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ISSN 0374-7859
THe
GARDENS’ BULLETIN
SINGAPORE
Volume 36
(1983)
A periodical reflecting the interests and
activities of the Botanic Gardens
Singapore
To be purchased at the Botanic Gardens
Cluny Road, Singapore 1025
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Singapore 1025
Printed by Amsterdam Type Printers, Singapore
CONTENTS
Volume 36
PART 1 — Ist June 1983 PAGES
ethan INGI@sacas sade os edo Roeb566 SO BS ANC Tene ae eee ee 1
VINCENT, J. R., AND P. B. TOMLINSON:
Architecture and Phyllotaxis of Anisophyllea disticha (Rhizophoraceae).............. 3-18
WAPISs K-:
Amesiodendron and Litchi (Sapindaceae): new records for the Malay Peninsula ...... 19-24
BRIGE. Mz G.:
SCG! URI TAI EIGN EINg DI EVAG) a ai eon Go oo te sO ome COS CR eee eee erie ieee 25-29
CHIN. -S;C.:
MEM BESLONCHEAIOOLalOn Wialaval(LW) ss trercienciere Siete ere ncasins no Deemed oe tees soe 31-91
STONE, BENJAMIN C.:
Studies in Malesian Rutaceae. III. Melicope suberosa,
a new species and new generic record for the Malayan flora........................ 93-100
STONE, BENJAMIN C-.::
Contributions to the Flora of the Solomon Islands.
PEI esneWICOIMDINALIONS INPATAIIACEAC ne acre tec a cln wienciers = oss wrevajuiels odreteys ne nea ae 101-102
KENG, HSUAN:
AMmNOMdtedulist.on Scedublantsiof sinpapore (VIN) =. ocnccs croc once ces oes cle ances 103-124
VELDKAMP, J. F., AND J. C. VAN DER HAVE:
The Genus Trisetum (Gramineae) in Malesia and Taiwan .......................4.- 125-135
KONING, R. DE, M. S. M. SOSEF AND J. F. VELDKAMP:
A Revision of Heteropholis and Thaumastochloa (Gramineae)...................... 137-162
PART 2 — Ist December 1983
JARZEN, DAVID M.:
he Rossi PollensRecord, of the: Pandanaceae... <2 <2... cece cee oes sie eee eee 163-175
NG, F. S. P., AND M. JACOBS:
A Guide to King’s ‘‘Materials for a Flora of the Malayan Peninsula” ............... 177-185
KOCHUMMEN, K. M.:
Notes on the Systematy of Malayan Phanerogams. XXX. Anacardiaceae............. 187-196
KURATA, SHIGEO:
A new Species of Nepenthes from Sulawesi, Indonesia................---.2.+-+-05: 197-200
WONG, KHOON MENG, AND AH LAN LIM:
On the Nature of Leaf-opposed Inflorescences in Aidia cochinchinensis.............- 201-204
STONE, BENJAMIN C.:
Some New and Critical Pandanus Species. 1. Supplement to Revisio Pandanacearum . . 205-212
LATIFF, A.:
Studies in Malesian Vitaceae. VII. The Genus Tetrastigma in the Malay Peninsula .... 213-228
MAXWELL, J. F.:
New and interesting Plant Records for Singapore, I]..................--.0---eeeee 229-232
GRUEZO, WILLIAM SM.:
Lobaria clemensiae Vain. (Lobariacea, Lichenes) on Halmaheira Island, Indonesia .... 233-236
BHATTACHARJEE, S. K.:
Influence of Growth-regulating Chemicals on Hippeastrum hybridum hort............ 237-242
WTI. ~ co ou cece Ga5 60d. Me Gc ee eee ee Oe ee eee ete 243-251
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1M ARNOLD ARBORETUM
3 SjAN 25 1984 ISSN 0374-7859
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( VOL. 36 (Part 1) Ist June 1983
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A PAGES ¢y
, ES age a Se ee ee 1 &
§2 VINCENT, J.R. AND P. B. TOMLINSON: ()
C) Architecture and Phyllotaxis of Anisophyllea disticha (Rhizophoraceae) ............... 3-18 a\ 4
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\ Amesiodendron and Litchi (Sapindaceae): new records y
aaa AU BES CETSN UNA oraren tod a diana svae iii crete = © vieye: die #3, <jo.3 wisn 2 89 @ brevs ese a elee ae oe 19-24
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PