

Hotta, M. 1986. Species list and cited specimens of the genus Homalomena

(Araceae) in Malesia, pp. 73-120. In: Hotta, M. (ed.), Diversity and

Dynamics of Plant Life in Sumatra. Sumatra Nature Study (Botany),

Kyoto University.

Hotta, M. 1993. Homalomena monandra, a new species of aroid from West

Sumatra. Acta Phytotaxonomica Geobotanica 44(2): 93-96.

Merrill, E.D. 1921. Araceae, pp. 86-109. In: Bibliographic Enumeration of

Bornean P1. Journal of Straits Branch Royal Asiatic Society (Special

edition, September 1921).

Miquel, F.A.W. 1856a. Flora van Nederlandsch Indié. vol. 3. Van der Post,

Amsterdam.

Miguel, F.A.W. 1856b. Orontiaceae, Aroideae, pp. 148-153. In: W.H. de

Vriese, Plantae Indiae Batiae Orientalis. Brill, Leiden.

Ridley, H.N. 1905. The Aroids of Borneo. Journal of Straits Branch Royal

Asiatic Society 41: 169-188.

Schott, H.W. 1860. Prodromus Systematis Aroidearum. Typis congregationis

mechitharisticae, Vienna. 602 pp.

Sulaiman, B. and P.C. Boyce. 2005. A remarkable new species of

Homalomena (Araceae: Homalomeneae) from Peninsular Malaysia.

Gardens’ Bulletin Singapore 57: 7-11.

(oe)

jam

Gardens’ Bulletin Singapore 60 (1): 31-36. 2008

Hapaline celatrix (Araceae: Caladieae)

— A New Record for Sarawak, Malaysian Borneo

PETER C. BOYCE ' AND WONG SIN YENG ”

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

No. 18, Jalan Tunku Abdul Rahman,

93100 Kuching, Sarawak, Malaysia

>Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan, Sarawak, Malaysia.

Abstract

Hapaline celatrix P.C.Boyce, a species known from only two collections

and hitherto considered endemic to Brunei, has recently been collected

in Gunung Mulu NP. An expanded description for the species, additional

ecological notes and the first ever published field photographs are

presented.

Introduction

Studies on the predominantly Indo-Chinese genus Hapaline Schott in recent

years have resulted in a published generic revision (Boyce, 1996) including

the description of a second Bornean endemic, Hapaline celatrix P.C.Boyce,

based on two collections from the Setap Shales of eastern Brunei, and the

rediscovery of Sarawak-endemic, H. appendiculata Ridl., after a period of

nearly 30 years, and then only the fourth collection (Boyce et al., 2005).

Yet still more recent fieldwork by the authors in Gunung Mulu N.P.

has located extensive populations of H. ce/atrix along the Sungai Licat, a

stream draining into the Sungai Tutoh along the SW flank of the park. These

collections represent the first ever record for Sarawak and are furthermore a

new species record for Malaysia. Additionally the new discovery expands

the information concerning species variability, not least variation in leaf

shape and leaf markings.

Hapaline celatrix P.C.Boyce

Kew Bull. 51: 70 (1996). Type: Boyce 417 (holotype K!; isotypes: BRUN!,

L!, K! + Kew spirit coll. no. 57283!, SING!). Plates 1 & 2.

Gard. Bull. Singapore 60 (1) 2008

Plate 1. Hapaline celatrix P.C.Boyce. A. Colony at Sg. Licat; B & C. Inflorescences at

male anthesis. Note the short appendix in B & C compared with that of H. appendiculata;

D. Hapaline appendiculata; E. Variation in leaf markings of H. celatrix at Sg. Licat.

Hapaline celatrix (Araceae: Caladieae)

Plate 2. Hapaline celatrix P.-C.Boyce. A & B. Range of variation in leaf shape and markings

present at Sg. Licat; C. Adaxial venation.

34 Gard. Bull. Singapore 60 (1) 2008

[Hapaline celator P.C. Boyce in Hay ef al. in Blumea suppl. 8: 68 (1995),

nom. nud. |

Diminutive, weakly tuberous or/or moderately rhizomatous evergreen

perennial herb up to 15 cm tall. Stem with globose tuber, 7-20 x 7-13 mm;

stolons terete, 1-16 cm x 2-4 mm, enclosed by several sub-fleshy, later

papery, later still decomposing cataphylls. Roots rather few, ca 0.2 mm in

diam. Prophyll of leaf linear-triangular, up to 10 cm x 5 mm, rather weakly

2-keeled, acute to briefly apiculate; cataphyll linear-triangular, up to 12 cm

x 9 mm, attenuate to rather blunt; petiole 8-17 cm x 0.5-1.5 mm, longer

petioles with up to half of the length buried; leaf blade hastato-sagitatte to

ovato-sagittate to cordiform, 5.5-16 x 2.7-10.5 cm, thinly to rather thickly

coriaceous or sub-succulent, dark green to variously and variably spotted

and blotched with pale green, yellow green or grey, margins smooth to

minutely crispulate, apex acuminate, posterior lobes rounded, divergent

to sub-parallel; mid-rib prominently raised abaxially, impressed adaxially;

_ primary lateral venation arising at ca 75° to the mid rib, prominent abaxially,

slightly impressed adaxially, running to a prominent (abaxially), impressed

(adaxially), brochidodromous intramarginal collecting vein; interprimary

veins much less prominent, running into intramarginal collecting vein;

secondary venation reticulate, moderately raised abaxially, slightly

impressed adaxially; tertiary venation reticulate, weakly visible abaxially,

invisible adaxially in fresh material, barely visible in dried specimens,

reticulate. Inflorescence solitary to several together; peduncle 3-9 cm x 0.5-

2 mm, longest peduncles with much of the length buried; spathe 1.5-2 cm

long; spathe limb elliptic, 1.5-2 cm x 6-10 mm, apex acute, base decurrent

into lower spathe; lower spathe margins clasping, 1.5-3 cm x 2 mm; spadix

lcm x 2.5-3 mm, free portion narrowly conic, ca 8 mm long, tapering

apically in to a stout conical appendix composed of fused synandrodes.

Flowers synandria, irregularly elongate in plane view, 2-7 x 0.75-1.5 mm;

ovaries ellipsoid, 2-2.5 x 0.5-1 mm, 2-4 in a single row; stigma slightly

prominent, ca 0.2 mm diam; style absent. Infructescence on declinate to

reflexed peduncle, partially enclosed by the persistent lower spathe, | cm x

3.5 mm, 2-3-berried; berries more or less globular, ripening pale green, ca

2.4 mm in diam, stigmatic remains not prominent. Seed ellipsoid, ca 1.5 x

3 mm, glossy pale brown with a conspicuous white oily rhape.

Other specimens seen: SARAWAK: Miri Division, Mulu, Long Lama,

Mulu N.P., Sg. Licat, trail to Long Lansat, 04° 00’ 03.5”; 114° 48’ 49.8”, 9

Aug 2007, PC. Boyce, S.Y.Wong et al., AR-1972 (SAR, SING). BRUNEI.

Temburong: Sg. Temburong at Kuala Belalong, 4°32’ N, 115°9’ E, 20 June

Hapaline celatrix (Araceae: Caladieae)

1989, P.C.Boyce 358 (BRUN!, K!); Sg. Temburong at Kuala Belalong,

banks of Sg. Belalong. 4°32’, 115°10° E, 24 June 1989, PC Boyce 417

(BRUN, K+K spirit 57283, L, SING).

Distribution: Sarwawak (Miri Division); Brunei (Temburong District).

Ecology: Disturbed lowland mixed dipterocarp forest on river banks in

deep leaf litter overlying soils derived from Setap shales. 20-32 m asl.

Notes: Hapaline celatrix differs from all other species of Hapaline by the

combination of evergreen habit, small inflorescences borne beneath the

leaves and a short sterile spadix appendix. It is most similar to H. brownii

Hook.f. in Peninsular Malaysia and S Thailand, and H. appendiculata. It

may be distinguished from H. brownii by its evergreen habit and overall

lesser stature and the smaller inflorescence carried beneath the leaves.

Additionally, H. brownii is restricted to Karst limestone. Hapaline celatrix

differs from Hapaline appendiculata by its evergreen habit, more coriaceous

leaves and short sterile appendix.

The habitat of H. celatrix is briefly seasonally-dry riverine forest

on shale. All known localities of H. ce/atrix are more exposed (higher light

levels) and less humid for at least part of each day than the known habitats

of H. appendiculata.

The specific epithet comes from the Latin ce/ator, ‘the concealer’,

in allusion to the manner in which the foliage obscures the inflorescences,

a character otherwise unknown in the genus.

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee and the

Forest Research Centre (Kuching), notably L.C.J. Julaihi & Lucy Chong.

Thanks are due to Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme

Brown and Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for their

support and funding of fieldwork in Sarawak. This study is funded by

the Ministry of Higher Education, Malaysia under fundamental research

grant scheme No. FRGS/01(04)/609/2006(42) and permitted by Sarawak

Forestry department Research Permit No. NPW.907.4.2(II)-80 and Permit

to enter Park No. 66/2007 valid until 10th October 2008, on a yearly

renewal basis.

36 Gard. Bull. Singapore 60 (1) 2008

References

Boyce, P.C. 1996. The genus Hapaline (Araceae: Aroideae: Caladieae).

Kew Bulletin 51: 63-82.

Boyce, P.C., Jeland ak Kisai and Jipom ak Tisai 2005. Hapaline

appendiculata (Araceae: Caladieae) Rediscovered. Gardens’ Bulletin

Singapore 57: 13-18.

Gardens’ Bulletin Singapore 60 (1): 37-43. 2008 37

New Records for the Flora of Peninsular Malaysia,

Family Orchidaceae 1. Appendicula floribunda,

Bulbophyllum elevatopunctatum, Cymbidium sigmoideum

and Dendrochilum bandaharaense

M. JUTTA, P.T. ONG & S.N. PHOON

Forest Research Institute Malaysia, 52109 Kepong, Selangor, Malaysia

Abstract

Four orchids, Appendicula floribunda (Schltr.) Schltr., Bulbophyllum

elevatopunctatum J.J.Sm., Cymbidium sigmoideum J.J. Sm., and

Dendrochilum bandaharaense J.J. Wood & J.B. Comber are new records

for the Orchidaceae of Peninsular Malaysia.

Introduction

Expanding infrastructure and development projects have made many areas of

forest in Peninsular Malaysia accessible which in recent years has led to the

discovery of new records, especially of taxa that were previously recorded

from surrounding regions (Jutta, 2004; Jutta & Faridah, 2005; Wong et ail.,

2002). Four such species are reported here, namely Appendicula floribunda

(Schltr.) Schltr. from Terengganu, Bu/bophyllum elevatopunctatum J.J.Sm.

from Selangor and Johor, and Cymbidium sigmoideum J.J. Sm. and

Dendrochilum bandaharaense J.J. Wood & J.B. Comber, from Pahang.

The presence of these species in Peninsular Malaysia is no surprise, given

their confirmed distribution in neighbouring countries.

1. Appendicula floribunda (Schltr.) Schltr.

Repert. Spec. Nov. Regni. Veg. Beth. 1 (1912) 355; Op. Bot. 89 (1986) 138;

Comber, Orchids of Sumatra (2001) 532. Basionym: Podochilus floribundus

Schltr. Mem. Herb. Boiss. 21 (1900) 58. Fig. 1A-F, Plate 1A, B.

Specimens examined: Terengganu.Taman Negara, Tasik Kenyir, on low-

canopy trees overhanging the banks of a shallow stream M. Jutta FRI 59560,

3 Aug 2007 (KEP); M. Jutta KBG 20070621 (living collection FRIM).

Gard. Bull. Singapore 60 (1) 2008

Imm

| E

ee P|

H I J /

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Me a4

i +

y %,

2mm

lem

AS eee

op)

2mm

Figure 1. Appendicula floribunda. A. Dorsal sepal; B. Lateral sepal; C. Petal; D. Lip; E.

Column; F. Floral bract. Bulbophyllum elevatopunctatum. G. Dorsal sepal; H. Petal; I.

Lateral sepal; J. Lip (adaxial); K. Lip (abaxial). L. Column. (A-F from FRI 595560, G-L

from FRI 58820).

New Records for the Flora of Peninsular Malaysia

Plate 1. Appendicula floribunda. A. Inflorescence; B. Close-up of a single flower.

Bulbophyllum elevatopunctatum. C. Flower heavily visited by fruit flies; D. Petal and

lateral sepal removed to show tooth on column. (Photographs by P.T.Ong)

40 Gard. Bull. Singapore 60 (1) 2008

The plant, whose collection was made possible by the building of the

Kenyir hydro-electric dam, was identified using Seidenfaden (1986). From

other Peninsular Malaysian Appendicula (Seidenfaden & Wood, 1992), the

species is easily distinguished by the prominent row of long, retrorse hairs

fringing the base of the three-pointed callus (Fig. 1D). Our material has

white flowers, turning yellow with age and with some purple markings on

the column. Comber (2001) mentioned ‘some purple’ on the lip but this

was not observed by us.

2. Bulbophyllum elevatopunctatum J.J. Sm.

Bull. Jard. Bot. Buit 3 (1920) 99; Comber, Orchids of Sumatra (2001) 771.

Fig. 1G-M, Plate 1C, D.

Specimens examined: Selangor. On an old rubber tree trunk Ong P-T. FRI

58820 2006 (KEP); Ong PT. KBG 20070191 (living collection, FRIM).

_ Johor. Ong P-T: FRI 58826 14 Dec 2007 (KEP).

Bulbophyllum elevatopunctatum, previously known only from Sumatra, is

most similar to B. membranifolium Hook.f. among Peninsular Malaysian

species within sect. Sestochilus, but differs in having relatively thick-

textured leaves, sepals and petals, leaves with shorter stalk, base of lip

with strips of papillae, fimbriate side lobe margins, and a dark red flower.

The species is, however, most closely related to the Bornean endemic, B.

vinaceum Ames & C.Schweinf., separated only by two strips of papillae at

the base of the lip and a distinct curved tooth along each lower margin of

the column (Fig. 1L; Plate 1D), which are absent in B. vinaceum. Presence

or absence of teeth along the lower margins of the column is considered a

specific character within sect. Sestochilus (Vermeulen, 1991). The flowers

examined here differed from Comber’s description in being consistently

larger in size. Flowers, as soon as the buds opened, were heavily visited by

fruit flies of the genus Bactrocera Macquatt.

3. Cymbidium sigmoideum J.J.Sm.

Bull. Dept. l’ Agric. Indes Néerl. (1907) 52; Comber, Orchids of Java (1990)

381, Orchids of Sumatra (2001) 235. Plate 2A, B.

Specimen examined: Pahang. Cameron Highlands, Phoon & Ong FRI

60439 (KEP).

New Records for the Flora of Peninsular Malaysia 4]

Plate 2. Cymbidium sigmoideum. A. Plant habit; B. flower. (Photographs by P.T.Ong)

The plant was identified using Comber (1990, 2001). Among Peninsular

Malaysian Cymbidium it is most closely related to C. roseum J.J.Sm. Both

belong to sub-genus Cyperorchis that is distinguished by the base of the

lip being fused to the column. However, it is clearly set apart from C.

roseum by the midlobe of the lip that is narrowly oblong, tongue-like and

strongly recurved, with an acute tip, also a column that is longer than the

lip. Comber (2001) mentioned a callus at the base of the midlobe but this

was not observed here. We also observed that anthesis commenced with

flowers at the apex and continued towards the base of the inflorescence.

4. Dendrochilum bandaharaense J.J. Wood & J.B. Comber

Lindleyana 10 (1995) 57; Comber, Orchids of Sumatra (2001) 398.

Specimen examined: Pahang. Cameron Highlands, Ong PT. et al. FRI

57305, 24 Oct 2007 (KEP).

The plant was identified using Comber (2001). Dendrochilum bandaharaense

J.J. Wood & J.B. Comber was until now regarded as endemic in Gunung

Bandahara in the Gunung Leuser Nature Reserve, Aceh Province of

42 Gard. Bull. Singapore 60 (1) 2008

Sumatra. Among the Peninsular Malaysian species, it is most similar to D.

odoratum and D. simile that have entire margins (a character not mentioned

in Seidenfaden & Wood, 1992) and from which it differs in having serrate

margins on the hypochile of the lip. In addition, it has pale salmon-creamy

flowers and the apical wing has more pronounced teeth.

Acknowledgements

The authors are grateful to Kueh Hock Leng for assistance in the field,

Abdul Aziz bin Mohd. Top, Johor National Parks Corporation, for

access to their ex situ collection and to Tai Ngong Chiang from Cameron

Highlands, Pahang. This work was in part made possible through the Flora

of Peninsular Malaysia Project (Project No. 01-04-01-0000 Khas) funded

by the Ministry of Science, Technology and Innovation (MOSTI). Jaap J.

Vermeulen provided valuable advice and reference materials while Ruth

. Kiew assisted with the manuscript.

References

Comber, J.B. 1990. Orchids of Java. Royal Botanic Gardens, Kew,

England.

Comber, J.B. 2001. Orchids of Sumatra. Natural History Publications, Kota

Kinabalu, Malaysia.

Jutta, M. 2004. Phaius indigoferus Hassk.: A new record for Malaysia.

Folia Malaysiana 5, 1: 17-20.

Jutta, M. & Q.Z. Faridah. 2005. Malaxis inexpectata and Habenaria

paradiseoides (Orchidaceae), new records for Peninsular Malaysia.

Gardens Bulletin Singapore 57: 263-267.

Seidenfaden, G. 1986. Orchid genera in Thailand 13. Thirty three

epidendroid genera. Opera Botanica 89: 1-216.

Seidenfaden, G. & J.J. Wood. 1992. The Orchids of Peninsular Malaysia

and Singapore. Olsen & Olsen, Denmark.

eS)

New Records for the Flora of Peninsular Malaysia

Vermeulen, J.J. 1991. Orchids of Borneo Vol. 2. Natural History Publications,

Kota Kinabalu, Malaysia.

Wong, K.M., M. Sugumaran, J. Jumian & Zulkapli Ibrahim. 2002.

New and interesting records for Peninsular Malaysia: Bulbophyllum

lemniscatoides and Mischobulbum crassum (Orchidaceae). Malaysian

Nature Journal 55: 183-186.

Gardens’ Bulletin Singapore 60 (1): 45-54. 2008 A5

Studies in the Peristylus tentaculatus-complex (Orchidaceae)

in Thailand

H. KURZWEIL

The Herbarium, Singapore Botanic Gardens

1 Cluny Road, Singapore 259569

Abstract

A large number of Thai specimens belonging to the Peristylus tentaculatus-

complex were examined in the present study. No characters were found

to distinguish between the three previously recognised species, Peristylus

tentaculatus (Lindl.) J.J.Sm., P. tipuliferus (C.S.P.Parish & Rchb.f.) Mukerjee

and P. garrettii (Rolfe ex Downie) J.J. Wood & Ormerod, which are therefore

considered as conspecific.

Introduction

During the preparation of the treatment of the orchid genus Peristylus for

the Flora of Thailand a large number of specimens of the group comprising

the three very similar species, Peristylus tentaculatus (Lindl.) J.J.Sm., P.

tipuliferus (C.S.P.Parish & Rchb.f.) Mukerjee and P. garrettii (Rolfe ex

Downie) J.J.Wood & Ormerod, were examined with the aim of finding

distinguishing characters between them. Special attention has been paid

to the characters which have been identified as taxonomically informative

before (Seidenfaden, 1977). It was concluded that many specimens in this

group can indeed clearly be referred to one of the three species, but that they

are interconnected by a large number of intermediate forms. Although only

material from inside Thailand was examined here, the results of the present

study are nevertheless significant and indicate that the previously recognised

species are not sufficiently different from each other to be recognised as

separate species. Therefore they are here considered as belonging to a single

variable species P. tentaculatus.

The inclusion of the concept of Peristylus garrettii in P. tentaculatus

is in agreement with the original suggestion of Seidenfaden (1977, p. 41)

(as Habenaria garrettii Rolfe ex Downie). Peristylus tipuliferus and P.

tentaculatus have in the past been maintained as separate entities. A detailed

comparison of the two was presented by Seidenfaden (1977, p. 44), where it

46 Gard. Bull. Singapore 60 (1) 2008

was pointed out that the species differ mostly in their vegetative habit and in

the thickness of their lip spur. However, Seidenfaden himself admitted that

some of the Thai specimens of these two taxa are difficult to place, stating

that further research on them is needed.

In the present study measurements of 38 specimens from 19 different

collections were taken, and additional information was obtained from the

literature (particularly from Seidenfaden & Smitinand, 1959; Seidenfaden,

1977).

The total plant size, leaf number, leaf size, leaf insertion, as well as

distance from lowermost to uppermost leaf, are indicated in Table 1, and

various spur measurements are shown in Table 2. Previous identifications

are also given in the two tables where available, but it needs to be pointed

out that these need not necessarily be correct. Of particular interest are the

features in different plants of the same collection (either deposited in different

herbaria or in the same), indicating that the critical characters are sometimes

variable within the same population.

. Total plant height: An even increase of the stem height from 14 to 57

cm was found which is probably largely caused by environmental and age

factors. Specimens that have previously been referred to Peristylus tipuliferus

are mostly taller than specimens that have been identified as P. tentaculatus as

already pointed out by Seidenfaden (1977, p. 44), but no obvious taxonomic

grouping based on the plant size can be made as the distribution of this

character shows an entirely continuous pattern.

Leaf size: The size of the blade of the largest leaf varies extensively,

but also in this character the variation 1s continuous and does not reveal any

obvious grouping (Fig. 1). Leaves can be as short as 3 cm or as long as 16.2

cm, with their width ranging from 0.7 to 3 cm. It is, however, evident that

specimens previously referred to P. tipuliferus often have larger leaves than

specimens of P. tentaculatus sensu stricto (= s.s.), which was used as one of

the distinguishing features in the past (Seidenfaden 1977, p. 44).

Leaf arrangement: Also the arrangement of the leaves has previously

been regarded as critical in delimiting the two species Peristylus tipuliferus

and P. tentaculatus s.s. (Seidenfaden 1977, p. 44; see also references given

there). Typical specimens of P. tipuliferus have the leaves scattered or

clustered on the stem 10-15 cm above the ground while typical specimens of

P. tentaculatus s.s. have their leaves clustered near the soil surface. A large

number of intermediate forms between these two character states were found

in the present study, as the insertion of the lowest leaf ranges continuously

from 0.5 to well over 6 cm. A whole range of variation was also found in

the distance of the lowermost and uppermost leaf (0.2 to over 6 cm). Both

observations suggest that the two species cannot be separated on this basis.

Peristylus tentaculatus in Thailand 47

Table 1. Vegetative features of various specimens of Peristylus tentaculatus, indicating

total plant size, leaf number, leaf blade size, height of insertion of the lowermost leaf,

height of insertion of the uppermost leaf and distance from uppermost to lowermost leaf.

All measurements in centimetres. In collections which have been identified before their

previous name is given. [-] data not available.

| Distance

‘ of

Previously Total Number | Blade Insertion | Insertion lowermost

Collections referred to plant of largest lowermost | uppermost and

size leaves leaf | leaf | leaf uppermost

leaf

| |

Chanthanaoraphin s.n. | tentaculatus | 24 3 6x | | 2S | 4 1.5

(BCU)

T . ~

Chanthanaoraphin s.n. | tentaculatus | 51 5 [sxis 2:5 | 8.5 6

(BCU)

| | t | |

Garrett 61 (BKF) pleas ulatus | 26 13 PS eexelen 1.1 2.2 1.1

5 ~ + + = +

Garrett 61 (K) tentaculatus | 33 3 5.2x 1.4 l 2 2

7 i i i + +

Garrett 61 (K) | tentaculatus | 15 13 ye 2 2 1.2

1 -

Larsen & al 46277 | tentaculatus | 25 3 4x1 | es 0.5

(AAU)

4 ] } + +

Larsen & al 46277 tentaculatus | 30 3 16x 1.2 13 4 |

(AAU) | |

| iE | |

Larsen & al 46277 —_| tentaculatus | 14 2 14x0.7 |1.7 2.2 0.5

-TAI }

(SING) |

+ ; is 4 :

Larsen & al 46277 | tentaculatus | 26 13 15x] 2 3 l

(SING)

Larsen & al 46684 tentaculatus | 27 2 broken off | 2.8 $7 0.4

(AAU)

Maxwell 00-386 (L) | tentaculatus | 35 | 3 16.3x1.7 {1.5 2.2 0.7

+ + + + +

Maxwell 00-386 (L) | tentaculatus | 57 14 10.8 x 1.6 | 5.4 7.5 2.1

4 + + + +

Maxwell 74-776 (L) |? 147 13 18x2 3 54 24

T 1 + +

Maxwell 74-776 9 30 | 4 16x | 0.8 2.5 1.7

(AAU) |

t | | |

Maxwell 74-776 |? }40.5 | 4 Wea 2 3 1.8

(AAU) | |

| ) | }

Murata 15894 tentaculatus | 20.8 3 44x] 0.8 2 0.4

(Seidenf. 1977)

4 +

Parish 292 (Myanmar) | tipuliferus | - 4 16.2 x 2.2 |- = =

(Seidenf. 1977)

2 . | Hl | i > > 42 2

Pumicong 392 (QBG, | tentaculatus | 26 3 xe 3 4.3 1.3

SING)

Gard. Bull. Singapore 60 (1) 2008

Smitinand & Sleumer | tentaculatus |31 3 Als IL

8337 (BKF)

Smitinand & Sleumer | fentaculatus |23 3 55) eli 0.3

8337 (L)

Smitinand & Sleumer | fentaculatus | 38 3 8.5x 1.8 l

8337 (L)

Sorensen & al. 4728 | tipuliferus iixales 1.7

(BKF)

Sorensen & al. 4728 tipuliferus NS 2:3} 4 6.5 2,5)

(C)

Sorensen & al. 4857 | tipuliferus Orcs) |! 8 4

(C)

Sorensen & al. 4857 | tipuliferus 5x |] 1.8 DiS 0.7

(BKF)

Sorensen & al. 4857 | tipuliferus | 26 4 6.5.x 2.4

(BKF)

Sorensen & al. 4857 | tipuliferus | 35 3 WD

(BKF)

Sorensen & al. 4871 | tentaculatus |22 2 Sexe

(C)

Sorensen & al. 5109 | tipuliferus | 41 3 8x 2.8

(BKF)

Sorensen & al. 5109 | tipuliferus | 40 4 8.5.x 3

(BKF)

Sorensen & al. 5109 | tipuliferus |25 3 broken off

(C)

Sorensen & al. 5109 | tipuliferus | 45 4 W).3) 38 2S)

(C)

Sorensen & al. 5109 | tipuliferus | 33 3 MES xe229

(C)

Thaithong 1132 (BCU)| tipuliferus | 40 3 7.3.x 1.4

Thaithong 1262 (BCU)) tipuliferus | 44 5 Dexa

Thaithong 1495 (BCU)| tentaculatus | 31 - 4x 1.5

Watthana & Wongnak |

2153 (QBG, SING) ? 29 3 Oxo ley

Peristylus tentaculatus in Thailand 49

Table 2. Spur characters in various flowers of Peristylus tentaculatus. All measurements

in millimetres. In collections which have been identified before their previous name is

given. [-] data not available.

: : N

SIDE VIEW satis

Previously ]

Collections erred ie length | width length width depth

spur spur stalk | stalk | apical

| | incision

Cumberlege 610 (K spirit) garrettii 2.8 1.4 0.5 | 0.4 0.05

Cumberlege 610 (K spirit) | garrettii 3 12 | 0.7 | 0.6 10.1

Garrett 61 (K) tentaculatus | 3.9 2 0.8 0 -

Garrett 61 (K) | tentaculatus 3.4 1.5 I 0.5 -

Garrett 61 (K) tentaculatus | 2.8 1.3 105 0 0.3

Garrett 61 (K) ~ | tentaculats 3 1.4 10.6 0.4 [9.7

Larsen & al 46277 (AAU) tentaculatus | 3.4 2 0.5 | 0.8 (2

Larsen & al 46277 (SING) _| tentaculatus_ [2.9 1.9 04 [o7 {os

Larsen & al 46684 (AAU) tentaculatus | 3 1 .8--2 0.8 0.6 [01

Maxwell 00-386 (L) tentaculatus | 3.7 1.8 | 0.8 | 0.05

Maxwell 74-776 (L) 1? [33 2 [08 07. (03

Maxwell 74-776 (L) ? Sri | ON 05 08 (03

Maxwell 74-776 (L) ? 3.2 2.7 0.5 0.7 0.4

Maxwell 74-776 (AAU) ? 3 1.8 0.6 0.7 0.2

Maxwell 74-776 (AAU) |? 19 16 0.8 Ge (03

Pumicong 392 (SING) | tentac ulatus | 2.6 | 2.1 | 0.4 0.6 [0.6

Pumicong 392 (SING) | tentaculatus | 2.7 1.9 | 0.5 0.7 0.6

Pumicong 392 (SING) | tentaculatus | 2.3 1.5 {0.7 0.4 | 0.2

Seidenfaden & Smitinand 2676 garrettii 3.75 1.75 0.75 0.75 | 0

(Seidenf. 1977)

Seidenfaden & Smitinand 2676 | tipuliferus | 3.2 ; 10.8 ie lo2

(Seidenf. 1977)

Seidenfaden & Smitinand 3036 | tentaculatus | 3.3 ca.1.6 | 0.5 | ca. 0.5 lc 1. 0.16

(Seidenf. 1977) |

Smitinand & Sleumer 8337 (L) | fentac ulatus |4 1.8 | 0.4 | 0.6 [o 3

Smitinand & Sleumer 8337 (L) tentaculatus 3.6 1.9 0.5 | 0.8 10.2

Sorensen & al. 4728 (C) tipuliferus 3.7 | 2.2 [0 5 | 0.8 0.3

Sorensen & al. 4857 (C) | tipuliferus 3.3 2 10.8 [0.7 {0.3

Sorensen & al. 4871 (AAU)) tentaculatus | 3 2 | | 0.8 Ii

Sorensen & al. 5109 (C) tipuliferus_| 3 [2.2 | 0.6 | 0.6 [0

Sorensen & al. 5109 (C) tipuliferus | 3.5 2.2 0 0.7 0.1

Sorensen & al. 5109 (C) tipuliferus 3 1.7 0.7 | 0.6 103

Watthana & Wongnak 2153 (SING) | ? [4 Dias abs [ 08 [03

Watthana & Wongnak 2153 (SING) | ? 3.8 | 2.7 | 0.5 | 0.8 5

Watthana & Wongnak 2153 (SING) | ? 3.9 | 3 0.5 [08 [0.6

50 Gard. Bull. Singapore 60 (1) 2008

3.5

| a

la c= rT

| a

2 1.5 a . mA

on A

| 4

A a

1 a BS eBe 8 xX x

0.5 X indet

@ fentaculatus

A tipuliferus

2 4 6 8 10 12 14 16

length

Figure 1. Diagram showing the relationship of leaf length and width in two of the taxa

previously recognised in the Peristylus tentaculatus complex, namely P. tentaculatus

and P. tipuliferus, and in specimens previously not named (‘indet’). All measurements in

centimetres.

Floral characters: Also an examination of the flowers of the

Peristylus tentaculatus-complex did not reveal any characters which would

allow a distinction of separate taxa. This is largely in agreement with the

view of Seidenfaden (1977) who was unable to find any differences between

the species in their flower structure except in their spur width.

A feature of particular importance is the shape of the lip spur (Table

2). It ranges from 2.3 to 4 mm in total length and has generally a short stalk

measuring 0.3-1 x 0.4-0.8 mm. A wide range of variation was found in the total

width of the spur when seen from the side, ranging from 1.2 to 2.2(-3) mm;

there is also slight variation within the same collection. Both observations

suggest that also the spur width cannot be used to recognise taxa within the

complex. This is in contrast to Seidenfaden’s (1977, p. 44) observation who

found that the spur diameter 1s critical, with P. tentaculatus s.s. having spur

diameters of about 2 mm, as opposed to 1-1.5 mm in P. tipuliferus.

In a short taxonomic note Peristylus garrettii was separated from P.

tentaculatus on account of its bifid spur apex (Ormerod, 2003: p. 141). The

Peristylus tentaculatus in Thailand

basionym, Habenaria garrettii, had been reduced to synonymy by Seidenfaden

(1977, p. 41), but Ormerod resurrected the species based on the examination

of one Chinese and several Thai and Burmese specimens. However, in the

Thai material examined here a wide range of variation from clearly bifid

spurs, slightly bifid spurs to spurs with entire apex, could be observed, with

the apical spur incision ranging from none to 0.6(-0.7) mm (Table 2). The

species is therefore considered as conspecific with P. tentaculatus. Also the

observation that clearly bifid, slightly bifid and entire spurs are occasionally

also found in other species of the genus Peristylus (sometimes even within the

same inflorescence) raises doubt about its usefulness in the P. tentaculatus-

complex.

Peristylus tentaculatus (Lindl.) J.J.Sm.

Orch. Java (1905) 35; Seidenf., Dansk Bot. Ark. 31 (1977) 41. — Glossula

tentaculata Lindl., Bot. Reg. 10 (1824) t. 862. — Habenaria tentaculata

(Lindl.) Rchb.f., Otia Bot. Hamburg. (1878) 34. Type from S China.

Peristylus chloranthus auct. non Lindl.: Seidenfaden & Smitinand, Orchids

Thail. 1 (1959) 47, p.p.

Habenaria garrettii Rolfe ex Downie, Bull. Misc. Inform. Kew 1925 (1925)

418. — Peristylus garrettii (Rolfe ex Downie) J.J. Wood & Ormerod, Taiwania

48 (2003) 141, syn. nov. — Typus: Thailand, Chiang Mai Province, Doi

Suthep, Kerr /18 (holo, K).

Peristylus tipuliferus (C.S.P.Parish & Rchb.f.) Mukerjee, Notes Roy. Bot.

Gard. Edinburgh 21 (1953) 153; Seidenf., Dansk Bot. Ark. 31 (1977) 41.

— Habenaria tipulifera C.S.P.Parish & Rchb.f., Trans. Linn. Soc. London 30

(1874) 139, syn. nov. Type from Myanmar. Figs. 2 & 3.

Terrestrial herbs, entirely glabrous, (15-)25-70 cm tall. Basal

sheaths 2-3, tubular, enveloping the basal part of the stem up to 3 cm high;

uppermost sometimes with a sheath to 1.5 x 1.5 cm. Leaves 2-5, basal,

clustered or scattered in the lower half of the stem, lanceolate-oblong, acute,

mucronate, 5.2-17 x 1.2-2.9 cm, margins entire or papillose. Sterile bracts

(1-)2-6, suberect or spreading, oblong-lanceolate, acute or acuminate, 0.9-

2.3(-3) * 0.2-0.35(-0.5) cm, sheathing or not, margins entire or papillose.

Inflorescence lax or semi-dense, 7- to many-flowered; rachis (5-)8-24(-30)

cm long; bracts broadly ovate-lanceolate, acuminate, 4.5-10 = 1.5-3 mm,

margins entire. Flowers green or yellow-green, also reported as whitish.

Sepals obtuse, l-veined; median sepal erect, broadly elliptic-oblong,

3.2-5 x 1.5-2.2 mm; lateral sepals oblong-elliptic, reflexed, sometimes partly

rolled-in, 3.2-5.2 x 1-1.6 mm. Petals erect, forming a hood with the median

Gard. Bull. Singapore 60 (1) 2008

Figure 2. Peristylus tentaculatus in its habitat (Pumicong 392).

sepal, 1-veined, elliptic-lorate, subacute or obtuse, 3.5-5 =< 1.3-1.7 mm,

basally united with the lip. Lip 4-5.5 mm long, three-lobed with a united

basal part 1.9-2.5 mm long; midlobe lorate, 1.9-3 < 0.6-1 mm; side lobes

thread-like and largely pointing upwards, normally curled, 13-24 mm long;

spur a shortly stalked globular sac with an entire or bifid apex, 2.3-3.8(-4)

mm long with a thickness of 1.2-2.2(-3) mm. Gynostemium 1-1.5 mm long.

Ovary (including pedicel) 5-8 mm long, smooth.

Specimens examined: UNLOCALISED: Khao Yai National Park [comprising

parts of the provinces of Nakhon Ratchasima, Saraburi, Prachinburi and

Nakhon Nayok], Cumberlege 610 (K, spirit collection); without locality and

collector (BCU, spirit collection). NORTHERN: Mae Hong Son Province:

Peristylus tentaculatus in Thailand 33

Figure 3. Inflorescence of Peristylus tentaculatus (Pumicong 392).

Huai Pu Ling, 10 Sep 2006, Watthana & Wongnak 2153 (QBG, SING);

ibid.: Mae Sarieng, 30 Aug 2006, Pumicong 392 (QBG, SING); Chiang Mai

Province, BCU [5144] p.p. (BCU, spirit collection); ibid., Obchant Thaithong

1132 (BCU, spirit collection); ibid., Doi Suthep, 30 Oct 1905, Kerr 1/8

(K); ibid., Doi Suthep, 19 Sep 1995, Larsen & al. 46684 (AAU); ibid., Doi

Suthep, 4 Sep 1958, Sorensen & al. 4728 (BKF, C); ibid., Doi Suthep, 9 Sep

1958, Sorensen & al. 4857 (BKF, C); ibid., Doi Suthep, 9 Sep 1958, Sorensen

& al. 4871 (C), ibid., Doi Suthep, 18 Sep 1958, Sorensen & al. 5109 (BKF,

C); ibid., Doi Inthanon, 29 Sep 1910, Garrett 6] (BKF, K); Nan Province,

Doi Wao, 10 Sep 1995, Larsen & al. 46277 (AAU, SING); Tak Province,

10 Oct 1992, Obchant Thaithong 1262 (BCU, spirit collection). SOUTH

WESTERN: Prachuap Khiri Khan Province, Chanthanaoraphin s.n. (BCU).

54 Gard. Bull. Singapore 60 (1) 2008

CENTRAL: Nakhon Nayok Province, Khao Yai National Park, Khao Khieo,

11 Aug 1974, Maxwell 74-776 (AAU, L), ibid.: 14 Aug 2000, Maxwell 00-

386 (L), ibid.: 29 Aug 1963, Smitinand & Sleumer 8337 (BKF, L). SOUTH-

EASTERN: Prachin Buri Province, Obchant Thaithong 745 (BCU), ibid.:

Obchant Thaithong 1495 (BCU).

Illustrations: Seidenfaden & Smitinand (1959, p. 31): figs. 20a-e, as

Habenaria garrettii; Seidenfaden (1977, p. 42-43): figs. 17a—g, as Peristylus

tipuliferus, 18a—e; Chen Singchi & al. (1999, two colour photos on p. 346).

Habitat and flowering time: The species is found in (or on the edge of)

evergreen forest, grassy pine forest and oak forest from 100 to 1800 m. There

are also reports from rocky marshland. Flowering occurs between August

and November.

Distribution: NE India, Nepal, Myanmar, Thailand, southern China,

_ Cambodia, Vietnam.

Acknowledgements

I would like to thank the curators of the herbaria AAU, C, K, L and P for the

loan of their specimens, and the staff of the herbaria BKF, BCU, QBG and

SING for their assistance while examining their collections. The issuing of

a research permit by the National Research Council of Thailand is gratefully

acknowledged.

References

Chen, S.-C., Z.-H. Tsi and Y.-B. Luo. 1999. Native orchids of China in colour.

Science Press, Beijing and New York.

Ormerod, P. 2003. Orchidaceous Additions to the Floras of China and

Vietnam. Jaiwania 48: 139-146

Seidenfaden, G. 1977. Orchid genera in Thailand V. Orchidoideae. Dansk

Botanisk Arkiv 31: 1-149.

Seidenfaden, G. and Smitinand, T. 1959. The Orchids of Thailand — A

preliminary list. Part I. The Siam Society, Bangkok.

Gardens’ Bulletin Singapore 60 (1): 55-61. 2008

Habenaria mandersii (Orchidaceae) Newly Recorded

from Thailand with Notes on the H. hosseusii Group

H. KURZWEIL

The Herbarium, Singapore Botanic Gardens

1 Cluny Road, Singapore 259569

Abstract

The occurrence of Habenaria mandersii Collett & Hemsl. in Thailand is

newly reported. The species was previously only known in Myanmar and

the former French Indochina (Cambodia, Laos, Vietnam). It is very similar

to H. hosseusii Schltr. and it is here suspected that it has frequently been

mistaken for this species in the past. Short notes on the taxonomy of the

H. hosseusii—-mandersii—dentirostrata Tang & F.T.Wang group are also

given.

Although the orchid flora of Thailand is reasonably well known compared

with some adjacent countries (see for examples, Kamemoto & Sagarik,

1975; Seidenfaden, 1977, 1986, 1988; Nantiya Vaddhanaphuti, 2005)

new distribution records continue to be made, and particularly during the

current preparation of the Orchidaceae volume of the Flora of Thailand the

orchid flora of the country is being re-examined. Several specimens of H.

mandersii Collett & Hemsl. were recently found in herbaria in Thailand

and Singapore which is reported here. The species is known to occur in

adjacent countries but has not been recorded in Thailand before.

Habenaria mandersii Collett & Hemsl.

J. Linn. Soc., Bot. 28 (1890) 133; Seidenf., Dansk Bot. Ark. 31 (1977) 126;

Seidenf., Opera Bot. 114 (1992) 67. — Kraenzlinorchis mandersii (Collett

& Hemsl.) Szlach., Orchidee (Hamburg) 55 (2004) 58. — Typus: Burma,

Shan State, Manders s.n. (K; not seen). Figs. 1-2, Table 1.

Terrestrial herbs, entirely glabrous, except sometimes for the bract

margins, 300-550 mm tall. Basal sheaths 2-3, tubular, enveloping the

stem base; uppermost with an acute blade to 25 x 14 mm. Leaves 5,

scattered in the lower stem portion, lanceolate-elliptic, acute or subacute,

mucronate, 70-120 x 12-23 mm, margins papillose. Sterile bracts

Figure 1. Habenaria mandersii (Pumicong 386).

Figure 2. Tongue in front of the spur mouth of Habenaria mandersii. All perianth lobes

and the gynostemium removed; spur broken off. Bar =1 mm (from Pumicong 386).

longitudinal laminal flange

lip

spur

Gard. Bull. Singapore 60 (1) 2008

Habenaria mandersii in Thailand

Table 1. Distribution of various floral characters in the Habenaria hosseusii group (all

measurements in millimetres).

Character | H. hosseusii H. mandersii H. dentirostrata

Ovary length 21—26(-30) 17-22 ?

Median sepal | §.5—12(-14) x 4-5(-6) | 7.5-10.5 x 4-5.5 9.7-11.3 x ca. 5.5

Lateral sepals 8.2—-12.5(—14) x 34 8-11 x 3-4.6 11-11.3 x 4.5-5.5

(-5.5

Petals 8.5—12.2(-14) x 1.5—1.7| 8-11.5 « 1.5-2 1112.5 x 2.62.8

(-2.5)

Lip 10—17(—24) x 1-2(-3.5) | 11-16.5 x 2.5-3 18-19.3 x 2.6-3

Lateral outgrowths of the | teeth obscure rounded or teeth

tongue in front of the spur absent

entrance

Longitudinal flange in the | short stretching over much _ | short

middle of the tongue

of the length

Apex of tongue

obtuse or mucronate

entire or slightly

emarginate

retuse or bilobulate

Spur length

50-100

(28—)30-38(-47)

50-60

9-10, lanceolate, acuminate, 22-60 x 3-8 mm, sometimes papillose on the

surfaces, denticulate or papillose on the margins; apically grading into the

floral bracts. Inflorescence lax to semi-dense, 10- to 20-flowered; rachis

80-100 mm long; bracts lanceolate, acuminate, 20-25 x 3-6 mm, elongate-

papillose to glandular-hairy on the margins. Flowers white. Sepals oblong-

elliptic, subacute or obtuse, 3-veined; median sepal erect, 7.5-10.5 x 4-5.5

mm; lateral sepals spreading, oblique, 8-11 x 3-4.6 mm, basally united with

the petals and the lip. Petals oblong or oblong-elliptic, obtuse or subacute,

1- or 3-veined, 8-11.5 x 1.5-2 mm. Lip entire, oblong-spathulate, obtuse,

widest in the upper part, margins involute, 11-16.5 x 2.5-3 mm, with a 1.5-

3 mm long suberect tongue in front of the spur entrance, tongue often with

obscure rounded bulges on the sides, the longitudinal flange in the middle

of this tongue (facing the spur mouth) stretching over much of its length,

tongue apex entire or slightly emarginate; spur cylindric, (28-)30-38(-47)

mm long, slightly clavate, apex obtuse. Gynostemium 3-4 mm long, anther

with a subacute terminal connective process, anther canals 3-5 mm long,

curved or geniculately angled upwards, auricles conspicuous and ca |

mm in diameter, stigmatic processes 4-5 mm long, bent upwards. Ovary

(including pedicel) 17-22 mm long, papillose, curved, with a narrow neck.

58 Gard. Bull. Singapore 60 (1) 2008

Specimens examined: NORTHERN: Chiang Mai Province, Omkoi, Thung

Jum Roen, 29 Aug 2006, Pumicong 386 (QBG, SING); EASTERN:

Ubon Ratchathani Province, Po Sai District, Ban Sang Kung, 7 Sep 1997,

Somran Suddee 801 (BKF, spirit collection); SOUTH EASTERN: Prachin

Buri Province, | Oct 1988, Obchant Thaithong 504 (BCU, dry and spirit

collection); Prachin Buri Province, 30 Oct 1977, Amorn Ubolcholakhate

s.n. (BCU, spirit collection, material largely disintegrated); WITHOUT

LOCALITY (bought at a market in Bangkok): 25 Oct 1989, Obchant

Thaithong 772 (BCU, spirit collection).

Illustrations: Seidenfaden (1977, p. 128): fig. 80a—c [flower and floral

details]; Seidenfaden (1992, p. 65): fig. 34a-e [plant, flower and floral

details].

Habitat and flowering time: The few available detailed collector’s notes

indicate that the plants are found in dry deciduous dipterocarp forest. A

_small colony in Chiang Mai Province (collection Pumicong 386), growing

in a wet area in a grassy clearing in heavily grazed dipterocarp forest, was

visited by the author of this paper. A dense clump of about ten plants was

seen, with a further few individuals scattered around in the area. Generally,

specimens of Habenaria mandersii have been collected at altitudes ranging

from 280 to 1025 m. Flowering occurs between August and October.

Distribution: Myanmar, Thailand, Laos, Cambodia and Vietnam.

Notes: Habenaria mandersii is very similar to the Thai endemic H. hosseusii

Schltr. The most obvious differences of the latter are the much longer lip

spurs (50-100 mm; as opposed to 28-47 mm in H. mandersii) and details

of the tongue in front of the spur mouth (lateral teeth, obtuse or mucronate

apex, short laminal flange; as opposed to merely obscure rounded lateral

bulges or no lateral outgrowths at all, entire or slightly emarginate apex,

long laminal flange in H. mandersii) (see also Table 1). In view of the

striking similarity it is here suggested that H. mandersii has in the past

frequently been mistaken for H. hosseusii in the field.

Notes on the Habenaria hosseusii group

The Habenaria hosseusii group comprises the three species H.

hosseusii, H. mandersii and the little-known H. dentirostrata Tang &

F.T.Wang. The species are very similar in many respects and are probably

Habenaria mandersii in Thailand 59

closely related. All three share a similar vegetative habit with three to eight

glabrous lanceolate-elliptic leaves scattered in the lower stem portion. The

white flowers, borne in few-flowered lax to semi-dense inflorescences, are

very characteristic. Pedicel and ovary measure 21-26(-30) mm in length

in H. hosseusii and 17-22 mm in H. mandersii and usually have a narrow

neck in both species; no data are available in H. dentirostrata. In all three

species the sepals and petals are oblong or oblong-elliptic and measure 7.5-

12(-14) mm in length. Unlike in the majority of Habenaria species the lip

is unlobed in the H. hosseusii group. It is oblong-spathulate in shape with

the widest part in the distal third, has involute margins and measures about

10-17(-24) mm in length. A very distinct feature is the presence of a 1.5-

3 mm long suberect tongue in front of the spur entrance, and its detailed

structure is used to differentiate between the three species (Seidenfaden,

1977). The tongue has mostly lateral outgrowth in its basal part which may

take the shape of prominent teeth (H. hosseusii, H. dentirostrata) and are

obscure rounded bulges or absent in H. mandersii. On the blade of this

tongue (facing the spur mouth) there is a longitudinal flange which is very

short in H. hosseusii and H. dentirostrata but occupies much of its length

in H. mandersii. Apically, the tongue is entire or slightly emarginate in

H. mandersii, obtuse or mucronate in H. hosseusii and clearly retuse or

bilobulate in H. dentirostrata. Also the spur length differs among the three

species, with (28-)30-38(-47) mm long spurs found in H. mandersii and

longer spurs found in the other two species (ca 50-60 mm in H. dentirostrata,

50-100 mm in H. hosseusii). The structure of the 3-4 mm long gynostemium

is rather uniform in the group, with its 3-6 mm long anther canals curved

or geniculately angled upwards and its similarly upwards-bent 3-5 mm

long stigmatic processes. The distribution of several of these characters is

indicated in Table 1.

The examination of a large number of Thai specimens during the

present study suggests that Habenaria hosseusii and H. mandersii appear

to be distinct, the former characterised by long spurs and lateral teeth

on the tongue, the latter by rather short spurs and the lack of such teeth.

Hardly any material of H. dentirostrata was examined here as there is no

definite Thai specimens (but see below), and according to illustrations this

species is nearly identical with H. hosseusii. Although Seidenfaden (1977)

considered it as distinct, a detailed future examination of its material from

Laos and Myanmar may well show that it is conspecific with the latter.

All three species are distributed in the region stretching from

Myanmar to Vietnam. Habenaria hosseusii is currently considered endemic

in Thailand and is rather common in the northern and western parts of the

country. Its occurrence has also been reported in Laos but the respective

60 Gard. Bull. Singapore 60 (1) 2008

specimens are now referred to H. dentirostrata (Seidenfaden, 1977: p.

130). H. mandersii is the most widespread species in the group, having

been recorded from Myanmar, Thailand, Laos, Cambodia and Vietnam. H.

dentirostrata is currently only known in Laos and Myanmar. A part of a

plant referable to this species was found in the spirit collection at BCU

(Obchant Thaithong 768). However, it was bought at the Sunday Market in

Bangkok and it is therefore not known whether it originated from Thailand

or was brought over the border from Laos or Myanmar.

Habenaria mandersii was l\isted by Grant in his account of the

orchids of Burma (1895, p. 338). This is possibly a misidentification as

the plant which he describes appears to share a critical character with H.

dentirostrata (which was not known at the time). Grant’s comments on the

tongue in front of the spur mouth, “.....large obtuse ligule with revolute

sides, the convex face (towards the mouth) has 2 short conical spurs side

by side...”, apparently allude to its lateral teeth which are found in H.

dentirostrata (and H. hosseusii) but are absent in H. mandersii. However,

_ this is contrasted by the short spurs of only “1-2 inches” (25.4-50.8 mm)

which are not known in H. dentirostrata.

Acknowledgements

I would like to thank the staff of the Queen Sirikit Botanic Garden (QSB),

Forest Herbarium, Bangkok (BKF), and the Herbarium of the Botany

Department of Chulalongkorn University (BCU). I am particularly grateful!

to Dr. Somran Suddee, Dr. Obchant Thaithong and Suchada Wongpakam

for much useful advice and help during my herbarium visits. The issuing of

a research permit by the National Research Council of Thailand 1s gratefully

acknowledged.

References

Grant, B. 1875. The orchids of Burma. Hanthawaddy Press, Rangoon.

Kamemoto, H. & Sagarik, R. 1975. Beautiful Thai Orchids. The Orchid

Society of Thailand, Bangkok.

Nantiya Vaddhanaphuti. 2005. A Field Guide to the Wild Orchids of

Thailand, ed. 4. Silkworm Books, Chiang Mai.

Habenaria mandersii in Thailand 61

. Seidenfaden, G. 1977. Orchid genera in Thailand V. Orchidoideae. Dansk

| Botanisk Arkiv 31: 1-149.

Seidenfaden, G. 1986. Orchid genera in Thailand XIII. Thirty-three

epidendroid genera. Opera Botanica 89: |—216.

Seidenfaden, G. 1988. Orchid genera in Thailand XIV. Fifty-nine vandoid

genera. Opera Botanica 95: |—398.

Seidenfaden, G. 1992. The orchids of Indochina. Opera Botanica 114:

1-502.

Gardens’ Bulletin Singapore 60 (1): 63-67. 2008 63

Polyalthia saprosma (Annonaceae),

a New Species from Borneo

I.M. TURNER

Research Associate

Singapore Botanic Gardens. Singapore

Correspondence: P.O. Box 20 Winchelsea

East Sussex TN36 4WA, UK

Abstract

Polyalthia saprosma |.M. Turner, sp. nov. (Annonaceae), is described from

material hitherto confused with P. cinnamomea Hook.f. & Thomson. The

new species is recorded from Sabah (Malaysia) and Kalimantan (Indonesia)

on the island of Borneo.

While working on an account of the Annonaceae for the Tree Flora of

Sabah and Sarawak, it became clear that most of the material from Borneo

assigned to Polyalthia cinnamomea Hook.f. & Thomson was not that

species and actually represented an undescribed species.

Polyalthia saprosma |.M. Turner, sp. nov.

A Polyalthia cinnamomea Hook.f. & Thomson petalis intus villosis differt.

-Typus: Malaysia, Sabah, Beluran District, Suah Tingguan, K.B. Dev.

Camp, Sungai Sapi, 24 April 1963, SAN 363/5 [holotype, K (barcode:

K000380547), isotype, SAN]. Figs. 1 & 2.

Treelet or tree to 20 m tall. Twigs densely brown tomentose when young,

glabrous with age, drying with longitudinal wrinkles, often pale and corky.

Leaves tomentose beneath when young, becoming glabrous with age, thinly

chartaceous to subcoriaceous, drying dark to grey-brown above, brown to

green-brown below, narrowly oblong elliptic to obovate, 8-16 < 3-7 cm,

base obtuse, apex apiculate to shortly acuminate, midrib slightly sunken

above in dry leaves with crest of red-brown hairs, midrib prominent beneath,

secondary veins obscure from above, slightly raised beneath in dry leaves,

14-16 pairs, looping within margin. Petioles 4-5 mm long, 1-2 mm thick,

densely brown pubescent. Inflorescences single-flowered, subopposite

leaves. Flowers malodorous, pedicels 3-4 mm long, densely tomentose,

sepals triangular, 6 x 4 mm, densely hairy outside, petals yellow/orange

64 Gard. Bull. Singapore 60 (1) 2008

ROYAL BOTANIC GARDENS KEW

K000380547

Figure 1. The holotype of Polyalthia saprosma |.M. Turner. |

Polyalthia saprosma (Annonaceae) 65

ROYAL BOTANIC GARDENS KEW

000380548

Figure 2. Fruiting specimen (Aban Gibot SAN 32512) of Polyalthia saprosma 1.M.

Turner.

66 Gard. Bull. Singapore 60 (1) 2008

brown, linear to 8 cm long, 4 mm wide, apex acute, densely covered with

long brown hairs which tend to be paler on the inside, glabrous near the base

inside, stamens many, ca | mm long, carpels many. Fruiting pedicels 5-8

mm long, ca 2 mm thick, sepals often persisting, monocarps to 50 or more,

globose or ellipsoidal to 12 mm long, 6-8 mm in diameter, drying black or

dark brown, covered with dark brown tomentum, becoming glabrous with

age, stipe 2-8 mm long. Seeds 1-2.

Specimens seen: Malaysia, Sabah: Kelumpang Balong, Tawau, A. Bakar, 9

Jun 1961, (SAN 18508); Tawau, Oct 1922-Mar 1923, A.D.E. Elmer 21475;

Mile 12 M. Road Kalabakan, Tawau, Aban Gibot (SAN 30571), 27 Jul 1962;

Ulu Balong, Tawau, Aban Gibot (SAN 32512), 10 Nov 1962; east of Sepilok

Camp, Sandakan, Aban Gibot (SAN 78649), 11 Dec 1973; Kota Merudu,

Aban Gibot (SAN 99586), 11 Aug 1983; Mile 32, Ulu Dusun, Sandakan,

Aloysius Simbut (SAN 78558), 12 Jul 1973; Kretam For. Res., Sandakan,

Amin & Soinin (SAN 96706), 20 May 1983; Danum Valley, Ulu Segama,

_E.J.F. Campbell et al. (SAN 110701), 30 May 1986; Kalabakan, Fedilis (SAN

94723), 6 Apr 1982; Sandakan, H.P. Nooteboom 1613, 17 Mar 1970; Ulu

Segama, Lahad Datu, L. Madani & Ismail P. (SAN 108717), 29 Mar 1985;

Madai Forest, Lahat Datu, L.E. Teo & B. David (T. & P. 1113) (KL 36/3),

25 Sep 1986; Bettotan, Orolfo 2871, 23 Feb 1933. Indonesia, Kalimantan:

Batu Badinging, Bukit Raya, J.F! Veldkamp 8546, 5 Feb 1983.

The flowers are reported to smell of rotten meat which is reflected in the

chosen epithet (Greek, sapros = rotten, putrid; osme, osma = fragrance,

odour). The scent of decay and furry, dark flowers suggest sapromyophily

as the pollination syndrome, with some sort of carrion-visiting insects the

pollinators.

Vegetatively P. saprosma is more likely to be confused with

Polyalthia motleyana (Hook.f.) Airy Shaw than P. cinnamomea. One

fairly consistent distinction is that in PR: motleyana the dry leaves below

have venation that contrasts with the lamina by being paler whereas in P.

saprosma the veins tend to be darker than the lamina.

Flowering specimens of P. saprosma are often collected before the

corolla is fully expanded. The dark brown tomentum on the relatively short

petals has led to confusion of some specimens with Polyalthia chrysotricha

Ridl., though this species is readily distinguished by the cordate leaf bases,

dense tomentum on the undersides of the leaves and the broader petals.

Differences between P. saprosmaand P. cinnamomeaare summarised

in Table |. Flowering and fruiting specimens of P. saprosma are illustrated

in Figs. | and 2.

Polyalthia saprosma (Annonaceae) 67

Table 1. Summary of differences between Polyalthia saprosma and Polyalthia

cinnamomea.

Character Polyalthia saprosma Polyalthia cinnamomea

Leaves Typically obovate, apiculate Typically ovate lanceolate,

drying grey-brown or olive acuminate, drying warm brown,

brown, fringe of red-brown midrib glabrous above

hairs along midrib above

Flowers Petals villose outside and upper _ Petals tomentose outside,

part of inner face glabrous inside

Fruits Monocarps globose or ellipsoidal Monocarps pyriform, 20-25 mm

to 12 mm long, 6-8 mm in long, 12-15 mm diameter, rusty

diameter, glabrescent tomentose

Acknowledgements

Financial support for this research has come through the Arnold Arboretum

of Harvard University. Professors Robert Cook and Peter Ashton are

thanked for facilitating this support. Prof. Simon Owens permitted access

to the facilities of the herbarium and library at Royal Botanic Gardens Kew.

Dr J.F. Veldkamp (L) kindly provided assistance with the Latin diagnosis.

Gardens’ Bulletin Singapore 60 (1): 69-71. 2008 69

Sarcotheca lunduensis (Oxalidaceae), a New Species from

W. Sarawak, Malaysia

J.F. VELDKAMP

National Herbarium of The Netherlands, Leiden University

2300 RA Leiden, The Netherlands

In 1993 I saw a collection from Sarawak (Kessler PK 320) of a Sarcotheca

Blume (Oxalidaceae) which looked like S. ochracea Hallier f., but with

rather smaller leaves that were glabrous underneath, while the petals were

said to be white, which is very exceptional in the woody members of this

family, where they tend to have shades of red from pink to nearly blackish

red (i.s.). Also this came from the Lundu District in the Kuching Division

of Sarawak, while S. ochracea occurs mainly in the Bintulu Division, with

some collections from E Sibu and S Miri, and recently has been collected

twice in Kalimantan (N Kutai, see note below). Over the years another

four collections arrived from Lundu and Sematan (ca 1° 40-48’ N, 109° 40-

50’ E). An analysis with DELTA showed that they agreed with each other,

except that where flowers were mentioned they were said to be red.

The flowers of the un-named plant are distylous with a short-styled

form (SF: styles shorter than the stamens) and a long-styled form (LF: styles

longer than the stamens). The mid-styled form (MF: stigmas between the

two rows of anthers) so far is unknown in the genus.

A key to the distinction of the two species is as follows -

- Leaves underneath glabrous, waxy, cinnamon or grey. Pedicels lower

joint 1-3 mm long. Petals obovate, ca 2.7 mm long. Filaments in LF the

shorter 0.7-1 mm long, the longer 1.3-1.4 mm long. Pistil in SF ca 1 mm

long. Seeds ca 5 mm wide. Blades 5-10.5 cm long. Panicles 4-9.5 cm

long. W. Sarawak (Kuching Div.: Lundu, Sematan) ....... S. lunduensis

- Leaves underneath pubescent, not waxy, reddish brown. Pedicels lower

joint 4-8 mm long. Petals lanceolate to oblanceolate, 4.5-8 mm long.

Filaments in LF the shorter 1.5-2 mm long, the longer 2-2.5 mm long.

Pistil in SF 1.7-2 mm long. Seeds 3.25-4.5 mm wide. Blades 7.5--23.5

cm long. Panicles 7-70 cm long. C. Sarawak (Bintulu, S Miri, E Sibu

WD ies) se, eam att (INSK GICAL)... s:0ceccatevovevaseversnnernendsconaorns S. ochracea

70 Gard. Bull. Singapore 60 (1) 2008

Sarcotheca lunduensis Veldkamp, spec. nov.

Ab omnibus congeneribus calycibus extus ferruginee puberulis, petiolis 1-

1.5 mm latis, paniculis ferruginee velutinis, petalis obovatis ca 2.7 mm

longis, filamentis brevioribus in forma brevistyla (SF) ca 2.5 mm longis,

ovariis ca 1 mm latis differt. - Typus: Sarawak, Kuching Division, Sematan

Distr., Biawak, Ladang DAFA, S 56383 (M. Eja) (L, holo; K, KEP, SAN,

SAR, MO, iso).

Petiole 8-21 by 1-1.5 mm; petiolule 4-5.5 by 1-1.5 mm. Leaves unifoliolate,

subcoriaceous, elliptic, ovate to oblong, above glabrous, beneath midrib

ferruginous hairy, waxy (see note), cinnamon or grey, base broadly cuneate,

rounded, or emarginate, margins not parallel, apex acuminate or caudate,

venation above not very conspicuous, nerves 5-8 pairs. Leaflets 5-10.5 by

2.75-4.5 cm. Panicles shorter to longer than subtending leaf, 1-3 together,

erect, loosely branched, 4-9.5 cm long, ferruginous velvety; branches more

or less elongated, not forked at the top, 5.5-10 mm long. Pedicels lower

_joint 1-3 mm long, upper joint 0.3-1 mm long. Flowers distylous (SF +

LF). Calyx 2.25-3 mm high, outside brown puberulous, reddish brown, red,

or purplish, persistent in fruit. Sepals suborbicular, ovate, broadly obovate,

or elliptic, 2-2.3 by 1-2 mm, apex rounded to obtuse. Petals obovate, ca

2.7 by | mm, rounded. Filaments in SF the shorter ca 2.5 mm long, the

longer ca 3 mm long, LF the shorter 0.7-1 mm long, LF the longer 1.3-

1.4 mm long. Pistil puberulous, in SF ca 1 mm long, in LF ca 2.4 mm

long. Ovary subglobose, 0.8-1 by | mm. Fruit subglobose, 8-9 by 6-8 mm.

Rimae conspicuous, papillose. Seeds ca 6.5 by 5 mm. Testa transversally

rugose. Cotyledons ca 4 by 2 m. Radicle ca | by 0.4 mm.

Distribution: Sarawak, Kuching Div., Sematan Distr. [S 56383 (frts), S

56384 (fls) (M. Eja), S 65450 (frts) (Hj. Othman & Yahud et al.), Lundu

Distr. [Kessler PK 320 (fls & frts), S 62268 (frts) (Hj. Othman Ismawi et

al. |

Habitat: Riverine forest, heath forest (kerangas), up to 200 m alt.

Phenology: Flowers in end July, end December, fruits in March, early April,

end July, end December. S$ 56383 (frts) and S 56384 (fls) apparently were

collected close together.

Vernacular name: Piang (Iban), also used for S. glauca (Hook. f.) Hallier

Sarcotheca lunduensis (Oxalidaceae) al

Collector s notes: A straggling shrub or small tree, 3--5 m tall. Bark surface

smooth, grey. Leaves glabrous. Flowers borne at the end of the twig. Sepals

brown, red, fine brown hairy. Corolla red, white (once, see note). Fruit red,

bright red.

Uses: Fruits edible, sour to taste.

Notes: The lower surface of the leaves of Sarcotheca species may be waxy

(previously called “glaucous”). This may be checked in dry material by

putting a small flame gently at some distance underneath the leaf (lower side

up, and don’t burn it). When there is wax a quickly spreading discoloration

will be seen. This experiment can only be done once.

Kessler PK 320 states that the petals would be white. This is quite

exceptional for the genus, where whitish flowers have been reported for

S. laxa (Ridl.) Knuth and S. rubrinervis Hallier f., while other collections

of this and other species state that they have various shades of red: pink to

crimson with nearly blackish tips.

Sarcotheca ochracea Hallier f.

Specimens of this species have been collected in E. Kalimantan, which

is a new record for Indonesia. Although quite disjunct with the Sarawak

collections at the moment | cannot find any significant differences.

The Kalimantan specimens are: Sidiyasa et al. 1238, N Kutai, Batu

Ampar, PT Kiani Lestari, c. 1°N, 117° E, 26 November 1994 (L, WAN);

Slik BE4-287, N. Kutai, Berau Distr., 2° 5’ 45” N, 117° 17’ 30” E, 2 May

1997 (E):

The type specimen of this species, Haviland 2343, was collected in

Bintulu, Kalong, 1893. The Gazetteer of Sarawak (Mohizah et al/., 2006)

mentions only a Kalong from the Sri Aman Div. (former Simanggang or

2nd Div.), far outside the distributional area of the species. I am fairly

confident that Haviland knew where he was and that there were two places

by that name.

Reference

Mohizah, M., S. Julia & W.K. Soh. 2006. 4 Sarawak Gazetteer. ix, 181 pp

+ 13 illus. Sarawak Forestry Department & Forest Research Institute,

Malaysia.

Gardens’ Bulletin Singapore 60 (1):73-153. 2008 73

Thirty Two New Species of Bulbophyllum (Orchidaceae)

from Sulawesi

J.J. VERMEULEN ' AND P. O’BYRNE ”

' National Herbarium Netherlands, Leiden Branch

P.O. Box 9514, 2300 RA Leiden, The Netherlands

*11 Hume Avenue #06-01, Hume Park 1, Singapore 598723

Abstract

Thirty two new species of Bulbophyllum, all from Sulawesi, are described.

Names currently in use for various sections of Bulbophyllum are made

junior synonyms to older sectional names that have been ignored so far,

partly because their taxonomic content, as appears from the suite of species

listed with the original descriptions of these sections, was ambiguous. These

sectional names are lectotypified and re-instated.

Introduction

Thirty two new species of Bulbophyllum, all from Sulawesi, are

described. In addition, new definitions of some sections of Bulbophyllum

are proposed. Averyanov (1994) and Ormerod (2002) have found names

prior to those commonly used for several sections of Bulbophyllum. While

admitting that name changes are a nuisance to many, we felt it would be

appropriate to further pursue this line of investigation in order to install a

new set of sectional names that, hopefully, will remain stable because they

are based on the oldest names available. The oldest source of sectional

names (in the genus Diphyes Bl., = Bulbophyllum, Thou.) is Blume (1825).

Often, we were able to choose the taxonomic contents of these names

by lectotypification. We have done this in such a way that the number of

Blume’s names to disappear into synonymy is minimized, particularly

in cases where the taxonomic contents of our newly delimitated sections

does not coincide with the contents of the sections as accepted so far. For

instance, sect. Polymeres, below, is better not given a name that derives from

one of its constituent sections to avoid confusion, because those section

names have been accepted and used by consensus since their description by

Schlechter (1913).

74 Gard. Bull. Singapore 60 (1) 2008

Most data on the flowering periods of the species described below

derive from plants in cultivation.

Bulbophyllum sect. Brachystachyae Benth. & Hook.f.

(= genus Cochilia, sect. Globiceps, genus Osyricera, sect. Saurocephalum)

Bulbophyllum sect. Brachystachyae Benth. & Hook.f., Gen. Pl. 3 (1883)

504. — TYPE SPECIES (a lit. reference given only): Bulbophyllum

(“Bolbophyllum”’) repens Wall.

Cochlia Bl., Biydr. Fl. Ned. Ind. (1825) 320. — Bulbophyllum Sect. Cochlia

(BI.) Benth. & Hook.f., Gen. Pl. 3 (1883) 503. — TYPE SPECIES:

Cochlia violacea BI. (= B. salaccense Rchb.f.).

Bulbophyllum sect. Cylindracea_ Pfitz. in Engler & Prantl, Nat.

Pflanzenfam., ed. 2, 6 (1889) 179. — TYPE SPECIES: Bulbophyllum

(“Bolbophyllum’’) cylindraceum Lindl.

_ Bulbophyllum sect. Saurocephalum Schltr., Feddes Repert. 10 (1912)

184. — LECTOTYPE SPECIES (here designated): Bulbophyllum

saurocephalum Schltr.

Bulbophyllum sect. Globiceps Schltr., Feddes Repert. Beih., 1 (1913) 875. —

TYPE SPECIES: Bulbophyllum globiceps Schltr.

Osyricera Bl., Bidr. Fl. Ned. Ind. (1825) 307.— Bulbophyllum Sect. Osyricera

(BIL.) J.J. Sm., Bull. Jard. Bot. Buitenzorg, Ser. 2, 13 (1914) 35. —

TYPE SPECIES: Osyricera crassifolia Bl. (= B. osyricera J.J. Sm.).

Bulbophyllum sect. Diptychanthes sensu Rchb.f. in Bonplandia 5 (1857) 57. —

SPECIES MENTIONED: Bulbophyllum salaccense Rchb.f.

Not Diphyes sect. Diptychanthes Bl., Bydr. Fl. Ned. Ind. (1825) 311.—TYPE

SPECIES: Diphyes mutabilis Bl. [= B. mutabile (Bl.) Lindl., is sect.

Stachysanthes; see below].

Bulbophyllum ankylodon J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo pubifloro floribus in seriebus 4 circa rhachidem, labello

papilloso abaxialiter callo crasso differt. — Typus: Indonesia, Sulawesi,

central part, SBG-O 4713 (holo, SING). Fig. 1.

Roots spreading. Rhizome creeping or shortly straggling, 2.5-4 mm diam.,

sections between pseudobulbs ca 3 cm long, bracts not persistent. Pseudobulbs

distant, ovoid, 3.5-4.2 x 1-1.4 cm, distinctly but obtusely 3-angled. Petiole

1-1.4 cm long. Leaf blade elliptic to obovate, 10-12 x 2.6-3.3 cm, index

(length/width) 3.6-3.9; acute. Inflorescence patent, a rather dense raceme,

7.5-15.5 cm long, 30-50-flowered. Peduncle 2.5-4 cm, bracts 2 (1 basal),

New Species of Bulbophyllum (Orchidaceae) 5 hs)

ey yerowr Hr

atte re j \ a ae

(bes ee eee

= Sa ‘tg

Figure 1. Bulbophyllum ankylodon J.J. Verm. & P. O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepals, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 47/3 (spirit

sample).

76 Gard. Bull. Singapore 60 (1) 2008

§ap

the longest 5-9 mm long. Rhachis thickened, cylindrical, 5.8-11.5 cm long,

2.5-3.2 mm across. Floral bracts 3-4.2 mm, acute. Flowers not resupinate,

spirally arranged in 4 rows, not fully opening, many open simultaneously.

Pedicel and ovary ca 1.5 mm long, basal node about flush with the surface

of the rhachis. Median sepal approx. porrect, obovate, ca 4 x 2.6 mm, index

|.5-1.6; top cucullate, rounded, margins entire, papillose-ciliolate, base rather

broadly attached; rather thick, 3-veined, surface adaxially glabrous, abaxially

shortly papillose-hirsute. Lateral sepals adnate along the lower margins, basal

part recurved, top part incurved, oblique, triangular, ca 4.2 x 2 mm, index

ca 2.1, top flat, subacute, base broadly attached; 4-veined, otherwise as the

median sepal. Petals recurved, ovate-triangular, ca 3.2 x 0.8 mm, index ca

4; subacute, margins entire, papillose-ciliolate, base broadly attached; rather

thick, l1-veined, surface approx. glabrous. Lip slightly curved, general outline

subtriangular, ca 2 x 1.3 mm, index 1.5-1.6 (all without artificial spreading,

hastate with a drawn out, elliptic midlobe and deltoid, acute sidelobes when

spread); rounded, margins entire, coarsely papillose distally; thick; adaxially

_ slightly concave near the base, with 2 rounded ridges starting near the margin

at about '4 of the length of the lip and then converging and running up almost

to the tip of the lip, adaxial surface finely papillose distally; abaxially without

a ridge near the base, but with a rounded lump in the distal half, surface

approx. glabrous except for the papillose lump. Column ca 1.5 mm long,

stigma without a tooth at its base, column foot with 2 rounded swellings

along the margin near its base, and a slight swelling just above the ligament.

Stelidia minute, abruptly bent downwards but with an antrorse, approx.

acute tip, along the upper margin with a very large (far more conspicuous

than the stelidia themselves), antrorse, wedge-shaped wing with a broadly

rounded, erose top margin. Anther abaxially with a crest near the base, surface

somewhat papillose, front margin drawn out into a triangular, rounded beak,

margin coarsely papillose. Pollinia 4, the inner much flattened and about as

long as the outer, all ellipsoid-ovoid; no appendages present.

Colour: Vegetative parts green, not suffused with red. Rachis densely mottled

pinkish purple. Sepals and petals yellowish, veins and margins purplish red.

Lip orange yellow, particularly the adaxial surface stained with purplish red.

Column yellow.

Habitat & ecology: Epiphyte in mossy montane forest. Alt. ca 1000 m.

Flowering Feb, Jul-Sep.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

New Species of Bulbophyllum (Orchidaceae) 77

Notes: Bulbophyllum ankylodon differs from B. pubiflorum by having the

flowers in 4 (not 5) rows around the rhachis, and the papillose (not glabrous)

lip with a distinct thickening on the abaxial surface.

Bulbophyllum condylochilum J.J. Verm. & P.O’ Byrne, sp. nov.

A Bulbophyllis fraterno et B. illecebrum /abello distaliter abrupte

conspicueque inflato differt. - Typus: Indonesia, Sulawesi, central part,

LEI 20050892 (holo, L). Fig. 2.

Roots mainly below the pseudobulbs. Rhizome creeping, 2.5-3 mm diam.,

sections between pseudobulbs 0.8-1.2 cm long, bracts barely persistent.

Pseudobulbs loosely clustering, |.4-2.3 x ca 0.9 cm, round in section. Petiole

0.7-1.2 cm long. Leaf blade elliptic, 4.2-5.9 x 1.3-1.8 cm, index (length

width) 3.2-3.3; acute. Inflorescence a rather dense raceme, ca 14 cm long,

ca 18-flowered. Peduncle erect to patent, ca 10 cm, bracts ca 4, the longest

ca 8 mm long. Rhachis nodding at the base, pendulous, not thickened, ca

3.6 cm long. Floral bracts approx. triangular, 2.5-3 x ca 2.5 mm, acute.

Flowers not resupinate, not fully opening, many open simultaneously.

of the rhachis. Median sepal somewhat recurved, approx. elliptic, ca 5.5 x

3.5 mm, index 1.5-1.6; top cucullate, rounded, margins entire, very finely

papillose, base narrowly attached, almost unguiculate; rather thin, 3-veined,

surface glabrous. Lateral sepals adnate along the lower margins, recurved,

top part slightly incurved, oblique, ovate-triangular, ca 5.2 x 2.1 mm, index

2.4-2.5, top flat, acute, base broadly attached; thin but with a callus distally,

near the lower margin, otherwise as the median sepal. Petals somewhat

recurved, narrowly triangular, ca 4 x 1 mm, index ca 4; acute, margins entire,

the lower somewhat erose near the base, base narrowly attached, almost

unguiculate; thin, 1-veined, glabrous. Lip almost straight, general outline

subspathulate with a slightly widened base, ca 4.5 x 1.9 mm, index 2.3-2.4,

rounded, margins entire; proximally thick and distally very thick, rather thick

in between, surface glabrous but distally papillose towards the margins on

both sides; adaxially concave near the base, with a transverse wall connecting

the sides just above the ligament, top part proximally abruptly thickened,

with the margins abruptly turned upwards into rounded auricles, and convex;

abaxially without a ridge near the base, top part convex and with a rounded

median callus. Column ca 1.3 mm long, stigma without a tooth at its base,

column foot with two obtuse, patent lateral teeth near its base, connected by a

thick transverse ridge. Stelidia minute, abruptly bent downwards but with an

antrorse, approx. acute tip, along the upper margin with a very large (far more

conspicuous than the stelidia themselves), antrorse, somewhat obliquely ovate

Gard. Bull. Singapore 60 (1) 2008

Figure 2. Bulbophyllum condylochilum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepals, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 5827 (spirit

sample).

New Species of Bulbophyllum (Orchidaceae) 79

wing with an obtuse top margin. Anther abaxially with a crest near its base,

surface approx. glabrous towards the base, papillose towards the tip, front

margin drawn out into a long, concave, rounded, papillose beak. Pollinia

4, the inner much flattened and slightly shorter than the outer, ellipsoid; no

appendages present.

Colours: Sepals whitish near the base, with thick, blackish purple lines over

the veins, entirely blackish purple near the top. Petals whitish near the base,

increasingly stained blackish purple towards the top. Lip whitish, margins

stained with blackish purple.

Habitat & ecology: Epiphyte in montane forest on steep slopes. Alt. 1200—

1400 m. Flowering Jun, Aug, Sep.

Distribution: (NDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. condylochilum differs from B. fraternum and B. illecebrum J.J.

Verm. & P. O’Byrne in the distally abruptly and conspicuously swollen lip.

The leaves are also distinctly wider.

Bulbophyllum fraternum J.J. Verm. & P. O'Byrne, sp. nov.

A Bulbophyllo illecebrum sepalo mediano glabro, petalis ligulatis abrupte

coartantibus prope apicem papillosis differt.—Typus: Indonesia, Sulawesi,

central part, SBG-O 5827 (holo, SING). Fig. 3.

Roots mainly below the pseudobulbs. Rhizome creeping, 2-2.5 mm diam.,

sections between pseudobulbs 0.6-1.5 cm long, bracts barely persistent.

Pseudobulbs slightly distant, ovoid, 2-2.3 x 1.2-1.5 cm, only slightly and

obtusely angular. Petiole 0.8-1.2 cm long. Leaf blade elliptic, 12-13 x 0.8-0.9

cm, index (length/width) 13-17; acute. Inflorescence a rather dense raceme,

5.5-11 cm long, 9-12-flowered. Peduncle erect to patent, 2.5-5.5 cm, bracts

ca 3, the longest 5.5-8.5 mm long. Rhachis nodding at the base, pendulous,

not thickened, 3-5.5 cm long. Floral bracts approx. triangular, ca 3.3 x 3 mm,

acute. Flowers not resupinate, not fully opening, many open simultaneously.

Pedicel and ovary ca 2.5 mm long, basal node approx. flush with the surface

of the rhachis. Median sepal somewhat recurved, approx. elliptic, ca 6 x 3.5

mm, index cal.7; top cucullate, rounded, margins entire, approx. glabrous,

base rather narrowly attached; rather thin, 3-veined, surface glabrous. Lateral

sepals adnate along the lower margins, recurved, top part slightly incurved,

oblique, ovate-triangular, ca 6.2 x 1.8 mm, index 3.4-3.5, top flat, subacute,

base broadly attached; thin but with a slight callus distally, near the lower

80 Gard. Bull. Singapore 60 (1) 2008

Figure 3. Bulbophyllum fraternum J.J). Verm. & P.O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepals, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 5828 (spirit

sample).

New Species of Bulbophyllum (Orchidaceae) 81

margin, otherwise as the median sepal. Petals recurved to spreading, linear

with a slightly wider base, ca 4.4 x 0.9 mm, index 4.8-4.9; subacute, margins

entire, finely papillose distally, base broadly attached; thin, 1-veined,

glabrous but somewhat papillose distally. Lip almost straight, general outline

subspathulate with a widened base, ca 5 x 1.7 mm, index 2.9-3, rounded,

margins entire; proximally rather thick, distally very thick, glabrous; adaxially

concave near the base, with a transverse wall connecting the sides just above

the ligament, top part distinctly convex; abaxially without a ridge near the

base, top part slightly concave but with a rounded median callus. Column ca

2 mm long, stigma without a tooth at its base, column foot with two obtuse,

patent or slightly retrorse wings near its base. Stelidia minute, abruptly bent

downwards but with an antrorse, approx. acute tip, along the upper margin

with a very large (far more conspicuous than the stelidia themselves), antrorse,

approx. rectangular wing with a truncate, slightly erose top margin. Anther

abaxially with a crest near its base, surface approx. glabrous towards the

base, papillose-ciliate towards the tip, front margin drawn out into a distinct,

concave, triangular, obtuse, papillose beak. Pollinia 4, the inner much flattened

and slightly shorter than the outer, ellipsoid; no appendages present.

Colours: Median sepal and petals translucent white with thick purplish red

veins. Lateral sepals very pale greenish, with purplish red veins and some

staining in the same colour. Lip pale orange, top part suffused with some

purplish red. Column white, stained pinkish purple.

Habitat & ecology: Epiphyte in montane forest on steep slopes. Alt. 1100—

1200 m. Flowering Feb-May, Aug, Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. fraternum differs from B. illecebrum in the shape of the petals,

being strap-shaped over most of their length, abruptly narrowing and papillose

at the tip. In B. illecebrum they are gradually tapering towards the tip and

entirely glabrous. B. fraternum also differs in having a glabrous median

sepal. Vegetatively, the plant resembles B. centrosemiflorum J.J. Sm. (sect.

Hyalosema), from New Guinea, with its swollen, ovoid pseudobulbs and

very narrow leaves.

Bulbophyllum kiamfeeanum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo pubifloro sepalis abaxialiter glabris, labello ovato differt.

— Typus: Indonesia, Sulawesi, central part, SBG-O 4806 (holo, SING).

Fig. 4.

82 Gard. Bull. Singapore 60 (1) 2008

Roots spreading. Rhizome creeping or shortly erect in mature plants, 2-3 mm

diam., sections between pseudobulbs 1.2-1.8 cm long, bracts not persistent.

Pseudobulbs loosely clustering, ovoid, 2.5-3 x 1-1.2 cm, without distinct

angles. Petiole 0.2-0.3 cm long. Leaf blade elliptic, 10-13 x 1.8-2.2 cm, index

(length/width) 5.8-5.9; acute. Inflorescence patent, a rather dense raceme, ca

9.2 cm long, 18-flowered. Peduncle ca 4.7 cm, bracts ca 3, the longest ca 4 mm

long. Rhachis somewhat nodding, slightly thickened, cylindrical, ca 4.5 cm

long, ca 3 mm across. Floral bracts ca 1.6 mm, acute. Flowers not resupinate,

spirally arranged in 4 rows, not fully opening, many open simultaneously.

Pedicel and ovary ca 2 mm long, basal node approx. flush with the surface

of the rhachis. Median sepal approx. recurved, ovate, ca 4.2 x 2.9 mm, index

1.4-1.5; top cucullate, rounded, margins entire, papillose-ciliolate, base rather

broadly attached; rather thin, 3-veined, surface glabrous. Lateral sepals adnate

along the lower margins, basal part recurved, top part incurved, oblique,

triangular, ca 4.2 x 2 mm, index ca 2.1, top flat, obtuse, base broadly attached;

3-veined, otherwise as the median sepal. Petals recurved, triangular, ca 3.6

_x 0.9 mm, index ca 4; acute, margins entire, papillose-ciliolate, base broadly

attached; thin, 1-veined, surface glabrous. Lip straight, general outline ovate,

ca 3.2 x 1.5 mm, index ca 2.1 (all without artificial spreading); rounded,

margins entire, finely papillose; rather thick, surface glabrous; adaxially

slightly concave near the base, convex towards the tip, abaxially without a

ridge near the base. Column ca 1.8 mm long, stigma without a tooth at its base,

column foot with 2 slight, rounded swellings along the margin near its base,

deeply sunk in between the basal part of the lateral sepals without even the tip

protruding. Stelidia minute, abruptly bent downwards but with an antrorse,

obtuse tip, along the upper margin with a very large (far more conspicuous

than the stelidia themselves), antrorse, rhombic wing with a truncate, slightly

erose top margin. Anther abaxially with a crest near the base, surface approx.

glabrous, front margin drawn out into a triangular, rounded beak, margin

coarsely papillose. Pollinia 4, the inner much flattened and about as long as

the outer, all ellipsoid-ovoid; no appendages present.

Colour: Pseudobulbs olive green, petiole suffused with some red, leaf blade

dark green. Peduncle and rhachis densely mottled purplish red. Sepals

yellowish, somewhat pale green towards the tip, the median with the veins

and the margins purplish red especially in the proximal half, the laterals over

their entire length. Petals translucent white with a purplish red midvein and

margins. Lip yellow, with some red markings near the base. Column white,

stelidia pale green.

New Species of Bulbophyllum (Orchidaceae) 83

Figure 4. Bulbophyllum kiamfeeanum J.J. Verm. & P. O’Byme. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepals, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 4806 (spirit

sample).

84 Gard. Bull. Singapore 60 (1) 2008

Habitat & ecology: Epiphyte in mossy montane forest. Alt. ca 1450 m.

Flowering Oct.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Bulbophyllum kiamfeeanum is most similar to B. pubiflorum but

differs by the sepals, which are glabrous abaxially. The lip 1s ovate in general

outline (not hastate). Vegetatively it differs from other Sulawesi species of the

section with a generally creeping rhizome and similar flowers (B. ankylodon

J.J. Verm., B. pubiflorum Schlitr., B. trigonobulbum J.J. Sm.) by having a

much shorter rhizome with loosely clustered pseudobulbs.

Named after Mr. Kiam Fee (Paul) Leong at the Herbarium of Singapore

Botanic Gardens.

Bulbophyllum molle J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo osyriceroidi pedunculo rhachide circa aequilonga, rhachide

_circa 4 mm crassa, petalis truncatis, labello parte apicali recta differt.

— Typus: Indonesia, Sulawesi, central part, SBG-O 5286 (holo, SING).

Fig. 5.

Roots spreading. Rhizome creeping, 3.5-7 mm diam., sections between

pseudobulbs 0.9-1.8 cm long, bracts not persistent. Pseudobulbs distant,

minute, ovoid to ellipsoid, 0.4-0.8 x 0.4-0.8 cm, rounded. Petiole 2.5-6.5 cm

long. Leaf blade elliptic to obovate, 9.5-21 x 1.9-3 cm, index (length/width)

5-7; obtuse. Inflorescence erect to patent, a lax raceme, 18-28 cm long, 15-

38-flowered. Peduncle 10-15 cm, bracts 3, the longest 6-16 mm long. Rhachis

thickened, cylindrical, 8-13 cm long, ca 4 mm across. Floral bracts 3.5-4 mm,

acute. Flowers not resupinate, spirally arranged, not fully opening, several

open simultaneously. Pedicel and ovary ca 1.3 mm long, basal node approx.

flush with the surface of the rhachis. Median sepal somewhat recurved,

ovate, ca 8.2 x 2 mm, index ca 4.1; subacute, margins slightly and finely

erose distally, base broadly attached; rather thin but rather thick distally and

along the midvein, 3-veined, surface glabrous. Lateral sepals adnate along

the lower margins, approx. porrect, oblique, ca 8 x 3.2 mm, index ca 2.5,

acute, margins entire, base rather broadly attached; rather thin, 3-4-veined,

otherwise as the median sepal. Petals recurved, obovate-spathulate, ca 2.2 x

1.8 mm, index cal.2; truncate-rounded, margins finely erose, base narrowly

attached; rather thin but thick proximally, 1-veined, surface approx. glabrous.

Lip slightly curved about half-way, top part straight, general elliptic-ovate, ca

5.2 x 2 mm, index ca 2.6 (all without artificial spreading), rounded, margins

entire, surface finely papillose distally; thick; adaxially slightly concave near

New Species of Bulbophyllum (Orchidaceae) 8

| 4

| c aw ae WY 7 as ‘

L=-=->)

Figure 5. Bulbophyllum molle J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepals, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 5286 (spirit

sample).

86 Gard. Bull. Singapore 60 (1) 2008

the base, without a median slit, transition to the top part a sharp, narrowly

v-shaped fold with the legs starting at the margin at about 4 of the length of

the lip, converging and meeting at about '2 of the length of the lip, top part

convex; abaxially with a truncate ridge up to about 4 of the length of the lip,

median part slightly concave, top part slightly convex because of two flat

longitudinal calli meeting over the midvein. Column ca 2 mm long, stigma

without a tooth at its base, column foot without teeth. Stelidia 0.9 mm long,

truncate and shallowly 3-toothed, with an antrorse, long, slender, triangular,

acute tooth along the upper and the lower margin. Anther abaxially with a

crest near the base, surface somewhat papillose, front margin drawn out into

a rounded beak with a coarsely and irregularly papillose margin. Pollinia

4, the inner much flattened and about half as long as the outer, all ellipsoid-

ovoid; no appendages present.

Colour: Rhachis pale green. Sepals very pale greenish with dark purplish red

veins and some stains in the same colour. Petals translucent white, midvein

_ purplish red, margins almost black. Lip yellowish green, stained purplish red,

and blackish purple towards the base. Column very pale greenish.

Habitat & ecology: Epiphyte in mossy montane forest. Alt. ca 1000 m.

Flowering Jul.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. molle is similar to B. osyriceroides J.J. Sm., from Sumatra. It differs

in having a peduncle about as long as the rhachis (peduncle much shorter

than the rhachis in B. osyriceroides), a rhachis of approx. 4 mm thick (2

mm in B. osyriceroides), truncate petals (rounded with an apiculate tip in B.

osyriceroides), and a lip without a median slit and with a straight top part

(with median slit and with a slightly recurved top part in B. osyriceroides).

Bulbophyllum sect. Epicranthes (Bl.) Hook.f.

Epicranthes Bl., Bijdr. Fl. Ned. Ind. (1825) 307. — Bulbophyllum sect.

“Epicrianthes” (Bl.) Hook. F.,; FI. Brit: India 5 (1890) 753;

Bulbophyllum subg. “Epicrianthes” (Bl.) Schltr., Feddes Repert.

Beih. 1 (1913) 705. - TYPE SPECIES: Epicranthes javanica BI.

[Bulbophyllum epicrianthes (Bl\.) Hook.f. ].

Assuming that the name Epicranthes is derived from the Greek ‘epikron’,

New Species of Bulbophyllum (Orchidaceae) 87

sailyard, which is likely to refer to the vegetative shape of the type species

and many others (a hanging rhizome with patent leaves in two rows), there is

no reason to accept the later modification into ‘Epicrianthes’.

Most species of section Epicranthes are rare plants, and appear to occur

very locally. They are known from a single or a few collections only. Next

to this, the distinguishing characters of many species would be considered

of minor importance in other sections of the genus. B. spodotriche and B.

xanthomelanon, described below, certainly fall within this category. For the

time being, we continue to apply a narrow species concept in this section,

because so far it appears the most accurate description of morphological

patterns that we observe in this section.

Bulbophyllum brachytriche J.J. Verm. & P.O’ Byrne, sp. nov.

A Bulbophyllo vesiculoso sepalis angustioribus, labello rotundato, vesiculis

multo minoribus differt. — Typus: Indonesia, Sulawesi, central part, SBG-O

5052 (holo, SING). Fig. 6.

Roots most produced close to the base of the rhizome. Rhizome hanging, ca

13 cm long, 1.2-2 mm diam., sections between pseudobulbs 1.5-2.5 cm long,

bracts persistent. Pseudobulbs well spaced, ovoid to almost cylindrical, 1-1.5

x 0.25-0.4 cm. Petiole 0.2-0.3 cm long. Leaf blade ovate, 4.5-6 x 1.5-2.4

cm, index (length/width) 2.3-3; acute to acuminate. Inflorescence ca 1.5 cm

long, 1-flowered. Peduncle porrect, 0.4-0.5 cm, bracts 3, the longest ca 4 mm

long. Floral bracts tubular at the base, ca 3.5 mm, acute. Flowers opening

wide. Pedicel and ovary ca 7,5 mm long, basal node on a ca 2.5 mm-long

stump. Median sepal spreading, ovate, ca 9.5 x 2.5 mm, index ca 3.8; acute,

margins entire, base broadly attached; thick, glabrous. Lateral sepals ca 9.5 x

2.7 mm, index ca 3.5; otherwise as the median sepal. Petals porrect, approx.

lyriform, ca | x 1.5 mm excluding appendages, index 0.6-0.7; margins with 7

appendages more or less regularly spaced along the front margin, base broadly

attached; thin, glabrous; appendages ovate, 1-1.3 x 0.2-0.25 mm, gradually

narrowing into a stalk of 0.5-0.8 mm, obtuse, fleshy, finely papillose. Lip

slightly recurved near the base, elliptic-ovate but constricted half-way, ca 2 x

1 mm, index ca 2 (all without artificial spreading); rounded, margins entire;

thick; adaxially deeply concave proximally and with two distinct, rounded

ridges starting at the base and running parallel, close together up to % of the

length of the lip, slightly over half way along the lip they are clasped between

the margins which converge over the adaxial surface of the lip and almost

meet the ridges, surface convex distally, glabrous; abaxially with a wide,

retuse ridge up to about half-way along the lip, surface coarsely papillose

to vesiculate distally along the sides, leaving a glabrous strip in the middle.

88 Gard. Bull. Singapore 60 (1) 2008

Figure 6. Bulbophyllum brachytriche J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Appendage of

petal; E. Lip, left: abaxial side, right: adaxial side; F. Column and lip, lateral view; G. Anther,

above: abaxial side, below: adaxial side; H. Pollinia, left: two pairs, right: a single pair. All

from SBG-O 5052 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 89

Column ca 1.7 mm long, stigma triangular, without a tooth at its base, column

foot without teeth near the tip. Stelidia porrect, triangular, ca 0.5 mm long,

acute, with an erose upper margin, with a distinct, deltoid, subacute tooth

along the lower margin, and a minute tooth in front of this. Anther abaxially

with a distinct, papillose crest, surface otherwise finely papillose, front

margin drawn out into a retuse beak with approx. entire margins. Pollinia 2,

ellipsoid, with a short, fleshy appendage.

Colour: Leaves suffused with purple. Sepals pale creamy yellow with

numerous red dots. Petals creamy white, appendages greyish green. Lip

white, dark red at the tip.

Habitat & ecology: _Epiphyte in montane forest, on branches of medium-

sized trees. Ait. ca 1200-1400 m. Flowering Feb-May, Jul.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Bulbophyllum vesiculosum J.J. Sm. also has petal appendages of about

1 mm long. It differs from B. brachytriche in having an acute lip with much

larger vesicles, and in having wider sepals (index 1.4-2). Sterile specimens

can be recognized by the pale purplish leaves, which are thinner than usual

in this section.

Bulbophyllum spodotriche J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllis chlororhopalon et B. decatriche ac B. psilorhopalon

appendicibus in petalis in fasciculis duobus ordinatis, inter fasciculos sine

appendice minuta differt.— Typus: Indonesia, Sulawesi, central part, SBG-O

493] (holo, SING). Fig. 7.

Roots most produced close to the base of the rhizome. Rhizome hanging, ca

14 cm long, 2-2.5 mm diam., sections between pseudobulbs 1.2-2 cm long,

bracts persistent. Pseudobulbs well spaced, ovoid to ellipsoid, 0.8-1.8 x 0.4-

width) 2.3-3.5; obtuse to acute. Inflorescence ca 2 cm long, 1|-flowered.

Peduncle porrect, ca 0.8 cm, bracts 3, the longest ca 4 mm long. Floral

bracts tubular at the base, ca 4 mm, acute. Flowers opening wide. Pedicel

and ovary ca 8,5 mm long, basal node on a ca 3.5 mm-long stump. Median

sepal spreading, elliptic-ovate, ca 10 x 4.5 mm, index 2.2-2.3; acute, margins

entire, base broadly attached; thick, glabrous. Lateral sepals as the median

sepal. Petals porrect, approx. lyriform, ca | x 3 mm excluding appendages,

index 0.3-0.4; margins with 11 appendages arranged in two dense clusters

90 Gard. Bull. Singapore 60 (1) 2008

Figure 7. Bulbophyllum spodotriche J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Appendage of

petal; E. Lip, left: adaxial side, right: abaxial side; F. Column and lip, lateral view; G. Anther,

left: abaxial side, right: adaxial side; H. Pollinia, left: a pair, right: a single. All from SBG-O

493] (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 9]

each at one end of the front margin, base broadly attached; thin, glabrous;

appendages ovate-triangular, 5-8.5 x 0.3-0.4 mm, abruptly narrowing into

a stalk of 0.4-0.7 mm, gradually narrowing towards an acute tip, fleshy but

flat, very finely papillose. Lip hardly recurved, elliptic-ovate, ca 3 x 1.8

mm, index 1|.6-1.7 (all without artificial spreading); rounded, margins entire;

thick; adaxially deeply concave proximally and with two distinct, rounded

ridges starting at the base and running parallel, close together over most of

the length of the lip, about half way along the lip they are clasped between the

margins which converge over the adaxial surface of the lip, almost meet the

ridges and then spread out again, adaxial surface convex distally, glabrous:

abaxially with a wide, retuse ridge up to about half-way along the lip, surface

coarsely vesiculate along the sides, leaving a narrow glabrous strip in the

middle. Column ca 2.1 mm long, stigma obovate, without a tooth at its base,

column foot without teeth near the tip. Stelidia porrect, triangular, ca 0.6 mm

long, acute, with a denticulate upper margin, with a distinct, deltoid, subacute

tooth which has the front margin folded inwards along the lower margin,

and a minute tooth in front of this. Anther abaxially with a distinct crest,

surface approx. glabrous, front margin drawn out into a retuse beak with

erose margins. Pollinia 2, ellipsoid, with a short, fleshy appendage.

Colour: Leaves dark green, slightly suffused with purple. Sepals yellowish,

densely spotted and suffused with ochrish red. Petals yellow, appendages

ash-grey near the base, white towards the tip. Lip white near the base, dark

red towards the tip. Column yellow, spotted pale red.

Habitat & ecology: Epiphyte in montane forest. Alt. ca 1000 m. Flowering

Feb, Mar, Jul-Sep, Dec.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Similar to B. chlororhopalon Schitr., B. decatriche J.J. Verm., and B.

psilorhopalon Schitr., but different in having the appendages on the petals

arranged in two clusters; the species mentioned have the appendages either

more or less regularly spaced along the front margin of the petal, or at least |

appendage in between the clusters.

Bulbophyllum stenomeris J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo epicrianthes /abello loriformi apice obtuso petalorum

appendicibus breviore differt. — Typus: Indonesia, Sulawesi, central part,

SBG-O 5220 (holo, SING). Fig. 8.

92 Gard. Bull. Singapore 60 (1) 2008

Roots most produced close to the base of the rhizome. Rhizome hanging,

ca 20 cm long, ca 3 mm diam., sections between pseudobulbs 1.5-2.5 cm

long, bracts persistent. Pseudobulbs well spaced, ovoid, 1-1.4 x 0.6-0.9 cm.

Petiole 0.1-0.3 cm long. Leaf blade elliptic, 3-4 x 1.8-2.3 cm, index (length/

width) 1.6-1.8; obtuse. Inflorescence ca 1.5 cm long, |-flowered. Peduncle

porrect, ca 0.3 cm, bracts (not seen). Floral bracts tubular at the base, ca 3.6

mm, acute. Flowers opening wide. Pedicel and ovary ca 7 mm long, basal

node on a ca 3.2 mm-long stump. Median sepal spreading, ovate, ca 9 x 3

mm, index ca 3; acute, margins entire, base broadly attached; thick, glabrous.

Lateral sepals as the median sepal. Petals porrect, approx. lyriform, ca |

x 2 mm excluding appendages, index ca 0.5; with 5 appendages more or

less regularly spaced along the front margin, base broadly attached; thin,

glabrous; appendages linear, 3.5-5 x 0.3-0.5 mm, abruptly narrowing into a

stalk of 0.5-1 mm, slightly narrowing towards an obtuse tip, fleshy, very finely

papillose-hirsute. Lip hardly recurved, ovate, ca 5 x 0.9 mm, index 5.5-5.6 (all

without artificial spreading); obtuse, margins entire; thick; adaxially deeply

_ concave proximally and with two distinct, rounded ridges starting at the base,

immediately converging and then running parallel, close together up to 4-'A

of the length of the lip, half-way along their length they are clasped between

the margins which converge over the adaxial surface of the lip and then fuse

to them, surface convex distally, very finely papillose; abaxially with a wide,

retuse ridge up to 4-’4 of the length of the lip, surface glabrous but with a

patch of coarse papillae towards the side at about 4 of the length of the lip,

leaving a glabrous strip in the middle. Column ca 2 mm long, stigma slitlike,

without a tooth at its base, column foot without teeth near the tip. Stelidia

porrect, deltoid, ca 0.4 mm long, acute, with a denticulate upper margin, with

a distinct, deltoid, subacute tooth which has the front margin folded inwards

along the lower margin, and a minute tooth in front of this. Anther abaxially

with a distinct, papillose crest, surface approx. glabrous, front margin drawn

out into a retuse, papillose beak with entire margins. Pollinia 4, all drop-

shaped, the inner about half as long as the outer, without appendage.

Colour: Leaves medium green, not suffused with purple. Sepals pale yellow,

with large, dark red spots near the base, smaller spots towards the tip and

the margins. Petals pale yellow, appendages dull brownish red. Lip dark red.

Column yellowish, stained with pale red.

Habitat & ecology: Epiphyte in remnant of dry, low forest on serpentinite

soil. Alt. 100- 400 m. Flowering Feb, Mar.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

New Species of Bulbophyllum (Orchidaceae) 93

Figure 8. Bulbophyllum stenomeris J.J. Verm. & P.O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Appendage of petal; E.

Lip, left: adaxial side, right: abaxial side; F. Column and lip, lateral view; G. Anther, left:

adaxial side, right: abaxial side; H. Pollinia, left: a pair, right: a single. All from SBG-O 5220

(spirit sample).

94 Gard. Bull. Singapore 60 (1) 2008

Notes: The only other species of sect. Epicranthes with a narrow, almost

glabrous top part of the lip is B. epicrianthes Hook.f. In this species, however,

the top part of the lip is ovate and acute, and the lip is relatively larger (about

twice as long as the petal appendages).

Bulbophyllum xanthomelanon J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo flavofimbriato stelidiis elongatioribus ala deltoidea acuta

inferne secus marginem inferiorem, antice dente minuto differt. — Typus:

Indonesia, Sulawesi, central part, SBG-O 5810 (holo, SING). Fig. 9.

Roots most produced close to the base of the rhizome. Rhizome hanging, ca

35 cm long, 2.5-3 mm diam., sections between pseudobulbs 1-2.2 cm long,

bracts persistent. Pseudobulbs well spaced, ellipsoid to obovoid, 1-1.5 x 0.5-

index (length/width) 1.8-2.3; acute. Inflorescence ca 2 cm long, 1-flowered.

Peduncle porrect, 0.35-0.6 cm, bracts 3, the longest ca 5 mm long. Floral

_ bracts tubular at the base, 3-3.8 mm, acute. Flowers opening wide. Pedicel

and ovary 9-9.5 mm long, basal node on a 3-4 mm-long stump. Median sepal

spreading, ovate, ca 8 x 4.5 mm, index 1.7-1.8; acuminate, margins entire,

base broadly attached; thick, glabrous. Lateral sepals as the median sepal.

Petals porrect, approx. lyriform, ca 1.5 x 2 mm excluding appendages, index

0.7-0.8; margins with 6 appendages more or less regularly spaced along the

front margin, base broadly attached; thin, glabrous; appendages ovate, |.3-3.8

x 0.3-0.35 mm, gradually narrowing into a stalk of 0.3-0.8 mm, obtuse, fleshy,

finely papillose. Lip hardly recurved, ovate, ca 2.7 x 1.7 mm, index 1.5-1.6

(all without artificial spreading); subacute, margins entire; thick; adaxially

slightly concave proximally and with two distinct, rounded ridges starting at

the base, and running parallel, close together up to ca half way the length of

the lip, there they are fused to the margins which converge over the adaxial

surface of the lip and then run parallel over another 3 of the length of the lip,

surface convex distally, glabrous; abaxially with a wide, retuse ridge near

the base, surface coarsely papillose or vesiculate except for a glabrous strip

around the midvein. Column ca 2.5 mm long, stigma obovate, without a tooth

at its base, column foot without teeth near the tip. Stelidia porrect, triangular,

ca 0.6 mm long, subacute, with a denticulate upper margin, with a distinct,

deltoid, subacute tooth along the lower margin, and a minute tooth in front

of this. Anther abaxially with a distinct crest, surface approx. glabrous, front

margin drawn out into a retuse beak with denticulate margins. Pollinia 4, all

drop-shaped, the inner about half as long as the outer, without appendage.

New Species of Bulbophyllum (Orchidaceae)

Figure 9. Bulbophyllum xanthomelanon J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, above: two pairs, below: a single pair. All from SBG-O 4902 (spirit

sample).

96 Gard. Bull. Singapore 60 (1) 2008

Colour: Leaves green, suffused with some red. Sepals yellow. Petals whitish

yellow, slightly suffused with pink; appendages blackish. Lip and tip column

blackish red.

Habitat & ecology: Epiphyte in montane forest. Alt. ca 1200 m. Flowering

all year round.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. xanthomelanon is most similar to B. flavofimbriatum J.J. Sm. It

differs in having more elongated stelidia, with a deltoid, acute wing along the

lower margin that is positioned further away from their tip. In front of this

wing a second, minute tooth is present, lacking in B. flavofimbriatum.

Bulbophyllum sect. Hybochilus Schltr.

Bulbophyllum sect. Hybochilus Schltr., Feddes Repert. 10 (1912) 96; Schltr.,

Feddes Repert. Beth. 1 (1913) 823.—TYPE SPECIES: Bulbophyllum

masarangicum Schltr.

As Schlechter (1913) says, this section differs only from sect. Monanthes (=

sect. Polyblepharon, see below) in having free, not fused, lateral sepals.

Bulbophyllum acutilobum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllis decurrentilobo et B. depresso petalis rotundatis, labelli

lobis lateralibus acutis differt. — Typus: Indonesia, Sulawesi, central part,

SBG-O 4912 (holo, SING). Fig. 10.

Roots below the pseudobulbs. Rhizome creeping, 0.8-1 mm diam., sections

between pseudobulbs 0.3-0.5 cm long, bracts not persistent. Pseudobulbs

spaced, ovoid, 0.3-0.5 x 0.3-0.4 cm. Petiole up to 0.1 cm long. Leaf blade

elliptic to ovate, ca 1.8 x 0.5-0.6 cm, index (length/width) 3-3.6; acuminate.

Inflorescence ca | cm long, 1-flowered. Peduncle patent, ca 0.45 cm, bracts

ca 2, the longest ca 1.7 mm long. Floral bract tubular, ca 2 mm, acuminate.

Flowers not fully opening. Pedicel and ovary ca 1.2 mm long, basal node

coinciding with the bract attachment. Median sepal recurved, triangular, ca 4

x 1.5 mm, index 2.6-2.7; acute, margins entire, base broadly attached; rather

thick, thickened towards the tip, glabrous. Lateral sepals free, oblique, ca 4 x

1.9 mm, index ca 2.1; otherwise as the median sepal. Petals porrect, elliptic-

obovate, ca 1.3 x 0.5 mm, index ca 2.6; rounded, margins erose distally,

Flower analysis, from left to nght: median sepal, petal, latera

side, right: abaxial side; E. Column and lip, lateral view

abaxial side: G. Pollinia, left: two pairs, nght: a singl

sample).

98 Gard. Bull. Singapore 60 (1) 2008

base broadly attached; thin, surface glabrous. Lip recurved about half way,

general outline approx. ovate, ca 1.9 x 0.9 mm, index ca 2.1 (all without

artificial spreading), margins entire; glabrous, three-lobed; midlobe approx.

ovate, slightly widened near the tip, truncate-rounded, thick; adaxially with

a distinct, retrorse, longitudinal, erect, triangular, obtuse tooth near the base,

surface towards the tip slightly convex; abaxially without a median ridge

near the base; sidelobes attached along the basal 2/5 of the length of the

lip, antrorse, oblique, obovate, acute, thin. Column ca 0.8 mm long, stigma

approx. circular, without keels inside, without teeth at its base, column foot

without teeth. Stelidia triangular, ca 0.4 mm long, obtuse. Anther abaxially

with a distinct, rounded crest, surface papillose, front margin drawn out into

a triangular beak. Pollinia 4, the inner slightly more than half as long as the

outer; obovoid, flattened, the outer obovoid; no appendages present.

Colour: Sepals whitish near the base, pale red with darker red veins up to

half-way their length, yellowish green towards the tip. Petals translucent

white with a red midvein. Lip purplish red.

Habitat & ecology: Found as an epiphyte in riverine forest. Alt. ca 1000 m.

Flowering Apr-Jun, Sep.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Bulbophyllum decurrentilobum J.J. Verm. & P. O’Byrne, and B.

depressum King and Pantling, are most similar. Both, however, have acute or

acuminate petals and rounded lateral lobes on the lip.

Bulbophyllum auritum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo masarangico /abello lobis lateralibus nonnihil antrorsis, lobi

mediani in parte distali sine carina differt. — Typus: Indonesia, Sulawesi,

central part, SBG-O 5740 (holo, SING). Fig. 11.

Roots below the pseudobulbs. Rhizome creeping, 1-1.5 mm diam., sections

between pseudobulbs 0.4-0.7 cm long, bracts not persistent. Pseudobulbs

well spaced, ovoid, 0.4-0.7 x 0.4-0.6 cm. Petiole up to 0.5 cm long. Leaf

blade ovate, 0.7-0.8 x 0.6-0.7 cm, index (length/width) 1.1-1.3; acute.

Inflorescence 0.8-1 cm long, 1-flowered. Peduncle patent, 0.4-0.5 cm,

bracts ca 3, the longest ca 1.5 mm long. Floral bract tubular, ca 1 mm, acute.

Flowers not fully opening. Pedicel and ovary ca | mm long, basal node

coinciding with the bract attachment. Median sepal recurved, ovate, ca 3.2

x 1.3 mm, index 1.7-1.8; acute, margins entire, base rather broadly attached;

New Species of Bulbophyllum (Orchidaceae) 99

Figure 11. Bu/bophyllum auritum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, left: a single pair, right: two pairs. All from SBG-O 5740 (spirit

sample).

100 Gard. Bull. Singapore 60 (1) 2008

rather thick, thickened towards the tip, glabrous. Lateral sepals free, oblique,

ca 3.3 x 1.5 mm, index ca 2.2; otherwise as the median sepal. Petals porrect,

obovate, ca 1.2 x 0.7 mm, index ca 1.7; shortly acuminate, margins erose

distally, base rather broadly attached; thin, surface glabrous. Lip slightly

recurved, general outline elliptic with a long drawn-out top part, ca 2.4 x 0.7

mm, index ca 3.4 (all without artificial spreading; when spread subhastate),

three-lobed; midlobe approx. linear, slightly widened towards the tip,

rounded, margins entire; thick; adaxially with a distinct, longitudinal, erect,

subtriangular, obtuse tooth near the base, surface elsewhere slightly convex,

very finely papillose locally, abaxially without a median ridge near the base,

surface approx. glabrous; sidelobes attached along the basal 2/5 of the length

of the lip, antrorse, oblique, ovate, rounded, front margin somewhat erose;

thin, surface glabrous. Column ca 0.7 mm long, stigma elliptic, without keels

inside, without teeth at its base, column foot with a distinct, rounded swelling

just above the ligament. Stelidia triangular, ca 0.4 mm long, obtuse, with a very

distinct, antrorse, semi-elliptic tooth protruding beyond the tip of the stelidia

_ along the lower margin. Anther abaxially with a distinct, broadly rounded

crest, surface approx. glabrous, front margin drawn out into a triangular beak.

Pollinia 4, the inner ca half as long as the outer; obovoid, flattened, the outer

ellipsoid; no appendages present.

Colour: Sepals yellowish towards the base, yellow towards the tip, in between

with a large purplish red patch around the midvein; midvein itself a darker

purplish red. Petals translucent white, midvein and top purplish red. Lip

blackish purple.

Habitat & ecology: Found as an epiphyte in isolated trees on a wind-swept

plateau. Alt. ca 1700 m. Flowering Apr.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Bulbophlylum masarangicum Schitr. is most similar to the new species,

but has a lip with a distinct median ridge over the top part of the midlobe, and

semi-elliptic, well-rounded sidelobes.

Bulbophyllum sect. Intervallatae Ridl.

Bulbophyllum sect. Intervallatae Ridl., J. Linn. Soc., Bot. 31 (1896) 276. —

LECTOTYPE SPECIES: Bulbophyllum tardeflorens Ridl. (= B.

attenuatum Rolfe).

New Species of Bulbophyllum (Orchidaceae) 10]

The two new species described below fit best into sect. /ntervallatae because

of their distichous inflorescence with flowers opening in succession. They are

most similar to a small number of species traditionally included in a separate

section, sect. Hymenobractea, a section to be merged with sect. /ntervallatae

(de Witte, in prep.).

The flower structure and the lip shape of both species are reminiscent

of S. American species such as B. micropetalum Barb. Rodr., and B. regnellii

Rchb.f. However, a close phylogenetic relationship between the Sulawesi and

the S. American species is unlikely.

Bulbophyllum allotrion J.J. Verm. & P. O’ Byrne, sp. nov.

Bulbophyllum allotrion J.J. Verm. & P. O'Byrne, inter congeneres asiaticos

euronotos inflorescentia laxissima necnon labello hastato singularis.

— Typus: Indonesia, Sulawesi, central part, SBG-O 5072 (holo, SING).

Fig. 12.

Roots mainly below the pseudobulbs. Rhizome creeping, 2-3 mm diam.,

sections between pseudobulbs 0.3-0.8 cm long, bracts not persistent.

Pseudobulbs close together, ovate, 1.5-2.8 x 0.7-1 cm, without angles.

Petiole 1.5-2.7 cm long. Leaf blade elliptic, 7-13 x 1.8-2.5 cm, index (length,

width) 3-6.9; acute. Inflorescence patent, a very lax raceme, 23-50 cm long,

up to 15-flowered. Peduncle 21.5-30 cm, bracts 6-7, the longest 5-8 mm long.

Rhachis not thickened, up to ca 20 cm long. Floral bracts 4—7 mm, acute.

Flowers not resupinate, distichous, fully opening, 1-2 at the time. Pedicel and

ovary 12—17 mm long, basal node on a ca 3 mm long stump; basal half of

the stump fused to the rhachis, top half sticking out perpendicularly. Median

sepal recurved, subtriangular, ca 8 x 5.5 mm, index 1.4-1.5; acute-acuminate,

margins entire, base narrowly attached; rather thin, glabrous. Lateral sepals

oblique, ovate-triangular, ca 12 x 4.5 mm, index 2.6-2.7, otherwise as the

median sepal. Petals aaprox. porrect, triangular with a distinctly widened

base, ca 4.8 x 4.2 mm, index 1.1-1.2; acute, margins entire, base broadly

attached; rather thin, glabrous. Lip slightly recurved about half-way, general

outline approx. hastate, ca 7.5 x 3.4 mm, index ca 2.2—2.4 (all without

artificial spreading); acute, margins entire; locally slightly and finely papillose;

3-lobed; midlobe approx. ovate, thick, adaxially slightly concave proximally,

with a rounded, not sharply delineated callus about half-way along the length

of the lip, slightly convex distally; abaxially with a wide, truncate ridge up

to about 4 of the length of the lip, distally slightly concave; lateral lobes

attached to the middle 4 of the length of the lip, antrorse, approx. seam-

like, gradually widening in the back, obtuse to subacute in front, rather thick.

Column ca 2.4 mm long, stigma elliptic, with 2 keels inside, without a tooth

102 Gard. Bull. Singapore 60 (1) 2008

Figure 12. Bulbophyllum allotrion J.J. Verm. & P.O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Column with anther and hamulus;

G. Anther, left: adaxial side, right: abaxial side; H. Pollinia, left: two pairs with appendage,

right: a single pair. All from SBG-O 5072 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 103

at its base, column foot elongated, without teeth near its tip. Stelidia ca |

mm long, triangular, acute, with a rather inconspicuous, deltoid, rounded to

obtuse wing along the lower margin. Anther abaxially with a high, triangular,

dorsoventrally flattened crest largely overtopping the front margin near the

tip and an obtuse ridge near the base, surface slightly papillose, front margin

not drawn out. Pollinia 4, the inner about as long as the outer, all ellipsoid-

ovoid; a long fleshy appendage present, fitting in between the wings along the

lower margin of the stelidia.

Colour: Plant medium green. Sepals and petals yellowish green with dark red

veins. Lip: basal part, including lateral lobes, pale greenish, locally suffused

with some pale red; top part yellow. Column yellowish green. Anther

yellow.

Habitat & ecology: Understory epiphyte (sometimes on thickly moss-clad

tree trunks near the forest floor) in mossy montane forest. Alt.1200- 1400 m.

Flowering Sep-Nov.

Distribution: I(NDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Uniquely identified among SE Asiatic Bulbophyllum by its very lax

inflorescence combined with the hastate lip.

Bulbophyllum cymbidioides J.J. Verm. & P. O° Bye, sp. nov.

A Bulbophyllo allotrion /abelli lobis lateralibus non antrorsis, antherae

operculo sine crista differt. — Typus: Indonesia, Sulawesi, Southwest part,

SBG-O 5028 (holo, SING). Fig. 13.

Roots mainly below the pseudobulbs. Rhizome creeping, 1.2-1.5 mm

diam., sections between pseudobulbs 0.3-0.6 cm long, bracts not persistent.

Pseudobulbs close together, ovate, 1.1-2.5 x 0.6-0.9 cm, without angles.

Petiole 0.5-1.4 cm long. Leaf blade elliptic to ovate, 2.9-8.2 x 0.8-1.2 cm,

index (length/width) 3.6-7.5; acute. Inflorescence patent, a very lax raceme,

7-18 cm long, up to 4-flowered. Peduncle 5.7-11 cm, bracts 4-5, the longest

3.5-4.5 mm long. Rhachis not thickened, up to ca 5.7 cm long. Floral bracts ca

4mm, acute. Flowers not resupinate, distichous, fully opening, 1-2 at the time.

Pedicel and ovary 12-14 mm long, basal node on a ca 5 mm long stump; basal

half of the stump fused to the rhachis, top half sticking out obliquely. Median

sepal recurved, subtriangular, ca 9.5 x 6.5 mm, index 1.4-1.5; acuminate,

margins entire, base narrowly attached; rather thin, glabrous. Lateral sepals

oblique, ovate-triangular, ca 12 x 7 mm, index ca 1.7, otherwise as the median

104 Gard. Bull. Singapore 60 (1) 2008

-- ———

--. om

SQs.------~

(ieee ee

Figure 13. Bulbophyllum cymbidioides J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, left: a single pair, right: two pairs, with appendage. — All from

SBG-O 5028 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 10

sepal. Petals approx. porrect, obliquely deltoid with a somewhat widened

base, ca 5 x 6.2 mm, index ca 0.8; subacute, margins entire, base broadly

attached; rather thin, glabrous. Lip recurved, general outline ovate, ca 7.2 x

4 mm, index ca 1.8 (all without artificial spreading); obtuse, margins entire;

glabrous; 3-lobed; midlobe approx. ovate, rather thick, adaxially concave

proximally, with a transverse, antrorse deltoid, obtuse, thick median tooth

near the base, slightly convex distally; abaxially with a wide, truncate ridge

up to about 4 of the length of the lip, distally slightly concave; lateral lobes

attached to the proximal of the length of the lip, semi-circular. Column ca 3

mm long, stigma ovate, with 2 keels inside, without a tooth at its base, column

foot elongated, widened but without teeth near its tip. Stelidia ca 1 mm long,

triangular, acute, with a rather inconspicuous, deltoid, rounded to obtuse wing

along the lower margin. Anther abaxially virtually without a crest, surface

glabrous, front margin not drawn out. Pollinia 4, the inner slightly shorter

than the outer and distinctly flattened; a long fleshy appendage present, fitting

in between the wings along the lower margin of the stelidia.

Colour: Median sepal translucent greenish, veins dull red, margins white.

Lateral sepals whitish, greenish at the tip, with some dull red markings

following the veins near the base. Lip: top part white, lateral lobes greenish,

basal half spotted purple along the midvein and near its base

Habitat & ecology: Remnants of very wet, low and open montane woodland.

Understory epiphyte on bushes, but also in the crowns of emergent trees. Alt.

1600-2200 m. Flowering Jan-Apr, Oct.

Distribution: INDONESIA. Sulawesi, southwest part (1 specimen seen).

Notes: Differs from B. allotrion in the not-antrorse sidelobes of the lip, and

the lack of a crest on the anther cap.

Bulbophyllum sect. Leopardinae Benth. & Hook.f.

Bulbophyllum sect. Leopardinae Benth. & Hook.f., Gen. Pl. 3 (1883)

503. — TYPE SPECIES (a lit. reference given only): Dendrobium

leopardinum Wall. [= Bulbophyllum leopardinum (Wall.) Lindl.]

Bulbophyllumsect. Beccarianae Pfitz. in Engler & Prantl, Nat. Pflanzenfam., ed.

2, 6 (1889) 179. -TYPE SPECIES: Bulbophyllum (“Bolbophyllum’)

beccarii Rchb.f.

106 Gard. Bull. Singapore 60 (1) 2008

Bulbophyllum sect. Pahudia Schltr., Feddes Repert. 10 (1911) 93; Garay,

Hamer & Siegerist, Nord. J. Bot. 14 (1994) 630. - LECTOTYPE

SPECIES (designated by Garay, Hamer & Siegerist): Cirrhopetalum

pahudii De Vriese [= B. pahudii (De Vriese) Rchb.f. ].

Bulbophyllum sect. Pahudiella Garay, Hamer & Siegerist, Nord. J. Bot. 14

(1994) 629. — TYPE SPECIES: Bulbophyllum subumbellatum Rid.

Here, sect. Leopardinae is segregated from sect. Sestochilus (Breda) Benth.

& Hook.f., and sect. Stenochilus J.J. Sm., to accommodate a series of species

which have the node between the peduncle and the pedicel more or less

coinciding with its bract, or have the distance between this node and its bract not

exceeding 1.5 times the diameter of the pedicel at its base. In sect. Sestochilus

as well as sect. Stenochilus the distance between the node and its bract equals

or exceeds 2 times the diameter of the pedicel at its base. A selection of species

included in sect. Leopardinae can be found 1n Vermeulen (1991), under sect.

Sestochilus, but with the exclusion of the following: B. apheles J.J. Verm.,

_B. dearei Rchb.f., B. /obbii Lindl. (all sect. Sestochilus), and B. cheiri Lindl.,

B. macranthum Lindl., and B. patens King (all sect. Stenochilus). A revision

including the three sections mentioned is in preparation.

Among our Sulawesi material is one new species of sect.

Leopardinae:

Bulbophyllum deviantiae J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllis incisilabro et B. rugoso /abelli parte apicali distincte

papillosa, et a B. maculoso ovario jugis lateralibus distincte sinuosis

differt.— Typus: Indonesia, Sulawesi, central part, SBG-O 5057 (holo, SING).

Fig. 14.

Roots mainly below the pseudobulbs. Rhizome 2-2.5 mm diam., sections

between pseudobulbs 0.8-1.2 cm long, bract fibres hardly persistent.

Pseudobulbs loosely clustered, ovoid to almost cylindrical, 1-2.1 x 0.4-0.7

cm. Petiole 0.2-2.2 cm. Leaf blade elliptic to ovate, 4.5-8.6 x 1-1.4 cm, index

(length/width) 3.7-6.6; acute. Inflorescence erect to patent, ca 4 cm long,

1-flowered. Peduncle ca | cm, bracts ca 2, the longest 6-7 mm long. Floral

bracts elliptic to ovate, ca 9 x 6 mm, acute. Flowers resupinate, opening

wide. Pedicel and ovary ca 30 mm long, basal node approx. coinciding with

the attachment of the floral bract; ovary ribs with the crests sharply angular,

all distinctly sinuous near the flower except the rib opposing the median

sepal. Sepal recurved to spreading, elliptic-ovate, ca 18 x 7.5 mm, index

2.2-2.3; subacute, margins entire, base rather broadly attached; rather thick;

glabrous. Lateral sepals free, ovate-triangular, ca 18 x 8 mm, index 2.2-2.3,

New Species of Bulbophyllum (Orchidaceae) 107

==22n

Figure 14. Bulbophyllum deviantiae J.J. Verm. & P.O’ Bye. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side, below:

abaxial side. All from SBG-O 5057 (spirit sample).

108 Gard. Bull. Singapore 60 (1) 2008

acute; otherwise as the median sepal. Petals spreading with incurved tips,

ovate-oblong, ca 18 x 4 mm, index ca 4.5; subacute, margins slightly erose

distally, base broadly attached; rather thick, glabrous. Lip recurved towards

the base and towards the tip, ovate-oblong, ca 6.5 x 5 mm, index ca 1.3

(all without artificial spreading), rounded, margins entire proximally, from

about 4 of the length of the lip increasingly erose, thick; adaxially with a

distinct median furrow over its entire length, concave and glabrous towards

the base except for a irregularly verrucate strip bordering the median furrow,

convex and coarsely and distinctly verrucate towards the tip; abaxially with

a retuse median ridge over most of its length, surface glabrous except for a

finely papillose patch in the centre. Column 3 mm long, stigma inside with

3 keels, at its base without teeth, column foot not widened at the tip. Stelidia

inconspicuous, deltoid, ca 0.05 mm, rounded, with a distinct, patent, recurved

, Strap-shaped, obliquely truncate tooth along the lower margin, near the top

of the column. Anther abaxially colliculate and with a rounded crest; front

margin somewhat papillose. Pollinia (not seen).

Colours: Young leaves with numerous small red spots. Sepals and petals

yellow the sepals densely spotted and stained with brownish red, the petals

less densely so. Lip orange yellow, adaxially bright red towards the tip.

Column yellow, stained with some red.

Habitat & ecology: Understory epiphyte in montane forest. Alt.1200-1400

m. Flowering Jan-May, Aug, Sep.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: This is part of a small group of species around B. membranifolium

Hook.f.; it shares the spreading (not fused or at least contiguous) lateral sepals

with B. incisilabrum J.J. Verm. & P O'Byrne, B. rugosum Ridl., and a taxon

of doubtful status known as B. maculosum Ames. It differs from the first two

in having a distinctly papillose top part of the lip, from the latter in having

distinctly sinuous lateral juga on the ovary. The plants that we have seen of B.

deviantiae are smaller than any other of the species mentioned.

Named after Devianti, daughter of Tampang, Rantepao, Sulawesi.

Bulbophyllum sect. Macrocaulia (Bl.) Aver.

(= sect. Monilibulbus)

Diphyes sect. Macrocaulia Bl., Bijdr. Fl. Ned. Ind. (1825) 318.— Bulbophyllum

New Species of Bulbophyllum (Orchidaceae) 109

sect. Macrocaulia (Bl.) Aver., Identification Guide to Vietnamese

Orchids (1994) 279. - LECTOTYPE SPECIES (here designated):

Diphyes ovalifolia B\. [Bulbophyllum ovalifolium (B1.) Lindl.].

Diphyes sect. Diasperia Bl., Bijdr. Fl. Ned. Ind. (1825) 317. -LECTOTYPE

SPECIES (here designated): Diphyes cernua BI. [= B. cernuum (B1.)

Lindl.].

Bulbophyllum sect. Odoardiana Pfitz. in Engler & Prantl, Nat. Pflanzenfam..,

ed. 2, 6 (1889) 179. — TYPE SPECIES: Bulbophyllum odoardii

Rchb.f. & Pfitz.

Bulbophyllum sect. Monilibulbus J.J. Sm., Bull. Jard. Bot. Buitenzorg. Ser.

2, 13 (1914) 33. - LECTOTYPE SPECIES: Diphyes inaequalis BI.

[Bulbophyllum inaequale (Bl.) Lindl.].

Bulbophyllum torajarum J.J. Verm. & P. O’Byrne, sp. nov.

A Bulbophyllis catenario ef B. carunculilabrum /Jabelli forma sine

explicatione artificiali: ad apicem gradatim coarctatus sine constrictione

distincta pare ubi margines convenent supra paginam abaxialem, sepalo

mediano triangulari differt. — Typus: Indonesia, Central Sulawesi, SBG-O

3205 (holo, SING). Fig. 15.

Roots below the pseudobulbs. Rhizome creeping, 0.8-1 mm diam., sections

between pseudobulbs 0.4-0.7 cm long, bracts not persistent. Pseudobulbs

ovoid, basal half prostrate on and fused to the rhizome, so that the new

pseudobulbs arise c. half-way up the old, 0.5-0.7 x 0.2-0.4 cm. Petiole 3-4

mm long. Leaf blade elliptic, 0.8-1.5 x 0.25-0.7 cm, index (length/width) 2.1-

3.7; obtuse. Inflorescence 3-4 cm long, 1-flowered. Peduncle erect to patent,

1.8-2.8 cm, bracts 2, the longest ca 1.8 mm long. Floral bracts tubular, 1.2-1.8

mm, acute. Flowers fully opening. Pedicel and ovary 6-8 mm long, basal

node on a 0.8—1 mm-long stump. Median sepal approx. porrect or somewhat

recurved, triangular, ca 4.5 x 1.4 mm, index ca 3.2; acute-acuminate, margins

entire, finely papillose-ciliolate, base widely attached: rather thin, adaxially

glabrous, abaxially with papillose-ciliolate veins. Lateral sepals recurved,

free, oblique, approx. elliptic, ca 6.7 x 3.1 mm, index 2.1-2.2; acute; rather

thick, otherwise as the median sepal. Petals porrect, elliptic-obovate, ca

1.7 x 0.9 mm, index 1.8-1.9; subacute, margins entire, base rather narrowly

attached; thin, glabrous. Lip recurved near the base, ovate in general outline

with the margins folded back over the abaxial side and touching one another,

ca 3 x 1.8 mm, index 1.6-1.7 (all without artificial spreading), slightly

truncated, margins somewhat erose; thick; adaxially concave near the base,

with 2 short ridges starting near the margins at about 4 of the length of the lip,

converging and continuing up to about 2 the length of the lip, their distal end

110 Gard. Bull. Singapore 60 (1) 2008

Figure 15. Bulbophyllum torajaraum J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, above: abaxial side,

below: adaxial side; G. Pollinia, above: a single pair, below: two pairs. All from SBG-O

3205 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 111

clasping a small, rounded callus, lip surface convex towards the tip and with

large, conical verrucae, abaxially with a rounded median ridge near the base,

surface glabrous. Column ca 1.2 mm long, stigma without a tooth at its base,

column foot without teeth. Stelidia narrowly triangular-subulate, ca 0.9 mm

long, acute. Anther abaxially with a widely rounded crest, surface papillose,

front margin not drawn out. Pollinia 4, the inner ca % as long as as the outer,

flattened, all ellipsoid-ovoid; no appendages present.

Colour: Median sepal translucent yellowish with orange veins. Lateral sepals

orange towards the base, yellow towards the tip. Petals yellow with a darker

midvein. Lip orange red towards the base, orange towards the tip.

Habitat & ecology: Understorey epiphyte in montane forest, at 2000-2500

m alt. Flowering Mar-May, Jul-Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: In the general flower shape B. torajarum is most similar to B. catenarium

Ridl. and B. carunculaelabrum Carr. All three have a lip that, when spread

out, widens abruptly at 4 —'% of its length. The margins are folded backwards

and meeting over the adaxial surface. B. torajarum differs from the other two

in the shape of the lip without spreading: it narrows gradually towards the tip,

without a distinct constriction at the level where the margins meet over the

abaxial surface, as in the other two. It furthermore differs from both in having

a triangular (not an elliptic) median sepal.

Bulbophyllum sect. Minutissima Pfitz.

(= sect. Nematorhizis)

Bulbophyllumsect. Minutissima Pfitz. in Engler & Prantl, Nat. Pflanzenfam., ed.

2, 6 (1889) 180. — TYPE SPECIES: Bulbophyllum (“Bolbophyllum’)

minutissimum V. Muell.

Bulbophyllum sect. Nematorhizis Schltr., Feddes Repert. Beih. 1 (1913) 701

& 790. - LECTOTYPE SPECIES (here designated): Bulbophyllum

nematorhizis Schltr.

Bulbophyllum insipidum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo perpusillo foliis 3.2—4.8 cm longis a B. keekee labello ad

apicem sine callo minuto differt. — Typus: Indonesia, Sulawesi, central part,

SBG-O 4804 (holo, SING). Fig. 16.

2 Gard. Bull. Singapore 60 (1) 2008

Roots along the entire rhizome. Rhizome creeping, 2-2.5 mm diam., sections

between pseudobulbs 0.3-0.5 cm long, bracts little persistent. Pseudobulbs

close together, ellipsoid to ovoid, 0.7-1.2 x 0.4-0.7 cm. Petiole 0.4-0.8 cm

long. Leaf blade elliptic to ovate, 3.2-4.8 x 0.6-1.1 cm, index (length/width)

3.6-5.3; acute. Inflorescence 2.2-3.5 cm long, 1-flowered. Peduncle patent,

1.5-1.7 cm, bracts ca 3, the longest ca 3.5 mm long. Floral bract tubular, ca

3 mm, acute. Flowers hardly opening. Pedicel and ovary 13-14 mm long,

basal node on a ca 1 mm long stump. Median sepal porrect, ovate, ca 2.7 x

1.5 mm, index ca 1.8; subacute, margins entire, base broadly attached; rather

thick, glabrous. Lateral sepals free, oblique, ovate-triangular, ca 2.7 x 2.7 mm,

index ca 1; otherwise as the median sepal. Petals porrect, elliptic, ca 2 x 0.6

mm, index ca 3.3-3.4; obtuse, margins entire; base rather broadly attached;

thin, surface approx. glabrous. Lip strongly recurved about half-way, general

outline obovate-subhastate, ca 2 x 1.2 mm, index 1.6-1.7 (all without artificial

spreading), obtuse, margins approx. entire, thin proximally, thick distally,

surface slightly papillose distally; adaxially somewhat concave up to % of its

_ length, with two slight, obtuse ridges near the margin slightly over half-way

its length, top 3 convex; abaxially with a slight, rounded ridge up to half-way

along the length of the lip. Column ca 1.5 mm long, stigma subtriangular,

without keels inside, without teeth at its base, column foot tapering towards

its tip, without lateral teeth. Stelidia triangular, ca 0.7 mm long, subacute.

Anther abaxially with a rounded crest, surface glabrous, front margin drawn

out, concave, obtuse. Pollinia 4, ovoid, the inner about % as long as the outer,

distinctly flattened; no appendages present.

Colour: Sepals whitish, stained with pale pinkish red, yellowish towards the

tip.

Habitat & ecology: Found as an understory epiphyte in secondary forest near

a stream. Alt. ca 1450 m. Flowering Apr-Sep.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: The general shape of the flowers, including the tapering column foot,

fits well in sect. Minutissima. The densely crowded pseudobulbs are very

unusual in the section. B. perpusillum Ridl., from Sarawak, grows in similar

tufts, but is a much smaller plant of slightly over one cm high, with acute-

acuminate sepals and petals and an acute lip. B. keekee Halle, from New

Caledonia, has similarly shaped flowers, but with a suborbicular lip with a

small callus near the tip.

New Species of Bulbophy llum (Orchidaceae) ] | 3

Figure 16. Bulbophyllum insipidum J.J. Verm. & P.O’ Byme. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip: D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, nght: abaxial

side; G. Pollinia, left: two pairs, right: a single pair. All from SBG-O 4804 (spirit sample).

114 Gard. Bull. Singapore 60 (1) 2008

Bulbophyllum sect. Monanthaparva Ridl.

Bulbophyllum sect. Monanthaparva Ridl., J. Linn. Soc., Bot., 32 (1896)

269; Van Royen, Alpine Fl. New Guinea 2 (1979) 207, sub sect.

Schlechteria, see note below. — LECTOTYPE SPECIES (here

designated): B. striatellum Ridl.

Bulbophyllum sect. Scyphosepalum Schlitr., Feddes Repert. Beih. 1 (1913)

701, 793; Van Royen, Alpine Fl. New Guinea 2 (1979) 217. —

LECTOTYPE SPECIES (here designated): Bulbophyllum nuruanum

Schltr.

Van Royen (l.c.) synonymised sect. Monanthaparva with sect. Schlechteria

(= sect. Micromonanthe, here synonymised with sect. Polymeres, see below),

without indicating a lectotype from among the species listed by Ridley with

the original description of the section. These species would now be included

in various sections; we choose B. striatellum Ridl., as a lectotype so that the

_ sectional name can be used for a group of species identified below. Doing

so, the taxonomic contents of the section differs from that implied by Van

Royen.

The species included here are most similar to sect. Hybochilus

Schltr, but differ in having the basal node of the pedicel 0.4-4 mm above the

attachment of the floral bract (level with the attachment of the floral bract in

sect. Hybochilus). Additional differences include: pedicel usually elongated

(versus short to virtually absent); lip entire (versus obscurely or distinctly

three lobed); median ridge on the lip absent or inconspicuous (versus usually

distinct, often with an erect tooth proximally).

Marked similarity also exists between sect. Monanthaparva and sect.

Polymeres (see below), but sect. Monanthaparva lacks the widened column

foot (usually with a protrusion on each side of the ligament) that characterizes

sect. Polymeres. The delimitation to sect. Hybochilus may still need some

adjustments, but the following species are included in sect. Monanthaparva,

with the species of which material has been checked for the diagnostic

character marked with an asterisk:

B. ascochiloides J.J. Sm. B. lipense Ames

B. camptochilum J.J. Verm.* B. lordoglossum J.J. Verm.*

B. cavipes J.J. Verm.* B. marudiense Carr*

B. ciliatum (Bl.) Lindl.* B. membranaceum Teysm. & Binnend.*

B. comberi J.J. Verm.* B. menglunense Tsi

B. delicatulum Schltr. B. pachyneuron Schltr.*

B. furcatum Aver. B. papillatum J.J. Sm.*

New Species of Bulbophyllum (Orchidaceae) 115

B. furcillatum J.J. Verm.* B. sensile Ames

B. grudense J.J. Sm.* B. striatellum Ridl.*

B. hemiprionotum J.J. Verm.* B. truncatum J.J. Sm.*

B. iterans J.J. Verm. & P. O’Byrne®* and B. trichorhachis J.J. Verm. &

P. O’Byrne* can also be included, but have a racemose inflorescense.

Our investigation has yielded three new species:

Bulbophyllum clinopus J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo pachyneuron fo/iis ellipticis obtusis, inflorescentia multo

breviore (2.5—3 cm longa in B. pachyneuron) differt. — Typus: Indonesia,

Sulawesi, central part, cult. Jongejan 387 (holo, L). Fig. 17.

Roots scattered along the rhizome. Rhizome long creeping, 1-1.5 mm

diam., sections between pseudobulbs 1.3-3 cm long, bracts not persistent.

Pseudobulbs widely spaced, patent, ovoid, 0.8-0.9 x 0.6-0.7 cm. Petiole up to

0.2-0.3 cm long. Leaf blade elliptic, 1.3-2 x 0.6-0.7 cm, index (length/width)

2.1-2.9; obtuse. Inflorescence ca | cm long, 1-flowered. Peduncle patent, ca

1.5 mm, bracts ca 2, the longest ca 1.8 mm long. Floral bract tubular, ca 1.8

mm, acute. Flowers not fully opening. Pedicel and ovary strongly curved, ca

4.5 mm long, basal node on a ca 0.5 mm long stump. Median sepal approx.

porrect, ovate, ca 3 x 1.7 mm, index 1.7-1.8; apiculate, margins papillose

towards the tip, base broadly attached; rather thick, surface glabrous. Lateral

sepals free, recurved, oblique, elliptic, ca 4.5 x 2.5 mm, index ca 1.8, margins

glabrous; otherwise as the median sepal. Petals porrect, oblong, ca 1.8 x 0.6

mm, index ca 3; obtuse, margins slightly erose towards the tip; base broadly

attached; rather thin, surface glabrous. Lip approx. straight, general outline

elliptic-ovate, ca 2 x 1.2 mm, index 1.6-1.7 (all without artificial spreading),

rounded, margins papillose; thick, surface glabrous but papillose towards the

margins; adaxially concave towards the base, with two inconspicuous, obtuse

ridges starting near the base, converging towards the tip and continuing over

about % of the length of the lip; abaxially with a weak, retuse ridge over most

of its length. Column ca 1.8 mm long, stigma elliptic, without keels inside,

with a very slight callus at its base, column foot not widened, hardly thickened,

without teeth. Stelidia triangular, ca 0.7 mm long, acute. Anther abaxially

with a rounded crest towards the base and a flat beak towards the tip, surface

somewhat papillose, front margin hardly drawn out, rounded, somewhat

erose. Pollinia 4, the inner somewhat shorter than the outer, flattened.

Colour: Sepals and petals whitish, suffused with purplish red, veins dark

116 Gard. Bull. Singapore 60 (1) 2008

Figure 17. Bulbophyllum clinopus J.J. Verm. & P. O’Byrme. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, left: abaxial side, right: adaxial

side; G. Pollinia, above: two pairs, below: a single pair. All from Jongejan 387 (spirit

sample).

New Species of Bulbophyllum (Orchidaceae) ia

purplish red. Lip dark purplish red, whitish near base. Column white with a

few pinkish specks, column foot stained dark red.

Habitat & ecology: No information available.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Most similar to B. pachyneuron Schltr, also from Sulawesi, but

easily distinguished by the elliptic, obtuse leaves (not ovate and acute, as

in B. pachyneuron), and the much shorter inflorescence (2.5-3 cm long in

B. pachyneuron).

Bulbophyllum ecristatum J.J. Verm. & P. O’Byrne, sp. nov.

A Bulbophylio ciliato petalis brevioribus latioribusque (2.5—3 mm

longis, longitudinis / latitudinis indice 3—4 in B. ciliato), labello ovato

(sine explicatione artificiali) stelidiis brevioribus acutis differt. — Typus:

Indonesia, Sulawesi, central part, SBG-O 4399 (holo, SING). Fig. 18.

Roots below the pseudobulbs. Rhizome creeping, 0.8-1 mm diam., sections

between pseudobulbs 0.8-1.1 cm long, bracts not persistent. Pseudobulbs

well spaced, ovoid, 0.4-0.6 x 0.3-0.5 cm. Petiole up to 0.05 cm long. Leaf

blade elliptic, 0.9-1.2 x 0.4-0.5 cm, index (length/width) 2.2-2.4; obtuse.

Inflorescence ca 1.4 cm long, 1-flowered. Peduncle patent, ca 0.8 cm, bracts

ca 3, the longest ca 1 mm long. Floral bract tubular, cal.2 mm, acute. Flowers

not fully opening. Pedicel and ovary ca 1.9 mm long, basal node on a ca

0.8 mm long stump. Median sepal recurved, triangular, ca 4.3 x 1.2 mm,

index ca 3.6; acute, margins entire, base rather broadly attached; rather thin,

somewhat thickened towards the tip, adaxially finely papillose towards the

tip. Lateral sepals free, oblique, ca 6 x 1.7 mm, index ca 3.5; otherwise as the

median sepal. Petals porrect, triangular, ca 1.2 x 0.7 mm, index ca 1.7; acute,

margins erose distally, base broadly attached; thin, surface glabrous. Lip

(all without artificial spreading; when spread hastate), three-lobed; midlobe

ovate, rounded, margins entire, about half-way along the length of the lip with

flat, elliptic, obtuse paleae ca 0.1 mm long; rather thick; adaxially concave,

surface aaprox. glabrous, abaxially with a rounded median ridge near the

base, surface elsewhere convex, papillose towards the tip; sidelobes attached

along the basal 4 of the length of the lip, slightly retrorse, oblique, triangular,

obtuse, front margin with paleae as above, back margin entire; rather thin,

surface glabrous. Column ca 0.9 mm long, stigma obovate, without keels

inside, without teeth at its base, column foot with a slight, rounded swelling

118 Gard. Bull. Singapore 60 (1) 2008

Figure 18. Bulbophyllum ecristatum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side. All from SBG-O 4399 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 119

just above the ligament. Stelidia triangular, ca 0.7 mm long, acute. Anther

abaxially with a slight, rounded crest, surface aaprox. glabrous, front margin

drawn out into a triangular beak. Pollinia (not seen).

Colour: Sepals whitish near the base, pale purplish red around the veins,

entirely pale purplish red towards the tip. Petals translucent, with a purplish

red midvein. Lip purple, darker at the base.

Habitat & ecology: Found as an epiphyte in the crown of small trees in

riverine forest. Alt. 1400-1500 m. Flowering Jan, Feb, Aug.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. ciliatum (B1.) Lindl., is most similar, but differs in having 2.5-3 mm

long, narrower petals (index 3-4), an elliptic or slightly obovate lip (without

artificial spreading) and more slender, acuminate stelidia.

Bulbophyllum hemisterranthum J.J. Verm. & P. O’Byre, sp. nov.

A Bulbophyllo pachyneuron foliis gradatim basin versus descrescentibus,

labello glabro differt. — Typus: Indonesia, Sulawesi, central part, SBG-O

5046 (holo, SING). Fig. 19.

Roots scattered along the rhizome. Rhizome long creeping, 1.5-2.5 mm

diam., sections between pseudobulbs 2.5-17.5 cm long, bracts not persistent.

Pseudobulbs widely spaced, patent, ovoid-cylindrical, 2.8-5 x 0.5-0.8 cm.

(length/width) 6-8.2; acute. Inflorescence 6-8.5 cm long, 1-flowered.

Peduncle patent, 3.5-5 cm, bracts ca 2, the longest 4-5 mm long. Floral bract

tubular, 3.5-4 mm, acute. Flowers not fully opening. Pedicel and ovary 18-

25 mm long, basal node on a 3.5-4 mm long stump. Median sepal approx.

porrect, ovate, ca 7 x 3.8 mm, index 1.8-1.9; apiculate-shortly acuminate,

margins entire, base broadly attached; rather thick, glabrous. Lateral sepals

free, recurved, oblique, ca 9 x 4 mm, index 2.2-2.3, base rather broadly

attached; otherwise as the median sepal. Petals porrect, ovate-triangular, ca

2.8 x 1.7 mm, index 1.6-1.7; rounded and shortly apiculate, margins entire;

base broadly attached; rather thin, glabrous. Lip recurved, general outline

ovate, ca 3 x 2 mm, index ca 1.5 (all without artificial spreading), rounded,

margins entire; thick; adaxially concave and furrowed over most of its

length, with two rather distinct, obtuse ridges starting close to the margin

and converging distally and running up to close to the tip where they meet,

surface glabrous; abaxially with a weak, retuse ridge over most of its length,

120 Gard. Bull. Singapore 60 (1) 2008

Figure 19. Bulbophyllum hemisterranthum J.J. Verm. & P. O’Byme. A. Habit; B. Flower;

C. Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left:

adaxial side, right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial

side, below: abaxial side; G. Pollinia, above: a single pair, below: two pairs. All from SBG-O

5046 (spirit sample).

eo ee eee

surface locally very finely papillose Column ca 3.1 mm long, stigma elliptic,

without keels insi ne with a very slight callus at its base, column foot narrow,

not thickened, without teeth. Stelidia triangular, ca 1.5 mm long, acute, upper

margin slightly erose or not. Anther abaxially with a rounded crest towards

the tip, surface glabrous, front margin drawn out, rounded, papillose. Pollinia

4. the inner as long as the outer, flattened.

Colour. Leaves green, adaxially with a thin, dull blackish purple midvein.

Sepals dark red, pale green along the margins. Petals pale green wi ith a

blackish spot at the tip. Lip orange. Column pale green.

Habitat & ecology: In mossy montane forest, epiphyte in thick moss pads on

trees bordering a stream, also terrestrial in mossy riverbank. Alt. ca 12

1400 m. Flowering Feb.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: This is best placed in sect. Monanthaparva because of its general

habit, the narrow column foot, combined with the somewhat fleshy, at least

not diaphanous flowers. Among the Sulawesi species, it is most similar

to B. pachyneuron Schltr, which differs in having wider leaves that are

abruptly narrowed at their base, and a partly papillose lip.

Bulbophyllum sect. Monanthes (Bl.) Aver.

(= sect. Polyblepharon)

Diphyes sect. Monanthes B1., Biydr. Fl. Ned. Ind. (1825) 311.— Bulbophyllum

sect. Monanthes (Bl.) Aver., Identification Guide to Vietnamese

Orchids (1994) 279. — TYPE SPECIES: Diphyes tortuosa Bl. [=

Bulbophyllum tortuosum (B1.) Lindl.]

Bulbophyllum sect. ese gape Schltr oe Repert. 10 (1912) 177:

Feddes Repert. eee 13) 701. 794; Van Royen, Alpine Fl. New

Guinea 2 (1979) 217. — we: TOTYPE SPECIES (des signated by Vz

Royen): ere polyblepharon Schltr.

Bulbophyllum consimile J.J. Verm. & P. O’Byme, sp. no1

A Bulbophyllo membranaceo sepali median marginibus papilloso-

ciliolatis - labell 1G) pagina adc wiali i sine crista me rik fel f di Oorum SeCcus

margines inferiores sine dente deltoideo differt dif = iyi Indonesia,

Sulawesi, southwest part SBG-O 5006 (holo, SING). "Fig, 20.

122, Gard. Bull. Singapore 60 (1) 2008

Roots below the pseudobulbs, spreading. Rhizome creeping or shortly

straggling, 1.5-2 mm diam., sections between pseudobulbs 4-5 cm long,

bracts not persistent. Pseudobulbs ovoid, (obliquely) erect, 1.4-1.5 x 0.6-0.8

cm. Petiole up to 2.5-4 mm long. Leaf blade ovate, 3.6-4 x 2.1-2.2 cm, index

(length/width) 1.7-1.8; acute. Inflorescence ca 1.4 cm long, 1-flowered.

Peduncle porrect, ca 0.6—0.9 cm, bracts 2.5- 4 mm long. Floral bracts

tubular, 1.8-2.5 mm, acute. Flowers not fully opening. Pedicel and ovary

ca 2 mm long, basal node coinciding with the attachment of the floral bract.

Median sepal slightly recurved, elliptic, ca 5.2 x 2.7 mm, index 1.9-2; acute-

acuminate, margins entire, finely papillose-ciliolate, base widely attached;

rather thick, surface glabrous. Lateral sepals approx. porrect, fused along

the lower margin, oblique, triangular, ca 6 x 2.4 mm, index ca 2.5; acute;

margins entire; glabrous; otherwise as the median sepal. Petals porrect,

ovate-triangular, ca 2.1 x 1.3 mm, index ca 1.6; acuminate, margins finely

erose, base widely attached; thin, surface glabrous. Lip recurved about half-

way, general outline ovate, ca 2.3 x 1.1 mm, index ca 2.1; acute-apiculate,

margins ciliate in the top % of its length; rather thick over its entire length,

surface glabrous; adaxially slightly concave towards the base, slightly convex

towards the tip, without a median ridge; abaxially virtually without a ridge

near the base. Column ca 1.4 mm long, stigma without basal tooth, column

foot thickened near the tip. Stelidia curved, triangular, ca 0.7 mm long, acute.

Anther abaxially with a slight, rounded, papillose crest, surface somewhat

papillose, front margin drawn out into a deltoid beak with approx. glabrous

margins. Pollinia 4, drop-shaped, the inner somewhat flattened and about

half as long as the outer, a fleshy stipes is present.

Colours: Median sepal and petals dull purple. Lateral sepals idem, but pale

yellow along lower margin. Lip purple.

Habitat & ecology: Understory epiphyte in moss cushions in low, dense, very

wet montane forest, also in trees in gardens, at 1600-2200 m alt. Flowering

Jan.

Distribution: INDONESIA. Sulawesi: Sulawesi, southwest part (1 specimen

seen).

Notes: B. consimile is very similar to the widespread B. membranaceum

Teysm. & Binnend., at first sight, but differs in details of the flowers: the basal

node of the pedicel is level with the atachment of the floral bract, the median

sepal has papillose-ciliolate margins (not glabrous), the lip margins are ciliate

(not papillose), the lip is about equally thick (in lateral view) over most of its

New Species of Bulbophyllum (Orchidaceae)

Figure 20. Bulbophyllum consimile J.J. Verm. & P.O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side, below:

abaxial side; G. Pollinia, above: two pairs with appendage, below: a single pair. All from

SBG-O 5006 (spirit sample).

124 Gard. Bull. Singapore 60 (1) 2008

length (not distinctly thicker near the base), and lacks a median ridge, and the

stelidia lack a deltoid tooth along the lower margins.

Bulbophyllum oncopus J.J. Verm. & P. O’Byrne, sp. nov.

A Bulbophyllo oligochaete sepalis ciliatis, lateralibus obtusis, petalis

triangularibus obtusis, stelidiis secus marginem inferiorem sine dente

differt. — Typus: Indonesia, Sulawesi, SBG-O 3947 (holo, SING). Fig. 21.

Roots all produced close to the base of the rhizome. Rhizome hanging down

perpendicularly, up to 10 cm long, ca 1 mm diam., bracts not persistent.

Pseudobulbs ovoid-cylindrical, prostrate on and fused to the rhizome, so that

the new pseudobulbs arise from half-way the length of the old, 0.6-1.2 x 0.15-

0.25 cm. Petiole up to 1 mm long. Leaf blade ovate, 2.7-4.8 x 0.5-1.1 cm,

index (length/width) 4.3-7; acute. Inflorescence ca 0.8 cm long, 1-flowered.

Peduncle porrect, ca 0.15-0.2 cm, bracts 1. 2-2.5 mm long. Floral bracts

tubular, ca 2.5 mm, acute. Flowers not fully opening. Pedicel and ovary ca

1.3 mm long, basal node coinciding with the attachment of the floral bract.

Median sepal recurved, elliptic, ca 3.3 x 1.8 mm, index 1.8-1.9; rounded,

margins entire, ciliate, base widely attached; rather thick, surface glabrous.

Lateral sepals porrect, fused along the lower margin up to % of their length,

oblique, ca 4 x 1.3 mm, index 3-3.1; obtuse; upper margin ciliate-fimbriate;

otherwise as the median sepal. Petals porrect, ovate, ca 0.8 x 0.7 mm, index

1.1-1.2; obtuse, margins slightly erose towards the tip, base widely attached;

rather thin, glabrous. Lip almost straight, general outline ovate with a drawn

out, narrow top half because the margins are folded backwards, ca 1.7 x 0.7

mm, index 2.4-2.5; obtuse, margins ciliate in the basal half, papillose in the

top half; thin, surface glabrous; adaxially slightly concave towards the base,

as well as near the tip, without a median ridge; abaxially without a ridge near

the base. Column ca 1.2 mm long, stigma without basal tooth, column foot

distinctly widened and thickened and with a flat somewhat angular swelling

near each margin. Stelidia curved, triangular, ca 0.6 mm long, acute. Anther

abaxially with a distinct, rounded, papillose crest, surface somewhat papillose,

front margin drawn out into a rounded beak with approx. glabrous margins.

Pollinia 4, drop-shaped, the inner flattened and less than half as long as the

outer.

Colours: Median sepal pinkish purple, whitish towards the base, with slighty

darker veins. Lateral sepals pinkish purple, with a white patch near the base.

Petals translucent white. Lip pinkish purple, but white along the midvein.

Column white.

New Species of Bulbophyllum (Orchidaceae) 125

! PecpCerh ch vCal

rpg ENT

May,

ALAA yy

Figure 21. Bulbophyllum oncopus J.J. Verm. & P.O’ Byrne. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side, right:

abaxial side; E. Column and lip, lateral view; F. Anther, above: abaxial side, below: adaxial

side; G. Pollinia, left: one pair, right: two pairs. All from SBG-O 3947 (spirit sample).

126 Gard. Bull. Singapore 60 (1) 2008

Habitat & ecology: Montane forest on limestone soil, at 1500 m alt. Flowering

Jan-Mar, Oct-Dec.

Distribution: INDONESIA. Sulawesi: Sulawesi, central part (1 specimen

seen).

Notes: B. oncopus shares the general shape of the lip, with recurved margins

in the top half, with B. oligochaete Schltr, from New Guinea. It differs in

having ciliate sepals, the lateral sepals obtuse rather than broadly round,

triangular, obtuse petals and stelidia without a tooth along the lower margin.

B. loxophyllum Schltr, also from New Guinea is even more similar in the

shape of the flower parts, but has eciliate lateral sepals and the margins of the

lip not recurved.

Bulbophyllum sect. Polymeres (Bl.) J.J. Verm. & P. O’Byrne, comb. nov,

. (= sect. Fruticicola, sect. Epibulbon, sect. Leptopus, sect. Megaloglossum,

sect. Rhizocaulon)

Diphyes sect. Polymeres Bl., Bidr. Fl. Ned. Ind. 7 (1825) 318.-LECTOTY PE

SPECIES (here designated): Diphyes tenuifolia Bl. [Bulbophyllum

tenuifolium (BI.) Lindl.].

Bulbophyllum sect. Micromonanthe Schltr., Feddes Repert. Beih. 1 (1913)

701 & 783. — Bulbophyllum sect. Schlechteria Van Royen, Alpine FI.

New Guinea 2 (1979) 207.— LECTOTYPE SPECIES (designated by

Van Royen): Bulbophyllum neoguineense J.J. Sm.

Bulbophyllum sect. Leptopus Schltr., Feddes Repert. Beth. 1 (1913) 702 & 828;

Van Royen, Alpine Fl. New Guinea 2 (1979) 223. - LECTOTYPE

SPECIES (designated by Van Royen): Bulbophyllum leptopus Schltr.

Bulbophyllum sect. Rhizocaulon Schltr., Feddes Repert. Beih. 1 (1913) 702

& 831. —- LECTOTYPE SPECIES (here designated): Bulbophyllum

dictvoneuron Schltr.

Bulbophyllum sect. Fruticicola Schitr., Feddes Repert. Beih. 1 (1913) 702 &

833; Van Royen, Alpine Fl. New Guinea 2 (1979) 227.—LECTOTY PE

SPECIES (designated by Van Royen): Bulbophyllum fruticicola

Schltr.

Bulbophyllum sect. Epibulbon Schltr., Feddes Repert. Beih. 1 (1913) 703

& 845. LECTOTYPE SPECIES (here designated): Bulbophyllum

epibulbon Schltr.

)

Bulbophyllum sect. Megaloglossum Carr, J. Malayan Branch Roy. Asiat.

Soc. 11, 1 (1933) 90.— TYPE SPECIES: Bulbophyllum brastagiense

Carr (= B. crepidiferum J.J. Sm.).

This section includes a large number of species, all characterized by the

Sie of lateral teeth or wings on the column foot, next to the ligament.

Additional characters present in many species include: flowers fairly small (1

cm long or less) sepals and petals thin, petals much shorter than the sepals, lip

thick and spongy, with a tuft of hairs or papillae on the abaxial side, column

with distinct, acute stelidia, column foot elongated.

Vegetatively, the species display a wide array of growth forms,

which have been used in the past to arrange the species into sections: long,

thin, creeping rhizomes with widely spaced, erect pseudobulbs (section

Megaloglossum), patent or pendulous rhizomes with slender porrect

pseudobulbs (section Leptopus), patent or pendulous rhizomes with short.

porrect, distichous pseudobulbs (section Fruticicola), patent or pendulous

rhizomes with the pseudobulbs fused to it, one placed on top of the other

(section Epibulbon). It is difficult, however, to uphold these groupings: too

many species appear intermediate between groups. The new Sulawesi species

here described include several examples of such intermediates. Provisionally.

we prefer to include them all in one section. The grouping given below is to

fit Sulawesi material only and has no practical value outside the

—

area.

Group a - Species with creeping rhizomes, more or less distant pseudobulbs

of substantial size, and inflorescences much shorter than the pseudobulb plus

leaf they arise from.

The flowers are often as in section Fruticicola, a section ger ay

characterized by a patent or pendulous rhizome. Bulbophyllum valeryi J/./.

Verm. & P. O'Byrne from Sulawesi belongs here too.

Bulbophyllum cymbochilum J.J. Verm. & P.O’ Byme, sp. nov.

Bulbophyllum cymbochilum J.J. Verm. & P. O'Byrne, in sectione Polymeres

labellum concavum leviter sursum flexum singularis. — Typus: Indonesia,

Sulawesi, central part, SBG-O 5239 (holo, SING). Fig. 22.

Roots prams below the pseudobulbs. Rhizome creeping, 1.2-1.8 mm

diam., sections between pseudobulbs 0.5-0.8 cm long, bracts not persistent.

Pseudobulbs spaced, obliquely erect, ovoid, 0.7-0.9 x 0.4-0.5 cm. Petiole

up to 0.3 cm long. Leaf blade elliptic to ovate, 2.2-2.8 x 0.7-0.9 cm, index

(length/width) 2.3-3.3; acute. Inflorescence ca 0.9 cm long, 1-flowered.

Peduncle patent, 0.18-0.25 cm, bracts ca 2, the longest ca 1.4 mm long. Floral

128 Gard. Bull. Singapore 60 (1) 2008

Figure 22. Bulbophyllum cymbochilum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C. |

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: abaxial side, right:

adaxial side. All from SBG-O 5239 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 129

bract tubular, ca 1.7 mm, acute. Flowers opening wide. Pedicel and ovary ca

2.2 mm long, basal node on a ca 0.8 mm long stump. Median sepal spreading,

elliptic, ca 5.8 x 2.2 mm, index 2.6-2.7; acuminate, margins entire, slightly

papillose distally, base rather narrowly attached; rather thin, surface glabrous.

Lateral sepals free, spreading to reflexed, oblique, ca 6 x 2.5 mm, index ca

2.4; otherwise as the median sepal. Petals porrect, elliptic, ca 2.1 x 1.8 mm,

index |.1-1.2; rounded, margins slightly erose and very finely papillose; base

shortly clawed; thin, adaxially slighty and very finely papillose towards the

tip. Lip approx. straight or slightly incurved at the tip, general outline approx.

elliptic, ca 2.9 x 1.3 mm, index 2.2-2.3 (all without artificial spreading),

rounded, margins entire slightly papillose towards the base; very thick and

soft, surface slightly papillose locally; adaxially entirely concave, with two

rounded ridges starting at the base, near the margin, gradually converging up

to about 2/5 of the length of the lip, in between these ridges, near the base of

the lip, a transverse ridge consisting of two fused teeth, leaving a small cavity

in between on their front side; abaxially with a wide, retuse ridge over most

of the length of the lip. Column ca | mm long, stigma elliptic, without keels

inside, without teeth at its base, column foot with thin, ovate, obtuse lateral

teeth just above the ligament. Stelidia triangular, ca 0.4 mm long, acute.

Anther abaxially with a rounded crest, surface approx. glabrous, front margin

drawn out, rounded, papillose. Pollinia (not seen).

Colour: Plant green. Sepals cream coloured with purplish red veins and

stains. Petal translucent white with large blackish blotches. Lip dark purplish

red, white near the base.

Habitat & ecology: Found as an understory epiphyte in mossy montane

forest. Alt. ca 1200-1400 m. Flowering Feb-Apr, Jun-Aug, Nov.

Distribution: INDONESIA: Sulawesi, central part (1 specimen seen).

Notes: Uniquely identified within section Polymeres by its concave, slightly

upturned lip.

Bulbophyllum gamandrum J.J. Verm. & P. O’Byrne, sp. nov.

A Bulbophyllo bowkettae /abello ovato, stelidiis rectis, et a B. sarcochilum

sepalis longioribus, labello ad basin plus distincte concavo differt.—Typus:

Indonesia, Sulawesi, central part, SBG-O 51/2 (holo, SING). Fig. 23.

Roots scattered along the rhizome. Rhizome creeping, 1.5-2 mm diam.,

sections between pseudobulbs 1.2-1.3 cm long, bracts not persistent.

130 Gard. Bull. Singapore 60 (1) 2008

Figure 23. Bulbophyllum gamandrum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C. i

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: abaxial side, right:

adaxial side. All from SBG-O 5/12 (spirit sample).

> |

New Species of Bulbophyllum (Orchidaceae) 131

Pseudobulbs spaced, prostrate for most of their length, ovoid, 0.9-1 x ca 0.6

cm. Petiole up to 0.15 cm long. Leaf blade elliptic, ca 1.9 x 0.9 cm, index

(length/width) ca 2.1; obtuse. Inflorescence ca 1.9 cm long, 1-flowered.

Peduncle patent, ca 0.9 cm, bracts ca 4, the longest ca 2.8 mm long. Floral

bract tubular, ca 2.5 mm, acute. Flowers not fully opening. Pedicel and ovary

ca 6mm long, the latter slightly costulate, basal node on aca 2 mm long stump.

Median sepal approx. porrect, ovate, ca 7 x 2.5 mm, index ca 2.8; acute,

margins entire, base broadly attached; rather thin, glabrous. Lateral sepals

free, recurved, oblique, ca 7.3 x 3 mm, index 2.4-2.5; otherwise as the median

sepal. Petals porrect, elliptic, ca 2.8 x 2 mm, index ca 1.4; obtuse, margins

slightly erose; base narrowly attached; thin, surface approx. glabrous. Lip

slightly curved, general outline approx. ovate with a slightly drawn-out top

part, ca 3.5 x 2.3 mm, index 1.5-1.6 (all without artificial spreading), obtuse,

margins entire; thick, glabrous but slightly and finely papillose near the tip;

adaxially concave and slightly furrowed in the basal half, surface slightly

convex near the tip; abaxially with a weak, retuse ridge up to about half-way

its length. Column ca 1.5 mm long, stigma approx. circular, without keels

inside, with a distinct callus at its base, column foot widened and thickened,

with thick, retrorse, semi-elliptic, obtuse lateral teeth just above the ligament.

Stelidia triangular, ca 0.7 mm long, acute. Anther fused to the top of the

column, abaxially with a inconspicuous crest, surface approx. glabrous, front

margin drawn out, obtuse. Pollinia (not seen).

Colour: Plant green. Sepals pale reddish with darker veins. Lip purplish red.

Habitat & ecology: Epiphyte in open, low, secondary woodland of thin pole

trees on a steep slope. Alt. 800-1000 m. Flowering Feb, Apr.

Distribution: (NDONESIA. Sulawesi, central part (1 specimen seen).

Notes: B. bowkettae F.M. Bail., from Australia, is most similar, but has an

oblong lip and upturned stelidia. B. sarcochilum J.J. Verm. & P. O’ Byrne, is

vegetatively similar but has much shorter sepals and a lip that is concave only

near the base.

Bulbophyllum rhodophyllum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo ciliipetalo foliis minoribus (versus circa 10 cm_ longa),

inflorescentia breviore (versus 3 cm longa), labello obtuso sepalis breviore, a

B. ruguloso petalis acutis differt. — Typus: Indonesia, Sulawesi, central part,

SBG-O 5066 (holo, SING). Fig. 24.

132 Gard. Bull. Singapore 60 (1) 2008

Roots below the pseudobulbs. Rhizome creeping, |.2-1.5 mm diam., sections

between pseudobulbs 0.7-1.5 cm long, bracts not persistent. Pseudobulbs

spaced, prostrate towards the base, ovoid, 0.6-1.2 x 0.4-0.7 cm. Petiole up to 0.3

cm long. Leaf blade elliptic to ovate, 1.3-2.8 x 0.5-0.8 cm, index (length/width)

2.6-3.6; acute. Inflorescence ca 1.7 cm long, 1-flowered. Peduncle patent, ca

0.25 cm, bracts ca 2, the longest ca 1 mm long. Floral bract tubular, ca 2.5 mm,

acute. Flowers opening wide. Pedicel and ovary ca 6 mm long, costulate, basal

node on a ca 1.5 mm long stump. Median sepal spreading, elliptic, ca 9 x 2.2

mm, index ca 4.1; acute, margins entire, base rather broadly attached; rather

thin, surface glabrous. Lateral sepals free, oblique, ovate, ca 9 x 3.2 mm, index

ca 2.8; otherwise as the median sepal. Petals porrect, elliptic, ca 3 x 1 mm, index

ca 3; acute, margins entire, papillose with elongated papillae; base rather broadly

attached; rather thin, adaxially very finely papillose towards the margins. Lip

slightly recurved near the base, general outline ovate, ca 5 x 1.4 mm, index 3.5-

3.6 (all without artificial spreading), obtuse, margins entire, slightly papillose

towards the base; thick and soft, surface slightly verrucate distally; adaxially

. concave and somewhat furrowed towards the base, slightly convex towards the

tip; abaxially with an inconspicuous ridge near the base, surface with a papillose-

shortly hairy patch in the basal half of the lip. Column ca 1.5 mm long, stigma

obovate, without keels inside, without teeth at its base, column foot widened

and thickened, with thick, triangular, obtuse lateral teeth just above the ligament.

Stelidia triangular, ca 0.5 mm long, acute. Anther abaxially with a rounded crest,

surface papillose, front margin drawn out, rounded, papillose. Pollinia 4, ovoid,

the inner almost as long as the outer, flattened.

Colour: Pseudobulbs purplish, leaves dark green. Sepals pale ochrish with

purplish red veins. Petals whitish with hyaline papillae along the margins. Lip

purplish red near base, orange red towards the tip.

Habitat & ecology: Found as an understory epiphyte in mossy montane

forest. Alt. ca 1200-1400 m. Flowering Feb, Apr-Aug, Nov, Dec.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Most similar to B. ciliipetalum Schlitr., which differs in having larger

leaves (about 10 cm long), a longer inflorescence (about 3 cm long), and an

acute lip which is almost as long as the sepals. B rugulosum J.J. Sm., differs

in being a larger plant, and in having rounded petals.

New Species of Bulbophyllum (Orchidaceae) 133

Figure 24. Bulbophyllum rhodophyllum J.J. Verm. & P. O’Byme. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side,

below: abaxial side; G. Pollinia, left: a single pair, right: two pairs. All from SBG-O 5066

(spirit sample).

134 Gard. Bull. Singapore 60 (1) 2008

Bulbophyllum sarcochilum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllis bowkettae ef B. johnsonii sepalis brevioribus et latioribus

(indice 2—3 in speciebus ambabus australiis), labello plus carnoso ad

basin leviter concavo tantum differt. — Typus: Indonesia, Sulawesi, central

part, SBG-O 5783 (holo, SING). Fig. 25.

Roots scattered along the rhizome. Rhizome creeping, ca 1.5 mm diam.,

sections between pseudobulbs 1.5-2 cm long, bracts not persistent.

Pseudobulbs spaced, prostrate towards the base, ellipsoid to obovoid, 1.1-

1.2 x ca 0.6 cm. Petiole up to 0.05 cm long. Leaf blade ovate to elliptic, ca

1.7 x 0.8-0.9 cm, index (length/width) 1.9-2.1; obtuse. Inflorescence ca 1.6

cm long, 1-flowered. Peduncle patent, ca 0.9 cm, bracts ca 3, the longest ca 2

mm long. Floral bract tubular, ca 2.5 mm, acute. Flowers not fully opening.

Pedicel and ovary ca 4.5 mm long, the latter slightly costulate, basal node on

aca 2 mm long stump. Median sepal approx. porrect, ovate, ca 3.2 x 2 mm,

index ca 1.6; shortly acuminate, margins finely papillose towards the tip, base

- broadly attached; rather thick, surface glabrous. Lateral sepals free, recurved,

oblique, ca 3.5 x 2.3 mm, index 1.5-1.6, upper margin glabrous; otherwise

as the median sepal. Petals porrect, elliptic-ovate, ca 2 x 1.3 mm, index 1.5-

1.6; rounded, margins finely papillose-ciliolate; base narrowly attached; thin,

surface approx. glabrous. Lip approx. straight, general outline approx. elliptic,

ca 2.2 x 1.2 mm, index 1.8-1.9 (all without artificial spreading), rounded,

margins entire; thick and soft; adaxially slightly concave and furrowed

near the base, with two low, obtuse ridges starting close to the margin and

converging distally and running up to ’/,-'/, of the length of the lip, surface

convex distally, adaxial surface glabrous; abaxially with a weak, retuse ridge

and a papillose surface near the base. Column ca 1.4 mm long, stigma circular,

without keels inside, with a slight callus at its base, column foot widened and

thickened, with thick, deltoid, obtuse lateral teeth just above the ligament.

Stelidia triangular, ca 0.7 mm long, acute. Anther abaxially with a rounded

crest, surface approx. glabrous but papillose towards the tip, front margin

drawn out, rounded, papillose. Pollinia 4, the inner slightly shorter than the

outer, flattened.

Colour: Sepals yellow, with slightly darker veins. Petals translucent yellow,

with a blackish red midvein and top. Lip bright yellow, suffused with red near

the base.

Habitat & ecology: Epiphyte in montane oak forest. Alt. ca 1200 m. Flowering

Apr.

oS)

‘Nn

New Species of Bulbophyllum (Orchidaceae) 1:

Figure 25. Bulbophyllum sarcochilum J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: adaxial side, right:

abaxial side; G. Pollinia, left: two pairs, right: a single pair. All from SBG-O 5783 (spirit

sample).

136 Gard. Bull. Singapore 60 (1) 2008

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Vegetatively, this species looks similar to B. bowkettae F.M. Bail., and,

to a lesser extent, B. johnsonii T.E. Hunt, both from Australia. It shares the

half-prostrate pseudobulbs on a long-creeping rhizome with both species;

it differs in having shorter and wider sepals (index 2-3 in both Australian

species). Also, the lip is much more fleshy, and only slightly concave near

the base.

Bulbophyllum semiindutum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllo valeryi petalis subacutis, stigmatis basin sine dentibus,

a B. caeco et B. pisibulbo sepalis latioribus ( indice plus quam 2 in B.

pisibulbo), pedicellis brevioribus (pedicellum ovariumque plus quam 12

mm longum) differt. — Typus: Indonesia, Sulawesi, central part, SBG-O

5051 (holo, SING). Fig. 26.

- Roots below the pseudobulbs. Rhizome creeping, |.2-1.5 mm diam., sections

between pseudobulbs 0.5-0.8 cm long, bracts not persistent. Pseudobulbs

spaced, obliquely erect, ovoid, 0.6-1 x 0.3-0.5 cm. Petiole up to 0.2 cm long.

Leaf blade elliptic, 1.7-2.7 x 0.4-0.5 cm, index (length/width) 4.2-5.4; acute.

Inflorescence ca | cm long, 1-flowered. Peduncle patent, 0.3-0.4 cm, bracts

ca 2, the longest ca 1.5 mm long. Floral bract tubular, ca 1.8 mm, acute.

Flowers opening wide. Pedicel and ovary ca 3.7 mm long, costulate, basal

node on a ca |.5 mm long stump. Median sepal recurved, ovate, ca 4.3 x

2.3 mm, index 1.8-1.9; acute, margins entire, base rather broadly attached;

rather thin, adaxial surface slightly fine papillose locally. Lateral sepals free,

spreading, oblique, ovate-triangular, ca 4.2 x 3 mm, index ca 1.4; otherwise

as the median sepal. Petals porrect, elliptic-obovate, ca 2.3 x 1.4 mm, index

1.6-1.7; subacute, margins entire, papillose; base narrowly attached; thin,

approx. glabrous. Lip recurved, general outline elliptic, ca 2.5 x 1.2 mm,

index ca 2.1 (all without artificial spreading), round, margins entire; thick

and soft; adaxially with a wide, deep cavity near the base, this separate from

the concave and somewhat furrowed middle part of the lip which is bordered

by two rounded ridges situated close to the margin and converging distally

and running up to 3/5 of the length of the lip, surface convex distally, entire

adaxial surface glabrous except for papillose patches close to the margins

about halfway along the length of the lip; abaxially with a retuse ridge up to

of the length of the lip, surface papillose except near the tip and near the base.

Column ca 1.3 mm long, stigma circular, without keels inside, without teeth

at its base, column foot widened and thickened, with thick, triangular, obtuse

lateral teeth just above the ligament. Stelidia triangular, ca 0.4 mm long,

oc SRN |

New Species of Bulbophyllum (Orchidaceae) 137

Figure 26. Bulbophyllum semiindutum J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side,

below: abaxial side. All from SBG-O 505/ (spirit sample).

138 Gard. Bull. Singapore 60 (1) 2008

acute. Anther abaxially with a rounded crest, surface approx. glabrous but

papillose towards the tip, front margin drawn out, retuse, papillose. Pollinia

(not seen).

Colour: Sepals dull pale yellowish, with dark red veins. Petals pale yellow

with a blackish red tip. Lip whitish around the ligament, purplish red up to

approx. half-way its length, top part orange.

Habitat & ecology: Found as an understory epiphyte in mossy montane

forest. Alt. ca 1200-1400 m. Flowering Feb, Jul, Sep-Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Most similar to B. valeryi J.J. Verm. & P. O’Byrne, which differs in

having ovate, acute petals and two large teeth at the base of the stigma. B.

caecum J.J. Sm., and B. pisibulbum J.J. Sm., are also similar, but differ in

. having narrower sepals (index larger than 2) and longer pedicels (pedicel and

ovary 12 mm or longer).

Group b - Species with creeping or more or less patent rhizome, clustered

pseudobulbs of reduced size, and inflorescences approx. as long as or longer

than the vegetative shoot they arise from. As in the previous group, the

flowers are as in section Fruticicola, a section generally characterized by

a patent or pendulous rhizome. However, J.J. Smith (1926, p. 59) , under

the description of B. caecum J.J. Sm., a species that would also fit in group

b, has expressed the opinion that such species are best included in section

Fruticicola nevertheless.

Bulbophyllum amauroloma J.J. Verm. & P. O’ Byrne, sp. nov.

Bulbophyllum amauroloma J.J. Verm. & P. O'Byrne, in sectione Polymeres

combinatione pseudobulborum fastigiatorum, labelli pagina elliptica

singularis. -Typus: Indonesia, Sulawesi, central part, SBG-O 5244 (holo,

SING). Fig. 27.

Roots below the pseudobulbs. Rhizome creeping or straggling at some

distance from the substrate, 1.5-2 mm diam., sections between pseudobulbs

0.2-0.4 cm long, bracts not persistent. Pseudobulbs rather close together,

obliquely erect, ovoid, 0.5-0.6 x 0.25-0.3 cm. Petiole up to 0.4 cm long.

Leaf blade elliptic, 2-3.5 x 0.35-0.45 cm, index (length/width) 6.4-9.7; acute.

Inflorescence 5-6 cm long, |-flowered. Peduncle patent, 3-3.5 cm, bracts ca

2, the longest ca 3.5 mm long. Floral bract tubular, ca 2.5 mm, acute. Flowers

140 Gard. Bull. Singapore 60 (1) 2008

not fully opening. Pedicel and ovary ca 12 mm long, basal node on a ca

1.7 mm long stump. Median sepal recurved, ovate, ca 7.5 x 2.8 mm, index

2.6-2.7; acuminate, margins entire, slightly and very finely papillose, base

rather narrowly attached; thin, surface glabrous. Lateral sepals free, oblique,

elliptic-obovate, ca 9 x 5 mm, index ca 1.8; base rather broadly attached;

otherwise as the median sepal. Petals porrect, elliptic-obovate, ca 3.2 x 1.5

mm, index 2.1-2.2; subacute, margins entire, finely papillose; base rather

broadly attached; thin, surface approx. glabrous. Lip recurved at approx.

2/5 of its length, consisting of a short, rather narrow basal part and a wide,

elliptic blade, ca 3.5 x 2.7 mm, index ca 1.3 (all without artificial spreading),

rounded, margins entire, glabrous except for a short, ciliate stretch just above

the base; thick and soft; adaxially concave near the base, with two rather

high and narrow ridges starting at the base and running up, while slightly

converging, to of the length of the lip, top “% approx. flat, adaxial surface

glabrous; abaxially with a retuse ridge up to '4-'4 of the length of the lip,

surface glabrous except for a hairy patch just in front of the ndge. Column ca

1.5 mm long, stigma elliptic, without keels inside, without teeth at its base,

column foot widened, with obtuse lateral teeth as well as a median cavity

just above the ligament. Stelidia triangular, ca 0.7 mm long, acute. Anther

abaxially with a large, rounded crest, surface finely papillose, front margin

drawn out, round, papillose. Pollinia (not seen).

Colour: Sepals and petals yellow, the latter with red margins. Lip yellow in

centre, red towards the margins. Column yellow.

Habitat & ecology: Found as an understory epiphyte in mossy montane forest.

Alt. ca 1200-1400 m. Flowering Feb, Mar, Aug, Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Uniquely identified within section Po/ymeres by the combination of

the clustered pseudobulbs and by the wide, elliptic blade of the lip.

Bulbophyllum croceodon J.J. Verm. & P.O’ Byrne, sp. nov.

A Bulbophyllo stylocoryphe sepalis lateralibus et petalis ac labello latioribus,

labelli parte apicali extensa, sed non contracta ante dilationem in apice

globoso differt. — Typus: Indonesia, Sulawesi, central part, SBG-O 4539

(holo, SING). Fig. 28.

Roots below the pseudobulbs. Rhizome creeping, 1.5-2 mm diam., sections

between pseudobulbs 0.2-0.4 cm long, bracts not persistent. Pseudobulbs

New Species of Bulbophyllum (Orchidaceae) 141

E 20

t ed ————— —_———

Figure 28. Bulbophyllum croceodon J.J. Verm. & P.O’ Byme. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side, below:

abaxial side. All from SBG-O 4539 (spirit sample).

142 Gard. Bull. Singapore 60 (1) 2008

rather close together, almost porrect to obliquely erect, ovoid, 0.4-0.6 x 0.2-

0.3 cm. Petiole up to 0.2 cm long. Leaf blade elliptic to (ob-)ovate, 1.7-2.7 x

0.3-0.35 cm, index (length/width) 5-9; subacute. Inflorescence ca 4 cm long,

|-flowered. Peduncle patent, ca 2.7 cm, bracts 3, the longest ca 3 mm long.

Floral bract tubular, ca 2.8 mm, acute. Flowers not fully opening. Pedicel and

ovary ca 13 mm long, basal node on a ca 2 mm long stump. Median sepal

recurved, ovate, ca 5.5 x 2 mm, index 2.7-2.8; shortly acuminate, margins

entire, base rather broadly attached; rather thin, glabrous. Lateral sepals free,

oblique, ovate-triangular, ca 5.5 x 2.5 mm, index ca 2.2; acute, base broadly

attached; upper margin finely papillose; otherwise as the median sepal. Petals

porrect, obovate, ca 2.2 x 1.2 mm, index ca 2; rounded, margins finely erose

and finely papillose; base rather narrowly attached; thin, surface glabrous. Lip

recurved near its base, obovate, with a drawn-out ovate top part, ca 3.4 x 1.6

mm, index 2.1-2.2 (all without artificial spreading); obtuse, margins entire,

papillose except near the top part; thick and soft; adaxially concave near the

base, with two rather high and narrow ridges starting at the base, near the

. Margins, and running up, while slightly converging, to c. 4 of the length of the

lip, and ending with an small, erect, rounded tooth, middle '4 approx. flat, top

/3 distinctly convex, adaxial surface glabrous; abaxially with a retuse ridge up

to '4 of the length of the lip, surface glabrous except for a long-papillose patch

approx. half-way along the length of the lip. Column ca 1.6 mm long, stigma

elliptic, without keels inside, without teeth at its base, column foot widened,

with ovate, obtuse lateral teeth as well as a median swelling just above the

ligament. Stelidia triangular, ca 0.8 mm long, acute, with a slight, deltoid,

obtuse wing along the lower margin. Anther abaxially with a large, rounded

crest, surface finely papillose, front margin drawn out, rounded, papillose.

Pollinia (not seen; anther under-developed in the flower available).

Colour: Flowers yellow. Front part of the ridges on the adaxial side of the lip

orange.

Habitat & ecology: Found as an understory epiphyte in montane forest with

small pole trees, only thinly mossy. Alt. ca 1000 m. Flowering Jun, Jul, Oct.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Similar to B. stylocoryphe, mainly different in the wider lateral sepals,

petals and lip. Also, the top part of the lip is drawn out, but not contracted

before widening into a globose tip, as in B. stylocoryphe.

ae wena eee

Bulbophyllum stylocoryphe J.J. Verm. & P. O° Byrne. sp. nov.

A Bulbophyllis croceodon, B. elodeifloro, B. gilvo et B. ischnopodus /abelli

apice elongata, quod adducens ante dilatatione in apicem globosam difjert.

— Typus: Indonesia, Sulawesi, central part, SBG- O 5054 (holo, SING).

Fig. 29.

Roots below the pseudobulbs. Rhizome (short-) creeping or straggling

at some distance from the substrate, 1.2-1.5 mm diam., sections between

pseudobulbs 0.2-0.3 cm long, bracts not persistent. Pdendobalhs rather close

together, almost porrect to obliquely erect, ovoid to almost cylindrical, 0.6-0.9

x 0.2-0.25 cm. Petiole up to 0.3 cm long. Leaf blade elliptic, 2-3.5 x 0.25-

0.35 cm, index (length/width) 8-10; obtuse. Inflorescence ca 5 cm long.

1-flowered. Peduncle patent, ca 3.2 cm, bracts ca 2, the longest ca 3 mm long.

Floral bract tubular, ca 2.8 mm, acute. Flowers not fully opening. Pedice/ and

ovary c. 12 mm long, basal node on a ca 1.3 mm long stump. Median sepal

recurved, ovate, ca 6.5 x 2.2 mm, index 2.9-3; shortly acuminate, margins

entire, base rather narrowly attached; rather thin, glabrous. Lateral sepals free,

oblique, triangular, ca 6.5 x 2.5 mm, index ca 2.6; acute, base athe: broadly

attached: Se patiese finely papillose: gel ise as the median sepal. Petals

porrect, obovate, ca 2 x 1 mm, index ca 2; rounded, margins finely erose: base

rather broadly attached; rather thin, surface glabrous. Lip recurved near rue

base, elliptic-obovate, narrowed distally but with a swollen, globose tip, ca

4.1 x 1.4 mm, index 2.9-3 (all without artificial ; spreading), rounded, margins

entire, glabrous except for a short, ciliate stretch just above the base; thick

and soft; adaxially concave near the base, with two rather high and narrow

ridges starting at the base, near the margins, and running up, while slightly

converging, to '-'s of the length of the lip and ending with an erect, deltoid,

subacute tooth, top % slightly convex but tip distinctly convex, adaxial surface

glabrous; abaxially with a retuse ridge up to of the length of the lip, surface

glabrous except for a hairy patch in front of the ndge. Column ca 1.4 mm

long, stigma elliptic, without keels inside, without teeth at its base, column

foot widened, with triangular, obtuse lateral teeth as well as a median swelling

just above the ligament. Stelidia triangular, ca 0.8 mm long, acute, with a

slight, deltoid, obtuse wing along the lower margin. Anther abaxially with a

large, rounded crest, surface finely papillose, front margin drawn out, rounded,

papillose. Pollinia 4, ovate; the inner slightly shorter than the outer, flat; a

fleshy appendage is present.

Colour: Flowers yellow. Crest of the ridges on the adaxial side of the lip

orange yellow.

144 Gard. Bull. Singapore 60 (1) 2008

“wee eee =

Figure 29. Bulbophyllum stylocoryphe J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: abaxial side, right:

adaxial side; G. Pollinia, left: two pairs, right: two pairs with appendage, below: a single pair

with appendage. All from SBG-O 5054 (spirit sample).

New Species of Bulbophyllum (Orchidaceae) 145

Habitat & ecology: Found as an understory epiphyte in mossy montane forest.

Alt. ca 1200-1400 m. Flowering Feb, Apr. Jul, Oct, Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: In general habit and flower shape this is most similar to B. e/odeiflorum

J.J. Sm., B. gilvum J.J. Verm., and B. ischnopus Schltr., but none of these

species have the globose lump at the tip of the lip.

Group c - Species with a creeping rhizome, clustered or distant pseudobulbs

of substantial size, and inflorescences approx. as long as the vegetative shoot

they arise from.

The group agrees with sect. Megaloglossum. Other species found in

Sulawesi to be included here are B. tenuifolium (B/.) Lindl., and B. spissum

J.J. Verm.

Bulbophyllum entobaptum J.J. Verm. & P. O’ Byrne, sp. nov.

A Bulbophyllis aestivali et B. cubico /abello parte apicali angustiore, carinis

in pagina adaxiali e basi ad 3/5-plo labelli longitudine et a B. graciliscapum

petalis obtusis differt. — Typus: Indonesia, Sulawesi, central part, SBG-O

5050 (holo, SING). Fig. 30.

Roots below the pseudobulbs. Rhizome creeping, 1-1.5 mm diam., sections

between pseudobulbs 0.2-0.6 cm long, bracts not persistent. Pseudobulbs

clustered, approx. erect, ovoid, 0.7-1.4 x 0.4-0.8 cm. Petiole 0.2-0.5 cm long.

Leaf blade elliptic to ovate, 3.3-4.3 x 0.7-1.1 cm, index (length/width) 3.4-

4.8, acute. Inflorescence ca 6 cm long, |-flowered. Peduncle erect, 5-5.5 cm,

bracts ca 3, the longest ca 3 mm long. Floral bract tubular, ca 1.5 mm, acute.

Flowers not fully opening. Pedicel and ovary ca 10 mm long, basal node on a

ca 0.6 mm long stump. Median sepal approx. porrect, ovate, ca 8.5 x 4.5 mm,

index 1.8-1.9; acute, margins ciliate, base rather broadly attached; rather thin,

glabrous. Lateral sepals free, recurved, oblique, ovate-triangular, ca 8.5 x 3.5

mm, index 2.4-2.5, margins entire, glabrous, base broadly attached; otherwise

as the median sepal. Petals porrect, ovate, ca 5.5 x 2.1 mm, index ca 2.6; obtuse,

margins approx. entire, ciliate; base narrowly attached; thin, surface approx.

glabrous. Lip somewhat recurved approx. half-way along its length, general

outline ovate, ca 6 x 1.8 mm, index 3.3-3.4 (all without artificial spreading),

obtuse, margins papillose in the proximal half, thick; adaxially slightly concave

near the base, with two narrow ridges starting near the margins at the base, then

converge up to about 4 of the length of the lip, then diverge a little towards

their end at about 3/5 of the length of the lip, top 2/5 convex, adaxial surface

146 Gard. Bull. Singapore 60 (1) 2008

Figure 30. Bulbophyllum entobaptum J.J. Verm. & P. O'Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial |

side, right: abaxial side; E. Column and lip, lateral view; F. Anther, left: abaxial side, right:

adaxial side; G. Pollinia, left: a single pair, right: two pairs. All from SBG-O 5050 (spirit

sample).

eEenotn or tne

IViis tii Vi Lilt

400 m. Flov ering Feb. Apr-Nov.

. - - -

1Wawec) 2 |nira 2)

ulaWeSsl1, Cental palit (i

—— 4 - +

at Eee = es sae es

OF Similar looki 1g species W1in more O

= ae ee a ae! eS Bey E a Pe

al exists. all originating from the Philippines aes ale

ownrsas a29 = + tA =m »*h cnortor as]

Ames have a lip with much shorter Keel

a wider top part: B. graciliscapum Am

acute-acuminate petals and a glabrous lip.

aes, onae J.J. Verm. & P.O’ Byme, sp. nov

A Bulb oph

Roots below the pseudobulbs. Rhizome creeping, I-15 mm a sections

alct Pls eee oe a ee Ins tite >) ae) Pa

clustered, approx. erect, ovoid, 0.7-0.8 x 0.4-0.5 cm. Petiole 0.2-0.3 cm long.

Se = ee 959 x 7 254

Leaf Dlade Clliptic tO Ovale, - »-2.6 X Ca U Cm, index { Pea

acute. Inflorescence ca 3.5 cm long. 1-flowered. Peduncle erect. 2-3 cm

GLULL . Cc ta J.J Lili IVIiK,. i-LiUWOICU CUULILIC CIOL 27 Lill,

rte pn 2 +} ] 5 ee ie ~ | = 5 hr + toh, ar («wt —_ mm 2 ta

bracts ca 3, the longest ca 2.5 mm long al bract tubular, 1.5-2 mm, acute

Flowers not fully opening. Pedicel and ovary

a ca 0.5 mm long stump. Median sepal appre

148 Gard. Bull. Singapore 60 (1) 2008

4 mm, index 1.7-1.8, base broadly attached; otherwise as the median sepal.

Petals porrect, ovate, ca 4 x 2.5 mm, index ca 1.6; rounded, margins slightly

erose towards the tip, sparsely ciliolate towards the base; base narrowly

attached; thin, surface approx. glabrous. Lip somewhat recurved about half-

way along its length, general outline oblong, ca 5 x 1.8 mm, index ca 2.7-2.8

(all without artificial spreading), obtuse, margins sparsely and shortly ciliolate

in the proximal half, thick, surface glabrous; adaxially distinctly concave

in the proximal % of its length, bordered by two rather high and narrow

ridges starting at the base, and running up, close to the margin and slightly

converging, to about ” of the length of the lip, top 4 convex; abaxially with a

truncate ridge up to 2 of the length of the lip. Column ca 3 mm long, stigma

semi-elliptic, without keels inside, with a thick callus at its base, column foot

widened and thickened, with thick, retrorse, semi-elliptic, rounded lateral teeth

just above the ligament. Stelidia curved, subulate, ca 1.8 mm long, acute, with

an antrorse, triangular, obtuse tooth along the upper margin. Anther abaxially

with a rounded crest near its base, surface papillose, front margin drawn

-out, widely round, papillose. Pollinia 4, ellipsoid; the inner 4 as long as the

outer.

Colour: Sepals and petals pale yellow, adaxially densely spotted with purplish

red. Lip orange red, white near the base.

Habitat & ecology: Found as an understory epiphyte in montane forest near

a stream. Alt. ca1000 m. Flowering Jan.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: Morphologically, B. fionae seems most similar to B. elassoglossum

Siegerist. This has, however, much larger flowers (sepals 24 mm long, petals

10 mm long) and a lip which is papillose abaxially. B. entobaptum differs in

having a ciliolate median sepal, longer and narrower, long-ciliate petals and

a partly papillose lip.

Named after Fiona, granddaughter of Yunus, driver in Rantepao,

Sulawesi.

Bulbophyllum sect. Schistopetalum Schltr.

Bulbophyllum sect. Schistopetalum Schltr., Feddes Repert. Beth. 1 (1913) 700

& 763. LECTOTYPE SPECIES (here designated): Bulbophyllum

schistopetalum Schltr.

New Species of Bulbophyllum (Orchidaceae) 149

Figure 31. Bulbophyllum fionae J.J. Verm. & P. O’Byme. A. Habit; B. Flower; C. Flower

analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial side,

right: abaxial side; E. Column and lip, lateral view; F. Anther, above: adaxial side, below:

abaxial side; G. Pollinia, left: two pairs, mght: a single pair. All from SBG-O 4345 (spirit

sample).

150 Gard. Bull. Singapore 60 (1) 2008

Bulbophyllum barbasapientis J.J. Verm. & P. O’ Byrne, sp. nov.

Bulbophyllum barbasapientis J.J. Verm. & P. O’Byrne, ab omnibus sect.

Epicranthes petalorum appendices quae texturam eadem atque in petalis

propriis, labelli parte apicali dense pubescenti, ab omnibus speciebus sect.

Schistopetali stelidiis brevibus differt.—Typus: Indonesia, Sulawesi, central

part, SBG-O 4920 (holo, SING). Fig. 32.

Roots mostly produced close to the base of the rhizome. Rhizome patent to

hanging, ca 10 cm long, 1-2 mm diam., sections between pseudobulbs 0.7-1.6

cm long, bracts persistent. Pseudobulbs well spaced, ellipsoid to obovoid,

0.7-1.4 x 0.2-0.6 cm. Petiole 0.1-0.2 cm long. Leaf blade elliptic, 2.7-3.8 x

0.9-1.7 cm, index (length/width) 1.8-3.8; round to obtuse. Inflorescence ca

1.3 cm long, 1-flowered. Peduncle porrect, ca 0.4 cm, bracts 3, the longest ca

4 mm long. Floral bracts tubular at the base, ca 3 mm, acute. Flowers opening

wide. Pedicel and ovary ca 5 mm long, basal node on aca 1.5 mm-long stump.

Median sepal spreading, ovate, ca 5.5 x 3 mm, index 1.8-1.9; acute, margins

_ entire, base narrowly attached; thick, glabrous. Lateral sepals ca 5.5 x 3.3

mm, index 1.6-1.7; base broadly attached; otherwise as the median sepal.

Petals porrect, approx. lyriform, ca 1.4 x 1.6 mm excluding appendages, index

0.8-0.9; margins with 7 appendages, base broadly attached; thin, glabrous;

appendages linear, |.3-1.8 x ca 0.1 mm, gradually tapering towards an acute

tip, otherwise as the petals themselves. Lip slightly but abruptly recurved near

the base, approx. triangular, ca 3.5 x 1 mm, index ca 3.5 (all without artificial

spreading); top slightly drawn out, tip subacute, margins entire, ciliate along

a section near the base; thick; adaxially deeply concave proximally and with

two distinct, rounded ridges starting at the base, immediately converging and

then running parallel, close together up to 4-4 of the length of the lip, where

they are clasped between, and fused to the margins which converge over

the adaxial surface of the lip, then running close together and parallel over

another 3 of the length of the lip, surface convex distally, with densely placed,

woolly, long hairs; abaxially with a wide, retuse ridge up to 4-4 of the length

of the lip, surface glabrous but with long hairs towards the sides. Column ca

1.7 mm long, stigma obovate, without a tooth at its base, column foot without

teeth near the tip. Stelidia porrect, triangular, ca 0.4 mm long, acute, with a

denticulate upper margin, with a distinct, deltoid, subacute tooth which has

the front margin folded inwards along the lower margin, and a minute tooth

in front of this. Anther abaxially with a weak crest, surface otherwise finely

papillose towards the tip, front margin drawn out into a retuse beak with

approx. entire margins. Pollinia 2, ellipsoid, without appendage.

New Species of Bulbophyllum (Orchidaceae) |

—

Figure 32. Bulbophyllum barbasapientis J.J. Verm. & P. O’Byrne. A. Habit; B. Flower; C.

Flower analysis, from left to right: median sepal, petal, lateral sepal, lip; D. Lip, left: adaxial

side, right: abaxial side; E. Column and lip, lateral view; F. Column and lip, lateral view; G.

Anther, left: adaxial side, right: abaxial side; H. Pollinia, above: a single, below: a pair. All

from SBG-O 5052 (spirit sample).

152 Gard. Bull. Singapore 60 (1) 2008

Colour: Leaves dark green, suffused with purplish red. Sepals and lip purplish

red. Petals white, tips of filaments yellow. Column pale greenish.

Habitat & ecology: Epiphyte in montane riverine forest. Alt. ca 1000 m.

Flowering Jun, Aug-Nov.

Distribution: INDONESIA. Sulawesi, central part (1 specimen seen).

Notes: While fitting perfectly into section Epicranthes in most aspects, B.

barbasapientis is included in sect. Schistopetalum because the appendages

on the petals have the same texture as the petal itself. In sect. Epicranthes the

appendages have a texture differing from that of the petal. The configuration

of the keels on the lip in B. barbasapientis is typical for sect. Schistopetalum;

in section Epicranthes a similar but less distinct structure occurs. Within

section Schistopetalum, B. barbasapientis combines the lip shape of B.

schistopetalum Schltr with the short sepals of B. fissipetalum Schitr. It differs

from all species so far attributed to sect. Schistopetalum by its short stelidia

(as in sect. Epicranthes).

B. barbasapientis 1s amorphological link between section Epicranthes

and section Schistopetalum. The latter includes species with a more or less

pendulous rhizome (as in section Epicranthes) as well as species with a

creeping rhizome. The petals are deeply divided (as in sect. Epicranthes)

in most species, but with only 3 short appendages in B. trigonidioides J.J.

Sm., and with a single caudate tip in B. filamentosum Schltr. Species like B.

hassalii Kores, B. distichobulbum Cribb, and B. aristopetalum Kores, make

complete the morphological grading of section Epicranthes into the genus

Bulbophyllum in a wide sense.

Bulbophyllum sect. Stachysanthes (B1.) J.J. Verm. & P.O’ Byrne, comb. nov.

(= sect. Aphanobulbon)

Diphyes sect. Stachysanthes Bl., Bijdr. Fl. Ned. Ind. (1825) 312. —

LECTOTYPE SPECIES, designated here: Diphyes gibbosa BI. [=

B. gibbosum (BI.) Lindl.].

Diphyes sect. Diptychanthes Bl., Biydr. Fl. Ned. Ind. (1825) 311. — TYPE

SPECIES: Diphyes mutabilis Bl. [= B. mutabile (Bl.) Lindl1.].

Bulbophyllum sect. Aphanobulbon Schlitr., Feddes Repert. 10 (1912) 181;

Feddes Repert. Beth. 1 (1913) 704, 874. - LECTOTYPE SPECIES

(here designated): Bulbophyllum gibbosum (BI.) Lindl.

New Species of Bulbophyllum (Orchidaceae) 153

Not Bulbophyllum sect. Diptychanthes Rchb.f. in Bonplandia 5 (1857) 57. —

TYPE SPECIES: Bulbophyllum salaccense Rchb.f.

Further synonymy, as well as a selection of species included can be found in

Vermeulen (1991), under sect. Aphanobulbon.

Acknowledgements

The authors gratefully acknowledge the help of Dr. J.F. Veldkamp (L),

who prepared the Latin diagnoses of the species. We also thank the staff of

Singapore Botanic Gardens, especially Mr. Kiam Fee (Paul) Leong, who

has flowered many of the new species described here.

References

Averyanov, L.V. 1994. Identification guide to Vietnamese Orchids

(Orchidaceae Juss.). St. Petersburg.

Blume, C.L. 1825. Bijdragen tot de Flora van Nederlandsch Indié, 7.

Batavia.

Ormerod, P. 2002. Orchidaceae Fragmentae 4. Oasis 2,2: 4-7 & 10.

Schlechter, R. 1913 (1911-1914). Die Orchidaceen von Deutsch-Neu-

Guinea. Repertorium Specierum Novarum Regni Vegetabilis, Beihefte

1: I-LXVI & 1-1079.

Smith, J.J., 1926. Orchidaceae novae Malayensis 11. Bulletin du Jardin

Botanique de Buitenzorg series 3, 8, 1: 35-70.

Vermeulen, J.J. 1991. Orchids of Borneo 2. Kota Kinabalu and Kew.

Witte, L.C. de (in prep.). A taxonomic revision of the Bulbophyllum

species attributed to section Hymenobractea (Orchidaceae). Blumea:

in preparation.

Gardens’ Bulletin Singapore vol. 60 (1): 155-163. 2008 155

Studies on Schismatoglottideae (Araceae) of Borneo III:

Schismatoglottis confinis, a Putative Sister Taxon to

Schismatoglottis bauensis from Sarawak, Malaysian Borneo

WONG SIN YENG !' AND PETER C. BOYCE ”

'Faculty of Resource Science and Technology, Universiti Malaysia Sarawak,

94300 Samarahan, Sarawak, Malaysia

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

No. 18, Jalan Tunku Abdul Rahman,

93100 Kuching, Sarawak, Malaysia

Abstract

Schismatoglottis confinis S.Y.Wong & P.C.Boyce is described and illustrated

as a new species closely related to Schismatoglottis bauensis A.Hay &

C.Lee. An expanded description of S. bauensis is also presented together

with a key to separate the two species. Both species are illustrated.

Introduction

Schismatoglottis bauensis A.Hay & C.Lee is a lithophyte in humus and

litter pockets on boulders and cliffs of the Bau limestones, Kuching

Division (West Sarawak), where it is locally endemic and coexists with

Schismatoglottis nervosa Ridl., also endemic. Schismatoglottis bauensis

is placed in the Multiflora group sensu Hay & Yuzammi (2000) defined

by pleionanthic shoots and the adnate portion of the petiolar sheath short

but the remainder extended into long ligular portion. Schismatoglottis

bauensis is readily distinguished from the rest of the Multiflora group by

the absence of an appendix, always with pendent leaf laminae and striking

pinkish innovations.

While undertaking a survey of the limestones and adjacent

sandstones in the Serian (Sri Aman Division) and Padawan (Kuching

Division) areas the authors became aware of a taxon related to S. bauensis

present on both these limestones and, curiously, given the high level of

limestone-endemism displayed by Schismatoglottis in Sarawak, also on the

sandstones of Gunung Ampungan, Serian (Southeast Sarawak) (Sri Aman

Division). Detailed study of these collections revealed a species that while

clearly allied to S. bauensis, is separable on a number of key characters

156 Gard. Bull. Singapore 60 (1) 2008

as well as ecologically and geographically. It is herewith described as

Schismatoglottis confinis S.Y.Wong & P.C.Boyce.

Schismatoglottis bauensis A. Hay & C. Lee

Telopea 9(1) (2000): 84. — Typus: Malaysia, Sarawak, Kuching Division,

Bau, Gua Peri-peri (Fairy Caves), ca 7 km from Bau, 10 Mar 1994, PC.

Boyce 790 {holo - 2 sheets, K; iso, SAR (n.v.) ]. The isotype of S. bauensis

was not located during a search of SAR. Plate 1.

Medium to robust herbs to ca 75 cm tall. Stem pleionanthic, condensed

when young, later more-or-less creeping-ascending, ca 2-2.5 cm thick, bright

red internally, internodes to ca 0.5 cm long. Leaves to ca 8 together; petiole

terete, 30-55 cm long, always tinged reddish towards the base, glabrous,

sometimes densely scabrid, drying rust-brown, sheathing only at the extreme

base, the sheath extended into a bicarinate narrowly lanceolate free ligular

portion to 15-22 cm long, this drying dark brown; laminae ovate, 15-30 cm

- long x 7-15 cm wide, always pendent, glossy dark green adaxially, abaxially

paler and never glaucous, base obtuse and slightly decurrent, never cordate,

tip acute, acuminate for up to ca 3 cm; midrib raised abaxially (dry), adaxially

flush with the lamina, 1.5-5 mm wide, with 18-26 primary lateral veins on

each side, irregularly alternating with interprimary veins, diverging at 60°-

70°; secondary venation rather obscure, arising from the midrib and from

the bases of the primary veins; tertiary venation not visible. Inflorescences

1-4 together, pendant, each subtended by lanceolate prophyll resembling

the ligular leaf sheaths; peduncle to 10 cm long, not exceeding the prophyll.

Spathe 8-13 cm long; lower spathe, obliquely inserted to peduncle, straight,

3.5-5 cm long, green, differentiated from the limb by a weak constriction level

with the top of the spadix interstice; limb 5 cm long, green at first, becoming

white at anthesis, widely elliptic to oblongo-lanceolate, caducous, with a

tubular mucro up to 8 mm long. Spadix 6.5-11 cm long, subcylindric; female

zone 3-4.5 cm long, '/, of spadix length, adnate to the spathe in the lower °/,,

widest at side (ca 7 mm wide), narrowest at ventral and dorsal (ca 6 mm),

the free part slightly conoid, apically ca 4 mm diam.; pistils numerous and

crowded, subcylindric, ca 0.4 mm diam.; stigma sessile, about the diameter

to slightly wider than the ovary, button-like, papillate, staining light brown

in alcohol; interpistillar staminodes present, less than five, small, ca 4 mm

diam., slightly taller than pistils, staminodes or pistillodes (?) confined to a

single row along the spathe/spadix adnation, about the height and diameter

of the pistils, subcylindric, flat-topped, remaining white in alcohol; sterile

interstice ca 6 mm long, somewhat obconoid, white when fresh but staining

brown in alcohol, distally 6 mm diam., basally with pistillodes in more-or-

Studies on Schismatoglottideae (Araceae) 157

Plate 1. Schismatoglottis bauensis A.Hay & C.Lee. A. Overall habitat; B. Leaf laminae

with obtuse to slightly decurrent base; C. Emerging pinkish innovations; D. Inflorescence

with limb abcissed; E. Limb gaping at anthesis; F. Female anthesis.

158 Gard. Bull. Singapore 60 (1) 2008

less two rows and distally with staminodes, pistillodes of equal size and

height to pistils, staminodes of interstice crowded, irregularly polygonal, 0.5-

1 mm diam., flat-topped; male zone 4-6 cm long, 2 of spadix length, finger-

like, lower zone remaining white in alcohol, upper zone stained brown in

alcohol, basally isodiametric with top of interstice, tapering to a blunt point in

the upper half; stamens truncate, flat-topped, 0.5-0.7 mm across, 1.2-1.6 mm

long, somewhat irregularly rectangular with the connective wide, elevated;

pore punctiform, on the narrower edges of the stamen, ca 0.15 mm diam. with

fine tissue protruding from the inner most surface; appendix absent. Fruiting

spathe narrowly urceolate, 4 cm long, immature, ripening green and splitting

into irregular strips.Fruits white-green.

Other specimens seen: SARAWAK: Kuching Division: Bau, Jambusan,

Bukit Batu, 6 Apr 2004, P.C. Boyce & Jeland ak Kisai AR-20 (SAR + spirit);

Bau, Krokong, Gua Peri-peri, 01° 22’ 51.9”; 110° 07’ 09.3”, 29 Oct 2003,

P.C.Boyce & Jeland ak Kisai AR-146.1 (SAR); Bau, Kampung Bogag,

. Gunung Tibugai, 01° 21’ 31.1”; 110° 03’ 48.7”, 6 Jan 2005, P.-C. Boyce

& Jeland ak Kisai AR-949 (SAR); Bau, Kampung Duyoh, Sungai Duyoh,

01° 20’ 45.6”; 110° 02’ 36.9”, 8 Jun 2005, P.C.Boyce & Jeland ak Kisai

AR-1205 (SAR); Bau, Krokong, Gua Peri-peri, 01° 22” 51.9”; 110° 07’

09.3”, 15 Nov 2006, P.C.Boyce & S.Y.Wong AR-2053 (SAR spirit only);

Bau, Kampung Bogag, Gunung Tibugai, 01° 21° 31.1”; 110° 03’ 48.7”,

12 Jan 2005, P.C.Boyce et al. AR-961 (SAR); Bau, Krokong, Kampung

Tringeus, 01° 15° 40:2”; 110° 05° 35:97, 19: Feb 2005, P € Boyce ema:

AR-997 (SAR); Bau, Segong, Gunung Opar, 01° 27’ 07.3”; 110° 04’ 00.5”,

9 Nov 2005, P.-C. Boyce et al. AR-1503 (SAR + spirit); Bau, Jambusan, 8

Dec 2004, M.Gibernau AR-843 (SAR); Bau, Gunung Noka, 11 Oct 2004,

Jeland ak Kisai AR-725 (SAR); Bau, Kampung Jugan, 19 Jun 2004, Jeland

ak Kisai & Jipom ak Tisai AR-475 (SAR); Bau, Gunung Poing, 23 Sep

2001, Julia S. et al. SBC 346 (SAR); Bau, Bengoh Range, Logging Road,

Pangkalan Tebang, 6 Jul 1996, M.Mohizah, Yahud et al. S. 73891 (SAR);

Bau, | mile west of Bau, 6 Aug 1961, Dan H. Nicolson 1300 (SAR); Ibid.,

6 Aug 1961 Dan H. Nicolson 1303 (SAR).

Distribution: Borneo: Sarawak - endemic, known only from the vicinity of

Bau, Kuching Division, West Sarawak.

Habitat: Lithophytic in humus and litter pockets on limestone boulders and

cliffs at ca 10-100 m asl.

Notes: In the three inflorescences of S$. bauensis that the authors investigated

Studies on Schismatoglottideae (Araceae) 159

there is a zone of pistillodes at the base of interstice that is followed by

staminodes distally. However, the authors are undecided whether the sterile

zone at the basal of the spadix insertion is comprised of staminodes or

pistillodes. There is an additional character for S. bauensis, where the pores

of the stamens uniformly appear to have a fine tissue protruding from inner

most surface.

Schismatoglottis confinis S.Y.Wong & P.C.Boyce, sp. nov.

AbS. bauensis foliis subtus glaucis, laminae foliae basi semper decurrenti,

veneris laminorum seconadariis abaxialiter prominentis et nervis tertiariis

abaxialiter aliquantum tessellatis; poris antherae oblongo ca 0.3 mm diam.

ad paginae interiories antherae positis; connectivo planis, inforescentia

mascula cum alcoholis brunneus.—Typus: Malaysia, Sarawak, Samarahan

Division: Serian, Pichin, Tubih Durud, Ampon Siribu, 15 Dec 2004, Simon

Kutuh ak Paru AR-926 (holo, SAR). Plate 2.

Medium to moderately robust herbs to ca 70 cm tall. Stem pleionanthic,

condensed when young, later more-or-less creeping-ascending. Leaves to

ca 8 together; petiole terete, sometimes slightly D-shaped towards the base

of leaf laminae, 17-33 cm long, always tinged reddish towards the base,

densely scabrid or sometimes glabrous, drying rust-brown, sheathing only at

the extreme base, the sheath extended into a bicarinate narrowly lanceolate

free ligular portion to 15 cm long, this drying dark brown; laminae elliptic

to obovate, sometimes oblong, 20-24 cm long x 7-13 cm wide, dark green

and glossy adaxially, abaxially paler and glaucous, base always decurrent,

tip acute and acuminate for up to ca 2 cm; midrib raised abaxially (dry),

sometimes densely scabrid, adaxially flush with the lamina, 2-5 mm wide,

with 15-19 primary lateral veins on each side, these irregularly alternating with

interprimary veins and diverging at 60°-70°; secondary venation prominent,

sometimes discontinuous, arising from the midrib and from the bases of

the primary veins; tertiary venation often tessellate but sometimes obscure.

Inflorescences 1-4, pendant, together subtended by lanceolate prophyll

resembling the ligular leaf sheaths; peduncle to ca 8.5 cm long, not exceeding

the prophyll. Spathe 7-13 cm long, lower spathe, obliquely inserted from

peduncle, straight, 2.5-4 cm long, thickly coriaceous, green, differentiated

from the limb by a weak constriction level with the base of the interstice; limb

4.5-9 cm long, becoming white, caducous, oblongo-lanceolate, with a tubular

mucro up to 1.2 cm long. Spadix 5.5-10 cm long, subcylindric; female zone

2-2.8 cm long, '/; of spadix length, adnate to the spathe in the lower */,, widest

at side, narrowest at ventral and dorsal, the free part slightly cylindrical;

pistils numerous and crowded, subcylindric, ca 0.7 mm diam. x 1.2 mm long;

160 Gard. Bull. Singapore 60 (1) 2008

Plate 2. Schismatoglottis confinis Wong S.Y. & P.C.Boyce. A. Overall habitat; B. Leaf

lamina with decurrent base; C. Leaf abaxial surface glaucous; D. Pinkish red innovations

with dark red to deep purple petioles; E. Emerging inflorescence.

Studies on Schismatoglottideae (Araceae) 161

stigma sessile, about the diameter to slightly wider than the ovary, button-

like, papillate, staining light brown in alcohol; interpistillar staminodes, less

than five, small, ca 0.4 mm diam., slightly taller than pistils, staminodes or

pistillodes (?) confined to a single row along the spathe/spadix adnation, about

the height and diameter of the pistils, remaining white in alcohol, subcylindric,

flat-topped; sterile interstice 2.5 mm long, somewhat obconoid, distally 6 mm

diam., with pistillodes basally and staminodes distally, pistillodes of equal

size and height to pistils, white but staining brown in alcohol, staminodes

of interstice crowded, irregularly polygonal, flat-topped 0.5-1 mm diam.,

remaining white in alcohol; male zone 2.5-5 cm long, 2 of spadix length,

rectangular, basally isodiametric with top of interstice, tapering to a blunt

point in the upper half, staining brown in alcohol; stamens crowded, truncate,

flat-topped, somewhat irregularly rectangular with the connective wide, flat,

0.6 mm across x 1.4 mm long, the pores small, oblong, deep, on the inner

surface of stamens, ca 0.3 mm across; appendix absent. Infructescence not

observed.

Other specimens seen: SARAWAK: Kuching Division, Padawan, Kampung

Sadir, 2 Feb 2006, Simon Kutuh ak Paru AR-1695 (SAR); Padawan,

Kampung Sadir, Simpang Banyak, | May 2005, Simon Kutuh ak Paru AR-

1816 (SAR); Samarahan Division: Serian, Gunung Ampungan, 01° 09°

08.2”; 110° 37° 21.2”, 21 Nov 2003, PC. Boyce & Jeland ak Kisai AR-

146.2 (SAR spirit only); Serian, Gunung Ampungan, 01° 09° 10.1”; 110°

37° 26.2”, 28 Aug 2006, PC.Boyce & S.¥. Wong AR-2004 (SAR); Serian,

Pichin, Tubih, Tahang Sipukam, 01° 07’ 16.6”; 110° 26’ 51.2”, 26 Jul 2005,

P.C. Boyce et al. AR-1305 (SAR); Ibid., 26 Jul 2005, P.C.Boyce et al. AR-

1307 (SAR); Serian, Mongkos, Kampung Batuh, Gunung Selabur, 00°57’

26.2"; 110° 30° 15.8”, 15 Mar 2006, P-C.Boyce et al. AR-1732 (SAR);

Serian, Pichin, Sungai Bombo, 25 Nov 2004, Simon Kutuh ak Paru AR-

761 (SAR).

Distribution: West Sarawak, Kuching & Samarahan Divisions, endemic to

the Padawan/Serian areas.

Habitat: Always terrestrial mostly under full shade in deep soil on limestones

and sandstones, sometimes not in full shade. 250-568 m asl.

Notes: Schismatoglottis confinis appears to be closely allied to S. bauensis

but can be distinguished by the leaf abaxial surface, which is glaucous in S.

confinis but not in S. bauensis. The leaf base is always decurrent in S. confinis

but obtuse to slightly decurrent in S. bauwensis, while secondary venation is

162 Gard. Bull. Singapore 60 (1) 2008

prominent in S. confinis but obscure in S. bauensis. The tertiary venation of S.

confinis 1s often tessellate. Schismatoglottis confinis has pores that are oblong

(ca 0.3 mm), deep and located on the inner surface of stamens as compared

punctiform pores (ca 0.15 mm) with protruding tissue from innermost surface

and located at the edges of stamens in S. bauensis. The anther connective is

flat in S. confinis but elevated in S. bauensis while the male zone of S. confinis

stains brown in alcohol but remains white basally and stains brown distally

in S. bauensis.

Although S. confinis can be found on limestones it always occurs

terrestrially in deep soilas compared to S. bauensis which occurs lithophytically

in humus and litter pockets on limestone boulders and cliffs. Schismatoglottis

confinis can be found as well on the sandstones of Gunung Ampungan; the

distance between this locality and Padawan/Serian limestones is only ca 30

km. Additionally, based on known collections, S. confinis occurs at much

higher altitudes (near to 600 m asl) as compared to S. bauensis which occurs

at most at 100 m asl.

Schismatoglottis confinis and S. bauensis are most similar to

Schismatoglottis monoplacentaM.Hotta but differ among other characters by

lacking a spadix appendix. Schismatoglottis bauensis and S. monoplacenta

share a similar habitat; both occur lithophytically on vertical limestone.

Etymology: The specific epithet is derived from the Greek confinis, meaning

adjacent or adjoining, in allusion to the morphological similarity (and we

speculate phylogenetic closeness) of S. confinis to S. bauensis.

Key to the species

Schismatoglottis confinis can be fitted into the key to Bornean

Schismatoglottis (Hay & Yuzammi 2000) as follows:

lla. Male zone subcylindric; pollen sacs opening through a common pore

In, CaCH thea. .cs..scsedeceacashastescereecee sauce reetenert tee tare eee aen: coe ere 12

l1b. Male zone clavate to ellipsoid; pollen sacs opening through paired

pores m each thecal, 205.35. sccomereee erent Care reene acer eee eee 13

12a. Leaf lamina abaxially not glaucous, base obtuse to slightly decurrent,

secondary and tertiary venation obscure; pore at edge of stamen,

punctiform (ca 0.15 mm) with protruding tissue from innermost surface,

connective elevated, male zone remained white basally and stained

brown distally in alcohol. Lithophytic on limestones, Bau, Kuching

4 7 : a]

Studies on Schismatoglottideae (Araceae) 163

LU ISUTD A eee S. bauensis

12b. Leaf lamina abaxially glaucous, leaf base always decurrent, secondary

venation prominent, tertiary venation sometimes pronouncedly

tessellate; pore at inner surface of stamen, oblong (ca 0.3 mm),

connective flat, male zone completely stained brown in alcohol.

Terrestrial in deep soil on limestones and sandstones, Padawan, Kuching

Division and Serian, Samarahan Division ...............::::000+ S. confinis

13a in key = 14a, etc.

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular Datin Eileen Yen Ee Lee and

the Forest Research Centre (Kuching), notably L.C.J. Julaihi & Lucy

Chong. Thanks are due to Datuk Amar (Dr) Leonard Linggi Tun Jugah,

Graeme Brown & Dr Timothy Hatch of Malesiana Tropicals Sdn Bhd for

their support and funding of fieldwork in Sarawak. Many thanks to Simon

Kutuh Ak Paru who arranges our field works in Pichin, Serian. The first

author is grateful for the support provided by Faculty of Resource Science

and Technology, UNIMAS. This study is funded by the Ministry of Higher

Education, Malaysia under fundamental research grant scheme No.

FRGS/01(04)/609/2006(42).

References

Hay, A. and Yuzammi. 2000. Schismatoglottideae in Malesia I —

Schismatoglottis. Telopea 9: 1-178.

Musa yamiensis C. L. Yeh & J. H. Chen (Musaceae),

a New Species from Lanyu, Taiwan

CHING-LONG YEH ', JE-HUNG CHEN *, CHUAN-RONG YEH *

SHU-YING LEE *. CHIO-WEI HONG *, TSAN-HSIU CHIU °

AND YING-YU SU -

closely related to M. insularimontana Hayata, but differs from the latter

in subhorizontal inflorescence. vellow-green with pink at apex bracts. 4

flowers in a bract in | row. and the size and structure of flowers

The Musaceae as three genus, namely Musa L., Ensete Bruce and

Musella (Franchet) C.Y.Wu ex H.W.Li (Wu and Kress, 2000). No one

dee = gay

L .

: =< - 1 ace —s

knows for Sure the bis cise numbvde

apy eS rities now ¢ sh eee tae eee 2S 4

record, most authorities now five the num D O VIuUSaA SPECIES aS d-

7 S 4 - 1 = wy Pe a is

ico na the Niimpyer ot cneric \t 7 f 2 ta 7 y 1<

4? in 4 sections and the number of species OI Ensete as tO 9. .Musetida 1S

‘ = = 0 = See ee eee d ee ee

monotypic (Constantine, 2006). There is only one genus, Musa, recorded

in Taiwan (Ying, 2000). Recently, Ensete glaucum (Roxburg) Cheesman

was fou am in the mountain areas of Tainan and Pingtung

d Musa species are generally grouped into four sections. They

are me aw 1. Seer Musa and Rhodochlamys (Hakkinen. 2006).

Two species of Musa are recorded in Taiwan, M. formosana (Warb.) Hayata

and M. insularimontana Hayata (Yang et a/.. 2001

Recently a new species of Musa was found at Lanyu Island, Taitung

(Fig. 1). To honour the traditional name of the Lanyu people (the Yami).

166 Gard. Bull. Singapore 60 (1) 2008

we named this plant Musa yamiensis. This paper provides the description,

information on the habitat and photos of M. yamiensis, and a new key to the

species of Musa in Taiwan.

ung- Ai Br

eal

Lanyu

Taitung County E V2) "35

Figure 1. Distribution of Musa yamiensis in Taiwan.

Materials and Methods

The fresh materials were collected first by the 3rd and 4th authors on Isl.

Lanyu on 20 Jul, 2006. Its habitat and floral characteristics were recorded

with reference to the INIBAP Musa Description List (IPGRI-INIBAP/

CIRAD, 1996) by the Ist author. In order to get more information about

this new species, the authors took the second collection trip on | Aug, 2006.

Since it is difficult to obtain and prepare adequate herbarium materials of

species of Musa, especially the massive inflorescences (Constantine, 2006),

and due to the insufficient references on Musa taxonomy in our possession,

we gathered taxonomic papers of Musa mainly through the internet and

discussed the new species status of our newly collected specimens from |

Taiwan with Dr. Hakkinen, a world-famous Musaceae taxonomic specialist |

from Helsinki University Botanic Garden.

Musa yamiensis, a New Species from Lanyu, Taiwan 167

Key to species of Musa in Taiwan

Because of the rare field observed data of M. insularimontana Hayata,

our key to the species below follows the character treatment of M.

insularimontana in Ying’s (2000).

1. Male bract yellow-green with lightly pink at apex; male flowers 4

per bract in | row; male flowers’ compound tepal 2.5 cm long; fruits

cylindrical, slightly curved, 5.5-7 cm long; rachis glabrous...................

- cone sgzecceceh EEE Re ae M. yamiensis

1. Male bract yellowish red or dark purplish red; male flowers 8 or more per

OS SE Spe 2 es ee 2

. Male compound tepals about 3.5-4.3 cm long; fruits fusiform, 6-7 cm

ieee TELESIS PICCICE! PUDESCEDE «.2......2..----.2ccedee--eeeennreoee M. formosana

. Male compound tepals 3-3.5 cm long; fruits obliquely tetragonal, 8-11

cm long; fruiting pedicel densely pubescent ........ M. insularimontana

Taxonomic treatment

Musa yamiensis C.-L.Yeh & J.-H.Chen, sp. nov.

Caulis 2.5 m altus. Folia anguste oblongus, 138 cm longa, 40 cm /atus.

Inflorescentia glabra prope horizontalis. Spatha viridis, 27 cm longa, 5 cm

latus, lanceolatus, apex caudatus, glabra; bractea flavovirens, apex rutilus,

lanciolatus. Flores masculini in quoque bractea 4, seriatus. Polypetalus

pentalobus; longia polypetalus ad dimidium gamopetalus attingens.

— Typus: Taiwan: Lanyu Island, 20 July 2006, C_R. Yeh & C.W. Hong 4096

(holo, PPI). Figs. 2 & 3.

Pseudostems green, about 2.5 m high, base not swollen. Leaf blades

narrow-oblong, 138-150 cm long, 40 cm wide, rounded and asymmetric

at base; petioles 60 cm long, margins erect and basally with wings.

Inflorescence subhorizontal, about 46 cm or longer, | cm in diameter,

densely angled, glabrous, green; spathe green, 27 cm long, 5 cm wide,

lanceolate, apex caudate, glabrous; male bracts revolute, persistent, ovate-

lanceolate, 6-7 cm long, 3 cm wide, adaxially yellowish-green, lightly pink

at their rounded tip, abaxially green. Male flowers 4 per bract in a row,

cylindrical, peduncle white, 0.6 cm long, glabrous; compound tepal 2.5

cm long, | cm wide, white, 5-lobed, apex of outer 2 lobes with a short

hook-like appendage, inner 3 lobes almost as long as outer lobes or shorter;

168 Gard. Bull. Singapore 60 (1) 2008

Figure 2. Musa yamiensis C. L. Yeh & J. H. Chen. A. Inflorescence; B. Habitat; C. Male

bud and flowers; D. Bract, showing the color; E. Male flower; F. Fruits; G. Fruits and

seeds. Bar A = 10 cm; bar B = 2 cm; bar C, D, F, G= 5 cm; bar E= 1 cm.

Musa yamiensis, a New Species from Lanyu, Taiwan 169

50 cm

lcm

Figure 3. Musa yamiensis C. L. Yeh et J. H. Chen. 1. Habit; 2. Petiole, cross-section;

3. Bract; 4. Male flowers; 5. Compound tepal of male flower; 6. Free tepal of male flower;

7. Stamen of male flower; 8. Reduced gynoecium; 9. Fruit, cross-section; 10. Seeds.

(drawn from fresh materials).

170 Gard. Bull. Singapore 60 (1) 2008

free tepal membranous, 1.3 cm long, 0.7 cm wide, about 1/2 as long as

compound petal, boat-shaped, sulcate at outer side, (ovate and cordate

at base when pressed), keeled in the central inner side, apex apiculate;

stamens 5, anthers white, linear, flat, 1 cm long; filaments white, 1.2 cm

long; reduced gynoecium white, 2.2 cm long, not clavate. Fruits 8 per

cluster, green, 5.5-7 cm long, 1.5 cm wide, cylindrical, apex bottle-necked,

slightly curved, 3-4 angled, glabrous. Seeds numerous, black, irregularly

angled, smooth, 0.5 cm in diameter, 0.3 cm thick.

Other specimen examined: Taitung Co.: Jung-Ai Bridge, Lanyu, C. R. Yeh

& C.W. Hong 4100, Jul 20, 2006.

Distribution: In shaded ravine, 150-250 m, southern part of Lanyu Island

(Orchid Island or Botel Tabago).

Flowering and fruiting season: June to July.

Note: We found this musa in southern Lanyu’s rainforest where there are

two canopy strata. The upper canopy is dominated by Pometia pinnata,

Bischofia javanica, Pisonia umbellifera, and Ficus septica. The understory

is dominated by Angiopteris palmiformis, Elatostema edule, Donax

canniformis, Schismatoglottis kotoensis and Alocasia macrorrhiza.

Discussion

Despite its importance as a source of valuable food plants, the Musaceae

family is still incompletely known (Constantine, 2006). In the family,

taxonomic literature is insufficient and inconsistent. Take M. basjoo var.

formosana for example, the male bud is described as purplish red by Ying

(2000), but it is described as bronze yellow by Wu and Kress (2000).

According to our field observation, there actually exist these two kinds of

wild bananas in Taiwan main island. The wild populations can be separated

by taxonomic treatment into two or more species.

In Lanyu, so far, we could not find M. insularimontana. Due to its

rarity, some researchers even doubted its existence in Taiwan even though

that it has been described in publications. More and extensive wild banana

explorations in Taiwan would be needed.

The distinguishing characteristics of Taiwanese Musa species are

shown in Table 1. The specimens of Musa formosana examined are as

follows — Taipei, Wulai Co., C.R. Yeh, s.n. (1 Sep 2006); Pingtung, Latyi

Musa yamiensis, a New Species from Lanyu, Taiwan 17]

Co., C.R. Yeh, s.n., (17 Aug 2006) and Pingtung, Shoka, C.R. Yeh, s.n. (6

Sep 2006). The characters for Musa insularimontana are obtained from

Ying (2000).

Table 1. Differences among three species of Musa in Taiwan.

Character Musa formosana Musa Musa yamiensis

insularimontana

Rachis pubescent pubescent glabrous

Color of male bud _ yellowish red dark purplish red yellow-green,

more or less

tinged with pink

Number of male 10-13 in 2 rows 8 in 2 rows 4in 1 row

flowers in a bract

Color of male pale yellow unknown white

flowers

Compound tepals

3-3.5 cm long

2.5 cm long

Free tepals

1.5-2 cm long

1.3 cm long

Fruits

fusiform, 6-7 cm

long

obliquely tetragonal,

stipitate, 8-11 cm

long

cylindrical, slightly

curved, 5.5-7 cm

long

Fruiting pedicel

pubescent

densely pubescent

glabrous

Acknowledgements

We are grateful to Dr. Hakinnen for providing excellent research papers and

precious comments. We also appreciate H.-L. Chiu, National Plant Genetic

Resources Center, and Y.-C. Chen, Miaoli District Agricultural Research

and Extension Station, for assisting in our field study in Lanyu.

172 Gard. Bull. Singapore 60 (1) 2008

References

Constantine, D. 2006. The Musaceae: An Annotated List of the Species

of Ensete, Musa and Musella. See www.users.globalnet.co.uk/~drc/

Introduction.htm

Hakkinen, M. 2006 Musa lawitiensis Nausution & Supard. (Musaceae) and

its intraspecific taxa in Borneo. Adansonia 28: 56.

IPGRI-INIBAP/CIRAD. 1996 Description for Bananas (Musa _ spp.).

International Plant Genetic Resources Institute, Italy/International

Network for the Improvement of Banana and Platain, Montpellier, France/

Centre de Cooperation Internationale en Recherche Agronomique pour

le Developpement, Montpellier, France.

Wu, D-L.and W.J. Kress. 2000. Musaceae. Flora of China 24: 297-313.

Yang, Y.-P, H.-Y. Liu and T.-P. Lin. 2001. Musaceae. In: Manual of

Taiwan Vascular Plants, vol. 5: 201. The Council of Agriculture, Taipei,

Taiwan.

Ying, S.-S. 2000. Musaceae. In: Huang, T.-C. (ed.), Flora of Taiwan. 2"

ed. 5: 704.

Gardens Bulletin Singapore 60 (1): 173-174. 2008 173

Book Review: Maxwell, J.F. 2006. Vascular Flora of Ko Hong Hill,

Songkla Province, Thailand. Thai Studies in Biodiversity No. 6,

Biodiversity Research and Training Centre, Bangkok, Thailand. pp.472.

Price: US$9.80 or 350 Baht.

This attractive and inexpensive volume includes introductory chapters on

the ecology of Ko Hong Hill, a small low hill in the Haad Yai District,

peninsular Thailand. The main body of the text includes keys and

descriptions of the families, genera and 637 species of vascular plants

found there, references, and some line drawings and colour photographs.

There is a dearth of information on the flora of southern Thailand so this

volume is a welcome contribution.

The great strength of the book lies in author’s first-hand knowledge

of Ko Hong Hill, the meticulous collections he makes, and the care with

which they are identified. This makes for accurate and detailed descriptions.

The families are arranged in a natural order, which facilitates checking

identification. The descriptions are obviously based on author’s own

observations: they are detailed and accurate and include information often

lacking in floras, such as unripe and ripe fruit colours. The inclusion of

local names will make the work more accessible to Thai users. During the

course of the project one new species, Barringtonia rimata Chant., was

discovered. It is indeed a solid piece of original work.

In addition, it should appeal to a wider audience than those interested

in the flora of peninsular Thailand. The locality has been disturbed in the

past so its flora includes many of the widespread secondary forest species

from most stages in succession. This means that the selection of species

is relevant throughout, not only Thailand, but the region. In this respect

the key to the 130 families is particularly useful as rather few references

include this basic aid. Being based on the author’s own experience, it is

practical and straightforward to use.

In some cases, the author shows a reluctance to use current names

so Syzygium still appears as Eugenia, Chromolena as Eupatorium and

Spermacoce as Borreria, Dianella (Hemerocallidaceae) is still included in

the Liliaceae and Dracaenaceae as Agavaceae.

While the colour photographs are useful and enhance the

attractiveness of the book, the line drawings are sketchy and do not include

details of inflorescences and flowers, and why are they bunched together

at the end of the book instead of being placed next to the description of the

species in the text?

One serious error is that the Gesneriaceae is missing from the text

(although it is referred to in the Index) and this has resulted in errors in the

Gard. Bull. Singapore 60 (1) 2008

Index in page numbers of families close to where the Gesneriaceae should

have been. Other small errors include repeating Figure 21 twice and the

reference to Maxwell (1983) on page 289 being missing from the References.

Can Maxwell produce a book without including a Maxwellism? Who but

Maxwell (p. 134) would describe differences of opinion on genera limits

in terms of ‘outrageously taxonomically brutalized’, and some people

consider taxonomy boring - in some families it certainly sizzles!

This book is strongly recommended, not only to Thai students

and professionals, but also those in the Indo-Chinese region, who need

to identify plants. The Thai flora numbers over 8,000 species, far too

large to enable the beginner to begin to tackle naming plants and, in any

case, many families have not yet been revised for the Thai Flora Project.

Maxwell’s book is therefore an excellent introduction, as well as serving

as a basic reference. It deserves to be widely used. It is available from BRT

Secretariat, 5 Floor, NSTDA Building, 73/1 Rama VI Road, Rajdhevee,

Bangkok 10400, Thailand.

Ruth Kiew

Forest Research Institute Malaysia,

Kepong, Selangor

Malaysia

The Gardens’ Bulletin

Singapore

VOL.60 (2) 2009 . | ISSN 0374-7859

AWE

ie I, My, ‘ah a ON

A WV f ai ' “a

‘Sineapoee Botanic Gardens gi.

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:@2 Van

Singapore Botanic Gardens

(Editor)

DresP ye Tan

Singapore NParks Board

(Assistant Editor)

Dr. Jana Leong-Skorniékova

Singapore Botanic Gardens

(Assistant Editor)

Ms. C. Soh

Singapore Botanic Gardens

(Journal Business Manager)

EDITORIAL BOARD

Dr. S.C. Chin

Singapore Botanic Gardens

Dr. M.J.E. Coode

Royal Botanic Gardens

Kew, U.K.

Prof. Sir P. Crane

University of Chicago

Chicago, USA

Dri Re Corlett

National University of Singapore

Singapore

Dr. W.J. Kress

Department of Botany, NMNH

Smithsonian Institute

Washington DC, USA

Dr. M.C. Roos

National Herbarium Netherlands

Leiden University, The Netherlands

Dr. E. Soepadmo

Forest Research Institute Malaysia

Kepong, Malaysia

Prof. T. Stuessy

University of Vienna

Austria

Dr. W.K. Tan

National Parks Board, Singapore

Dr. I.M. Turner

Research Associate

Singapore Botanic Gardens

The Gardens’ Bulletin is published twice yearly by the National Parks Board, Singapore.

Neither the National Parks Board nor the Editorial Board is responsible for the opinions or

conclusions expressed by the contributing authors.

The annual subscription for the Gardens’ Bulletin is Singapore $100.00 including postage.

Overseas subscribers are required to make payment in the form of bank drafts or international

money orders in Singapore currency payable to National Parks Board, Singapore.

Instructions for contributing authors are found on the inside back cover.

[Cover photo: Livistona lanuginosa (see p. 302); photo by J.L. Dowe]

JUN 13 2009

ARNOLD ARBORETUM

UBRARY

The Gardens’ Bulletin

Singapore

VOL. 60 (2) 2009 ISSN 0374-7859

CONTENTS

Bogner, J. and P.C. Boyce

Studies on the Schismatoglottideae (Araceae) of Borneo VI: A new

Schismatoglottis species from Sarawak, Malaysian Borneo...............::ceccceseeeereeeteeeees 175

Dowe, J.L.

AMtaxonomie account On livistona REBT (ATECACEAC)) cccccc.c.--e0cc-sc00csceee.ess0.seenc-cacncocese 185

Ho, B.-C. and B.C. Tan

Does the moss genus Lepidopilum (Brid.) Brid. (Pilotrichaceae) occur in Asia? ........ 345

Kiew, R. and J. Sang

Seven new species of Begonia (Begoniaceae) from the Ulu Merirai and

Bukiisaranealimestone areas mi Sarawake BOGMCO) ccessec--.sec-eeseceerooceceocccesecetesceceevecrese-s SOI

Kurzweil, H., S. Chantanaorrapint and A.Buakhlai

A new species of Habenaria (Orchidaceae) from Southeast Asia...............::cccceeeeeeee 373)

Leong-Skornickova, J., M. Sabu and M.G. Prasanthkumar

Transfer of Amomum fenzlii, a Nicobar Islands endemic, to Etlingera

(ZLISIEROSTICSAS)s costteicedseecedeecBe see d8a00 3 cOe EE BSCE? ASSET Er EOE ee Oe 381]

Leong-Skorni¢kova, J. and A.K. Nura

WII @r En Moan@ultivataRe oisityyalecseeces:ceetecseeeestectees se ceeseecSoscs-tceceecscces nceesesseeeocuevsecceees 389

Wilson, P.G.

Conspectus of the genus Eugenia (Myrtaceae) in the Philippines ..................e ee 399

Book Review

Monkey Puzzle Man, Archibald Menzies, Plant Hunter by J. McCarthy

CS UA LES ees ee ree Nese ee ee ne SnD ee ee Nate Penaeus aes dacecunnsit vs dees stesiuebe pavtaese 41]

Date of publication: February 2009

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Photoplates Pte Ltd

Gardens’ Bulletin Singapore 60 (2): 175-183. 2009 IS

Studies on the Schismatoglottideae (Araceae) of Borneo VI:

A New Schismatoglottis Species from Sarawak,

Malaysian Borneo

J. BOGNER' AND P.C. BOYCE?

' Augsburgerstrasse 43a, D-86368 Gersthofen, Germany

> Malesiana Tropicals, Level 5, Tun Jugah Tower,

No. 18, Jalan Tunku Abdul Rahman, 93100 Kuching, Sarawak, Malaysia

Abstract

Fieldwork in Sarawak continues to reveal further novelties in the genus

Schismatoglottis Zoll. & Moritzi, notably in the Asperata and Multiflora

Groups (sensu Hay & Yuzammi, 2000). The second author collected a new

Schismatoglottis species in Sarawak belonging to the Multiflora Group.

This is here described and illustrated and included in additions to the key to

Bornean Schismatoglottis published by Hay & Yuzammi (2000).

Introduction

In recent years Schismatoglottis Zoll. & Moritzi has been the subject of a

major alpha-taxonomic revision (Hay & Yuzammi, 2000), which, together

with subsidiary papers describing additional Bornean novelties (Hay, 2002;

Hay and Herscovitch, 2003; Boyce and Wong, 2007; Wong, 2007; Wong and

Boyce, 2007), and with the caveat that numerous tropical Asian novelties

remain yet to be described, provides an alpha-taxonomy that is sufficiently

stable to facilitate the description of further novelties, as well as higher

level (molecular phylogenetic) and multidisciplinary (notably ecological

and phenological) studies. Fieldwork in Sarawak continues to reveal

further novelties in the genus Schismatoglottis, notably in the Asperata and

Multiflora Groups (sensu Hay & Yuzammi, 2000).

Schismatoglottis hayana Bogner & P.C. Boyce, sp. nov.

Petiolus brevior quam lamina, vagina ligula librera; lamina foliorum

lanceolate vel elliptica, apica longa acuminata vel caudata (1.5—2.5 cm

longa); 12-19 nervi laterales primarii in utraque parte nervi mediani.

Pedunculi breviores quam petioli; spadix subcylindrica, exappendiculata;

ambo sacci pollinis thecae apertura poro separata; connectivum planum.

176 Gard. Bull. Singapore 60 (2) 2009

— Typus: Malaysia, Sarawak, Kuching, Sematan, Teluk Selabang, 01° 58’

40.09”; 109° 38° 56.00”, 6 Oct. 2004, P.-C. Boyce, Jeland ak Kisai, Jipom

ak Tisai & Mael ak Late AR 703 (holo, SAR; iso, SING). From the same

collection: Bogner 2906 (para, M). Plates 1-3.

Medium herbs to ca 25 cm tall. Stem pleionanthic, erect, condensed, short

2-3 cm long, up to 3 cm in diam., upper part densely covered with several

glabrous leaves. Leaves to ca 8 together; petiole terete (slightly flattened

on upper side at the base) (8)10-12 cm long and 0.3-0.5 cm in diam., green,

tinged slightly reddish at base; sheathing only at the extreme base for 0.7-

1.0 cm, the sheath extended into a free ligular portion, to (5) 7-9 cm long,

this long-triangular, membranaceous when fresh, very soon becoming

dry and dark brown; laminae lanceolate to elliptic, 18-23 cm long x (4)5

-7 cm wide, dark green adaxially, lighter green abaxially, base cuneate,

apex long acuminate to caudate with a 1.5-2 (2.5) cm long tip; venation

parallel-pinnate, midrib strong and very prominent abaxially, slightly raised

-adaxially, primary lateral veins 12-19 on each side of the midrib, these

prominent abaxially, sunken adaxially; secondary and tertiary veins thinner,

all veins merging into a prominent marginal vein ca Imm in from the and

a further, more obscure, marginal vein between the thicker one and the

margin, abaxially this thicker vein often light brownish. Cataphylls 5-7 cm

long, membranaceous, becoming dry and brown soon. Inflorescences 1-5

together in a synflorescence, the spathe bent and held sub-horizontal (apex

of peduncle and base of spadix bent); the whole synflorescence subtended

by a long-triangular longitudinally two-keeled prophyll 5-7 cm long and

each subsequent inflorescence subtended by a long-triangular cataphyll

5-7 cm long, these membranaceous when fresh, soon becoming dry and

brown; peduncle shorter than petiole, 6-8(9) cm long, ca 4 mm cm in diam.,

terete, green, reddish at base. Spathe 8-9 cm long, lower spathe obliquely

inserted on the peduncle, glossy mid-green with irregular, slightly raised

longitudinal areas, 3-3.5 cm long x 2.3-1.4 cm diam., differentiated from

the limb by a moderately pronounced constriction level with the top of the

spadix interstice; limb 4-4.5 cm long, at first green, turning greenish white

at anthesis, oblong-lanceolate, inflating and then caducous, but occasionally

persisting in a decaying state until after the onset of male anthesis; apex

acuminate with a 0.8-1 cm long tubular mucro. Spadix sub-cylindric,

fertile to apex, 6-7 cm long; female zone adnate basally on one side to the

spathe for a length of ca 2 cm (2/3 to 4/5 of its length), female zone a little

thinner than the male zone, slightly obconic, 2.5-2.8 cm long, at adnation

0.8-0.9 cm in diam., apically 0.6-0.7 cm long in diam., at the base of female

flowers adjacent to the adnation sometimes a few pistillodes, these longer

Studies on the Schismatoglottideae (Araceae) of Borneo VI e/a

Plate 1. Schismatoglottis havana Bogner & P.C. Boyce. A. Mature plants at the type

locality on shale rocks; B. Plant in cultivation with almost mature inflorescence.

(Photo: 1A, Malesiana Tropicals Sdn Bhd)

178 Gard. Bull. Singapore 60 (2) 2009

Plate 2. Schismatoglottis hayana Bogner & P.C. Boyce. A. Plant in cultivation with

inflorescence at female anthesis (note the inflated spathe limb); B. Detail of the inflated

and open spathe limb at female anthesis; C. Plant with inflorescences at various sequential

stages. Note the near-most post-anthetic inflorescence has shed the spathe limb and spadix

above the female portion. (Photos: J. Bogner)

Studies on the Schismatoglottideae (Araceae) of Borneo VI 179

Plate 3. Schismatoglottis hayana Bogner & P.C. Boyce. A. Inflorescence at late male

anthesis. Note the strings of pollen and the decaying spathe limb; B. Inflorescence

just post anthesis. Spathe limb now fallen; C. Inflorescence with the persistent lower

portion artificially opened to reveal the female flowers and associated basal pistillodes.

Schismatoglottis multiflora Rid\.; D. Spadix artificially removed from spathe to show the

differences between it and that of S. hayana. Note particularly the larger, fusiform spadix

in S. multiflora. Schismatoglottis hayana Bogner & P.C. Boyce; E. pollen grain.

(Photos: 3A-C, G. Gerlach; 3D, Malesiana Tropicals Sdn Bhd; 3E, M. Hesse)

180 Gard. Bull. Singapore 60 (2) 2009

and slightly more robust than, but otherwise resembling, the gynoecia;

between the female and male zone a ring of rings 0.6-0.7 cm long and 0.6-

0.7 cm in diam., with 6-7 rows of staminodes; male part cylindric, fertile

to the tip, 3.5-4 cm long x 0.9 cm in diam., blunt. Female flowers densely

arranged; gynoecium sub-cylindric, |.1-1.3 cm long; ovary 1-1.2 mm long

and 0.5-0.7 mm in diam. (in the middle), whitish, placentation parietal,

ovules many and anatropous; stigma sessile, disk-like, 0.4-0.5 in diam.,

yellow, later becoming brownish; staminodes ca 1.2 mm tall, truncate,

yellowish, irregularly rectangular in view from above, ca 0.8 mm long

and ca 0.4 mm wide; stamens free, truncate, cream, |.2-1.3 mm tall and

ca. 0.5 mm in diam. at lower part, apically somewhat broader, irregularly

rectangular in view from above, ca 0.8 mm long and ca 0.4 mm wide, each

pollen sac opens with an own pore (2 pores for each theca), therefore, 4

pores on each stamen, connective flat. Pollen grains inaperturate, ellipsoid

to sub-reniform (one side more or less straight 17-19 um long and ca 13 um

wide, exine psilate (smooth). Fruits unknown.

Ecology: Rheophytic on shale stream banks in lowland evermoist riverine

forest, ca 40 m asl.

Etymology: Schismatoglottis hayana is named after Dr Alistair Hay,

formerly of the Royal Botanic Gardens, Sydney, who among his prolific

publications on Asian aroids co-authored with Yuzammi (BO) forming the

current standard revision of the genus Schismatoglottis of Malesia.

Distribution: Sarawak, only known from the type locality.

Notes: Schismatoglottis hayana belongs to the Multiflora Group (sensu Hay

& Yuzammi, 2000) by reason of the free-ligular leaf sheath, pleionanthic

shoots and the inflorescence held sub-horizontal by the bent peduncle,

and caducous spathe limb. The lack of a spadix appendix (1.e., the spadix

fertile to the tip) is shared within the group with S. bauensis A.Hay, S.

erecta M.Hotta, S. multiflora Ridl., S. roseospatha Bogner and S. schottii

Bogner & Nicolson, while thecae each dehiscing via two pores occurs in S.

nicolsonii A.Hay and S. mayoana Bogner & M.Hotta.

Vegetatively S. hayana most closely resembles S. multiflora (West

Sarawak & NW Kalimantan: obligately rheophytic ona variety of geologies),

S. bauensis (West Sarawak & NW Kalimantan: lithophytic/chasmophytic

on limestones), and an as yet undescribed putative sister species to S.

bauensis from SW Sarawak (Wong and Boyce, 2008). Schismatoglottis

hayana shares with these species slightly glossy leaves with primary lateral

Studies on the Schismatoglottideae (Araceae) of Borneo VI 181]

veins adaxially prominently impressed, a white spathe limb and a spadix

interstice with conspicuous staminodes.

Schismatoglottis hayana is readily distinguished from both S.

bauensis andS. multifloraby the thecae, each withtwo pores. Schismatoglottis

hayana is further distinctive from S. bauensis by the preference of shale

(vs. limestone) and rheophytic (vs. lithophytic/chasmophytic) ecology,

while S. hayana differs additionally from S. multiflora by the inflated (vs.

only loosened/unfurled) spathe limb, the proportionately shorter, thicker

cylindrical spadix and the larger pistillodes situated at the base of the female

flowers.

Thecae each with two pores also occur in S. nicolsonii and S.

mayoana (both in West Sarawak) but these differ from S. hayana by the

presence of a sterile appendix and in having leaves adaxially matte (vs.

glossy in S. hayana). Schismatoglottis nicolsonii and S. mayoana are

lithophytic on sandstones.

Schismatoglottis hayana is immediately distinguished from S.

erecta and S. schottii by the condensed stems with larger plants producing

few to several offshoots at the base (vs. internodes elongated and stems

loosely erect and little or not offsetting basally: S. erecta and S. schottii),

flat connective (vs. raised: S. schottii; grooved: S. erecta) and cylindrical

(vs. clavate) spadix.

Schismatoglottis roseospatha is distinguishable by the slightly

raised connective and much narrower and much glossier leaves in which

the primary lateral veins are flush with the adaxial surface (conspicuously

sunken in S. hayana). To date, S. roseospatha has been found only in the

Rejang valley.

Almost all species in the Multiflora Group so far observed (including

S. hayana) have the spathe inflating at male anthesis. However, S. multiflora

is unusual in the Multiflora Group in having a spathe that only loosens

at anthesis. Research on S. multiflora at the type locality by the second

author, co-workers and students, has revealed while vegetatively similar

in appearance to S. hayana, several unique floristic characteristics suggest

that S. multiflora maybe somewhat isolated in the Multiflora Group (Li ef

al, in prep).

Schismatoglottis hayana can be fitted into the key to Bornean

Schismatoglottis (Hay & Yuzammi 2000) as follows:

8a. Appendix absent or very reduced (a few terminal staminodes) ............. 9

cil Ee SACO PeTCIES el (0) 2 2 Un 14

182 Gard. Bull. Singapore 60 (2) 2009

9a. Plants rheophytic-2e5 eee a ee ee eee 10

9b. Plants not rheophytic-e =. ee ee ee 12

10a. Spathe inflated at anthesis; widespread in Bomeo .............:.sceseeeeeeees 1]

10b. Spathe unrolling slightly at anthesis but not inflated; Sarawak & W

K alutnncamitatils “ eiett feasedie eee te sao aR cca ee es 41. S. multiflora

lla. Spadix ca. 4.5 cm long; leaves abaxially very glossy; Rejang Valley

val a See ee es UE Ie 45. S. roseospatha

1lb. Spadix ca. 6-7 cm long; leaves abaxially slightly glossy; Sematan

FR ene ery ena ee En Foon sccbachcupaoccou. S. hayana

12a. Male zone sub-cylindric; pollen sacs opening through a common pore in

each theca; lithophytic on limestone; Bau, Sarawak ............. 34. S. bauensis

12b. Male zone clavate to ellipsoid; pollen sacs opening through paired pores

in €ach theca tonnes tial ...05 2.2.2. 20s dveetenceneaesreeeseeeeees eee tee ee 13

‘(13a in key = 14a, etc.)

Acknowledgements

The collaboration and support of the Sarawak Forestry Department, the

Sarawak Biodiversity Centre, in particular former Acting Director Datin

Eileen Yen Ee Lee and the Forest Research Centre (Kuching), notably L.C.J.

Julaihi and Lucy Chong is gratefully acknowledged. Thanks are due to

Datuk Amar (Dr) Leonard Linggi Tun Jugah, Graeme Brown & Dr Timothy

Hatch of Malesiana Tropicals Sdn Bhd for their continued support and

funding of fieldwork in Sarawak. We thank very much Professor Michael

Hesse, Wien, Austria, for the SEM micrographs of the pollen grains & Dr

H. Roessler, Miinchen for translation of the Latin diagnosis.

References

Bogner, J. and A. Hay. 2000. Schismatoglottideae (Araceae) in Malesia

I] - Aridarum, Bucephalandra, Phymatarum and Piptospatha. Telopea

9(1): 179-221.

Boyce, P.C. and S.Y. Wong. 2006. Studies on Schismatoglottideae (Araceae)

of Borneo I: A trio of new Schismatoglottis from Sarawak, Borneo.

Gardens’ Bulletin Singapore 58: 7-18.

Studies on the Schismatoglottideae (Araceae) of Borneo V1 183

Hay, A. 2002. A New Bornean Species of Schismatoglottis (Araceae).

Aroideana 25: 67-69.

Hay, A. and C. Herscovitch. 2003. A new species of Schismatoglottis

(Araceae) from Sabah. Gardens’ Bulletin Singapore 55: 27-30.

Hay, A. and Yuzammi. 2000. Schismatoglottideae (Araceae) in Malesia, I —

Schismatoglottis. Telopea 9(1): 1-177.

Wong, S.Y. 2007. Studies on Schismatoglottideae (Araceae) of Borneo, V:

Preliminary ecological observations of Schismatoglottis on the Matang

Massif, Sarawak, Malaysian Borneo. Aroideana 30: accepted.

Wong, S.Y. and P.C. Boyce. 2008. Studies on Schismatoglottideae

(Araceae) of Borneo, III: Schismatoglottis confinis, a putative sister

taxon to Schismatoglottis bauensis. Gardens’ Bulletin Singapore 60(1):

155-163.

Gardens’ Bulletin Singapore 60(2): 185-344. 2009 185

A Taxonomic Account of Livistona R.Br. (Arecaceae)

J.L. DOWE

Australian Centre for Tropical Freshwater Research,

James Cook University, Townsville, Qld 4811, Australia

Abstract

A taxonomic account of the palm genus Livistona is presented. Thirty-

SIX species are recognised. Taxonomic and nomenclatural changes are

applied to a number of species. Livistona rotundifolia (Lam.) Mart. is

treated as a highly variable species, with L. rotundifolia var. luzonensis

Becc., L. rotundifolia var. microcarpa (Becc.) Becc., L. rotundifolia var.

mindorensis (Becc.) Becc. and L. robinsoniana Becc. placed as synonyms;

L. fengkaiensis X.W.Wei & M.Y.Xiao is placed under L. speciosa Kurz;

Livistona chinensis var. subglobosa (Hassk.) Becc. is placed under L.

chinensis (Jacq.) R.Br. ex Mart.; Livistona tonkinensis Magalon is placed

under L. saribus (Lour.) Merr. ex A.Chev.; and Livistona kimberleyana

A.N.Rodd is placed under L. /orophylla Becc. A neotype is proposed for

L. saribus, and lectotypes are chosen for L. altissima Zoll., L. beccariana

Burret, L. hoogendorpii Hort. ex Teysm. & Binn. ex Miq., L. olivaeformis

(Hassk.) Mart., L. subglobosa (Hassk.) Mart., L. tonkinensis Magalon, L.

woodfordii Ridl., and Chamaerops biroo Siebold.

Introduction

Livistona R.Br. is one of the most ecologically diverse and widespread

genera of palms. It is distributed in the Horn of Africa and southern Arabia,

throughout southeastern and eastern Asia, Malesia and Australia (Fig.

1). Because of this extensive distribution, the genus has received uneven

taxonomic attention at the species level. In southeastern and eastern Asia, and

Malesia, there appeared to be an over-description of species, with a number

of taxa based on political boundaries (Dransfield, 1999). Considering that

the genus is of major horticultural and economic importance, as well as

ecologically diverse, it is timely to provide a revision of the genus over its

entire distributional range.

Livistona is included in the subfamily Coryphoideae, within the

tribe Trachycarpeae, and is one of five genera in the subtribe Livistoninae

(Dransfield et a/., 2008). Genera in the Livistoninae possess the unique

186 Gard. Bull. Singapore 60 (2) 2009

2.000 3,000

Figure 1. Distribution of Livistona, based on herbarium specimen data.

character of three basally free carpels that are united by their styles.

As circumscribed by Uhl and Dransfield (1987, p. 192), Livistona is

distinguished by: “...gvnoecium of three carpels connate only by their

styles, by united sepals, by petals with internal grooves, by the usually

small fruits with apical stigmatic remains and basal carpel remains, by

seed with homogeneous endosperm, and a large intrusion of seed coat.”

The closest related genus, Pholidocarpus Blume, is distinguished by a thick

conspicuous androecial ring that is almost free at the base (in Livistona

the androecial ring is scalloped apically and epipetalous), a top-shaped

gynoecium (in Livistona it is widest above the locules), and large fruit to 6

cm diameter at maturity (in Livistona usually 1-4 cm) usually with a corky-

warted epicarp (in Livistona it is always smooth). With regard to other

closely related genera, Livistona is distinguished from Licuala Wurmb

by lanceolate, apically tapered rather than wedge-shaped apically broad

leaf segments, while Johannesteijsmannia H.E.Moore has unsegmented

diamond-shaped leaves, an inflorescence with several inflated peduncular

bracts (in Livistona they are tightly tubular to loosely sheathing) and a

warty epicarp on the fruit (in Livistona it is always smooth). The generic

A Taxonomic Account of Livistona R.Br. (Arecaceae) 187

boundaries of Livistona based on morphological characteristics are well

defined (Tomlinson, 1990; Rodd, 1998). However, relationships inferred

from molecular data, of some genera within the Livistoninae and with

related groups, are not congruent with traditional classification (Asmussen

and Chase, 2001), and are presently unresolved (Dransfield et a/., 2008).

Taxonomic history

Livistona was established by Robert Brown (1810), and named for Patrick

Murray (born 1634?, died 1671), Baron Livingston, whose plant collection

at Livingston Estate near Edinburgh, was incorporated into the founding

collection of the present-day Royal Botanic Gardens, Edinburgh, established

in 1670 (Hyam and Pankhurst, 1995). The genus was first described with

two species, L. humilis R.Br. and L. inermis R.Br., from northern Australia.

Livistona humilis was subsequently chosen as the lectotype for the genus

(Moore, 1963a) and the specimen collected by Brown from Morgans Is., in the

Gulf of Carpentaria, was designated as the type specimen for that species.

Brown (1810) established the genus on the following characters:

flowers hermaphroditic; perianth in two-parts, each part with three

segments; stamens six, filaments distinct, basally broad; ovaries three;

style connate; stigma undivided; fruit single-seeded; endosperm with a

ventral cavity; embryo lateral; leaves palmate, and segment apices bifid.

Although Brown’s broad circumscription of Livistona can be applied to

other genera in the Livistoninae, Livistona, as it is presently understood, is

readily distinguished from related genera (Uhl and Dransfield, 1987).

Prior to establishment of Livistona in 1810, other species that would

eventually be included in Livistona had been named. The first was a pre-

Linnaean mononomial Saribus, described by Rumphius (1741), and which

was later to be recognised as Livistona rotundifolia (Lam.) Mart. The first

name in the Linnaean system was that provided by Lamarck (1786) who

described Corypha rotundifolia Lam. This species was transferred to

Livistona by Martius (1838). Corypha saribus Lour., described by Loureiro

(1790), was to become L. saribus (Lour.) Merr. ex A.Chev., while Latania

chinensis Jacq., described by Jacquin (1801), was to become Livistona

chinensis (Jacq.) R.Br. ex Mart.

Blume (1838) established the genus Saribus to include three species,

including the lectotype, Saribus rotundifolius (Lam.) Blume, based on

Corypha rotundifolia Lam. (= L. rotundifolia (Lam.) Mart.). Shortly after

Blume’s work was completed in 1836 (though not published until 1838,

Dransfield and Moore (1982)), Martius (1838) amended the description

of Livistona, subsumed Brown’s Corypha species, and Blume’s Saribus

species as well as some Corypha species of other authors. Subsequently

188 Gard. Bull. Singapore 60 (2) 2009

Martius (1849) also included the Livistona species described by Griffith

(1845) from India and Malaysia. The circumscription provided by Martius

firmly established the identity of Livistona, and was accepted by most

taxonomists (Miquel, 1855; Wendland and Drude, 1875; Hooker, 1883).

Treatments that proposed taxonomic changes were provided by Devansaye

(1875), Kuntze (1891) and Drude (1893).

Odoardo Beccari made a significant contribution to Livistona

taxonomy by describing many species in Australia, southeastern Asia and

Malesia. Beccari’s (1931) most detailed treatment of Livistona appeared in

Asiatic Palms - Corypheae.

A unique chapter in Livistona taxonomy was provided by herbarium-

based palm specialist, Max Burret (b.1883-d.1964), who described most

specimens received by him in Berlin as new taxa (Henderson, 1999), and

many of which have since been synonymised (Potztal, 1958), as they were

narrowly described on limited materials. Although never having visited

New Guinea, Burret (1935, 1939) described four Livistona species from

there, three of which were recently synonymised (Rodd, 1998). Burret

(1943) also established the genus Wissmannia, with the only species W.

carinensis (Chiov.) Burret, based on Hyphaene carinensis Chiov. from

Somalia (Chiovenda, 1929). This taxon was subsequently placed in

Livistona (Dransfield and Uhl, 1983a), thus providing an extension of the

genus both in geographical range and morphological diversity.

More recently, Rodd (1998), in his revision of the genus in Australia,

described five new species. One species was described for Vietnam (Nguyen

and Kiew, 2000), and four species for New Guinea and one for Australia

(Dowe and Barfod, 2001; Dowe and Mogea, 2004). Dowe and Jones (2004)

have recently applied nomenclatural changes to some Australian species.

Conservation

The WCMC (1993) report on palm conservation listed 69 entries for

Livistona, based on populations of 34 species defined by political

boundaries. The difference between the number of entries and the number

of species is because some widespread species were included in more than

one political region. Of the entries, 27 (39%) were of unknown status, 14

(20%) were not threatened, 16 (23%) were rare, and there were four entries

(6%) in each of the intermediate, vulnerable and endangered categories.

The high percentage of taxa in the unknown category indicated a lack of

information regarding the populations of many species. In Australia, all

Livistona species have been allocated a conservation rating (Rodd, 1998),

and Livistona mariae, from Palm Valley, Northern Territory, has been

the subject of a conservation action plan (Leach, 1992). In southeastern

A Taxonomic Account of Livistona R.Br. (Arecaceae) 189

Asia and Malesia, various species have received conservation ratings and

protection plans (Basu, 1991; Kiew, 1991; Madulid, 1991; Mogea, 1991:

Lim, 2002, 2004; Lim and Chan, 2002). For Malaysia, Kiew (1989) applied

a vulnerable/endangered status to L. saribus and rare but not threatened to

other Livistona species for that area. The endangered status of L. carinensis

in the Horn of Africa and Yemen has been highlighted by Lucas and Synge

(1978), Hedberg (1979), Bazara’a et al. (1990), Welch and Welch (1998,

1999), and Ford and Bealy (2004).

A conservation status is herein applied to all species based on IUCN

categories (IUCN, 2006). Some species of Livistona have been included in

the most recent Red List of Threatened Species, and these are appropriately

referenced. Established ratings have been reviewed and where necessary, a

revised status has been suggested.

Materials and Methods

The morphological species concept by Stuessy (1990) and Kanis ef

al. (1999) was used in this treatment to define species limits. Although

systematists differ as to the precise definition of this concept, here it is

based on the recognition of basic taxonomic units determined by readily

observable morphological discontinuities that ultimately reflect biological

limits and genetic divergence.

Abbreviations for publications and journals are based on Lawrence

et al. (1968) and Stafleu and Cowan (1976-1988). Herbarium acronyms are

according to Holmgren ef a/. (1990), and author abbreviations are based

on Brummitt and Powell (1992). Nomenclatural rules as in McNeill er ai.

(2006) were followed. In the taxonomic citations, pre-Linnaean names are

placed in square brackets.

About 840 specimens were examined from the following herbaria:

A, AD, AAU, B, BM, BO, BR, BRI, CAHUP, CANB, CANT, DNA, FI, FT,

FTG, HN, JCT, K, KEP, L, LAE, LBC, M, MA, MAK, MEL, NSW, NY, P,

PERTH, PNH, QRS, S, SING, UC, US. Appropriate parts and organs were

measured and the data incorporated into the species descriptions. To obtain

data that were not available from herbarium specimens, fieldwork was

undertaken in Australia, Papua New Guinea, the Philippines and Thailand.

To further expand the descriptive database, reference was also made to

species descriptions in the available literature. Distribution maps and ranges

were based on the localities recorded on the labels of the specimens that

were examined and other localities reported in literature, but this excluded

cultivated specimens.

190 Gard. Bull. Singapore 60 (2) 2009

Morphology

Stem

All Livistona species have a single, more or less, erect stem, except where

damage has induced either aerial branching or basal branching. Rodd

(1998) suggested that at least one species, L. inermis, had basally clustered

stems, but field observations indicate that this situation is caused by

damage, possibly from fire, and is not the natural condition for that species.

Stem heights range from 5 m tall in L. exigua (Plate 2A) to 45 m tall in L.

rotundifolia (Plate 3A) and with a diameter at breast height of from 2.5 cm

in L. exigua to ca 60 cm in L. nasmophila and L. saribus. Stem surfaces may

be smooth, as in L. rotundifolia (Plate 3B), or have longitudinal fissures of

varying width and depth, as in L. australis (Plate 21A) and L. chinensis

(Plate 9C). Leaf scars, where leaves were formerly attached to the stem,

are annular around the stem, usually of irregular width, and may be close

together and appearing contiguous, as in L. fu/va (Plate 22D), or up to 30

cm apart, as in L. decora (Plate 20A). Leaf scars may be raised, or flush

~with the internodes. In some species such as L. chinensis, L. humilis and

L. muelleri (Plate 15A), the leaf scars are roughened, retaining some leaf-

base tissue. In some species, such as L. humilis (Plate 14A), L. muelleri

(Plate 1SA) and L. tahanensis (Plate 11A), the petiole base can be retained

as a stub. This is very pronounced in L. benthamii (Plate 18B), where the

petiole base and the proximal half of the petiole is retained on the stem,

only to be removed by weathering, fire or old age. In many species, the

petiole stubs remain persistent on the stem up to the position that flowering

commences, and from that point upward the stubs are not retained and the

stem is more or less smooth.

Leaf

Leaves are produced in globose, hemispherical, elongate, vertically extended

or compact crowns. Leaves have a clasping leaf-base, a long petiole and a

segmented blade radiating from a hastula (Fig. 2). Leaf-bases are not split

at the base, as in some other Coryphoid palms such as Corypha, but are

entire. Fibres of various textures and thickness form ligule-like appendages

on either side of the leaf-base. These fibres can be diagnostic in some

species, whereas in many they are undifferentiated and/or variable and of

little diagnostic value.

The petiole, up to 300 cm long in L. concinna (Plate 18E, 18F), and

as less as 40 cm long in L. exigua (Plate 2B, 2C), narrows toward the leaf

blade, is either convex or concave on the abaxial surface, and convex on the

adaxial surface. In some species there may be a stripe extending along the

abaxial surface, this being yellowish-green in L. australis (Plate 21B) and

A Taxonomic Account of Livistona R.Br. (Arecaceae) 19]

apical cleft

lamina outline

free portion

central

segment

lateral

segment

segment length

hastula

Figure 2. Composite half-leaf of Livistona. Three segments are shown — central, lateral

and basal - indicating from where on the central segment the length, free portion and apical

cleft were measured. Both the depth of the free portion and the depth of the apical cleft are

determined, for comparative purposes, as a percentage of the length of the segment.

reddish-purple in L. saribus (Plate 1B). The petiole base in L. drudei (Plate

19C) is characteristically purple. The petiole margins are armed with spines

in most species. Spines are of varying shapes, but mostly shark-tooth like,

or retrorsely recurved. In some species, such as L. saribus (Plate 1C) and L.

192 Gard. Bull. Singapore 60 (2) 2009

speciosa (Plate 12C), the spines resemble the shape of the traditional Thai

dagger, the kris. Spines are usually ca 20 mm long, but may reach up to 60

mm long in L. saribus (Plate 1C). In L. inermis, the spines are reduced to

bluntened calli up to 5 mm long, but usually only ca | mm long (Plate 16E).

In some species, such as L. chocolatina, only juveniles have spines, but are

not produced, or are very reduced, in mature individuals. Spines are most

often congested in the proximal portion of the petiole in most species, but

may extend for the full length in others, such as L. rotundifolia.

The petiole terminates in a hastula, from which the leaf segments

radiate in most species. In some species, such as L. decora (Plate 20B) and

L. inermis (Plate 16E) the segments extend along the costa producing a

pseudo-pinnate appearance. The hastula may have a vertically erect flange

of tissue, such as in L. rotundifolia, or the flange may be rudimentary as in

L. halongensis (Plate 9B).

The lamina is costapalmate in most species, and is palmate in

only L. exigua (Plate 2A, 2B). The costa, a much thickened rib that is

“morphologically an extension of the abaxial surface of the petiole, may be

strongly arcuate as in L. Januginosa (Plate 25A), or almost straight as in L.

muelleri (Plate 15B). The outline of the lamina may be circular, or nearly

so, as in L. rotundifolia (Plate 3B), to oblong or ovate in most species.

The length of the lamina is interpreted here as the length of the midleaf

segment when measured from the hastula to the apex of the segment, 1.e.,

in a line extending directly outward from the petiole, and parallel to it.

Segments are induplicate. The number of segments ranges from 25 or less

in L. brevifolia (Plate 4C, 4D), up to 30 in L. exigua (Plate 2B), and to 90 or

more in L. australis (Plate 21B), L. lanuginosa, L. papuana (Plate 4B) and

L. surru (Plate 8B) among others. Segments are arranged in either of two

ways: firstly in a regular arrangement 1n which the segments are widest in

the mid section and become decreasingly less wide and usually longer or

infrequently shorter toward the outer-leaf, and with the point of disjunction,

herein termed divisions, between the segments terminating at more or less

the same position in the lamina relative to the length of the segments. The

second type of arrangement, which occurs in only two species, L. saribus

(Plate 1B) and L. exigua (Plate 2B), has the segments of similar width

and dimensions as mentioned above, but the segments are arranged in

groups, and there is a deep division between the groups that extends almost

to the hastula. In the first mentioned arrangement of segments, divisions

between the segments may extend for up to 98% of the lamina length, as

in L. lorophylla (Plate 17B), and as shallow as 34% in L. /anuginosa (Plate

25A). In this revision, the depth of the division between the segments and

the depth of the apical cleft are recorded as percentages of the length of the

A Taxonomic Account of Livistona R.Br. (Arecaceae) 193

segment, rather than absolute ranges of measurements, as this allows greater

direct comparison between species in what is a good diagnostic character in

mature leaves. In leaves that have many deep divisions between segments,

the apical portion of the segment may be pendulous, such as in L. chinensis

(Plate 9D), L. concinna (Plate 18D), L. decora (Plate 20B) and L. drudei

(Plate 19B). The segment apex is bifurcate, with the depth of the cleft as

shallow as 1% of the segment length as in L. brevifolia (Plate 4D), to as

deep as 84% in L. inermis (Plate 16E). Species that have deep apical clefts

tend to have pendulous segment apices, although those in L. inermis (Plate

16E) have somewhat rigid apices because of the relatively thick midribs.

Venation of a segment consists of a dominant central midrib,

accompanied by a series of parallel veins of more or less equal number each

side of the midrib, and transverse veins of various thickness and lengths, that

connect, girdle or traverse the parallel veins (Fig. 3). The number of parallel

veins is more or less consistent for a species. There are as many as 38 in L.

carinensis (Fig. 3A), and as few as nine in L. chinensis (Fig. 3D). There is a

relationship between the number of parallel veins and environment: species in

moist, humid environments tend to have fewer parallel veins, whilst species

in semi-arid to arid environments have relatively more.

Figure 3. Venation patterns in Livistona. Example of the variation for the genus present

in four species: a. L. carinensis — a large number of parallel veins and very few transverse

veins; b. L. mariae — a moderate number of parallel veins, though very prominent, and a

moderate number of transverse veins; c. L. australis — less prominent parallel veins and

prominent transverse veins; d. L. chinensis — less prominent parallel veins but numerous

transverse veins.

Inflorescence

Inflorescences are interfoliar, initiated in the axil of each leaf at the onset

of reproductive maturity. Leaves and inflorescence buds are produced

more or less regularly throughout the year. Leaf growth is continuous but

inflorescence buds remain dormant within the leaf axil until the onset of the

flowering season. With the onset, buds that were previously initiated develop

rapidly to facilitate more or less synchronised or closely sequential acropetal

inflorescence production. From | to 15 inflorescences can be produced in

194 Gard. Bull. Singapore 60 (2) 2009

a flowering event, with a period of a few weeks between maturation of the

proximal and distal inflorescences. Most species flower seasonally at more

or less the same time each year. Some species may flower aseasonally.

There are two types of inflorescences in Livistona. Type 1 has a

single dominant axis (Fig. 4a), whilst Type 2 is trifurcate, or occasionally

bifurcate, with three (or sometimes only two) collateral equi-dominant

axes that emerge from a single base and have a common prophyll (Fig.

4b). Partial inflorescences are positioned along the length of each axis, at

more or less regular intervals. The partial inflorescences may be interpreted

as discrete panicles that terminate in flower-bearing rachillae, and reduce

in size and complexity toward the inflorescence apex. There are 11 or

more partial inflorescences in species such as L. carinensis (Plate 13B),

L. decora (Plate 20A), L. lanuginosa (Plate 25A) and L. mariae, and with

the average number across all species being ca 6. There 1s one species, L.

humilis, which has only a single partial inflorescence, but only in female

plants (Plate 14C); male plants have numerous partial inflorescences (Plate

-14B). The structure of the partial inflorescences is similar in most species.

Inflorescences may be branched up to 5 orders, when the primary axis (Fig.

4a) is considered as ‘0’. The surfaces of the partial inflorescences may be

variously glabrous, papillose, or tomentose. The rachillae of L. carinensis

(Plate 13D) are unique within the genus as they are covered in sparse long

unbranched hairs. In L. chocolatina, there is a dense chocolate-brown

tomentum in the proximal portion of the partial inflorescence branches.

The prophyll is bicarinate in all species, but of various lengths and

widths, and the outer surfaces are furnished with various types and densities

of tomentum, or are glabrous. There is a single peduncular bract in 14

species, including L. boninensis, L. carinensis, L. eastonii, L. fulva and L.

mariae, among others; and as many as 4-8 in L. chocolatina, L. halongensis

and L. humilis. The peduncular bract may also be lacking, as in 16 species,

such as L. australis, L. benthamii, L. chinensis and L. saribus, among

others. Peduncular bracts do not subtend a partial inflorescence, and are

confined to the portion of the peduncle that is proximal to the emergence of

the lowest partial inflorescence. Each partial inflorescence is subsequently

subtended by a single rachis bract that is, otherwise, morphologically

similar to the peduncular bracts. Peduncular bracts are tightly tubular to

loosely sheathing, and glabrous or variously tomentose. In L. /anuginosa

(Plate 25B), the tomentum is very dense and fur-like. Rachis bracts are

tightly tubular to loosely sheathing, glabrous or variously tomentose.

Rachillae are straight to moderately flexuose, pliable to rigid,

glabrous or variously tomentose, papillose, pubescent or pruinose. They

are as short as | cm long in L. inermis (Plate 16F, 16G), and up to 45 cm

A Taxonomic Account of Livistona R.Br. (Arecaceae) 195

a partial inflorescence

primary axis

Figure 4. Inflorescence types in Livistona: a. Type | — inflorescence has an unbranched

primary axis with partial inflorescences placed at + regular intervals and reducing in

size and complexity toward the distal end; b. Type 2 — inflorescence has a trifurcate or

infrequently bifurcate axis with + identical collateral axes that share a common prophyll

but each axis with an individual peduncular bract. The individual axes otherwise have a

Type | structure.

long in L. saribus, and 2.5 mm diameter in L. exigua (Plate 2D) and to 9

mm diameter in L. chinensis (Plate 9E, 9F).

Floral morphology

Flowers are arranged spirally on the rachillae, solitary or in pairs, or in

sympodial clusters of 3-8, widely spaced or congested, and either sessile

or on pedicels. They are bisexual or functionally unisexual with little

dimorphism, a situation that will be discussed below under sexuality.

There are three triangular sepals fused at the base, glabrous or with long

hairs in a single species L. carinensis (Plate 13D); three free petals that

are longer than the sepals, triangular, valvate, glabrous or hairy as only

in L. carinensis (Plate 13D). There are six epipetalous stamens that are

196 Gard. Bull. Singapore 60 (2) 2009

shorter than the petals, and are basally connate to form a narrow fleshy

ring. Filaments are ‘shouldered’ and narrow abruptly into the connective.

Anthers are didymous, roundish to ovoid, medi-fixed, versatile, and open

latrorsely. The gynoecium in tri-carpellate; carpels are wedge-shaped and

basally separated, and connate distally to form a common slender style.

The style is shorter than the stamens. The stigma is small and 3-lobed. The

ovule is basally attached and anatropous.

The colour of sepals and petals in most species 1s white to cream to

yellow; however both sepals and petals can be red, as exclusively in some

New Guinea species such as L. brevifolia (Plate 4E) L. chocolatina, L.

tothur (Plate 7B) and L. woodfordii; sepals can be maroon as in L. muelleri

(Plate 15C, 15D); and petals can be bright yellow as in L. merrillii and L.

muelleri (Plate 15D). The length of sepals ranges from 0.2 mm to 4 mm,

and for petals 0.4 mm to 4 mm.

Septal nectaries have been reported in L. humilis (Schmid, 1983).

The nectaries are situated at the base of the partitions between the carpels.

‘Nectaries have not been reported in other species.

Pollen is monosulcate, ellipsoidal, and in those species that have

been studied, dimensions fall within a range of the long axis of 154m to

Figure 5. Pollen micrographs of Livistona species: a. L. chinensis; b. L. decora; c. L.

lanuginosa; d. L. muelleri. Scale bar = 10um.

A Taxonomic Account of Livistona R.Br. (Arecaceae) 197

44 um, and the short axis of 9um to 37um (Dowe, 2001) (Fig. 5). Tectum

ranges from finely perforate to reticulate. Livistona pollen is binucleate at

the time of shedding (Mahabale, 1967).

Fruit

The fruits contain a single seed, and are globose, subglobose, ellipsoidal,

depressed-globose, obovoid, reniform, pyriform, or obpyriform. The fruit

may be as small as ca 9 mm long, as in L. benthamii, L. exigua (Plate 2D),

L. victoriae (Plate 17D) and L. woodfordii (Plate 5F); to 40 mm long in L.

alfredii; 36 mm in L. lanuginosa (Plate 25E), 43 mm in L. tothur (Plate 7D);

and the largest to 65 mm in L. surru (Plate 8E). The epicarp is thin or thick,

smooth or with scattered pores or mildly rugose. The stigmatic remains are

apical to subapical and there is a suture line extending longitudinally from

the stigmatic remains toward the base in most species or sometimes this is

not readily visible. Colour at maturity can be shades of green, orange, red,

blue, purple, brown or black. The mesocarp is dry or fleshy, and with or

without embedded fibres.

Seed

The seed is globose, subglobose, ovoid, ellipsoid or reniform. The

endosperm is homogeneous, and is intruded by the testa from one side

with spongy/crystalline tissue to displace a moderate or large proportion

of the endosperm. The hilum is broad, circular to elongate, and the raphe

branches are few or lacking. The embryo is small, and positioned supra- to

sublateral.

Sexuality

Sexual expression in Livistona is not well understood. Uhl and Dransfield

(1987, pp. 190, 191) described the genus as “hermaphroditic (rarely

dioecious)” and “where dioecious, anthers or ovules not developing

but otherwise as in the hermaphroditic’. In a review of pollination in

palms, Henderson (1986) reported that Livistona flowers possessed septal

nectaries and that bee pollination was possible. Rodd (1998, pp. 72, 73)

stated that “Some of the Australian species display clear signs of sexual

differentiation among the palms of a population, although the nature of

this sexuality is still unclear”, and that “it is possible, indeed, that this is

a feature of all the Australian species, or even of the genus as a whole. It

remains an interesting area for further study”.

Dowe (2001) concluded that there was a range of sexual

functionality in the genus, from hermaphroditic species such as L.

rotundifolia and L. chinensis, to functional dioecy, such as in L. decora

198 Gard. Bull. Singapore 60 (2) 2009

and L. /anuginosa, and dioecy, such as in L. concinna and L. humilis.

Where there is reasonable evidence available of sexuality, it is noted in

the species descriptions, but pending further study sexuality is otherwise

noted as hermaphroditic.

Phylogeny and species relationships

Based on phenetic similarity, Rodd (1998) proposed four informal groups

for the Australian species of Livistona:

e Mariae Group: L. /anuginosa, L. mariae, L. nasmophila

and L. rigida: palms of alluvial flood-plains in semi-arid

environments.

e East Coast Group: L. australis, L. benthamii, L. decora, L.

drudei, L. fulva and L. nitida: mesomorphic palms of moist

habitats.

e Arafura Group: L. eastonii, L. humilis and L. muelleri: smallish

palms of monsoonal climates.

e Northwestern Group: L. alfredii, L. inermis, L. lorophyllaand L.

victoriae: small to moderate palms in semi-arid environments.

Dowe (2001) performed cladistic analyses of all Livistona species

based on morphological characters. In summary, three moderately supported

lineages within the genus were identified:

e = L. exigua/L. saribus Group

e =Asian/Malesian Group (with two sublineages)

o L. chinensis subgroup (L. boninensis, L. chinensis, L.

endauensis, L. halongensis, L. jenkinsiana, L. tahanensis)

o L. rotundifolia subgroup (L. chocolatina, L. merrillii,

L. papuana, L. rotundifolia, L. surru, L. tothur, L.

woodfordii)

e = African/Australian Group (with two sublineages)

o L. mariae subgroup (L. alfredii, L. australis, L. carinensis,

L. decora, L. lanuginosa, L. mariae, L. nasmophila, L.

nitida, L. rigida, L. victoriae)

L. humilis subgroup (L. benthamii, L. concinna, L. drudei,

L. eastonii, L. fulva, L. inermis, L. humilis, L. lorophylla, L.

muelleri)

More recently, Isagi (pers. comm. and unpublished data, 2008)

established a molecular phylogeny of the Australian species and a small

number of extra-Australian species using AFLPs, cpDNA sequences and

nuclear DNA sequences. Preliminary results indicated that the Australian

species form a well-supported monophyletic group in all analyses, and

the extra-Australian species a less well-supported group. Some of the

A Taxonomic Account of Livistona R.Br. (Arecaceae) 199

arrangements based on phenetic and morphological analyses as noted

above were in most part confirmed by molecular studies, such as Rodd’s

(1998) Mariae and Northwestern groups, and Dowe’s (2001) L. mariae and

L. humilis groups. In addition, the closest relationship of L. carinensis was

variously with extra-Australian species rather than the L. mariae group as

predicted by Dowe (2001).

Taxonomy

Livistona R.Br., Prodr. (1810) 267; Martius, Hist. Nat. Palm. (1838) 238:

Miquel, Fl. Ned. Ind. 3 (1855) 57; Hooker, Gen. PI. 3 (1883) 880; Drude,

Natur. pflan. 2 (3) (1887) 35; Beccari, Ann. Roy. Bot. Gard. (Calcutta) 13

(1931) 43; Uhl & Dransfield, Gen. Palmarum (1987) 190; Rodd, Telopea 8

(1998) 51. — Type species: L. humilis R.Br. (fide Moore, 1963a).

-Saribus Blume, Rumphia 2 (1838) 48. — Type: S. rotundifolius (Lam.)

Blume.

[Saribus Rumph., Herb. Amboin. | (1741) 42].

-Corypha L., Juss., Gen. Pl. (1789) 39, pro parte.

-Livistona sect. Eulivistona Drude, Bot. Jahr. 39 (1893) 12. — Type: L.

leichhardtii F.Muell. (= L. humilis R.Br.).

-Livistona sect. Gregorya Drude, Bot. Jahr. 39 (1893) 12.—Type: L. alfredii

F.Muell.

-Wissmannia Burret, Bot. Jahr. 73 (1943) 184.—Type: Wissmanniacarinensis

(Chiov.) Burret (= L. carinensis (Chiov.) J.Dransf. & N.W.Uhl).

Solitary, short to tall, erect, pleonanthic, hermaphroditic or dioecious or

functionally dioecious palms. Trunk slender to robust, cylindrical, broader

at the base, ringed with leaf scars; internodes narrow to broad. Leaves

palmate or costapalmate, when senescent leaves sometimes forming a

persistent skirt; leaf-bases forming a fibrous network, sometimes forming an

appendage either side of the petiole base; petioles long, adaxially concave,

flat or ridged, abaxially rounded, margins smooth or spiny; adaxial hastula

present; lamina regularly segmented, or irregularly segmented with segments

grouped and divisions between each group deeper into the lamina than

the divisions between individual segments; segments induplicate, narrow,

lanceolate, single-fold, tapered to the apex, apices shallowly or deeply cleft,

rigid or pendulous; segment with a prominent midrib, few to many parallel

veins each side of midrib; transverse veins equal thickness or thinner than

parallel veins. Inflorescences interfoliar, usually held within the limits of

the crown, infrequently extending beyond the crown, either a single axis not

200 Gard. Bull. Singapore 60 (2) 2009

branched at the base but with partial inflorescences positioned along its length,

or basally trifurcate or occasionally bifurcate with + similar collateral axes,

which have a common prophyll, and with partial inflorescences positioned

along the length of each axis; partial inflorescences | (L. humilis only) or

numerous per axis; inflorescences branched 2 to 5 orders when the primary

axis of the inflorescence is regarded as ‘0’; prophyll 2-keeled; peduncular

bracts lacking or if present then loosely or tightly tubular; rachis bracts loosely

or tightly tubular, subtending each partial inflorescence, secondary rachis

bracts that subtend subsequent branches of the partial inflorescence may

be lacking or present; rachillae straight or flexuose, glabrous or tomentose,

papillose, pubescent, hairy or pruinose (L. carinensis only). Flowers bisexual,

or unisexual with little dimorphism, solitary or in sympodial pairs or clusters,

spirally arranged, variously coloured; sepals triangular, fused at the base with

3 apical lobes, glabrous, or hairy (L. carinensis only); petals 3, free, longer

than the sepals, triangular, valvate, glabrous, or hairy (L. carinensis only);

stamens 6, epipetalous, shorter than the petals; anthers didymous, globose to

- ovoid, medi-fixed, versatile, latrorse; pollen monosulcate, ellipsoidal; tectum

finely perforate to reticulate; carpels wedged-shaped, basally separated;

styles united distally, slender, shorter than the stamens; stigma 3-lobed.

Fruit globose, subglobose, ellipsoid, depressed-globose, obovoid, reniform,

pyriform, or obpyriform, variously coloured, 1-seeded; stigmatic remains

apical to subapical; epicarp thin or thick, smooth or with scattered pores or

mildly rugose; suture line extending longitudinally from stigmatic remains

toward the base or sometimes not readily visible; mesocarp dry or fleshy, with

or without embedded fibres. Seed globose, subglobose, ovoid, ellipsoid or

reniform; endosperm homogeneous, intruded by the testa from one side with

spongy/crystalline tissue to displace a moderate or large proportion of the

endosperm; hilum broad, circular to elongate; raphe branches few or lacking;

embryo small, supra- to sublateral. Eophyll simple, with longitudinal ribs.

= 18 (Sharma and Sarkar, 1956).

A genus of 36 accepted species distributed in Dyibouti, Somalia,

Yemen, NE India, Assam, Sikkim, Bangladesh, Myanmar, Thailand,

Malaysia, Taiwan, China, Ryukyu Islands, southern Japan, Bonin Islands,

Vietnam, Kampuchea, Laos, the Philippines, Brunei Darussalam, Indonesia,

Papua New Guinea, Solomon Islands, and Australia; grows in rainforest,

montane forest, swamp forest, monsoon forest, coastal forest, semi-closed

to open woodlands and savanna, semi-arid woodlands, and riparian and

riverine forest, on various soils, including limestone and peat, in permanently

or seasonally wet situations, or in semi-arid to arid situations where usually

associated with permanent ground water or at the base of cliffs where

subsurface moisture is seasonally available.

A Taxonomic Account of Livistona R.Br. (Arecaceae) 201

Key to Livistona species

la. Lamina with the segments grouped, with divisions between each group

of segments deeper into the lamina than the divisions between individual

Sait RO BE ee Oe Saree AN oA SPY los ton dow ddneeseeenshei ces scecadnesveweclex 2

lb. Lamina with leaf segments not grouped, with divisions between

adjacent segments to more or less the same depth into the lamina ........... 3

2a. Large canopy palms to 40 m tall; lamina costapalmate; mid-leaf

segments to 200 cm long; partial inflorescences 4-9; fruit blue to purple at

LEVEICIIN' 2 scpeet ae ANE lee Mica Acie eae me 1. L. saribus

2b. Small understorey palms to 5 m tall; lamina palmate; mid-leaf

segments to 50 cm long; partial inflorescences 3-4; fruit purplish-green at

nN a oo ca yench nc ntavennewmeaiinsvowsestesssccesess Ds Le CXIQHA

3a. Inflorescences trifurcate, or very infrequently bifurcate, branched at

the base, with the 3 (or 2) collateral axes more or less similar and all sharing

a common prophyll but with each axis bearing its own peduncular bract(s)

PESSIL) cepscen te Stee ser oe en ee 4

3b. Inflorescences not branched at the base and composed of a single

Se a I Oe a Ses SATA eke 22S S oa cae db dace thivand oentesdoevese 11

4a. Lamina with 25 or less segments; depth of apical cleft 1-4% of the

ees OREM SCO INCI: «5a, ye passiod. cee iesecneaszcstcheaweecacedss..ccs D. L. brevifolia

4b. Lamina with 45 or more segments; depth of apical cleft 4-40% of the

SEU COUTE SES TSIT pee op eer nae ey 5

See BPI ADE o YS TERETE NOVERD 2882283002 5n0 choc so ceoe< edd dcécdadeosesveoeedssseceecssebecieveesssss 6

oo. PRIDE SSS CT) 0) 0g ra Ce eet ee ee iT

6a. Lamina with abaxial surface silvery glaucous; segment apices rigid;

rachillae 6-12 cm long, glabrous; flowers solitary; fruit 35-43 mm

ODE seceuscodheccesceaacec costo ene oe 9. L. tothur

6b. Lamina with abaxial surface green; segment apices pendulous, hanging

+ vertically; rachillae 14-24 cm long, pubescent; flowers in clusters of 2-4;

EE STU ESS: aa ra TY 00oe 10. L. surru

7a. Inflorescences branched to 4 orders ........................ 3. L. rotundifolia

Seralnnonescenees Branched (0) 3 OLGers. ....52.2.5.cs22.ccsdeeuc.nseboiscseethendecdecessdeds 8

Pass eet StATIOL (NG LARS YC INOW 2 bose 2h. cc ade ceceanncendetancnncndesccescsedesencdccsaveads 9

202 Gard. Bull. Singapore 60 (2) 2009

8b, Sepals and petals ted) i222... ee ee 10

9a. Petiole usually lacking spines, or if present then confined to the very

proximal part; depth of apical cleft 30-40% of the length of the segment;

flowers.3:0-4:5 nomi long; init dark rede eeee eee ee 6. L. merrillii

9b. Petiole armed throughout, or with spines infrequently confined

to proximal portion or very infrequently lacking; depth of apical cleft

5-11% of the length of the segment; flowers to 1.2 mm long; fruit

Orangested vaisidinn ioc eee eee 4. L. papuana

10a. Segment apices semi-pendulous, hanging ca. 45° or more to the

vertical; lamina deeply undulate, angles of undulations 90° or less; rachillae

4-6 cm long, | mm thick, basally with brown-purple tomentum, distally

@labrOous \ccsscnccseedssceteecee eee ee ee 7. L. woodfordii

10b. Segment apices rigid; lamina moderately undulate, angles of

undulations 91° or more, usually much greater; rachillae 8-12 cm long, 2-3

~ mm thick, basally with chocolate brown tomentum, distally with cream-

efeen tomenmbumy s.cesenssh ee eee 8. L. chocolatina

lla. Inflorescences held vertically above the crown, with at least 30% of

their length extending beyond the limit of the crown ...............:::ccceeees 12

11b. Inflorescences held obliquely, horizontally, or below the horizontal,

confined to within the crown, or if extending beyond the crown then by no

more than 10% of their lempthy ris. cve eee ee is

12a. Fruit globose, glossy dark green at maturity; plants hermaphroditic;

inflorescences with 6-7 partial inflorescences .............. 11. L. halongensis

12b. Fruit ellipsoidal or pyriform to obovoid, shiny purple-black at

maturity; plants dioecious; inflorescences on male plants with 4-7

partial inflorescences and the female plants with a single terminal partial

INfLOTESCEMCE 2. saline whd naedeaeeeeR eeee eee ee 19. L. humilis

13a. Inflorescences extending beyond the limit of the crown by no more

than 10% of their length; rachillae, sepals and petals with scattered long

WAITS wccnaccusscdacsscqeebieelemesedee Stee cae eeeeneen aeeene ee eee 18. L. carinensis

13b. Inflorescences not extending beyond the limit of the crown; rachillae,

sepals and petals lacking lomgshairs) 22.2) o-cseeeee cece eee ee 14

4a. Inflorescences lacking peduncular bractsieeee se 15

14b. Inflorescences with | or more peduncular bracts ..................eee 27

- - >

A Taxonomic Account of Livistona R.Br. (Arecaceae) 203

Weer ec eAT HES WIELD BIER APICES 2 cos oon sccacak cde e cee seteesennseveocencndeaustesveceess 16

RSE tes WILE PENGUIOUS APICES 2.22... 22. sec esececcceceeneesececneetcoes 22

l6a. Flowers with maroon sepals, yellow petals, and pink to maroon

PDE Sahoo car 20. L. muelleri

16b. Flowers with sepals, petals and carpels of a single similar colour,

en IDMNNE ADEM OME EN YN 92 22 20252252228 Seek a eas oaks dns sbacdacncecesdsccenceecs 17

02. PESTS SSS Es GSTs 20 a Go oees enero eee ee 18

RIPEN EIEHIAE CIDE ORCCID AL SETALENEILY 22203 < 2 conc cnna2anoscacnce nace sencestiietacdsesscennenessscs 19

18a. Lamina with 60-70 segments; abaxial surface green; stems with

persistent petiole stubs only at the base; inflorescences to 150 cm long;

RECEIVE 10 PYEUOCI 2ooo2. ccs ceeise dese. e cid eceecdseassteesseee 14. L. endauensis

18b. Lamina with 40-50 segments; abaxial surface = os: green; stems with

persistent petiole stubs for most of its length; inflorescences to 91 cm long;

fruit globose to depressed-globose ...............::cceeeeeeeeeees 15. L. tahanensis

19a. Mid-leaf segments no more than 70 cm long: lamina with divisions

between segments more than 80% of the length of the lamina; apical cleft

in the segment more than 70 % of the length of the se cement; fruit glossy

SLE EA oa eT ee ee 2. L. inermis

19b. Mid-leaf segments not less than 110 cm long; lamina with divisions

between segments less than 75% of the length of the lamina; apical cleft

in the segment less than 48% of the length of the segment; fruit purple-

black or leaden blue to dark bluish-purple or greenish-blue to blue at

ESO cee 2 ee eo 20

20a. Fruit Hohe lamina with less than 60 segments; depth

of apical cleft in the segment more than 40% of the length of the

SS TUT SLLD ae es ae ce cee 33. L. nasmophila

20b. Fruit leaden blue to dark bluish-purple or greenish-blue to blue at

maturity; lamina with more than 70 segments; depth of apical cleft in the

segment less than 10% the length of the segment ..............eeeeeeeeeee 21

21a. Lamina adaxially shiny green, abaxially subglaucescent grey or bluish;

inflorescence branched to 3 orders; rachillae puberulous; fruit globose to

reniform, leaden blue to dark bluish-purple .................. 16. L. jenkinsiana

21b. Lamina adaxially green, abaxially glaucous grey; inflorescence

branched to 4 orders; rachillae glabrous; fruit obovate, obpyriform to ovoid,

greenish-blue to light-blue at maturity «0.0.0.0... eee 17. L. speciosa

204 Gard. Bull. Singapore 60 (2) 2009

22a. Petiole stubs long due to the abscission of leaves occurring in about

the middle of the petiole; stubs persistent on the trunk for most of its length,

unless decayed by age or fire S24... ccccee etree eran ee CITI ATE

22b. Petioles stubs lacking the persistent remnant portion of petiole; stubs

persistent on the trunk only im the basal portiontean---ce eee ZS

23a. Rachis bracts glabrous, tightly tubular, not disintegrating into open

fibres) butremaining intactwitlh age ress ene 26. L. concinna

23b. Rachis bracts tomentose or scaly, loosely tubular, disintegrating into

open fibres with age. :.s.20-cs..sSeeemaceses eae eee ee 24

24a. Fruit blue-green to bright green, glossy, epicarp ceramic-like at

TOT: TAT 9 | been RR RE Rees etre tse ci oceracsoccingouoseqonabode 12. L. chinensis

24b. Fruit reddish-brown to black, variously dull, glaucous, glossy or shiny

but not ceramic=like*at matumity 2 cere essere eet ere ZS

25a. Lamina with divisions between segments more than 82% of the length

of the seemient fruntyshimnys blacks eee ence eee 28. L. decora

25b. Lamina with divisions between segments less than 70% of the length

Of the segment 225... ibsccck secs ee eee 26

26a. Inflorescences branched to 5 orders; fruit dull, reddish-brown to dull

black, or occasionally glaucous) <...1.:cciccceee-ccceseeeeee cere tee EES

26b. Inflorescences branched to 4. orders; fruit glossy jet

6) E16) inna ne re Fee BARBER OR i cca cgonsciigyi0os 30. L. nitida

27a. Lamina with coppery brown floccose tomentum on the abaxial

SUTLACE a ccccvissassdsacenseeduegoeate cee cece eeeaae eee ane ECO R nee en eee 31. L. fulva

27b. Lamina not with coppery brown floccose tomentum on abaxial surface,

but variously green OF Grey ..steecesccecceese epee eee tenet ne eee nee ee 28

28a. Inflorescences branched to 3 Orders ere eeetes ee eee eee 29

28b.' Inflorescences branched:tor4ordess treet ree 32

29a. Fruit bught greeniat maturity eee 13. L. boninensis

29b. Fruit not green at maturity, but variously brown to

9) Ec]. qin PnP pe eee ce re orcs anda dgccncJucasevdocnauigdocncoasc0s 30

30a. Petiole purple in basal portion) -.25.22ceeeree ere

30b. Petiole green in basal portion {i eeece ee ee eee 31

——————

A Taxonomic Account of Livistona R.Br. (Arecaceae) 205

31a. Fruit 25 mm or greater in length, globose; lamina segments 50-66;

divisions between segments less than 70% of the length of the segment;

Facuillae wiite pruinose, PapillOSe <....2...c.22-2...-00cc.s000cceseseees 32. L. alfredii

31b. Fruit 14 mm or less in length, obovoid-pyriform; lamina segments

34-50; divisions between segments more than 85% of the length of the

Beomenterachillae SIADLOUS .2.212¢.-c--c+-22c<2.0e-0+2-er-0002---00e2e- 23. L. lorophylla

oe aeknuit diameter 25 mmy OF PTeater ..2....:..2..000000..-.000-- 36. L. lanuginosa

PAO MHPHGOLATMCLEE NS MINT OLUCSS: &.2.5..b.2..25-ceccedascesceeaeste.cebsctevewsceassnteceee 33

33a. Moderate palms to 15 m tall, dbh less than 15 cm; less than 20 leaves

SMMC ICLOM NM MUL ODOVOIG 22522000: testcasesteevensossnocncsvsstecsseeees 21.0. eastonii

33b. Tall palms more than 15 m tall; dbh more than 15 cm; more than 25

leaves im the crown; fruit globose to ellipsoidal ...........2.2......scs::ccseeceeees 34

34a. Lamina not more than 100 cm long; divisions between segments more

than 55% of the length of the segment; rachillae not more than 3 cm long;

MG@RMEES SOMGAIYV AOL MM PAINS sc on¢.c+-c02 --desneescteeocr-avcceseosencoene-ee, 24. L. victoriae

34b. Lamina more than 100 cm long; divisions between segments less than

55% of the length of the segment; rachillae more than 3 cm long; flowers in

eee Kem SUOMI ORC TOWELS 4:5.82s02.00dscrdacteensacaseavecdeososedassasccetssassecesseseece SD

35a. Lamina to 220 cm long, moderately undulate with undulations > 91°;

segment apices pendulous hanging + parallel to the vertical; peduncular

bracts with dense white-grey scales; rachis bracts with dense white scales;

Raemmlac PiABle HO MEKUOSE *:2isecrc42.24...0dscoevsdssses sacaceseecsesncess 34. L. mariae

35b. Lamina to 170 cm long, strongly undulate with undulations < 90°;

segment apices rigid or semi-pendulous; peduncular bracts with moderate

to dense white-grey scales; rachis bracts with sparse white scales; rachillae

Descriptions of species *

[* In the following species descriptions, species are arranged informally in

morphological groupings. This arrangement does not suggest phylogenetic

relationships, but rather that similar species are placed close to each

other. |

|. Livistona saribus (Lour.) Merr. ex. A.Chev., Bull. Econ. Indo-Chine.

ser. 2, 22 (1919) 501. -Corypha saribus Lour., Fl. Cochinch. 1 (1790) 212;

Gmelin, Syst. Nat. 2 (1791) 569. — Type: Pierre 4837, Cay Ki, Vietnam,

206 Gard. Bull. Singapore 60 (2) 2009

July 1866 (neotype, here designated L; isoneotypes, A, BO, K, P, UC).

Plate 1A-E.

-Livistona spectabilis Griff., Calcutta J. Nat. Hist. 5 (1845) 336; Palms

Brit. E. Ind. (1850) 130, pl. 226C; Miquel, Fl. van Ned. Ind. 3 (1855)

58 (as L. rotundifolia); Wendland, in Kerchove, Palmiers (1878) 250,

prop. syn., L. rotundifolia; Beccari, Webbia 5 (1921) 21, prop. syn., L.

chinensis. — Type: Malaysia. Malacca, 1843, Lewis s.n. (holotype, BR).

-Livistona cochinchinensis (Blume) Mart., Hist. Nat. Palm. 3 (1849) 242;

Beccar1 & Hooker, FI. Brit. India 6 (1892) 434. -Saribus cochinchinensis

Blume, Rumphia 2 (1838) 49. Type not designated.

-Livistona hoogendorpii Hort. ex Teysm. & Binn. ex Miq., Palm. Archip.

Ind. (1868) 14 (as L. hoogendorpi); Teysmann & Binnendijk, Cat. ’slands

Plant. Buitenzorg (1866) 71, nomem; André, Ill. Hort. 21 (1874) 108, 121,

t. 174; Kerchove, Palmiers (1878) t. 33. -Saribus hoogendorpii (Hort. ex

Teysm. & Binn. ex Miq.) Kuntze, Revis. Gen. Pl. 2 (1891) 736. — Type:

Indonesia. Cultivation, Bogor Botanic Gardens, May 1878, Beccari

's.n. (lectotype, here designated, FI, Beccari collection, sheets 11330 &

11330-B).

-Livistona hasseltii (Hassk.) Mig., Palm. Archip. Ind. (1868) 14 (as

L. hasselti); Wendland, in Kerchove, Palmiers (1878) 250; Backer &

Bakhuizen van den Brink, Fl. Java 3 (1968) 174. -Saribus hasseltii Hassk.,

Flora 25 (Beibl. 2) (1842) 16. — Type: Indonesia. Batam Prov., Tjikoi and

Tjilakatan, 1820-23, van Hasselt s.n. [type not located at either L or U (N.

Sol, pers. comm. )].

-Livistona inaequisecta Becc., Philipp. J. Sci. C. Botany 4 (1909) 616. —

Type: Philippines. Luzon. Laguna Proy., Santa Maria Mavitae, Feb 1908,

Curran 10079 (holotype, PNH).

-Livistona tonkinensis Magalon, Contr. étude Palmiers Indoch. (1930) 54;

Feddes Repert. Spec. Nov. Regni Veg. 28 (1930) 239; Feddes Repert.

Spec. Nov. Regni Veg. 29 (1931) 23; Gagnepain, Fl. Indo-Chine 6 (1937)

983. — Type: Vietnam. Rivieré Noire and the middle region of the Tonkin

R., 25 Jan 1930, Magalon s.n. (lectotype, here designated, P).

Hermaphroditic palm. Trunk to 40 m tall, 15-65 cm dbh, leaf scars raised,

internodes broad, petiole stubs persistent in the basal 2 m or so. Leaves 25-

30 ina+ globose crown; petiole arching, 100-200 cm long, to 12 cm wide in

proximal portion, to 15 mm wide in distal portion, adaxially flat to slightly

ridged, glabrous, green to green-purple to green-red, glossy, frequently with

a reddish-purple longitudinal stripe; margins with large, single, retrorsely

recurved, green to brown spines 10-60 mm long, largest and closer in the

proximal portion; leaf-base fibres moderately fibrous, coarse, persistent;

A Taxonomic Account of Livistona R.Br. (Arecaceae) 207

lamina costapalmate, irregularly segmented, with segments grouped and

with divisions between each group of segments deeper into the lamina

than the divisions between individual segments, subcircular, 80-200 cm

long, 150-170 cm wide, adaxially green, abaxially a similar green; lamina

divided into groups of 2-12 segments separated from adjacent groups by a

deep split that almost reaches the hastula; lamina divided for 37-78% of its

length, with 80-90 segments, depth of apical cleft 19-50% of the segment

length, apical lobes pendulous; parallel veins 6-7 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, 60-230 cm long, not extending beyond the limit of the crown,

curving, branched to 4 orders; partial inflorescences 4-9, 45-60 cm long:

prophyll not seen; peduncular bract(s) lacking; rachis bracts loosely

sheathing, glabrous; rachillae 15-45 cm long, drooping, yellow, glabrous.

Flowers in clusters of 3-5, 1.5-1.75 mm long, yellow; sepals suborbicular:

petals broadly triangular, obtuse. Fruit globose, or ellipsoid to reniform,

11-25 mm long, 10-18 mm diam., often bilobed, glossy blue to purple;

epicarp thin with scattered lenticellular pores; suture line extends for full

length of the fruit, marked with lip-like structures; mesocarp fleshy, 1.0-

1.5 mm thick; endocarp crustaceous; pedicel 1-3 mm long, ca 3 mm wide.

Seed globose to ellipsoid, 9-24 mm long, 9-10 mm diam., apically pointed;

endosperm intruded for ca half its width; embryo lateral. Eophyll 6-8-

ribbed.

Specimens examined: VIETNAM: West Tonkin. Bonn 6184 (P); Cay

Ki, Pierre 4837 (A, BO, K, L (neotype), P, UC); Cay Ki, Poilane 1222

(BO, P); Tonkin. Balansa 4366 (K); Tonkin. Rivieré Noire, 25 Jan 1930,

Magalon s.n. (P); Location not specified, 3 Oct. 1936, Poilane s.n. (K);

Location not specified, Jun 1918, Lecomte s.n. (P). KAMPUCHEA: Foret

de Phnom Penh, 16 Feb 1934, Bejaud s.n. (K, P). THAILAND: Chiang

Mai Prov. Road to Doi Chang Dao, 19°21°N, 98°47°E, 900-1100 m alt.,

Barfod 45208 with Pooma & Burholt (AAU, K, NY): Phitsanulok, 7 km E

of Tung Salaeng Luang, 600 m alt., Larsen 792 with Smitinand & Warncke

(AAU, BKF); Loei, 10 km W of Loei, Dowe 56] & 562 (JCT); Kabin

Buri, 45 km N of Kabin Buri, Ban Numklae village, Dowe 564 (JCT);

Prachinburi, Krabin, Bupram, 500 m alt., Kerr 9836 (AAU, BK, K, NY); 1

km NW of Wang Chan, 30 km from Klaeng, Dowe 570 (JCT); Chumphon,

Bang Son, 100 m alt., Kerr 1/342 (AAU, BM, K); Chumphon, 20 km N of

Thai Sae, Dowe 567 & 568 (JCT); Ranong, Bang Mon Kapur, Klong Na

Kha Wildlife Sanctuary, Smith 46 with Sumawong (K); Ranong, Klaeng,

Nam Groi (outside N. P.), farmland on boundary of park, Smith 126 & 132

with Sumawong (AAU, K); Ranong. Pluak Daeng, Sukhaphiban Rd No.

208 Gard. Bull. Singapore 60 (2) 2009

9, 100 yards, Smith 144 with Sumawong (K); Trang, Muang, Tran Airport,

Smith 78 with Sumawong (K); Songkhla, Kha Kok Hng, Hatyai, 07°00’N,

100°20’E, 150-250 m alt., Larsen 41280 (AAU); Nakhon Si Thammarat,

Ta Sumet, less than 50 m alt., Kerr 14313 (BK, BM, K, NY). MALAYSIA:

Malacca, 1843, Lewis s.n. (BR); Sabah. Abai, Kinabatangan, Keith 3429

(K); Perak. Gunong Pondok, ca 700 ft alt., 7 Jun 1930, Henderson s.n.

(BO, SING); Selangor. Tanjong Karang, Sungai Ting1, low alt., Nur 34149

(A, BM, L, K, SING, US); Selangor. 5 Jul 1902, Burn-Murdoch s.n. (K);

Selangor. Jugua, Burn-Murdoch 11352 (BM). INDONESIA: Kalimantan.

Djaro Dam, Muara Uja, 100 m alt., Dransfield & Sauerudin 2141 & 2145

(BO, K); South Sumatra. Rasau, Waikambas Reserve, 10 m alt., Dransfield

1254 (BO); Java. Track to Tjibunar, Udjang Kulai, Dransfield 1459 (BO);

Java. Bantam, Tjimura, Koorders 6123/3 (BO); Java. Koorders 35163 (BO).

PHILIPPINES: Luzon. Nueva Vizcaya Prov., ca 350 m alt., Fernando 7286

(K); Luzon. Nueva Vizcaya Prov., Diadi, Magat, 350 m alt., Fernando

EF671 (LBC); Luzon. Tarlac Prov., Vidal 1951 (K); Luzon. Zambales

-Prov., Pannubuan, hills between San Marcelino and Mt Pinatubo, Bartlett

14236 (K); Luzon. Zambales Prov., Dinahyihan, Loher 1390 (K); Luzon.

Laguna Prov., Santa Maria Mavitae, Curran 10079 (PNH); Luzon. Laguna

Prov., Tical, Cavinti, Loher 7508 (K).

Specimens from cultivated material: BRAZIL: Gardens, Piracicaba, anon.

s.n.(K); CHINA: Yunnan Institute of Tropical Botany, Chen San-yvang 18831

(K); Dominican Republic: Location not known, anon. 13 (K); INDONESIA:

Bogor Botanic Gardens, May 1878, Beccari s.n. (FI); Mauritius. Botanic

Gardens, 1881, Horne s.n. (K); PHILIPPINES: Luzon, Laguna Prov., Los

Banos Forestry Campus, Fernando 7359 (LBC); SINGAPORE: Singapore

Botanic Gardens, Lawn K, 22 Jun 1929, Nur s.n. (K); Singapore Botanic

Gardens. Furtado 34149 (US); SRI LANKA: Royal Botanic Gardens,

Peradeniya, S-16, Rutherford 54 with Bandara (K); THAILAND: Trang,

Khao Chong Gardens, Whitmore 3153 (K); Trang, Khao Chong, ca 15 km

E of Trang, Peninsula Botanic Garden, Barfod 41588 with Ueachirakan

(AAU, PSU); Trang, Khao Chong, Peninsula Botanic Garden, Barfod 737

with Pongsattayapipat (AAU); VIETNAM: Saigon Botanic Garden, 3 Oct

1936, Poilane s.n. (P, PNH, UC).

Local and vernacular names: Serdang, Sar (Trengganu), Tarao (Cagayan

Prov., Philippines).

Habitat and ecology: In rainforest or swampforest as scattered individuals

or in small to very large colonies, and occasionally in watercourses and

A Taxonomic Account of Livistona R.Br. (Arecaceae) 209

adjacent slopes, at 0-600 m alt. Occurs in peatforest and mangroves in

central Sumatra (Dransfield, 1974; Laumonier, 1997). Flowers Mar.-July:

fruits June-Sept.

Conservation status: Vulnerable.

Distribution: Laos, Vietnam, Kampuchea, Thailand, Malaysia, Indonesia

and the Philippines. Widespread, but rare in Laos, Vietnam, Kampuchea,

Thailand and Malaysia because of forest clearance. In Indonesia confined

to Batam, Sumatra and Kalimantan, and in the Philippines on Luzon Is.,

in Cagayan, Nueva Vizcaya, Tarlac, Zambales and Laguna Provinces

(Fig. 6).

130°E 140°E 150°E 160°E

— i i

= . Legend Lo”

° °

i c

+4 L. bre\ a

Ez &. CAIGUG

Fs el =

=) A L. sanb ro

N N

10°N

Oo i ~

eae :

—~»s i ~ |

=e - y

~ = : iv -

75) SS BESD et = = 2)

Oo} 0 500 -1.000 1.500 2 ea e —S yo

ee ek “£0 J L = e

sae f 4 ;

100°E 110°E 120°E 130°E 140°E 150°E 160°E

Figure 6. Distribution of Livistona saribus (A), L. exigua (m), L. rotundifolia (@), L.

papuana (©), and L. brevifolia (a).

Notes: Livistona saribus is a variable species with some distinct regional

forms, mainly expressed in fruit size, but otherwise unable to be reliably

taxonomically differentiated on that basis. Dransfield (1999) noted the

over-description of palm taxa based on political boundaries, with particular

210 Gard. Bull. Singapore 60 (2) 2009

reference to L. saribus and its former synonyms, with L. hasseltii and

L. hoogendorpii originally named for Java, and L. cochinchinensis for

Peninsula Malaysia, Indochina and the Philippines. The taxonomy of L.

saribus is the most complex for all the species of Livistona. There 1s also a

paucity of type specimens, with only four of the eight names applied to the

species having been typified.

The mononomial Saribus was attributed by Rumphius (1741) to

the vernacular name used by the Macassans of eastern Indonesia. Livistona

saribus was first named as Corypha saribus (Loureiro, 1790) for plants

collected in Vietnam, based on the specimen Loureiro s.n. but not able to

be located and presumed lost (M. Pignal, pers. comm.). Because of the loss

of the holotype, the neotype Pierre 4837 has been chosen.

Livistona_ cochinchinensis was first named as _ Saribus

cochinchinensis by Blume (1838), who based it in part on Loureiro’s

Corvpha saribus and Brown’s C. australis, and suggested that S.

cochinchinensis was the appropriate name for the species occurring in

-Cochinchina (Vietnam). It was transferred to Livistona by Martius (1838)

who similarly related it to Loureiro’s C. saribus but also, in part, to

Griffith’s L. jenkinsiana. Miquel (1855) suggested that L. cochinchinensis

was a synonym of L. chinensis, but did not elaborate. Beccari (1886)

clearly recognised the conspecifity of L. cochinchinensis and L. saribus but

refrained from making the appropriate combination based on the earliest

name, stating “// nome di L. cochinchinensis, é€ per questo da preferirsi,

come piu antico.” This scheme was continued by Beccari (1921, 1931),

despite the formalisation of Livistona saribus (Chevalier, 1919; Merrill,

1925). However, most taxonomists have recently listed it as a synonym

(Burret, 1936, 1941; Moore, 1963b; Pei ef a/., 1991).

Livistona spectabilis was described by Griffith (1845) for plants

collected from Malacca, Malaysia, possibly with the name referring to the

remarkable height attained by the palm. The specimen cited by Griffith, “Mr.

Lewes”, correctly W.T.Lewis, is extant in BR. Wendland (1878) included

the name as a synonym under L. rotundifolia; subsequently, Beccari (1886)

included it as a synonym of L. cochinchinensis (= L. saribus); but later as

a synonym of L. chinensis (Beccari, 1921). The protologue, the illustration

included in Griffith (1850) and the specimens in BR confirm its correct

synonymy under L. saribus.

The name L. hoogendorpii was first used by Teysmann & Binnendijk

(1866) in a list of palms cultivated in Bogor Botanic Gardens, with no

explanation for the choice of specific epithet. Miquel (1868) provided the

first description thus formalising the name, and André (1874) described and

illustrated juveniles in cultivation in Europe. Kerchove (1878) provided

A Taxonomic Account of Livistona R.Br. (Arecaceae jp

AT 4 t of Livistona R.Br. (A ) Pil

a brief description and an illustration of a juvenile plant. Beccari visited

Bogor Botanic Gardens in May 1878 and collected specimens from the

plants upon which Teysmann & Binnendijk established the name, and

those specimens, sheets 11330 and 11330-B in FI are here chosen as the

lectotype. Although the origin of the species remained unknown, Blatter

(1926) noted the habitat as the “/ndian Archipelago”. Moore (1963b) was

the first to designate L. hoogendorpii as a synonym of L. saribus.

Livistona hasseltii was described by Hasskarl (1842) as Saribus

hasseltii from a collection by van Hasselt s.n., from Batam Province in

Indonesia, and named for the collector, the Dutch botanist, J.C. van Hasselt

(1797-1823). Miquel (1868) provided the transfer to Livistona. The name

has been applied to plants growing in Bogor Botanic Gardens, and cited in

Backer & Bakhuizen van der Brink’s (1968) Flora of Java, although with

the proviso that it “...may be only a form of continental L. saribus...”.

Livistona inaequisecta was described by Beccari (1909) from the

collection Curran 10079, from Luzon in the Philippines, and named for the

“unequally parted’ leaves. Beccari (1919a) placed it as a synonym of L.

cochinchinensis. Livistona tonkinensis was applied to a population in the

Tonkin region of Vietnam by Magalon (1930) based on his own collection

now in P, and is synonymised under L. saribus.

Livistona saribus is a canopy palm to 40 m tall; leaves are large

with grouped segments and deep divisions between the groups; segment

apices are pendulous; there are long prominent spines on the petiole; the

inflorescence is unbranched, not extending beyond the limit of the crown,

and with up to 9 partial inflorescences; flowers are yellow; and fruit are

globose to ellipsoid to reniform, to 25 mm long, to 18 mm diam., often

bilobed, and glossy blue to purple at maturity.

2. Livistona exigua J.Dransf., Kew Bull. 31 (1977) 760, fig. 2. — Type:

Brunei. Belait District, Ulu Ingei, ca 60 m alt., Aug 1959, Ashton BRUN

5513 (holotype, K; isotypes, BH, BO, L, NY, SAR). Plate 2A-D.

Hermaphroditic palm. Trunk to 5 m tall, 1.5-2.5 cm dbh, leaf scars

raised, irregular in width, internodes narrow, irregular in length, pale

brown, petiole stubs not persistent. Leaves 16-20 in a vertically ovoid

crown; petiole 40-60 cm long, slightly arching, proximally ca 6 mm wide,

tapered to ca 3 mm wide distally, adaxially flat, armed on the margins

in the lower one-third, upper margins sharp, slightly winged; petiole

surface with scattered caducous pseudo-parenchymatous peltate scales

on both surfaces; spines retrorsely recurved, brown, to 4 mm long, bases

swollen, regularly placed at ca 5 mm apart; leaf-base fibres prominent,

PWV? Gard. Bull. Singapore 60 (2) 2009

coarse, closely woven, dull dark brown, persistent; appendage to ca 30

cm long; lamina palmate, irregularly segmented, with segments grouped

and with divisions between each group of segments deeper into the

lamina than the divisions between individual segments, circular, ca 50

cm wide, adaxially midgreen, abaxially lighter green, adaxially glabrous,

scattered brown scales abaxially; lamina divided for ca 42% of its length,

with 16-30 segments that are single or grouped into 2-6 segments, depth

of apical cleft ca 8% of the segment length; segments to 25 cm long,

widest at mid segment, 1.0-1.5 cm wide, acuminate toward both apex

and base, apical lobes rigid, finely toothed; parallel veins 5-6 each side

of midrib; transverse veins + same thickness as parallel veins; hastula

conspicuous, to 5 mm wide and 2-3 mm high. Inflorescences unbranched

at the base, 15-40 cm long, not extending beyond the limit of the crown,

straight to slightly curving, branched to 2 orders; partial inflorescences

3-4; peduncle 20-25 cm long, elliptical in cross-section; prophyll tubular,

to 18 cm long, dark brown with pale brown scurfy indumentum along

the keels, minutely longitudinally striate, the apices acute; peduncular

bract(s) lacking; rachis bracts loosely tubular; partial inflorescence axes

covered densely with indumentum and papillae; rachillae 6-10 cm long,

rigid, to ca 2.5 mm diameter. Flowers solitary or paired, sessile, purplish,

subtended by a fimbriate bract ca 0.2 mm high, ca | mm diam.; sepals

fused for ca half their length to form a short tube ca 0.3 mm high, lobes

ca 0.2 mm high, rounded, fimbriate, abaxially densely covered with

indumentum; petals broadly triangular, glabrous, ca 0.4 mm high, basally

fused for ca one-quarter their length; stamens ca 2 mm high, basally

fused, swollen; carpels glabrous, style ca 0.2 mm high. Fruit globose, ca

9 mm diam., purplish-green; epicarp smooth; suture line extending the

full length of the fruit; mesocarp ca 1.5 mm thick with numerous tannin

cells; endocarp crustaceous, ca 0.3 mm thick; pedicel 1-2 mm long. Seed

globose. Eophyll not seen.

Specimens examined: BRUNEI DARUSSALAM: Belait District, Ulu Ingei,

ca 60 m alt., Ashton BRUN 5513 (BH, BO, L, K holotype, NY, SAR); Ulu

Ingei, Bukit Batu Patam, 04°5’N, 114°42’E, 200 m alt., ridge top kerangas

forest, transitional to mixed Dipterocarp forest, abundant miniature serdang

forming thickets on ridge tops, Dransfield JD6568 with Wong Khoon Meng,

Dransfield & Boyce (A, K, SING); Ulu Ingei, 150 m alt., Jan 1982, Morgan

s.n. (K, SING).

Habitat and ecology: In forest transitional between lowland Dipterocarp

forest and kerangas (heath forest) on sandstone, at 60-260 m alt.

A Taxonomic Account of Livistona R.Br. (Arecaceae) 213

Conservation status: Near threatened.

Distribution: Brunei. Belait District, Ulu Ingei (Fig. 6).

Notes: Livistona exigua was described by Dransfield (1977) based on the

collection Ashton BRUN5513 collected from the Belait District, Brunei, in

1959, and named with reference to the palm’s small stature. This collection

had remained as unidentified material in Kew, but with examination by

Dransfield was recognised as a new species. It has provided a considerable

ecological extension for the genus, as a rainforest understorey palm in

kerangas heath, which was previously not recorded in Livistona.

Livistona exigua is an understorey palm to 5 m tall; leaves are small,

and have grouped segments and deep divisions between the groups; segment

apices are rigid; the inflorescence is unbranched, not extending beyond the

limit of the crown, and with up to four partial inflorescences; flowers are

purplish; fruit are globose, to 9 mm diam., and purplish green at maturity.

3. Livistona rotundifolia (Lam.) Mart., Hist. Nat. Palm. 3 (1838) 241;

Miquel, Fl. Ned. Ind. 3 (1855) 58; Palm. Archip. Ind. (1868) 12; Beccari,

Ann. Roy. Bot. Gard. (Calcutta) 13 (1931) 74. -Corypha rotundifolia Lam.,

Encycl. 2 (1786) 131; Willdenow, Sp. pl. 2 (1797) 201; Hayne, Term. Bot.

tab. 11, fig. 1 (1807); Sprengel, Syst. Veg. 2 (1825) 138; Schultes, Syst.

Veg. 7 (2) (1830) 1309. -Saribus rotundifolius (Lam.) Blume, Rumphia 2

(1838) 49, pl. 95, 96. — Type: Illustration in Rumphius, Herb. Amboin. |

(1741) t. 8 (lectotype, fide Merrill, 1917). Plate 3A-D.

-[Saribus Rumph., Herb. Amboin. | (1741) 42].

-Corypha umbraculifera auct. non L., Sp. Pl. (1753) 1187.

-Livistona umbraculifera Hort. ex Vilm., Blumengartn. ed. 3, 1 (1895)

1149, in syn.

-Livistona altissima Zoll., Nat. Tijdschr. Ned. Indié 14 (1857) 150. Saribus

sp. n. ? ‘Sadjeng’, Zollinger, Syst. Verz. (1854) 78; Miquel, Palm. Archip.

Ind. (1868) 14. — Type: Lectotype here chosen. Indonesia. Cultivation.

Bogor Botanic Gardens, undated, Zo/linger 2684 (lectotype, BM).

-Livistona robinsoniana Becc., Philipp. J. Sci. 6C (1911) 230; Philipp. J.

Sci. 14 (1919) 341. — Type: Philippines. Polillo Is., Quezon Prov., Aug.

1909, Robinson 9265 (holotype, FI; isotypes, K, US).

-Livistona rotundifolia var. microcarpa (Becc.) Becc., Philipp. J. Sci.

14 (1919) 341; Leafl. Philipp. Bot. 8 (Aug. 25) (1919) 3024. -Livistona

microcarpa Becc., Philipp. J. Sci. 2 (1907) 231. — Type: Philippines.

Palawan, Carafiugan R., 28 Feb. 1906, Curran 3784 (holotype, FI; isotypes,

BM, NY, P).

214 Gard. Bull. Singapore 60 (2) 2009

-Livistona rotundifolia var. mindorensis (Becc.) Becc., Philipp. J. Sci. 14

(1919) 341. -Livistona mindorensis Becc., Philipp. J. Sci. 4 (1909) 615.

— Type: Philippines. East Mindoro Prov., Mindoro, Bongabong R., May

1906, Merritt 4108 (holotype, FI).

-Livistona rotundifolia var. luzonensis Becc., Leafl. Philipp. Bot. 8 (Aug.

25) (1919) 3025; Philipp. J. Sci. 14 (1919) 340; as Livistona rotundifolia

(Lam.) Mart., Leafl. Philipp. Bot. 2 (1909) 647; Merrill, Fl. Manila (1912)

122. — Type: Philippines. Luzon. Quezon (Tayabas) Prov., Lucban, Mt

Banahao, May 1907, Elmer 9293 (holotype, FI; isotypes, BM, BO, K, L,

NY, US).

Hermaphroditic palm. Trunk to 45 m tall, 15-25 cm dbh, leaf scars

obscure to prominent, light green to white, internodes broad, green to

grey, smooth or infrequently with longitudinal fissures, petiole stubs not

persistent. Leaves 20-50 in a globose crown; petiole slightly arching, 90-

210 cm long, 15 cm wide proximally, ca 2 cm wide distally, adaxially flat

or moderately ridged, margins with retrorsely recurved black spines 1-20

mm long throughout or proximally only, with largest proximally, distally

becoming smaller and more widely spaced, or very infrequently with spines

lacking in mature plants; leaf-base fibres moderately prominent, coarse,

in a criss-cross pattern, brown, persistent, appendage triangular; hastula

very prominent, 2 cm high; lamina costapalmate, circular to subcircular,

regularly segmented, 75-150 cm long, adaxially semi-glossy dark green,

abaxially lighter subglaucescent green; lamina divided for 38-62% of its

length, with 60-90 segments, depth of apical cleft 4-25% of the segment

length, apical lobes usually erect, but pendulous in segments with deeper

clefts; mid-leaf segments ca 5 cm wide where the segments diverge; parallel

veins 6-9 each side of midrib; transverse veins equal thickness or thinner

than parallel veins. Inflorescences trifurcate with + similar collateral axes,

branched to 4 orders, 90-150 cm long, not extending beyond the limit of the

crown, arching; partial inflorescences ca 10, longest to ca 30 cm; prophyll

to 30 cm long, glabrous, straw coloured; peduncular bracts lacking or 1, and

then tightly tubular; rachis bracts tightly tubular, reddish brown, glabrous,

apically truncate, remaining intact with age; rachillae 3-20 cm long, 1-1.5

mm thick, straight, yellowish, glabrous. Flowers solitary or in clusters of

2-4, to 2-3 mm long, yellowish, sessile on small pulvini; sepals broadly

ovate, very obtuse, dorsally carinate; petals less obtuse, yellowish; ovary |

glabrous; style subulate, acute, very short. Fruit globose to subglobose,

11-25 mm diam., at first yellow, ripening though to orange-red to red or to |

dark violet or bluish-black; stigmatic remains inconspicuous; epicarp thin, |

smooth or with scattered lenticellular pores; suture line for full length of

. )

—_

7A)

A Taxonomic Account of Livistona R.Br. (Arecaceae) 72) Ve

fruit; mesocarp ca 1.5 mm thick, slightly fibrous to gritty; endocarp very

thin; pedicel 2-3 mm long. Seed globose, 10-13 mm diam., endosperm

intruded for two-thirds to almost full width of endosperm: hilum broad,

orbicular; embryo lateral, 2-2.4 mm long. Eophyll 5 ribbed.

Specimens examined: INDONESIA: anon. s.n. (BO); Ternate and Sulawesi

(Woka), Reinwardt 60 (L): North Maluku. Halmahera Is., Jailol District,

Kampung Pasir Putih, Zay/or 605 (US); Maluku. Halmahera, Ekor, Bukit

Dowora Ina, de Vogel 3236 (BO, K, L); Maluku. Halmahera, Akelamo

Oba, 00°34’°N, 127°36’E, ca 50 m alt., de Vogel 4436 (K, LAE); North

Sulawesi, Amoerang, undated, collector unknown (BO); North Sulawesi,

Manado, Koorders 18424 (BO); North Sulawesi, Koorders 18425 (BO):

Sulawesi. near Manado, Forman 409 (A, BO, LAE, PNH): Sulawesi.

Bolaang Mongondow, Dumoga Bone NP, Toraut Dam, de Vogel 6894

with Vermeulen (K); Sulawesi. Belaang Mongondow, Pindool, Lelak,

Dransfield & Mogea JD 3831 (BO, K); Ceram, undated, de Vriese &

Teysmann s.n. (L); Papua. Sorong. Raja Ampat Islands, Waigeo Is.,

Waifoi Village, Maturbongs 501 (K). PHILIPPINES: Calayan Is. Cagayan

Prov., 150 m alt., Quisumbing & del Rosario 61-369 (PNH); Luzon.

Cagayan Prov., Linao, Tuguegarno, Rocero 220 (PNH); Luzon. Cagayan

Proy., Claveria, Mt Taggat, Reynoso 11903 (K); Luzon. La Union Prov.,

Contilla, Loher 7070 (K):; Luzon. Tarlac Prov., Vidal 1952 (K):; Luzon.

Zambales Prov., Mt Pinatubo, Villar, 350 m alt., Fox 26 (PNH); Luzon.

Quezon (Tayabas) Prov., Lucban, Mt Banahao, E/mer 9293 (BM, BO, FI,

K, L, NY, US); Polillo Is. Quezon Prov., Fox 8996 (A): Polillo Is. Quezon

Prov., Robinson 9265 (FI, K, US); Polillo Is. Quezon Prov., Barangsay

Sibulan, 4 km N of Polillo town, Dowe 700 with Romero (K, PNH):

Polillo Is. Quezon Prov., Karlagan, Fox 6b (PNH); Polillo Is. Quezon

Prov., Lukutaw, Fox 223 (PNH); Luzon. Manila, Paco, Bartlett 13203

(K); Luzon. Laguna Prov., Mt Makiling, Gates 5700 (CAHP); Luzon.

Laguna Prov., Mt Makiling, 450 m alt., Gates 093 (CAHP): Luzon,

Laguna Prov., Los Bafios (Mt Makiling), E/mer 18054 (BM, K, L); Luzon.

Laguna Prov., Mt Makiling, Pancho 3488 (CAHP); Luzon. Laguna Prov.,

Mt Makiling, 100 m alt., Fernando 7115 (LBC): Luzon. Laguna Prov.,

Mt Makiling, 100 m alt., Fernando 745] (LBC); Luzon, Laguna Prov.,

Mt Makiling, Mabesa 26228 (K, US); Luzon. Batangas Prov., San Jaun,

Laiya, Hernaez 3625 (CAHP); Luzon. Camarines Prov., Curran 10631

(BM, L, US); Luzon. Camarines Sur Prov., Albay, Tabacco, Hernaez 3870

(CAHP); Marinduque Is. Marinduque Prov., Vidal 1943 (K); Mindoro.

Mt Yagaw, 400 m alt., Conklin 535 (PNH): Mindoro. Bongabong R..,

Merritt 4108 (Fl); Mindanao. District of Davao, Todayo (Mt Apo), E/mer

216 Gard. Bull. Singapore 60 (2) 2009

11967 (A, BM, BO, K, L, NY, UC); Samar. Llorente, Mt Apoy, 200-

300 m alt., Guiterrez 367 (PNH); Palawan. Quezon, Panitian, ca 50 m

alt., Dransfield JD6208 (K); Palawan. Mt Tagburos, vicinity of Puerto

Princesa, Ebalao 648 (K); Palawan, Carafugan R., Curran 3784 (BM, FI,

NY, P); No locality: 1904, carpological collections 1712, Burck s.n. (L).

Specimens from cultivated material: INDIA: Madras, 26 May 1900,

A.G. Bourne s.n. with Lady Bourne (K); INDONESIA: Bogor Botanic

Gardens, May 1878, Beccari s.n. (FI); Java. Puger, Koorders 6128 (BO);

Java, Koeli, Koorders 6129 (BO); North Sumatra, Lérzing 12131 (BO);

Bogor Botanic Gardens, Zollinger 2654 (BM). MALAYSIA: Penang Hill,

Viaduct Rd, Methodist Centre, Whitmore FRIO291 (L); SINGAPORE:

Singapore Botanic Gardens, Lawn B near main gate, Furtado 29216 (K);

Singapore Botanic Gardens, near main gate, B area, introduced as sp.

from Negros, Furtado 29393 (A, K, SING); Singapore Botanic Gardens,

Furtado 29394 (A); Singapore Botanic Gardens, near main gate, B area,

’ Furtado 40542 (SING); PAPUA NEW GUINEA: Lae Botanic Gardens,

Croft 71100 (LAE); PHILIPPINES: Luzon, Manila Botanic Gardens,

Loher 1391 (K); Luzon, Pasay City, Mona Lisa Steiner’s garden, Steiner

685 (PNH); Luzon, Los Banfios, College of Agriculture Campus, University

of Philippines, Pancho 2405 (CAHP); SRI LANKA: Royal Botanic

Gardens Peradeniya, S-7, Rutherford 51 with Bandara (K); TRINIDAD

AND TOBAGO: Trinidad, 1884, anon. s.n. (K); UNITED KINGDOM:

Royal Botanic Gardens Kew, Palm House, anon. 000-73.12587 (K);

Royal Botanic Gardens Kew, anon. 000-73.1258 (K).

Local and vernacular names: Footstool Palm, Balla (Mindanao), Luzon

Livistona, Anahao (Tagalog), Bulus (Albay Province), Mindoro Livistona,

Panobao (Tagalog), Pilig (Tagalog); Sergand (Sumatra), Woka (North

Sulawesi).

Habitat and ecology: Locally abundant in swampforest, seasonally dry

swampforest, mangrove margins, rainforest, moist evergreen forest, along

rivers and secondary forest at 0-300 m alt. Flowers all year; fruits all year.

Conservation status: Least concern.

Distribution: Indonesia, Malaysia and the Philippines. In Indonesia on Java

(though possibly escaped from cultivation only), Kalimantan, Sulawesi,

Maluku, and Raja Ampat Islands; in Malaysia, in northern Sabah and

nearby islands; and throughout the Philippines (Fig. 6).

. |

A Taxonomic Account of Livistona R.Br. (Arecaceae) DiGi

Notes: Livistona rotundifolia is treated here as a variable species. Previous

taxonomy, which included a number of taxa that are herein synonymised,

is otherwise difficult to support. The morphological diversity within L.

rotundifolia encompasses both leaf and fruit characteristics; the former

with segment apices erect to semi-pendulous, and the latter with size, of

11-25 mm diam., and colour, at first yellow, then ripening though to red

or to dark violet or bluish-black. These variable characters appear to occur

more or less randomly throughout the entire population.

Livistona rotundifolia was the first species in the genus to be

taxonomically recognised, and named by Rumphius (1741) in the pre-

Linnean publication Herbarium Amboinense, as the mononomial Saribus.

Linnaeus (1753) included Rumphius’ Saribus as part of his broadly

circumscribed Corypha umbraculifera. Subsequently, Lamarck (1786)

extracted Saribus from that taxon and used it as the basis for his Corypha

rotundifolia, which is the first use of the specific epithet, and named for

the round leaves: “... Coryphe a feuilles rondes....”._ Merrill (1917, p.

111) noted that “Saribus Rumph. is the whole basis of Corypha rotundifolia

Lam., which in turn typifies Livistona rotundifolia Mart.”, and therefore

proposed the illustration in Herbarium Amboinense, tab. 8 (Rumphius,

1741), to be the lectotype. Moore (1963a) proposed S. rotundifolius as the

lectotype for the genus Saribus.

The entity of Saribus Rumph. was implicated in other taxa, with

Loureiro (1790) partly basing his Corypha saribus on it. To clarify the

identity of the species, Blume (1838) established the genus Saribus,

utilising the name of Rumphius’ mononomial as his genus name, to include

C. rotundifolia and other taxa, and thus made the combination Saribus

rotundifolius. Soon after, Martius (1838) provided the first synopsis of

Livistona, subsuming Blume’s Saribus and some Corypha species by

various authors, resulting in the currently accepted combination Livistona

rotundifolia. Martius’ account clearly established the relationship of L.

rotundifolia in regards to other taxa, including Linnaeus’ C. wmbraculifera,

Blume’s Saribus, and the versions of C. rotundifolia provided in the

works of Willdenow (1799), Sprengel (1825) and Schultes and Schultes

(1829). In the latter two references, Loureiro’s C. saribus was placed as a

synonym of C. rotundifolia. Martius, however, excluded C. saribus from

his L. rotundifolia, but included it as a synonym of L. cochinchinensis,

thus aligning C. saribus and L. cochinchinensis and establishing the

disassociation of C. saribus from L. rotundifolia. See Notes under L. saribus

for further discussion about this.

Livistona altissima was described by Zollinger (1857) for palms

cultivated at Bogor Botanic Gardens (Miquel, 1868) with a “...trunco

218 Gard. Bull. Singapore 60 (2) 2009

altissimo gracili...” but otherwise resembled L. rotundifolia but lacked

petiolar spines, “...frondibus habitu et conglomeratione L. rotundifolia

Mart. petiolis subrecurvis inermibus....” The undated collection Zollinger

2684 (BM) from Java, is here chosen as the lectotype. Livistona altissima

was first synonymised under L. rotundifolia by Beccari (1931).

Livistona robinsoniana was described by Beccari (1911) based

on Robinson 9265 from Polillo Is (Robinson, 1911), and named for the

collector, Canadian botanist, C. B. Robinson (1871-1913). Beccari related

L. robinsoniana to L. rotundifolia, but distinguished it on fruit colour, being

orange-reddish rather than bluish-black, and in the depth to which the testa

intruded into the endosperm, it being much deeper than in L. rotundifolia.

However, fruit colour in L. rotundifolia as interpreted here, is variable, with

fruit maturing when orange, red, crimson or nearly black.

Beccari (1919a, 1919b) ultimately recognised three subspecies

of L. rotundifolia in the Philippines, L. rotundifolia var. microcarpa, L.

rotundifolia var. mindorensis, and L. rotundifolia var. luzonensis, all of

- which cannot be separated from L. rotundifolia sensu lato. The characters

that Beccari used to delimit the Philippine subspecies were narrow and can

be accounted for in the overall variation that would be expected to occur

in a widespread species. Beccari (1931, p. 76) wrote of L. rotundifolia

that it was “.... a palm of wide geographical distribution and subject,

for that reason, to vary more or less, but easily grouped around one well

characterised type...”.

Regarding fruit colour in L. rotundifolia: it appears that individuals

have uniformity in mature fruit colour, and that variation occurs in individuals

within and between populations. Some of the original designations of fruit

colour for L. rotundifolia included Lamarck (1786), “orange, then red’;

Blume (1838), “yellowish to atro-coerulescentes”; and Beccari (1907)

for L. microcarpa “shining vermilion red, ultimately wine red or nearly

black’.

Livistona rotundifolia is one of a distinct group of closely related

species that has its distribution in Malesia, including the Philippines.

The group is characterised by a trifurcate, or very infrequently bifurcate

inflorescence, and fruit maturing through an orange-red phase to be fully

mature at orange, red, crimson, dark red or black. The group consists of

L. rotundifolia (Indonesia, Philippines), L. merrillii (Philippines), L.

brevifolia, L. chocolatina, L. papuana, L. surru, L. tothur (New Guinea),

and L. woodfordii (New Guinea and Solomon Islands).

Livistona rotundifolia is a variable canopy palm to 45 m tall; leaves

are large and regularly segmented; segment apices are rigid or pendulous,

and with a bifurcate cleft 4-25% of the segment length; the inflorescence is

TE _————

32 But

A Taxonomic Account of Livistona R.Br. (Arecaceae) 219

basally trifurcate or infrequently bifurcate, not extending beyond the limit

of the crown, and with up to 10 partial inflorescences; bracts are tightly

tubular: flowers are yellowish; fruit are globose, to 25 mm diam., and

orange-red to red, dark violet to bluish black at maturity.

4. Livistona papuana Becc., Malesia | (1877) 84. -Saribus papuanus

(Becc.) Kuntze, Revis. Gen. Pl. 2 (1891) 736. — Type: Indonesia. Papua,

Miosnom Is., Apr 1875, Beccari s.n. (holotype, Fl). Plate 4A-B.

Hermaphroditic palm. Trunk to 30 m (50 m7) tall, 12-30 cm dbh, leaf scars

raised, internodes broad, grey. Leaves 17-40 in a globose crown; petiole

111-200 cm long, ca 15 mm wide at the apex, whitish with a thin, flakily

deciduous waxy coating, adaxially flat to slightly ridged, margins armed

throughout, or with spines infrequently confined to proximal portion or

very infrequently lacking; margins sharp when unarmed; spines retrorsely

recurved, to 20 mm long; leaf-base fibres soft, reddish, partly disintegrating:

lamina costapalmate, regularly segmented, subcircular, 90-180 cm long,

100-150 cm wide, adaxially shiny or greyish green, abaxially lighter green,

rigid, waxy glaucous; lamina divided for 23-69% of its length, with 45-

90 segments, depth of apical cleft 5-11% of the segment length; apical

lobes acuminate, rigid; parallel veins 7-8 each side of midrib: transverse

veins thinner than parallel veins. Inflorescences with 3 + similar collateral

axes, branched to 3 orders, each axis 100-225 cm long, not extending

beyond the limit of the crown, arched-nodding, the central axis slightly

longer than the laterals; partial inflorescences 5-10; prophyll to 25 cm long,

to 8 cm wide, glabrous, basally brown, distally yellowish, disintegrating

fibrous; peduncular bracts 1-4, 30-50 cm long, tightly tubular, deeply bifid

and lacerate at the apex, glabrous; rachis bracts tubular-elongate, tightly

sheathing, margins acute; rachis bracts tubular, subterete or very slightly

flattened, mouths truncate, entire or lacerate-fibrous, and extended on one

side into bi-toothed tips; rachillae 3-12 cm long, ca 1 mm thick, straight,

reddish-brown pubescent, most dense underneath and immediately outside

of enclosing bracts, whitish-green on exposed parts. Flowers solitary or

in clusters of 2-4, ca 1.2 mm long, sessile on superficial pulvinuli, with an

inconspicuous bracteole; sepals fused, lobes long, triangular, ca 1 mm long,

apically acute, longitudinally nerved, yellow: petals triangular, obtuse,

apically acute, ca 1 mm long, ca 1.2 mm wide at the base, yellow; stamens

fused for ca 2 the length of the petal, shoulders flat, filament very thin,

ca 0.5 mm high; anthers ca 0.1 mm long: carpels ca 0.8 mm long: stigma

pointed. Fruit globose to obovoid-obpyriform, 14-25 mm long, 5-20

mm diam., orange-red, apex rounded, tapered to a narrow base, stigmatic

220 Gard. Bull. Singapore 60 (2) 2009

remains slightly subapical; epicarp smooth with longitudinally arranged

yellow dots or short lines; mesocarp fibrous, with fibres embedded in the

endocarp; endocarp bony to woody, 1.5-2 mm thick; pedicel 1-5 mm long.

Seed globose, 20-25 mm diam.; embryo sublateral. Eophyll not seen.

Specimens examined: INDONESIA: Papua. Misool Is, Pulalongkakalong,

Heatubun 363 with Iwanggin (K, MAN); Manokwari District, Bintuni

Subdistrict, near Tanah Merah Village, Maturbongs 730 with Siwana &

Kamisopa (BO, K, MAN); Manokwari District, Bintuni Subdistrict, near

Saengga Village, Maturbongs 731 with Siwana & Areta (AAU, BO, K,

LAE, MAN); Miosnom Is., Apr 1875, Beccari s.n. (FI, holotype); Yapen

Is, N side, Sewenui village, Arisoi HA] (MAN); Yapen Is, N side, Sewenui

village, Arisoi HA2 (MAN); Fakfak. Timika, path E at 50 mile on road to

Tembagapura, 04°17.18’S, 137°1.3’E, 540 m alt., Baker WJB851 (AAU,

BH, BO, BRI, K, MAN).

~ Local and vernacular names: Woka (Papuan name), Wanna (Poparo

language).

Habitat and ecology: In rainforest at 200-540 m alt.

Conservation status: Near threatened.

Distribution: Indonesia. In Papua on Miosnom, Biak and Yapen islands,

and the Timika area (Fig. 6).

Notes: Livistona papuana was described by Beccari (1877) from his own

collection from Miosnom Is in Cenderawasih Bay, Papua. He suggested

some resemblance to L. rotundifolia, but from which it was distinct by the

lack of armature and obovoid/obpyriform rather than globose fruit. Beccari

did not describe flowers or mature fruits. Recent collections from Yapen Is,

such as Arisoi HAI (MAN), have yellow flowers and orange-red fruits.

Livistona papuana is a canopy palm to 30 m tall; leaves are large

and regularly segmented; segment apices are rigid, and with a bifurcate

cleft 5-11% of the segment length; the inflorescence is basally trifurcate,

not extending beyond the limit of the crown, and with up to 10 partial

inflorescences; bracts are tightly tubular; flowers are yellow; fruit are

globose, to 25 mm diam., and orange-red at maturity.

5. Livistona brevifolia Dowe & J.P.Mogea, Palms 48 (2004) 201, fig. 6.

— Type: Indonesia. Papua, Kepulauan Raja Ampat, West Waigeo, Kawe Is.,

_— ——

A Taxonomic Account of Livistona R.Br. (Arecaceae) 221

17 Nov. 2002, Mogea 8171 with Takeuchi, Neville & Liuw (holotype, BO;

isotypes, K, L, MAN, NY). Plate 4C-E.

Hermaphroditic palm. Trunk to 22 m tall, ca 12 cm dbh, leaf scars slightly

raised, light grey, internodes narrow, petiole stubs not persistent. Leaves

16-40 in a globose crown; petiole ca 110 cm long, slightly arching, green,

proximally 29-42 mm wide, distally 12-13 mm wide, adaxially slightly

concave, glabrous, with deciduous white waxy scales on the adaxial

surface, abaxially rounded, margins lacking spines; leaf-base fibres coarse,

woven in one layer, brown, persistent until leaf fall then deciduous;

appendage 12-25 cm long; lamina costapalmate, regularly segmented,

semi-circular, moderately undulate, rigid, 55-62 cm long, 45-55 cm wide,

adaxially midgreen, abaxially light green; lamina divided for 17-53% of

its length, with 22-25 segments, depth of apical cleft 1-4% of the length

of the segment, 2-2.5 cm wide where the segments diverge, apical lobes

rigid; hastula poorly developed, strongly asymmetric, lobed; parallel

veins 5-7 each side of midrib; transverse veins thinner than parallel veins.

Inflorescences trifurcate with + similar collateral axes, branched to 3

orders, ca 60 cm long, not extending beyond the limit of the crown, but

with central axis slightly longer and slightly more robust than the lateral

axes; each axis with 2-3 partial inflorescences; prophyll 35-45 cm long,

2.5-3.5 cm wide, papyraceous, entire at the apex, glabrous; peduncle of

central axis subterete to laterally compressed, 18-20 mm diam.; peduncle

of lateral axes terete, 8-12 mm diam.; peduncular bract(s) lacking; rachis

bracts 15-25 cm long, tightly tubular, papyraceous, remaining intact at the

apex with maturity; bases of partial inflorescences with green tomentum;

rachillae 4-9 cm long, straight, subterete to angular, ca 0.5 mm thick,

pubescent, red at floral anthesis. Flowers solitary or in clusters of 2-4;

sepals basally fused, red, with lobes to 1.0-1.2 mm long; petals broadly

triangular, ca 2 mm long, basally connate for ca half their length, thick,

fleshy, red, apical margins recurved; stamens much shorter than petals,

basal part fused to petal, shoulders flat; filament very short; anthers 0.2 mm

long, pink. Fruit globose, 10-12 mm diam.; epicarp thin, smooth, drying

minutely tuberculate with scattered lenticels; stigmatic remains apical;

mesocarp thin, nonfibrous; endocarp thin, crustaceous; pedicel 2-3 mm, ca

1 mm thick. Seed globose, 8-10 mm diam.; endosperm intruded by the testa

to about three quarters across the seed, intrusion of soft tissue irregularly

shaped with minor intrusions extending to the outer edge of the endosperm;

embryo supra-lateral, ca | mm long. Eophyll not seen.

Specimens examined: INDONESIA: Papua, Raja Ampat, Kawe Is., Mogea

222, Gard. Bull. Singapore 60 (2) 2009

8171 with Takeuchi, Neville & Liuw (BO holotype, K, L, MAN, NY);

Papua. Raja Ampat, Kawe Is., Mogea 8224 with Takeuchi, Neville & Liuw

(BO, K).

Habitat and ecology: Forming colonies in open coastal forest on ultrabasic

rocks at 10-20 m alt.

Conservation status: Near threatened.

Distribution: Indonesia. Papua, Raja Ampat, Kawe Island and Gag Island

(Fig. 6).

Notes: Livistona brevifolia was described by Dowe and Mogea (2004) based

on a specimen collected during an ecological survey of the Raja Ampat

Islands (Takeuchi, 2003). Livistona rotundifolia also occurs on Kawe, but

the two species occupy different habitats: L. brevifolia on well-drained

slopes and low ridges, whilst L. rotundifolia occupies moist depressions

and swampforest.

Livistona brevifolia is a moderate canopy palm to 22 m tall; leaves

are smallish and regularly segmented; segment apices are rigid, and with

a bifurcate cleft 1-4% of the segment length; the inflorescence is basally

trifurcate, not extending beyond the limit of the crown, and with up to 3

partial inflorescences; flowers have all parts red, and the anthers pink; fruit

are globose, to 12 mm diam. at maturity.

6. Livistona merrillii Becc., in Perkins, Fragm. Fl. Philipp. 1 (1904) 45;

Webbia | (1905) 339; Philipp. J. Sci. 14 (1919) 340. — Type: Philippines.

Luzon. Tayabas Prov., Guinayangan, Jan. 1903, Merrill 2071 (holotype,

FI). Plate 5A-C.

-Corypha minor Blanco non Jacq., Fl. Filip. (1837) 229.

-Livistona whitfordii Becc., Webbia 1 (1905) 341; Philipp. J. Sci. 4 (1909)

615. — Type: Philippines. Luzon. Quezon Prov., Atimonan, 23 Aug 1904,

Whitford 731 (holotype, FI; isotype, PNH).

-Livistona blancoi Merr., Sp. Blancoan. (1918) 84. — Type: Philippines.

Luzon, Quezon (Tayabas) Prov., Unisan, 11 May 1916, Merrill 919

(holotype, PNH; isotypes, A, BM, BO, K, L, NY).

Hermaphroditic palm. Trunk to 20 m tall, 15-30 cm dbh, leaf scars

prominent, light green to grey in the upper portions aging to grey throughout,

internodes broad, dark green to purple-green in the upper portion, aging to

grey, petiole stubs not persistent. Leaves 40-50 in a + globose crown; petiole

>a 2

A Taxonomic Account of Livistona R.Br. (Arecaceae) 223

90-200 cm long, slightly arching, proximally ca 15 cm wide, distally ca 2.5

cm wide, adaxially flat, with deciduous tomentum, margins most frequently

unarmed in leaves from mature plants or if present the spines confined to

the very proximal part with single curved black spines 6-20 mm long, then

reduced to scattered rudimentary tubercules toward the apex; leaf-base

fibres prominent, coarse, in a woven pattern, persistent after leaf fall and as

intermittent encircling masses in the upper portions of the stem; appendage

long; lamina costapalmate, regularly segmented, circular, 100-150 cm long,

adaxially green, abaxially slightly lighter green; lamina divided for ca 34% of

its length, with ca 70 segments, depth of apical cleft 30-40% of the segment

length, apical lobes acuminate, rigid when depth of apical cleft is shallow,

pendulous when depth of apical cleft is deep; hastula prominent, irregularly

curved, lobed; parallel veins 5-6 each side of midrib; transverse veins thinner

than parallel veins. Inflorescences trifurcate with + similar collateral axes,

branched to 3 orders, 100-150 cm long, not extending beyond the limit of the

crown; each axis with 5-10 partial inflorescences, longest to 40 cm; prophyll

to 45 cm long, 7 cm wide, glabrous, thinly coriaceous, apices acuminate,

caudate; peduncle ca 2.5 cm wide near the base; peduncular bract 1, elongate-

tubular, to 50 cm long, closely longitudinally striate, bicarinate, glabrous,

apices acuminate caudate; rachis bracts tubular, brown, broadening slightly

toward the apex, apices acute, glabrous; rachillae 4-10 cm long, yellowish,

straight, glabrous. Flowers solitary, sessile, 3-4.5 mm long, ovate-acute

in bud, on large raised pulvini; sepals fused basally, fleshy, lobes broadly

subtriangular, thick, yellow; petals elongate-triangular, thick, acute, slightly

spreading at anthesis, bright yellow; stamens with basally fused filaments,

thick and short, contracted into short apices, connective very short, narrow.

Fruit globose to depressed-globose, tapered to the base, 16-23 mm diam.,

dark red, shiny; epicarp smooth; mesocarp 2-3 mm thick, grainy; endocarp

thin, crustaceous; pedicel 2-5 mm long. Seed globose, 11-15 mm diam.;

intruded by the testa to about halfway to almost fully across the seed; embryo

lateral. Eophyll not seen.

Specimens examined: PHILIPPINES: Luzon. Zambales Prov., Curran 5834

(US); Polillo Is. Quezon Prov., Hernaez 3622 (CAHP); Polillo Is. Quezon

Prov., Panukulan, Sitio Kitian, Fernando 8019 (LBC); Luzon. Laguna

Prov., Covinti, Loher 7056 (K); Luzon. Quezon (Payabas) Prov., Curran

10189 (US); Luzon. Quezon Prov., Atimonan, Whitford 731 (FI, PNH);

Luzon. Quezon (Tayabas) Prov., Unisan, Merrill 919 (A, BM, BO, K, L,

NY, PNH); Luzon. Camarines Prov., Paracale, Ramos & Edano 33759 (K,

US); Luzon. Camarines Prov., Curran 10409 (A, UC, US); No location

given, Llanos 221 (MA).

224 Gard. Bull. Singapore 60 (2) 2009

Specimens from cultivated material: Philippines: Luzon, Laguna Prov., Los

Bafios Forestry Campus, Fernando 7337 (LBC).

Local and vernacular names: Ballang (Cagayan Prov., Luzon), Telsis

(Zambales Province).

Habitat and ecology: Forming colonies in Shorea forest and on rocky

limestone hills near the seashore at 60-200 m alt.

Ethnobotany: Leaves used for thatching and for raincoats, stems for bows

and flooring.

Conservation status: Vulnerable.

Distribution: Philippines. Luzon, in Cagayan, Camarines, Tayabas and

Zambales Provinces, and on Polillo Is (Fig. 7).

Notes: Livistona merrillii was first described by Beccari (1904) based on a

100°E 110°E 120°E 130°E 140°E 150°E 160°E

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, L. chocolatina

BL. menillii

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a ff L. woodfordii [TS

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oe Fs ‘,

wes ® =

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= uy 6 ~

= is ace + a” 4 o

<i fe 500 °1,000 1,500 a =

eens |)

i aly

100°E 110°E 120°E 130°E 140°E 150°E 160°E

Figure 7. Distribution of Livistona merrillii (m), L. woodfordii ( &), L. chocolatina (0), L.

tothur (@), and L. surru (©).

Sy SRR ARERR 6

A Taxonomic Account of Livistona R.Br. (Arecaceae) Pipe

specimen from Guinayangan, Luzon, Merri// 2071, with unarmed petioles,

and cherry-like fruit, although only seen in the immature state, and named

for the American botanist, E.D. Merrill (1876-1956), who worked in the

Philippines. Blanco (1837) used the name Corypha minor in his treatment

of Philippine palms, However, the name Corypha minor was first used by

Jacquin (1776) and is the basionym of Sabal minor (Jacq.) Pers. Blanco’s

use of the same name for the Filipino palm constituted a misapplication of a

name already taken, and Beccari’s creation of anew name, L. merrillii, was

the correct procedure, according to Article 58.1 (Tokyo Code). Confusingly,

Blanco’s description of C. minor clearly differentiated it from S. minor

describing his C. minor as “eleva tanto como los cocos y las Arecas” (as

tall as the coconut and Arecas), which contrasts with Sabal minor, i.e.

Jacquin’s C. minor, which is trunkless. Correctly, it should be C. minor

Blanco non Jacq. when used in synonymy.

Beccari (1905), ina subsequent description of L. merrillii, suggested

that C. minor was related to L. whitfordii, a species that he newly described

following his treatment of L. merrillii, but was distinguished from it by the

shallower segment apical clefts. In subsequent descriptions of L. whitfordii,

a species named for American botanical collector, H. N. Whitford (1872-

1941), Beccari (1909) concluded that the petiole margins were basally armed

with spines in contrast to the protologue, which described them as unarmed.

Beccari (1919b) placed L. whitfordii as a synonym of L. merrillii.

Merrill (1918), in reviewing the species established by Blanco,

proposed a new name, L. blancoi, for L. merrillii. It is not clear if Merrill

considered L. blancoi to be a new species or whether it was simply a matter

of providing what he considered to be a more appropriate epithet. The taxon

was named for the Spanish Augustinian friar, F.M. Blanco (1780-1845),

who provided various accounts of the Philippine flora. However, Merrill

distinguished L. blancoi from L. merrillii by “shorter leaf-segments and in

its much more slender petioles” thus implying that he considered that they

were distinct species. In addition, Merrill maintained that L. merrillii, L.

whitfordii and his L. blancoi, had unarmed petioles. The type for L. blancoi

is Merrill 919.

Subsequently, Beccari (1919b, 1931) reiterated the synonymy of

C. minor under L. merrillii, but did not make any reference to Merrill’s L.

blancoi. Merrill (1925), however, reappraised his former placement of C.

minor and L. blancoi and included them as synonyms of L. rotundifolia var.

luzonensis, but still retained L. merrillii as a separate taxon.

Livistona merrillii is a moderate canopy palm to 20 m tall; leaves

are moderate to large and regularly segmented; segment apices are rigid

to pendulous, and with a bifurcate cleft 30-40% of the segment length; the

226 Gard. Bull. Singapore 60 (2) 2009

inflorescence is basally trifurcate, not extending beyond the limit of the

crown, and with up to 10 partial inflorescences; bracts are loosely tubular;

flowers are yellow; fruit are globose to depressed-globose, to 23 mm diam.,

and dark red at maturity.

7. Livistona woodfordii Ridl., Gard. Chron. ser. 3 (1), 23 (1898) 177 (as

L. woodfordi). — Type: Solomon Islands. San Cristobal Is., (1897?) 1898,

Micholitz s.n. (lectotype here designated, BM; isolectotypes, FI, K, SING).

Plate 5D-F.

-Livistona beccariana Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15 (1941)

326. — Type: Papua New Guinea. Milne Bay Prov., Louisiade Archipelago,

1888, W. MacGregor s.n. (lectotype here, designated, BM; isolectotypes,

Fl, MEL).

Hermaphroditic palm. Trunk to 16 m tall, 12-20 cm dbh, leaf scars slightly

raised, irregular in width, light grey, internodes broad, greyish-brown

to grey with age, petiole stubs not persistent. Leaves 30-60 in a globose

to broadly conical crown; petiole to ca 110 cm long, 13 mm wide in the

distal portion, slightly arching, adaxially slightly ridged, flat or slightly

concave, covered with a deciduous white powder, margins unarmed or with

single, small, curved, green spines, confined to the proximal half; leaf-base

fibres moderately prominent, coarse, persistent, reddish-brown; lamina

costapalmate, regularly segmented, subcircular to circular, 60-170 cm long,

45-90 cm wide, rigid, adaxially shiny midgreen, glaucous, abaxially lighter

green with fine powdery wax; lamina divided for 51-75% of its length, with

60-70 segments, depth of apical cleft 5-23% of the segment length, apical

lobes acuminate, semi-pendulous, hanging ca 45° or more to the vertical;

lamina deeply undulate, angles of undulations 90° or less; hastula raised ca 10

mm; parallel veins 5-7 each side of midrib; transverse veins equal thickness

or thinner than parallel veins. Inflorescences with + similar collateral axes,

branched to 3 orders, 120-270 cm long, not extending beyond the limit of

the crown, slightly curving; partial inflorescences 5-10, longest to 45 cm

long; prophyll to 14 cm long; peduncular bracts lacking; rachis bracts

chartaceous, tubular, glabrous, apex acute; tertiary rachis bracts subtend all

minor branches, tubular, glabrous, chartaceous, apex acute; rachillae 4-6 cm

long, | mm thick, straight, basally with brown-purple tomentum, distally

glabrous. Flowers in clusters of 2-6, sessile, subglobose in bud, 1-1.5 mm

long; sepals obtuse, 0.2 — 0.3 mm long, imbricate, red; petals deltoid, 0.8-1

mm long, 0.7-0.8 mm wide, red; filaments with a very narrow connective;

style sharply tapered, ca the same height as the anthers. Fruit globose, 7-12

mm diam., reddish orange to reddish brown; epicarp smooth, shiny, with

scattered lenticellular pores; suture line extends for ca '2 the length of the

fruit, marked with lip-like structures; mesocarp 0.7-1.7 mm thick fieshy

fibrous: endocarp cartilaginous, shiny, yellowish: pedicel

Seed globose, 6-9.5 mm diam.; endosperm intruded by th

the way through to form a central cavity filled with brownish tissue: embryo

lateral. Eophyll not seen.

ia?)

Specimens examined: SOLOMON ISLANDS: Location not given, anon.

H840-14 (K): eae Islands), Woodford 651-14 (K): Tulagi Is..

abundant in wet laterite soils, 90-110 m alt, Dennis BSIP7944 (K, L, LAE,

SING); Tulagi Is., Whitmore s collectors 4427 | K, L, SIN ere US); Tulagzi Is.,

Whitmore 18101 & collectors (K, LAE): Tulagi Is., (N° gela), ca 50 m alt.,

Brass 3517 (A, L; photo BRI); Tulagi Is., W end of Fbaalie 0-50 m alt., Zona

661 (K); San Cristobal Is., (1891?) 1898, Micholitz s.n. (BM lectotype, FI.

K, SING). PAPUA NEW GUINEA: Milne Bay Prov. Cape Vogel Pearically

Menapi, 120 malt ee ee ANB, LAE, US); a ogel Peninsula,

near Tapio village, 9°38°S, 149°53’E, alt. 10 m, Hoogland 4284 (A, CANB.

L, LAE); Rossel Is.. a 50 m alt., Brass 2828] (K, L. LAE, PNH., US):

Rossel Is., Abeleti, 11°20°S, 154°10°E, 50 ft alt. es 27041 (L, LAE):

Rossel Is., Pambwa area, Yeleamba, 50 m alt.. Banka 2015 with Dowe (BRI.

K, LAE): Sudest Is., ca 5 km NW of East Point, 60-70 m alt., Banka 2019

with Dowe (BRI, K, LAE).

Specimens from cultivated material: SINGAPORE: sine nies Botanic

Gardens, Lawn O, 7 Mar 1930. Fi irtado s.n. (K): Singapore Botanic Gardens,

Garden O, Furtado 21175 (K, SING): Singapore Botanic Gardens, —( ) towards

M & Q, probably progeny of the type collection, Furtado 29223 (SING):

Singapore Botanic Gardens, K, near Office. Furtado 29 (BH. K. L, SING).

Local and vernacular names: Boda (Wedau language), Filu (Kakabai

language).

Habitat and ecology: Coastal forest on limestone or lateritic soils at 0-120 m

alt., in high rainfall areas. Flowers Jan- Apr: fruits May-July.

Conservation status: Vulnerable (IUCN, 2006).

Distribution: Papua New Guinea and Solomon Islands. In the Solomon

Islands on Tulagi (Nggela) and San Cristobal Islands. In Papua New ones

in Milne Bay Prov., on Rossel and Sudest Islands and the mainland nea

Cape Vogel and East Cape (Fig

228 Gard. Bull. Singapore 60 (2) 2009

Notes: Livistona woodfordii was described by Ridley (1898) from the

collection Micholitz s.n. from “...insula Polynesiae” and annotated as “S.

S. Islands’ on the specimen labels, and named for the English naturalist,

Charles Morris Woodford (b. 1852, d. 1927), who collected natural history

specimens in the Solomon Islands (1886-1914) and was the colony’s first

Acting High Commissioner (1896-1915) (Woodford, 1890; Golden, 1993).

There are discrepancies regarding the collection date of the type specimen.

On the type it is written as 1898, but the itinerary of Micholitz shows he was

in the Solomon Islands in October 1897, and in the Louisiade Archipelago,

Papua New Guinea, in Apr/May 1898 (van Steenis-Kruseman, 1950). The

date on the type could relate to the time that Ridley received the specimen

rather than its collection date. An expanded description was provided

by Beccari (1931), and was in part based on an inflorescence and fruit

collection, Macgregor s.n., in the Louisiade Archipelago, most probably in

1888 (Thomson, 1889; van Steenis-Kruseman, 1950). Beccari(1931) related

that the MacGregor specimen was sent to him by Mueller prior to 1889 (in

1888, via Hermann Wendland, according to Burret (1941)). Burret (1941),

providing taxonomic and descriptive notes on some species of Livistona,

concluded that the Louisiade collection was a different species and named

it L. beccariana. As Burret’s description was based on the Macgregor

collection, it is the type. Burret provided a detailed description of the

inflorescence and fruit fragments, but nothing of the palm’s habit or leaves.

Livistona beccariana is hereby placed as a synonym of L. woodfordii.

Livistona woodfordii 1s a moderate canopy palm to 16 m tall; leaves

are moderate and regularly segmented; segment apices are semi-pendulous,

and with a bifurcate cleft 5-23% of the segment length; the inflorescence is

basally trifurcate, not extending beyond the limit of the crown, and with up

to 10 partial inflorescences; bracts are tubular; flowers (sepals and petals)

are red; fruit are globose, to 12 mm diam., and reddish orange to reddish

brown at maturity.

8. Livistona chocolatina Dowe, Palms 48 (2004) 199, figs 2, 3. — Type:

Papua New Guinea. Central Prov., Kuriva Mission area, 4 km north of

Haritano Hwy along forestry road, 9°00.821°S, 147°07.815’E, 300 m alt.,

4 Mar 2000, Barfod 466 with Banka, Dowe & Kjaer (holotype, AAU;

isotypes, BRI, CANB, K, LAE). Plate 6A-C.

Hermaphroditic palm. Trunk to 22 m tall, 16-18 cm dbh, leaf scars slightly

raised, internodes narrow, light grey, petiole stubs not retained. Leaves 30-

40 in a globose crown; petiole 110-155 cm long, slightly arching, green,

proximally ca 3 cm wide, distally ca 2 cm wide, adaxially flat, glabrous

7 |

A Taxonomic Account of Livistona R.Br. (Arecaceae) 229

with a cover of deciduous white waxy powder, margins usually spineless

in mature plants, or with small single spines to 5 mm long only in the very

basal portion in juvenile plants; leaf-base fibres course, brown, persistent

until leaf fall then readily deciduous; appendage short; hastula ca 1 cm

high, 5 cm across with a central division; lamina costapalmate, regularly

segmented, subcircular, flat, rigid, 100-120 cm long and wide, adaxially

mid grey green, abaxially light grey green, glaucous waxy; lamina divided

for ca 44% of its length, with 45-60 segments, depth of apical cleft ca 4% of

the segment length, apical lobes rigid; lamina moderately undulate, angles

of undulations 91° or more usually much greater; mid-leaf segments 4-5 cm

wide where the segments diverge; parallel veins 7-8 each side of midrib;

transverse veins thicker than parallel veins. Inflorescences trifurcate with +

similar collateral axes, 195-225 cm long, not extending beyond the limit of

the crown, branched to 3 orders, but with central axis slightly more robust

than the lateral axes; each axis with 6-10 partial inflorescences; prophyll

22-37 cm long, 8-15 cm wide, glabrous, chartaceous, lacerate-fibrous at the

apex, basally brown, distally yellow; peduncle of central axis subterete, to

2.8 cm diam.; peduncle of lateral axes terete, to 1.6 cm diam.; peduncular

bracts 2-4; peduncular bracts glabrous, tubular, lacerate at the apex; rachis

bracts 40-45 cm long, tightly tubular, fibrous, disintegrating at the apex

with maturity, pubescent throughout but more densely so toward the apex,

light reddish brown; bases of partial inflorescences with dense chocolate

brown tomentum; rachillae 8-12 cm long, straight, subterete to angular, 2-3

mm thick, basally with chocolate brown tomentum, distally with cream-

green tomentum. Flowers solitary or in clusters of 2-4, tightly aggregated

in bud and during anthesis, ca 1.2 mm high; sepals basally fused, lobes

long, triangular, ca | mm long, red, apically acute, longitudinally nerved;

petals triangular, obtuse, red, apically acute, ca 1 mm long, ca 1.2 mm

wide at the base; connective very thin, ca 0.5 mm long; anthers ca 0.1

mm long; carpels ca 0.8 mm high, stigmas pointed. Fruit globose, ca 25

mm diam., shiny orange-red; epicarp with scattered lenticellular dots and

light 3 mm long lines pointing in longitudinal direction toward the apex;

stigmatic remains apical to slightly subapical; mesocarp fleshy, fibres thick,

distributed throughout but more densely aggregated toward the endocarp

and shallowly embedded in the surface of the endocarp; endocarp to |

mm thick, bony; pedicel 4-5 mm long, 2 mm thick, jointed, green, with

prominent scars of fallen flowers. Seed globose; endosperm intruded by the

seed coat to ca two-thirds across, intrusion broadly kidney-shaped; embryo

lateral. Eophyll 5-ribbed.

Specimens examined: PAPUA NEW GUINEA: Morobe Prov. Lababia,

230 Gard. Bull. Singapore 60 (2) 2009

Bulili Ridge, 400 m alt., Barfod 514 with Kjaer & Magun (AAU, LAE):

Gulf Prov. Vailala R., hills inland, Lane-Poole 332 (BRI); Central Prov.

Kuriva Mission, Ferrero 980080, 980081, 980083 (LAE); Central Prov.

Kuriva Mission area, 4 km north of Haritano Hwy along forestry road,

9°00.821°S, 147°07.815°E, 300 m alt., Barfod 466 with Banka, Dowe &

Kjaer (AAU holotype, BRI, CANB, K, LAE).

Local and vernacular name: Manganau (Lababia language).

Habitat and ecology: Grows in isolated colonies, sometimes locally

common, on slopes with calcareous or clayey soils, at 300-400 m alt.

Flowers Jan-Feb; fruits Mar-May.

Conservation status: Vulnerable.

Distribution: Papua New Guinea. In Central Province in the Kuriva area,

in Gulf Province on hills near the Vailala R., and in Morobe Province near

Lababia on Bulili Ridge (Fig. 7).

Notes: Livistona chocolatina was first collected by Lane-Poole in 1922 as

‘Livistona sp. No. 332’ (Lane-Poole, 1925) from “hills inland from Vailala

River”. It was collected again in 1998 by M. Ferrero, and then again in

2000 by Barfod ef a/., and from whose collections the type Barfod 466 was

chosen. The species is distinguished by the usually spineless or only mildly

spined petiole that initially has a thick coating of white waxy powder,

smallish rigid leaves, a trifurcate inflorescence with each axis having

multiple peduncular bracts, distinctive chocolate brown tomentum on the

proximal surfaces of the partial inflorescences and rachillae, and globose

fruit to 25 mm diameter that mature orange-red.

9. Livistona tothur Dowe & Barfod, Austrobaileya 6 (2001) 171, fig. 3. —

Type: Papua New Guinea. West Sepik Prov., Oenake Mts, on road to Niau

Kono from Vanimo, 2°45.89’ S, 141°04.06’ E, 500 m alt., 26 Nov 1996,

Damborg 418 with Barfod (holotype, AAU; isotype, BRI, K, LAE). Plate

6D-E, Plate 7A-D.

Hermaphroditic palm. Trunk to 20 m tall, 15-20 cm dbh, leaf scars

slightly raised, narrow, dark grey, internodes broad, grey. Leaves 24-40

in a globose crown; petiole 150-200 cm long, proximally ca 10 cm wide,

distally ca 1.5 cm wide, adaxially flat, arching, glabrous, green, margins

with single, recurved, green spines 1-2 mm long throughout its length, but

A Taxonomic Account of Livistona R.Br. (Arecaceae) 23)

larger and more closely spaced in the proximal portion, margins lacerate

fibrous in extreme basal portion; leaf-base fibres prominent, in 2 layers

of more or less equal coarseness, persistent in sheets, chestnut brown;

appendage to 60 cm long; hastula raised, small, semi-circular; lamina

costapalmate, regularly segmented, subcircular, 150-200 cm long, 120-

150 cm wide, adaxially bluish-green, abaxially silvery glaucous; lamina

divided for 62-85% of its length, with 60-75 segments, depth of apical cleft

1-3% of the segment length, apical lobes rigid, but becoming pendulous

with age or damage, segment 3-4 cm wide where the segments diverge;

parallel veins 6-7 each side of midrib; transverse veins thinner than parallel

veins. Inflorescences trifurcate with + similar collateral axes, branched to

3 orders, each axis ca 200 cm long, not extending beyond the limit of the

crown, with 5-6 partial inflorescences; prophyll to 30 cm long, 10.5 cm

wide, yellow, coriaceous basally, brown chartaceous distally, glabrous apart

from ferruginous woolly tomentum along the carinae; peduncle with one

tubular, papery and loosely sheathing peduncular bract, densely scaled at

the apex otherwise glabrous; peduncle of individual axes basally subterete,

becoming dorsi-ventrally compressed distally, edges angular, furfuraceous

in parts enclosed in bracts, otherwise patchily furfuraceous or glabrous on

exposed surfaces; rachis bracts tubular, papery, loosely sheathing, glabrous

apart from some irregular pubescence toward the apex, apices more or less

nonfibrous showing only minor disintegration; rachillae 6-12 cm long, to

3 mm diam., straight, green-red, glabrous. Flowers solitary; sepals basally

fused, red, with lobes to 1-1.2 mm long; petals broadly triangular, ca 2

mm long, basally connate for ca half their length, thick, fleshy, red, apical

margins recurved, outer surface minutely warty; stamens much shorter

than petals, basal part fused to petal, shoulders flat; filament very short;

anthers 0.2 mm long, cream. Fruit globose with a basal constriction, 35-

43 mm diam., semi-glossy orange-red, stigmatic remains apical; epicarp

with scattered lenticellular dots, glossy, longitudinal stripe of suberised

epidermal tissue usually visible for the full length of the fruit; mesocarp

13-15 mm thick, softly fibrous, mealy, orange; endocarp to 2 mm thick,

bony; pedicel 2.5-5 mm high, ca 2 mm wide. Seed globose, 22-28 mm

diam., endosperm deeply intruded by orange pulpy tissue; embryo lateral.

Eophyll not seen.

Specimens examined: PAPUA NEW GUINEA: West Sepik Prov. Oenake

Mts, Apol area, on road to Niau Kono from Vanimo, 2°45.89S, 141°04.06E,

500 m alt., Dowe 516 with Ferrero (JCT, LAE); Oenake Mts, Niau,

2°46.276°S, 141°03.611°E, 425 m alt., Barfod 510 with Banka & Kjaer

(AAU, BRI, JCT, K, LAE); Oenake Mts, on road to Niau Kono from

232 Gard. Bull. Singapore 60 (2) 2009

Vanimo, 2°45.89°S, 141°04.06’E, 500 m alt., Damborg 418 with Barfod

(AAU holotype, BRI, K, LAE).

Local and vernacular names: Tot-hur and Yu Bbraal (Bewani language).

Habitat and ecology: Rainforest on ridges, limestone and metamorphic

rocks, at 400-600 m alt. Flowers Nov-Jan; fruits Feb-Mar.

Ethnobotany: Roofs and umbrellas are made from the leaves, bows from

the split trunk, and salt is obtained from the ash of burned petioles.

Conservation status: Near threatened.

Distribution: Papua New Guinea. In West Sepik Prov., Oenake Mts (Fig.

‘Notes: The first recognition of Livistona tothur in the literature was an

informal note by Essig & Young (1981) who described a palm seen from

their helicopter as “a large Livistona bearing red fruit’. Upon landing

nearby, they were only able to find juveniles that they photographed, and

which closely resembled those that are now known in populations of L.

tothur. Subsequently, field work in New Guinea was undertaken by Ferrero

(1997) who investigated reports of palm populations throughout Papua

New Guinea and was to eventually collect and record the populations of

this species in the Oenake Mountains of West Sepik Prov. Further field

work undertaken by Barfod and Damborg resulted in collections, including

the type, Damborg 418, upon which this taxon was described. The species

name was taken from the vernacular, tot-hur, used in the Bewani area, West

Sepik Prov., Papua New Guinea.

Livistona tothur is a moderate canopy palm to 20 m tall; leaves

are moderate and regularly segmented; segment apices are rigid, and with

a bifurcate cleft 1-3% of the segment length; the inflorescence is basally

trifurcate, not extending beyond the limit of the crown, and with up to 6

partial inflorescences; bracts are loosely sheathing; flowers (sepals, petals

and carpels) are red; fruit are globose, to 25 mm diam., and shiny orange-

red at maturity.

10. Livistona surru Dowe & Barfod, Austrobaileya 6 (2001) 169, fig. 2.

— Type: Papua New Guinea. West Sepik Prov., Miwaute, 03° 25’S, 142°

07°S, 950-1000 m alt., 20 Nov 1996, Barfod 390 with Ferrero & Damborg

(holotype, AAU; isotypes, BRI, K, LAE). Plate 8A-E.

di Sia eRe UR eer sb

oS)

4 Taxonomic Account of Livistona R.Br. (Arecaceae)

Hermaphroditic palm. Trunk to 20 m tall, 18-25 cm dbh, usually covered

by crustaceous lichens, leaf scars slightly raised, internodes narrow, light

grey, petiole stubs not retained. Leaves 17-29 in a + globose crown:

petiole 140-180 cm long, slightly arching, green, ca 25 mm wide in the

middle, adaxially flat, glabrous except for scattered lepidote scales that are

brown in the centre and grey at the margin, more densely so on the abaxial

surface, margins with single or grouped black spines 5-10 mm long, larger

and more closely inserted in the proximal portion, becoming smaller and

wider spaced in the distal portion; leaf-base fibres in 2 layers, the outer

with thick fibres, the inner with thin coir-mat like fibres, reddish brown,

persistent until leaf fall then deciduous in sheets; appendage to | m long,

to 10 mm thick; hastula very prominent, 2 cm high, 5 cm across; lamina

costapalmate, regularly segmented, subcircular to ovate, undulate, | 80-224

cm long, 143-160 cm wide, adaxially mid green, abaxially similar green;

lamina divided for 45-80% of its length, with 70-90 segments, depth of

apical cleft ca 6% of the segment length: apical lobes pendulous, hanging

+ vertically; mid-leaf segments 4.5-7 cm wide where the segments diverge:

parallel veins 5-6 each side of midrib; transverse veins thinner than parallel

veins. Inflorescences trifurcate with + similar collateral axes ca 120 cm

long, not extending beyond the limit of the crown, branched to 3 orders:

each axis with 5-7 partial inflorescences; prophyll 37-42 cm long, 12.5-15

cm wide, glabrous, lacerate-fibrous at the apex; peduncle of individual axes

subterete, to 3 cm wide, peduncular bract(s) lacking: rachis bracts 40-45

cm long, loosely tubular, fibrous, disintegrating at the apex with maturity,

pubescent throughout but more densely so toward the apex; rachillae 14-24

cm long, subterete to angular in cross section, pubescent with dense long

coarse red appressed scales in the proximal portion, distally with long white

scales, less dense to glabrous in the extreme distal portions. Flowers in

clusters of 2-4 (only decayed ones seen). Fruit globose to obovoid, 55-65

mm long, 50-55 mm diam., orange-red; epicarp with scattered lenticellular

dots and light 3 mm long lines pointing in longitudinal direction toward the

apex; stigmatic remains apical; longitudinal stripe of suberised epidermal

tissue usually visible for full length of fruit; mesocarp fleshy, fibres thick,

distributed throughout but more densely aggregated toward the endocarp

and shallowly embedded in the endocarp; endocarp to 2 mm thick, bony;

pedicel 6-12 mm long, 3 mm thick, green, with prominent scars of fallen

flowers. Seed globose to subglobose; endosperm intruded by the seed coat

to ca two-thirds across, intrusion broadly kidney-shaped, orange; embryo

lateral. Eophyll not seen.

Specimens examined: PAPUA NEW GUINEA: West Sepik Prov., Miwaute,

234 Gard. Bull. Singapore 60 (2) 2009

03°25’S, 142°07’°E, 950-1000 m alt., Barfod 399 with Ferrero & Damborg

(AAU, LAE); West Sepik Prov., Miwaute, 03° 25’S, 142° 07’S, 950-1000 m

alt., Barfod 390 with Ferrero & Damborg (AAU holotype, BRI, K, LAE);

West Sepik Prov., Upper Freida R., Mt Ekwai, 1200-1300 m alt., Ferrero

980029 (LAE). Madang Prov., Goinbang, near Bosmun 2, mouth of Ramu

R., on road from Bogia, Baker 582 with Utteridge (BH, FTG, K); Madang

Prov., Bosmun Village at mouth of Ramu R., 04°07’S, 144°43’E, Damborg

354 with Ferrero & Barfod (AAU).

Local and vernacular names: Surru, Bop or Tim (Olo language at

Miwaute).

Habitat and ecology: In rainforest and swampforest, 10-1300 m alt. Flowers

all year; fruits all year.

Ethnobotany: Leaves are used for roof thatch and umbrellas, stem portions

for axe handles and house frames, and leaf sheath fibres for brooms and

sago strainers.

Conservation status: Near threatened.

Distribution: Papua New Guinea. In Madang Province near Bosmun village

at the mouth of the Ramu R., and in West Sepik Province in the Miwaute

area and Mt Ekwai (Fig. 7).

Notes: The first mention of Livistona surru was by Hay (1984, p. 208) who

noted a Livistona at Ramu R.: “ ... here the Livistonas are growing in a

remarkable rain forest dominated almost to the exclusion of dicotyledonous

trees by palms of the genera Actinorhytis, Rhopaloblaste, Cyrtostachys,

Ptychococcus, Orania, Gulubia, Caryota and Livistona”’. Subsequently,

this report was investigated by Ferrero (1997) and by Barfod who collected

the type specimen, Barfod 390. The specific name was taken from the

vernacular surru, from the Olo language used in the Miwaute area of West

Sepik Prov., Papua New Guinea.

Livistona surru 1s a moderate to large canopy palm to 20 m tall;

leaves are large and regularly segmented; segment apices are pendulous,

and with a bifurcate cleft to 6% of the segment length; the inflorescence is

basally trifurcate, not extending beyond the limit of the crown, and with up

to 7 partial inflorescences; bracts are loosely tubular; fruit are globose, to

65 mm diam., and orange-red at maturity.

A Taxonomic Account of Livistona R.Br. (Arecaceae) 235

11. Livistona halongensis T.H.Nguyen & Kiew, Gard. Bull. Singapore 52

(2000) 198, pl.la; Wild Pl. Ha Long Bay (2000) 35, pl. 24.— Type: Vietnam.

Quang Ninh Prov., Ha Long Bay, 12 May 1999, Nguyen, Averyanov &

Nguyen NTH2630 (holotype, HN; isotype, SING). Plate 9A-B.

Hermaphroditic palm. Trunk to 10 m tall, ca 20 cm dbh, leaf scars

conspicuous, internodes narrow. Leaves ca 40 in a + globose crown;

petiole 125-130 cm long, ca 2 cm wide at the base, ca 1.5 cm wide distally,

adaxially flat, green with a light yellow band along the margin, margin with

irregularly spaced triangular hooked dull orange spines along entire length

but more aggregated proximally; leaf-base fibrous, appendage triangular,

rich brown, fibrous, distally tattered into ribbons; hastula papery, to 2.5 cm

high; lamina costapalmate, regularly segmented, circular, 75-80 cm long,

undulate, adaxially dark green, abaxially lighter green; mid-leaf segments

ca 4 cm wide where the segments diverge; lamina divided for 42-73% of its

length, with 45-64 segments, depth of apical cleft ca. 40% of the segment

length, apical lobes rigid to semi-pendulous; parallel veins 11-13 each side

of midrib; transverse veins thinner than parallel veins; basal part of outer

segment with a 2 cm long row of fine teeth on the margin. Inflorescences

unbranched at the base, ca 340 cm long, erect, projecting vertically above

the crown and extending well beyond the limit of the crown, branched

to 3 orders; partial inflorescences 6-7, longest to 95 cm; peduncle ca 160

cm long, 2 cm diam.; prophyll to 33 cm long, basally 5 cm wide, distally

to 3 cm wide; peduncular bracts 5, each ca 50 cm long, tightly tubular,

thick, leathery-fibrous; rachis bracts ca 27 cm long, otherwise similar to

peduncular bracts; rachillae 1.0-1.5 mm diam., cream, velvety tomentose.

Flowers in pairs on pedicel ca 0.3 mm long, ovoid in bud, pale cream-

yellow; sepals basally fused with lobes ca 1 mm long, glabrous; petals ca

2 mm long; stamens ca | mm long, subulate, staminal tube rich brown,

filament very short; anthers white, ca 0.3 mm long; carpel obovoid, deeply

ridged, dark reddish brown, ca | mm long; style pale brown, ca 0.3 mm

long; stigma minute. Fruit globose, 10-12 mm diam., epicarp smooth,

glossy dark green; mesocarp leathery; endocarp crustaceous, brittle, ca

0.75 mm thick; endosperm intruded by seed coat to ca halfway through;

pedicel 2-5 mm long. Eophyll not seen.

Specimens examined: VIETNAM: Quang Ninh Prov., Ha Long Bay,

Nguyen, Averyanov & Nguyen NTH2630 (HN holotype, SING); Quang

Ninh Prov., Ha Long Bay, 15 Jul 1999, Ngwyen & Kiew s.n. (HN); Ha Long

Bay, 11 Apr 2000, Nguyen & Kiew s.n. (HN); Hai Phong Prov., Cat Ba NP,

limestone hill, ca 300 m alt., Nguyen Tien Hiep NTH 4853 (HN).

236 Gard. Bull. Singapore 60 (2) 2009

Habitat and ecology: Confined to limestone islands, in soil-filled crevices

on rocky limestone substrate, sometimes gregarious, 100-300 m alt. Flowers

May-Jun; fruits July.

Conservation status: Lower risk, conservation dependent.

Distribution: Vietnam. Islands in Ha Long Bay (Fig. 8).

100°E 110°E 120°E 130°E 140°E 150°E

40°N

30°N

20°N

Legend

Ryne Dia Soe)

AN = ast L. boninensis

z i be Ss ~ @ L. chinensis <

° } \ XS [Sd

-| @ @ L. halongensis | —

= t > tt r

100°E 110°E 120°E 130°E 140°E 150°E

Figure 8. Distribution of Livistona halongensis (@), L. chinensis (m), and L. boninensis

(Q).

Notes: Livistona halongensis was described by Nguyen and Kiew (2000)

from limestone islands in Ha Long Bay, Gulf of Tonkin, Vietnam, based

on Nguyen Tien Hiep NTH 2630. This is the only Livistona known to be

restricted to limestone. It was previously mentioned by Yang (1996) as an

“unidentified species of Livistona” from Cat Ba Is.

Livistona halongensis is a smallish sub-canopy palm to 10 m tall;

leaves are small and regularly segmented; segment apices are rigid to

semi-pendulous, and with a bifurcate cleft to 40% of the segment length;

OGLE ee |

A Taxonomic Account of Livistona R.Br. (Arecaceae) D3i7/

the inflorescence is unbranched, extending vertically above the crown by

about | m, and with up to 7 partial inflorescences; bracts are tightly tubular;

flowers are cream-yellow; fruit are globose, to 12 mm diam., and glossy

dark green at maturity.

12. Livistona chinensis (Jacq.) R.Br. ex Mart., Hist. Nat. Palm. 3(7) (1838)

240. -Latania chinensis Jacq., Fragm. Bot. (1801) 16, t. 11, fig. 1. -Saribus

chinensis (Jacq.) Blume, Rumphia 2 (1838) 49. -Livistona sinensis Griff.,

Palms Brit. E. Ind. (1845) 131, ortho. var. — Type: Illustration in Jacquin,

Fragm. Bot. (1801) t. 11, fig. 1, (lectotype, fide Moore, 1979). Plate 9C-F.

-Latania borbonica auct. non Lam., Encyclop. 3 (1792) 427.

-Livistona mauritiana Wall. ex Mart., Hist. Nat. Palm. 3 (1838) 240,

nomem.

-Chamaerops biroo Siebold, Syn. Pl. Oecon. Jap. (1830) 11; Martius, Hist.

Nat. Palm. 3 (1838) 242. — Type: Lectotype here chosen. Japan. undated,

Siebold s.n. (lectotype, L).

-Livistona olivaeformis (Hassk.) Mart., Hist. Nat. Palm. 3 (1850) 319;

Miquel, Fl. van Ned. Ind. 3 (1855) 59; Palm. Archip. Ind. (1868) 13.

-Saribus olivaeformis Hassk., Tijdschr. Natuurl. Gesch. Physiol. 9 (1842)

176. -Latania olivaeformis (Hassk.) Devansaye, Rev. Hort. 47 (1875) 34,

nom. inval. — Type: Indonesia. Cultivation, Bogor Botanic Gardens, ‘nel

viale presso la chieta’, May 1878, Beccari s.n. (lectotype, here designated,

FI, sheets 1131, 1131-B & 1131-C).

-Livistona subglobosa (Hassk.) Mart., Hist. Nat. Palm. 3 (1850) 319;

Miquel, Fl. van Ned. Ind. 3 (1855) 59; Palm. Archip. Ind. (1868) 13; Nakai,

J. Jap. Bot. 11 (1935) 224. -Saribus subglobosus Hassk., Tijdschr. Natuurl.

Gesch. Physiol. 9 (1842) 177; Cat. Hort. Bot. Bog. (1844) 65. -Livistona

chinensis var. subglobossa (Hassk.) Becc., Webbia 5 (1921) 2, 16. — Type:

Indonesia. Cultivation, Bogor Botanic Gardens, May 1878, Beccari s.n.

(lectotype, here designated, FI, sheets 11333 & 11333-B).

-Livistona japonica Nakai ex Masam., (as Livistonia japonica), Prelim.

Rep. Veg. Yak. (1929) 50; Masamune & Suzuki, Ann. Rep. Taihoku Bot.

Gard. 1933, 3 (1933) 73. Type not designated.

Hermaphroditic palm. Trunk to 15 m tall, 20-30 cm dbh, leaf scars obscure,

roughened and with remnant tissue, light coloured, internodes narrow,

irregular, brown to grey with age, petiole stubs not persistent, longitudinal

fissures prominent. Leaves 40-60 in a globose to ovoid crown; petiole

slightly arching, to 180 cm long, ca 15 cm wide in the middle, adaxially

flat or slightly ridged, surface glabrous; margins with single curved spines

in the proximal one-half or less or sometimes absent, distal margins

238 Gard. Bull. Singapore 60 (2) 2009

otherwise sharp and slightly winged; spines 2-20 mm long, subulate, green

to black; leaf-base fibre not prominent, coarse, and disintegrating; lamina

costapalmate, regularly segmented, subcircular to reniform, 120-200 cm

long, 120-180 cm wide, light yellowish green on both surfaces, dull to

glossy, nonwaxy; lamina divided for 45-55% of its length, with 50-90

segments, depth of apical cleft ca 13% of the segment length, apical lobes

pendulous; lower segments 2-5 mm wide, filamentous, parallel veins 8-9

each side of midrib; transverse veins thinner than parallel veins; hastula

ca 3 cm high. Inflorescence unbranched at the base, 100-120 cm long,

not extending beyond the limit of the crown, slightly curving, branched

to 3 orders; 6-7 partial inflorescences; rachillae 10-18 cm long, to 9 mm

diameter, glabrous; peduncular bract(s) lacking; rachis bracts tomentose or

scaly, loosely tubular, disintegrating into open chartaceous fibres with age,

brown, apex acute. Flowers in clusters of 4-7, subglobose in bud, 2-2.5

mm long, white to yellowish; sepals imbricate, ca 1.5 mm long, rounded,

margins hyaline, inner surface striate; petals triangular, apically pointed,

fleshy, ca 2.2 mm long, ca 2 mm wide; filaments united at the base, free

in the upper part, connective thickish, ca 0.5 mm long; anthers ca 0.5

mm long; style sharply tapered, ca the same height as the anthers. Fruit

globose, subglobose, ellipsoid or pyriform, 15-26 mm long, 9-18 mm wide,

glossy blue-green to bright green; epicarp ceramic-like; suture line extends

for full length of fruit, marked with lip-like structures; mesocarp ca 1.5

mm wide, fleshy, slightly oily, moderately fibrous, endocarp woody, ca 0.5

mm wide; pedicel 2-3 mm long. Seed globose, subglobose to ellipsoid,

variously reniform in longitudinal section, ca 14 mm long, ca 10 mm wide;

endosperm intruded by the testa to ca one-half to two-thirds through to

form a cavity filled with brown crystalline tissue; embryo supra-lateral to

lateral. Eophyll 7-ribbed.

Specimens examined: JAPAN: Yakushima, Ambo, Ohwi & Okamoto 995

(A, B, BM, BR, K, L, NY, UC, US); Ryukyu Islands (Loo-Choo Islands).

1853-56, Wright s.n. (A); Ryukyu Islands. Dharen Is., Wilson 8164

(K); Ryukyu Islands. Okinawa Is., Motobu Peninsula, Mt Awa, Walker

7636 (S); Ryukyu Islands. Yaeyama Islands, Mt. Kubura, Yonakuni,

Furuse 4570, 4577 & 4579 (K); No location, Siebold s.n. (L). TAIWAN:

Bankinsing Mts, Henry 827 (K, NY). CHINA: Canton. 1880, Hance s.n.

(K); Hainan Is. Loktung, Lau 27175 (A); Guangdong Prov., Levine 479

(US); Guangdong Prov., Jo Kang Ping 10956 (BO, NY); No location,

Burch 1591 (L).

Specimens from cultivated material: ANTIGUA: Collector name illegible

——————EeEeEeEe"

A Taxonomic Account of Livistona R.Br. (Arecaceae) 239

II (K); CHINA: Hong Kong, Victoria Is., Deep Water Bay, Shiu Ying Hu

7067 (K, US); Victoria Peak, Shiu Ying Hu 5452 (K); CUBA. Botanic

Garden Havana, Curtiss 656 (K); GUAM: Merizo, Fosberg 35668 (BH,

US); GUYANA: Henritta roadside, Essequibo, Harris 538 (K); INDIA: Ex

H. Bot. Sahar, 1887, Beccari s.n. (K); Tiruchirapalli, St Josephs College

House, Matthew 30953 (K); Tamilnadu, Villupura District, Auroville Shakti,

Matthew 72880 (K); INDONESIA: Bogor Botanic Gardens, May 1878,

Beccari s.n. (Fl, K); Bogor Botanic Gardens, ‘nel viale presso la chieta’,

May 1878, Beccaris.n. (FI, sheets 1131, 1131-B & 1131-C); Bogor Botanic

Gardens, May 1878, Beccari s.n. (FI, sheets 11333 & 11333-B). JAPAN:

Ryukyu Islands, Okinawa, Gesashi, 26°36’°N, 128°08.75°E, Morgan 5058

(UC, US); MALAYSIA: Sabah, Sandakan, Elopura, Me/grito 10029 (K);

NIGERIA: Calabar, Ndozi 15441] (K); PANAMA: Panama City, Curundu

District, Churchill 6023 with Churchill (K); PHILIPPINES: Luzon,

Laguna Prov., Mt Makiling Garden, Pancho 3684 (CAHP); Zamboanga

City, Zambowooe, Canadian Logging Concession, Hernaez 3663 (CAHP);

TRINIDAD: Holloway s.n. (BRI); SEYCHELLES: Insula Platte, Bojer s.n.

(K); SINGAPORE: Location not given, 24 Dec 1894, anon. s.n. (MEL);

Singapore Botanic Gardens, Lawn D, 22 Jun 1929, Nur s.n. (K); Singapore

Botanic Gardens, Furtado 29396 & 29397 (A, BRI, K); Singapore

Botanic Gardens, Lawn B, Furtado 29398 (K); MacRitchie Reservoir,

27 Aug 1970, Tambi s.n. (K, SING); SOCIETY ISLANDS: Moore 443

(BRI); SRI LANKA: Royal Botanic Gardens, Peradeniya, Rutherford 52

with Bandara (K); Royal Botanic Gardens, Peradeniya, Garden Q-237,

Rutherford 101 with Bandara (K); TANZANIA: Amani, Greenway 1042

(K); THAILAND: Pattani, Pattani University Campus, Barfod 40986 with

Ueacharakan (AAU); Trang, Khao Chong, ca 15 km E of Trang, Barfod

141431 with Ueacharakan (AAU); Trang, Khao Chong, Peninsula Botanic

Garden, Barfod 43885 (AAU, BKF); UNITED KINGDOM: Ex Herbarium

Benthamianum, Hort. Chil. May 1823, anon. s.n. (K); Ex Herbarium

Hookerianum 1867, anon. s.n. (K); Royal Botanic Gardens Kew, Feb 1880,

anon. s.n. (K); Royal Botanic Gardens Kew, 1880, anon. s.n. (K); Royal

Botanic Gardens Kew, Propagating House, 1880, anon. 210 (K); Royal

Botanic Gardens Kew, 29 Apr 1882, anon. s.n. (K); Royal Botanic Gardens

Kew, Temperate House, Dransfield 4463 (K); Royal Botanic Gardens Kew,

anon. 1982-5601 (K); UNITED STATES: Florida. Fairchild Tropical

Gardens, Balick 3388 (NY); Hawaii. University of Hawaii campus, 80 ft

alt., Krajina 6288 (NY); Kauai, Lihue District, Nuhou Grove Farm, Wood

2102 with Schleck (K).

Local and vernacular names: Chinese fan palm, Biro (Japanese).

240 Gard. Bull. Singapore 60 (2) 2009

Distribution: Japan, China and Taiwan. In Japan, on the Ryukyu Islands of

Okinawa, Iriomote and Aharen, and on Kyushu, Aoshima (Yoshida ef al.,

2000) and Shikoku (Horikawa, 1972; Suzuki, 1982). In Taiwan on Kisanto

Is., and in China on Hainan Is. and formerly recorded in Guangdong

Province but now apparently extinct there in the wild (Fig. 8).

Habitat and ecology: Coastal forests on various soils, often in sand,

sometimes in dense mono-specific colonies, otherwise in small isolated

colonies, 1-100 m alt. Flowers Feb-Apr; fruits Apr-Sept.

Conservation status: Vulnerable.

Notes: Livistona chinensis was first described as Latania chinensis (Jacquin,

1801), from plants cultivated and subsequently naturalised in Mauritius

and brought to Schoenbrunn Gardens, Vienna in 1788. It is lectotypified

by the illustration in Jacquin (1801), Tab. 11, Fig. 1. The species name was

taken from that used for the palm in Mauritius, “Latanier de la Chine”.

Bretschneider (1898) provided some evidence to suggest that the naturalist

and traveller Pierre Poivre was responsible for introducing the palm, during

the mid 1700s, to Mauritius where it soon became naturalised. Poivre had

made extensive collections of plants from southeast China and Indochina

during the period 1740-1767. Martius (1838) provided the transfer to

Livistona, based on Brown (1810) who suggested it should correctly have

been in Livistona, but without formal transfer.

Other names have been applied to the species, including Latania

mauritiana, L. borbonica and Chamaerops biroo (Martius, 1838); Saribus

chinensis (Blume, 1838), the orthographic variation L. sinensis by Griffith

(1845); and L. japonica (Nakai, 1928). Of these names, a lectotype is

here chosen for C. biroo (Siebold, 1830), it being Siebold s.n., a specimen

that was located in the Siebold collections at Leiden. Types for Latania

mauritiana and Livistona japonica have not been designated, while Latania

borbonica 1s a misapplied name.

Livistona olivaeformis and L. subglobosa were established by

Hasskarl (1842) from cultivated plants in Bogor Botanic Gardens as species

of Saribus, but placed as Livistona species by Martius (1849). The species

names were derived from the fruit shapes, being olive-like and subglobose

respectively. Livistona olivaeformis was synonymised under L. chinensis by

Beccari (1921), and the specimens collected by Beccari in May 1878 from

plants in Bogor Botanic Gardens that Hasskarl based the name upon are

here chosen as the lectotype. Those specimens are sheets 11331, 11331-B.

and 11331-C in Fl. However, L. subglobosa was retained by Beccari as a

ee ee

A Taxonomic Account of Livistona R.Br. (Arecaceae) 24]

valid taxon and subsequently made a subspecies. Livistona chinensis var.

subglobosa was described as differing from his L. chinensis forma typica

only by fruit shape (Beccari, 1921), it being subglobose to ovoid rather than

olive-like. Although the original description of L. swhglobosa was from

cultivated plants in Bogor Botanic Garden, Beccari (1931) applied the name

to specimens collected in the Ryukyu Islands of southern Japan. Beccari

visited Bogor Botanic Gardens in May 1878 and collected specimens from

the plants upon which Hasskarl established the name. These specimens,

sheets 11333 and 11333-B in FI are here chosen as the lectotype. The taxon

was elevated to specific status by Nakai (1935) and Kanehira (1936), but

it otherwise was referred to most commonly as a subspecies (Moore &

Fosberg, 1956; Li, 1963; Ohwi, 1965; Walker, 1976; Li, 1978; Suzuk:,

1982; Yoshida ef a/., 2000). Collections from the distributional range of L.

chinensis var. subglobosa (cf. Horikawa, 1972) exhibit such variation in

fruit shape that no reasonable taxonomic distinctions can be made between

L. chinensis forma typica and L. chinensis var. subglobosa. Accordingly, L.

chinensis var. subglobosa is placed as a synonym of L. chinensis.

A third subspecies, L. chinensis var. boninensis, was also described

by Beccari (1921, 1931) and its taxonomic history is discussed in Notes

under L. boninensis.

Livistona chinensis 1s a moderate sub-canopy palm to 15 m tall;

leaves are large and regularly segmented; segment apices are pendulous,

and with a bifurcate cleft to 13% of the segment length; the inflorescence

is unbranched, not extending beyond the limit of the crown, and with up to

7 partial inflorescences; bracts are loosely sheathing; flowers are white to

yellow; fruit are globose, subglobose, ellipsoid, to pyriform, to 26 mm long

to 18 mm wide, and glossy blue green to bright green at maturity.

13. Livistona boninensis (Becc.) Nakai, J. Jap. Bot. 11 (1935) 222; Nakai,

Sci. World (Japan) (Rigakkai) 26(4) (1928) 10, nomen; Bull. Biogeogr. Soc.

Japan 1(3) (1930) 255, nomen. -Livistona chinensis var. boninensis Becc.,

Webbia 5 (1921) 12, 16; Ann. Roy. Bot. Gard, Calcutta 13 (1931) 62; Moore

& Fosberg, Gentes Herb. 8 (1956) 435, fig. 130C. — Type: Japan. Bonin

Islands, Haha-jima, 23 Apr 1917, Wilson 8271] (lectotype, A; isolectotypes,

BM, FI, K, US; fide Moore and Fosberg, 1956). Plate 10A-B.

-Corypha japonica Kittlitz, Veg.-Ansicht. 48 (1844) t.14-16, nom. inval.

(Nakai, 1936).

Hermaphroditic palm. Trunk to 20 m tall, ca 30 cm dbh, leaf scars raised,

with remnant leaf-base fibres, internodes narrow, petiole stubs not persistent.

Leaves 40-60 in a globose to flattened crown; petiole adaxially moderately

242 Gard. Bull. Singapore 60 (2) 2009

ridged, margins with single curved green spines well spaced in proximal

portion of petiole; leaf-base fibres not prominent, very coarse, persistent;

lamina costapalmate, regularly segmented, subcircular, 120-200 cm long,

adaxially grey-green, abaxially lighter grey-green, nonwaxy; lamina divided

for 45-75% of its length, with 50-82 segments, depth of apical cleft ca

20% of the segment length; apical lobes pendulous; parallel veins 5-6 each

side of midrib; transverse veins thinner than parallel veins. Inflorescences

unbranched at the base, 130-220 cm long, not extending beyond the limit of

the crown, branched to 3 orders; 6-7 partial inflorescences; rachillae 4-16

cm long, glabrous; prophyll not seen; peduncular bract 1, tubular, glabrous;

rachis bracts tubular, glabrous. Flowers in clusters of 5-8, 2.0-2.8 mm long,

cream. Fruit globose to pyriform, 19-30 mm long, 14-28 mm diam., glossy

bright green; pedicel 3-4 mm long. Seed reniform. Eophyll not seen.

Specimens examined: JAPAN: Bonin Islands, Haha-jima, Wilson 8271

(Lectotype, A; isolectotypes, BM, K, US); Haha-jima, Okimura, Furuse

_ 7719 (K); Haha-jima, Koshin-dzuka, top of Mt. Sekimon, 200-400 m alt.,

18 Mar 1972, Momiyama, Kobayashi & Ono s.n. (MAK); Chichi-jima, 75

m alt., Fosberg 31545, 31547, 31548 (US); Chichi-jima, Mt. Mikazuki,

Fujita & Shimizu 23 (A); Chichi-jima, vicinity of Tsutsujryama, Murata

171, with Tabata, Tsuchiya & Takada (A); Chichi-jima, Futami Graveyard,

3 Jul 1976, Kanai & Ono s.n. (MAK); Chichi-jima, Mt Mikadzuki, S slope,

250 m alt., Ono & Kobayashi 79009 (MAK); Chichi-jima, enroute from

the northern valley of Mt Akahatayama to Mt Tori-yama, 50-150 m alt.,

6 May 1974, Ono, Kobayashi & Wakabayashi s.n. (MAK); Mei-jima, 24

Aug 1980, Ono, Kobayashi, Sugawara & Sugawara s.n. (MAK); (Bonin

Islands), 1853-56, Wright s.n. (US).

Local and vernacular names: Bonin Island Fan-palm.

Habitat and ecology: Coastal and near coastal forest, 50-500 m alt.

Flowering Apr—May; fruiting Oct-Dec.

Conservation status: Near threatened.

Distribution: Japan. Bonin (Ogasawara) and Volcano Islands (Fig. 8).

Notes: The first mention of Livistona boninensis was by Kittlitz (1844) in

an account of his voyage through the northwest Pacific, where he noted

*... eine grosse schéne Facherpalme (Coryphajaponica?)...”,and included

it in various illustrations of the Bonin Islands. Nakai (1936) argued that

4 = + a +

A Taxonomic Account of Livistona R.Br. (Arecaceae) 243

the name Corypha japonica was invalid. The first formal recognition was

by Beccari (1921) who described it as a subspecies of L. chinensis in a

diagnostic key to Livistona species, as L. chinensis var. boninensis. The

first use of the name raised to specific rank as L. boninensis, was by Nakai

(1928) in his Plants in Ogasawara Islands. Beccari (1931) maintained it

as a subspecies of L. chinensis, and provided an expanded description.

However, Nakai (1935) continued using the name at specific level, and

placed Beccari’s subspecies in synonymy. Moore & Fosberg (1956) re-

established it as a subspecies, provided comparative illustrations with other

subspecies, and established the type as Wilson 8271.

Recently collected specimens seen at MAK carry larger fruits

than originally described, up to 30 mm long as opposed to 25 mm long

in previous descriptions. Fruit are globose to subglobose with a tapered

base. These differ from those in L. chinensis where they are only up to 26

mm long in the largest specimens seen, varying from globose to ovoid but

lacking any basal tapering. A peduncular bract is present in L. boninensis

but is lacking in L. chinensis. There is also considerable difference in the

degree of tomentum carried on the inflorescence bracts. In L. chinensis,

the tomentum is dense and persistent, while in L. boninensis bracts are

glabrous or only mildly tomentose becoming glabrous at maturity. Thus

the taxon formerly known as L. chinensis var. boninensis is hereby

reassigned to specific status in accordance with the taxonomy of Nakai

(1935).

14. Livistona endauensis J.Dransf. & K.M.Wong, Malayan Nat. J. 41

(1987) 121, fig. 1. — Type: Malaysia. Johore, Gn. Janing, Ulu Endau, 16

Jun 1977, Dransfield JD5089 (holotype, K; isotype, KEP). Plate 10C-E.

Hermaphroditic palm. Trunk to 15 m tall, 12-20 cm dbh, leaf scars

obscure, internodes narrow, pale grey brown, petiole stubs at first persistent

at the base only, but deciduous with age, vertical fissures shallow. Leaves

ca 75 in a globose to conical crown; petiole ca 175 cm long, arching,

orange to reddish-brown, ca 15 mm wide by 12 mm wide, triangular in

cross-section, adaxially concave, covered in thin, caducous, grey-brown

indumentum, margins unarmed in the distal half, armed in the proximal

half with single retrorsely recurved black spines, regularly spaced, 10-

14 mm long, 6-15 mm apart, distally becoming more widely separated;

leaf-base fibres prominent, coarse, disintegrating: distally the fibres form

a triangular, chocolate-brown appendage to 35 cm long, to 10 cm wide;

lamina costapalmate, regularly segmented, circular to subcircular in outline,

ca 100 cm long, slightly undulate, adaxially bright green, abaxially lighter

244 Gard. Bull. Singapore 60 (2) 2009

green, glabrous on both surfaces; lamina divided for ca 61% of its length,

with 60-70 segments, depth of apical cleft ca 8% of the segment length,

apical lobes rigid to occasionally pendulous through damage or age; mid-

leaf segments ca 3 cm wide where the segments diverge; outer segments

ca 0.8 cm wide where the segments diverge; parallel veins 4-5 each side of

mid rib; transverse veins thinner than parallel veins; hastula conspicuous,

ca 10 mm high, triangular, crescent shaped. Inflorescences unbranched at

the base, to 150 cm long, not extending beyond the limit of the crown,

arching, branched to 3 orders; partial inflorescences ca 8; peduncle ca 60

cm long, laterally compressed at the base, 3 cm wide by 0.5 cm thick, ca

2.5 cm wide distally by 1.0 cm thick; prophyll tubular, 30 cm long by 2.5

cm thick, dark brown, glabrous except for scattered caducous rust coloured

indumentum; peduncular bract(s) lacking; rachis bracts tubular, glabrous;

partial inflorescences to ca 35 cm long; rachillae 6-10 cm long, ca 0.25

cm diameter, rigid, yellow, pubescent. Flowers solitary or in clusters of

2-3, sessile, globular, ca 1 mm high, 1.5 mm wide, golden yellow; calyx

- basally tubular, ca 0.3 mm high, glabrous, 3-lobed, lobes triangular to 0.4

mm long by | mm wide; petals broadly triangular, to 1 mm long by 0.8 mm

wide, glabrous, abaxially rugulate, adaxially striate; stamens basally fused

to form a tube ca 0.4 mm high; filaments very short, free; anthers ca 0.1 mm

diam.; carpels ca 0.6 mm high, style to ca 0.2 mm high, stigma punctiform.

Fruit obovoid to pyriform, ca 16 mm long, 14 mm wide, bluish green;

epicarp smooth, wrinkled on drying; mesocarp ca 2 mm wide; endocarp

crustaceous ca | mm wide; pedicel 1-2 mm long. Seed globose, ca 1 cm

diam., intruded shallowly by the testa. Eophyll 5-ribbed.

Specimens examined: MALAY SIA: Trengganu. Kuala Dungun, Bukit Bauk,

350 m alt., Dransfield 915 (K); Dungun, Bukit Bauk, ridge top, Dransfield

5184 (K); Dungun Bukit Bauk, Kamarudin 31445 (K); Kemaman, Compt.

4, Rasau Kerteh Ulu Chukai Forest Reserve, Meijer & Yong 94940 (K,

KEP, L, SING); Pahang, Lesong Forest Reserve, Ulu Sg. Teroh, Wong

32280 (K); location not given, ‘ridge-top vegetation’, Saw 34174 (K);

Johore. Kuala Jasin, S. Endau, 300 ft (c. 90 m) alt., 12 Nov 1972, Heaslett

& Ahmad Shukor s.n. (SING); Gunung Janing, Ulu Endau, Dransfield

JD5089 (K holotype, KEP); Gunung Janing via Kahang, 350 ft (ca 110 m)

alt., Heaslett, Shah, Samsuri & Shukor 2609 (SING); Gunung Janing Barat,

steep slope near summit, Kiew 19/2 (K); Hutan Simpan Labis, sandstone

plateau S of Gunung Janing, Wong 32461 (K).

Habitat and ecology: \n rainforest at 90-660 m alt. Flowers all year; fruits

all year.

ee eee

A Taxonomic Account of Livistona R.Br. (Arecaceae) 245

Conservation status: Lower risk, near threatened (IUCN, 2006).

Distribution: Malaysia. In Johore at Ulu Endau, on Gunung Janing, Gunung

Janing Barat and Bukit Peta, and the eastern hills of Trengganu at Bukit

Bauk (Fig. 9).

90°E 100°E 110°E 120°E 130°E

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Figure 9. Distribution of Livistona endauensis (Q), L. tahanensis (@), L. jenkinsiana (m),

and L. speciosa (©).

Notes: Livistona endauensis was described by Dransfield and Wong (1987)

from a population on Gunung Janing in the Ulu Endau area, Johore, Malaysia,

based on the collection Dransfield JD5089. In the protologue, a population

of a then unidentified Livistona in Trengganu was compared to this taxon,

though maintained as distinct pending further study. Subsequent examination

of collections from Trengganu confirmed it as the same taxon, and therefore

giving the species a disjunct range in eastern Peninsular Malaysia.

Livistona endauensis is a moderate sub-canopy palm to 15 m

tall; leaves are large and regularly segmented; segment apices are rigid

to pendulous, and with a bifurcate cleft to 8% of the segment length; the

inflorescence is unbranched, not extending beyond the limit of the crown,

246 Gard. Bull. Singapore 60 (2) 2009

and with about 8 partial inflorescences; bracts are loosely tubular; flowers

are golden yellow; fruit are obovoid to pyriform, to 16mm long to 14 mm

wide, and bluish green at maturity.

15. Livistona tahanensis Ridl. ex Becc., Webbia 5 (1921) 17; in Ridley, J.

Fed. Malay States Mus. 6 (1915) 189, nomen. — Type: Malaysia. Pahang,

Gunung Tahan, | Jun 1905, Wray & Robinson 5355 (holotype, BM; isotype,

FI, SING). Plate 11A.

Hermaphroditic palm. Trunk to 8 m tall, ca 12 cm dbh, leaf scars narrow,

internodes narrow, petiole stubs persistent for most of its length. Leaves

30-40 in a+ globose crown; petiole to ca 70 cm long, 12-15 mm wide in the

middle, 5-6 mm wide distally, adaxially flat, with scattered appressed hyaline

ciliate-margined scales, margins with short, flat, brown, blunt triangular

5-8 mm long spines throughout, distally reducing in size; leaf-base fibres

prominent, fine, disintegrating; appendage very long, membranous, bright

“mahogany red, polished on both surfaces; lamina costapalmate, regularly

segmented, subcircular, 57-76 cm long, ca 90 cm wide, rigid, flat to undulate,

adaxially dark green, abaxially grey-green; lamina divided for ca 58% of

its length, with 40-50 segments, depth of apical cleft ca 16% of the segment

length, apical lobes rigid; parallel veins 6-7 each side of midrib; transverse

veins thinner than parallel veins. Inflorescences unbranched at the base,

80-91 cm long, not extending beyond the limit of the crown, branched to 3

orders; partial inflorescences ca 4, longest to ca 23 cm long; prophyll not

seen; peduncular bracts lacking; rachis bracts loosely tubular; peduncle 2.5

cm wide at the base; rachillae 7-10 cm long, thin, green-red, tomentose.

Flowers very small; sepals ovate, cream tipped red; petals oblong, blunt, tip

thickened, incurved, cream; anthers white; style short, conic. Fruit globose

to depressed-globose, 12-14 mm long, glossy green; epicarp smooth; suture

line extends for the full length of the fruit, marked with lip-like structures;

pedicel 2-3 mm long. Seed not seen. Eophyll not seen.

Specimens examined: MALAYSIA: Pahang. Gunung Tahan, Wray &

Robinson 5355 (BM holotype, FI); Gunung Tahan, 3500-4000 ft alt., Nur

8006 (SING); Gunung Tahan, 3500-4500 ft alt., Holttum 20631 (BO,

K, SING, UC); Gunung Tahan, 3000 ft., Corner 36597 (BO, K, SING);

Gunung Tahan, 3500 ft, Jan 1923, Pendlebury s.n. (K); Gunung Tahan,

3500 ft, Jan 1923, Pendlebury & Kloss s.n. (BM); Gunung Tahan, Tahan

Woods near Wrays Camp, Dransfield 659 (K).

Local and vernacular names: Tahan Serdang, Daun Tau.

. |

oe a ie

4 Taxonomic Account of Livistona R.Br. (Arecaceae) 24]

Habitat and ecology: In moist montane forest, at 900-1500 m alt. Wild

elephants are reported to eat the cabbage (Kiew and = ison, 1989).

Conservation status: Lower risk, conservation dependent (IUCN, 2006).

Distribution: Malaysia. In Pahang, endemic to Gunung Tahan (Fig. 9

Notes: The name Livistona tahanensis was first used by Ridley (1915) in

an annotated list of plants of Gunung Tahan, Pahang, Malaysia. There was

no description provided but an indication that the species was pending

description by Beccari. Beccari (1921) provided the first formal description,

based on the collection Wray & Robinson 5355 from 1000 m on Gunung

Tahan.

Livistona tahanensis is a moderate canopy palm to 8 m tall, with

persistent petiole stubs on most of the stem: leaves are moderate and

regularly segmented; segment apices are rigid, and with a bifurcate cleft to

16% of the segment length: the inflorescence is unbranched, not extending

beyond the limit of the crown, and with about 4 partial inflorescences:

bracts are loosely tubular; flowers are cream with red tipped sepals; fruit

are globose to depressed globose, to 14 mm long to 12 mm wide, and glossy

green at maturity.

16. Livistona jenkinsiana Griff., Calcutta J. Nat. Hist. 5 (1845) 334; Palms

Brit. E. Ind. (1850) 128, figs 226A, 226B; Anderson, Journ. Linn. Soc. 11

(1871) 13. -Livistona jenkinsii Griff. ex Mart., Hist. Nat. Palm. 3 (1849)

242, orth. var. -Latania jenkinsiana (Griff.) Devansaye, Rev. Hort. 47

(1875) 34. -Saribus jenkinsii (Griff.) Kuntze, Revis. Gen. Pl. 2 (1891) 73

— Type: India. Assam, Nowgong, 1842, Jenkins s.n. (holotype, BR). Plate

11B-F.

Hermaphroditic palm. Trunk to 10 m tall, 15-23 cm dbh, leaf scars

prominent, narrow, roughened, light coloured, internodes narrow, dark

coloured, petiole bases persistent in the lower portion. Leaves 20-50 in a+

globose crown; petiole 130-200 cm long, 20-25 mm wide distally, adaxially

slightly concave, margins armed throughout with single or double retrorsely

recurved reddish to brown spines 15-20 mm long, 10-12 mm wide at the

base, base frequently swollen, spines reducing in size to tubercles toward

the apex; leaf-base fibres moderately prominent, coarse, disintegrating

or persistent; appendage to 25 cm long; lamina costapalmate, regularly

segmented, ovate-reniform in outline, 105-130 cm long, 150-200 cm wide,

adaxially shiny green, abaxially subglaucescent grey or bluish; lamina

248 Gard. Bull. Singapore 60 (2) 2009

divided for 72-75% of its length, with 70-80 segments, depth of apical cleft

8-16% of the segment length, 3-8 cm wide where the segments diverge,

apical lobes rigid; parallel veins 9-10 each side of midrib; transverse veins

thinner than parallel veins; hastula cordate. Inflorescences unbranched at

the base, 60-100 cm long, not extending beyond the limit of the crown,

branched to 3 orders; partial inflorescences 3-6; prophyll 30-45 cm long,

woody, keeled; peduncular bract(s) lacking; rachis bracts loosely tubular,

reddish-brown, glabrous, expanded distally into lanceolate acuminate

lobes, sometimes longitudinally split, scurfy to glabrous; rachillae 10-30

cm long, 3-4 mm wide, rigid, yellow-green, puberulous. Flowers in clusters

of 3-5, sessile, greenish cream, with inconspicuous bracteoles; sepals fused

basally, fleshy, 1.2-4 mm long, |.3-1.5 mm wide, lobes broadly ovate, with

thin and subhyaline margins; petals basally fused, deltoid, acute, 2.5-4 mm

long, 2.5-3 mm wide; stamens with basally fused filaments, thick and short,

contracted into elongate apices, connective very short, narrow; carpel

turbinate-obconical, yellow, distinctly sculptured, and contracted into a

short trisulcate, filiform style, stigma simple. Fruit globose to reniform, 19-

28 mm long, 16-25 mm diam., somewhat asymmetrical, apically rounded,

slightly tapered below to an acute base, leaden blue to dark bluish-purple;

epicarp very thin, with scattered lenticellular pores; suture line extends full

length of the fruit, marked with lip-like structures; mesocarp succulent,

moderately fibrous or lacking fibres; endocarp woody, brittle, cinnamon

brown inside, 0.5-1 mm wide; pedicel 3-6 mm long, 2.5-4 mm wide. Seed

globose, 17-20 mm diam., slightly flattened on one side; intruded broadly

and deeply by the testa; embryo sublateral. Eophyll 7-ribbed.

Specimens examined: INDIA: Assam. Nowgong, Jenkins s.n. (BR holotype);

Valley of the Mali Itka, Masum, PutooRd, 1000-2500 m alt., Kingdon-Ward

1288 (BM); Assam. Griffith 1205 (BM). CHINA: Yunnan. You-louh Shan,

Che-li Hsien, Wang 78114 (A); THAILAND: Mae Hong Son, Doi Nam

Bon Musir, Smith 18 (K); Mae Hong Son, Nai Sai Village, Smith 2] (K);

Chiang Mai, Doi Suthep, ca 1200 m alt., Kerr 3430 (AAU, K, NY); Chiang

Mai, Doi Suthep, ca 1500 m alt., Kerr 4686 (AAU, BK, BM, K, NY);

Chiang Mai, Suthep, Hue Me Pan, Doi Suthep, Rock 437 (K); Phetchabum,

Muanga Lom, Kao Keo Ngoi, ca 900 m alt., Kerr 576] (AAU, BK, BM,

NY); Chatyaphum, Pu Kio, ca 1000 m alt., Kerr 20241 (AAU, BK, BM,

NY); Nakhon Ratchasima, Khao Yai NP, near headquarters, Whitmore 3172

with Chamlong (K); Nakorn Ratchasima, Khao Yai NP, Phatabak, Smith 3

(K); Ranong, La-un, Kerr 16472 (AAU, BK, K); Suratthani, Kaw Tao, ca

200 m alt., Kerr 16033 (AAU, BK, K, BM); Suratthani, Ban Ta Khun,

Bang Chang, Klong Saeng Wildlife Sanctuary, Smith 66 with Sumawong

A Taxonomic Account of Livistona R.Br. (Arecaceae) 249

(K); Pattani, Kao Kalakiri, ca 900 m alt., Kerr 779] (AAU, BK, BM, K).

Specimens from cultivated material: (NDONESIA: Bogor Botanic Gardens,

May 1878, Beccari s.n. (Fl); SINGAPORE: Singapore Botanic Gardens,

Apr—May 1936, C.X. Furtado 31108 (L).

Distribution: India, Sikkim, Bangladesh, Myanmar, China and Thailand. In

northeast India in the Naga Hills, Khasia Hills, Jainita Hills, Nowgong,

Darjeerling Hills; in Siang, Lohit and Tirap Valleys in Arunachal Pradesh;

and Gubro Purbat in Assam. In Sikkim in the Teesta Valley; Bangladesh

in Chittagong; China, in Yunnan; and in northern and peninsular Thailand

(Fig. 9).

Habitat and ecology: \n high rainfall areas in moist evergreen forest mostly

on sandy loam with a laterite mixture, at 100-2500 m altitude. Datta and

Rawat (2003) observed foraging of mature fruit by Hornbills in northeast

India. Flowers Jul-Aug; fruits Oct— Nov.

Ethnobotany: Leaves are used for thatch and hats (Lepchas tribe of north-

east India and Sikkim). The endosperm, presumably prior to maturity, is

reported to be consumed.

Conservation status: Near threatened.

Notes: Livistona jenkinsiana was first described by Griffith (1845), based

on observations at Gubro and a collection made in 1836 from Assam by the

British Commissioner of Assam, Major-General F. Jenkins (1793-1866),

and named in his honour. The type is in BR. Martius (1849) published

the orthographic variation L. jenkinsii and provided a description of the

inflorescence and flowers based on information given to him by Griffith.

Griffith (1850) provided an expanded description, with illustrations (Plates

226A and 226B) of a leaf, a partial inflorescence, flowers and fruit.

Livistona jenkinsiana is a moderate canopy palm to 10 m tall;

leaves are large and regularly segmented; segment apices are rigid, and

with a bifurcate cleft to 16% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to

6 partial inflorescences; bracts are loosely tubular; flowers are greenish

cream; fruit are globose to reniform, to 28 mm long to 25 mm wide, and

leaden blue to dark bluish purple at maturity.

17. Livistona speciosa Kurz, J. Asiat. Soc. Bengal 43(2) (1874) 204, pl. 13

250 Gard. Bull. Singapore 60 (2) 2009

& 14. -Saribus speciosus (Kurz) Kuntze, Revis. Gen. Pl. 2 (1891) 736. —

Type: Burma (Myanmar). Pegu Yomas, Chounmenahchy, Feb 1871, Kurz

3330 & 3331 (holotype, B, destroyed; isotypes, BM, K). Plate 12A-D.

-Livistona fengkaiensis X.W.Wei & M.Y.Xiao, J.S. China Agric. Coll. 8(1)

(1982) 22, figs. | & 2. — Type: China. Guangdong, Fengkai, Heishi Ding,

450 m alt., 6 Dec 1981, Fengkai Exped. 1587 (holotype, CANT).

Hermaphroditic palm. Trunk to 25 m tall, 20-30 cm dbh, leaf scars lightly

raised, light grey, petiole stubs sometimes persistent, particularly at the

base, otherwise smooth, longitudinal fissures not evident. Leaves 40-50 in

a globose or oblong crown; petiole slightly arching, 140-159 cm long, 20-

25 mm wide distally, adaxially slightly concave to flat, abaxially convex,

margins armed throughout with single retrorsely recurved orange-brown

spines to 25 mm long, to 10 mm wide at the base, base frequently swollen,

apically acute, spines reducing in size to tubercles toward the apex; leaf-

base fibres moderately prominent, coarse, persistent; appendage to 25 cm

long; lamina costapalmate, regularly segmented, circular to subcircular

in outline, 150-200 cm long, to 200 cm wide, adaxially green, abaxially

glaucous grey; lamina divided for 25-75% of its length, with ca 100

segments, depth of apical cleft 3-4% of the segment length, 3.5-8 cm wide

where the segments diverge, apical lobes rigid; basal segments overlap

at the hastula; parallel veins 9-10 each side of midrib; transverse veins

thinner than parallel veins; hastula cordate. Inflorescences unbranched at

the base, 120-200 cm long, not extending beyond the limit of the crown,

branched to 4 orders; partial inflorescences 3-6, 40-60 cm long; prophyll

30-45 cm long, woody, keeled; peduncular bract(s) lacking; rachis bracts

loosely tubular, reddish-brown, glabrous, expanded distally into lanceolate

acuminate lobes, sometimes longitudinally split, scurfy to glabrous;

rachillae 10-20 cm long, 3-4 mm diameter, spreading to drooping or rigid,

yellow-green, glabrous. Flowers in clusters of 5-6, sessile, greenish cream,

carried on small, sessile tubercles, broadly ovate in bud, to 2.5 mm long;

sepals fused basally, fleshy, 1.2-2 mm long, 1.3-1.5 mm wide, semi-ovate,

subacute, with subhyaline margins; petals basally fused, thick, deltoid,

acute, 2.5-4 mm long, 2.5-3 mm wide; stamens with briefly basally fused

filaments, partly adnate to the petals, thick and short, contracted into

elongate apices, connective very short, narrow; anthers suboribicular-

didymous; carpel turbinate-cbconical, yellow, distinctly sculptured, and

contracted into a short trisulcate, filiform style to 2.5 mm long, stigma

simple. Fruit obovate, obpyriform, to ovoid, rounded apically, narrowed

basally, 25-35 mm long, 18-25 mm diam., greenish-blue to light-blue at

maturity; epicarp less than | mm wide, with scattered lenticellular pores;

. |

_—

A Taxonomic Account of Livistona R.Br. (Arecaceae)

suture line extends full length of the fruit, marked with lip-like structures:

mesocarp thinly woody, brittle, whitish within; endocarp woody, brittle, to

1 mm wide. Seed oblong-elliptical, rounded at both ends, to 17 mm long,

to 12 mm wide; intruded to more than half the width of the endosperm:

embryo lateral. Eophyll 7- ribbed.

Specimens examined: MYANMAR: Pegu Yomas, Chounmenahchy, Kurz

3330 & 3331 (BM, K): Tharapon, Meebold 14335, 14423 (M). CHINA:

Guangdong Prov., Fengkai, Heishi Ding, 450 m alt., Fengkai Exped.

1587 (CANT); Hainan. Liang 36627 (NY); Hainan. Mo-San-Leng, 3000

ft alt., Chun & Tso 44362 (A, K, NY, US). THAILAND: Chiang Mai.

Muang, Doi Suthep, E side above Rusee Cave, Song See 86 (PHCMU):

Chiang Rai. Ban Thing Sai, 8 km NW of Chiang Khong, Anderson 6042

(PHCMU); Lampoon. Mae Tah, Doi Kuhn Dahn NP, southside, Doi Hou

Chang, Maxwell 94978 (BKF, CMU); Loei, Phu Kradang, 16°53°54°N,

101°47°49”°E, 1150-1250 m alt., Murata 49918 with Phengklai, Mitsuta,

Yahara, Nagamusa & Natasan (A, BKF, L); Khao Kho, 600-700 m alt.,

Dowe 560 (JCT); Kho Yai NP, 20 km S of Main Rd, Dowe 565 (JCT):

45 km N of Thong Pha Phum, Goengavia village, Dowe 3566 (JCT):

Ranong Prov. 37 km NE of Ranong, less than 200 m alt., King 5586

(US); Petchaburi Prov., Kaeng Krachen NP, 12°54°N, 99°20°E, 500-700

m alt., Barfod 45194 with Ueacharakan (AAU); Petchburi Prov., Kaeng

Krachen NP, 2 km after Panoentung Camp, westward on Khao Panoen

Rd, Damborg 2 (AAU); Chumphon, Dowe 569 (JCT); Songkla, 1 km SW

of Songkhla, Ton Nga Waterfall, 20 km W of Hat Yai, 07°00°N, 100°00’E,

100-250 m alt., Barfod 141040 with Ueacharakan (AAU, K); Songkhla,

Hat Yai, Nathawi Rd, Whitmore 3100 with Charal & Chamlong (BKF, K):

Trang. Khao Chong, ca 15km E of Trang, 07°30°N, 99°45°E, 200 m alt.,

Barfod 141333 with Ueacharakan (AAU, PSU); Trang, Kantang, Kuan

Kantrang Reserved Forest, Luang Samarn 64 (BKF); Yala. Village of

Sungai Kolok, near Sirindhorn Waterfall at Ban Bala, 6 km N of turnoff,

Damborg 3 & 4 (AAU). MALAYSIA: Perak. Ulu Batang Padang, on

Pahang boundary, ca 6000 ft alt., Henderson 23665 (BO, BRI, K, SING,

NY).

Distribution: Bangladesh, Chittagong; Burma, Pegu Yoma and Tenasserim;

China, Guangdong, Fengkai, and Hainan Is. at Mo San; throughout Thailand

at 400-900 m alt.; and Peninsula Malaysia, on Langkawi Is., Gunung Inas

to Genting Sempah in the west and Gunung Stong and Gunung Mandi

Angin in the east at 700-1200 m alt. Moderately common in moist to wet

forest (Fig. 9).

DISD, Gard. Bull. Singapore 60 (2) 2009

Habitat and ecology: \n high rainfall areas in moist evergreen forest, at

200-3000 m altitude. Flowers Jul—Aug; fruits Oct— Nov.

Conservation status: Near threatened.

Notes: Livistona speciosa was described by Kurz (1874) from the mountains

of Pegu Yoma of central Myanmar, based on Kurz 3331. Although Kurz did

not specifically record himself as the collector of the specimen from Pegu

Yoma, it is hereby accepted as the holotype as reference to the Pegu Yoma

collection is sufficient to confirm the connection. This taxon’s similarity to L.

Jenkinsiana was noted by Kurz in the protologue, although Beccari (1931),

still noting a close relationship between the taxa, provided descriptions of

some characters that he considered readily separated the species.

Livistona fengkaiensis was the name given to a population occurring

in the mountains of the Fengkai area in Guandong, southern China, based

on the collection Fengkai Exped. 1587.

Livistona speciosa is a large canopy palm to 25 m tall; leaves

are large and regularly segmented; segment apices are rigid, and with a

bifurcate cleft to 4% of the segment length; the inflorescence is unbranched,

not extending beyond the limit of the crown, and with up to 6 partial

inflorescences; bracts are loosely tubular; flowers are greenish cream; fruit

are obovate, obpyriform to ovoid, to 35 mm long to 25 mm wide, and

greenish blue to light blue at maturity.

18. Livistona carinensis (Chiov.) J.Dransf. & N.W.Uhl, Kew Bull. 38 (1983)

200. -Hyphaene carinensis Chiov., Fl. Somala | (1929) 319. -Wissmannia

carinensis (Chiov.) Burret, Bot. Jahrb. Syst. 73 (1943) 184; Monod, Bull.

de VI.F.A.N. 17 (1955) 343, figs 3-23. — Type: Somalia. Carin, Oasis of

Uncud, 6 July 1924, Puccioni & Stefanini 1027 (holotype, FT). Plate

13A-D.

Hermaphroditic palm. Trunk to 40 m tall, ca 40 cm dbh, leaf scars

prominent, slightly oblique, irregular in width, internodes narrow, petiole

stubs persistent in the lower 1-2 m. Leaves 30-40 in a globose to conical

crown; petiole to 125 cm long, 5-8 cm wide proximally, 1-2 cm wide distally,

adaxially flat to shallowly concave, bright orange-yellow-green, margins

armed with large single or double, dark brown curved spines throughout, but

largest and closer in the proximal portion; spines conical, basally swollen,

apically acute, retrorsely recurved, brown-black, 7-25 mm long, 3-10 mm

wide at the base, 8-15 mm apart, reduced to tubercles in the distal portion

of the petiole, 4-6 cm apart; leaf-base fibres prominent, fine, persistent;

ee ee eee

A Taxonomic Account of Livistona R.Br. (Arecaceae) 3

appendage rigid, 6-7 cm long, brown to black; lamina costapalmate,

regularly segmented, subcircular in outline, 80-95 cm long, thick, both

surfaces waxy, glabrous, grey-green on adaxial surface, grey abaxially,

drying chartaceous; lamina divided for 75-85% of its length, with 50-

70 segments, depth of apical cleft 40-50% of the segment length, apical

lobes rigid, segment midrib very prominent; parallel veins 18-20 each

side of midrib; transverse veins thinner than parallel veins, inconspicuous

on the surface; segment margins thickened, with a deciduous filament,

the remains of which persist where the segments diverge from adjacent

segments; lamina anatomically isolateral, hypodermis |-layered below

each surface (not with a 2—layered adaxial hypodermis as in other species,

cf. Tomlinson, 1961). Inflorescences unbranched at the base, 200-240

cm long, extending beyond the limit of the crown by ca 20 cm, slender,

arching, eventually pendulous, branched to 3 orders; partial inflorescences

6-12; rachillae very thin, yellowish, with scattered long hairs; peduncular

bract 1, glabrous; rachis bracts tubular, brown red, striate, glabrous.

Flowers in clusters of ca 5, ca 2 mm long, yellow-green, abaxial surface

of perianth segments with scattered long hairs; sepals much shorter than

the petals, irregular, margins hyaline, scattered long hairs on the abaxial

surface near the apex; petals apically pointed, scattered long hairs near

the apex; filaments basally connate; carpels scarcely fused, similarly the

styles. Fruit globose, 5-20 (50) mm diam., dark brown to black; epicarp

thin, dull, shallowly rugose in the fresh state, deeply rugose in the dried

state; stigmatic remains apical; suture line extends for the length of the

fruit; mesocarp greenish with very large sclerenchymatous cells; mesocarp

very thin, adhering to the endocarp; pedicel narrow, 4-5 mm long. Seed

globose; intruded by the seed coat to displace most of the endosperm;

embryo sublateral. Eophyll 7-ribbed.

Specimens examined: SOMALIA: Carin. Oasis of Uncud, Puccioni &

Stefanini 1027 (FT holotype); Galgala, 10°58’°N, 49°02°E, Barbier 972

(K); Galgala, 25 km W of Carin, 10°59’N, 49°02’E, Lavranos & Carter

24835 (K). DJIBOUTI: Bankouale Wadi, Nov. 1985, Coghlan s.n. (K);

Bankouale, 2000, Welch s.n. with Welch (K).

Local and vernacular names: Bankouale Palm, Nakilto (Afar), Daban,

Madah (Somali), N,tug, somm (Arabic).

Habitat and ecology: Grows in or adjacent to intermittently flowing streams

or soaks in valley bottoms, but never fully dry soils, in semi-arid regions

with rainfall less than 400 mm per annum, at 200-975 m alt.

254 Gard. Bull. Singapore 60 (2) 2009

Conservation status: Vulnerable (IUCN, 2006), but here upgraded to

Endangered based on Welch & Welch (1999) and Ford & Bealy (2004).

Distribution: Djibouti, Somalia and Yemen. In the Horn of Africa, in

Djibouti in the Goda Mts, and in Somalia at Carin, Uncad, Galgala, Marajo,

Duud Shabeel and Xamur. In Yemen in the Hadramaut region at El Mintaq

and Wadi Hadjer (Fig. 10).

Legend

s L. carinensis

30°E 40°E 50°E 60°E

Figure 10. Distribution of Livistona carinensis (@).

Notes: The geographic isolation of Livistona carinensis, some 4000 km

from the nearest Livistona species in northeast India, introduces interesting

questions with respect to historical distribution and phylogeny. Based on

morphology it is similar to species in western and northwestern Australia,

but this appears to be an example of ecological convergence. Recent

molecular analysis (Isagi ef a/., in prep.) places it close to L. jenkinsiana.

Livistona carinensis was first described by Chiovenda (1929) as a

Borassoid palm and named Hyphaene carinensis with the specific name

derived from the site of its collection in the Carin region of Somalia.

The type specimen, Puccioni & Stefanini 1027, was collected in 1924.

ee ee ee

‘nN

A Taxonomic Account of Livistona R.Br. (Arecaceae)

The original description was based solely on leaf material as flowers and

fruit were not collected. Subsequently, the generic determination was not

soundly established, and the species true affinities were open to question.

Chevalier (1939) suggested that it was a species of Medemia: “... peut-

étre M. argun P.G. von Wurtemberg connu seulement en Nubie...”, a genus

from northeast Africa also in the Borassoid group of palms. However, it was

the German palm specialist, Burret (1943), in his treatment of the Arabian

palms, who correctly recognised it as a Coryphoid palm and assigned it

to a new genus, Wissmannia, named after the discoverer of the palm in

Yemen, H. von Wissmann, and thus established the name Wissmannia

carinensis (Chiov.) Burret. Burret only had leaves at his disposal, so there

still remained questions about the affinities of the species.

The first complete description, including flowers and fruit, was

provided by Monod (1955) based on specimens collected in Djibouti by E.

Chédeville. Monod noted the similarity of its flowers and fruit to Livistona,

but did not introduce any taxonomic changes and maintained the species

in Wissmannia. Tomlinson (1961), in a study based on leaf anatomy,

provided data that suggested that there was no single character that could

be used to separate Wissmannia from Livistona, but that by invoking a suite

of characters, all of which were otherwise shared with certain species of

Livistona, \t was a distinct genus. Moore (1973, 1977) and Langlois (1976),

similarly, indicated a close relationship with Livistona but maintained

taxonomic distinction. Based on this evidence and with further study,

Dransfield and Uhl (1983a) provided the formal transfer of Wissmannia to

Livistona, and established the name Livistona carinensis.

Livistona carinensis is listed as a critically endangered species.

Reports on the palm’s ecological status have been prepared by Welch

and Welch (1998, 1999) and Ford and Bealy (2004). Surveys in Djibouti,

Somalia and Yemen, resulted in the location of all adult palms, and with a

total of less than 1800 palms. Compared to reports on population numbers

taken 10-20 years previously, there had been a decline in numbers of 23-59%

across all locations. The primary causes of such rapid decline are the cutting

down of the palms for timber, or the clearance of land for agriculture. There

is no active program to conserve this species in either Somalia or Yemen,

but the Government of Djibouti has developed a conservation plan.

Livistona carinensis is a large canopy palm to 40 m tall; leaves

are large and regularly segmented; segment apices are rigid, and with a

bifurcate cleft to 50% of the segment length; the petiole is armed with very

large spines; the inflorescence is unbranched, extending beyond the limit of

the crown and pendulous, and with up to 12 partial inflorescences; bracts

are tubular; flowers are yellowish green with long unbranched hairs on

256 Gard. Bull. Singapore 60 (2) 2009

the sepals and petals; the rachillae are also covered with long unbranched

hairs; fruit are globose, to 5-20 (50) mm diameter, and dark brown to black

at maturity.

19. Livistona humilis R.Br., Prodr. (1810) 268. -Saribus humilis (R.Br.)

Kuntze, Revis. Gen. PI. 2 (1891) 736. — Type: Australia. Northern Territory,

Gulf of Carpentaria, Blue Mud Bay, Morgans Is (Brown’s island ‘s’), 20

Jan 1803, Brown (Bennett no. 5796) (lectotype, BM; isolectotypes, FI, K).

Plate 14A-D.

-Livistona leichhardtii F.Muell. ex F.Muell., Fragm. 8 (1874) 221; Fragm.

5 (1865) 49, nomen. — Type: Australia. Northern Territory, MacAdam Ra.,

1855, Mueller s.n. (lectotype, MEL) (fide Rodd, 1998).

-Livistona humilis var. minutiflora Becc., Ann. Roy. Bot. Gard. (Calcutta) 13

(1931) pl. 7, fig. 3a, nom. inval. Corrected to L. humilis R.Br. in Errata.

Dioecious palm. Trunk to 7 m tall, 5-8 cm dbh, leaf scars raised, roughened

and with remnant tissue, internodes narrow, grey, petiole stubs persistent

or deciduous with extreme age or fire. Leaves 8-15 in a globose crown;

petiole 40-70 cm long, 6-14 mm wide, margins with small, single, curved

dark red spines; leaf-base fibres not prominent, coarse, persistent; lamina

costapalmate, regularly segmented, circular, 30-50 cm long, chartaceous,

adaxially dark green, abaxially lighter green, glossy; lamina divided

for 60-87% of its length, with 30-44 segments, depth of apical cleft

35-89% of the segment length, apical lobes acuminate, rigid; parallel

veins 6 each side of midrib; transverse veins thinner than parallel veins.

Inflorescences unbranched at the base, sexually dimorphic, vertically

erect in both sexes, extending well beyond the limit of the crown; those

on fruiting plants (functionally female) straight, to 230 cm long, branched

to 3 orders, with a single terminal partial inflorescence; peduncular bracts

5-8, scurfy pubescent; inflorescences on nonfruiting plants (functionally

male) arcuate, to 180 cm long, branched to 3 orders; peduncular bract(s)

lacking; partial inflorescences 4-7; rachis bracts are tightly tubular, scurfy

pubescent; rachillae 3-12 cm long, pubescent. Flowers, male and female

similar in gross morphology, in clusters of 2-4, globose, 1.5-1.8 mm long,

yellow; sepals broadly ovate, 1-1.3 mm long, membranous, cuspidate;

petals broadly ovate, 1.5-1.8 mm long, fleshy, acute; stamens ca 1.3 mm

long. Fruit ellipsoidal or pyriform to obovoid, 11-19 mm long, 8-10 mm

diam., shiny purple-black; epicarp with scattered lenticellular pores; suture

line extends for full length of the fruit, marked with lip-like structures;

mesocarp fleshy; endocarp thin; pedicel 0.5-1 mm long. Seed ellipsoid, 7-9

mm long. Eophyll 3-ribbed.

ey mien smn sc RO ES 5

A Taxonomic Account of Livistona R.Br. (Arecaceae) 257

Specimens examined: AUSTRALIA: Northern Territory. Gulf of

Carpentaria, Blue Mud Bay, Morgans Is, 20 Jan1803, Brown (Bennett no.

5796) (BM lectotype, FI, K); Melville Is, Jul-Aug 1911, Spencer s.n. (MEL,

NSW); Coburg Peninsula, 13.5 miles S of Danger Point, Chippendale 8220

(MEL); ca 2 miles NW of Caiman Ck, 11°12’S, 132°12’E, Byrnes 1014

with Maconochie (DNA, K, LAE, NSW, PERTH); 19 miles NW of Mount

Norris Bay, Chippendale 8164 (BRI); Port Denier, Hill 47 (K); Wessell

Islands, Latz 3470 (CANB, DNA); Elcho Is, Maconochie 2179 (CANB,

DNA, K, NSW); Arnhem Land, 10.5 km NNE of Murgenella, 11°26’S,

132°58°E, Ross 3306 (MEL); 160 km from Pine Ck, Oenpelli Rd, Gittens

2646 (BRI): Arnhem Land, Schultz 7 (MEL); Arnhem Land, undated (ex

Herbarium Hookerianum 1867), Mueller s.n. (K); MacAdam Ra., 1855,

Mueller s.n. (MEL); 5 km S of Gove Airport, 12°19°S, 136°48°E, Eurell

78714 (L); 21 km from Gove Airport on Katherine Rd, Forster PIF6028

(BRI); Gove Peninsula, half way between Gove Airport and Nhulanbuy,

Rodd 2926, 2927 & 2928 (CANB, DNA, K, NSW); Yirrkala, 12°12’S,

136°47°E, Specht 901 (A, BRI, K, L, MEL, PERTH, US); Munmarlary Stn..,

Latz 3729 (BRI, CANB, DNA, K); 36.3 miles N of Oenpelli, Chippendale

NT8&108 (DNA, NSW); Oenpelli, 12°18’S, 133°40’E, Specht 1221 (AD, BRI,

K, L, LAE, US); ca 80 km E of Oenpelli, 12°20’°S, 133°56°E, Maconochie

1598 (AD, BRI, DNA, PERTH); ca 20 km SE of Maningrida, Rodd 2923

& 2924 (BH, DNA, K, NSW); Gunn Point, Dunlop 3035 (BRI); Near

Darwin, ironstone ridges near coast, Allen 158 (K); Port Darwin, Schultz

372 (K); Cannon Hill Airstrip, Martensz 617 & 618 with Schodde (BRI,

CANB, DNA, K, L); Darwin, CSIRO Research Stn, /rvine 1674 (QRS);

5 km NWW of Humpty Doo, McKean 39 (K); MacMillan Rd, 8 miles SE

of Darwin, Chippendale NT4391 (DNA, NSW): Palmerston Escarpment

Reserve, 12°29°09”S, 130°59°24”E, Smith 4334 (DNA, JCT); Palmerston,

Stapleton, Nov 1915, Hill s.n. (BM); Palmerston, 10 Jun 1886, Lea s.n.

(BM); 35-40 miles from Darwin, on turnoff to Berry Springs, Moore 9227

(BRI); 11 miles S of Berry Springs, turnoff on Stuart Hwy, Beauglehole

10997 (MEL); 2 miles SSW of Mudginberri Stn, Adams 2725 (CANB, K);

34 miles SE of Midginberry HS, 12°45’S, 133°20°E, Lazarides 7795 (K);

30 km W of Jabiru on Kakadu Hwy, Kakadu NP, Dowe 203 (BRI, FTG):

Kakadu NP, Craven 5572 (L, MEL); Kakadu NP, 6 km N of Barramundi Ck,

Dowe 207 (JCT); Kakadu NP, Lazarides 9090 (MEL); 3 km S of Kakadu

NP Headquarters, 12°40’S, 132°43°E, Munir 5590 (NY); 10 miles W of Jim

Jim, Byrnes 1725 (CANB, DNA); Obiri Rock, Maloney 2 (NSW); Between

Rum Jungle and Batchelor, Shaw 868 (BRI); ca 35 km N of Batchelor, Stuart

Hwy, Eichler 17887 & 17888 (AD); ca 160 km SSE of Darwin, Stuart Hwy,

Jackson 1050 (AD); 4 miles E of Lake Evella turnoff, 12°45°S, 135°51’E,

258 Gard. Bull. Singapore 60 (2) 2009

Maconochie 1513 (DNA, MEL, NY, PERTH); ca 15 km W of BHP Camp,

near Caledon Bay, 12°47’S, 136°22’E, Maconochie 1560 (CANB, DNA,

K, NSW); Cox Peninsula, 10 km E of Finniss R Stn turnoff, Dowe 214

(JCT); Cox Peninsula, 1.5 km S of Belyuen, Dowe 215 (JCT); Daly Ra.,

14 km S of Manton Rd, on Stuart Hwy, Rodd 2910 & 2911 (BH, DNA, K,

NSW); Dorisvale Rd, 41 km W of Stuart Hwy, Dowe 2/3 (BRI, DNA); 140

miles ESE of Darwin, Everist 9267 (BRI); Macdonnell Airstrip, 67 miles

N of Katherine, Maconochie 1300 (B, BRI, CANB, DNA, L, K); 16 miles

NE of Tipperary Homestead, Lazarides 6687 (BRI, CANB, K, NSW, US);

near Grove Hill, 13°28’S, 131°35’E, Blake 16347 (A, BRI, K, L); 3 miles S

of Adelaide R., Trapnel/ 99 (BRI); Daly R. Rd, 30 km SSW of Adelaide R.

township, 13°30’S, 131°01°E, 150 m alt., Streimann 8956 with Curnow (L);

Adelaide R., 18 Jun 1943, Tyack Bake s.n. (BRI); Stuart Hwy, 11 km NW

of Pine Ck, 13°46’S, 131°45’E, 240 m alt., Streimann 8955 with Curnow

(L); 22 miles NW of Pine Ck, Perry 1948 (BRI); ca 4 miles W of Pine

Ck, Lazarides 227 & Adams (CANB); 36 miles N of Wilton R., Bumen

‘Crossing, 13°55’S, 134°35’E, Maconochie 1457 (BRI, CANB, DNA, K,

L, NY); Liverpool R., Gulliver s.n. (MEL); Castlereagh Bay, Gulliver

s.n. (MEL); (Darwin), Leanyar, Bosshart 1 (QRS); (Port Darwin), 1855,

Mueller s.n. (BO); Wood Is., 1855, Mueller s.n. (BO, MEL); Liverpool R.,

1855, Mueller s.n. (BO); MacAdam Ra., 1855, Mueller s.n. (MEL); Strauss

Airstrip, 28 miles S of Darwin, Must 957 (BRI, CANB, DNA, K, L); Port

Darwin, Schultz 373 (MEL); Woodah Is, Apr 1922, Tindale (US); N of Pine

Ck on Stuart Hwy, Walter 3494 (B); (Northern Territory), 1892, Holtze s.n.

(MEL); (Port Darwin), Holtze 332 (MEL).

Local and vernacular name: Sand Palm.

Habitat and ecology: Most common in open forest and woodland growing

in deep sandy soils, mostly at low elevation, 0-240 m alt.; occasionally on

rocky ironstone or granite ridges. Usually occurs in extensive populations.

Flowers May-Dec; fruits Nov-May.

Conservation status: Least concern.

Distribution: Australia. Northern Territory. From near Fitzmaurice R.

across the Top End to Cape Arnhem and islands in north-western Gulf of

Carpentaria, and as far inland as Katherine (Fig. 11).

Notes: Livistona humilis was one of two species described by Brown (1810)

when establishing Livistona. Moore (1963a) chose it as the lectotype for

A Taxonomic Account of Livistona R.Br. (Arecaceae) 259

110°E 120°E 130°E 140°E 150°E

” o

° ro

no on

oO o

N N

” (a)

o °o

ae) a)

Legend

no L. eastonii ‘ ae me

? Lt humilis 0 500 1,000 1,500 eS E

L. muelleri Os ee ’

~A*

T T T T T

110°E 120°E 130°E 140°E 150°E

Figure 11. Distribution of Livistona humilis (m), L. muelleri (©), and L. eastonii (0).

the genus. Although Brown only provided a meagre description, Martius

(1838) clearly established its identity, albeit the illustrations accompanying

his description appear to be based on L. inermis or a combination of the

two taxa.

Bentham (1878) cited a specimen of L. rigida (Mueller s.n., Albert

R.) as the one on which he based his description of L. humilis, but his

description is certainly of L. humilis. Beccari (1931) provided a very

thorough appraisal of the species, but however included two varieties, L.

humilis var. sclerophylla from north-east Queensland and L. humilis var.

novoguineensis from Merauke, Indonesia, that are attributable to L. muelleri

(see Notes under that species).

Mueller (1874b) described L. Jeichhardtii based on a collection

made from MacAdam Ra., Mueller s.n., Northern Territory. Kuntze (1891)

provided the combination Saribus humilis.

Livistona humilis is a small sub-canopy dioecious palm to 7 m

tall; leaves are small and regularly segmented; segment apices are rigid,

and with a bifurcate cleft to 89% of the segment length; the inflorescence

is unbranched in both male and female plants, extending vertically to

260 Gard. Bull. Singapore 60 (2) 2009

sub-vertically well beyond the limit of the crown; male plants have

up to 8 partial inflorescences, and female plants a single distal partial

inflorescence; bracts are tightly tubular; flowers are yellow; fruit are

ellipsoid, pyriform, to obovoid to 19 mm long, and shiny purple black at

maturity.

20. Livistona muelleri F.M.Bailey, Queensl. Fl. 5 (1902) 1683. — Type:

Australia. Queensland, Cairns, Oct 1900, Cowley s.n. (holotype, BRI;

isotype, FI). Plate 15A-F.

-Livistona humilis R.Br. ‘var.’, F.M.Bailey, Queensland Agric. J. 2 (1898)

130, prop. stat.

-Livistona humilis var. sclerophylla Becc., Webbia 5 (1921) 20. — Type:

Australia. Queensland, Bloomfield R., Bauer 3 (lectotype, FI) (fide Rodd,

1998).

-Livistona humilis var. novoguineensis Becc., Webbia 5 (1921) 20. — Type:

Indonesia. Papua, Merauke Division, Merauke R., Apr 1901, Jaheri s.n.

(holotype, FI; isotype, BO).

-Livistona brassii Burret, Notizbl. Bot. Gart. Berlin-Dahlem 12 (1935) 309.

— Type: Papua New Guinea. Western Prov., Dagwa, Oriomo R., Feb 1934,

Brass 5950 (holotype, A; isotypes, BM, BO, BRI, NY).

-Livistona crustacea Burret, J. Arnold Arbor. 20 (1939) 189. — Type: Papua

New Guinea. Western Prov., Middle Fly R., Lake Daviumbo, Sep 1936,

Brass 7668 (holotype, A; isotypes, BO, BM, BRI, L; photos, BRI).

Functionally dioecious palm. Trunk to 12 m tall, 15-25 cm dbh, leaf scars

narrow, raised, roughened and with remnant tissue, internodes narrow, grey,

petiole stubs persistent, or deciduous with extreme age or fire. Leaves 25-

35 in a globose crown, held erect; petiole 70-100 cm long, 14-20 mm wide,

adaxially concave, margins with single curved black spines 2-12 mm long

throughout, larger and closer spaced in the proximal portion; both adaxial

and abaxial surfaces with rows of corky scales, persistent, at first red-

brown aging to grey; leaf-base fibres not prominent, fine, disintegrating;

lamina costapalmate, regularly segmented, circular, 60-90 cm long, rigid,

flat, chartaceous, adaxially olive green to grey green, abaxially dull bluish

green, glabrous except for a few scales on ribs; lamina divided for 50-65% of |

its length, with 48-60 segments, depth of apical cleft 5-14% of the segment

length; apical lobes acute, rigid; parallel veins ca 8 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, not sexually dimorphic, 80-160 cm long, not extending beyond

the limit of the crown, branched to 4 orders; partial inflorescences 5-10;

peduncular bract(s) lacking; rachis bracts loosely tubular, with silver scales,

A Taxonomic Account of Livistona R.Br. (Arecaceae) 261

splitting and disintegrating with age, but margins not lacerate; rachillae

2-13 cm long, papillose, maroon to red. Flowers solitary or in clusters of

2-3, 1.3-1.6 mm long; sepals broadly triangular, 0.8-1 mm long, maroon,

fleshy, cuspidate; petals ovate, 1.3-1.6 mm long, bright yellow, subacute;

stamens ca 1.4 mm long, yellow; carpels pink to maroon. Fruit ellipsoid,

10-12 mm long, 8.5-10 mm diam., powdery blue, reddish black or bluish

black; epicarp smooth, pruinose; suture line extends for full length of the

fruit, marked with lip-like structures; mesocarp thin, dry; endocarp very

thin, brittle; pedicel 0.5-1 mm long. Seed globose, 8-9 mm wide. Eophyll

3-ribbed.

Specimens examined: AUSTRALIA: Queensland. Torres Strait, Saibai Is.,

Stocker 1376 (QRS); Torres Strait, Badu Is, Cameron 2737 (QRS); Torres

Strait, Horn Is, Cameron 2031 (QRS); Torres Strait, Horn Is, Cameron

2246 (QRS); Torres Strait, Horn Is, 11 July 1943, Tvack Bake s.n. (BRI);

Torres Strait, Prince of Wales Is, Cameron 20244 (QRS); Torres Strait,

Prince of Wales Is, Cameron 20335 (QRS); Lockerbie Scrub, Dowe 407

(JCT); Jardine R., N bank, near crossing of Peninsula Dev. Rd, Hind 576

(NSW); 10 km S of Jardine R., bypass road, Dowe 378 (JCT); ca 24 km

W of Peninsula Dev. Rd, on track to west coast between Pt Musgrave and

Skardon R., Clarkson 9157 (BRI, QRS); 11.5 km N of Weipa Mission,

Specht 120 (BRI); 31 km E of Weipa-Batavia Rd, Maconochie 2699 (BRI);

4 km NNW of Beagle North Camp, ca 41 km NNE of Aurukun, Clarkson

4359 (K); between Nesbit and Chester Rivers, Hyland 6834 (QRS); Silver

Plains Stn, 8 Apr 1958, Wassell s.n. (BRI); Silver Plains-Goanna Ck Rd,

Webb 3186 (BRI); 2 miles S of Scrubby Ck, /rvine 72 (BRI, QRS); Upper

Massey Ck, 15 miles S of ENE of Coen, Smith 11916 (BRI); Flinders Is,

Elsol 754 (BRI); Flinders Is, Clarkson 2262 (BRI); Edward R. Aboriginal

Reserve, 6.8 km S of Musgrave-Edward R road, Clarkson 3538 (BRI); E

of Edward R. Community, Clarke WCC1082 (BRI); Musgrave Stn, Hind

749 (NSW); Starke Homestead, along track 7.5 km N, 14°59’S, 145°08’E,

ca 40 m alt., Kanis 1947 (BRI, L); Durkam Stn, 10 km NE of Cooktown,

McDonald 1541 (BRI); 11 km WNW of Cooktown, N side of Endeavour

R., near Jones Lagoon, Rodd 3159 & 3160 with Jacobs (BH, BRI, K,

NSW); Near Cooktown, Webb 6159 (BRI); Helensvale-Bloomfield Rd,

ca 3 km N of Ayton, Rodd 3170 & 3171 (BH, BRI, K, NSW); Bloomfield

R., Bauer 3 (FI); Daintree R., Hind 216 (QRS); Near Port Douglas, Blake

19722A (BRI); Craiglea, Cook Hwy near Port Douglas, 38.5 miles N

of Cairns, Moore 9244 (BRI); Craiglea, Cook Hwy near Port Douglas

turnoff, Hind 226 (NSW); NW of Walsh R., 7.2 km NW of Nolans Ck,

16°45’S, 144°06’E, 200 m alt., Dowe 608 (BRI); Trinity Beach, 16°47’S,

262 Gard. Bull. Singapore 60 (2) 2009

145°42°E, Smith & Pedley 10001 (BRI, L); Trinity Beach, N of Cairns,

Maconochie 1717 (DNA); Captain Cook Hwy, W side of road, 2.5 km N

of the Cook Hwy and Kennedy Hwy junction, /rvine 1776, 1777, 1778 &

1779 (QRS); Cairns, Oct. 1900, Cowley s.n. (BRI holotype, FI); Cairns,

Flecker 3251 (QRS); Cairns, Scott St, 8 May 1941, Wright s.n. (QRS);

Cairns, Port Rd, 2 km E of Mulgrave Rd, Dowe 271 (JCT); Few kms S

of Cairns opposite Golf Links on the Bruce Hwy, Moriarty 1168 (BRI);

PAPUA NEW GUINEA: Western Prov. Middle Fly R., Lake Daviumbo,

Brass 7668 (A, BO, BM, BRI, L); Western Prov. Daua (Daru) Is, 09°05’S,

143°10°E, 70 m alt., Gillison 45 (LAE); Western Prov., Dagwa, Oriomo

R., Brass 5950 (A, BM, BO, BRI, NY). INDONESIA: Papua. Merauke

Division, road from Mopa Airstrip to Manggatrikke, ca 10 m alt., van

Royen 4545 (A, BO, K, L, LAE); Merauke Division, Jaheri 197 (BO);

Merauke Division, Jaheri s.n. (BO); Merauke Division, Merauke R., Apr

1901, Jaheri s.n. (BO, FI).

Local and vernacular name: Cairns Fan Palm.

Habitat and ecology: Grows in grassy open forest, woodland, moist

sclerophyll forest, and less commonly on the margins of vine thickets,

0-300 m alt., and most common in areas that have a strongly seasonal

rainfall pattern. Flowers Sep-Apr; fruits Nov-May.

Conservation status: Least concern.

Distribution: Australia, Indonesia and Papua New Guinea. In north

Queensland from the Torres Strait islands to near Innisfail. In Papua New

Guinea in the Western Province; and in Indonesia, in Papua, Merauke

Division (Fig. 11).

Notes: Livistona muelleri was first recognised as distinct and described as

‘L. humilis R.Br. ‘var’ by Bailey (1898), and then as L. muelleri in 1902

based on the collection Cowley s.n. from Cairns, Queensland. It was named

for Ferdinand von Mueller (1825-1896), Victorian Government botanist,

1853-1896. Previously, Solander (1768-1771) had included it as another

manuscript name in his hand-written Plantae novae hollandiae, based on

plants he saw near the Endeavour R., Queensland, but this name was never

published. Previous to Bailey’s account, Wendland, in a contribution to

Kerchove (1878), used the name Livistona miilleri in a list of Australian

palms, but this was merely a name in a list without a description or

reference to specimens and therefore a nomem nudum. Bailey (1898) made

a ee eee

no reference to Wendland’s name and there appears to be no connection in

Bailey's choice of the name, as in reference to L. benthamii noted that he

named both of these species to honour the authors of Flora Australiensis.

Beccari (1921) described two varieties of L. humilis that are

attributable to L. muelleri: L. humilis var. sclerophylla, based on Bauer 3

from the Bloomfield R., Queensland, and L. humilis var. novoguineensis,

based on Jaheri s.n., from Merauke, Papua, Indonesia.

Burret (1935, 1939) described L. brassii and L. crustacea from

Western Prov., Papua New Guinea, based on the collections Brass 3950 and

Brass 7668 respectively. Both taxa were synonymised under L. muelleri by

Rodd (1998).

Although the relationships of L. muelleri are obscure, it most closely

resembles L. eastonii from the Kimberleys, Western Australia. However.

this may be an example of ecological convergence, and its true relationships

may possibly be with moist forest species such as L. benthamii, with which

it shares at least a close association in distribution. Livistona muelleri also

bears morphological resemblance to L. fulva of central Queensland, with

similarity in both leaf and inflorescence.

Livistona muelleri is a moderate sub-canopy palm to 12 m tall:

leaves are moderate and regularly segmented: segment apices are rigid,

and with a bifurcate cleft to 14% of the segment length: the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to 10

partial inflorescences; bracts are loosely sheathing: flowers have maroon

sepals, yellow petals and mauve carpels, on maroon to red rachillae; fruit

are ellipsoid, pyriform, to obovoid, to 12 mm long to 10 mm wide, and

powdery blue, reddish black to bluish black at maturity.

21. Livistona eastonii C.A.Gardner, Forest Depart. Bull. Western Australia

32 (1923) 36 (as L. eastoni). — Type: Australia. Western Australia, Broome

Bay, Napier, Lower King Edward R., 22 Aug 1921, Gardner 1544 (lectotype,

PERTH; isolectotypes, B, BH, CANB, K, MEL: fide Rodd, 1998). Plate

16A-C.

Functionally dioecious palm. Trunk to 15 m tall, 10-15 cm dbh, leaf scars

raised, internodes narrow, grey. leaf-bases often persistent. Leaves 10-20 in

a globose crown; petiole 50-200 cm long, 1.4-2 cm wide, arcuate, margins

with single or double curved brown spines throughout its length but larger

and closer in the proximal portion: leaf-base fibres moderately prominent,

coarse, persistent; lamina costapalmate, regularly segmented, subcircular,

60-90 cm long, coriaceous, adaxially pale greyish green, abaxially lighter

green, yellowish with age, pruinose, waxy; lamina divided for 50-90% of

264 Gard. Bull. Singapore 60 (2) 2009

its length, with 40-50 segments, depth of apical cleft 49-63% of the segment

length, apical lobes acute, rigid; parallel veins 5-9 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, not sexually dimorphic, 100-200 cm long, not extending beyond

the limit of the crown, branched to 4 orders; partial inflorescences 5-6;

prophyll not seen; peduncular bract 1, tubular; rachis bracts tubular with

scattered appressed scales; rachillae 1-9 cm long, glabrous. Flowers solitary

or in clusters of 2-4, 1.6-1.9 mm long, campanulate, cream to yellow;

sepals broadly ovate, 0.2-1 mm long, fleshy, acute; petals narrowly ovate,

0.5-1.9 mm long, thick, acute; stamens ca 1.5 mm long. Fruit obovoid,

infrequently approaching ellipsoid, 12-16 mm long, 8-9 mm diam., glossy

purple-black; epicarp smooth; pedicel 0.5-2 mm long. Seed ellipsoid, 10-

13 mm long. Eophyll not seen.

Specimens examined: AUSTRALIA: Western Australia. 15 miles S

of mouth of Mitchell R., 20 Jan 1973, Schultze s.n. (PERTH); Mitchell

' Plateau, Beard 6992 (PERTH, NSW); Mitchell Plateau, 15 miles (ca 23

km) N of Amax Bauxite Camp, Maconochie 1281 (BRI, CANB, DNA,

K, L, PERTH); Mitchell Plateau Airfield, 14°47°S, 125°48°E, Hnatiuk

MP28 & MP36 (K, PERTH); 2.9 km SE of Mitchell Plateau mining camp,

14°50’S, 125°51°E, George 14503 (CANB, LE, PERTH); Mitchell Plateau,

Bauxite Camp, 18 km SSW of head of Port Warrender Bay, Montagu,

14°47’S, 125°49’°E, 300 m alt., Rodd 2889 (K); Mitchell Plateau, Bauxite

Camp, 20 km SSW of head of Port Warrender Bay, 14°49’°S, 125°50’E,

300 m alt., Rodd 2888 (BH, K, NSW, PERTH); Mitchell Plateau, 25 km

SSW of head of Port Warrender Bay, 14°51°S, 125°49’E, 270 m alt., Rodd

2887 (BH, K, NSW, PERTH); 15 miles S of Admiralty Gulf, Gardner 972

(PERTH); Broome Bay, Napier, Lower King Edward R., Gardner 1544 (B,

BH, CANB, K, MEL, PERTH lectotype); 75 miles NNW of Gibb R Stn,

Speck 4949 (CANB); 10 km SE of mining camp on track to Mitchell Falls

Homestead, 14°53’S, 125°55’E, Fryxell & Craven 4044 (K, MEL, PERTH,

UC); 100 miles S of Kalumburu, Hutchinson 84 (PERTH); Kalumburu

Rd, 40 km N of Drysdale Stn, 15°21’S, 126°13’E, Dowe 360 (BRI);

Kalumburu Rd, 40 km N of Drysdale Stn, 15°21°S, 126°13’E, Dowe 359 &

361 (JCT); 11 km N of Doongan Homestead, 15°21’S, 126°13’E, George

15220 (AD, CANB, K, NSW, PERTH); 8 km N of Doongan Homestead,

15°22’S, 126°13’E, George 15221 (CANB, K, NSW, PERTH); Doongan

Stn, Crosland Ck at Gibb R., 15°23’S, 126°17’E, 380 m alt., Telford 6096

with Butler (CBG, NSW, PERTH); 15 km S of Doongan Stn, 34 km N of

Drysdale R. Crossing, 15°25’S, 126°18’E, Forbes 2293 (L, MEL, PERTH);

25 miles N of new Drysdale Stn, Maconochie 1228 (CANB, DNA, LAE,

A Taxonomic Account of Livistona R.Br. (Arecaceae 265

MEL, NSW, NY, PERTH); 40 km N of Drysdale HS on Kalumburu Rd.

Dowe 360 (BRI): 40 km N of Drysdale Stn crossing, 15°25°S, 126°18°E, 7

June 1984, Willis s.n. (MEL, PERTH): ca 12 km N of Drysdale R crossing,

Kalumburu Rd, Symon 10254 (NSW): Kimberley District, 4 May 1922,

Kessell s.n. (MEL).

Habitat and ecology: Grows in a strongly monsoonal climate in open

woodland, usually on flat sites or depressions and forms extensive colonies,

100-380 m alt. Soils are lateritic in origin. Flowers Apr— Sep: fruits Dec-

June.

Distribution: Australia. Western Australia. In the Kimberley Region

confined to the Mitchell Plateau to as far south as Doongan Stn (Fig. 11).

Notes: Livistona eastonii was described by Gardner (1923) from Lower

King Edward R. in Western Australia, and named for William R. Easton,

who directed the Kimberley Exploration Expedition in 1912 of which

Gardner was the botanist. Gardner cited no specimens in the protologue,

and Rodd (1998) chose Gardner 1544, a specimen collected from th

Lower King Edward R. in Western Australia, as the lectotype.

Livistona eastonii most closely resembles L. humilis, but it

distinguished by its larger size, and by not having sexually dimorphi

inflorescences. Wilson (1992) described L. eastonii as dioecious, but

provided only a description of ‘male flowers’, and noted that the ‘female

flowers’ were not seen, however providing an illustration of a ‘female

inflorescence’.

Livistona eastonii is a moderate sub-canopy palm to 15 m tall:

leaves are moderate and regularly segmented: segment apices are rigid,

and with a bifurcate cleft to 63% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to 6

partial inflorescences; bracts are tubular: flowers are cream to yellow; fruit

are obovoid to ellipsoid, to 16 mm long to 9 mm wide, and glossy purple

black at maturity.

iq?)

ie .

22. Livistona inermis R.Br., Prodr. (1810) 268. -Saribus inermis (R.Br.)

Kuntze, Revis. Gen. PI. 2 (1891) 736.— Type: Australia. Northern Territory,

Gulf of Carpentaria, Sir Edward Pellew Group, North Is. (Brown’s island

‘h’), 16? Dec 1802, Brown (Bennett no. 5795) (holotype, BM: isotypes, FI,

K). Plate 16D-G.

Functionally dioecious palm. Trunk to 10 m tall, 6-10 cm dbh, leaf scars

266 Gard. Bull. Singapore 60 (2) 2009

prominently raised, internodes narrow, grey, persistent petiole stubs in the

lower portion only. Leaves 10-30 in a globose crown; petiole arching, 60-

90 cm long, 6-10 mm wide, adaxially flat or shallowly concave, margins

with small single curved reddish to black spines to 5 mm long restricted

to the proximal portion, distally smooth or with widely spaced small

reddish to black calli to | mm long; leaf-base fibres moderately prominent,

coarse, persistent; lamina costapalmate, regularly segmented, circular to

subcircular, 30-70 cm long, coriaceous, adaxially light green to green-

grey, abaxially lighter green-grey glossy to lightly pruinose, segments

extended along the costa; lamina divided for 80-97% of its length, with

24-48 segments, depth of apical cleft 70-84% of the segment length,

apical lobes acuminate to filiform, rigid to semipendulous; parallel veins

8-10 each side of midrib; transverse veins thinner than parallel veins.

Inflorescences unbranched at the base, not sexually dimorphic, 40-90 cm

long, not extending beyond the limit of the crown, branched to 3 orders;

partial inflorescences ca 3; the most basal partial inflorescence about as

long as the remainder of the inflorescence; prophyll 12-28 cm long, 2-3 cm

wide, papyraceous, glabrous; peduncular bract(s) lacking; rachis bracts

loosely tubular, papyraceous, glabrous, slightly pubescent toward the apex;

rudimentary bracts subtend the distal partial inflorescences; rachillae 1-9

cm long, glabrous. Flowers solitary or in clusters of 2-3, funnel-shaped,

1.5-2.3 mm long, white to cream or yellow; sepals narrowly triangular,

0.7-1.4 mm long, membranous, acute; petals triangular to broadly ovate,

1.5-1.9 mm long, thick, acute to apiculate; stamens 1.5-1.6 mm long;

anthers bright yellow. Fruit obovoid to pyriform, 10-13 mm long, 6-7

mm diam., glossy black; epicarp smooth; suture line extends the length of

the fruit, marked with lip-like structures; mesocarp fleshy; endocarp thin.

Seed ellipsoid, 8-9 mm long. Eophyll 3-ribbed.

Specimens examined: AUSTRALIA: Northern Territory. Melville Is.,

Jul-Aug 1911, Baldwin-Spencer s.n. (MEL); 2 km W of West Alligator R.,

Arnhem Hwy, Benson 952 (NSW); 72 km E of Kakadu NP boundary on

Kakadu Hwy, Dowe 200 (BRI, FTG); Kakadu NP, Craven 6053 (MEL);

Kakadu NP, 6 km from Mt Brockman, Telford 8064 & Wrigley (CBG,

NSW); Kakadu NP, 2.5 km NW of Koongarra Saddle, Telford 814 &

Wrigley (CBG, NSW); Nourlangie, 220 km ESE of Darwin, Beauglehole

58707 (MEL); Banks of Liverpool R., Aug 1819, Cunningham s.n. (M);

W of Liverpool R., June 1869, Gulliver s.n. (MEL); headwaters of

Liverpool R., 12°46’S, 133°44’E, Craven & Wightman 8410 (A, MEL);

Munmarlary Stn, Latz 3728 (CANB, DNA, QRS); 21 miles NW of

Katherine, Lazarides 6627 (B, CANB, L); Katherine Gorge, 15 miles E of

ae ee ee eee Oe

A Taxonomic Account of Livistona R.Br. (Arecaceae) 267

Katherine, Lazarides 7040 (BRI, DNA, K, L); 90 miles S of Maningrida,

12°51’S, 134°32’E, Maconochie 1581 (CANB, DNA, K); El Sharana

Mining Camp, Martensz 384a with Schodde (B, BRI, CANB, DNA, K,

L, NSW, US): 8 km NW of El Sharana, Benson 994 (NSW): 13 miles

E of El Sharana Mining Camp, Lazarides 7974 (K); Katherine Gorge,

ca 0.5 km SE of NP HQ, Rodd 2931 & 2932 (BH, DNA, K, NSW);

Katherine Gorge, Must 955 (BRI); Katherine Gorge, 15 miles NE of

Katherine, Lazarides 6990 (CANB, NSW); 90 km from Pine Ck, Gittens

2590 (BRI, DNA, NSW); Nitmiluk, above visitor centre, Evans 3544

(CANB, DNA, K); Pine Ck-Oenpelli Rd, near Mudginberri Stn, Symon

10347 (NSW); South West Is., 15°41’S, 136°40’°E, McKey 101 (DNA, K,

NSW); Maria Is., 14°52’S, 135°44’E, Dunlop 2836 (AD, BRI, CANB,

DNA, DWL, NSW); Groote Eylandt, 25 km W of Umbakumba on main

road, Fosberg 62325 (BRI); Sir Edward Pellew Islands, North Is., Brown

(Bennett no. 5795) (BM holotype, FI, K); Sir Edward Pellew Islands,

Centre Is., Rice 2037 (BRI, NSW); Sir Edward Pellew Islands, Centre

Is., Craven 3841 (CANB); Wollogorang Stn, Echo Gorge, Thomson

802 (DNA, NSW). Queensland. Wentworth-Troutbeck Stn, Gasteen 20

(BRI); Westmoreland Stn, near Burketown, 17°26’S, 138°10°E, 16 June

1991, Birch s.n. (BRI); Stockyard Camp, 17°29’S, 138°13’E, Melville

1122 (BRI); 50 km E of Wollogorang on road to Burketown, Halford

605 (BRI).

Habitat and ecology: Grows in open forest and woodland and in sandstone

gorges and outcrops, 2-300 m alt. Flowers Aug—Mar; fruits Jun—Sept.

Conservation status: Least concern.

Distribution: Australia. Northern Territory and Queensland. In the northern

part of the Northern Territory from the headwaters of the Daly R. across

the Top End inland to Katherine. In northwest Queensland, including near-

shore islands in the Gulf of Carpentaria (Fig. 12).

Notes: Livistona inermis was one of two species described by Brown

(1810) in establishing Livistona, based on the collection Brown s.n. from

Sir Edward Pellew Islands, Gulf of Carpentaria, Australia in 1802, and

named for the unarmed petioles. The other species was L. humilis. The

identity of L. inermis has been confused by many taxonomists. For example,

Wendland and Drude (1875) established their description of L. inermis on

specimens of what is now known to be L. decora, while Bentham (1878)

suggested that “...it may prove to be a variety only of L. humilis”. Drude

268 Gard. Bull. Singapore 60 (2) 2009

110°E 120°E 130°E 140°E 150°E

D 7p)

oA] °o

on | 79)

ai °

N N

oO 7p)

o7 °o

oO o

Legend ine:

D | L. inermis ae cs Sar Ly

° j ie % °

= ® L. lorophyila ; = fond a a. 3

ot oe ee g (

L. victoriae km fae

T T T T fe:

110°E 120°E 130°E 140°E 150°E

Figure 12. Distribution of Livistona inermis (O), L.lorophylla (m), and L. victoriae (©).

(1893) placed it as a possible synonym of L. leichhardtii (= L. humilis).

Beccari (1931) applied considerable discussion on the true identity of L.

inermis and provided the first account that succinctly characterised the

species, thus profoundly separating it from L. humilis. Saribus inermis was

a combination proposed by Kuntze (1891).

Subsequent records of L. inermis in various flora and regional

accounts have been somewhat variable. It was correctly recorded and

described by Ewart and Davies (1917) and Blake (1954), while Specht

and Mountford (1958) confused it with L. humilis and Chippendale (1972)

identified it as L. /orophylla. Gardner (1930) named the Western Australian

populations of L. /orophylla as L. inermis.

Rodd (1998) described L. inermis with a “sometimes branching”

trunk. Although the stem may be damaged and form ‘spontaneous branching’

(‘freak branching’ cf. Tomlinson, 1990), it is not a naturally occurring

event. Spontaneous branching is usually the result of physical damage |

to the apical meristem, with the result that more than one growing point

can develop. During fieldwork, populations of L. inermis were observed

in which individual plants were tightly grouped and may be incorrectly

A Taxonomic Account of Livistona R.Br. (Arecaceae) 269

interpreted as ‘basally branching’. However, examination of these palms

revealed that ‘branches’ were indeed separate plants or individuals that had

been damage by fire or termites.

Livistona inermis is a moderate sub-canopy palm to 10 m tall;

leaves are small and regularly segmented; segment apices are rigid to semi-

pendulous, and with a bifurcate cleft to 84% of the segment length; the

petiole is unarmed, or with very small spines or calli; the inflorescence

is unbranched, not extending beyond the limit of the crown, and with up

to 3 partial inflorescences; bracts are loosely tubular; flowers are white to

cream; fruit are obovoid to pyriform, to 13 mm long to 7 mm wide, and

glossy black at maturity.

23. Livistona lorophylla Becc., Webbia 5 (1921) 18 (as L. loriphylla).

— Type: Australia. Western Australia, Cambridge Gulf, 1885, Johnston s.n.

(holotype, FI; isotypes, NSW, MEL). Plate 17A-B.

-Livistona kimberleyana A.N.Rodd, Telopea 8 (1998) 121, figs. 4g, 17d.

— Type: Australia. Western Australia, Kimberley Region, Durack Ra.,

SW base of Mt King, 24 Oct 1974, Rodd 2866 (holotype, NSW; isotypes,

CANB, K, PERTH).

-Livistona sp. ‘A’, Wilson, Fl. Kimberley Region (1992) 1250, prop. stat.

-Livistona alfredii auct. non F.Muell., Fitzgerald, J. & Proc. Roy. Soc.

Western Australia 3 (1918) 24; Gardner, Forest Depart. Bull. Western

Australia 32 (1923) 36.

Functionally dioecious palm. Trunk to 15 m tall, 8-20 cm dbh, leaf scars

raised, internodes narrow, grey, persistent petiole stubs in the lower portion

only. Leaves 25-40 in a globose crown; petiole arching, 70-200 cm long,

10-17 mm wide, adaxially flat, margins with scattered single curved black

spines up to ca 10 mm long; leaf-base fibres moderately prominent, coarse,

persistent; lamina costapalmate, regularly segmented, circular, 60-100 cm

long, coriaceous, glossy mid green to pruinose grey on both surfaces; lamina

divided for 85-98% of its length, with 34-50 segments, depth of apical

cleft 55-78% of the segment length, apical lobes acuminate to filiform,

semi-pendulous; parallel veins 8-10 each side of midrib; transverse veins

thinner than parallel veins. Inflorescence unbranched at the base, not

sexually dimorphic, 20-160 cm long, not extending beyond the limit of

the crown, branched to 3 orders; partial inflorescences 4-8; prophyll not

seen; peduncular bract 1; rachis bracts loosely tubular, sparsely tomentose;

rachillae 1-6 cm long, glabrous. Flowers solitary or in clusters of 2-4,

broadly funnel-shaped, 1.2-3 mm long, cream; sepals triangular, 0.7-1.8

mm long, membranous, acute; petals triangular to broadly ovate, 1.2-3 mm

270 Gard. Bull. Singapore 60 (2) 2009

long, acute to cuspidate, cream; stamens ca 1.6 mm long. Fruit obovoid

to pyriform, 8-14 mm long, 6-9 mm diam., pruinose dull black; epicarp

with scattered lenticellular pores; mesocarp thin, fibrous; endocarp thin,

crustaceous. Seed ellipsoid, 7-9 mm long. Eophyll not seen.

Specimens examined: AUSTRALIA: Western Australia. S slopes of Sir

Graham Moore Is. facing the Geranium Islands, Willing 293 (PERTH);

Sir Graham Moore Is., 13°56’S, 126°33°E, Wilson 1/222 (PERTH):

Stewart R., ca 68 km NNE of Derby, 14 km from Kimbolton Stn, 7e/ford

6404 with Butler (CBG, NSW, PERTH); North Kimberley, Speck 4916

(CANB, PERTH); Kalumburu, Crawford 58 (PERTH); ca 3 km ESE of

Kalumburu Mission, Rodd 287] (BH, K, NSW, PERTH); 19 km W of

Cape Rulhiers, 13°56’05”S, 127°11’10”E, Kenneally 10145 (PERTH);

Kalumburu Mission, Brigden DNA6885 (CANB); Kalumburu, 18 Jan

1975, Lullfitz s.n. (PERTH); Kalumburu, 3 Jul 1960, Douglas & Mees

s.n. (PERTH); 190 km W of Wyndham, Gibb R.-Kalumburu MissionRd,

~1.5 km S of Doongan R., Beauglehole 51787 (PERTH); Cambridge Gulf,

1885, Johnston s.n. (FI holotype, NSW, MEL); Mitchell Plateau, N of

mining camp, Beauglehole 59046 with Errey 2746 (PERTH); Drysdale R.

NP, Fern Gully, 14°39°S, 126°57’E, Kenneally 4557 (PERTH); Drysdale R.

NP, 14°40’S, 123°00’E, George 13758 (PERTH, NSW); Drysdale R. NP,

Planigale Ck, 14°43’S, 126°54’E, Kenneally 4447 (PERTH); Drysdale R.

NP, Cracticus Falls, 14°47°S, 127°0S’E, Kenneally 4153 (PERTH, NSW);

Drysdale R. NP, Carson Escarpment Plateau, 2 km N of Face Point (at

Laryoo), 14°50°30”S, 126°48°30”, 240 m alt., Forbes 2341S (MEL);

Brunswick Bay, Unwins Is., 15°08’S, 124°48°E, Wilson 11450 (PERTH);

West Kimberleys, upper reaches of Roe R., 15°12’S, 125°32’E, Kenneally

9991 (PERTH); Prince Regent R. Reserve, Marigui Promontory, 15°20’S,

124°56°E, Kenneally 2162 (NSW, PERTH); King Leopold Ranges, Sale

R. 29.2 km WSW of Mt French, 15°59°15”S, 124°38°30”E, Alford 1402

(PERTH); Chamberlain R. Gorge, 7 km W of El Questro Stn Homestead,

16°00’S, 127°55’E, Kenneally 10978 (PERTH, NSW); El Questro Stn,

Cambridge Gulf, 100 m E of Pentecost R., Dowe 362 (BRI, FTG); Sale

R., 15.5 km W of Mt Lochee, 16°04’S, 124°45’45’E, 80 m alt., Kenneally

10464 (PERTH); West Kimberleys, gorge of unnamed creek running

W of Sale R. 30 km ESE of mountain, Kenneally 9652 (PERTH); West

Kimberleys, 6.5 km NW of Kimbolton Homestead on mid Stewart R.,

16°41°S, 123°50°E, Hastings B2 (PERTH); West Kimberley, 1901, House

s.n. (PERTH); Kimberleys, 54.7 km SW of turnoff to Beverley Springs

Homestead, Gibb R. Rd, Kenneally 9793A (PERTH); King Leopold |

Ranges, 2 km W of March Fly Glen towards Derby on Gibb R. Rd

—————

ae Ce ee ee

A Taxonomic Account of Livistona R.Br. (Arecaceae) 27M

near Mt Bell, 17°10’°S, 125°17’E, Hastings 1 (PERTH); Teroni Gorge, 12

miles NNW of Elgie Cliffs Stn, Lazarides 6453 (CANB); King Leopold

Ranges, 26 km NE of Inglis Gap, 17°12’S, 125°32’E, Dowe 356, 357

& 358 (BRI, JCT); SW base of Mt King, Durack Ra., Rodd 2867 &

2868 (CANB, K, NSW, PERTH, QRS); King Leopold Ranges, Lenard

R. Gorge, ca 130 km ENE of Derby, Carr 3978 & Beauglehole 47756

(PERTH); King Leopold Ranges, Mt Herbert, May 1905, Fitzgerald s.n.

(PERTH); King Leopold Ranges, 11 miles NE of Inglis Gap, Maconochie

1296 (L); ca 26 km NE of Inglis Gap, Symon 10154 (NSW): Durack

Ra., SW base of Mt King, Rodd 2866 (CANB, K, NSW, PERTH); Mt

Broome, Fitzgerald 815 (PERTH); Mt Leake, Fitzgerald 1206 (PERTH).

Northern Territory. Victoria R. area, 15°16’S, 129°35’E, Dunlop 8203 &

Leach (NSW).

Habitat and ecology: Grows in open forest and woodlands, and on sandstone

outcrops and gorges, 10-300 m alt. Flowers Aug-Dec; fruits Dec-July.

Conservation status: Least concern.

Distribution: Australia. Kimberley region of Western Australia, from Sir

Graham Moore Is. to Sale R. and Cambridge Gulf, and inland to the King

Leopold and Durack Ranges, and just within Northern Territory in the

Victoria R. area (Fig. 12).

Notes: Livistona lorophylla was described by Beccari (1921) based on

Johnston s.n. collected at Cambridge Gulf, Western Australia, and named

for the resemblance of the leaf segments to leather straps.

Livistona kimberleyana was described by Rodd (1998) based on

Rodd 2866 collected from Mt King, Western Australia. This taxon was

distinguished from L. /orophylla primarily by grey pruinose rather than

green leaves. However, in some populations individuals exhibit intermediate

leaf colour. As noted by Rodd: “... /t is possible even that further study may

show it to represent an arid race of L. lorophylla, but for the meantime it

seems preferable to treat it as a distinct species.” Other than leaf colour,

there are no other characters to distinguish the taxa and L. kimberleyana

is therefore placed as a synonym of L. lorophyilla. The original spelling

provided by Beccari (1921) was L. loriphylla. This was corrected to L.

lorophylla by Rodd (1998).

Some populations now regarded as L. /orophylla had previously

been assigned to other taxa. Fitzgerald (1918) misidentified it as L.

alfredii, occurring in the King Leopold and Durack Ranges (= Rodd’s L.

Die. Gard. Bull. Singapore 60 (2) 2009

kimberleyana), while Gardner (1923) noted it at the same locations, also

under L. alfredii, as well as throughout the presently known distribution

of L. lorophylla throughout the Kimberleys. Gardner (1930) also referred

to L. lorophylla as L. inermis in a later systematic account. Conversely,

some populations of L. inermis in the Northern Territory have been

incorrectly named as L. /orophylla (Chippendale, 1972).

Livistona lorophylla is a moderate sub-canopy to canopy palm to

15 m tall; leaves are small and regularly segmented; segment apices are

semi-pendulous, and with a bifurcate cleft to 78% of the segment length;

petiole margins are strongly spined in the basal portion; the inflorescence

is unbranched, not extending beyond the limit of the crown, and with up

to 8 partial inflorescences; bracts are loosely tubular; flowers are cream;

fruit are obovoid to pyriform, to 14 mm long to 9 mm wide, and pruinose

dull black at maturity.

24. Livistona victoriae A.N.Rodd, Telopea 8 (1998) 123, figs. 1b, 4f, 17c.

_— Type: Australia. Northern Territory, Victoria R. Crossing, Katherine-

Kununurra Rd, 3 Nov 1974, Rodd 2934 (holotype, NSW; isotypes, DNA,

K, PERTH). Plate 17C-D.

-Livistona sp. “‘B’, Wilson, Fl. Kimberley Region (1992) 1250, prop. stat.

Functionally dioecious palm. Trunk to 17 m tall, 15-30 cm dbh, leaf scars

narrow, internodes narrow, grey, petiole stubs not persistent. Leaves 25-40

in a globose crown; petiole arching, 80-200 cm long, pruinose, adaxially

flat, margins with single, curved, blunt, black thorns, confined to the

proximal portion; leaf-base fibres moderately prominent, fine, persistent;

lamina costapalmate, regularly segmented, subcircular, 80-100 cm long,

coriaceous, adaxially grey green or bluish green, strongly pruinose, dull,

abaxially similar; lamina divided for 55-65% of its length, with 40-56

segments, depth of apical cleft 55-70% of the segment length, apical

lobes attenuate, rigid; parallel veins 9-11 each side of midrib; transverse

veins thinner than parallel veins. Inflorescences unbranched at the base,

not sexually dimorphic, 50-150 cm long, not extending beyond the limit

of the crown, branched to 4 orders; partial inflorescences 5-9; prophyll

not seen; peduncular bract 1; rachis bracts loosely sheathing, fibrous

with age, densely tomentose; rachillae 1-3 cm long, glabrous. Flowers

solitary or paired, funnel shaped, 2-4 mm long, cream to pale yellow;

sepals triangular, 0.7-0.8 mm long, membranous, acute; petals narrowly

ovate, 1.2-1.5 mm long, acute; stamens ca 0.9 mm long. Fruit globose

to ellipsoid, 8-15 mm long, ca 10 mm diam., dull dark reddish brown

to black; epicarp smooth; suture line extends for most of the length of

A Taxonomic Account of Livistona R.Br. (Arecaceae) /3

the fruit; mesocarp thin, fibrous; endocarp thin, brittle; pedicel 0.5-1 mm

long. Seed globose, ca 9 mm wide. Eophyll 3- ribbed.

Specimens examined: AUSTRALIA: Western Australia. Kimberleys, 16

km NE of Bungle Bungle Outcamp, 17°13°S, 128°26°E, Kenneally 9212

(PERTH): Bungle Bungle Ra., 3 km SE of Outcamp, Kenneally 9260 (K,

PERTH); between Njitparriya and Dilmariyu, 3 km SE of Bungle Bungle

Outcamp, 17°21°30°S, 128°21°30°E, Forbes 2587 (BRI, MEL): between

Njitparriya and Dilmariyu, 3 km SE of Bungle Bungle Outcamp, 17°21°30”S,

128°21°30°E, 320 m alt., Scarlett 315 (MEL); Kimberleys, 15 km SE

of Bungle Bungle Outcamp, Piccaninny Ck Gorge, 17°27°S, 128°25°E,

Blackwell BB419 (PERTH); Bungle Bungle Ra., gorge off Piccaninny Ck,

Briggs 9309 (NSW); Bungle Bungle Ra., Frog Hole Canyon, Briggs 9307

(NSW); Ord R basin, Koford & Dortch 120 (PERTH); East Kimberleys,

Carr Boyd Ranges, Cabbage Tree Ck, Rodd 2800 (K, NSW, PERTH): East

Kimberleys, Carr Boyd Ranges, Wyndham to Halls Ck Rd, 1978, Hand

s.n. (NSW): Thompson Spring, ca 40 km SE of Kununurra, 6 Jun 1979,

Ryan s.n. (PERTH, NSW); 7 miles E of Denham R. Stn, Perry 2529 (BRI,

CANB, K, MEL, NSW, US); near Lucky Hill, 23 km NNE of Dunham R

Stn, Lazarides 8552 (CANB, K, PERTH): Teroni Gorge, 12 miles NNW

of Elgie Cliffs Stn, Lazarides 6395 (CANB):; Tributary of Keep R., Nov

1978, Lullfitz s.n. (DNA, NSW). Northern Territory. Gregory NP, Victoria

R., ca 29 km NW of Bullita Outstn, Duretto 991a with Coles (MEL); Jasper

Gorge, Carr & Beauglehole ACB 46741 (DNA, MEL): N side of Jasper

Gorge, below sandstone cliffs, McGillivray 3804 with George (DNA,

NSW): Jasper Gorge, Parker 1065 (DNA, NSW); Jasper Gorge, 2 km S of

Charles Crossing, Dowe 208 (BRI, FTG): Victoria R. Crossing, Katherine-

Kununurra Rd, Rodd 2934 (DNA, K, NSW holotype, PERTH); 2 miles E

of Victoria R., N of Katherine-Wyndham Rd, Beauglehole 11044 (MEL):

ca 3.5 km NE of Victoria R., Carr & Beauglehole ACB 46751 (MEL):

Victoria R., Dunlop 3158 (DNA); Victoria R., Yambarran Ra., Leach 4554

with Walsh (BRI, DNA, MEL, NSW); 5 km E of Victoria R. Crossing,

Victoria Hwy, Maconochie 2493 (DNA, K, NSW).

Local and vernacular name: Victoria River Fan Palm.

Habitat and ecology: Grows in open forest in sandstone gorges, inescarpment

gullies and cliff bases and along streams and intermittent watercourses, 50-

320 m alt. Flowers Mar-Dec; fruits Dec-July.

Conservation status: Lower risk, conservation dependent.

274 Gard. Bull. Singapore 60 (2) 2009

Distribution: Australia. Northern Territory and Western Australia. In the

region between the Victoria R. basin and Bungle Bungle Ra (Fig. 12).

Notes: Livistona victoriae was described by Rodd (1998) based on

a collection from Victoria R. Crossing, Rodd 2934, and named with

references to the species occurring in the vicinity of the Victoria R. and also

to complete the trio of names of L. alfredii and L. mariae, species named

for Queen Victoria’s son and daughter-in-law respectively. Previously the

palm had informally been known as ‘Livistona sp. Victoria River’ (Jones,

1996).

Livistona victoriae 1s a moderate canopy palm to 17 m tall; leaves

are moderate and regularly segmented; segment apices are rigid, and

with a bifurcate cleft to 70% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to

nine partial inflorescences; bracts are loosely sheathing; flowers are cream

to pale yellow; fruit are globose to ellipsoid, to 15 mm long to 10 mm wide,

and dull dark reddish brown to black at maturity.

25. Livistona benthamii F.M.Bailey, Queensl. Fl. 5 (1902) 1683 (as L.

benthami). — Type: Australia. Queensland, Cape York Peninsula, Somerset,

Dec 1897, Jardine s.n. (holotype, BRI). Plate 18A-C.

-Livistona humilis R.Br., var., F.M.Bailey, Queensland Agric. J. 2 (1898)

130, prop.stat.

-Livistona holtzei Becc., Webbia 5 (1921) 18. — Type: Australia. Northern

Territory, Port Darwin, undated (1882?), Holtze & Holtze s.n. (holotype,

Fl);

-Livistona melanocarpa Burret, J. Arnold Arbor. 20 (1939) 190. — Type:

Papua New Guinea. Western Prov., Daru Is, 9 Mar 1936, Brass 6310

(holotype, BRI; isotypes, A, BM, BO, L).

Functionally dioecious palm. Trunk to 18 m tall, 13-20 cm dbh, leaf scars

raised, internodes narrow, grey, petiole stubs long because of the abscission

of leaves occurring in about the middle of the petiole; stubs persistent on

the trunk for most of its length, unless decayed by age or fire. Leaves 30-50

in a globose crown; petiole 120-200 cm long, 10-18 mm wide, adaxially

moderately ridged, margins with single curved black spines congested in

the proximal portion; leaf-base fibres prominent, coarse, persistent; lamina

costapalmate, regularly segmented, circular, 90-160 cm long, chartaceous,

adaxially dark green, abaxially lighter green, glossy, nonwaxy; lamina

divided for 60-75% of its length, with 50-80 segments, depth of apical

cleft 60-75% of the segment length, apical lobes acute, pendulous; parallel

A Taxonomic Account of Livistona R.Br. (Arecaceae) 275

veins 8 either side of midrib; transverse veins thinner than parallel veins.

Inflorescences unbranched at the base, not sexually dimorphic, 120-210

cm long, not extending beyond the limit of the crown, branched to 2 orders;

7-9 partial inflorescences; prophyll not seen; peduncular bract(s) lacking:

rachis bracts tightly sheathing, light brown, with sparse scurfy silver scales;

rachillae 5-12 mm long, patchily pubescent. F/owers solitary or in clusters

of 2-3, tubular, whitish/cream to pale yellow; sepals triangular, 0.8-2 mm

long, membranous, acute; petals broadly triangular, 1.0-1.3 mm long,

fleshy, acute; stamens ca | mm long. Fruit globose to obovoid or pyriform,

9-13 mm long, 9-11 mm diam., pruinose purple-black; epicarp smooth,

suture line extends for the length of the fruit; pedicel 2-3 mm long. Seed

ovoid, 8-9 mm long. Eophyll not seen.

Specimens examined: AUSTRALIA: Northern Territory. Port Darwin,

undated (1882?), Holtze & Holtze s.n. (Fl); Darwin, Black Jungle,

Trapnell 105 (BRI); Koolpinyah, Bankers Jungle, Dowe 220 (JCT); S of

Koolpinyah, 13°32’S, 131°14’E, Blake 16967 (BRI, K, L); Howard R.,

Must 1150 (CANB, DNA); Kakadu NP, Cooinda, Home Billabong, Dowe

206 (BRI); Elcho Is., Warangaiyu Lagoon, 11°57’S, 135°43’E, Latz 6273

(BRI, CANB, DNA, K, L); Elcho Is., Maconochie 2119A & 2119B (CANB,

DNA, NSW); Howard Springs, ca 3 km N of Howard Springs, 23 km E of

Darwin, 12°25’S, 131°03’E, Rodd 2904 & 2905 (DNA, K, NSW, QRS);

Howard Springs, E of Stuart Hwy, Beauglehole 10970 (MEL); Fright Pt,

E of Darwin, Ginger Palmers Camp, Tracey 14045 (BRI); (Port Darwin),

Holtze 539 (MEL); Oenpelli, 12°18’S, 133°04’E, Specht 1187 (AD, BRI,

CANB, K, MEL, US); Jim Jim Ck, Jacobs 1790 (NSW); East Alligator

R., Obiri Rock, Maloney 1978 (NSW). Queensland. Torres Strait, Saibai

Is., Stocker 1397 (QRS); Lockerbie Scrub, Bamaga-Pajinka Rd, ca 15 km

S of Pajinka, 10°48’S, 142°29’E, 20 m alt., Dowe 406 with Smith (JCT);

Somerset, Dec 1897, Jardine s.n. (BRI holotype); Cape York, 7 miles NE

of Bamaga, Long Scrub, Webb 6122 with Tracey (BRI); Swamp Ck, /rvine

1887 (QRS); Jardine R., N bank, near Peninsula Development Rd crossing,

Hind 377 (NSW); Archer R., gallery forest, Hyland 7604 (QRS); Archer

Bend, near junction of Archer R. and Coen R., Webb 13470 (BRI); Archer

Bend NP, Smyth s.n. (BRI). INDONESIA: Papua, Merauke Division. Wasur

NP, Kali Maro, Widjaja, Wally & Subari EW6031 (BO); Papua, Merauke

Division. Wasur Village 2, Maturbongs MAT661 with Wanggai & Wally

(AAU, BO, CANB, K, MAN). PAPUA NEW GUINEA: Central Prov. ca

7 miles W of Kanosia Plantation, Kairuku, ca 10 ft alt., Darbyshire 685 (A,

BO, BRI, CANB, K, L, LAE); Waigani Swamp, 9°30’S, 147°10°E, Gillison

22018 (A, BRI, CANB, L); Brown R., 9°15’S, 147°20’E, 100 ft alt., Millar

276 Gard. Bull. Singapore 60 (2) 2009

38385 (BRI, CANB, K, L, LAE); Brown R., ca 6 miles W of Karema, 50

ft alt., Schodde 2677 (BO, BRI, CANB, K, LAE). Western Prov., Daru

Is., Brass 6310 (A, BM, BO, BRI, L); Upper Wassi Kussa R., left branch,

Brass 8645 (A, BM, BO, BRI, L); Bula village at mouth of Morehead R..,

Pullen 7027 (CANB, L, LAE); Daru Is., Brass 6406 (A, BM, BRI, L);

Daru Is., 9°05’S, 143°10’E, 2 m alt., Streimann & Lelean 18472 (L, LAE);

Balimo District, 20 km NW of Daru, Paijmans 1523 (CANB).

Habitat and ecology: Grows in swamp forests, on alluvial flats, on the

leeward side of mangroves and in moist to wet areas in monsoonal thickets

and gallery forests at low elevations, 0-100 m alt. Flowers Sep-Jan; fruits

Nov-Apr.

Conservation status: Least concern.

Distribution: Australia, Indonesia and Papua New Guinea. In Australia in

the Northern Territory from Daly R. across the Top End to east Arnhem

Land, and in Queensland from Moa Is. in Torres Strait through Cape

York to MclIlwraith Ra. In Papua New Guinea in the Western and Central

Provinces, and in Indonesia in Papua, Merauke Division (Fig. 13).

Notes: As with L. muelleri, Bailey (1898) first described L. benthamii as

a possible new variety of L. humilis, but subsequently described it based

on the specimen Jardine s.n. (Bailey, 1902). Bailey noted “... / place this

and a palmate-leaved palm from Cairns as varieties of Dr. Robt. Browns

Livistona humilis for the present: but hope to have full and complete

material for a more satisfactory arrangement when working up the order

for my work now in hand on the Queensland Flora’. In naming both L.

benthamii and L. muelleri, Bailey (1902) wrote: “... give them specific rank

bearing the names of the authors of the Flora Australiensis”.

For the population occurring in the Northern Territory, Australia,

Beccari (1921) provided the name L. ho/tzei based on Holtze s.n. and for

New Guinea, Burret (1939) provided the name L. melanocarpa based on

Brass 6310 collected from Western Prov., Papua New Guinea. Both of

these taxa are attributable to L. benthamii.

Livistona benthamii is a large canopy palm to 18 m tall, with long

retained petiole stubs; leaves are moderate to large and regularly segmented;

segment apices are pendulous, and with a bifurcate cleft to 75% of the

segment length; the inflorescence is unbranched, not extending beyond the

limit of the crown, and with up to 9 partial inflorescences; bracts are tightly

sheathing; flowers are whitish cream to pale yellow; fruit are globose to

ae ney

A Taxonomic Account of Livistona R.Br. (Arecaceae) Pe

110°E 120°E 130°E 140°E 150°E

o |

o7 be

a | oF, - =

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wo a ie

Fe es :

9) w

on; Sas fe 14 ro

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Legend — a =

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Oo} « L decora ; me sie ee ~~ Te

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a eee

110°E 120°E 130°E 140°E 150°E

Figure 13. Distribution of Livistona benthamii (c), L. concinna (m), L. drudei (©), and

L. decora( A).

obovoid to pyriform, to 13 mm long to 11 mm wide, and pruinose purple

black at maturity.

26. Livistona concinna Dowe & Barfod, Austrobaileya 6 (2001) 166, fig.

1. — Type: Australia. Queensland, ca 5 km N of Cooktown Airport, Barrett

Ck, 15°25’S, 145°11°E, 5 m alt., 17 Oct 2000, Dowe 607 (holotype, BRI:

isotypes, AAU, K, QRS). Plate 18D-G.

Dioecious palm. Trunk to 30 m tall, 24-35 cm dbh, leaf scars raised:

internodes narrow; grey, petiole stubs not persistent. Leaves 50-65 in a

globose to conical crown: petioles 120-300 cm long, glabrous, green

throughout, 5-11 cm wide proximally, 2.2-3.5 cm wide in mid area, ca

2.8 mm wide distally, triangular in cross-section, adaxially moderately

ridged, margins with solitary symmetric or retrorsely or latrorsely curved,

black spines 3-5 mm long congested in the proximal portion, distal

margin unarmed, sharp, slightly winged: leaf-base fibres not prominent,

coarse, persistent; lamina costapalmate, regularly segmented, subcircular

in outline, glabrous, adaxially midgreen, abaxially slightly lighter green,

278 Gard. Bull. Singapore 60 (2) 2009

glossy on both surfaces, nonwaxy, subcircular in outline, 155-165 cm long,

ca 200 cm wide, undulate; lamina divided for ca 60% of its length, with

60-78 segments, depth of apical cleft ca 41% of the segment length, apical

lobes acuminate, filamentous, pendulous; hastula raised, sharp, papery

on the margins; parallel veins 9-10 each side of midrib; transverse veins

thinner than parallel veins. Inflorescences unbranched at the base, sexually

dimorphic; nonfruit bearing (functionally male) inflorescences 120-180 cm

long, not extending beyond the limit of the crown, branched to 4 orders,

slightly curved with 8-9 partial inflorescences; peduncle 25 mm wide, 10

mm thick, glabrous; fruit bearing (functionally female) inflorescences

160-250 cm long, branched to 5 orders, slightly curved, held horizontal to

semi-pendulous with 8-9 partial inflorescences; peduncle ca 30 mm wide;

peduncular bract(s) lacking; prophyll 27-35 cm long, glabrous; rachis

bracts 30-50 cm long, tightly tubular, not disintegrating into open fibres but

remaining intact with age, glabrous, apically acute, margins entire, lateral

splits uneven, one ca twice as deep as the other; rachillae 5-20 cm long,

glabrous. Flowers, male and female similar in gross morphology, solitary

or in clusters of 2-4, 1.6-2 mm high and 1.5-2 mm wide, white to cream;

sepals basally fused, cupular, 3-lobed, lobes triangular, margins hyaline, to

ca 1.5 mm high, apices acute; petals triangular, slightly asymmetric, 2.0-

2.2 mm long, 1.8-2 mm wide at the base, apex acute; stamen ca 1 mm high,

filament subulate, connective 0.2 mm long; carpels ca 1 mm high. Fruit

globose, 9-12 mm diam., shiny black; stigmatic remains subapical; epicarp

smooth, with scattered lenticellular pores, drying slightly rugose; suture

line extends for about *%4 the length of the fruit; mesocarp ca | mm wide,

moist, oily, gritty; endocarp thin, crustaceous, light brown, tessellate, 0.1—

0.2 mm wide; pedicel to 2 mm long. Seed globose to subglobose; intrusion

extends to half or less of the width of the seed, contorted; embryo lateral to

sublateral, ca 2 mm long. Eophyll 5-ribbed.

Specimens examined: AUSTRALIA: Queensland. Lakefield NP, Kennedy

Bend, 15°50’S, 144°15’E, 50m, Dowe 415 with Smith (JCT); Lakefield NP,

Twelve Mile Waterhole, Dowe 606 with Barfod (BRI, K, JCT); Kennedy

R., 5.5 km N of New Laura Stn, 15°27’S, 144°18’E, 50 m alt., riparian

forest, Irvine 2204 & 2205 (QRS); near Cooktown airport, Barrett Ck,

Gray 2764 (QRS); ca 5 km N of Cooktown Airport, Barrett Ck, 15°25’S,

145°11°E, 5 malt., Dowe 607 (AAU, BRI holotype, K, QRS); 3.3 km NE

of Cooktown Airport, Barrett Ck, NW boundary, 15°25’S, 145°12°E, 0 m

alt, riparian evergreen mesophyll vine forest bordering mangroves, /rvine

2178, 2179 & 2180 (QRS); 16 km N of Cooktown, Barrett Ck, Dowe 604

with Barfod (BRI, JCT, K); Endeavour R., opposite the SW corner of the

A Taxonomic Account of Livistona R.Br. (Arecaceae) 279

Cooktown Airport, 15°27’S, 145°11’E, 3 m alt., Irvine 2181] (QRS); 3.3

km off main road at N side of Cooktown Airport, Hind 4594 with Hill &

Healy (NSW); Cooktown, N side of Endeavour R., near airport, Dowe

253, (ICT).

Specimens from cultivated material: AUSTRALIA: Queensland, Cooktown,

Anzac Park, Dowe 252 (BRI, FTG, JCT); Queensland, Townsville,

Anderson Park Botanic Gardens, Dowe s.n. (JCT).

Local and vernacular names: Cooktown Livistona, Kennedy River

Livistona.

Habitat and ecology: \n seasonally moist open forest, seasonally inundated

Melaleuca swamp, along creek and river banks, and at mangrove margins

in non-saline environments at low elevations, 0-100 m alt. Soils are usually

alluvial; grows with Corypha utan in the Kennedy R. area and with L.

muelleri in the vicinity of Cooktown. Flowers Dec-Mar; fruits Apr-Oct.

Fire regularly occurs in most populations.

Conservation status: Vulnerable.

Distribution: Australia. Queensland. On Flinders Is, Kennedy R. and

tributaries, Barrett Ck, north of Cooktown, and Archer Point (Fig. 13).

Notes: This taxon was placed tentatively under L. drudei by Rodd (1998),

who cited three specimens, /rvine 2204 & 2205, and Hind 4594 that were

collected in the distributional range of L. concinna. The latter consists of a

fallen dead leaf and old infructescence. Dowe and Barfod (2001) described

L. concinna based on Dowe 607, and named it in reference to the neat

and regular appearance of the petioles in the crown. The species had been

variously known as “Livistona sp. Cooktown’, ‘Kennedy River Livistona’

and ‘Cooktown Fan Palm’ in informal accounts (Irvine, 1984: Jones, 1996;

Tucker, 1988).

Livistona concinna is a large dioecious canopy palm to 30 m tall;

leaves are large and regularly segmented; segment apices are pendulous, and

with a bifurcate cleft to 41% of the segment length; the inflorescences are

unbranched, not extending beyond the limit of the crown; male inflorescences

to 180 cm long, branched to 4 orders, and with up to 9 partial inflorescences;

female inflorescences to 250 cm long, branched to 5 orders, and with up to 9

partial inflorescences; bracts are tightly tubular; flowers are white to cream;

fruit are globose, to 12 mm diam., and shiny black at maturity.

CC ry

280 Gard. Bull. Singapore 60 (2) 2009

27. Livistona drudei F.Muell. ex Drude, Bot. Jahrb. Syst. 39 (1893) 12;

Mueller, Fragm. 11 (1878) 55, nom. provis.; Fragm. 11 (1878) 147, nom. in

index; Beccari, in Webbia 5 (1921) 15, 19. — Type: Australia. Queensland,

Rockingham Bay, Sep 1865, Dallachy s.n. (lectotype, MEL, fide Rodd,

1998). Plate 19A-D.

-Livistona inermis auct. non R.Br., Wendland & Drude, Linnaea 39 (1875)

229, pro parte.

Functionally dioecious palm. Trunk to 28 m tall, 15-20 cm dbh, leaf scars

narrow, pale grey; internodes broad. Leaves 30-60 in a globose or conical

crown; petiole 150-230 cm long, 15-25 mm wide, adaxially ridged, purple

in basal portion, green distally, margins with single, curved reddish spines

larger and closer in the proximal portion; leaf-base fibres moderately

prominent, coarse, persistent; lamina costapalmate, regularly segmented, +

circular, 100-150 cm long, coriaceous, adaxially dark green, abaxially lighter

_green, glossy; lamina divided for 60-70% of its length, with 60-84 segments,

depth of apical cleft ca 60% of the segment length, apical lobes attenuate,

pendulous; parallel veins 7 each side of midrib; transverse veins thinner

than parallel veins. Inflorescences unbranched at the base, not sexually

dimorphic, 150-300 cm long, not extending beyond the limit of the crown,

branched to 3 orders; 7-8 partial inflorescences; peduncle subterete, 15-25

mm diam.; prophyll glabrous; peduncular bract 1, tightly tubular, glabrous;

rachis bracts tightly tubular, glabrous, apices with acuminate lobes; rachillae

1-8 cm long, pubescent. Flowers solitary or in clusters of 2-5, funnel-shaped,

1.7-2.2 mm long, cream to yellow; sepals narrowly triangular, 1.3-1.5 mm

long, fleshy, subacute; petals broadly ovate, 1.7-2 mm long, fleshy, apex

thickened; stamens ca 1.5 mm long. Fruit globose to pyriform, 10-12 mm

long, 10-11 mm diam., semi-glossy purple-black; epicarp with scattered

lenticellular pores; suture line extends for ca 2 length of fruit, marked with

lip-like structures; mesocarp thin, fibrous; endocarp thin, brittle; pedicel 1-2

mm long. Seed ellipsoid, 8-9 mm long. Eophyll 5-ribbed.

Specimens examined: AUSTRALIA: Queensland. Kurrimine Beach, | km

N of Maria Ck, Dowe 28] (BRI, FTG); Tully Heads Rd, 1 km W of Hull

Heads turnoff, Dowe 290 (JCT); Rockingham Bay, Sept. 1865, Dallachy

s.n. (MEL lectotype); Hinchinbrook Is., Webb 1/170 (BRI); Herbert R., }

1886, Dittrich s.n. (MEL); 14.5 km NW of Ingham, N side of Herbert R., at |

SW base of Mt Leach Ra., Rodd 3135 with Jacobs (BRI, K, NSW); Mouth j

of Herbert R., Webb 11176 with Tracey (BRI); Braemeadows, near Ingham, :

27 Nov 1959, Thomas s.n. (BRI); Halifax, 2 km along road to Lucinda,

Irvine 1832 (BRI, NSW, QRS); Little Crystal Ck, near Youth Camp, 22

“i J

A Taxonomic Account of Livistona R.Br. (Arecaceae) 281

Jan 1982, Irvine s.n. (K); Kinduro, Hen Camp Ck, near Bruce Hwy, /rvine

1833 & 1834 (BRI, NSW, QRS); Kinduro, Hen Camp Ck, near Bruce Hwy,

Irvine 1841 (BRI, QRS); Conway Beach, 20 km from Proserpine, 20°28’S,

148°45’E, Dowe 329 (BRI, FTG).

Specimens from cultivated material: Australia: Queensland. Brisbane,

CSIRO, Long Pocket, Tracey 15264 (BRI); CSIRO, Long Pocket, Tracey

15296 (BRI).

Habitat and ecology: Grows on the landward side of mangroves, in fore-

dune swales, open forest and swamp forest on coastal plains and along the

margins of streams and estuaries at low elevations, 0-100 m alt. Flowers

Aug-Dec; fruits Dec-June.

Conservation status: Endangered (IUCN, 2006; Dowe, 2007).

Distribution: Australia. Queensland. Coastal and near-coastal areas between

Kurrimine Beach and Townsville, and with a disjunction to Conway Ra.

near Proserpine (Fig. 13).

Notes: Livistona drudei was first mentioned informally by Mueller (1878)

as “... palme nunc pro speciei vel varietatis distinctione nomen Doctoris

Oscaris Drude impono...”, followed by the use of the name L. drudei in the

index of the same volume. Drude (1893) reviewed this taxon and applied

a brief description, thus validating the name. However no specimens were

directly linked to either Mueller’s or Drude’s treatments, although Mueller

(1878) mentioned a specimen from ‘Rockingham’s Bay’ that Wendland

& Drude (1875) had used, incorrectly, as reference in their treatment of

L. inermis. The lectotype chosen by Rodd (1998) is the “Rockingham s

Bay” specimen Dallachy s.n.. The mention of the” valley of the Mackay”

on a slip accompanying the type is a reference to present-day Tully R..,

of which Mackay R. was an earlier name; a Mt Mackay still exists at this

location though. This does not refer to the present-day Mackay R., which is

associated with the city of Mackay in central Queensland.

Beccari (1931) provided a thorough description based on additional

specimens, but did not cite the Dallachy specimen. Rodd (1998) interpreted

Beccari’s (1931) description as formalising the name. However, as the

Dallachy specimen was not cited by Beccari, I have chosen Drude’s (1893)

treatment as the valid place of publication of the name.

Livistona drudei is a tall canopy palm to 28 m tall; leaves are large

and regularly segmented; segment apices are pendulous, and with a bifurcate

282 Gard. Bull. Singapore 60 (2) 2009

cleft to 60% of the segment length; the petiole bases are distinctively

purple; the inflorescence is unbranched, not extending beyond the limit of

the crown, and with up to 8 partial inflorescences; bracts are tightly tubular;

flowers are cream to yellow; fruit are globose to pyriform, to 12 mm long

by 11 mm wide, and semi-glossy purple black at maturity.

28. Livistona decora (W.Bull) Dowe, Austrobaileya 6 (2004) 979.

-Corypha decora W.Bull, Catal. (1887) 10; Watson, in Kew Bull. 3 (1889)

294. — Type: Australia. Queensland, Yepoon-Emu Park Rd, ca 11 km S of

Yepoon, 12 May 1976, Rodd 3069 with Jacobs (neotype, BRI; isoneotypes,

BH, K, NSW). Plate 20A-D.

-Livistona decipiens Becc., Webbia 3 (1910) 301, fig. 1, tav. 2. — Type:

France. Cultivation, Nice, May 1908, ‘sheet with older inflorescence’,

Robertson-Proschowsky (lectotype, FI; fide Rodd, 1998).

-Livistona inermis auct. non R.Br., Wendland & Drude, Linnaea 39 (1875)

229, pro parte.

_-Livistona australis auct. non (R.Br.) Mart., Bentham, Fl. Austral. 7 (1878)

146, pro parte.

-Livistona decipiens var. polyantha Becc., Webbia 5 (1921) 15, 18. — Type:

Indonesia. Cultivation, Bogor Botanic Gardens, undated, Beccari s.n.

(holotype, FI; isotype, BO).

Functionally dioecious palm. Trunk to 18 m tall, 25-30 cm dbh, leaf

scars narrow, internodes broad to 30 cm wide, smooth, brown in newly

exposed parts, grey with age, with shallow vertical fissures, petiole stubs

frequently retained at the base of the trunk. Leaves 30-60 in a + globose

crown; petiole 150-280 cm long, 18-24 mm wide, adaxially flat or slightly

concave, green throughout, margins with single curved black spines to 20

mm long in the proximal portion; leaf-base fibres moderately prominent,

coarse, disintegrating; lamina costapalmate, regularly segmented, + circular

in outline, 120-185 cm long, coriaceous, adaxially bright green, abaxially

lighter green, segments extended along the costa; lamina divided for 82-

88% of its length, with 70-84 segments, depth of apical cleft 44-54% of

the segment length, apical lobes pendulous; parallel veins 6-7 each side

of midrib; transverse veins thinner than parallel veins. Inflorescences

unbranched at the base, not sexually dimorphic, 100-350 cm long, not

extending beyond the limit of the crown, branched to 4 orders; 8-13 partial

inflorescences; longest partial inflorescence to 110 cm, each with a single

rachis bract; prophyll 37 cm long, 11-16 cm wide, woody; peduncular

bract(s) lacking; rachis bracts glabrous to sparsely papillate; primary rachis

bract loosely sheathing, apically fibrous-lacerate, bracts subtending partial

4 Tax : f Livistona R.Br (A 283

fi nat agra gee hr = ms ll 41} ar hecominc Se oe eS ane

inhiorescence Drancnes Dasa \ udula VELUILLIITI = ~OCU UilStall V apex

’ = ies Ee = a eer, ~t a (iy RS Ne

pointed: in the most distal partial inflorescence there are 2 vestigial bracts

at the base: peduncle more or less terete, 6 cm wide at the base, glabrous:

— : Ey | ~ 1] ~ ary - - + ~ F

rachillae 5-20 mm long, papillose. Flowers solitary o clusters of 2-6

obtuse: stamens ca 1.5 mm long. Fruit globose, 12-18 mm diam... shiny

black: epicarp with scattered lenticellular pores; suture line extends ca

length of fruit; mesocarp 1-3 mm thick, spongy, moist, fibres aggregated

toward the endocarp: endocarp thin, brittle: pedicel to 2 mm ong: Seed

globose, ca 10 mm wide: intrusion %-*% across endosperm. Eophyll

Specimens examined: AUSTRALIA: Queensland. Magnetic Is, Gustav

Ck, on track 1.5 km from Mandalay Rd. Dowe 250 & 251 (BRI, FTG):

Magnetic Is., Sandercoe 943 (BRI); Cape Cleveland, 1 km from eamacalh e

Ck towards AIN

Ck towards AIMS,

Hwy, Briggs 20 neat wy, Irvine 181

1818 (QRS): West es atta a Ck ruce soe between Ayr and ee Rodd

3117(NSW): East Barratta Ck. near Bruce Hwy, /rvine 1815 & 1816 (QRS):

N of St Margaret Ck, Bruce Hwy between Ayr and Tow Satie Moriarn

67 (CANB, BRI): 51 miles S of Townsville, adjacent to creek, Trapnell

305 (BRI, K); 27 miles N of Ayr, Bruce Hwy . Moriarty s.n. (BRI): Hom

Hill District, 1962, Wyatt s.n. (BRI); Cement Ck, tributary of Campaspe eS

Dowe 708 (BRI, JCT): 35 km NNE of oe ine, E of Dingo Beach Rd. Oo

Earlando Beach turnoff, Rodd 3077 with Jacobs (BRI. ee NSW); NE of! Mt

Dryander near coast, I5 Apr. 1978, Byrnes s.n. (BRI): e Hillsborough

Rd, 4.4 km from Seaforth Rd, Jrvine 1840 (BRI. QRS): nee Jane Ck,

Irvine 1812, 1813 & 1814 (QRS): Mackay, pasture lands, /rvine 1748

ES): | 10 km W of Mackay on bypass road, 75 (NSW); Mackay,

Jun 1970, V.K. Moriarty s.n. (BRI); Farleigh, TE 6 (BRI); Denham

Ra.., ee of Sandy Ck, 520 m alt., 21 5°40.6°S, 148°06°01.6"E,

Dowe 742 (BRI, JCT); Between Byfield and Maryvale, White 8197 (BRI):

North Keppel Is., NE of Yeppoon, Thomas 183 (BRI ); Yeppoon, Byfield,

Flecker 11493 (BRI, QRS): Yepoon-Emu Park Rd, ca 11 km S of Yepoon,

Rodd 3069 with Jacobs, (BH, BRI neotype, K, NSW); Yeppoon-Emu Park

Rd, 11 kmS . Yeppoon, Rodd 3070 with Jacobs (BRI, K); Yeppoon-Emu

Park Rd. ca 11 km S of Yeppoon, Rodd 3069 (BH, BRI. K. NSW): Near

Emu Park, Ps 19968 (BRI): Kinka Beach, Yepoon-Emu Park Rd, 2 km

] (FTG); Cape P25

284 Gard. Bull. Singapore 60 (2) 2009

S of Kinka town, 23°20’S, 150°50’E, Dowe 312b (FTG); Joskeleigh, 15

miles E of Rockhampton, Everist 7634 (BRI, K, L); (Queensland) Moore’s

Ck Ra., (pre-1875), Thozet s.n. (MEL); Palm Ck, 8 km S of Miriam Vale,

Hind 257 (BRI); Eurimbula, near Bustard Bay, Jul 1966, Cooper s.n. (BRI);

Just off Round Hill Head on inland of Peninsula, 27 Mar 1970, Everist s.n.

(BRI); Between Agnes Waters and Round Hill Head, S of Bustard Bay,

28 Mar. 1970, Everist s.n. (BRI, K); Agnes Waters, 40 km E of Miriam

Vale, McDonald 1224X & Batianoff (BRI); Wide Bay, Woodgate Reserve,

25°10°S, 152°30*, Dowe 31l & 312a(FVG, JCT); Reedy Ck Sikmese

of Lowmead on Bundaberg Rd, Rodd 3045 with Jacobs (BRI, K, NSW);

Cooloola SF, ENE of Gympie, Moriarty 668, 668A & 669 (BRI); Tewantin,

Blake 23786 (BRI, CANB, K).

Specimens from cultivated material: AUSTRALIA: Brisbane, Long

Pocket, CSIRO, Tracey 15295 (BRI); FRANCE: Nice, May 1908,

‘sheet with older inflorescence’, Robertson-Proschowsky (FI); Cannes,

Villa Valetta, as ‘Copernicia cerifera’, 12 Dec 1889, anon. s.n. (K);

INDONESIA: Java, Bogor Botanic Gardens, Garden No. XII.E.91,

Furtado 30948 (BRI, K); Java, Bogor Botanic Gardens, undated, Beccari

s.n. (BO, FI); SRI LANKA: Royal Botanic Gardens, Peradeniya, Q-291,

Rutherford 119 with Bandara (K); UNITED KINGDOM: Royal Botanic

Gardens Kew, 4 Dec 1889, “as Livistona inermis... being the true plant

of R.Br., Wendland s.n. (K)”.

Local and vernacular names: Weeping Cabbage Palm, Ribbon Fan Palm.

Habitat and ecology: Most common in coastal and near-coastal lowlands in

large dense colonies or in scattered groups, in open forest, littoral rainforest

and dry rainforest at low to moderate elevations, and with isolated inland

populations at moderate elevations associated with permanent springs or

seasonal streams, 0-550 m alt. Flowers Jul-Dec; fruits Dec-Apr.

Conservation status: Least concern.

Distribution: Australia. Queensland. From Magnetic Is. near Townsville,

south to Tewantin and Rainbow Beach (Fig. 13).

Notes: The name Corypha decora was first used by Bull (1887) for plants

introduced into horticulture from his nursery at Chelsea, and subsequently

discussed by Watson (1889) in a paper on the cool cultivation of tropical and

subtropical plants. Bull’s (1887, p.10) description was: “Corypha decora: An

A Taxonomic Account of Livistona R.Br. (Arecaceae) 285

elegant and ornamental species introduced from Queensland. It early develops

characterized leaves, which are fan-shaped in outline and divided almost to

the petiole into linear lanceolate segments each about half-an-inch in breadth.

The petioles are sparsely furnished with small hooked prickles. This species

will undoubtedly be found one of the most useful of greenhouse palms”.

Beccari (1910) was apparently unaware of the use of the name

Corypha decora when he named Livistona decipiens from the collection,

Robertson-Proschowsky s.n. taken from a cultivated plant at Nice, France,

and named for the deception that it had caused as to its true identity: “... i/

nome di L. decipiens, per gli errori di cui é stata causa....”. In cultivation in

the Riviera during the mid to late 1800s it had incorrectly been referred to by

local horticulturists as “Copernicia cerifera’, and it was with the intention

of clarifying the species’ identity that Beccari provided the description and

established the name. Although unaware of its origin, but suggesting that

it came from eastern tropical Australia, Beccari (1931) related it to other

Australian species, primarily L. australis. Despite it being well known

and widely cultivated throughout the world, its origin continued to remain

speculative until relatively recent times. For example, Bailey (1976)

noted (with reference to the name L. decipiens): “... described from cult.,

supposedly Australian....”. The first unequivocal application of the name

to natural populations was by Johnson (1981) who recorded distribution

from Miriamvale to Townsville, Queensland.

Through the lack of understanding of the true identity of L. inermis,

and the incorporation of misapplied specimens into descriptions, L. decora

was inadvertently identified as that species. Both Wendland and Drude

(1875) and Bentham (1878) cited the specimen Thozet s.n., from Moore’s

Ck near Rockhampton, central Queensland, now known to be of L.

decora in their accounts of L. inermis and L. australis respectively. Many

specimens in both the wild and in cultivation with a deeply segmented

leaf were subsequently identified as L. inermis sensu H.Wendl. (Hill, 1873,

1875). It is highly probable that L. inermis R.Br. was never in cultivation

in Europe as it is an exceedingly difficult plant to propagate and maintain

while L. decora is exceptionally easy to germinate, and one of the fastest

growing species of Livistona.

Beccari (1921) described a variety, L. decipiens var. polyantha,

from plants cultivated in Bogor Botanic Gardens based on Beccari s.n., but

this is attributable to L. decora.

Livistona decora is most closely related to L. australis, but is readily

distinguished by the deeply segmented leaves and pendulous segment

apices. The range of these species overlaps in the Fraser Is./Rainbow

Beach area of southeastern Queensland, and it may be that hybridisation

286 Gard. Bull. Singapore 60 (2) 2009

has occurred as some individuals in that area are difficult to assign to either

species.

Livistona decora is a large canopy palm to 18 m tall; leaves are

large and regularly segmented; segment apices are pendulous, and with a

bifurcate cleft to 54% of the segment length; the inflorescence is unbranched,

not extending beyond the limit of the crown, and with up to 13 partial

inflorescences; bracts are loosely sheathing; flowers are yellow; fruit are

globose to 18 mm diam., and shiny black at maturity.

29. Livistona australis (R.Br.) Mart., Hist. Nat. Palm. 3(7) (1838) 242.

-Corypha australis R.Br., Prodr. (1810) 267. -Saribus australis (R.Br.)

Kuntze, Revis. Gen. Pl. 2 (1891) 736. — Type: Australia. New South Wales,

Port Jackson, undated, Brown s.n. (holotype, BM; isotype, FI). Plate

21A-E.

-Livistona inermis auct. non R.Br., Wendland & Drude, Linnaea 39 (1875)

229.

Functionally dioecious palm. Trunk to 25 m tall, 25-40 cm dbh, leaf scars

raised, internodes broad, brown, longitudinal fissures prominent, petiole

stubs retained in the lower 2 m or so, otherwise deciduous. Leaves 35-60

in a globose crown; petiole 150-250 cm long, 10-20 mm wide, adaxially

moderately ridged, margins smooth or with short single black curved spines

congested in the proximal portion, a thin yellowish-green longitudinal

stripe present on the abaxial surface; leaf-base fibres prominent, finely

woven, sheet-like, persistent; lamina costapalmate, regularly segmented,

+ circular, 100-130 cm long, chartaceous, adaxially dark green, glossy,

abaxially slightly lighter green, nonwaxy; lamina divided for 49-69% of its

length, with 80-100 segments, depth of apical cleft 50-63% of the segment

length, apical lobes acute, pendulous; parallel veins 8 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, not sexually dimorphic, 140-250 cm long, not extending beyond

the limit of the crown, branched to 5 orders; partial inflorescences 6-9;

peduncular bract(s) lacking; rachis bracts loosely sheathing, densely

floccose; rachillae 5-25 mm long, glabrous. Flowers solitary or in clusters

of 2-4, funnel-shaped, white to cream to pale yellow; sepals triangular,

1.2-2 mm long, fleshy, acute; petals triangular to ovate, 2-3.5 mm long,

fleshy, subacute to obtuse; stamens ca 2.5 mm long. Fruit globose, 12-

22 mm diam., dull reddish-brown to black, occasionally glaucous; epicarp

with scattered lenticellular pores; suture line extends almost the length of

the fruit; pedicel 2-3 mm long. Seed globose, 10-16 mm wide. Eophyll

5-ribbed.

A Taxonomic Account of Livistona R.Br. (Arecaceae) 287

Specimens examined: AUSTRALIA: Queensland. Paluma Ra., 6 km NW

of Paluma Dam on forestry road, Dowe 29/ (JCT); Paluma Ra., Dotswood

Holding, /rvine 907 (QRS); Paluma Ra., Dotswood Holding, 22 km W of

Bluewater Forestry Rd, 19°13’S, 146°25’E, 620 m alt., Irvine 1928, 1929

& 1930 (K, MEL, NSW, QRS); Paluma Ra., Dotswood Holding, 22 km

W of Bluewater Forestry Rd, 19°13’S, 146°25’E, 620 m alt., /rvine 1929

(K); Bluewater Ck Forest Rd, 22 km from Bruce Hwy, Rodd 3758 with P.

Lavarack (NSW); Bluewater Ra., 0.5 km past SF sign, Dowe 171 (JCT):

Bluewater Ra., | km past Rangers Hut, Dowe 520 (JCT); Cathu State Forest,

Clarke Ra., North Rd, 16 km along road to Palm Tree Lookout, 20°45’S,

138°35’E, 800 m alt., Dowe 3/3 (BRI, FTG, JCT); Eungella Ra., SFR 679,

Irvine 1839 (QRS); Broken R. NP, Webb 3322 (BRI): Eungella Ra., ca

2 km SE of Eungella on Broken R. Rd, Rodd 3073 with Jacobs (NSW);

Eungella Ra., ca 3.5 km S of Broken R. crossing, S of Eungella, Rodd 3798

& 3799 (BRI); Fraser Is, E side inland from Happy Valley, Baxter 90]

(BRI); Tin Can Bay, 20 Oct 1946, Clemens s.n. (A, K); 13 km E of Cooroy

on Tewantin Rd, Rodd 3030 & 3031 with Jacobs (BRI, K, NSW, QRS);

Nambour, Tuckers Ck, /rvine 1893, 1894, 1895 & 1896 (QRS): 4.8 km N of

Nambour, Moriarty 671 & 672 (BRI); Forest Glen, 2 km SW of Buderim,

26°48°S, 153°00°E, Dowe 310 (FTG); 18 km from Landsborough towards

Nambour, Carroll 1140 & Telford (CANB); Yandina, 1 Mar 1891, anon.

s.n. (BRI); Mooloolaba, 70 miles N of Brisbane, Moore 9229 with Everist

(BRI); Between Mt Glorious and Mt Nebo, Blake 21484 (BRI); Cabbage

Tree Ck, | Mar 1888, Simmonds s.n. (BRI). New South Wales. Seal Rocks,

32 km E of Bulahdelah, 1 Aug 1964, Briggs s.n. (NSW); N of Ferny Ck, W

of Wallis Ck, Salasoo 3322 (NSW); Warrah, Pearl Beach, Rodd 2672 (BRI,

NSW, QRS); Bayview, Cabbage Tree Rd, W of golf course, Rodd 3615

(NSW); Myall Lakes, Aug 1934, Osborn s.n. (K); Kuring-gai NP, McCarrs

Ck, Rodd 2671 (NSW); Royal NP, above Garie Beach, 34°10’S, 151°03’E,

60 malt., 28 Aug. 1975, Rodd s.n. (K, NSW, QRS); Upper Kangaroo Valley,

Rodd 2398 (NSW); Port Jackson, undated, Brown s.n. (BM holotype, FI);

near Sydney, anonymous s.n. (MEL); Botany Bay, Kurnell, May 1906,

Boorman s.n. (NSW); near Mt Dromedary, 1881, Bates s.n. (MEL);

(Australia), Nov 1806, Caley s.n. (BM); (New South Wales), 1864, Daemel

s.n. (MEL); (New South Wales), Johnson s.n. (MEL); Kiama, 1872, Henry

s.n. (MEL); Illawarra, Kirton s.n. (MEL); Mt Cambewarra, NW of Nowra,

7 Dec 1950, Constable s.n. (NSW); 13 km from West Bodalla, Evans 96

(NSW); Tanja, ENE of Bega, Jul 1954, anon. s.n. (NSW); Ash Is., 25 Nov

1842, Leichhardt s.n. (MEL); (Australia), undated, Lhotsky s.n. (BM);

Royal NP, Meebold 2631] (M). Victoria. Gippsland, Cabbage Tree Ck, near

Marlo, Webb CBG010319 (CANB, NSW); Palm Ck, S of Cabbage Tree,

EEE

288 Gard. Bull. Singapore 60 (2) 2009

Beauglehole 34001 with Beaton (MEL); Palms Reserve, 5.4 km S from

Princes Hwy along Palm Track on Palm Ck, 37°44’45”S, 148°39E’, 40

m alt., Forbes 3222 with Worboys (MEL); East Gippsland, 1888, French

s.n. (MEL); East Gippsland, 1889, French s.n. (MEL); East Gippsland, Jan

1889, Searle s.n. (MEL); East Gippsland, Weindorfer 2396 (MEL); NE of

Marlo, Brodribb R., Curlip Jungle, Melville 2933 with Wakefield & Hunter

(K); 16 km E of Marlo, Cabbage Tree Ck on Cabbage Palms Track, Carroll

264 (CANB).

Specimens from cultivated material: AUSTRALIA: Queensland, Brisbane

Botanic Gardens, White 7764 (A, BRI, K); UNITED KINGDOM: Royal

Botanic Gardens Kew, 1891, anon. s.n. (K); Royal Botanic Gardens Kew,

Feb 1862, anon. s.n. (K); Royal Botanic Gardens Kew, Temperate House,

Dransfield 392-26-23904 (IK); INDIA: Madras, 4 Jun 1900, Bourne s.n.

(K); Locality not provided, Mar 1900, Kerchove de Deterghem s.n. (K).

Local and vernacular names: Cabbage Palm, Cabbage Tree.

Habitat and ecology: Grows in moist areas of open forest, swamp forest,

moist sclerophyll forests, along stream banks and in rainforest, 0-1000 m

alt. Often locally common and may grow in large colonies. In the north

confined to high elevation above 400 m and in central and southern parts in

coastal and near-coastal lowland areas. Flowers all year; fruits all year.

Conservation status: Least concern.

Distribution: Australia. Queensland, New South Wales and Victoria. In

the northern part of its range in Queensland confined to high elevation on

Paluma Ra., Seaview Ra., Mt Elliot, Mt Abbott, and Eungella Ra. with a

major disjunction to Fraser Is., and then more or less continuously at low

to moderate elevation south through south-eastern Queensland, through

coastal New South Wales and with a minor disjunction to the Brodribb R.

in eastern Victoria (Fig. 14).

Notes: Brown (1810) described L. australis, as Corypha australis, from a

specimen that he collected from Port Jackson, New South Wales, and which

he distinguished from his Livistona species by the character of embryo

position. Martius (1838) was the first to place it in Livistona. Previously, \

Solander (1768-1771) had included the species as another manuscript

name in his hand-written and unpublished Plantae Novae Hollandiae,

based on plants he saw at Botany Bay, the specimens of which are located

A Taxonomic Account of Livistona R.Br. (Arecaceae) 289

110°E 120°E 130°E 140°E 150°E

10°S

20°S

30°S

Legend

L. alfredii

o @ L. australis

4 L. fulva

A L. nitida

110°E 120°E 130°E 140°E 150°E

alfredii (2).

in BM. This species is the first palm to be mentioned in European accounts

of Australia. Banks (Beaglehole, 1962) noted during Cook’s Endeavour

voyage of the Australian coast in 1770 in southern New South Wales: *

trees were not very large and stood separate from each other without the

least underwood; among them, we could discern many cabbage trees, but

nothing else which we could call by any name...”. These “cabbage trees’

refer to L. australis.

Livistona australis is a tall canopy palm to 25 m tall; leaves are

large and regularly segmented; segment apices are pendulous, and with

a bifurcate cleft to 63% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to

9 partial inflorescences; bracts are loosely sheathing; flowers are white to

cream to pale yellow; fruit are globose to 22 mm diam., and dull reddish

brown to black at maturity.

30. Livistona nitida A.N.Rodd, Telopea 8 (1998) 96, figs 2a-b, 3h, 8d,

9b-c. — Type: Australia. Queensland, NW of Cracow, junction of Dawson

—_<$ re

290 Gard. Bull. Singapore 60 (2) 2009

R. and Delusion Ck, 9 May 1976, Rodd 3055 with Jacobs (holotype, NSW;

isotypes, BH, BRI, K). Plate 22A-C.

Functionally dioecious palm. Trunk to 35 m tall, 25-40 cm dbh, leaf scars

raised; internodes narrow, grey; petiole stubs persistent in the basal | m or

so, otherwise deciduous. Leaves 35-50 in a globose crown; petiole 170-

200 cm long, 20-26 mm wide, adaxially ridged, margins distally smooth,

proximally with single, curved, dark red spines; leaf-base fibres moderately

prominent, coarse, disintegrating; laminacostapalmate, regularly segmented,

+ circular in outline, 160-190 cm long, coriaceous, adaxially dark green,

glossy, abaxially lighter green; lamina divided for 63-70% of its length,

with 68-80 segments, depth of apical cleft 60-73% of the segment length,

apical lobes pendulous; parallel veins ca 7 each side of midrib; transverse

veins thinner than parallel veins. Inflorescences unbranched at the base,

not sexually dimorphic, 150-200 cm long, not extending beyond the limit

of the crown, branched to 4 orders; partial inflorescences 8-12; peduncular

-bract(s) lacking; rachis bracts loosely sheathing, densely scaly; rachillae

5-20 cm long, glabrous to papillose. Flowers solitary or in clusters of 2-5,

funnel-shaped, 2-3.2 mm long, cream to yellow; sepals triangular, ca 1.5

mm long, fleshy, subacute; petals broadly ovate, 2.0-2.2 mm long, thick,

fleshy, acute; stamens ca 1.6 mm long. Fruit globose, 13-20 mm diam.,

glossy jet black; epicarp with scattered lenticellular pores, suture line

extends for ca 2 the length of the fruit, marked with lip-like structures;

mesocarp fibrous, dry; endocarp thin, brittle; pedicel to 0.5 mm long. Seed

globose, 10-12 mm wide. Eophyll 5-ribbed.

Specimens examined: AUSTRALIA: Queensland. Carnarvon Gorge, White

11374 (BRI); Carnarvon Gorge, Dec 1939, Waldron s.n. (BRI); Carnarvon

Gorge NP, Carnarvon Ck, Rodd 1/112 (NSW); Delusion Ck, Cracow-

Theodore Rd, 12 miles N of Cracow, Johnson 847 (BRI); Delusion Ck,

Theodore-Cracow Rd, 25°11’S, 150°11’E, Dowe 30/ (BRI, FTG); Junction

of Dawson R. and Delusion Ck, NW of Cracow, Rodd 3051, 3052 & 3054

with Jacobs (BH, BRI, K, NSW); NW of Cracow, junction of Dawson R.

and Delusion Ck, Rodd 3055 with Jacobs (BH, BRI, K, NSW holotype);

Cracow to Theodore Rd, Delusion Ck, 17.5 km W of Cracow, /rvine

1898, 1899, 1900, 1901 & 1902 (QRS); 61 km W of Eidsvold at start of

Isla-Delusion Rd, 21 Apr 1986, Ladiges, Humphries & Bean s.n. (BM);

Robinson Ck, NW of Taroom, Speck 1986 (BRI, CANB); Robinson Ck,

13 miles NW of Taroom, Johnson 2764 (BRI); Palm Tree Ck, 12 miles

N of Taroom, Johnson 1671 (BRI); Palm Tree Ck, 12 km from Taroom,

Moriarty 1280 (BRI); Palm Tree Ck, 19 km N of Taroom, Telford 5648

A Taxonomic Account of Livistona R.Br. (Arecaceae) 29]

(CBG, NSW): Palm Tree Ck, 10 miles (ca 16 km) N of Taroom, Reeve //

(CANB); Taroom, 16 Sep 1960, Stiller s.n. (BRI).

Local and vernacular names: Carnarvon Gorge Cabbage Palm. Dawson

River Fan-Palm.

Habitat and ecology: Abundant along the margins of streams and flood

plains, but also associated with sandstone cliffs and gorges, 100-650 m alt.

Flowers Sep-Dec; fruits Nov-Mar.

Conservation status: Lower risk, conservation dependent.

Distribution: Australia. Queensland. In the catchments of the Dawson,

Comet and Burnett Rivers, and Carnarvon Gorge (Fig. 14).

Notes: Livistona nitida was described by Rodd (1998) based on Rodd 3055

collected from Delusion Ck near Cracow, Queensland, and named for its

highly glossy jet black fruits. The taxon had previously been informally

known as the ‘Carnarvon Gorge Fan Palm’ and ‘Livistona sp. Carnarvon’

(Irvine, 1984; Jones, 1996). An early account was provided by Leichhardt

(1847) during his transcontinental expedition of 1845, when near the

Dawson R., Queensland, he wrote in his diary: “....a creek with Corypha

palms, growing to a height of 25 or 30 feet” and “several rocky gullies were

passed, that were full of palm trees. The valley of Palm-tree Creek extends

about nineteen miles from west to east” and “the Corypha-palm provided a

good supply of cabbage’’.

Livistona nitida is a large canopy palm to 35 m tall; leaves are

large and regularly segmented; segment apices are pendulous, and with a

bifurcate cleft to 73% of the segment length: the inflorescence is unbranched,

not extending beyond the limit of the crown, and with up to 12 partial

inflorescences; bracts are loosely sheathing; flowers are cream to yellow;

fruit are globose to 20 mm diam., and glossy jet black at maturity.

31. Livistona fulva A.N.Rodd, Telopea 8 (1998) 103, figs 4a, 12, 13a. —

Type: Australia. Queensland, Blackdown Tableland, 11 May 1976, Rodd

3062 & Jacobs (holotype, NSW; isotypes, BH, BRI, K). Plate 22D-E.

Functionally dioecious palm. Trunk to 13 m tall, 20-25 cm dbh, leaf scars

raised, internodes narrow, grey or brown, petiole stubs deciduous. Leaves

25-35 in a globose crown; petiole 150-250 cm long, 12-15 mm wide,

adaxially moderately ridged, basal margins armed with single, curved

EEE

292 Gard. Bull. Singapore 60 (2) 2009

black spines; leaf-base fibres moderately prominent, fine, persistent; lamina

costapalmate, regularly segmented, circular in outline, 90-100 cm long,

coriaceous, adaxially greyish green to glaucous, coppery brown floccose

tomentum abaxially; lamina divided for 50-55% of its length, with 60-66

segments, depth of apical cleft 3-5% of the segment length, apical lobes

acute to acuminate, rigid; parallel veins 8-9 each side of midrib; transverse

veins thinner than parallel veins. Inflorescences unbranched at the base,

not sexually dimorphic, 100-230 cm long, not extending beyond the limit

of the crown, branched to 4 orders; partial inflorescences 7-9; peduncular

bract 1, loosely sheathing, with orange-brown scales; rachis bracts loosely

sheathing, with orange-brown scales; rachillae 5-16 cm long, papillose.

Flowers solitary or in clusters of 2-3, funnel-shaped, yellow, 1.6-2 mm

long; sepals narrowly triangular, 1.0-1.3 mm long, membranous, acute;

petals broadly ovate, 1.6-2 mm long, thick, acute; stamens ca. 1.6 mm

long. Fruit globose, 12-16 mm diam., pruinose dull black; epicarp smooth;

suture line not obvious; mesocarp thin, fibrous; endocarp thin, crustaceous.

Seed globose, 10-13 mm wide. Eophyll 5-ribbed.

Specimens examined: AUSTRALIA: Queensland. Blackdown Tableland,

Rodd 3062 & Jacobs (BH, BRI, K, NSW holotype); slopes below N

escarpment, Blaxell 921 with Johnson (NSW); Blackdown Tableland,

Irvine 1903, 1904, 1905 & 1906 (QRS); N escarpment, Jones 6338 &

Jones (CANB); Mimosa Ck, 23°05’S, 149°00’E, Henderson 1180 (BRI,

K, NSW); 32 km SE of Blackwater, Henderson 876 (BRI); ca 35 km SE

of Blackwater, 23°05’S, 149°00°E, Durrington & Sharpe 1180 (MEL); just

below escarpment at NE end (roadpass), Rodd 3063, 3064 & 3065 (BH,

BRI, K, NSW); below escarpment, 2 km from top, 23°43’S, 149°07°E,

Dowe 300 (FTG).

Local and vernacular name: Blackdown Fan Palm.

Habitat and ecology: Grows in moist sites in open forest and woodland,

in gullies and gorges near streams and waterfalls at cliff bases; confined to

sandstone areas, 400-900 m alt. Flowers Sep— Feb; fruits Dec—May.

Conservation status: Lower risk, conservation dependent.

Distribution: Australia. Queensland. Endemic to Blackdown Tableland

(Fig. 14).

Notes: Livistona fulva was described by Rodd (1998) based on Rodd 3062

: are

A Taxonomic Account of Livistona R.Br. (Arecaceae) 293

from Blackdown Tableland, and named for the coppery brown floccose

tomentum on the abaxial surface of the leaf. The relationships of L. fulva

are unclear, but it is most similar to L. muel/leri in having a flat rigid lamina,

to L. decora in inflorescence size and morphology, and to L. victoriae in

overall size and fruit morphology.

Livistona fulva is a moderate sub-canopy palm to 13 m tall: leaves

are moderate and regularly segmented, and with coppery brown floccose

tomentum abaxially; segment apices are rigid, and with a bifurcate cleft

to only 5% of the segment length; the inflorescence is unbranched,

not extending beyond the limit of the crown, and with up to 9 partial

inflorescences; bracts are loosely sheathing; flowers are yellow; fruit are

globose to 16 mm diam., and pruinose dull black at maturity.

32. Livistona alfredii F.Muell., Victorian Naturalist 9 (1892) 112 (as L.

alfredi); Bot. Centralbl. 53 (1893) 28, verbatim exemplar. —Type: Australia.

Western Australia, Hamersley Ra., Millstream, June 1878, Forrest s.n.

(holotype, MEL). Plate 23A-B.

-Livistona mariae auct. non F.Muell., Mueller, Fragm. 11 (1878) 54.

-Livistona mariae auct. non. F.Muell., Gardner, Forest Depart. Bull. Western

Australia 32 (1923) 36.

Functionally dioecious palm. Trunk to 12 m tall, 20-50 cm dbh, leaf scars

prominent, internodes narrow, pale grey, petiole stubs persistent in basal

1 m, otherwise deciduous. Leaves 25-30 in a globose crown; petiole 90-

130 cm long, 25-30 mm wide, adaxially flat, margins with curved single

black spines congested in the proximal portion; leaf-base fibres prominent,

coarse, persistent; lamina costapalmate, regularly segmented, subcircular

in outline, 90-140 cm long, rigid, adaxially pale green-grey to glaucous,

waxy, dull; abaxially light green-grey, waxy; lamina divided for 60-70% of

its length, with 50-66 segments, depth of apical cleft 60-75% of the segment

length, apical lobes attenuate, rigid; parallel veins 6-8 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, not sexually dimorphic, 80-270 cm long, not extending beyond

the limit of the crown, branched to 3 orders; partial inflorescences 5-7;

prophyll not seen; peduncular bracts 1-2, loosely sheathing, sparsely to

moderately scaly; rachis bracts loosely sheathing, sparsely to moderately

scaly; rachillae to 13 cm long, to 2 mm diameter, white pruinose, papillose.

Flowers solitary or in pairs, cylindrical in bud; sepals triangular, 0.8-1

mm long, acute, cream to yellowish; petals triangular, 2-3 mm long, acute

to mucronate, cream to yellowish; stamens ca 2 mm long. Fruit globose,

25-40 mm diam., dark brown to black; epicarp with scattered lenticellular

294 Gard. Bull. Singapore 60 (2) 2009

pores; suture line extends to ca ’2 way to base; pedicel to 3 mm long. Seed

globose, 17-20 mm wide. Eophyll 3-ribbed.

Specimens examined: AUSTRALIA: Western Australia. Hamersley Ra.,

Millstream, Jun 1878, Forrest s.n. (MEL holotype); On the Mill-stream,

Fortescue R., Forrest 301 (MEL); Hamersley Ra., 19 Feb 1879, Forrest

s.n. (MEL); Millstream, Hamersley Ra., Forrest s.n. (MEL); Millstream,

Hamersley Ra., 1892, anon. s.n. (MEL); Millstream R., 15 Oct 1915, Thorpe

s.n. (K); Longreach, Millstream, Fortescue R., 21 Aug 1932, Gardner s.n.

(PERTH); Pilbara, Millstream-Chichester NP, Fortescue R., Crossing Pool,

20°50’S, 116°35’E, Dowe 354 (BRI); Pilbara, Millstream-Chichester NP,

Fortescue R., Crossing Pool, 1 km downstream of Dawson Ck junction,

Dowe 355 (JCT); Millstream-Chichester NP, 28 Jul 1990, Leyland s.n.

(PERTH); Miullstream-Chichester NP, Fortescue R., below lookout

overlooking Crossing Pool, 21°33’12”S, 116°58°5”E, Bromilow LA2, LA5

(BRI, PERTH); Millstream Stn, 21°35’S, 117°04’E, 300 m alt, Rodd 2847,

2849 & 2850 (BH, K, NSW, PERTH); Fortescue R., Millstream, Aug 1974,

Beauglehole s.n. (PERTH); Fortescue R., Brooker 2064 (PERTH); Fortescue

R., Millstream, George 3523 (PERTH); Fortescue R., Crossing Pool near

Millstream Homestead, 13 Aug 1974, Willis s.n. (MEL); Millstream R.,

Brooker 2064 (PERTH); Millstream/Yarraloola Rd crossing the Fortescue

R., 21°37°58”S, 117°07’08”E, Bromilow LA] (BRI, PERTH); Fortescue R..,

400 m E of Robe R. iron rail bridge, 21°28°35”S, 116°49°14”E, Bromilow

LA6 (BRI, PERTH); ca 32 km W of Mt Brockman Stn, Duck Ck, 3 May

1975, Wark A & B (NSW); Nichol Bay, 1879, McRae s.n. (MEL); Cape

Ra., North West Cape, 22°22’09”S, 113°54’03”E, 29 Sept. 1988, Waldock

sn (PERGE):

Local and vernacular name: Millstream Palm.

Habitat and ecology: Grows adjacent to watercourses and along drainage

lines in well-drained sites, 50-560 m alt. Flowers Sep—Jan; fruits Dec-

May.

Conservation status: Lower risk, conservation dependent (IUCN, 2006).

Distribution: Western Australia. In the Hamersley Region mainly in the

upper reaches of the Fortescue, Robe and Ashburton Rivers. A small

disjunct and apparently senescent population occurs in the Cape Ra., North

West Cape Peninsula, on the western side of Exmouth Gulf (Humphreys er

al., 1990) (Fig. 14).

A Taxonomic Account of Livistona R.Br. (Arecaceae) 295

Notes: Specimens relating to L. alfredii were first cited by Mueller (1878) as

part of the distribution of L. mariae. The protologue of L. alfredii (Mueller,

1892) was an informal account in Victorian Naturalist. It included some

manner of distinguishing it from L. mariae, and discussed leaf colour and

fruit size. Mueller named it in honour of Alfred, Duke of Edinburgh. The

type, Forrest s.n. is mentioned in the account of L. mariae. Drude (1893)

found L. a/fredii distinct enough to place it in its own section, Gregorya.

Beccari (1921) similarly provided only a meagre description, but in

1931 provided a complete description based on the Forrest and McRae

specimens.

Livistona alfredii is the most geographically isolated species of

Livistona in Australia.

Livistona alfredii is a moderate canopy palm to 12 m tall; leaves

are large and regularly segmented; segment apices are rigid, and with a

bifurcate cleft to 75% of the segment length; the inflorescence is unbranched,

not extending beyond the limit of the crown, and with up to 7 partial

inflorescences; bracts are loosely sheathing; flowers are cream to yellow;

fruit are globose to 40 mm diam., and dark brown to black at maturity.

33. Livistona nasmophila Dowe & D.L.Jones, Austrobaileya 6 (2004) 980.

-Livistona mariae subsp. occidentalis A.N.Rodd, Telopea 8 (1998) 81, figs

3c, 5d. — Type: Australia. Western Australia, Durack Ra., SE base of Mt

King, Rodd 2868 (holotype, NSW; isotypes, BH, K, PERTH, QRS). Plate

23C-E.

Functionally dioecious palm. Trunk to 30 m tall, 30-65 cm dbh, leaf

scars raised, internodes broad, grey, petiole stubs persistent in basal | m

or so, otherwise deciduous. Leaves 35-55 in a globose crown; petiole

165-230 cm long, 4-10 cm wide, adaxially flat, margins with single or

double retrorsely recurved reddish spines confined to the proximal portion,

margin distally smooth; leaf-base fibres moderately prominent, coarse,

persistent; lamina costapalmate, regularly segmented, circular, 130-175 cm

diam., undulate, rigidly coriaceous, adaxially glossy grey-green, abaxially

dull grey to glaucous; lamina divided for ca 48% of its length, with 52-

58 segments, depth of apical cleft ca 48% of the segment length, apical

lobes at first rigid, becoming semi-pendulous with age or damage; parallel

veins 7-9 each side of midrib; transverse veins thinner than parallel veins.

Inflorescences unbranched at the base, not sexually dimorphic, 260-300

cm long, not extending beyond the limit of the crown, branched to 5 orders;

partial inflorescences 9-11; peduncular bracts | or lacking; rachis bracts

loosely tubular, with scattered long scales, becoming fully glabrous with

EE re

296 Gard. Bull. Singapore 60 (2) 2009

age, not disintegrating or becoming marginally lacerate; rachillae 5-9 cm

long, glabrous. Flowers in clusters of 4-6, globose, angular in bud, 1.4-1.5

mm long, cream to yellow; sepals broadly ovate, ca 1.2 mm long, thin, acute;

petals triangular, 1.4-1.5 mm long, obtuse; stamens ca 0.9 mm long. Fruit

globose, 11-14 mm diam., purple-black; epicarp with scattered lenticellular

pores; mesocarp fibrous, dry; endocarp thin, crustaceous; pedicel to | mm

long. Seed globose, 7-11 mm wide. Eophyll not seen.

Specimens examined: AUSTRALIA: Western Australia. Northeast

Kimberleys, Bens Springs, 3 km E of El Questro Homestead, 16°01’S,

128°01°E, Kenneally 10965 (PERTH, NSW); Cambridge Gulf, El Questro

Stn, Zebedee Springs, Dowe 363 (BRI, FTG); Durack Ra., SW base of Mt

King, 17°20’S, 127°23’E, Rodd 2869 (PERTH); Durack Ra., SE base of Mt

King, Rodd 2868 (BH, K, NSW holotype, PERTH, QRS); King Leopold

Ranges, Mt Ord Gorge, 20 miles SSW of Mt House Stn, Lazarides 6453

(CANB, K, MEL); King Leopold Ranges, southern end of Eva Ck Gorge,

Sands 4620 & 4630 (K); SE Kimberleys, base of Osmond Ra. on Swamp

Ck, 0.8 km N of Samim mining camp, Forbes 25353 (MEL, PERTH).

Specimens from cultivated material: AUSTRALIA: Queensland, Townsville

Palmetum (ex seed from King Leopold Ranges, Mt Gladys, May 1987,

Willing s.n.), Dowe 117 (NSW).

Habitat and ecology: Along intermittent or permanent water courses, in

open forest, occurs in very large colonies in some areas, 50-200 m alt.

Flowers Jul-Oct; fruits Oct-Dec.

Conservation status: Lower risk, conservation dependent.

Distribution: Western Australia. In the Durack Ra. and Cambridge Gulf

area (Fig. 15).

Notes: Rodd (1998) established this taxon as L. mariae subsp. occidentalis,

based on the collection Rodd 2868, from Mt King, Western Australia. In

the protologue, Rodd wrote: “Recognition of this population as a separate

subspecies on the basis of a single wild collection (and one from cultivation)

is arguably rather premature”. Following subsequent fieldwork in the

Kimberleys, with collections of fruit and flowers from both wild and

cultivated sources, it became apparent that this taxon was distinct enough

from both L. mariae and L. rigida to be raised to species rank. However,

the correct nomenclatural procedure would have been to take Rodd’s

- CT osxyic ole wi

A Taxonomic Account of Livistona R.Br. (Arecaceae) 297

ios]

p»

(

”

= =

” =

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2 | @ L mariae ; Pe *%

¢ L. nasmophila c as es Lie ® ae re

4 L figida ==, a

L we

T T T = =

110°E 120°E 130°E 140°E 150°E

L. lanuginosa (3).

varietal name ‘occidentalis’ and use it as the specific epithet, but the name

‘Livistona occidentalis’ had previously been used by Hooker (1884) as a

synonym for Brahea dulcis Mart. and was therefore unavailable. The new

name, ‘nasmophila’ was chosen to illustrate the palm’s habit of occupying

permanent watercourses fed by springs through much of its range.

Livistona nasmophila is a large canopy palm to 30 m tall; leaves are

large and regularly segmented; segment apices are rigid to semi-pendulous,

and with a bifurcate cleft to 48% of the segment length; the inflorescence

is unbranched, not extending beyond the limit of the crown, and with up to

11 partial inflorescences; bracts are loosely tubular; flowers are cream to

yellow; fruit are globose to 14 mm diam., and purple black at maturity.

34. Livistona mariae F.Muell., Fragm. 11 (1878) 54; Fragm. 8 (1874) 283,

nom. prov.; Giles, Geo. Trav. Central Austral. (1875) 222, nom. ined. -Saribus

mariae (F.Muell.) Kuntze, Revis. Gen. Pl. 2 (1891) 736. -Livistona mariae

subsp. mariae A.N.Rodd, Telopea 8 (1998) 80, figs 3a, 5a. — Type: Australia.

Northern Territory, Macdonnell Ranges, undated, Giles s.n. (holotype, MEL

67699, 67700, 67701; fide Rodd, 1998, in part). Plate 24A.

EEE

298 Gard. Bull. Singapore 60 (2) 2009

Functionally dioecious palm. Trunk to 30 m tall, 30-40 cm dbh, leaf scars

stepped, raised, internodes broad, grey, petiole stubs persistent only in the

basal | m or so. Leaves 30-50 in a globose crown; petiole erect to arching,

150-250 cm long, 20-45 mm wide, adaxially ridged, margins with small,

single, curved, black spines in the proximal portion, smooth distally; leaf-

base fibres moderately prominent, coarse, persistent; lamina costapalmate,

regularly segmented, circular in outline, 100-220 cm diam, rigidly

coriaceous, adaxially grey green, glossy, abaxially lighter grey green, waxy

pruinose; lamina divided for 45-55% of its length, with 50-86 segments,

depth of apical cleft 45-65% of the segment length, apical lobes attenuate,

pendulous, proximal margins of outer segments with small spines; parallel

veins 7-9 each side of the midrib; transverse veins thinner than parallel

veins. Inflorescences unbranched at the base, not sexually dimorphic, 125-

250 cm long, not extending beyond the limit of the crown, branched to

4 orders; partial inflorescences 10-14; peduncular bract 1, loosely tubular

with dense white-grey scales; rachis bracts loosely tubular with dense

white scales; rachillae 3-8 cm long, pliable to flexuose, glabrous. Flowers

in clusters of 3-6, campanulate, 1.0-1.8 mm long, greenish cream to yellow;

sepals ovate, 0.8-1 mm long, membranous, bluntly acute; petals broadly

oblong, 1.2-1.5 mm long, acute; stamens ca 1.2 mm long. Fruit globose,

12-18 mm diam., semi-glossy black; epicarp with scattered lenticellular

pores; suture line extends for ca % the length of the fruit, marked with lip-

like structures; mesocarp fibrous; endocarp thin, crustaceous. Seed globose,

8-12 mm wide. Eophyll 3-ribbed.

Specimens examined: AUSTRALIA: Northern Territory. Macdonnell

Ranges, undated, Giles s.n. (MEL holotype); Palm Valley, 1897, Stirling

s.n. (K); Palm Valley, 17 Aug 1929, Cleland s.n. (AD); Palm Valley,

Colliver 5281 (QRS); Palm Valley, Lothian 315 (AD); Palm Valley, ca 8

miles SSW of Hermannsburg Mission, 16 Sep 1965, Willis s.n. (MEL);

Palm Valley, Beauglehole 10460 (MEL); Glen of Palms (Palm Valley),

Beauglehole 23850 (MEL); Palm Valley, Beauglehole 27516 (DNA,

MEL, NSW); Palm Valley, Mitchell 84 (DNA); Palm Valley, Rodd 3215

with Jacobs (DNA, QRS); Palm Valley, Rodd 3216 (K); Palm Valley, 4

Aug 1978, Latz s.n. (QRS); Palm Valley, Oct 1951, anon s.n. (BRI); Palm

Valley, Maconochie 1665 (BRI); Finke Gorge NP, Palm Ck, 2 km W of

carpark, 24°02’S, 132°41’E, Dowe 352 (DNA); Finke Gorge NP, Palm Ck,

2 km W of carpark, Dowe 353 (JCT).

Specimens from cultivated material: AUSTRALIA: New South Wales.

Royal Botanic Gardens Sydney, Rodd 136879 (DNA, K).

A Taxonomic Account of Livistona R.Br. (Arecaceae) 299

Local and vernacular names: Central Australian fan palm, Central

Australian cabbage palm, Palm Valley Livistona.

Habitat and ecology: Grows along creek lines and watercourses with a

permanent shallow water supply, and in sandstone gorges, at 600-650 m

alt. The total population includes about 2000 mature individuals confined

to Palm Valley and Little Palm Ck. Flowers Jul-Dec; fruits Nov-Feb.

Conservation status: Lower risk, conservation dependent (IUCN, 2006).

Distribution: Australia. Northern Territory. Endemic to the Finke R. system

in the Macdonnell Ranges (Fig. 15).

Notes: The name Livistona mariae was first used by Mueller (1874b)

in notes under L. /eichhardtii, with which he considered that it may be

conspecific. Previously, Mueller (1874b) had mentioned it only as the

palm from the “Glen of Palms’ collected by Gilles. These instances had

no descriptions and in the former were considered as a provisional name

only. A complete description in Mueller (1878) provided validation of the

name. He named it in honour of the “... Grandi-principi Mariae, Ducissae

Edinensi, cultus hortorum nobilioris tam in Russia quam nunc in Brittania

patronae imperiali’. However, Mueller’s description included elements of

another species (“... Mill-stream fluminis Fortescue-River satis numerosa,

F. Gregory, J. Forrest...”), later described as L. alfredii from Western

Australia (Mueller, 1892).

Subsequently, the identities of L. mariae and L. alfredii remained

unclear to some botanists, with Gardner (1923) referring to the Millstream

Palm (i.e. L. alfredii), as L. mariae and applying L. alfredii to yet another

taxon in the Kimberleys. However, other taxonomists distinguished the two

taxa (Drude, 1893; Beccari, 1931).

The closeness of L. mariae to L. rigida has been recognised by

some authors. Johnson (1981) wrote of the relationship of L. rigida:

. very Closely related to the Fan-Leaved Palm (L. mariae) of central

Australia and may even be conspecific...”. Rodd (1998) resolved this by

placing L. rigida as a subspecies of L. mariae, but noting his action as

a temporary measure pending a thorough investigation. Considering the

above, and while acknowledging that L. mariae and L. rigida are closely

related, I have reinstated the latter to specific status. The leaves of L. rigida

are comparatively smaller, the folding into a ‘v’ along the axis of the costa

is considerably more pronounced and the segment apices are most often

rigid rather than semi-pendulous. The fruit are also comparatively smaller,

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300 Gard. Bull. Singapore 60 (2) 2009

although occasionally L. mariae produces smaller than average fruits and

L. rigida larger than average fruits.

Livistona mariae is a large canopy palm to 30 m tall; leaves

are large and regularly segmented; segment apices are pendulous, and

with a bifurcate cleft to 65% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to

14 partial inflorescences; bracts are loosely tubular and covered with dense

white-grey scales; flowers are greenish cream to yellow; fruit are globose

to 18 mm diam., and semi-glossy black at maturity.

35. Livistona rigida Becc., Webbia 5 (1921) 19. -Livistona mariae subsp.

rigida A.N.Rodd, Telopea 8 (1998) 80, figs 3b, 5b-c, 6. — Type: Australia.

Queensland, Gulf of Carpentaria, Albert R., Aug 1855, Mueller s.n.

(holotype, K; isotype, FI). Plate 24B-E.

Functionally dioecious palm. Trunk to 28 m tall, 30-40 cm dbh, leaf scars

raised, internodes narrow, grey, petiole stubs persistent in the basal | m or

so, otherwise deciduous. Leaves 30-50 in a globose crown; petiole 150-250

cm long, 20-35 mm wide, adaxially concave, margins with single curved

reddish-black spines to 6 mm long in the proximal portion, otherwise

smooth; leaf-base fibres moderately prominent, coarse, persistent; lamina

costapalmate, regularly segmented, circular in outline, 125-170 cm long,

100-150 cm diam., rigidly coriaceous, adaxially grey to glaucous, midgreen,

semi-glossy, abaxially lighter green, glaucous; lamina divided for 50-55%

of its length, with 50-78 segments, depth of apical cleft 30-63% of the

segment length, apical lobes acuminate, rigid or semi-pendulous; parallel

veins 5-7 each side of midrib; transverse veins thinner than parallel veins.

Inflorescences unbranched at the base, not sexually dimorphic, 100-250

cm long, not extending beyond the limit of the crown, branched to 4 orders;

partial inflorescences 9-14; prophyll 25-38 cm long, 8-9 cm wide; peduncular

bract 1, loosely tubular with moderate to dense white-grey scales; rachis

bracts loosely tubular with sparse white scales; rachillae 3-8 cm long, rigid,

glabrous. Flowers in clusters of 3-8, globose, ca 1.8 mm long, cream to

yellow; sepals broadly ovate, ca 2 mm long, thin, acute; petals triangular,

|.0-1.4 mm long, obtuse; stamens ca 1.2 mm long. Fruit globose, 12-14

mm diam., semi-glossy black; epicarp with scattered lenticellular pores;

suture line extends for ca 2 the length of the fruit, marked with lip-like

structures; mesocarp fibrous; endocarp thin; pedicel ca 2 mm long. Seed

globose, 9-11 mm wide. Eophyll 5-ribbed.

Specimens examined: AUSTRALIA: Northern Territory. Mataranka, Must

—_

A Taxonomic Account of Livistona R.Br. (Arecaceaeé

1499 (DNA): Kapalga, Dunlop 5052 with McKean (CANB, DNA, NSW):

Goyder R., Dunlop 7259 (DNA, NSW); Roper Hwy, 20 k E of Stuart Hwy.

Latz 7828 (DNA, NSW): Near Roper R. crossing, Salt Ck, Dowe 209

& 211 (JCT): Mataranka Stn, Salt Ck, 5 Jul 1964, Schneider s.n. (AD):

Roper R., road crossing 1.5 km W of Mataranka Homestead, 14°55’S,

133°06°E, 120 m alt., Dowe 210 & 212 (JCT):; S of Mataranka, Warlock

Ponds Bridge, Symon 10358 (NSW); Elsie Ck, 20 km SE of Mataranka,

Beauglehole 58320 with Errey (MEL). Queensland. Burketown-Camoweal

Rd, Gregory R. between crossing of Beames Brook and Planet Downs

Stn, 17 Aug 1973, Williams s.n. (BRI); Gregory R., 10 km N of Gregory

Downs, Irvine 1856, 1857, 1858 & 1859 (QRS): Lawn Hill Ck, SW of

Burketown, Jun 1963, Gittens s.n. (BRI); Lawn Hill NP, Jan 1988, O'Keefe

s.n. (BRI); Lawn Hill Ck, 3 km above Lawn Hill Homestead, Rodd 2937

& 2938 (BRI, K, QRS, NSW); Lawn Hill Stn, Babbling Brook, Latz 1619

(BRI, CANB, DNA): Adels Grove, de Lestang 233 (QRS): Adels Grove,

Lawn Hill Ck, 500 m W of camping grounds, 18°41°55°S, 138°31°06°E,

Dowe 630 with Kazandjian (BRI, JCT, K); 34 miles S of Burketown, Webb

10634 (BRI); Albert R., Aug 1855, Mueller s.n. (K holotype): 30 km E

of Highland Plains, 18°43”S, 138°28”°E, Maconochie 1656 & 1657 (BRI):

Palm Springs, junction of Police Ck and Leichhardt R., ca 40 km NNE of

Mt Isa, Farrell TF815 (BRI); Lake Julius Dam, 12 Feb 1977, Schmid s.n.

(BRI); Dugald R., tributary of Flinders R., 20°12°08.8"S, 140°14°14.07E,

178 malt., Dowe 749 with Martin (BRI, JCT).

Local and vernacular names: Mataranka Palm, Gregory R. Cabbage

Palm.

Habitat and ecology: Grows as a riparian element along stream margins,

on seasonally inundated banks, in creek lines and watercourses, sometimes

with intermittent flow, but with a permanent shallow water supply, 2-300 m

alt. Flowers Jun-Dec; fruits Nov-May.

Conservation status: Lower risk, conservation dependent.

Distribution: Australia. Queensland and Northern Territory. In north-western

Queensland in the catchments of the Gregory, Leichhardt, Nicholson, Albert

and Flinders Rivers, and in the Northern Territory on the Roper R., South

Alligator Rivers and east Arnhem Land (Fig. 15).

Notes: Livistona rigida was described by Beccari (1921), based on a

collection, at that time by an unknown collector, from Albert R. in the Gulf

302 Gard. Bull. Singapore 60 (2) 2009

of Carpentaria, and named for the “... spadix rigidus”. This specimen,

conserved in K, is most likely one that was collected by Mueller during

the 1855 North-Australian Expedition and sent to Kew. Mueller (1858)

reported on the expedition, in which he noted: “... Livistona inermis and

an allied species supplied us occasionally with palm cabbage...”, but

whether the ‘allied species’ relates to L. rigida cannot be verified. But it

is known that in August of 1855 Mueller traversed the area in which L.

rigida is now known to occur and collected extensively (Mueller, 1857).

Bentham (1878) included this same specimen as one of his reference

specimens in his description of L. humilis, citing Mueller as the collector.

However, his description certainly does not account for the size of the

leaf in the Mueller specimen, as his description placed the radius of the

leaf at “about 1/2 ff’ (ca 45 cm) whereas the leaf of L. rigida is 150-170

cm long. Beccari (1931), not knowing the size or habit of L. rigida as he

described it on the unannotated Mueller specimen, suggested that Plate

145, Fig. 4 and Plate 146 in Martius (1838) could belong to L. rigida

based on the apparent large size of the leaf. Beccari correctly determined

that the leaf of L. inermis, the species that the illustrations were supposed

to represent, were much smaller that those illustrated.

Livistona rigida is a large canopy palm to 28 m tall; leaves are large

and regularly segmented; segment apices are rigid to semi-pendulous, and

with a bifurcate cleft to 63% of the segment length; the inflorescence is

unbranched, not extending beyond the limit of the crown, and with up to

14 partial inflorescences; bracts are loosely tubular, with the peduncular

bract with moderate to dense white-grey scales and the rachis bracts with

sparse white-grey scales; flowers are cream to yellow; fruit are globose to

14 mm diam., and semi-glossy black at maturity.

36. Livistona lanuginosa A.N.Rodd, Telopea 8 (1998) 82, figs 3d, 7, 9a.

— Type: Australia. Queensland, 75 km SE of Ravenswood, Glenroy Ck,

20°35’S, 147°10E, 25 Aug. 1978, Irvine 1912 (holotype, NSW; isotypes,

BH, BRI, K, MEL, QRS). Plate 25A-E.

-Livistona mariae auct. non F.Muell., F.M.Bailey, Queensl. fl. 5 (1902)

1684.

Functionally dioecious palm. Trunk to 18 m tall, 25-35 cm dbh, leaf

scars raised, compressed, internodes broad, grey-brown, base with

persistent petiole stubs. Leaves 35-45 in a globose crown; petiole 159-

200 cm long, 30-35 mm wide, adaxially flat, margins with small single,

curved black spines confined to the proximal portion; leaf-base fibres

moderately prominent, coarse, persistent; lamina costapalmate, costa

A Taxonomic Account of Livistona R.Br. (Arecaceae) 303

arcuate, regularly segmented, + circular in outline, 130-190 cm long,

coriaceous, adaxially pale grey-green, abaxially lighter grey-green, waxy,

strongly pruinose; lamina divided for ca 34% of its length, with 70-92

segments, depth of apical cleft ca 24% of the segment length, apical

lobes acuminate, semi-pendulous; parallel veins 8 each side of midrib;

transverse veins thinner than parallel veins. Inflorescences unbranched at

the base, not sexually dimorphic, 140-220 cm long, not extending beyond

the limit of the crown, branched to 4 orders; partial inflorescences 8-12;

prophyll densely tomentose; peduncular bracts 1-2, loosely sheathing,

densely lanuginose; rachis bracts loosely sheathing, not disintegrating,

densely lanuginose; rachillae 3-12 cm long, glabrous. Flowers solitary or

in pairs, funnel-shaped, 2.8-3 mm long, cream to yellow; sepals narrowly

triangular, ca 1.5 mm long, fleshy, acuminate to aristate; petals triangular,

2.8-3 mm long, thick, acute; stamens ca 2 mm long. Fruit globose, 25-

36 mm diam., purple-brown to black with scattered white flecks; epicarp

with large lenticellular pores; suture line extends for ca '2 the length of

the fruit, marked with lip-like structures; mesocarp fibrous, dry; endocarp

2-3 mm wide; pedicel to | mm long. Seed globose, 18-22 mm wide.

Eophyll 5 ribbed.

Specimens examined: AUSTRALIA: Queensland. Campaspe R.,

Hellenslie Stn, Jul 1900, Berney s.n. (BRI); Liontown Stn, Oaky Ck,

20°23754.3”S, 146°03’35.4”E, 290 m alt., Dowe 751 (JCT); Windsor

Sin, Oaky Ck, 20°19’20.8”S, 146°03’58.7”E, 300 m alt., Dowe 752

(JCT); Cement Ck, tributary of Campaspe R., Dowe 707 (JCT); 75 km

SE of Ravenswood, Glenroy Ck, 20°35’S, 147°10E, /rvine 19/2 (BH,

BRI, K, MEL, NSW holotype, QRS); Glenroy Ck, ca 3 km from Glenroy

HS, McLain 2 (QRS); Glenroy Ck, ca 11 km N of Burdekin Falls, Rodd

3759 with Jackes (NSW); Glenroy Ck, | km N of junction with Stones

Ck, 20°33’S, 147°06’E, Dowe 292 & 293 (BRI, FTG); Middle Burdekin

Valley near Burdekin Falls, 1978, McLain s.n. (NSW); Nosnillor Stn,

Campaspe R., on Clermont Rd, 112 km S of Charters Towers, /rvine

1912, 1913 & 1914 (QRS, NSW); Nosnillor St, Muckinbulla Waterhole,

Dowe 706 (JCT); Cattle Ck, 7 km E of Harvest Home, Thompson 238

(BRI); Longton Stn, Garden Ck, $21° 04.482’, E145° 36.185’, 290 m, 28

Oct 2005, Dowe s.n. (JCT); Longton Stn, Amelia Ck, 9.8 km NW of Cape

R. crossing on Pentland road, Dowe 705 (JCT).

Specimens from cultivated material: AUSTRALIA: Queensland.

Townsville Palmetum (ex Burdekin R. near Burdekin Dam wall, 1986,

Tucker s.n. (NSW); ibid, Dowe 111 (NSW).

304 Gard. Bull. Singapore 60 (2) 2009

Local and vernacular names: Cape River Fan-palm, Burdekin Livistona,

Woolly Palm.

Habitat and ecology: Occurs as a riparian element along the margins of

streams and in seasonal gullies and associated floodplains with high

watertables, at 140-270 m alt. Flowers Mar-Nov; fruits Nov-Jan.

Conservation status: Endangered (Dowe, 2007).

Distribution: Australia. Queensland. Restricted to the Cape, Campaspe,

Rollston, Burdekin and Belyando Rivers and tributaries (Fig. 15).

Notes: Livistona lanuginosa was described by Rodd (1998) based on /rvine

1912 collected from Glenroy Ck, a tributary of the Burdekin R., Queensland, |

and named for the densely lanuginose inflorescence bracts. Bailey (1902) |

-had previously determined the population to be L. mariae: “... I have

received a portion of a leaf from FL. Berney, of Hellenslie, Campaspe

R., which in all probability belongs to this inland palm’. Recently, the

species was informally known as ‘Livistona sp. Cape River’ (Jones, 1996).

Livistona lanuginosa most closely resembles L. mariae and L. rigida, but

is readily distinguished by its much larger fruit and the densely lanuginose

prophyll and inflorescence bracts.

An interesting historical account was provided by Leichhardt (1847)

who noted this palm in the diary of his transcontinental expedition of 1845:

. (in the Burdekin R. valley on 25 March 1845)... at the junction of

the creek, a great number of small Corypha palms were growing, and my

companions observed the dead stems of some very high ones, whose tops

had been cut off by the natives, probably to obtain the growing shoot’.

Livistona lanuginosa is one of Australia’s most endangered palms

(Dowe, 2007). Pettit and Dowe (2003) undertook a population study and

estimated that the total population consisted of less than 1000 mature

individuals, all of which occur on private property within the Burdekin R.

catchment. Primary threats are trampling and browsing of seedlings and

juveniles by cattle, which prevents regeneration. |

Livistona lanuginosa is a large sub-canopy to canopy palm to 18 m

tall; leaves are large and regularly segmented; segment apices are rigid to

semi-pendulous, and with a bifurcate cleft to 63% of the segment length; the |

inflorescence is unbranched, not extending beyond the limit of the crown, |

and with up to 12 partial inflorescences; bracts are loosely sheathing and |

distinctively densely tomentose/lanuginose; flowers are cream to yellow;

fruit are globose to 36 mm diam., and purple brown to black at maturity.

" | |

A Taxonomic Account of Livistona R.Br. (Arecaceae 305

Excluded and uncertain names

Livistona subgen. Livistonella Becc., Ann. Roy. Bot. Gard. (Calcutta) 13

(1931) 53. -Type: Livistona rupicola Ridl. = Maxburretia rupicola (Ridl.)

Furt.

Livistona ? bissula Mart., Hist. Nat. Palm. 3 (1850) 319. Bissu/a Rumph.

Herb. Amboin. | (1741) 85. -Licuala ? bissula Mig., Fl. Ned. Ind. 3 (1855)

57. -Saribus bissula Blume ex Pfister, Beitr. Vergl. Anat. Sabaleenblat.

(1892) 25. — Bissula was the name applied by Rumphius (1741) to a fan

palm collected in Sulawesi. Rumphius’ description does not allow for a

positive identification, and therefore its identity and taxonomic position

has been variously interpreted. Blume (1838) was the first to discuss the

name, including it in his account of Saribus, and suggesting a possible

relationship with Licuala excelsa Zipp. Martius (1838) similarly included

it initially only in the mononomial form and placed it at the conclusion of

his account, but was later (Martius, 1849) to combine it with Livistona,

though with a question mark. Miquel (1855) used it in combination with

Licuala, including Martius’ Livistona bissula as asynonym. Pfister (1892

used the name Saribus bissula Blume in his study of the anatomy of the

Sabai alliance, and concluded that it was near Livistona. Merrill (1917)

noted that it “may be neither a Livistona nor a Licuala; the description

of the fruits is suggestive of Pholidocarpus”, while Beccari (1921,

1931) placed it as a synonym of Licuala celebica Mig. However, Miquel

(1868) in his protologue of that species did not mention any relationship

to Rumphius’ Bissu/a, but rather placed it under Livistona with the

suggestion that “forsan ad L. rotundifoliam referenda”. As Bissula is a

pre-Linnean mononomial, the name is otherwise inadmissible under the

ICBN.

Livistona diepenhorstii Hassk., Bonplandia 6 (1858) 180. Basionym of

Pholidocarpus diepenhorstii (Hassk.) Burret, Notizbl. Bot. Gart. Berlin-

Dahlem 15 (1941) 327. = Pholidocarpus ihur (Giseke) Migq., Fl. Ned. Ind.

3 (1855) 47.

Livistona dournowiana Hort. ex Taylor, Bailey Stand. Cycl. Hort. (1943)

1896, nom. inval.— This name was used for a palm in horticulture and

described as “a new palm with lvs. resembling those of Latania borbonica,

which is Livistona chinensis, Hab. (?). — L. muelleri, Bailey.” However, the

fruit is further described as “oval, nearly 6 in. long” which excludes it from

either of those species.

306 Gard. Bull. Singapore 60 (2) 2009

Livistona enervis Hort. ex anon., Ill. Gart.-Zeitung 16 (1891) 346. — This

name was applied to a palm from the nursery collection of William Bull.

The leaves were described as delicately textured and with narrow segments.

The description is inadequate for correct identification. This could be a

typographical error for L. inermis R.Br., which otherwise was the name

incorrectly used for L. decora (W.Bull) Dowe in many informal accounts

and nursery catalogues of that era.

Livistona eocenica Ettingsh. & J.S.Gardner, Proc. Roy. Soc. London 29

(1879) 393. — A fossil taxon described from Sheppey, United Kingdom.

Reid and Chandler (1933) synonymised it under Hightea elliptica Bowerb.

(Myrtaceae?) pro parte and genus?, Cornaceae sect. Mastixioideae pro

parte, but apparently not a palm. Type not designated.

Livistona erecta fide Moore (1963b), nom. inval. — Moore listed this name

in his annotated list of cultivated palms as ‘a name without botanical

standing’.

Livistona filamentosa Hort. Petrop. ex R.Pfister, Beitr. Vergl. Anat.

Sabaleenblat. (1892) 25. = Washingtonia filifera (Linden ex André)

H.Wendl. ex de Bary (Beccari, 1921).

Livistona filifera Hort. ex H.Wendl., in Kerchove, Palmiers (1878) 250,

nomen. — A name that appeared in a list of palms in the ‘Index Général’,

without a description or reference to specimens. It was noted as a synonym

of L. inermis R.Br., but without a description or reference to specimens, the

name remains invalid.

Livistona (?) gaudichaudii Mart., Hist. Nat. Palm. 3 (1849) 242. -Type:

Hawaii. 1836, Gaudichaud-Beaupreé s.n. (FI, P) = Pritchardia gaudichaudii

(Mart.) H.Wendl. = Pritchardia martii (Gaudich.) H.Wendl., Bonplandia

10 (1862) 199.

Livistona hoffmanni Hort. ex. André, Rev. Hort. 77 (1905) 527, nomen.

— This name was applied to juvenile palms exhibited at a meeting of the

Royal Horticultural Society of England by M.R.Hoffmann, of Streatham,

England. André quoted that M.M.Watson, in the Gardeners’ Chronicle (no |

publication details provided) suggested that they were plants of Livistona |

mariae F.Muell. However, without a description or specimens this cannot |

be supported, and the name cannot be placed. |

A Taxonomic Account of Livistona R.Br. (Arecaceae) 307

Livistona humilis R.Br. var. bloomfieldiensis, in Cuccuini & Nepi, Quad. Bot.

Amb. Appl., 17/1 (2006) 104, 192, nom. inval.—This name was included in a

list of specimens conserved in FI. Cuccuini and Nepi (2006) suggested it was a

synonym of Livistona humilis var. sclerophylla Becc. and Livistona muelleri.

It is an invalid name that was used on a herbarium specimen sheet.

Livistona humilis var. leichhardii (F.Muell.), in Cuccuini & Nepi, Quad.

Bot. Amb. Appl., 17/1 (2006) 104, nom. inval. — This name was included in

a list of specimens conserved in FI. It is an invalid name that was used on

a herbarium specimen sheet.

Livistona kingiana Becc., Malesia 3 (1889) 199. -Type: Malaysia. Perak,

Dr King s.n. (holotype, Fl) = Pholidocarpus kingianus (Becc.) Ridl., Mat.

Fl. Malay. Penins. 2 (1907) 167 (Dransfield & Uhl, 1983b).

Livistona macrophylla Bruder, Lotos. Jahrbuch fiir Naturwissenschaften 10

(1890) 39; Bot. Centralbl. 41 (1890) 297. — A fossil taxon described by G.

Bruder from the Tertiary deposits of freshwater sandstone from Tuchorschitz,

Sasser Region in B6hmen (present-day south-east Germany), collected by

Carl Ihl prior to 1888. Current location of the type specimen is not known.

Livistona macrophylla Roster in Bull. Soc. Tose. Ort. 29 (1904) 82, nomen.

— This is a name that appeared in a list of cultivated plants and appears not

to be associated with L. macrophylla Bruder (see previous entry).

Livistona martii Gaudich., Voy. Bonité, Atlas (1842) t. 58, 59. Livistona

(?) martiana Gaudich. ex Mart., Hist. Nat. Palm. 3 (1849) 242, ortho. var.

—Lectotype: Illustrations in Gaudichaud, Voy. Bonité, Atlas (1842) t. 58,59.

= Pritchardia martii (Gaudich.) H.Wendl., Bonplandia 10 (1862) 199.

Livistona minima Reid & Chandler, London Clay FI. (1933) 109. —A fossil

taxon described from a single seed from southern England. -Type: United

Kingdom. Sheppey, Bowerbank V. 22021 (BM).

Livistona moluccana Hort. ex. H.Wendl., in Kerchove, Palmiers (1878) 250,

nomen. — A name that appeared in a list of palms in the ‘Index General’,

without a description or reference to specimens. Although there are a

number of specimens in FI (sheets 1132G & 1132G-A) that are labelled as

L. moluccana, in Beccari’s handwriting, the name remains invalid as it has

not been validly published. Beccari (1921, 1931) included the name in the

‘Sinonimi e specie escluse’ section in his treatment of Livistona.

308 Gard. Bull. Singapore 60 (2) 2009

Livistona miilleri Hort. ex H.Wendl., in Kerchove, Palmiers (1878) 250,

nomen. — A name that appeared in a list of palms in the ‘Index Général’

without a description or reference to specimens.

Livistona occidentalis Hort. ex Hook.f., Report on the progress and

condition of the Royal Gardens at Kew 1882 (1884) 64. = Brahea dulcis

(Kunth) Mart. — This name was used by Hooker in a list of palms cultivated

in the Royal Gardens, Kew. He listed both L. occidentalis and Thrinax

tunicata (Hort. ex Verschaff.) Hook.f. as synonyms of Brahea dulcis.

Livistona okinawensis Hort. ex L.H.Bailey, Hortus third (1976) 674, nom.

inval. — A name used in horticulture and suggested to be a synonym of L.

chinensis var. subglobosa. This name cannot be placed as there were no

description or specimens cited.

Livistona ovaliformis Hort. ex. R.M.Grey, in Rep. Harvard Botanic Garden,

Cienfuegos, Cuba (1927) 66, nom. inval. — A name of horticultural origin,

and which lacks a description and specimens. Otherwise, it may possibly

be a misspelling of Livistona olivaeformis (Hassk.) Mart.

Livistona ramsayi F.Muell., Fragm. 7 (1874) 221. -Type: Australia.

Queensland, Rockingham’s Bay, Ramsay s.n. (holotype, MEL). = Licuala

ramsayi (F.Muell.) Domin.

Livistona rupicola Ridl., Jour. Straits Branch Roy. Asiat. Soc. 41 (1904)

41. -Type. Malaysia. Selangor, Gua Batu, Ridley 8285 (lectotype, SING). =

Maxburretia rupicola (Ridl.) Furt.

Livistona ternatensis Hort. ex Salomon, Palmen (1887) 174, nomen. — A

name without a description or reference to specimens and cannot be placed.

Livistona vidalii Becc., Webbia | (1905) 343. -Type: Philippines. Luzon,

Pampanga, Arayat, 27 Feb. 1903, Garcia 63 (holotype, PNH; isotype, FI)

= Corypha elata Roxb. = Corypha utan Lam. (Beccari, 1919b). — Beccari

(1919b) related the illustration, t. 93 in Vidal (1883), as this taxon. Although

the leaves somewhat resemble those of L. saribus, the flowers and fruits are

of C. utan.

Livistona vogamii H.Sander ex Becc., Webbia 5 (1921) 22 (also as ]

‘voganii’, Beccari, 1931). — A name attributed to Sander, possibly from

horticulture, but without a description or specimens, cannot be placed with

- a owed 5]

A Taxonomic Account of Livistona R.Br. (Arecaceae) 309

certainty. Beccari (1921, 1931) questionably placed it as a synonym of L.

cochinchinensis.

Livistona zollingeri Hort. ex Devansaye, Rev. Hort. 47 (1875) 34, nom.

inval. -Livistona zollingeriana Blume, Salomon, Palmen 142 (1887) nom.

inval. -Saribus zollingerianus Hassk., Salomon, Palmen (1887) 142, nom.

inval. — This name was first used by Devansaye without a description or

reference to specimens and cannot be placed with certainty. However,

Zollinger (1857), in the protologue of L. altissima, stated that the name

‘Zollingerianus’ had been used on the herbarium label for that species.

Previously the taxon had been designated as Saribus sp. by Zollinger (1854)

and Miquel (1855), and subsequently as Livistona zollingeri (Devansaye,

1875) and the orthographic variation Livistona zollingeriana Salomon

(1887).

Future work

There are a number of issues with regards to Livistona that are presently

unresolved. A phylogenetic assessment of the entire genus has yet to be

undertaken. Additionally, the relationships of Livistona to other genera

have yet to be fully investigated and resolved. Detailed studies of pollen

morphology and sexuality are lacking for many species. In addition, some

records of unidentified species from unusual habitats have to be investigated.

There is reference to a possible Livistona species on the karst limestone

island of Palawan, and populations of unidentified Livistona species

occurring in Sumatra. Within Australia, the full extent of the distribution of

some species has yet to be ascertained, such as species in the Kimberleys

of Western Australia and Cape York Peninsula in Queensland.

Acknowledgements

I offer sincere thanks to Associate Professor Betsy Jackes, Dr Leone

Bielig and Dr Anders Barfod for their support during my PhD candidature

and project (from which this revision was largely taken) at James Cook

University, Townsville. For assistance at Aarhus University, Denmark, |

thank Annie Sloth and again, Dr Anders Barfod. For general herbarium

assistance, I am indebted to J.Mogea, Irawati, K.Kramadibrata and

C.Heatubun (BO): G.Guymer, A.Bolan, P.Forster and R.Henderson (BRI):

E.Fernando (CAHUP, LBC); P.Cuccuini and M.Tardelli (FT); N.G.Hiep

Nd ——————

310 Gard. Bull. Singapore 60 (2) 2009

(HN); B.Baker, J.Dransfield and M.Harley (K); L.G.Saw (KEP); M.Thijsse

(L); O.Gideon and R.Banka (LAE); H.Ehara, R.Nobe, H.Kato, M.Ono and

N.Yoshida (MAK); R.Maturbongs (MAN); C.Coles, P.Milne and B.Conn

(MEL); K.Hill (NSW); P.Pignal (P); D.Madulid (PNH); and R.Kiew

(SING). For assistance with field work and other logistical support, I thank

A.Cairns, D.Dixon, P.Gadek and A.Kazandjian (James Cook University);

S.van Leeuwen (CALM, WA); G.Kendrick (WA Museum); D.Greenwood

(VUT); J.Conran (University of Adelaide); A.K.Irvine (CSIRO); and

K.Tansasha, P.Vatcharakorn and M.Ferrero (Thailand). Mirjam Maughan

(ACTER) is thanked for preparing the distribution maps. Lucy T. Smith

provided the line drawings of Livistona carinensis. Dr John Dransfield is

thanked for assistance in the neotypification of Livistona saribus. | also

thank the reviewers who provided constructive comments for improvements

to this revision. Funding was provided by Australian Biological Resource

Study; Australia Pacific Science Foundation; Pacific Biological Foundation;

_and James Cook University (Doctoral Merit Research Scheme and Internal

Research Supplementary Funding).

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320 Gard. Bull. Singapore 60 (2) 2009

Plate 1. Livistona saribus: A. In habitat, palm swamp forest, Khum Cha-not, northeast

Thailand; B. Leaf detail; C. Petiole with armature, cultivated plant, Cairns Botanic Gardens,

Queensland, Australia; D. Portion of rachilla with mature flowers; E. Fruit, whole and one

dissected to reveal seed coat intrusion. (Photos: J.L.Dowe; 1B, 1D and 1E from cultivated

plant, Townsville Botanic Gardens, Queensland, Australia).

A Taxonomic Account of Livistona R.Br. (Arecaceae) 321

Plate 2. Livistona exigua: A. In habitat, at ridge top kerangas forest at 150 m alt.; B.

Crown detail; C. Inflorescence and leaf-base fibres; D. Rachillae and immature but full-

size fruit. (Photos: J.Dransfield; voucher specimen, Dransfield et al. 6568, from Bukit

Batu Patam, Ulu Ingei, Brunei Darassalam).

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Gard. Bull. Singapore 60 (2) 2009

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Plate 3. Livistona rotundifolia: A. Habit, avenue in Bogor Botanic Gardens, Indonesia;

B. Crown with infructescence, cultivated plant, Florida, USA; C. Crown with petiole

bases and leaf-base fibres, cultivated plant, Townsville Botanic Gardens, Queensland,

Australia; D. Fruit, whole and with one dissected, cultivated plant, Florida, USA. (Photos:

J.L.Dowe).

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A Taxonomic Account of Livistona R.Br. (Arecaceae)

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Plate 4. Livistona papuana: A. In habitat, rainforest at 540 m alt.; B. Leaf detail. (Photos:

W.J.Baker; voucher specimen, Baker WJB85/], from Fakfak, Timika, Papua, Indonesia).

Livistona brevifolia: C. In habitat, coastal forest; D. Leaf; E. Inflorescence with mature

flowers. (Photos: C.D.Heatubun, from Gag Island, Raja Ampat, Papua, Indonesia).

Gard. Bull. Singapore 60 (2) 2009

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Plate 5. Livistona merrillii: A. Habit; B. View into crown; C. Fruit. (Photos: J.L.Dowe,

cultivated plant at Bogor Botanic Gardens, Indonesia). Livistona woodfordii: D. In habitat,

in coastal forest at sea-level, West Point, Rossel Island, Milne Bay Province, Papa New

Guinea; E. Crown with infructescence (voucher specimen, Banka et al. 2015, from 50 m

alt. at Yeleamba, Rossel Island, Milne Bay Province, PNG); F. Fruit (voucher specimen, .

Banka et al. 2019, from East Point, Sudest Island, Milne Bay Province, PNG). (Photos:

J.L.Dowe).

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A Taxonomic Account of Livistona R.Br. (Arecaceae)

Plate 6. Livistona chocolatina: A. \n habitat, rainforest. (Photos: J.L.Dowe, taken at 300 m

alt., Kuriva, Central Province, Papua New Guinea; voucher specimen, Barfod et al. 466-

holotype); B. Leaf and inflorescence. (Photos: A.Kyjaer, taken at 400 m alt., Bulili Ridge,

Lababia, Morobe Province, PNG; voucher specimen, Barfod et al. 514); C. Inflorescence.

(Photo: J.L.Dowe, from Kuriva, Central Province, PNG; voucher specimen, Barfod et al.

466). Livistona tothur: D. Leaf, adaxial view; E. Leaf, abaxial view. (Photos: A.Damborg,

taken at 500 m alt., Oenake Mts, West Sepik Province, PNG; voucher specimen, Damborg

et al. 418-holotype).

326 Gard. Bull. Singapore 60 (2) 2009

Plate 7. Livistona tothur: A. Yaflorescence with mature flowers; B. Flowers; C.

Infructescence. (Photos: A.S.Barfod, taken at 500 m alt., Niau Kono, Oenake Mts, West

Sepik Province, PNG; voucher specimen, Barfod et al. 510); D. Fruit. (Photo: J.L.Dowe,

taken at S00 m alt., Niau Kono, Ampol, Oenake Mts, West Sepik Province, PNG; voucher

specimen, Dowe et al. 516).

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A Taxonomic Account of Livistona R.Br. (Arecaceae) By)

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Plate 8. Livistona surru: A. In habitat, rainforest; B. Leaf; C. Inflorescence with rachis

bract detail and immature fruit. (Photos: W.J.Baker, taken from Bosmun, Ramu River,

Madang Province, PNG; voucher specimen, Baker et al. 582); D. Infructescence with

immature fruit; E. Fruit. (Photos: A.S.Barfod, taken at 950 m alt., Miwaute, West Sepik

Province; voucher specimen, Barfod et al. 390-holotype).

328 Gard. Bull. Singapore 60 (2) 2009

Plate 9. Livistona halongensis: A. In habitat, forest on limestone; B. Leaf. (Photos:

G. Dahme, from Cat Ba Island, Halong Bay, Vietnam). Livistona chinensis: C. Habit,

cultivated plants, Townsville Botanic Gardens, Queensland, Australia; D. Leaf detail,

cultivated plants, Cairns Botanic Gardens, Queensland, Australia. (Photos: J.L.Dowe);

E. Flowers, cultivated plant, Peninsula Botanical Garden, Khao Chong, Trang, Thailand.

(Photo: A.S.Barfod); F. Infructescence with mature fruit, cultivated plants, Townsville

Botanic Gardens, Queensland, Australia. (Photo: J.L.Dowe).

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A Taxonomic Account of Livistona R.Br. (Arecaceae)

Plate 10. Livistona boninensis: A. Illustrated in habitat [as Corypha japonica] in Kittlitz,

F.H. (1844), Veg.-Ansicht. t.15; B. In habitat, remnant forest, Chichi-jima, Bonin Islands,

Japan. (Photo: H.Kato). Livistona endauensis: C. In habitat, rainforest at Bukit Bauk,

Trengganu, Malaysia; D. Leaf detail, cultivated plant, Singapore Botanic Gardens; E.

Infructescence with mature fruit, cultivated plant, Singapore Botanic Gardens. (Photos:

J.L.Dowe).

330 Gard. Bull. Singapore 60 (2) 2009

Plate 11. Livistona tahanensis: A. In habitat, submontane forest, near Wrays Camp,

Gunung Tahan, Pahang, Malaysia (voucher specimen, Dransfield 659). (Photo:

J.Dransfield). Livistona jenkinsiana: B. In habitat, rainforest, Khao Chong, Thailand. }

(Photo: J. Dransfield); C. Leaf detail. (Photo: K.Anker); D. Armature, from cultivated plant |

at Cairns Botanic Gardens, Queensland, Australia. (Photo: J.L.Dowe); E. Infructescence

with mature fruit; F. Mature fruit. (Photos 11C, 11E, and 11F by K.Anker are based on

specimen (Barfod et al. 710) at Doi Phu Ka National Park, Thailand).

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A Taxonomic Account of Livistona R.Br. (Arecaceae) 331

Plate 12. Livistona speciosa: A. In habitat, rainforest. (Photo: A.S. Barfod, from 900 m

alt., Doi Chiang Dao, Thailand): B. Crown: C. Armature, cultivated plant, Thailand: D.

Mature fruit. (Photos 12B. 12C and 12D: J.L.Dowe, based on Dowe 560 at Khao Ko, 600-

700 m alt., Thailand).

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Gard. Bull. Singapore 60 (2) 2009

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Plate 13. Livistona carinensis: A. \n habitat, wadi in semi-desert, Toha, Djibouti. (Photo:

G. & H. Welch); B. Crown with mature inflorescences, cultivated plant, Florida, USA.

(Photo: H.Ford); C. Armature, cultivated plant, Nong Nooch, Thailand. (Photo: J.L.Dowe);

D. Diagnostic illustration: a. flowers on rachilla; b. flower from above; c. flower side

view; d. flower, longitudinal section; e. sepal; f. fruit; g. seed, longitudinal section. Scale

bar: a = 2.5 mm; b, c, d, e = | mm; f, g = 4 mm. Figs. a-e based on Dowe s.n. collected

fromTownsville Palmetum, Queensland, Australia, dated 18 Mar 2000; Figs. fand g based

on cultivated plant at Miami, Florida, USA (line illustrations: L.T.Smith).

____ ——— ee

is)

eS)

ws)

A Taxonomic Account of Livistona R.Br. (Arecaceae)

Plate 14. Livistona humilis: A. In habitat, open tropical woodland; B. Male plant with

mature inflorescence; C. Female plant with infructescence and mature fruit: D. Fruit.

(Photos: J.L. Dowe; voucher specimen (Dowe 203) from Kakadu National Park, Northern

Territory, Australia).

o>)

1S)

Gard. Bull. Singapore 60 (2) 2009

Plate 15. Livistona muelleri: A. In habitat, from coastal moist forest at Smithfield, Cairns,

Queensland, Australia; B. Leaf detail; C. Crown with inflorescence and mature flower; D.

Flowers; E. Infructescence with full size but slightly immature fruit; F. Mature fruit, and

seed in longitudinal section to reveal seed coat intrusion and embryo. (Photos: J.L. Dowe,

ISB-ISE from Barrett Creek, Cooktown, Queensland, Australia; 1SF from Lockerbie

Scrub, Cape York Peninsula, Queensland, Australia).

19S)

ies)

A Taxonomic Account of Livistona R.Br. (Arecaceae)

Plate 16. Livistona eastonii: A. In habitat, open tropical woodland; B. Infructescence with

mature fruit; C. Fruit. (Photos: J.L.Dowe:; voucher specimen, Dowe 360, from Kimberley

Plateau, Western Australia). Livistona inermis: D. In habitat, open woodland at Nitmiluk,

Katherine Gorge, Northern Territory, Australia. (Photo: D. Gorton); E. Leaf detail,

cultivated plant, Townsville Botanic Gardens, Queensland, Australia; F. Inflorescence with

mature flowers; G. Flowers. (Photos: J.L.Dowe, 16F and 16G from Mt Bundy, Northern

Territory, Australia; voucher specimen, Dowe 200).

eS)

wy)

Gard. Bull. Singapore 60 (2) 2009

Plate 17. Livistona lorophylla: A. In habitat, open woodland; B. Leaf detail. (Photos:

J.L.Dowe; voucher specimen, Dowe 362, from Pentecost River, Western Australia).

Livistona victoriae: C. In habitat, monsoonal open woodland, Victoria River, Northern

Territory, Australia. (Photo: D.Gorton); D. Fruit and seed in longitudinal section to reveal

seed coat intrusion and embryo. (Photo: J.L.Dowe; voucher specimen, Dowe 208, from

Charles Crossing, Jasper Gorge, Northern Territory, Australia).

eS)

A Taxonomic Account of Livistona R.Br, (Arecaceae)

Plate 18. Livistona benthamii: A. In habitat, seasonally inundated riparian forest, Home

Billabong, Cooinda, Northern Territory, Australia (voucher specimen, Dowe 206); B.

Trunk detail; C. Leaf detail. (Photos: J.L.Dowe; voucher specimen, Dowe et al. 406, 18b

and 18c from Lockerbie Scrub, Cape York Peninsula, Queensland, Australia). Livistona

concinna: D. In habitat, upper mangrove forest at 5 m alt.; E. Crown detail with petiole

bases and leaf-base fibres (voucher specimen, Dowe et al. 606, from Kennedy River,

Twelve Mile Waterhole, Lakefield National Park, Queensland, Australia); F. Leaf; G.

Fruit, and seed in longitudinal section to reveal seed coat intrusion and embryo. (Photos:

J.L.Dowe; 18D, 18F, 18G based on Dowe 607 (holotype) from Barrett Creek, Cooktown,

Queensland, Australia).

o>)

ios)

ioe)

Gard. Bull. Singapore 60 (2) 2009

Plate 19. Livistona drudei: A. In habitat, coastal semi-open forest, Clemant State Forest,

Queensland, Australia; B. Crown, Balgal Beach, Queensland, Australia; C. Petiole bases

and leaf-base fibres; D. Fruit and endocarps. (Photos: J.L.Dowe; 19C & 19D from plant at

Hen Camp Creek, Queensland, Australia).

A Taxonomic Account of Livistona R.Br. (Arecaceae)

Plate 20. Livistona decora: A. \n habitat, riparian forest at Black Gully, Haughton River,

Queensland, Australia: B. Leaf detail; C. Flowers, Cape Cleveland, Queensland, Australia;

D. Fruit. (Photos: J.L.Dowe; 20B & 20D from plant at Cocoa Creek, Cape Cleveland,

Queensland, Australia).

340 Gard. Bull. Singapore 60 (2) 2009

Plate 21. Livistona australis: A. Ir habitat, rainforest at Eungella Range, Mt McCartney,

Queensland, Australia; B. Leaf detail (voucher specimen, Dowe 520, at Bluewater Range,

Queensland, Australia); C. Inflorescence with mature flowers; D. Flowers; E. Fruit

(voucher specimen, Dowe 171, from Seaview Range, Queensland, Australia. (Photos:

J.L.Dowe; 21C & 21D from plant (Dowe 3/3) at 800 m alt., Clarke Range, Cathu State

Forest, Queensland, Australia).

= - o = )

A Taxonomic Account of Livistona R.Br. (Arecaceae) 34]

prt

Plate 22. Livistona nitida: A. In habitat, riparian forest; B. Leaf; C. Crown with

inflorescence and mature flowers. (Photos: J.L.Dowe, from Robinson Creek, Queensland.

Australia). Livistona fulva: D. In habitat, semi-open woodland at base of sandstone

escarpment; E. Leaf abaxial view. (Photos: J.L.Dowe: voucher specimen, Dowe 300, from

Blackdown Tableland, Queensland, Australia).

342 Gard. Bull. Singapore 60 (2) 2009

Plate 23. Livistona alfredii: A. In habitat, riparian forest with scribbly gums; B. Juvenile

with petiole bases and leaf-base fibres. (Photos: J.L.Dowe; voucher specimen, Dowe

354, from Millstream-Chichester National Park, Crossing Pool, Fortescue River, Western

Australia). Livistona nasmophila: C. In habitat, permanent spring; D. Crown with mature

inflorescence; E. Leaf detail. (Photos: J.L.Dowe; voucher specimen, Dowe 363, from

Zebedee Springs, Western Australia).

oS)

A Taxonomic Account of Livistona R.Br. (Arecaceae) 34

Plate 24. Livistona mariae: A. In habitat, open woodland in semi-arid zone at Finke Gorge

National Park, Palm Creek, Northern Territory, Australia. (Photo: J.L.Dowe). Livistona

rigida: B. In habitat, riparian semi-closed forest at Lawn Hill Gorge, Queensland, Australia;

C. Crown detail with petiole bases and leaf-base fibres (voucher specimen, Dowe et al.

630, from Adels Grove, Lawn Hill Creek, Queensland, Australia); D. Fruit, and seed in

longitudinal section to reveal seed coat intrusion, (voucher specimen, Dowe 2/0, from

Roper River, Mataranka, Northern Territory, Australia); E. Seedling, with distinctive

red colouration (voucher specimen, Dowe 209, from Salt Creek, Roper River, Northern

Territory, Australia). (Photos: J.L.Dowe).

344 Gard. Bull. Singapore 60 (2) 2009

Plate 25. Livistona lanuginosa. A. \n habitat, semi-open riparian forest at Deep Creek; B.

Crown with petiole bases, leaf-base fibres and peduncular bracts; C. Infructescence with

mature fruit; D. Flowers; E. Fruit, with longitudinal sections of entire fruit, and seed to

reveal seed coat intrusion and embryo (voucher specimen, Dowe 292, from Glenroy Creek,

Queensland, Australia). (Photos: J.L.Dowe, 25A-25D from Campaspe River, Queensland,

Australia).

Gardens’ Bulletin Singapore 60 (2): 345-349. 2009 34

Does the Moss Genus Lepidopilum (Brid.) Brid.

(Pilotrichaceae) Occur in Asia?

B.-C. HO' AND B.C. TAN?

' Nees-Institut ftir Biodiversitat der Pflanzen,

Rheinische Friedrich-Wilhelms-Universitat Bonn,

Meckenheimer Allee 170, 53115 Bonn, Germany

E-mail: calyptrochaeta@yahoo.com

> Herbarium, Singapore Botanic Gardens, | Cluny Road, Singapore 259569

Abstract

The Asian species of Lepidopilum (Brid.) Brid. are reviewed. Hitherto,

L. novae-guineae E.B. Bartram is the only remaining Asian species in

the predominantly neotropical genus. However, examination of the type

specimen of this taxon proves it to be a synonym of Dimorphocladon

borneense Dixon.

Introduction

The genus Lepidopilum (Brid.) Brid. (Pilotrichaceae sensu Buck et al.,

2005) is a predominantly neotropical moss genus, with a few reported

outlier representatives in the paleotropics. Although the checklist of mosses

(Crosby et al., 1999) listed 61 accepted valid species names known in the

world, Churchill (1992) had estimated that about 35-40 will remain as good

species.

The main distinguishing characters of the genus include: (1)

prostrate primary axes with ascending secondary axes; (2) distinct double

costae extending at most to midleaf; (3) median laminal cells smooth and

narrowly hexagonal; and (4) leaf border indistinct (Welch, 1962, 1966). In

addition, Lepidopilum is traditionally separated from the gametophytically

similar Lepidopilidium (Mill. Hal.) Broth. solely by its papillose peristome

teeth, which are neither striolate nor furrowed. Buck ef a/. (2005) had

remarked that the true delimitation between these two genera is yet to be

resolved with certainty.

Due to our research interest on the Hookeriaceae sensu /ato in Asia,

we noted that only a single species of Lepidopilum is accepted today from

Asia. Hitherto, 11 Asian species names had been published either originally

as, or transferred later to the genus Lepidopilum (Wijk et al., 1964, 1969).

346 Gard. Bull. Singapore 60 (2) 2009

According to the present day taxonomy, all, except one taxon, have been

referred to other Hookeriaceous genera (Table 1). Remarkably, many of

them are, in fact, the synonyms of Hookeriopsis utacamundiana (Mont.)

Broth. [syn. Thamniopsis utacamundiana (Mont.) W.R. Buck], a very

variable species in the same family Pilotrichaceae. It is also interesting to

note that the two known species of Actinodontium Schwagr. in tropical Asia

were also once considered by Brotherus (1907) to belong to Lepidopilum,

but no one seems to have accepted this view today. The only remaining

Lepidopilum species left in Asia is L. novae-guineae E.B. Bartram from

Papua New Guinea, a species that is known only from the type collection.

Its generic affinity needs re-examination at present.

When Bartram (1961) described L. novae-guineae, he had noted

the narrow leaf outline, the elongate laminal cells, and the smooth seta, as

characters that do not fit into Lepidopilum. He had, nevertheless, decided to

tentatively place it in this genus based on the well-developed double costae,

the calyptral ornamentation, and the zig-zag median line of the peristome

> teeth:

Material and Method

To be able to ascertain the true identity of L. novae-guineae, we have

requested for a study of the holotype material from the Farlow Herbarium

of Cryptogamic Botany (FH).

Results and Discussion

In addition to Bartram’s (1961) observations on the above stated “non-

Lepidopilum characters”, other striking features of the type material

include: (1) prorate laminal cells, (2) the bi-geminate teeth at leaf margins,

and (3) the dimorphic branches, namely laxly foliated, short complanate

ones and densely foliated, longer penicillate ones. All together, this species

stands clearly out of place in Lepidopilum. Moreover, the type specimen

is an epiphyllous moss, a habitat rather uncommonly seen in species of

Lepidopilum.

After considering all the distinctive morphological and ecological

features, we have come to the conclusion that L. novae-guineae belongs

to the genus Dimorphocladon Dixon (Symphyodontaceae sensu Buck &

Goffinet, 2000). Upon comparing the type of L. novae-guineae with the

description and authentic specimens of Dimorphocladon borneense Dixon

Does the Moss Genus Lepidopilum Occur in Asia? 347

(1922), also an epiphyllous moss, we find no significant difference in their

morphology, thus, we are proposing the following new synonymy.

Dimorphocladon borneense Dixon, J. Bot. 60 (1922) 109, t. 564: f. 5 a-f.

— Type: Borneo. Upper Sarawak, A.H. Everett, mixed with Taxithelium sp.

(holo, NY-Mitten n.s.)

Lepidopilum novae-guineae E.B. Bartram, Brittonia 13 (1961) 373, syn.

nov. -Chaetomitrium novae-guineae (E.B. Bartram) S.P. Churchill, Rev.

Moss Gen. Lepidopilum (1988) 185, nom. inval. — Type: Papua New

Guinea. Sepik District: Wewak-Angoram Area, near Nagipem village,

Prince Alexander Range, Maprik-But track, on palm frond, rain forest,

1500 ft. 28 Jul 1959, Robbins 1990 (holo, FH!). Figs. 1 & 2.

Figure 1. Habit of Lepidopilum novae-guineae E.B. Figure 2. Habit of lateral,

Bartram showing the two kinds of branches, one penicillate branches of Lepido-

with elongate penicillate tip (based on Robbins 1990, pilum novae-guineae (Robbins

holotype at FH). (Photo: Y.H.Lim) 1990, FH) showing the two

kinds of leaves seen also in plant

specimen of Dimorphocladon

borneense. (Photo: Y.H.Lim)

With the inclusion of L. novae-guineae, the genus Dimorphocladon

still remains a monotypic taxon. The taxon is known today from New

Guinea, Seram, Philippines (Palawan), Borneo, Sumatra, Malay Peninsula,

and Thailand (Akiyama, 1997; Dixon, 1922, 1932, 1935; Schultze-Motel,

1965; Tan, 1993).

348 Gard. Bull. Singapore 60 (2) 2009

Churchill (1988), in his unpublished doctoral thesis on the revision

of Lepidopilum, had already excluded L. novae-guineae from the genus.

In fact, he made a new combination for this species in Chaetomitrium

Dozy & Molk. as C. novae-guineae. Although, this new binomial was not

effectively published according to the current ICBN rules, his taxonomic

interpretation of the New Guinean material as a species of Chaetomitrium

is rather close to the correct identity of this taxon.

We concur with Dixon (1922) that Dimorphocladon 1s closely allied

to Chaetomitrium. Furthermore, we agree with Tan and Robinson (1990)

and in part with Buck & Goffinet (2000) in placing them together with

Chaetomitriopsis M. Fleisch. in the same family, Symphyodontaceae, close

to the Hypnaceae.

Table 1. Asian moss species that were once placed in Lepidopilum.

Previously in Lepidopilum Currently accepted

L. adscendens (Schwagr.) Broth. => Actinodontium adscendens Schwagr.

L. furcatum Thwaites & Mitt. => Lepidopilidium furcatum (Thwaites & Mitt.)

Broth.

L. macropus Bosch & Sande Lac. — => Hookeriopsis utacamundiana (Mont.) Broth.

L. novae-guineae E.B. Bartram => Dimorphocladon borneense Dixon [proposed

here]

L. purpuratum Mitt. => Hookeriopsis utacamundiana (Mont.) Broth.

L. rhaphidostegum (Mill. Hal.) => Actinodontium rhaphidostegum (Mill. Hal.)

Broth. Bosch & Sande Lac.

L. secundum (Griff.) Mitt. => Hookeriopsis utacamundiana (Mont.) Broth.

L. spinosum (Mill. Hal.) A. Jaeger ==> Cyathophorum spinosum (Mill. Hal.) M.Fleisch.

L. sumatranum Bosch & Sande Lac. => Hookeriopsis utacamundiana (Mont.) Broth.

L. thwaitesianum Mitt. => Hookeriopsis thwaitesiana (Mitt.) Broth.

L. utacamundianum (Mont.) Mitt. => Hookeriopsis utacamundiana (Mont.) Broth.

References

Akiyama, H. 1997. Taxonomic studies of mosses of Seram and Ambon

(Moluccas, East Malesian) collected by Indonesian-Japanese Botanical

Expeditions VIII. Meteoriaceae, Hookeriaceae, and Trachypodaceae.

Nature and Human Activities 2: 9-31.

Bartram, E.B. 1961. Low altitude mosses from northeastern New Guinea.

Brittonia 13: 368-380.

Brotherus, V.F. 1907. Bryales. In: A. Engler & K. Prantl. (eds.). Die

Gardens’ Bulletin Singapore 60 (2): 351-372. 2009 3511

Seven New Species of Begonia (Begoniaceae)

From the Ulu Merirai and Bukit Sarang Limestone Areas

in Sarawak, Borneo

R. KIEW' AND J. SANG?

' Forest Research Institute Malaysia, Kepong, Selangor, Malaysia

* Sarawak Herbarium, Forest Research Centre, Kuching, Sarawak

Abstract

Seven new species are described from Sarawak, Borneo: two, Begonia

payung S.Julia & Kiew and B. sarangica Kiew & S.Julia, are endemic to

Bukit Sarang in Ulu Sungai Kakus, Tatau District; and five, B. juliasangii

Kiew, B. meriraiensis S.Julia & Kiew, B. nagaensis Kiew & S.Julia, B.

piring Kiew & S.Julia and B. rhodochaeta S.Julia & Kiew, are endemic in

the Ulu Merirai area in the Tatau District.

Introduction

Limestone hills in Sarawak, Borneo, are found majorly in the Kuching

District (the Bau, Penrissen, and Padawan-Serian areas), and, distant from

these, the Subis limestone (including the famous Niah Cave), the Mulu

limestone of which Gunung Api rises to about 1,700 m and is the highest

limestone peak in Malaysia, and the largely unknown limestone hills in the

Tatau District, at Bukit Sarang and the Ulu Merirai area.

In Borneo, begonias that grow on limestone are diverse and

endemism is high. For example, in Sabah, two thirds of the 18 species

occur on just one limestone hill and almost a further third on hills within the

same area (Kiew, 2001). Only one species, Begonia gueritziana Gibbs, is

widespread and it is the only species that grows on other rock types besides

limestone. Because of the many microhabitats available on limestone,

frequently several begonias can be found on a single hill each occupying

a different niche. In Sabah, up to four species were found on a single hill

(Kiew, 1998).

A similar pattern is emerging for begonia diversity and distribution

in Sarawak. In fact, limestone begonias are proving to be more species-rich

than in Sabah. Already 28 species are recorded from Sarawak compared

with 18 for Sabah. As in Sabah, begonia species are confined to one hill

352 Gard. Bull. Singapore 60 (2) 2009

or a group of hills. For example, five begonia species are recorded from

the Subis limestone (Pearce, 2005); six from the Bau limestone (Kiew &

Connie, 2003); and eight from each of the Penrissen and Padawan-Serian

areas (Kiew & Julia, 2007). There are no begonia species in common

between the Subis limestone and the rest of Sarawak. Of the 21 species in

the Kuching District, only B. speluncae Ridl. occurs in the Bau, Penrissen,

and Padawan-Serian areas, a third of the species are found in two of these

three areas, but the majority (60%) are restricted to one of the three areas.

As in Sabah, individual hills usually support more than one species (Kiew

& Julia, 2007) with five species on Gunung Kawa (Bau limestone) and

four each on G. Manok (Padawan-Serian limestone) and G. Bah (Penrissen

limestone).

The surveys of the Bukit Sarang and Ulu Merirai begonias show

similar high endemism and restricted distributions. The two begonias that

grow on Bukit Sarang are found nowhere else as are the five from the Ulu

Merirai.

. Among begonias in Borneo, section Petermannia is the most

speciose and the limestone begonias are no exception. The cane-like and

creeping species belong to this section. Of the 28 species described from

Sarawak limestone, 20 belong to this section. They grow at the base of

the hills, or in gullies or on ledges on limestone derived soil, rather than

on bare rock. One group among them is distinct for the strongly corrugate

leaves, and is represented on all the hills, each with its own species — B.

congesta Ridl. in Bau, B. corrugata Kiew & S.Julia in Padawan-Serian and

Penrissen, B. niahensis K.G.Pearce on Subis, B. sarangica Kiew & S.Julia

on Bukit Sarang, and B. nagaensis Kiew & S.Julia on Ulu Merirai.

The next most speciose section on limestone with six species is

sect. Reichenheimia Group | of Doorenbos ef a/. (1998) that includes the

peltate or orbicular-leaved species. These are characteristic of shaded

sheer vertical cliff faces or around cave mouths. In the Kuching District,

B. speluncae Ridl. is the most common and widespread; B. payung S.Julia

& Kiew occurs on Bukit Sarang; and B. juliasangii Kiew on Ulu Merirai.

(The Subis flora does not include a peltate begonia). B. rhodochaeta S.Julia

& Kiew also belongs to this section even though it does not have peltate

leaves.

Only two limestone species belong to sect. Dip/oclinium Group I

of Doorenbos et al. (1998). One is B. calcarea Ridl. from the Kuching

Division; the other is B. piring Kiew & S.Julia from Ulu Merirai. The latter

is less usual for this section in having peltate leaves.

os)

is)

Seven New Species of Begonia (Begoniaceae)

Bukit Sarang

Bukit Sarang (02° 39° 12”N, 113° 03° 05”E) lies in Ulu Sungai Kakus in

the Tatau District and includes two adjacent low hills: Batu Anyi (140 m at

the summit) and Batu Lebik that is both smaller and slightly lower. Unlike

most karst limestone, there are no sheer cliff faces, the hills being broken by

narrow gullies deeply shaded and with a thick litter layer. These two hills

are also unique for Bornean limestone, in being surrounded by freshwater

swamp. Until our survey in October 2004 (Kiew ef a/., 2007), the flora was

very little known botanically having been visited only by J.A.R. Anderson

in 1965, where in a day, he collected 25 herbarium specimens, including

both the peltate and cane-like begonias, but which until this account have

remained undescribed. Both species are endemic to the Bukit Sarang

limestone and with just two species, this area is the least biodiverse of

Sarawak limestone.

Ulu Merirai

We surveyed the flora of the Ulu Merirai limestone in the Tatau District

(02° 46° 13.7°N, 113° 39° 02.9"E) because it was said that Gunung Lumut,

the largest hill in the area, was limestone. In fact it proved to be sandstone

and the limestone in the area, rather than consisting of tower karst, was

represented by exposed cliff faces on steep slopes, most below the tree

canopy and only a few, Gua Aki Bukok, Gua Naga and Gua Tiang, rising

sheer above the tree canopy to 430 m. Many of the caves are underground

with their mouths more or less at ground level. During this trip, a total of

nine limestone outcrops were surveyed.

This area proved to be more diverse than Bukit Sarang with five

species discovered, all new species described below. All are endemic to the

Ulu Merirai limestone. Only B. nagaensis was restricted to a single hill.

Most hills supported just two species, only Gua Tiang had three.

Most striking were the two peltate species (B. piring Kiew & S.Julia

with remarkable whitish green succulent circular leaves with a raised

crimson rim, and B. juliasangii Kiew whose peltate leaves are unusual in

being narrowly oval) and B. nagaensis Kiew & S.Julia, a cane-like begonia

that had iridescent peacock-blue leaves.

354 Gard. Bull. Singapore 60 (2) 2009

New Species

|. Begonia juliasangii Kiew, sp. nov.

Section: Reichenheimia

A Begonia baramensi Merr. habitu rhizomatoso, alis fructus angustioribus

2.5-6 mm latis (nec 10-15 mm) et placentis in quoque loculo singulis differt.

— Typus: Borneo, Sarawak, Tatau District, Ulu Merirai, trail to Gua Tiang,

6 Jul 2005, Julia et al. S 94658 (holo, SAR; iso E, KEP, SAR). Plate 1A-C.

Creeping herb climbing vertically for a short distance up cliffs and rocks,

on soil decumbent and rooting at the nodes. Stems rhizomatous, glabrous,

succulent, reddish brown, leafy stem up to 15 cm long and in life ca 1.3 cm

thick; internodes 0.5-10 cm long; unbranched; without a tuber. Stipules

greenish to pale reddish brown, glabrous, except for the sparsely hairy

margin and outer surface of the midrib, ovate, 10-23 x 6-10 mm, margin

- entire, apex setose, persistent. Leaves spirally arranged, distant, laminas

not oblique, pendent; petioles dark crimson to reddish brown or green,

7.5-13 cm long, terete, covered by translucent hairs up to 5 mm long;

laminas plain matt dark green above, pale green beneath, young leaves

bronze or pinkish above, pinkish beneath, glabrous, in life fleshy and

succulent, drying papery, peltate, elliptic to obovate, slightly asymmetric

with one side straight and the other slightly rounded, (15-)20-29 x (5-)7-9

cm long, broad side 3.5-5.5 cm wide, base rounded, peltate base very

short, 5-6(-8) mm long, margin very shallowly and distantly crenate in

the lower half, sometimes distinctly scalloped in upper half to one third

of the lamina with a pointed lobe extending from the middle vein, apex

acuminate; venation palmate-pinnate with a basal pair and 2-3 pairs along

the midrib with 2 minute veins in the peltate base, prominent on both

surfaces, main veins concolorous above, paler green and sparsely hairy

beneath. Inflorescences axillary, crimson, glabrous, erect, very slender,

2.5-4.5 cm long and shorter than petioles, a simple or once branched

cyme, in twos from the lower leaf axils; male flowers 4 or 9 and female

flowers 2 at base; peduncles 2-2.5 cm long, male branches ca 7.5 mm

long, female branch 5-13 mm long. Bracts clustered at the base, crimson,

lanceolate, 5-7 x 2-3 mm, glabrous, margin entire, apex setose, persistent;

bracteoles in pairs, similar to bracts but smaller, 2-3 x 1-2 mm. Male

flowers with a crimson pedicel, 23-26 mm long; tepals 4, white at the

base, deep pink along margin towards the apex, almost equal in size,

glabrous, ovate, margin entire, apex acute; outer tepals 11-13 x 9-10 mm,

inner tepals similar but slightly smaller, 9-10 x 6-7 mm; stamens 30-35,

ws)

¥ re ae f 7) oe 2

Seven New Species of Begonia (Begoniaceae)

et aL

Plate 1. Begonia juliasangii Kiew. A. Habit; B. Male flowers; C. Female flowers. Begonia

payung S.Julia & Kiew. D. Habit. (Photos: J. Sang).

EEE

356 Gard. Bull. Singapore 60 (2) 2009

cluster hemispherical, ca 4 mm diam., stalk ca 1 mm long; filaments pale

yellow, ca 1.2 mm long; anthers pale yellow, ellipsoid, ca 1.2 mm long,

apex slightly emarginate, dehiscing along lateral slits. Female flowers

with glabrous, rosy red pedicels 2-5 mm long; ovary light crimson, broadly

obovate, 8-10 x 9-10 mm, wings 3 (rarely 4), equal, locules 3, placentas |

per locule; tepals 5, isomorphic, white at the base and rosy red along the

margin, elliptic, ca 7 x 5-6 mm, margin entire, apex slightly acute; styles

and stigmas ca 2 mm long, styles 3, yellow, bifurcating and U-shaped;

stigmas forming a continuous twisted band. Fruits on the lower part of

the stem, pendent; peduncle and pedicel slender 1.7-3 cm long; capsules

deep crimson, glabrous, ovoid, 16-18 x 10-19 mm, wings 3, equal, narrow,

2.5-6 mm wide and tapering to the base and apex, thinly fibrous, dehiscing

between locules and wings. Seeds barrel-shaped, ca 0.3 mm long, collar

cells ca 4/5 seed length.

_ Other specimens examined: Borneo. Ulu Merirai. Gua Tiang, Malcom et al.

S 95535 (SAR); above Gua Pak Danum, 12 Jul 2005, Leong et al. PL 314

(SAR, SING).

Habitat: Restricted to limestone, on the base of vertical, deeply shaded

cliffs or within cave mouths or sometimes rooting in soil at the cliff base.

Distribution: Endemic in Borneo, Sarawak; known only from the Ulu

Merirai area in Tatau District.

Notes: This is a very distinct begonia, quite unlike any other Bornean

limestone species in its elliptic, scarcely peltate leaves. Other limestone

peltate species have almost rotund leaves with the petiole attached towards

the centre of the lamina. In its elliptic to obovate peltate leaves, it most

resembles Begonia baramanesis Merr. but otherwise it is quite different

because B. baramensis is a cane-like begonia to 60 cm tall (not creeping),

the leaves are shorter and distinctly oblique with petioles 2.5-4.5 cm long

and lamina 10-17.5 cm long. In addition, the peltate base is more pronounced

being 2.5-4.5 cm long. The fruit too is very different in being large (16-25 x

28 mm) and having downward pointing wings 10-15 mm wide. In addition,

B. baramensis is placed in a different section, sect. Petermannia, because

it has three locules each with two (not one) placentas. (Doorenbos ef al.

were doubtful as to its placement in sect. Petermannia but dissection of the

ovary of S 3/503 confirms its position).

The species is named for Julia Sang, who first discovered this

begonia.

¥ co - - = VE

Seven New Species of Begonia (Begoniaceae) 357

2. Begonia meriraiensis S Julia & Kiew, sp. nov.

Section: Petermannia

A Begonia paoana Kiew & S.Julia inflorescentiis brevioribus 9-10 cm longis

(nec 11-25 cm), fructibus angustioribus 14-20 mm latis (nec 21-25 mm)

et pedicellis in fructu 5-10 mm longis (nec 10-23 mm) differt. — Typus:

Borneo, Sarawak, Tatau District, Ulu Merirai, Gua Tiang 6 Jul 2005 Julia

et al. S 94657 (holo SAR; iso E, KEP, SAN, SAR, SING, K, L).

Shrubby cane-like begonia, densely hairy, hairs 3 mm long. Stems

ferruginous, erect, woody, to 75 cm tall, in life 8 mm thick, 4 mm thick

when dried; internodes 3-5.7-8.5 cm long: without a tuber. Stipules

light green, broadly triangular, 17 x 8-10 mm, margin entire, apex acute,

caducous. Leaves spirally arranged, distant, held horizontally: petioles

ferruginous, 1-1.7(-3.2) cm long, terete; laminas scarcely oblique, plain

glossy dark green above, light green below, in life corrugate and thinly

succulent, papery when dried, broadly oval to slightly ovate on the broad

side, scarcely asymmetrical, (14-)18.5-20(-25.5) x (8-)10-12(-15) cm, broad

side (5-)6.5-7.5(-9.5) cm, base unequal, basal lobe rounded sometimes

auriculate, 1-2.5 cm long, margin minutely and distantly toothed at the

veins ending, apex caudate, acumen 0.5-2 cm long; venation palmate-

pinnate, veins concolorous, ferrugineous beneath, 1-2 pairs at the base,

4-5(-6) pairs along the midrib, 2 veins in the basal lobe, branching three

times, in life deeply impressed above, slightly raised when dried, prominent

below. Inflorescences axillary from the upper leaf axils, racemose, hairy,

erect, up to 9-10 cm long, beginning to flower at ca 2 cm long; peduncles

stout, ca 2.5 mm thick, reddish to brownish, 2-3 cm long, branches up to

0.5 cm long and 1-2 cm apart, above male flowers many, below female

flowers 2, protogynous. Bracts ferrugineous, 10-13 x 5-8 mm; bracteoles

yellowish green. Male flowers with densely and minutely hairy pedicels,

3-8 mm long; tepals 2, outside rosy red and densely and minutely hairy,

inside greenish cream and glabrous, oval to round, 4-6 x 4-6 mm, recurved

when open, margin entire, apex acute to rounded; stamens ca 35, stamen

cluster conical, subsessile, stalk ca 0.5 mm long; filaments 0.5-0.7 mm

long; anthers dull yellow, obovate, ca 0.7 mm long, apex rounded, dehiscing

along lateral slits. Female flowers with minutely hairy pedicels, 4-7 mm

long; ovary pale green, minutely hairy, ellipsoid, ca 18 x 10-12 mm, wings

3, equal, locules 3, placentas 2 per locule: tepals 5 almost isomorphic,

pale green, pinkish at base, minutely hairy outside, ovate, 9-12 x 7-9 mm,

toothed towards the apex; styles 3, lemon-yellow, 5-6 mm long, bifurcating,

stigmas forming a spiral papillose band. Fruits pendent on thick stalks 7-10

358 Gard. Bull. Singapore 60 (2) 2009

mm long, capsules narrowly oblong, narrowed to base, 20-28 x 14-20 mm,

wings 3 (rarely 4), equal, truncate distally, 5-10 mm wide, glabrous, thinly

fibrous, dehiscing between the locules and wings. Seeds barrel-shaped, 0.3

mm long, collar cells more than half the seed length.

Other specimens examined: Borneo. Sarawak, Ulu Merirai. Gua Aki Bukok,

Malcom et al. S 95606 (E, KEP, SAR, SING); ibid, Malcom et al. S 95631

(SAR); Gua Naga, 11 Jul 2005, Leong et al. PL 278 (SAR, SING).

Habitat: Extremely common from the lower to upper slopes in limestone

forest, on calcareous soil as well as on litter-covered limestone rocks.

Distribution: Endemic in Borneo, Sarawak; known only from the Ulu

Merirai area in Tatau District.

_ Notes: Begonia meriraiensis is the most common and widespread limestone

species found in Ulu Merirai area, so is named for this locality. It is a cane-

like begonia similar to B. paoana Kiew & S.Julia from limestone in the

Kuching Division, but from which it can be distinguished by the following

characters (see Table 1).

Table 1. A comparison of Begonia meriraiensis and B. paoana.

B. meriraiensis B. paoana

Stem densely hairy (hairs to 3 mm long) minutely pubescent

-Stem diam. | 8mm 3-4 mm

Stipule triangular obovate

Lamina 14-25.5 x 8-15 cm 15-24 x 4.5-11 cm

(length:width = <2x) (length:width = >2x)

Basal lobe rounded, sometimes auriculate rounded

1-2.5 cm long 0.5-1.3 cm long

Apex caudate acuminate

Veins ferrugineus glabrous/minutely pubescent

Inflorescences 9-10 cm long 11-25 cm long

Bracts ferrugineus, 10-13 x 5-8 mm glabrous, 7 x 3 mm

Male flower pedicel pubescent pedicel glabrous

tepals minutely hairy tepals glabrous

Female flower ovary minutely hairy ovary glabrous

-Tepals 7-9 mm wide 4-6 mm wide

-Style 5-6 mm long 3-4 mm long

Fruits pedicels 5-10 mm long pedicels 10-23 mm long

capsule 14-20 mm wide capsule 21-25 mm wide

Seven New Species of Begonia (Begoniaceae) 359

3. Begonia nagaensis Kiew & S.Julia, sp. nov.

Section: Petermannia

A Begonia corrugata Kiew & S.Julia foliis cyaneis (colore pennarum

pavonis) 8.5-9 cm latis (nec 9-16 cm), inflorescentiis brevioribus 4-6 cm

longis (nec 13.5-23 cm) et fructibus ca 14 X 12 mm (nec 9-16 mm) differt.

— Typus: Borneo, Sarawak, Tatau District, Ulu Merirai, Gua Naga, 11 Jul

2005, Leong et al. PL 290 (holo, SAR; iso, SING). Plate 2A-B.

Cane-like begonia. Stems dull green, erect with a few branches, up to 75 cm

tall, 4 mm thick, bristly, bristles red when young, brown when old, woody,

sometimes thicker at the nodes, internodes 5.5-6 cm long; without a tuber.

Stipules pale green; densely hairy along the midrib, lanceolate, 9-25 x 8-10

mm, midrib densely hairy, margin entire, apex setose with the apical hair up

to 10 mm long, caducous. Leaves alternate, distant, slightly pendent; petioles

reddish with long bristles, 2-2.5 cm long, grooved above; laminas slightly

oblique to oblique, plain iridescent blue and velvety above with a red patch at

the base of the midrib, young leaves purple green above and purple beneath,

thin and corrugate in life, elliptic, asymmetric, 17.5-19 x 8.5-9 cm, broad side

5.5-6 cm wide, base cordate with rounded basal lobes 1.5-2 cm long, margin

minutely dentate, each tooth tipped by a hair, apex acuminate, acumen 075-1

cm long; venation palmate-pinnate, main veins slightly reddish and hairy

beneath, 2 pairs at the base, 3-5 pairs along the midrib and 3 veins in the

basal lobe, equal-sized, branching towards the margin, impressed above and

prominent beneath. Inflorescences axillary, reddish or reddish brown with

bristly red hairs ca 2 mm long, racemose with cymules of male flowers

spaced along the rachis and with | or 2 female flowers below, erect, longer

than the petioles, 4-6 cm long, sessile or with peduncles up to 1.3 cm

long, protogynous. Bract pair pale green, ovate, ca 10 x 6 mm, margin

entire, apiculate; bracteoles pale green, hairy, elliptic, ca 3 x 1.5 mm,

margin entire, apex apiculate, caducous. Male flowers with a glabrous, red

pedicel 2-6 mm long; tepals 2, reddish with red hairs outside, light pink

inside, almost round, 4-5 x 3.5-4 mm, margin entire and recurved, apex

rounded; stamens ca 30, stamen cluster sessile, globose, 2.5-3 mm across;

filaments ca 0.75 mm long; anthers pale yellow, obovate, ca 1 mm long,

apex emarginate, dehiscing along lateral slits. Female flowers with a red

to dark red, densely hairy pedicel ca 12 mm long; ovary fleshy, reddish

green, locules densely covered with dark red hairs on the locules (not on the

wings), broadly ellipsoid, ca 20 x 15 mm, wings 3, equal, 2.5-3 mm wide,

locules 3, placentas 2 per locule; tepals 5, covered with red hairs outside

and on the margin, light pink, elliptic, outermost ca 15 x 8 mm, innermost

Gard. Bull. Singapore 60 (2) 2009

360

Plate 2. Begonia nagaensis Kiew & S.Julia. A. Habit; B. Female flowers. Begonia

rhodochaeta S.Julia & Kiew. C. Habit; D. Male flower; E. Female flower. (Photos: J.

Sang).

ca 10 x 4 mm, margin entire, apex acute; styles 3, bifurcating, ultimate

branches spirally twisted, ca 5 mm long: stigmas forming a papillose spiral

twisted band. Fruits | or 2 per infructescence, pendent on a pale green,

stiff, minutely bristly stalk 12-13 mm long, capsules green with long red

hairs especially dense on the locule wall; ellipsoid, ca 14 x 12 mm, wings

3, equal, tapered to the base, slightly angled distally, ca 3 mm wide, thinly

fibrous, dehiscing between the locules and wings. Seeds barrel-shaped.

brown, ca 0.25 mm long, collar cells about 4/5 of the seed length.

Specimen examined: Known only from the type specimen.

Habitat: In limestone forest on a ledge above a steep slope at the base of

cliff face, on soil in light shade.

Distribution: Endemic in Borneo, Sarawak: known only from Gua Naga

above Sungai Bekuyat in Ulu Merirai area, Tatau District.

Notes: Begonia nagaensis is very local and was found at only one locality,

Gua Naga, for which it is named. The other begonias were found at two or

more localities.

This species can be easily recognised by its striking iridescent leaves

that like peacock feathers change from blue to bright green depending on

the angle of the light. It also has conspicuous dark red hairs on the male

and female flowers.

It is a cane-like begonia with corrugated leaves that, apart from its

peacock-blue leaves and capsule with red hairs, can be distinguished from

B. corrugata Kiew & S. Julia from the Kuching Division by its broader

lamina (8.5-9 cm vs. 9-16 cm), shorter inflorescences (4-6 cm not 13.5-23

cm long) and smaller capsule (14 x 12 mm not 15-20 x 12-15 mm).

4. Begonia payung S.Julia & Kiew, sp. nov.

Section: Reichenheimia

A Begonia speluncae Rid. petiolis longioribus 10-18 cm longis (nec usque 7

cm), foliis latioribus 6-11 cm latis (nec 4(-6) cm) et fructibus alis carentibus

differt.— Typus: Borneo, Sarawak, Tatau District, Ulu Sungai Kakus, Bukit

Sarang, Batu Anyi, 5 Oct 2004, Lee et al. SL 38 (holo, SAR; iso, SING).

Fig. 1, Plate 1D.

Small, creeping begonia, whole plant glabrous. Stems rhizomatous and

rooting at the nodes, young stem succulent and pale green becoming

362 Gard. Bull. Singapore 60 (2) 2009

4mm 2mm 2mm 4mm

Figure 1. Begonia payung S.Julia & Kiew. A. Habit; B. Male flower bud; C. Male flower;

D. Stamen; E. Female flower; F. Styles and stigmas; G. Transverse section of ovary; H.

Fruit. (Drawing by J. Pao).

Seven New Species of Begonia (Begoniaceae) 363

brown and woody, little branched, up to 11 cm long, in life 5-6 mm thick;

without a tuber. Stipules reddish, narrowly lanceolate, 5-7 x 4 mm, margin

slightly toothed, apex setose, persistent. Leaves up to 4-5 mm apart,

spreading over the cliff face; petioles pale green, 10-18 cm long, terete;

laminas sometimes bronzy green when young, mature leaves plain matt

dark green, in life thickly succulent, papery when dried, peltate, broadly

ovate, almost symmetric, 5-8.5 x 6-11 cm, broad side 2.7-5.5 cm wide,

base rounded and 1.5-2.5 cm long, margin crenate, apex acute; venation

palmate-pinnate, 3 pairs at the base with one pair along the midrib, and one

pair in the peltate base, branching ca two thirds of the way to the margin,

slightly raised above, impressed beneath, pale green, sometimes bronzy

green. Inflorescences axillary, whitish green, erect, longer than the leaves,

cymose panicles, (12.5-)15.5-20 cm long with two main branches 0.5-1

cm long, male flowers 5 to 11, female flowers 2 or 4, protandrous. Bracts

brownish, ovate, apex pointed, ca 10 x 3 mm, margin entire, persistent,

bracteoles similar but smaller, ca 5 x 3 mm. Male flowers with pale pink

pedicels 10-16 mm long; tepals 4, margin entire, apex rounded or slightly

acute, outer two, rosy pink outside, pale pink inside, rotund, 6-9 x 6-10 mm,

inner two pale pink inside and out, narrowly obovate or oval, (4-)7-10 x 2-3

mm; stamens ca 25, stamen cluster fan-shaped, 2.5-5 x 4-5 mm, stalk ca |

mm long, filaments ca 1.5 mm long, anthers pale yellow, narrowly obovate,

ca 1.5 mm long, apex emarginate, dehiscing along lateral slits. Female

flowers with reddish pedicels ca 5 mm long; ovary pale pink, rhomboid,

7-8 x 5-9 mm, 3-angled in cross section, without wings but with a deep

pink rib ca 0.5 mm wide along the angle of the ovary, locules 3, placenta

1 per locule; tepals 5, margin entire, apex rounded, outer two rotund, deep

pink outside, pale pink inside, 7-8 x 6 mm, inner three oblong, white, ca 4

x 2 mm; styles 3, bifurcating and U-shaped, styles and stigmas pale yellow,

3-4.5 mm long, stigmas forming a spiral papillose band. Fruits dangling

on fine and hair-like stalks 3-4 mm long, capsules 7-10 x 5-6 mm, locules

3, three-angled but without wings, ribs ca 1 mm wide, splitting between

the locules and the rib. Seeds brown, barrel-shaped, ca 0.25-0.3 mm long,

collar cells more than half the seed length.

Other specimen examined: Borneo. Ulu Sungai Kakus, Bukit Sarang, 14

Mar 1965, Anderson S 20958 (SAR).

Habitat: Vertical limestone cliffs in light or deep shade, common.

Distribution: Endemic in Borneo, Sarawak, known only from Ulu Sungai

Kakus, Bukit Sarang (Batu Anyi and Batu Lebik) in the Tatau District.

364 Gard. Bull. Singapore 60 (2) 2009

Notes: Begonia speluncae Ridl., from the Kuching Division, also has

peltate leaves and belongs to sect. Reichenheimia. However, it differs from

B. payung in being a diminutive plant with orbicular leaves up to 4 (rarely

6) cm across and with petioles up to 7 cm long. In addition, the wingless

capsules of B. payung are most unusual among Bornean begonias that in

general have winged capsules or exceptionally have fleshy indehiscent

berries. In B. payung, the wings are reduced to a narrow rib less than | mm

wide at the extremities of the angular fruit. In contrast, B. speluncae has

capsules with distinct wings.

We have named this species Begonia payung (payung is the Malay

word for umbrella) in reference to its round peltate leaf. [Anderson S20958

(SAR) was annotated as Begonia laxiflora by Irmscher, but this name was

never published].

5. Begonia piring Kiew & S.Julia, sp. nov.

_ Sectiom: Diploclinium

A Begonia payung S.Julia & Kiew internodiis longioribus 12-35 mm longis

(nec 4-5 mm), marginibus folii coccinei, fructibus subsessilibus trialatis

et placentis bifidis differt. — Typus: Borneo, Sarawak, Tatau Destrict, Ulu

Merirai, Sungai Bekuyat, Gua Aki Bukok, 10 Jul 2005, Julia et al. S 91200

(holo, SAR, iso, AAU, E, K, L, MO, SAN, SAR, SING). Plate 3A-E.

Rhizomatous, glabrous begonia. Stems creeping and rooting on vertical

cliff faces, succulent, pale green, almost translucent, brittle in life, up to

30 cm long and 5 mm thick, internodes 1.25-3.5 cm long; without a tuber.

Stipules pale green; broadly triangular, 8-16 x 4-6 mm, margin entire, apex

acute to apiculate, persistent. Leaves slightly spirally arranged, distant,

pendent; petioles pale green, translucent, 10.5-17.5 cm long, grooved

above; laminas oblique, plain whitish green, surface scintillating, succulent

and brittle in life, tissue paper thin when dried, peltate, broadly elliptic,

slightly asymmetrical, 7-10 x 8-10 cm, broad side 5-6.5 cm, base rounded,

projecting 0.5-1.5 cm beyond petiole attachment, margin crimson, undulate

or scalloped and acute at the vein endings, apex acute; venation palmate-

pinnate, main veins slightly paler than the lamina, 2 pairs at the base of and

4 pairs along the midrib, | pair in the base, plane above, slightly prominent

beneath. Inflorescences axillary, crimson, cymose panicles branched 4 or

more times, erect, as long as the petioles, 18.5-33.5 cm long, peduncle

succulent, 7.5-12 cm long and ca 5 mm thick, male flowers many on the

upper branches, female flowers 2 below on a branch 2.2-3.5 cm long,

protandrous. Bract pairs pale green, obovate, 10-11 x 4 mm, margin entire,

S = : aE

Seven New Species of Begonia (Begoniaceae) 365

Plate 3. Begonia piring Kiew & S.Julia. A. Habit; B. Leaf; C. Male flowers: D. Close up

of a male flower; E. Female flower. (Photos: J. Sang).

apex broadly truncate or rounded, caducous; bracteole pairs similar to

bracts but smaller, 3-7 x 1.5-2 mm. Male flowers with a pinkish pedicel

5-6 mm long; tepals 4, white or pinkish with pink or red entire margin,

apex rounded, glabrous, outer two round, 4.5-6 x 5 mm, inner two narrowly

obovate, 4-5.5 x 1.5-2 mm; stamens ca 20; stamen cluster sessile, conical,

2-2.5 mm diam.; filaments ca 0.5 mm long; anthers pale yellow, narrowly

366 Gard. Bull. Singapore 60 (2) 2009

obovate, ca | mm long, apex emarginate, dehiscing along lateral slits.

Female flowers with a crimson pedicel ca 2 mm long; ovary crimson or

pinky white, ca 8 x 8 mm, wings 3, equal, locules 3, placentas 2 per locule;

tepals 5, reddish at the margin, whitish at the base, outer three oval, ca 10

x 7 mm, apex rounded, inner two ovate, ca 8 x 5 mm, apex acute; styles 3,

bifurcating with ultimate branches broadly U-shaped, styles and stigmas

pale yellow, ca 4 mm long. Fruits almost sessile or on a stiff stalk up to 3

mm long, pendent; capsules broadly obovoid, 13-17 x 12-14 mm, wings 3,

equal, tapered to the base, slightly angular distally, 2-3(-5) mm wide, thinly

fibrous, dehiscing between the locules and the wings. Seeds barrel-shaped,

brown, ca 0.3 mm long, collar cells three quarters the seed length.

Other specimens examined: Borneo. Sarawak, Ulu Merirai, Gua Kenyalang,

Julia et al. S 91194 (E, KEP, SAR); Gua Kenyalang, 9 Jul 2005, Leong et

al. PL 214 (SAR, SING).

- Habitat: On sheer limestone cliff faces or on the arch of cave mouths at or

above the level of the tree canopy, sometimes fully exposed to sunlight.

Distribution: Endemic in Borneo, Sarawak; known from Gua Kenyalang

and Gua Aki Bukok in the Ulu Merirai area, Tatau District.

Notes: This is an unusual begonia with almost transparent, whitish green,

very succulent stems and petioles and round leaves with a raised deep-red

broad margin, hence the name ‘piring’, which means saucer in Malay. The

inflorescences are crimson with attractive crimson flowers with a white

centre and the ovary is crimson or pinky white. Begonia piring can be

distinguished from the other peltate limestone, B. pavung by the following

characters (see Table 2).

6. Begonia rhodochaeta S Julia & Kiew, sp. nov.

Section: Reichenheimia

A Begonia stichochaeta K.G.Pearce petiolis longioribus 3.5-8 cm longis

(nec 0.7 cm), inflorescentiis brevioribus 1-2.2 cm longis (nec ca 3.4 cm),

Jructibus latioribus quam longioribus et placentis in quoque loculo singulis

differt. —Typus: Borneo, Sarawak, Tatau District, Ulu Merirai, trail to Gua

Spring, 8 Jul 2005, Julia et al. S 91169 (holo, SAR; iso, E, K, KEP, L, SAN,

SING). Plate 2C-E.

Rhizomatous begonia, the prostrate stem rooting at the nodes. Stems

Seven New Species of Begonia (Begoniaceae)

Table 2. A comparison between Begonia piring and B. payung.

§ é payun;

367

B. piring

B. payung

Stems

Internodes

Stipules

Lamina

Veins

Male flowers

-tepals

-outer tepals

-inner tepals

-stamen cluster

-filaments

Female flower

Fruit

succulent, translucent

12-35 mm long

broad triangular

8-16 x 4-6 mm

whitish green, scintillating

brittle when fresh

base projecting 5-15 mm

margin crimson

2 pairs at base, 4 pairs along

midrib

pedicels 5-6 mm long

white with red margin

4.5-6 x 5mm

4-5.5 x 1.5-2 mm

conical

0.5-1 mm long

pedicels 2 mm long

placentas 2 per locule

pedicels 3 mm long, stiff or

almost sessile

capsules 13-17 x 12-14 mm

wings 3, 2-5 mm wide

not succulent, not translucent

4-5 mm long

narrowly lanceolate

5-7 x 4mm

bronzy green when young to

mid-dark green

not brittle when fresh

base projecting 15-25 mm

margin concolorous

3 pairs at base, | pair along

midrib

pedicels 10-16 mm long

rosy pink

6-9 x 6-10 mm

4-10 x 2-3 mm

fan-shaped

1.5 mm long

pedicels 5 mm long

placenta | per locule

pedicels 3-4 mm long, fine

and dangling

capsules 7-10 x 5-6 mm

without wings, with narrow r

ibs

succulent, dark green, densely bristly, unbranched, erect part of the stem to

10 cm tall and 7-8 mm thick with leaves in a tuft at the top, lower internodes

to 1.4 cm long, upper ones touching; without a tuber. Stipules green or dark

red, narrowly lanceolate, 8-14 « 4-7 mm, midrib densely bristly, margin

entire, apex attenuate. Leaves spirally arranged, upstanding; petioles dark

red when young, densely bristly, bristles brownish, (3.5-)5.5-8 cm long,

grooved above; laminas not oblique, plain glossy green, paler beneath,

young leaves pink with a crimson midrib, upper surface with scattered

bristles between the veins, glabrous beneath, slightly asymmetric, obovate,

(13.5-)18-23(-27) x (5.5-)7-10(-13) cm, broad side 3-7 cm wide, concave

on the narrower side, base tapering or cuneate, rarely rounded, basal lobe up

to 3 mm long, margin red, serrate and scalloped toward the apex, distantly

ciliate, apex attenuate, acumen to 2-3 cm long, leathery when dried, margin

red; venation pinnate, veins (4-)5 pairs along the midrib, glabrous above,

prominent beneath with red bristles, impressed above. Inflorescences

axillary, deep red from the upper leaf axils, hairs red in life, erect, shorter

368 Gard. Bull. Singapore 60 (2) 2009

than the petioles, protogynous, at first with 2 female flowers produced in

succession, followed by slender lax cymes 1-2.2 cm long each comprised

of 3 male flowers. Bract pair dirty red, narrowly lanceolate, ca 7 x 2-3 mm,

midrib densely bristly, margin entire, apex setose, caducous; bracteole pairs

broadly ovate, 2-4 x 1.5-2 mm, margin entire, caducous. Male flowers with

dark red pedicel 10-26 mm long; tepals 4, white, outside pink toward the

base, round, glabrous, margin entire, outer two 10-12 x 7 mm, apex slightly

acute, inner two 10-11 x 6-9 mm, apex rounded; stamens 25-30, stamen

cluster sessile, globose, ca 3 mm diam.; filaments ca 1.5 mm long; anthers

bright yellow, narrowly obovate, ca | mm long, apex rounded, dehiscing

along lateral slits. Female flowers with a dark red pedicel 9-12 mm long;

ovary red, broadly ellipsoid, 11-13 x 9-12 mm, wings 3, equal, locules 3,

placenta | per locule; tepals 5, pinkish, glabrous, elliptic, margin entire, apex

rounded, outermost ca 7 x 4 mm, innermost similar but narrower, ca 7 x 2.5

mm; styles 3, bifurcating, ultimate branches broadly U-shaped, 2.5 mm long,

_ pale yellow. Fruits maturing on the prostrate rhizome, stalk slender, 12-25

mm long, capsules glabrous, campanulate in outline, 12-15 x 13-18 mm,

wings 3, equal, extremely thin, 2-6 mm wide, tapered to the base, angular

distally, dehiscing between the locules and the wings. Seeds barrel-shaped,

brown, ca 0.3 mm long, collar cells more than half the seed length.

Specimen examined: Borneo. Sarawak, Ulu Merirai, Pak Danum, Julia et

al. S 95783 (SAR, SING).

Distribution: Endemic in Borneo, Sarawak; known only from the Ulu

Merirai area, Tatau District.

Habitat: On slopes below the limestone hills on thick black soil or on

litter-covered limestone boulders in very wet shaded areas or sometimes

epiphytic on trees beside streams.

Notes: Superficially, in its non-oblique, obovate leaves it resembles Begonia

stichochaeta K.G.Pearce from the Subis limestone but it differs in its longer

petioles (3.5-8 cm long vs. 0.7 cm), shorter simple cymes 1-2.2 cm long

with 3 male flowers (vs. inflorescences to 3.4 cm long with a cluster of

many male flowers), longer pedicels of the male flower 10-26 mm long (vs.

7 mm long) and fruit stalks 12-25 mm long (not 5-9 mm long) and capsules

broader than long, 12-15 x 13-18 mm (not longer than broad, 29 x 18 mm).

While both have three locular ovaries and capsules, B. rhodochaeta has

one placenta per locule which places it in sect. Reichenheimia, while B.

stichochaeta has two and belongs to sect. Petermannia.

= i - . 2?

Seven New Species of Begonia (Begoniaceae) 369

It is named for its striking red, bristle-like hairs on the leaf

surfaces.

7. Begonia sarangica Kiew & S.Julia, sp. nov.

Section: Petermannia

A Begonia corrugata Kiew & S.Julia petiolis brevioribus 1.2-2.5 cm longis

(nec 4.5-6.5 cm), inflorescentiis brevioribus 4.3-7.5 cm longis (nec 13.5-

23 cm) et pedicellis in fructu 4-8 mm longis (nec 10-13 mm) differt.

—Typus: Borneo, Sarawak, Tatau District, Ulu Sungai Kakus, Bukit Sarang,

Batu Anyi, 5 Oct 2004, Lee et al. SL 39 (holo, SAR; iso, E, KEP, SING).

Fig. 2.

Cane-like begonia. Stems dark brown, woody, erect, 30-100 cm tall, stout

in life and 6-8 mm thick, little branched, nodes swollen, densely bristly with

bristles 1-2 mm long, at first greenish, becoming brownish; without a tuber.

Stipules sparsely bristly, obovate, ca 17 x 7 mm, margin entire, fringed by

long hairs, apex acute, caducous, stipular scar conspicuous. Leaves distant,

held horizontally, internodes 3.5-8.5 cm long; petioles dark brown, densely

dull, dark green with pale green hairs on the lamina, beneath pale green,

corrugate, thin in life, papery when dried, obovate, asymmetric, 11.5-20 x

10-15.5 cm, broad side 6-11 cm wide, base unequally cordate, basal lobes

rounded, the larger 1.3-3 cm long, margin minutely toothed, apex slightly

acute; venation palmate-pinnate, 5-6 pairs with another 3-4 veins in the basal

lobe, veins narrowly parallel, narrowly branching once less than halfway to

the margin and again about two thirds of the way to the margin, impressed

above, beneath prominent and concolorous tinged purple-brown towards

the petiole, densely bristly. Inflorescences axillary, reddish, densely bristly,

erect, shorter than the leaves, racemose, 4.3-7.5 cm long, peduncles 7-55

mm long, each few-flowered branch arising from a pair of foliose bracteoles,

branches 1-4 mm apart with the bracts and bracteoles overlapping the upper

ones, with up to 4 or 5 branches, male flowers many, female flowers one or

two only from the lowest branch, protogynous. Bracts and bracteoles foliose,

paired, pale green, ovate, bracts 15-18 x 8-11 mm, bracteoles 8-12 x 5-6

mm, decreasing in size towards the apex, dorsally hairy and with a distinct

midrib, margin entire, apex acuminate and setose. Male flowers with reddish

hairy pedicels 2-4 mm long, tepals 2, broadly oval to obovate, 5-6 x 3-5

mm, outside deep rosy red, inside pale yellow sometimes rosy red or rosy

red and greenish towards the base, minutely hairy outside, margin entire,

apex rounded; stamens ca 30, cluster sessile, globose or conical, ca 3 x 2-3

370 Gard. Bull. Singapore 60 (2) 2009

Figure 2. Begonia sarangica Kiew & S.Julia. A. Habit; B. Leaf surface upper; C. Leaf

surface lower; D. Inflorescence; E. Bract; F. Male flower bud; G. Male flower; H. Stamen;

I. Female flower; J. Styles and stigmas (top view); K. Styles and stigmas (side view); L.

Transverse section of the ovary; M. Fruit. (Drawing by J. Pao).

Seven New Species of Begonia (Begoniaceae) 371

mm, filaments sessile or up to 0.5 mm long, anthers pale yellow, narrowly

obovate, ca | mm long, apex rounded, dehiscing along lateral slits. Female

flowers with a reddish pedicel 2-4 mm long, ovary pale green, oblong, 13-

18 x 9-11 mm, wings 3, equal, locules 3, placentas 2 per locule; tepals 5,

pale cream to yellow green, oval to oblong, minutely hairy outside, margin

minutely toothed in the upper half, each tooth tipped by a hair, apex rounded

or slightly acute, outermost 12-13 x 7-8 mm, innermost similar but smaller

8-9 x 5-7 mm; styles 3, ca 5 mm long, joined for ca 1 mm at the base and

bifurcating above, styles and stigmas pale greenish yellow, stigmas in a spiral

papillose band. Fruits held horizontally or downwards at 45°, stalks stiff and

thick, 4-8 mm long, capsules (17-)20-23 x (10-)13(-17) mm, minutely hairy

on the locule wall, wings 3, equal, thin, 4-6 mm wide, splitting between the

locules and wings. Seeds barrel-shaped, brown, ca 0.3 mm long, collar cells

ca half to three quarters the seed length.

Other specimens examined: Borneo. Sarawak, Bukit Sarang, Batu Lebik,

Lee et al. SL 105 (E, KEP, SAR, SING); Bukit Sarang, Anderson S 20942

(K, L, SAR).

Habitat: On limestone derived soil high on the limestone hill, in deep shade

at the base of the summit cliff faces or in saddles, widespread on the two

hills but found only as scattered small populations of about 20 plants.

Distribution: Endemic in Sarawak, Tatau District; known only from Ulu

Sungai Kakus, Bukit Sarang (Batu Anyi and Batu Lebik).

Notes: This is one of the cane-like begonias with corrugated leaves and

resembles Begonia corrugata Kiew & S.Julia not only in its habit and

leaves but also in its very small male flowers with tepals that are minutely

hairy outside. However, B. sarangica differs from this species in its shorter

petioles, 1.2-2.5 cm long (not 4.5-6.5 cm long), shorter inflorescences, 4.3-

7.5 cm long (not 13.5-23 cm long), and shorter fruit stalk, 4-8 mm long

(not 10-13 mm long).

Begonia sarangica is named for the locality where it grows. It Is

unusual in having only one male flower of an inflorescence open at a time.

Acknowledgements

The first author thanks the Singapore Botanic Gardens for permission to

take part in the field surveys and for financing participation in the Ulu

—~_———EE——EEE

372 Gard. Bull. Singapore 60 (2) 2009

Merirai Survey; the second author thanks the Forest Research Institute

Malaysia through the Tree Flora of Sabah and Sarawak Project for

permission to participate and for financial support; to the Sarawak Forestry

Corporation for providing financial support for their staff who participated

in these surveys; to Lucy Chong for her strong support; to Grand Perfect

Sdn Bhd for sponsoring the botanical inventory of Bukit Sarang and to

R.B. Stuebing for organising the logistics; to Malcom Demies for arranging

the logistics for the Ulu Merirai Survey; to Malcom Demies, Rantai Jawa,

Serena Lee, Paul K.F. Leong, Tinjan Kuda, Shalih Rebi and Army Kapi

for their excellent support in the field; to Mark Coode for correcting the

botanical Latin and to Joseph Pao for the botanical plates.

References

_ Doorenbos, J., M.S.M. Sosef and J.J.F.E. de Wilde. 1998. The sections

of Begonia including descriptions, keys and species lists. Wageningen

Agricultural University Paper 98-2: 1-266.

Kiew, R. 1998. Niche partitioning in limestone begonias in Sabah, Borneo,

including two new species. Gardens Bulletin Singapore 50: 161-169.

Kiew, R. 2001. The limestone begonias of Sabah — flagship species for

conservation. Gardens Bulletin Singapore 53: 241-286.

Kiew, R. and C. Geri. 2003. Begonias from the Bau limestone, Borneo,

including a new species. Gardens Bulletin Singapore 55: 113-123.

Kiew, R. and S. Julia. 2007. Begonia (Begoniaceae) from limestone hills

in the Kuching Division, Sarawak, Borneo, including nine new species.

Gardens Bulletin Singapore 58: 199-232.

Kiew, R., S. Julia and S. Lee. 2007. The flora of Bukit Sarang: The odd-

man out, pp. 134-137. In: R. Stuebing ef al. (eds.), Proceedings of the

Regional Conference of Biodiversity Conservation in the Tropical Planted

Forests in Southeast Asia. Natural History Publications (Borneo), Kota

Kinabalu, Malaysia.

Pearce, K.G. 2003. Five new Begonia species (Begoniaceae) from the Niah

National Park, Sarawak, Malaysia. Gardens’ Bulletin Singapore 55: 73-

88.

Gardens’ Bulletin Singapore 60 (2): 373-379. 2009 373

A New Species of Habenaria (Orchidaceae)

from Southeast Asia

H. KURZWEIL’, S. CHANTANAORRAPINT?

AND A. BUAKHLAI

' Singapore Botanic Gardens, | Cluny Rd., Singapore 259569

E-mail: hubert_kurzweil@nparks.gov.sg

> Herbarium, Department of Biology, Faculty of Science,

rince of Songkla University, Hat Yai, Songkhla 90112, Thailand

E-mail: sahut.c@psu.ac.th

* Faculty of Natural Resources, Prince of Songkla University,

Hat Yai, Songkhla 90112, Thailand,

E-mail: amonrat.b@psu.ac.th

Abstract

Habenaria anomaliflora H. Kurzweil & S. Chantanaorrapint, sp. nov. from

Southeast Asia (Thailand and Laos) 1s described and illustrated.

Introduction

While revising the genus Habenaria Willd. for the Flora of Thailand the

first author of this paper came across herbarium material of an unusual

terrestrial orchid from Khao Yai National Park in central Thailand with

small Liliaceae-like flowers. A subsequent search revealed that the same

plant had been noted by Kerr (1969) in Laos and was later illustrated by

Seidenfaden (1992: 70). A third collection, also in central Thailand, was

made by the second and third authors of this paper. As the plant does not

match any currently known species it is here described as new.

Habenaria anomaliflora H. Kurzweil & S. Chantanaorrapint, sp. nov.

Habenaria humistrata Rolfe ex Downie, H. siamensis Schltr., H.

porphyricola Schltr. et H. reniformis (D.Don) Hook.f. affinis sed floribus

omnino regularibus et ecalcaris differt. - Typus: Thailand, Nakhon Nayok

Province, Khao Yai National Park, Mo Sing To Reservoir, ca. 700 m alt.,

14°25’50”N, 101°21°50”E, 25 Sep 2005, A. Buakhlai 158 (holo PSU!; iso

BCU!, BKF!). Fig. 1 & Plate 1.

374 Gard. Bull. Singapore 60 (2) 2009

5 mm

0.5 mm

Figure 1. Habenaria anomaliflora H. Kurzweil & S. Chantanaorrapint. A. Habit; B, C.

flower; D. Floral bract; E. Dorsal sepal; F. Lateral sepal; G. Petal; H. Lip; I. Column; J.

Seed (all from A. Buakhlai 158). (Drawn by S.Chantanaorrapint).

A new species of Habenaria (Orchidaceae) 375

Plate 1. Habenaria anomaliflora H. Kurzweil & S. Chantanaorrapint. A. Upper

part of the inflorescence; B. Flower; C. Column. (all from 4. Buakhlai 158). (Photos:

S.Chantanaorrapint). [— Scale bars: A= 5 mm, B—C = | mm]

376 Gard. Bull. Singapore 60 (2) 2009

Terrestrial herbs, 16—25 cm tall, entirely glabrous. Root tubers ovoid,

|—|.2 x 0.6—0.8 cm, brown, with few fibrous roots. Basal sheathing leaves

few, scale-like. Foliage leaves 2, radical, adpressed to the substrate, ovate-

acute, shortly pointed, cordate at the base, the two leaves unequal in size,

the larger one 2.5—4 x 2.4-3.8 cm, the smaller one 2—2.5 x 1.6—2 cm,

length of the basal sheath 5.7—7 mm. Sterile bracts of the flowering stem

3—6, erect in their basal part and spreading above, narrowly lanceolate,

acuminate, 9.4-12 x 3-4 mm, base shortly sheathing. Inflorescences

terminal, lax, (1—)5- to 8-flowered; rachis 4—8.1 cm long and to | mm thick;

floral bracts ovate-lanceolate, acuminate, 6.6—11 < 2.35—4 mm. Flowers

in most plants not opening very widely, subactinomorphic with the six

perianth lobes subequal, spurless, 4-6 mm in diameter; tepals green on

the outside, inside of petals and lip white. Ovaries and pedicels 10—14.5

mm long, not twisted, suberect, borne close to the rachis, with pronounced

ridges. All 6 tepals with entire margins, dorsal surface papillate above,

ovate-lanceolate, widest just above the base, acute or subacute, with 3

or 5 pronounced dark veins developed as slight keels or ridges, with the

edges and tips turned in. Dorsal sepal 5.6—7 < 2.2—5 mm, lateral sepals

5.5—6.5 x 2-3 mm. Petals 4.9-6.5 x 1.8-2.8 mm. Labellum 4.5—6.5 x

2—3.5 mm. Column short, dark green, 1.5—2 mm long and ca. 3 mm wide

across the top. Anther erect, bright green with vertical white stripe in the

middle, anther canals (?) stout and curving upwards, with white globules

(? viscidia) on their tips; pollinia not seen. Rostellum similar to the anther

in shape but smaller, green, 3-lobed with a short central lobe and two paler

stout lateral lobes. Stigmas 2, apparently largely united, ca. 1 mm long,

black-green, clavate-globular, slightly curved forwards. Fruits fusiform,

12-15 x 2.5—-3 mm, with 6 smooth ridges. Seeds fusiform, testa made up

of few longitudinally-elongate cells, ca. 0.73 mm long.

Distribution: Central Thailand and Laos. Apparently also found in Loe

Province in Northeastern Thailand (P. Tripetch, pers. comm.).

Flowering period: August—October.

Ecology: The type specimen was collected on sandy soil in secondary

forest near Mo Sing To Reservoir in Khao Yai National Park at ca 700 m

altitude. According to the detailed collector’s label of Maxwell 00-351 the

plants of this population were found in secondary, fire-damaged, evergreen

scrubland over sandstone bedrock at an altitude of approximately 825 m.

The specimens which were collected by A. D. Kerr (Kerr 2681) were

found ‘in a large population, in forest’ (Seidenfaden, 1992).

A new species of Habenaria (Orchidaceae) By 7/

Etymology: The name of this new species refers to the abnormal flowers.

Other specimens examined: CENTRAL THAILAND: Nakhon Nayok

Province, Khao Yai National Park, 12 Aug 2000, Maxwell 00-351, p.p.

(BKF!, CMU!, L!); excluding the specimen in the spirit collection at

CMU, which is Habenaria humistrata; Khao Yai National Park, Tripetch

A03 (QBG [spirit], only photographs seen). LAOS: Ban Muong Noi,

Vientiane Plains, flowering in Aug, Kerr 268/, (C! [flowers only],

illustrated by Seidenfaden, 1992: 70).

Notes: The new species is similar in its habit to the three Thai endemics

Habenaria humistrata Rolfe ex Downie, H. siamensis Schltr. and H.

porphyricola Schlitr., and the widespread H. reniformis (D.Don) Hook.f.

(Himalaya to China and Sumatra), all of which are slender plants with (1—)

2—4 small, cordate foliage leaves adpressed to the substrate (Seidenfaden,

1977). However, the new species differs from all of these by its

subactinomorphic perianth which lacks a lip spur.

In his treatment of the orchids of Indochina Seidenfaden (1992:

68-69) commented on the Laotian population and suggested that this

plant is a Habenaria-like monstrosity which is supported here. L. Garay

(pers. comm. to G. Seidenfaden) suggested that the Laotian specimen

(Kerr 2681) may be a peloric form of H. porphyricola but no reasons

for this suggestion were given (however, this would be unlikely as H.

porphyricola is currently considered endemic to Thailand). Seidenfaden

(1992) abstained from proposing a name but illustrated the plant. As

four different collections of the plant are now know it is obvious that

the abnormality is constant and apparently genetically fixed. Floral

abnormalities are not unknown in the genus Habenaria. H. dentata (Sw.)

Schltr., a widespread Asian species, has a tendency towards the formation

of abnormal flowers (inner perianth lobes aborted in some flowers, spurs

strongly reduced in some specimens), and the related H. malintana

(Blanco) Merr. is consistently peloric (perianth subactinomorphic and

spurless). Abnormal specimens in the genus Habenaria have also been

reported in a number of tropical African species (Summerhayes, 1968).

Kerr (1969: 187) suggested that this species (though unnamed at the

time) is close to Habenaria malintana, probably mainly on account of the

subactinomorphic perianth.

Due to the similar habit the new species might possibly be an

abnormal form of Habenaria humistrata, H. siamensis, H. porphyricola

or H. reniformis. Particularly the fact that it was once found growing

together in the same population with the former could suggest that it is

378 Gard. Bull. Singapore 60 (2) 2009

merely its peloric form (however, it is equally possible that it was growing

in a mixed population). In an age with advanced techniques available

to us where it is the standard procedure to obtain additional characters

including ultrastructural and molecular ones we consider it premature

to interpret two morphologically different taxa as mere forms of one

another based on mere suspicion and speculation. As (1) this abnormality

is constant, (2) there is currently no definitive evidence indicating that

this abnormality is a form of a particular species, we feel it is justified to

propose it as a new species.

Traditionally the main generic character of Habenaria is the column

with its freely projecting stigmatic processes (Pridgeon ef a/., 2001). This

character is not expressed in H. anomaliflora although this is also not

surprising in a monstrosity. Therefore questions with regard to its generic

position may arise. Currently the only argument supporting the inclusion

of this species in Habenaria is the habit which 1s similar to what is found

—in H. humistrata, H. siamensis, H. porphyricola and H. reniformis, and

it is hoped that future studies will address this problem. The fact that

the column of this new Habenaria species is untypical in its genus is

very much in contrast to H. malintana which has a ‘normal’ column with

elongate and freely projecting stigmatic processes.

Also the homologisation and the organophyletic origin of the

different parts of the strongly reduced column of the new species are

unclear and should be studied in detail in the future. Based on studies of

alcohol material by the second author we provisionally interpret the taller

of the two 3-lobed structures as the anther with has 2 globular viscidia

at the tips of the stout anther canals (although originally developed by

the rostellum these suspected viscidia are distant from it at the flowering

time). We further interpret the smaller of the three-lobed structures as the

rostellum and the two bulges in front of it as the lateral stigma lobes (as

seen by the position of the stigmatic channel). Developmental studies

would be very helpful regarding the exact interpretation of the structures,

but are probably difficult for practical reasons as young ontogenetic stages

will be hard to obtain.

Kerr (1969) points out a transparent coating over the foliage leaves.

This is reminiscent of a similar surface in some southern African Satvrium

and Holothrix species that have similar basal leaves adpressed to the

substrate, where it has been interpreted as a water-storing device (Linder

& Kurzweil, 1999: 27).

St eee =A

4 new species of Habenaria (Orchidaceae) 379

Acknowledgements

We would like to thank the curators of the herbaria BKF, C. CMU and L for

making their collections accessible to us, and Dr. Benito Tan for checking

the Latin diagnosis. The first author would also like to thank Dr. Somran

Suddee (BKF) for drawing his attention to the relevant literature, and the

National Research Council of Thailand for issuing a research permit.

References

Kerr, A.D. 1969. On a collection of orchids from Laos. Natural History

Bulletin of the Siam Society 23: 185-211.

Linder, H.P. and H. Kurzweil. 1999. Orchids of southern Africa. A.A.

Balkema, Rotterdam.

Pridgeon, A.M., P.J. Cribb, M.W. Chase and F.N. Rasmussen. 2001. Genera

Orchidacearum, vol. 2 (Orchidoideae, part 1). Oxford University Press,

New York.

Seidenfaden, G. 1977. Orchid genera in Thailand V. Orchidoideae. Dansk

Botanisk Arkiv 31: 1-149.

Seidenfaden, G. 1992. The orchids of Indochina. Opera Botanica 114:

1-502.

Summerhayes, V.S. 1968. Orchidaceae (part 1), pp. 1-235. In Milne-

Redhead, E. & Polhill, R.M.(eds), Flora of Tropical East Africa. Crown

Agents for Oversea Governments and Administrations, London.

Gardens’ Bulletin Singapore 60 (2): 381-387. 2009 381

Transfer of Amomum fenzlii, a Nicobar Islands endemic,

to Etlingera (Zingiberaceae).

J. LEONG-SKORNICKOVA|, M. SABU?

AND M. G. PRASANTHKUMAR?

'The Herbarium, Singapore Botanic Gardens, Cluny Road 1, 259569 Singapore

E-mail: jana_skornickova@nparks.gov.sg

> Department of Botany, University of Calicut, Calicut University P.O.

673 635, Kerala, India

Abstract

A new combination of Etlingera fenzlii (Kurz) Skorni¢ék. & M. Sabu

(Zingiberaceae) based on the basionym, Amomum fenzlii Kurz, is proposed.

Recent exploration on Great Nicobar Island resulted in the collection of this

endemic species. An amended description with a colour plate is provided

and a lectotype is designated.

Introduction

The genus Et/ingera(Zingiberaceae) was established by Giseke in 1792. Taxa

falling under Et/ingera today include species previously placed in Amomum

Roxburgh, Diracodes Blume and Hornstedtia Retzius. The taxonomic

and nomenclatural history of Et/ingera and related genera was elaborated

by Burtt & Smith (1986). Following the work of Holttum (1974), they

proposed Ev/ingera as the inclusive name for Achasma Griffith, Geanthus

Valeton and Nicolaia Horaninow, which share overlapping characters,

such as an involucre of sterile bracts (sometimes reduced in Geanthus),

elongate tubular bracteoles and a distinct androecial tube formed above the

insertion of the petals by the lower part of the labellum and filament. This

clarification resulted in the transfer of 57 taxa to Etlingera by Smith (1986),

eight taxa by Poulsen (2003) and one taxon by Lim (2000).

Etlingera now comprises over 100 species (Poulsen, 2006) and is

distributed from India and southwestern China into Myanmar, Thailand,

Malaysia, Indonesia, New Guinea and Australia to the Pacific islands with

center of diversity in Malaysia and Indonesia, notably in Borneo.

During our revisionary work on Indian Zingiberaceae the first

author came across several herbarium sheets of plant specimens from the

Nicobar Islands at CAL determined as Amomum fenz/ii and suspected that

382 Gard. Bull. Singapore 60 (2) 2009

they belonged to E¢lingera, but was unwilling to confirm their generic

identity without studying living material. In 2003 the third author collected

this interesting ginger from the lowland evergreen forest of Campbell Bay

in Great Nicobar.

Amomum fenzlii was originally described by Kurz (1876) based

on a collection from Kamorta in the Nicobar Islands, and the name was

subsequently used by Baker (1892). Schumann in his 1904 Monograph on

Zingiberaceae, transferred the species to Hornstedtia and as such, it has

been upheld in Indian regional floras (Jain & Prakash, 1995; Srivastava,

1998; Sinha, 1999). Our study of living specimens confirmed the presence

of sterile involucral bracts, tubular bracteoles, and a distinct androecial

tube just above the insertion of the petals, placing it clearly in the genus

Etlingera.

Kurz in his original 1876 description did not mention any collection

numbers and failed to designate a type. There are two sheets of Amomum

fenzlii collected by S. Kurz from Kamorta (Nicobar Islands) found among

the general plant collections of CAL. According to Index Herbariorum,

Kurz’s collections from the Nicobars are supposed to be duplicated also at

Land W. However, none of these herbaria houses duplicate of the collection

in question. We therefore believe the two sheets at CAL to represent the

only original materials of Kurz’s Amomum fenzlii and propose one of them

as the lectotype (sheet with inflorescence) and the other one as isolectotype

(sterile sheet). Since the original description appears in Latin, we include

below an amended English description of the taxon.

Etlingera fenzlii (Kurz) Skorni¢k. & M. Sabu, comb. nov.

Basionym: Amomum fenzlii Kurz, J. Asiat. Soc. Bengal. 45(3): 154, t.

12. 1876 = Hornstedtia fenzlii (Kurz) K. Schumann in Engler, Pflanzenr.

20:195. 1904. — Typus: India. Nicobar Islands, Kamorta Island, s.d., Kurz

s.n. (lectotype, designated here, CAL 467328; isolectotype, CAL 467329).

Plate 1.

Perennial erect herb growing in clumps, with 5-15 cm between shoots

(at the base of plant). Rhizome subterranean, thick, ca 1.5-2 cm diam.,

covered with reddish brown triangular scales, roots densely white villous.

Plate 1. Etlingera fenzlii (Kurz) Skorniék. & M. Sabu. A. Inflorescence of white-flowered

form (side view); B. Inflorescence of pink flowered form (top view); C. Inflorescence of

pink-flowered form (side view); D. Infructescence; E. Habit; F. Flower dissection and

fruit. (Photos: A-C, M. Sabu; D-E, V.P. Thomas; F, J. Leong-Skorniékova).

Transfer of Amomum fenzlii, a Nicobar Islands endemic

384 Gard. Bull. Singapore 60 (2) 2009

Leafy shoots 2-5 (6) m, leafless for 50-90 cm from the base with ca 18-22

leaves per frond complement, base of leafy shoot swollen, ca 5-6 cm diam.,

green. Sheaths mucronate, mucro 2-10 mm, obscurely striate, glabrous

with pubescent margin at apex, green; ligule 1.5-2 <1.8-2 cm, ovate,

slightly unequally twisted, green to brownish green (when older), glabrous

with villose margin, with 1.3 mm white hairs; petiole ca 15-20 mm, green,

glabrous; lamina oblong to ovate, (16-) 40-75 = (7.5-) 13-18 cm, glabrous,

dark green above, lighter below, acuminate, ca 1.5 mm, base obtuse or

rounded, slightly oblique, margin wavy, hyaline, slightly incurved, pinkish

(on young leaves) to light brownish green (when old), 0.2-0.3 mm broad,

densely cilliate, hairs light golden, 1.5-2 mm. Inflorescence radical; peduncle

to 15 cm, creeping or erect, ca 1.5 cm diam., covered with reddish brown

scales; sterile bracts 5-6 < 2-2.5 cm, pink to white; inflorescence head 6-10

x 6-8 cm, many-flowered (ca 80-150), 5 to 6 flowers open at a time, fertile

and sterile bracts compactly forming a rosette or strobilate structure; lower

_ Sterile bracts 22 to 25 as involucre, broadly ovate, outer ones ca 4-6 = 2-4

cm (inner ones gradually smaller), colour vary from pink to white, margin

hyaline, lanate, outer surface pubescent, apices slightly darker in colour;

fertile bracts, ca 3-4 = 1-1.5 cm, numerous, compactly arranged, pink to

white, size acropetally decreases, margin lanate or villous, outer surface

villous near to margin and apex, but almost glabrous in the center, subtending

a single flower; bracteole ca 20 mm, 5-6 mm diam., tubular, slightly curved,

broader toward apex, purple-red, pink or white, densely pubescent, with

white hairs ca I-1.5 mm, bracteole apex bilobed, rarely three lobed with

unilateral split 6-10 mm, villous at margins with white hairs, hair ca | mm.

Flower 3.5-4 cm; calyx ca 18-21 mm, slightly curved, apex deeply bilobed

with unilateral split ca 1 cm deep, purple or white, densely pubescent, with

ca | mm white hairs, villous at margins; corolla tube ca 11-13 mm, light

purple or white, sparsely pubescent outside, glabrous at internal base, but

densely pubescent in distal portion, hairs ca 1.2 mm, white; dorsal corolla

lobe, ca 13-15 x 6 mm, light purple to the base, dark purple to the tip, or

pink or white, slightly obovate, apex hooded, sparsely pubescent on dorsal

surface; lateral lobes ca 15-17 < 3-4 mm, narrowly oblong-spatulate, light

purple basally, darker purple apically, or light pink or white, dorsal surface

sparsely pubescent, apex slightly hooded, inserted slightly below (ca 0.5

mm) or with the dorsal lobe; labellum dark purple with white or yellowish

margins or white, 14-16 x 13-15 mm, rhomboid, obscurely three-lobed,

basal lobes folded toward anther, distance between insertion of lobes and

base of filament ca 6 mm; filament 1-2 x 2 mm, white, densely pubescent,

hair 1mm, white, externally rather appressed, internally less so; anther 6 x 4

mm at apex (2mm at base), apex emarginate, anther theca 5-6 mm x | mm,

Transfer of Amomum fenzlii, a Nicobar Islands endemic 385

white, dehiscing in upper half, densely pubescent toward base; ovary ca 5

x 3.5 mm, trilocular, ovules many, with axile placentation; style ca 25 mm,

filiform, white, densely pubescent, more so apically, white hairs 0.5-1 mm;

stigma triangular, capitate, deep purple or cream colour, glabrous, ciliate at

slit; epigynous glands two, ca 3 mm long, yellow, cylindrical, surrounding

style. Infrutescence capitate, + globose to 13 cm diam., deep scarlet or

greenish when mature; capsule irregularly obovoid, 2.8-4 = 1.8-2.2 cm (at

top), densely pubescent, apex flattened to depressed with persistent calyx

remnants. Seeds 5 x 3 mm, rounded triangular, brown-black, completely

embedded in a white, fleshy, sweet aril.

Additional specimens examined: INDIA: Andaman & Nicobar Islands.

South Nicobars, Great Nicobar Isl., 26" km on East-West Road, Hore

6771 (CAL, PBL); Great Nicobar Isl., Campbell Bay near jetty, Hamza

6712 (CAL, PBL): Great Nicobar 6" km on E-W Road, Balakrishnan 2960

(CAL, PBL); 33% km on E-W Road, Balakrishnan 3942 (CAL, PBL);

Great Nicobar 24" km on E-W Road, Sinha 1637] (PBL); Campbell Bay

35" km North-South Road, Prasanthkumar 92604 (CALI); 30" km on N-S

Road, Prasanthkumar 92605 (CALI); 9" km on E-W road, Prasanthkumar

92610 (CALI). North Nicobar, Arong, Car Nicobars, Nair 2631 (CAL,

PBL); Katchal Island, Nirman Nagar, Vasudeva Rao 7421 & 7422 (PBL).

Nicobars, Man s.n. (CAL); Nicobars?, sine coll. 89 (BM).

Vernacular name: Hami (in Nicobari tribal language).

Phenology: April — August.

Ecology: This species grows profusely in the evergreen forests of Nicobar

Islands, especially in open places near the streams and along roadsides.

Distribution: Endemic to the Nicobar Islands.

Uses: Juice of the rhizome is used in the treatment of cough, fever,

respiratory disorder and skin diseases by the Nicobari tribes (Srivastava,

1998), who also eat the sweet and fleshy white aril.

Notes: Variability among populations of this species growing in Great

Nicobar Island has been observed. Locals consider two different varieties,

the more common plants with rose-pink fertile and sterile bracts and scarlet

red capsules and the plants with white fertile and sterile bracts and greenish

capsules. We have observed that the form with white bracts further shows

386 Gard. Bull. Singapore 60 (2) 2009

variability in flower colour, which can be white or pink and that the

intensity of colour in plants with pink bracts varies considerably. We are

of the opinion that these characters are not enough to treat any of these as

distinct varieties.

In India Etlingera is represented by two species, E. loroglossa

(Gagnepain) R.M. Smith and E. /inguiformis (Roxburgh) R.M. Smith

(Prakash & Tripathi, 1998), both distributed in Northeastern part of the

country. Etlingera fenzlii can be readily distinguished from the above two

taxa by short labellum and well developed, showy bracts of involucre

(Nicolaia type), while the former two have long prominent labellum

(Achasma type).

Acknowledgements

The research grant provided by the Department of Science and Technology,

-(D.S.T) (SP/SO/A-20/99) Govt. of India, for the Revision of Indian

Zingiberaceae is gratefully acknowledged. We are thankful to the officers of

the Forest Department, Andaman & Nicobar Islands for granting permission

and providing necessary help for the field studies in the islands. Thanks are

due to curators of CAL, L, PBL and W for letting us to examine herbaria in

their care. We thank Dr. A. D. Poulsen (E) for the discussion on the genus

Etlingera and his comments on the manuscript and the two reviewers for

their comments.

References

Baker, J.G. 1892. Amomum, pp. 233-243. In: Hooker, J.D. Flora of

British India. Vol. V1. L. Reeve & Co. London.

Burtt, B.L. and R.M. Smith. 1986. Etlingera: The inclusive name for

Achasma, Geanthus and Nicolaia (Zingiberaceae). Notes from Royal

Botanic Garden, Edinburgh 43: 235-241.

Giseke, PD. 1792. Praelectiones in Ordines Naturales Plantarum,

Hamburg. Pp. 209, 229, 251.

Sinha, B.K. 1999. Zingiberaceae, pp. 447-449. In: Flora of Great Nicobar

Island. BSI, Calcutta.

Transfer of Amomum fenzlii, a Nicobar Islands endemic 387

Holttum, R.E. 1974. A Commentary on comparative morphology in

Zingiberaceae. Gardens Bulletin Singapore 27: 155-165.

Jain, S.K. and V. Prakash. 1995. Zingiberaceae in India: Phytogeography

and Endemism. Rheedea. 5(2): 154-169.

Kurz, S. 1876. A Sketch of the vegetation of the Nicobar Islands. Scitamineae.

Journal of Asiatic Society of Bengal, Part 2. Natural History 45: 154-

opr

Lim, C.K. 2000. Taxonomic Notes on Etlingera Giseke (Zingiberaceae) in

Peninsular Malaysia: the “Nicolaia” taxa. Folia Malaysiana 1(1): 1-12.

Poulsen, A.D. 2003. One new name and new combinations of Malesian

Zingiberaceae. Blumea 48: 523-527.

Poulsen, A.D. 2006. Etlingera of Borneo. Natural History Publications

(Borneo), Kota Kinabalu in association with Royal Botanic Garden

Edinburgh.

Prakash, V., and S. Tripathi. 1998. Studies on Zingiberaceae of N.E India:

II. Notes on Etlingera Giseke. Rheedea. 8(2): 173-178.

Schumann, K. 1904. Zingibeaceae, pp. 195-196. In: A. Engler (ed.), Das

Pflanzerich, Heft 20. Berlin.

Smith, R.M. 1986. New Combinations in Et/ingera Giseke (Zingiberaceae).

Notes from Royal Botanic Garden, Edinburgh 43: 243-254.

Srivastava, S.K. 1998. Zingiberaceae in Andaman and Nicobar Islands,

India. Higher Plant Indian Subcontinent 8: |-33.

Gardens’ Bulletin Singapore 62(2): 389-397. 2009 389

The Curcuma Cultivar Registry I.

J. LEONG-SKORNICKOVA AND A.K. NURA

Singapore Botanic Gardens, | Cluny Road, 259269 Singapore

E-mail: jana_skornickova(@nparks.gov.sg

nura_abdul_karim@nparks.gov.sg

Introduction

This is the inaugural list of Curcuma cultivar names accepted by the

Singapore Botanic Gardens, which acts as the International Cultivar

Registration Authority for the genus Curcuma.

The list provides the name of the registered cultivar, SBG cultivar

registration number under which the standard portfolio is accessioned and

lodged at the Gardens’ Herbarium (SING), the hybridiser, the parentage as

provided by the hybridisers, and basic diagnostic features of the cultivar

extracted from the standard portfolio, which consists of registration form,

herbarium specimen number and a photograph, unless otherwise stated

under additional information. The colour images of the inflorescence and

flowers are provided. Colours indicated in the descriptions follow RHS

Colour Chart (1995).

List of Cultivars

‘Blue Andaman’ — SBG R#001 Annop Ongsakul, Sulee Nursery

cv. of probably C. petiolata

DESCRIPTION: Rhizome pale yellow inwards. Leaf ca 47 x 20 cm, green;

midrib green. Inflorescence peduncle ca 18 cm, spike ca 22 x 7 cm; basal

bracts ca 25-42, recurved, uniform yellow-green (145B); coma bracts

10-12, wider than basal bracts, flat, uniform violet (85B). F/ower uniform

orange-yellow (21A). Additional information: Flowering period June —

August. Fig. 1A

‘Blue Moon’ — SBG R#002 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 20 x 5 cm,

green; midrib green. Inflorescence peduncle ca 25 cm, spike ca 15 x 5

cm; basal bracts ca 8-9, rounded, uniform yellow-green (145A); coma

bracts ca 5-7, slightly acute, erect, uniform violet (84D) with green tip.

390 Gard. Bull. Singapore 60 (2) 2009

Flower white with violet labellum (83A). Fig. 1B

‘Blue Sky’ — SBG R#003 Golden Lotus Nursery

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 23 < 4.5 cm,

green; midrib red. Inflorescence peduncle ca 28 cm, spike ca 10 x 4 cm;

basal bracts ca 6, rounded,uniform yellow-green (144B); coma bracts ca

8-9, pointed, erect, violet (85C) with yellow-green tip (144A), base paler

yellow-green (144D). Flower white with violet labellum (86A). Fig. 1C

‘Blushing Crown’ — SBG R#004 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 30 x 6 cm,

green; midrib green. Inflorescence peduncle ca 49 cm, spike ca 16 x

5.5 cm; basal bracts ca 9, rounded,uniform yellow-green (146D); coma

_bracts 8-11, acute, erect, almost white, turning pale purple (76C), tip with

large brown patch above and yellow-green below (148C) with marging of

tip red-purple (61C). Flower violet (81A). Additional information: Opens

white, turns a nice pink with age and bracts become more lax. Fig. 1D

‘Blushing Star’ — SBG R#005 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 21 x 6 cm,

green; midrib green. Inflorescence peduncle ca 35 cm, spike ca 16 x 6 cm;

basal bracts c.11-12, rounded, uniform green (143C); coma bracts ca 11-

13, erect, white when young, mature light red-purple (65B) with darker

red-purple patch (65A) and yellow-green tip (144B). Flower with violet

labellum (83A). Additional information: White changes to pale pink with

age. Fig. 1E

‘Cambodian Scarlet’— SBG R#006 Golden Lotus Nursery

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 25 x 4.5 cm,

green; midrib green. Inflorescence peduncle ca 13.5 cm, spike ca 15 x 5

Figure 1. Curcuma cultivars. For each, a side view of inflorescence, coma (top view) and

flower (front view), is presented: A. ‘Blue Andaman’; B. ‘Blue Moon’; C. ‘Blue Sky’;

D. ‘Blushing Crown’; E. ‘Blushing Star’; F.“Cambodian Scarlet’. (Photos: Koh Sin Lan,

SBG).

392 Gard. Bull. Singapore 60 (2) 2009

cm; basal bracts ca 10-11, rounded, uniform yellow-green (144A); coma

bracts ca 7, pointed, erect, purple (78B) with darker greyed-purple tip

(183B). Flower white with violet labellum (83C). Fig. 1F

‘Cherry Princess’ — SBG R#007 Chiangmai Setcon Co. Ltd

cv. of unknown parentage

DESCRIPTION: Rhizome information not recorded. Leaf ca 34 x 8.7 cm,

green; midrib green. Inflorescence peduncle ca 35 cm, spike ca 10 x 4.5

cm; basal bracts ca 12, rounded, light yellow-green (144D) with darker

yellow-green rim (144A); coma bracts ca 10-11, rounded, erect, purple

at base (75A), paler at upper part (75B) with yellow-green tip (144A).

Flower white with deep purple labellum (77A) fading to lighter purple

(77B). Fig. 2A

‘Emerald Pagoda’ — SBG R#008 Chiangmai Setcon Co. Ltd

_ev. of C. harmandii

DESCRIPTION: Rhizome information not recorded. Leaf ca 33 x 8 cm,

green; midrib green. Inflorescence peduncle ca 42 cm, spike ca 16 x 6.5

cm; basal bracts ca 15, uniform green (143B); coma bracts, flatter and

slightly longer than the basal bracts, uniform green (143C). Flower violet |

(83A). Fig. 2B

‘Lady Ann’ — SBG R#009 Golden Lotus

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 26 x 5.5 cm,

green; midrib green. Inflorescence peduncle ca 26 cm, spike ca 16.5 x 5

cm; basal bracts ca 10-11, rounded, uniform yellow-green (144B); coma

bracts ca 11-14, pointed, narrow, erect, white to pale pink with red-purple

tinge (73B) and light yellow-green tip (144C). Flower white with deep

violet labellum (83A). Fig. 2C

‘Paloma Blanca’ — SBG R#010 Golden Lotus, Nursery

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 25 = 6.5 cm,

green; midrib green. Inflorescence peduncle ca 22 cm, spike ca 14.5 x 6

Figure 2. Curcuma cultivars. For each, a side view of inflorescence, top view of coma

and front view of flower, is presented: A. ‘Cherry Princess’; B. ‘Emerald Pagoda’; C.

‘Lady Ann’; D. ‘Paloma Blanca’; E. ‘Phudin Pink’; F.“Phudin Red’. (Photos: Koh Sin

Lan, SBG).

The Curcuma Cultivar Registry I.

394 Gard. Bull. Singapore 60 (2) 2009

cm; basal bracts ca 9-10, rounded, uniform green (143C); coma bracts ca

8-9, pointed, erect, very pale purple to almost white (76D), lower coma

bracts with yellow-green tip (144C) on outer side, upper coma bract with

red tips (47B) on outer side. Flower large, white with violet labellum

(86B). Fig. 2D

‘Phudin Pink’ — SBG R#011 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia x C. rhabdota

DESCRIPTION: Rhizome information not recorded. Leaf c. 32 x 10 cm,

green; midrib green. Inflorescence peduncle c. 40 cm, spike c. 14 x 5.5

cm; basal bracts c.9, rounded, green (143B) with greyed-purple pattern

(185B) and two greyed-purple patches (186D); coma bracts c.9-11,

flatter than basal bracts, erect, light red-purple (69B) with yellow-green

tip (146C) with greyed-purple stripe on outer side (185A). Flower red-

purple (70A). Fig. 2E

‘Phudin Red’— SBG R#012 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia x C. rhabdota

DESCRIPTION: Rhizome information not recorded. Leaf ca 39 x 12 cm,

green; midrib red. Inflorescence peduncle ca 51 cm, spike ca 11 x 4.5

cm; basal bracts ca 8-12, rounded, green (143B) with two purple brown

patches (184B); coma bracts ca 7-10, erect, red-purple (74C), veins white

with yellow-green tip (148B). Flower deep violet (86A) fading to lighter

violet (86C) with red stripes and yellow patch at middle of throat. Fig.

*Pitcha Pink’ — SBG R#013 Supranee Kongpitchayanond,

Pitcha Nursery

cv. of C. aurantiaca

DESCRIPTION: Rhizome information not recorded. Leaf ca 64 x 17 cm,

green; midrib green. Inflorescence peduncle ca 30-42 cm, spike ca 19-22

< 6.5-8 cm; basal and coma bracts similar in shape ca 60, bracts at base

of the inflorescence light green with light red-purple edges (74C), middle

bracts light green at lower half and light red-purple (74C) at upper part

gradually changing into entirely light red-purple (74C) bracts at the top

Figure 3. Curcuma cultivars. For each, a side view of inflorescence, top view of coma and

front view of flower, is presented: A. ‘Pitcha Pink’; B. ‘Snowball’; C. ‘Sulee Sunrise’; D.

‘Sulee Sunshine’; E. ‘Sunrise’; F..Wipada White’. (Photos: Koh Sin Lan, SBG).

The Curcuma Cultivar Registry I

396 Gard. Bull. Singapore 60 (2) 2009

of inflorescence. The pink colour fades with age to 74D. Flower light

yellow-orange (14C) with darker labellum (14A). Additional information:

Seasonal June — October. Fig. 3A

‘Snowball’ — SBG R#014 Supranee Kongpitchayanond,

Pitcha Nursery

cv. of C. parviflora

DESCRIPTION: Rhizome information not recorded. Leaf ca 27.5 x 9 cm,

green; midrib green. Inflorescence peduncle ca 19 cm, spike ca 5.5-7 5-7

cm; basal and coma bracts similar, white with green tip (137C). Flowers

present only as sterile white remnants. Additional information: This

cultivar is formed by a ball of multiple sterile inflorescences. Seasonal

June — October. Fig. 3B

‘Sulee Sunrise’ — SBG R#015 Annop Ongsakul, Sulee Nursery

_ ev. of C. cordata = C. auratiaca

DESCRIPTION: Rhizome pale yellow inwards. Leaf ca 65 x 27 cm,

green; midrib green. Inflorescence peduncle ca 35-45 cm, spike ca 22-30

x 7-9 cm; basal bracts ca 25-55, recurved, greyed-orange (171C) with

green veins; coma bracts ca 9-12, larger and flatter than basal bracts,

light red-purple (69A) at base with green veins. F/ower uniform orange-

yellow (23A). Additional information: Flowering period June — August.

Fig. 3C

‘Sulee Sunshine’ — SBG R#016 Annop Ongsakul, Sulee Nursery

cv. of unknown parentage

DESCRIPTION: Rhizome pale yellow. Leafca 55 x 19 cm, green; midrib

green. Inflorescence peduncle c. 28-36cm, spike ca 25-27 < 5.5-6.5 cm;

basal bracts ca 21-35, recurved, base pale green with upper half coloured

(42B); coma bracts ca 7-9, broader than basal bracts, uniform light purple

(78C). Flower uniform orange-yellow (21C) with median band (21A).

Additional information: Flowering period June — August. Fig. 3D

‘Sunrise’ — SBG R#017 Golden Lotus Nursery

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 214 x 5 cm,

green; midrib green. Inflorescence peduncle ca 29 cm, spike ca 16 x 5.5

cm; basal bracts ca 8-10, rounded, uniform green (143C); coma bracts ca

10-12, pointed, flatter than basal bracts, erect, white tinged with pink, tip

reddish (60A). Flower white with deep violet labellum (83A) with white

longitudinal line. Fig. 3E

The Curcuma Cultivar Registry I. 397

*‘Wipada White’ — SBG R#018 Chiangmai Setcon Co. Ltd

cv. of C. alismatifolia

DESCRIPTION: Rhizome information not recorded. Leaf ca 40 = 8 cm,

green; midrib red. Inflorescence peduncle ca 50 cm, spike ca 17.5 * 6.5 cm;

basal bracts ca 9-13, rounded, uniform pale yellow-green (144B); coma

bracts ca 10, erect, very pale purple (76D) with pale green tip (143C).

Flower white with deep violet labellum (83A). Fig. 3F

Gardens’ Bulletin Singapore 60 (2): 399-410. 2009 399

Conspectus of the genus Eugenia (Myrtaceae)

in the Philippines

P.G. WILSON

National Herbarium of New South Wales, Royal Botanic Gardens, Sydney

NSW 2000, Australia

E-mail: peter.wilson@rbgsyd.nsw.gov.au

Abstract

This paper gives an overview of the genus Eugenia, with an emphasis

on species found in the Philippines. In particular, the generic segregate

Jossinia, adopted by Merrill for those species, is re-evaluated in the light

of more recent research. This research does not support inclusion of

species from the Asia-Pacific region in a segregate genus. Consequently,

the Philippines species are here assigned to Eugenia and two new names,

Eugenia uminganensis and E. inaequisepala, are proposed.

Introduction

The genus Eugenia was established by Linnaeus in 1735 with five species.

Of these five species, only Eugenia uniflora remained in the genus at the

time of publication of volume three of de Candolle’s Prodromus in 1828. De

Candolle himself transferred two of the species to the genus Jambosa (now

considered a synonym of Syzygium). The other two species had already

been reassigned to Barringtonia (Lecythidaceae). The number of species

then considered to belong in Eugenia had grown rapidly, so that de Candolle

included 194 species in it. McVaugh (1968) estimated that there were at

least 500 species in tropical America alone and van der Merwe ef al. (2005)

suggest there are approximately 1000 species worldwide, predominantly

in tropical America but including around 60 in Africa. In the Asia-Pacific

region, the highest number of species occurs in New Caledonia where over

60 species occur (J.W. Dawson, pers. comm., 2006), many of which are

undescribed.

Although Syzygium and other segregates were recognised by a

number of early workers, there was widespread adoption of a broad generic

concept of Eugenia that included the segregates. The foremost advocate of

this was Bentham (1862, 1869) who emphasised the nature of the embryo,

EE OE eT

400 Gard. Bull. Singapore 60 (2) 2009

with its thick, fleshy cotyledons, as a unifying feature of the group. As a

result, his view largely held sway, particularly in English-speaking parts

of the world, until well into the twentieth century. The resurgence of the

recognition of Syzygium as a genus separate from Eugenia was led by

Diels in the 1920s and was continued by Merrill and Perry in their various

publications in the 1930s. Merrill (1951) traced the history of this and

discussed some of the proposed segregates from Syzygium (ie, Jambosa

and Carvophyllus), which he did not recognise as distinct.

Schmid (1972) published a summary of his research on floral

anatomy in this group that showed that Syzygium could be distinguished

from Eugenia by a fundamental difference in vascular supply to the

placentas: in Ewgenia the vascular supply is trans-septal, while in Syzygium

it is axial. Wilson et al. (2005) have presented chloroplast sequence data

that has confirmed this separation and, furthermore, demonstrated that the

traditional concept of subfamily Myrtoideae is not monophyletic. They

segregated Syzygium into a tribe, Syzygieae, separate from the main group

of fleshy-fruited taxa, the Myrteae. More recently, Biffin et al. (2007) have

reached similar conclusions from their analysis of nuclear ITS data. Eugenia

is correctly placed in the Myrteae but the long-established division of this

group into three subtribes does not receive any support in phylogenetic

studies (Lucas et al. 2005, 2007).

Diels (1922: 376), in recognising the genus Jossinia Commers. ex

DC., states that he is following Blume who, he says, separated the genus on the

basis of the inflorescence type, the relatively large sepals, the broad staminal

disk and the aestivation of the stamens in the bud. Merrill (1951) followed

Diels in referring all Old World species of the Eugenia group to Jossinia and

transferred a number of native Philippine species to that genus.

The major issues surrounding Jossinia have been typification

of the genus and the application of some names. De Candolle recognised

eight species that he stated were from Mauritius and the Mascarenes. One

of the species, Jossinia cotinifolia (syn. Eugenia cotinifolia), was for a long

time a name of uncertain application. Some workers, for example Hyland

(1983), considered it quite likely that this name might apply to the species

in Australia, for which he used the later name Eugenia reinwardtiana (syn.

Jossinia reinwardtiana). It now seems certain that Eugenia cotinifolia is a

New World species (Scott 1979) so the name is not applicable to any Old

World taxon. Scott (1979) chose Jossinia tinifolia, a Mascarene species,

as lectotype of the genus. The recently published phylogeny of the Eugenia

group (van der Merwe et al., 2005) does support the monophyly of a group

of Mascarene species that includes not only the type of Jossinia but also a

member of the segregate genus Monimiastrum. However, the Australasian

Conspectus of the genus Eugenia (Myrtaceae) 40]

species Eugenia reinwardtiana is not a member of this clade and it seems

unlikely that any other Malesian or Pacific species that has been placed

in the genus Jossinia by earlier workers would be a member of this clade

either. While the phylogeny is inconclusive with regard to possible generic

segregates, it does not give any support for the recognition of Jossinia in

the broader sense adopted by Merrill and Diels. In a recent paper, Snow

(2008) made new combinations in Eugenia for all Monimiastrum species.

In the Philippines, two of the species described or recognised by

Merrill (1951) lack combinations in Eugenia and these are provided here.

The Philippine species are mostly still poorly known and the descriptions

given below are often deficient in some way. Three of the nine native

species are represented only by the type collection and a further three are

known from fewer than five collections. With one exception, the types of the

native species of Eugenia were collected before the war and, in all cases, the

holotypes of these species were lost when the Manila herbarium (PNH) was

destroyed during the battle for Manila in February 1945. Fortunately, almost

all have isotypes that are preserved in other herbaria.

Since the genus is under-collected in the Philippines, little or nothing

is known of the conservation status of the endemic species. It is reported

(Hay, 1998) that rainforest is being destroyed in the Philippines at a rate

second only to Madagascar’s, with current estimates suggesting that less

than three percent of the extensive primary forests that once covered the

archipelago remain intact.

Eugenia L.

Eugenia L., Sp. Pl. 1 (1753) 470. — Type species: Eugenia uniflora L.

(lectotype, vide N. L. Britton, Fl. Bermuda (1918) 261).

Jossinia Commerson ex DC., Prodr. 3 (1828) 237; Blume, Mus. Bot. | (1850)

119; Diels, Bot. Jahrb. 56 (1921) 531 & Bot. Jahrb. 57 (1922) 376; Mermill,

J. Amold Arbor. 31 (1950) 329 & Philipp. J. Sci. 79 (1951) 356. — Type

species: Jossinia tinifolia (Lam.) DC. [lectotype, vide A.J. Scott, Kew Bull.

34 (1979) 474].

Shrubs or small trees; twigs glabrous or pubescent when young. Leaves

opposite, petiolate; lamina gland-dotted, intramarginal vein visible.

Inflorescence axillary; flowers solitary, fascicled or rarely in triads; peduncles

pubescent: bracts pubescent, persistent. Flowers 4-merous, buds usually

turbinate; sepals rounded, sparsely pubescent; petals + orbicular, gland-dotted,

with ciliate margins. Stamens in multiple whorls on a broad staminal disc.

402

Gard. Bull. Singapore 60 (2) 2009

Ovary 2-locular, ovules several to many, radiating from a centrally-located

axile placenta; style about as long as the stamens or slightly longer; stigma

not dilated. Fruit a succulent berry; sepals persistent; seed 1(—2), cotyledons

of uniform texture, partly fused.

Key to Philippine species

Leaves + sessile, lamina rounded to cordate at base ...............:cceseeeee

Wiis sad naina iUauas tna da Rea Ne Oetker ee 2. E. uminganensis

Leaves with petioles > 2 mm long, lamina usually tapering to the

PetiOke 5. .1::si3 SR ee Re ee 2

Pedicels glabrous, usually > 15 mm long; hypanthium ribbed ...........

sib nats dabnusd a Lad SOgCURG eM Ea PORE SSS CASE Oe eee ae |. E. uniflora

Pedicels puberulent, to 12 mm long or flowers subsessile; hypanthium

not ribbed... nt ene ee eee ee 3

Lamina > 8 ciiwide! tis Soe eee 3. E. kamelii

Lamina<.7 Cm wide wie. ee 4

Petiole< 3: manivlomn@i:...cc22 eases ee 5

Petiole > 3 mmilongy..% one ee 8

Lamina > 6 cm lone iss eee 4. E. aherniana

Lamina-< 6:en long :4.0...8ih. ie See ee ee 6

Pedicels very Shottwx..teetentee ee ee 5. E. inaequisepala

Pedicels > 5 muvilomgs cnd0- cet eee i

Leaves micmbranous .c.ccecccrt.eseotee heme eee ee 6. E. loheri

Leaves chartaceous or subcoriaceous ..............:08 7. E. pasacaensis

Leaves > 12 cm long; young stems ferrugineous-villous..............00+

Ee re a a aac snee thc Leos so adbcentosae: 8. E. montalbanica

Leaves mostly < 12 cm long; young stems not villous ................... y)

Staminal filaments + 19 mm long; hypanthium sparingly pubescent

sannadasincd bees cis Ldds Mu dewesdodehwseswOneegte Peed ess te eeeaae mae ta ee 10

Staminal filaments up to 5 mm long; hypanthium densely white-

PUDESCONE .ssi.ci.cuccdsacevaceuaveedee Geena Cee eeree eee Een meee eae eee 1]

Conspectus of the genus Eugenia (Myrtaceae) 403

10 Petiole 2—3 mm long; lamina 2—3 cm wide ....... 5. E. inaequisepala

Petiole 4-10 mm long; lamina mostly 3—6.5 cm wide ...............0:5

1] Staminal filaments 4-5 mm long; intramarginal vein 2—-6(—8) mm

TEL LEIS ga SSA te et 4. E. aherniana

Staminal filaments 34 mm long; intramarginal vein < 2 mm from

SUE RECE eel eee cesar eo 10. E. sargentii

1. Eugenia uniflora L.

E. uniflora L., Species Plantarum 1: 470 (1753). — Type: Micheli, Nov. PI.

Gen. (1729) 226, t. 108 (lecto selected by McVaugh, 7axon 5: 140, 1956).

Shrub or small tree; branchlets glabrous. Leaves opposite; lamina thinly

coriaceous, ovate to broadly ovate, 2.5—5 cm long, 1.25—2.5 cm wide, apex

shortly acuminate, base rounded or broadly cuneate: petiole 2—5 mm long.

Inflorescence axillary; flowers solitary or paired; pedicels slender, (8—)15—

25(-40) mm long, glabrous. Hypanthium 2—3 mm long, ca | mm wide,

sparsely puberulent to glabrescent; sepals + equal, ovate to lanceolate, 3—S

mm long, persistent, puberulent; petals white, obovate to elliptical, 7-8 mm

long, ciliate; stamens numerous, 2.5—7 mm long. Ovary ribbed, 2-locular, style

5—7 mm long. Fruit usually orange-red, conspicuously 8-ribbed, 1—2(—3) cm

diam.; seed one, rarely 2 or 3.

Distribution: Native of Surinam, Guyana and French Guiana, Brazil and

Uruguay; now widely cultivated. Naturalised in Luzon.

2. Eugenia uminganensis Peter G. Wilson, nom. nov.

Garcinia heterophylla Merr., Philipp. J. Sci., Bot. 12 (1917) 285 ; Mermill,

Enum. Philipp. Pl. 3 (1923) 84. — Jossinia heterophylla (Merr.) Merr.,

Philipp. J. Sci. 79 (1951) 359. — Type: Luzon, Province of Nueva Ecija,

Mount Umingan, 13 Aug 1916, Ramos & Edano B.S. 26457 (holo, PNH7*;

iso, K—photo!, US—photo!).

Small tree; branchlets glabrous. Leaves opposite; lamina coriaceous,

lanceolate to ovate, 6—10cm long, 2—5 cm wide, shortly and obtusely acuminate,

base obtuse to cordate; petiole very short. Inflorescence axillary; flowers

apparently solitary; pedicels very short. Hypanthium and flower characters

unknown. Fruit + sessile, globose, 20-30 mm diam.; seed solitary.

404 Gard. Bull. Singapore 60 (2) 2009

Note: a new name is required due to the pre-existing name EF. heterophylla

A. Rich.

Habitat: “in forests, altitude about 300 meters”’.

Distribution: Luzon. ‘Mount Umingan’ is probably part of the Caraballo

mountains, located close to the town of Umingan. The municipality of

Umingan is today part of Pangasinan Province close to its border with the

province of Nueva Ecija.

3. Eugenia kamelii Merr.

Eugenia kamelii Merr., Philipp. J. Sci., Bot. 10 (1915) 219; Merrill, Enum.

Philipp. Pl. 3 (1923) 168. — Jossinia kamelii ( Merr. ) Merr., Philipp. J. Sci.

79: 359 (1951). — Type: Samar, Cauayan Valley, 26 Mar 1914, Ramos B.S.

17539 (holo, PNH*; iso, US—photo!, A-fragment).

Tall tree; branchlets glabrous. Leaves opposite; lamina coriaceous, elliptical,

15—20.cm long, 9-13 cm wide, shortly and obtusely acuminate, tapering rapidly

to the petiole, glands inconspicuous; petiole 14-18 mm long. Inflorescence

axillary; flowers fascicled, sessile or pedicels very short. Hypanthium to 10

mm long and 10 mm wide, slightly pubescent with scattered hairs; sepals

unequal, orbicular-reniform, 4-7 mm long, 5—6 mm wide; petals white,

obovate to elliptical, 10-12 mm long; staminal disk 2.7-3.5 mm wide;

stamens numerous, 8-10 mm long. Ovary 2-locular, fruit unknown.

Note. Merrill (1951) said that this “species is still known only from the

original collection ... there being an excellent specimen of this in the U.S.

National Herbarium”. As far as I can tell, this isotype seems to be the only

extant duplicate.

Local name: Damol (Northern Samar: a Central Visayan language).

Habitat: In the protologue, this is recorded as growing “in damp forests”.

Distribution: Known only from the type collection. The locality “Cauayan

Valley’ does not appear on the label of the type specimen, only in the

protologue. The Cauayan Valley is in the province of Northern Samar near

the town of Catarman.

4. Eugenia aherniana C.B. Rob.

-Eugenia aherniana C.B. Rob., Philipp. J. Sci. 4 (1909) 344; Merrill, Enum.

Conspectus of the genus Eugenia (Myrtaceae) 405

Philipp. Pl. 3 (1923) 157. — Jossinia aherniana (C.B.Rob. ) Merr., Philipp.

J. Sci. 79 (1951) 358. — Type: Mindanao, Province of Lanao del Sur, Lake

Lanao, Camp Keithley, Clemens 619 (holo, PNH7).

-Eugenia melastomoides Elmer, Leafi. Philipp. Bot. 4 (1912) 1429; Merrill,

Enum. Philipp. Pl. 3 (1923) 171. — Types: Mindanao, Davao, Todaya (Mt.

Apo), May 1909, Elmer 10750 (isosyn [flowering], A, GH, K—photo!, NSW!,

US—photo!); ibid, Jun 1909, E/mer 11013 (isosyn [fruiting], A, GH, K— photo!,

MO-photo!, NSW!, US—photo!).

Tree or small tree; branchlets pubescent, soon glabrescent. Leaves opposite;

lamina chartaceous, elliptical to obovate or orbicular, (5—)6.5—12 cm long,

(2.5—-)3—5(-6) cm wide, acuminate (sometimes only shortly and obtusely),

tapering gradually to the petiole, glands inconspicuous; petiole 3—7 mm long.

Inflorescence axillary; flowers solitary or few and fascicled; pedicels very

short, white-pubescent. Hypanthium ca 3 mm long, ca 5 mm wide, white-

pubescent; sepals in unequal pairs, 3.5—5 mm long; petals white, obovate to

elliptical, ca 6 mm long, ciliate; staminal disk 1 mm wide, covered with very

dense white hairs; stamens numerous, 4—5 mm long. Ovary 2-locular, style ca

6 mm long. Fruit subglobose, 15—20 mm diam.; seed solitary.

Note: the type specimen of EF. aherniana has not been located. There is a

specimen in the Munich herbarium, collected by Mary Clemens at Lake

Lanao in July 1907, that may be a type but it has no collecting number (H-J

Esser, pers. comm., 2008). The paratype (Surigao, Ahern 518) has not been

located, either.

Local names: Hangos (Surigao); Baring-oras, Magdan (Davao: Bagobo

language). The fruit is said to be edible.

Habitat: In Elmer’s protologue, he gives “in humid forests at 3500 feet

altitude” as habitat for the flowering syntype and “dry woods at 2500 feet”

for the fruiting syntype.

Distribution: Widespread, Luzon to Mindanao; also in Sulawesi.

5. Eugenia inaequisepala Peter G. Wilson, nom. nov.

Jossinia brachypoda Merr., Philipp. Journ. Sci. 79 (1951) 359, hom. illeg.

— Type: Mindanao, Zamboanga, Santa Maria, Sep-Oct 1912, Reillo B.S.

16403 (holo, GH; iso, BM, L—photo!, US—photo!).

Small tree: branchlets glabrous. Leaves opposite; lamina coriaceous,

406 Gard. Bull. Singapore 60 (2) 2009

elliptical or obovate-elliptical, 4-5.5 cm long, 2-3 cm wide, obtuse or

shortly and obtusely acuminate, tapering gradually to the petiole, glands

inconspicuous; petiole 2-3 mm long. Inflorescence axillary; flowers solitary

or paired; sessile or pedicels very short, somewhat pubescent. Hypanthium ca

3 mm long, ca 6 mm wide, lightly pubescent; sepals unequal, rounded, one

pair 5 x 6 mm, the other 6 x 7 mm, with scattered hairs on the outside; petals

white, size unknown; stamens numerous, ca 10 mm long. Ovary 2-locular,

style ca 10 mm long. Fruit unknown.

Note: a new name is required due to the pre-existing name, E. brachypoda

DC.

Distribution: Mindanao. Province of Zamboanga.

6. Eugenia loheri C.B. Rob.

Eugenia loheri C.B. Rob., Philipp. Journ. Sci. 4 (1909) 345; Merrill, Enum.

Philipp. Pl. 3 (1923) 169. —VJossinia loheri(C.B.Rob. ) Merr., Philipp. Journ.

Sci. 79 (1951) 360. — Type: Luzon, Province of Pampanga, Mount Arayat,

Jun 1896, Loher 2475 (holo, PNH*; iso, M).

Habit unknown; branchlets ? glabrous. Leaves opposite; lamina membranous,

lanceolate or narrow-lanceolate, rarely subelliptical, 2.5—5 cm long, 0.8—2 cm

wide, obtusely acuminate, tapering gradually to the petiole, glands dense;

petiole 1-2 mm long. Inflorescence in or above the axil; flowers solitary or

rarely paired; pedicels 5-10 mm long, villous. Hypanthium ca 2 mm long,

ca 3 mm wide, villous; sepals unequal, oblong, 3—4 mm long, ciliate; petals

white, broadly elliptical to orbicular, 2.5—3 mm long; stamens numerous,

1.5—-3 mm long. Ovary (1—) 2-locular, style ca 3 mm long. Fruit unknown.

Notes: Duplicates of the paratype, Cuming 1388, are held at BM and M, and

a fragment at A. Merrill (1951) considered that this species was known only

from the two original collections but it appears that Loher made a further

collection of this taxon in January 1913 (Rizal Province, Jan 1913, Loher

13920,) that is held at M (H-J Esser, pers. comm., 2008).

Distribution: Luzon: Provinces of Nueva Ecija and Pampanga. Mt Arayat is

an inactive volcano located ca 40 km ENE of Mt Pinatubo. The mountain is

today incorporated in the Mt Arayat National Park. Nueva Ecija is an adjacent

province to Pampanga.

Conspectus of the genus Eugenia (Myrtaceae) 407

7. Eugenia pasacaensis C.B. Rob.

Eugenia pasacaensis C.B. Rob., Philipp. J. Sci. 4 (1909) 346; Merrill, Enum.

Philipp. Pl. 3 (1923) 173. — Jossinia pasacaensis (C.B.Rob.) Merr., Philipp.

J. Sci. 79 (1951) 360. — Type: Luzon, Province of Camarines, Pasacao,

May 1908, Curran F-B. 10467 (holo, PNHT; iso, K—photo!, US-photo!, A—

fragment).

Small tree or shrub; branchlets pubescent, glabrescent. Leaves opposite:

cm wide, obtusely acuminate, tapering gradually to the petiole, glands dense

but inconspicuous; petiole 1—2.5 mm long. Inflorescence axillary (often in

the uppermost axils); flowers solitary or few and fascicled; pedicels 5—12

mm long, puberulent. Hypanthium ca 2 mm long, ca 3 mm wide, puberulent;

sepals unequal, oblong-orbicular, 2.5—3.5 mm long; petals white, broadly

oblong, ca 6 mm long; stamens numerous, 34 mm long. Ovary 2-locular,

style ca 6 mm long. Fruit unknown.

Habitat: Protologue indicates that the type specimen was found “growing on

limestone cliffs along the beach at an elevation of 100 m.”

Distribution: Only known from the type locality. Merrill (1951) knew of this

species only from the type collection but a probable further collection exists.

This specimen, van Wickle FB. 700, May 1904 (NSW!), agrees with the

protologue in having glandular branchlets but appears to represent juvenile

or sucker growth with leaves that are much longer and wider than mature

foliage: the lamina is broadly ovate to broadly elliptical, 5.5—9 cm long and

3.4-6 cm wide, tapering abruptly to the petiole.

8. Eugenia montalbanica Mert.

Eugenia montalbanica Merr., Philipp. J. Sci. 30 (1926) 417. — Jossinia

montalbanica (Merr.) Merr., Philipp. J. Sci. 79 (1951) 360. — Eugenia

diospyrifolia Merr., Philipp. J. Sci., 27 (1925) 39 [non E. diospyrifolia Wall.

ex Duthie]. — Type: Luzon, Rizal Province, Montalban, May 1913, Loher

13307 (holo, PNH*; iso, K—photo!, M).

Small tree; branchlets ferrugineous-villous. Leaves opposite; lamina

subcoriaceous, oblong to oblong-lanceolate, 12—17 cm long, 3—S cm wide,

shortly and obtusely acuminate, tapering gradually to the petiole, villous

below: petiole 5-12 mm long. Inflorescence axillary; flowers solitary or

few-flowered fascicles, sessile. Hypanthium ca 2 mm long, ca 4 mm wide,

pubescent; sepals unequal, ovate, ca 3 mm long, subacute; petals white,

408 Gard. Bull. Singapore 60 (2) 2009

obovate, ca 3 mm long; stamens numerous, short. Ovary 2-locular, style

short. Fruit globose, glabrous, ca 2 cm diam.; seed 1.

Habitat: as suggested by the name of the region, this species is also likely to

occur on limestone substrates.

Distribution: This species is known only from the type and two paratypes

(Loher 13328 & 14879), all from the Montalban region in Rizal Province.

9. Eugenia tulanan Mert.

Eugenia tulanan Merr., Philipp. J. Sci., Bot. 11 (1916) 201; Merrill, Enum.

Philipp. Pl. 3 (1923) 179. — Jossinia tulanan ( Merr. ) Merr., Philipp. J. Sci.

79 (1951) 360. — Type: Samar, Province of Northern Samar, Catubig River at

Pinipisakan, Mar 1916, Ramos B.S. 24453, (holo, PNH7; iso, A, K—photo!,

NSW!).

Tree up to 15 m high; branchlets glabrous. Leaves opposite; lamina +

coriaceous, elliptical to oblong elliptical, S—12.5 cm long, 3—6.5 cm wide,

shortly and obtusely acuminate, triplinerved at the base, tapering to the

petiole, glands inconspicuous; petiole 4-10 mm long. Inflorescence axillary;

flowers solitary or fascicled, pedicels stout, ca 1.5 mm long. Hypanthium

glandular, ca 4 mm long, 6-7 mm wide, sparsely pubescent; sepals glandular,

unequal, orbicular-ovate, 3-7 mm long, 34 mm wide; petals white, obovate

to elliptical, to ca 10 mm long; staminal disk 1.5 mm wide, densely covered

with golden-brown hairs; stamens numerous, to ca 10 mm long. Ovary and

fruit not described.

Local name: Tulanan (a Visayan language).

Habitat: At type locality, the plant is described as growing “in damp forests

at low altitudes”.

Distribution: In the protologue, Merrill cites two localities, Pinipisakan and

Palapag, both on the Catubig River in northern Samar. Pinipisakan, the type

locality, is a waterfall, further upstream from Palapag, the site of the paratype

collection (Ramos B.S. 24421). The species is fairly widespread and has

been recorded from Luzon (Rizal Province and Tayabas (Quezon) Province);

Bohol; and Mindanao (Province of Agusan).

10. Eugenia sargentii Merr.

Eugenia sargentii Merr., Philipp. J. Sci. 18 (1921) 290; Merrill, Enum.

Conspectus of the genus Eugenia (Myrtaceae) 409

Philipp. Pl. 3 (1923) 177. — Jossinia sargentii ( Merr. ) Merr., Philipp. J.

Sci. 79 (1951) 360. — Type: Luzon: Province of Cagayan, [Littoe, vicinity of]

Pefiablanca, 5 Jun 1917, Adduru 169 (holo, PNH*; iso, A, K—photo!).

Tree 5—12 m high; branchlets densely white-tomentose, glabrescent. Leaves

opposite; lamina + coriaceous, oblong-elliptical to ovate-elliptical, 4.5—9 cm

long, 24 cm wide, apex shortly and obtusely acuminate, retuse, base tapering

gradually to the petiole, glands scattered; petiole 5-10 mm long. Inflorescence

axillary (often in the uppermost axils); flowers fascicled, sessile. Hypanthium

ca 2 mm long, ca 4 mm wide, white-tomentose; sepals unequal, reniform to

orbicular-ovate, 2.5—3 mm long; petals white, obovate to elliptical, ca 5 mm

long; stamens numerous, 3—4 mm long. Ovary 2-locular, style ca 4 mm long.

Fruit ovoid, ca 15 mm diam., nearly glabrous; seed solitary.

Local names: Tulisayan and tumaluhu (Cagayan Province: Ibanag language);

Pandaraga (Ticao: probably Masbatefio, a Visayan language).

Distribution: Also recorded from other areas in Luzon (Provinces of Zambales,

Pampanga, Ilocos Norte and Rizal) and in the Bicol Region (Province of

Masbate: Ticao). The island of Ticao has substantial deposits of limestone

that are being mined for cement production.

Acknowledgements

I am indebted to Hans-Joachim Esser (M) for information about specimens

collected by August Loher in the Philippines, and to Jeremy Bruhl

(Australian Botanical Liaison Officer at Kew 2007-8) and John Boggan

and Ingrid Lin (US) for supplying photographs of type specimens.

References

Biffin, E., M.G. Harrington, M.D. Crisp, L.A. Craven and P.A. Gadek.

2007. Structural partitioning, paired-sites models and evolution of the

ITS transcript in Syzygium and Myrtaceae. Molecular Phylogenetics

and Evolution 43: 124-139.

Diels, L. 1922. Die Myrtaceen von Papuasien. Botanisches Jahrbuch fiir

Systematik 57: 356-426.

410 Gard. Bull. Singapore 60 (2) 2009

Hay, I. 1998. E. D. Merrill, From Maine to Manila. Arnoldia 54:11—19.

Hyland, B.P.M. 1983. A revision of Syzygium and allied genera (Myrtaceae)

in Australia. Australian Journal of Botany, Supplement Series 9:

1-164.

Lucas, E.J., S.R. Belsham, E.M. Nic Lughadha, D.A. Orlovich, C.M.

Sakuragui, M.W. Chase and P.G. Wilson. 2005. Phylogenetic patterns

in the fleshy-fruited Myrtaceae — preliminary molecular evidence. Plant

Systematics and Evolution 251: 35-51.

Lucas, E.J., S.A. Harris, F.F. Mazine, S.R. Belsham, E.M. Nic Lughadha, A.

Telford, P.E. Gasson and M.W. Chase. 2007. Suprageneric phylogenetics

of Myrteae, the generically richest tribe in Myrtaceae (Myrtales). Taxon

56: 1105-1128.

Merrill, E.D.‘1950’(1951). Readjustments in the nomenclature of Philippine

Eugenia species. Philippine Journal of Science 79(4): 351-430.

Schmid R. 1972. A resolution of the Eugenia-Syzygium controversy

(Myrtaceae). American Journal of Botany 59: 423-436.

Scott, A.J. 1979. Notes on Myrtaceae in the Mascarene with some

recombinations for taxa from Aldabra, Malaya, New Caledonia. Kew

Bulletin 34: 473-498.

Snow, N. 2008. Studies of Malagasy Eugenia (Myrtaceae) - I: Two

new species from the Masoala Peninsula and generic transfers from

Monimiastrum. Systematic Botany 33: 343-348.

Van der Merwe M.M., A.E. van Wyk and A.M. Botha. 2005. Molecular

phylogenetic analysis of Eugenia L. (Myrtaceae), with emphasis on

southern African taxa. Plant Systematics and Evolution 251: 21—34.

Wilson, P.G., M.M. O’Brien, M.M. Heslewood and C.J. Quinn. 2005.

Relationships within Myrtaceae sensu lato based on a matK phylogeny.

Plant Systematics and Evolution 251: 3-19.

Gardens’ Bulletin Singapore 60 (2): 411-414. 2009 4] ]

Book Review: McCarthy, James. 2008. Monkey Puzzle Man, Archibald

Menzies, plant hunter. Whittles Publishing & Royal Botanic Garden,

Edinburgh, Scotland. 223 pp. (paperbound). ISBN 978—1904445—61-6.

Paice: £25.

This is the first biography of Archibald Menzies, 1754-1842, a Scotsman

who traveled around the world with early British voyages of discovery and

commercial exploitation, botanizing in the new lands visited, and collecting

seeds and plants for introduction to Great Britain. His achievements are

many and his name deserves to be better known and for those reasons this

book is welcome and long overdue. But like the old story about the curate’s

egg, the book is not all that it could be and while it is ‘very good, in parts’

as the curate said to the bishop, there are some bits that are not so savory.

The book reads well — chapters are short and extensively referenced

through endnotes, linked to a bibliography of published and unpublished

sources. I found each chapter engaged my interest and I wanted to know

what would happen next — who knew the biography of a man dead for 165

years could be such a page-turner?

Menzies lived a fascinating life and was the first to record impressions

of peoples and lands that had scarcely been explored before. The extensive

passages quoted from his journal make for fascinating reading and it is

in these parts the book makes for engrossing reading. One problem with

the citation of “Menzies Journal” which is cited extensively in the chapter

endnotes is that, when you check the bibliography, it turns out the there

are several pieces of the journal (or perhaps several different journals

by Menzies), housed in six libraries and archives as distant as London,

Edinburgh, and Australia with no distinction made as to which of these is

being cited in the endnotes.

Menzies’ early life and education are covered briefly and then the

voyages around the world in more depth, but focusing primarily on the

Pacific Northwest coast of North America, first (1786-1789) with Capt.

James Colnett on fur-trading commercial expeditions, and later (1791—

1795) with Capt. George Vancouver of the Royal Navy over three years to

chart the Pacific Northwest coast, search for the fabled Northwest Passage,

and to make such terrestrial investigations as could be accommodated

in the course of what was primarily a naval hydrographic mission. The

last part of the book provides a synopsis of Menzies’ life after these great

adventures: later naval assignments in the Caribbean; declining health and

his retirement from the Navy; marriage and private practice as a medical

doctor in London; the struggle to write up his journals for publication (which

never happened), and an overview of the many plants Menzies introduced

Gard. Bull. Singapore 60 (2) 2009

to horticulture, some of which were to reshape the landscape of the British

Isles. Five appendices conclude the book, each summarizing some aspect

of his life, career, and contributions.

If one were to read quickly through the book just for the big picture

of Menzies’ life and achievements and then put it down, this would make

for an enjoyable story. But as has rightly been said, the devil is in the

details. And it is in the details that this book falls short. Errors of fact and

interpretation are to be found in the parts of the Menzies story best known

to me, and if that is so, then what does it imply about the other chapters,

dealing with regions and historical events less familiar?

For practical reasons, this review concentrates on the Hawaiian

portions of the Menzies story and the problems detected in the chapters,

endnotes, photographs, and bibliography concerning that epoch of his life.

It will be up to other readers, better versed in the histories of the Pacific

Northwest of America, Pacific exploration, and Scotland itself, to decide

whether those portions of the book are the good part of the curate’s egg or

not.

The photos, of which there are many black and white images

scattered throughout the book and eight unnumbered pages of color images

between pp. 98-99, are one area where some serious overhaul is needed.

A full list of criticisms has been sent to the publisher in the hope that the

worst howlers can be corrected in a future printing. A few points worth

noting: the photo on p. 99 of King Kamehameha 1s stated in the caption

to be the one at Kapa‘au, which is a small village located on the Island of

Hawai‘1. Yet the roofline clearly visible behind the statue 1s that of Ali‘iolani

Hale in downtown Honolulu, on the island of O‘ahu. The portrait of Queen

Kaahumanu (p. 104) is said to be “by an unknown artist of the time” but

is surely a copy based on the famous portrait by Louis Choris, artist on

von Kotzebue’s expedition from Russia. And sadly, the plant depicted on

p. 105 as “taro” (more correctly kalo in Hawaiian, scientifically Colocasia

esculenta) is another species entirely, most likely a kind of Xanthosoma.

Among the color images that of the double-hulled canoe has several

problems: firstly it is not attributed, but is almost certainly the work of a

living artist in Hawai‘i who is renowned for his contemporary paintings

of historical events. Did the author or the publishers receive permission

to reproduce this painting in the Menzies book? Secondly, the caption is

so muddled that it is hard to know what event the painting is meant to

represent. The caption reads: “A painting showing Chief [sic] Kamehameha

in a canoe, sailing out to meet Captain Cook on arrival in Hawaii in 1789.”

Well, for starters, Kamehameha was not a chief, in the Hawaiian sense

of that word; he was a noble who usurped power and battled his way to

Book Review: Monkey Puzzle Man, Archibald Menzies 413

becoming supreme ruler, and thereafter “King” of the Hawaiian Islands.

Secondly, Capt. Cook was killed at Kealakekua Bay in February 1779, so it

seems unlikely he was the one being received in 1789, unless there is a typo

in the date. But perhaps what the author meant is that Kamehameha was

sailing out to meet Capt. Vancouver in 1789, although this year does not

agree with the chronology to be found in Appendix 2. It really isn’t possible

to know what is intended here.

Turning to the text, it is a simplification, to put it mildly, to state that

“Kamehameha ...eventually overcoming the chiefs of all the other islands.

He did this through the power of firearms obtained from western ships...”

In fact, Kamehameha, having already bested his rivals on the largest island,

Hawai‘i, managed to battle his way to supremacy on the islands of Maui and

O‘ahu, the latter just barely, due to the rapid decimation of the Hawaiians by

diseases introduced by Europeans. But Kamehameha never managed to win

Kaua‘1 by military might. Instead, the wily chief of that island, Kaumuali‘i,

struck a bargain with Kamehameha and voluntarily ceded his island to him,

in return for which Kaumuali‘i was permitted to continue to govern his

traditional lands. A smaller point is that the Hawaiian personal and place

names are a mish-mash, sometimes rendered as Menzies spelled them (e.g.

Wha-ra-rai), other times given in the contemporary forms (Hualala1), but

usually without a cross-reference. The misspellings of the same name that

appear on one page (such as Kauai and Kaui on p. 107, for example) should

have been caught by a copy editor and corrected. Other spelling errors:

Wimea (for Waimea) Bay; kappa for kapa; Haiwai‘iloa (for Hawai‘iloa);

and Don Jose Mozino (for Mogifio).

Turning to a few botanical faux pas, the statement (p. 91, note 23)

that “...hepatics, which include the lichens.” would have old Menzies sitting

up in his grave and wailing in protest. As someone who had a lifelong

fascination with cryptogamous plants (e.g., the non-flowering plants such

as mosses, liverworts, and ferns that reproduce by spores), Menzies himself

appreciated that lichens are a completely separate group from hepatics,

and while the nature of lichens was not well understood in his time, we

know today that lichens are not plants at all but fungi that have developed

a symbiosis with one or more algae! The shaddock trees mentioned in the

text and depicted in a color photo are correctly known as Citrus maxima,

and not C. grandis, which is a synonym. It is a shame that the text was

not reviewed by one of the Royal Botanic Gardens botanists, who would

certainly have detected these and other botanical lapses and corrected

them.

These few examples suffice to demonstrate that this biography is not

the scholarly and rigorously researched tome that a man such as Archibald

414 Gard. Bull. Singapore 60 (2) 2009

Menzies deserves; instead this is more like the screenplay for a Hollywood

movie about the man that captures the broad brush strokes defining his

life but falls short in accurately delineating its details. Those who are nit-

pickers will no doubt find material not mentioned here to satisfy them,

and those who want a fast-paced read will find a tale that is surprisingly

enjoyable, if one can manage to suspend disbelief for just a little while.

While I can not offer my wholehearted congratulations to the author for the

book he produced, I can sincerely thank him for bringing Menzies’ life and

accomplishments into greater prominence. He has been long overlooked

and it is clear that there is more to be written about this remarkable Scot.

G. Staples’

The Herbarium

Singapore Botanic Gardens

' Prior to joining the Singapore Botanic Gardens in 2007, Dr Staples was the Botanist at the

Bishop Museum’s Herbarium Pacificum in Honolulu from 1988—2006. He has researched

and published extensively on Hawaiian and Pacific plants and has a special interest in the

history of botanical exploration in the region.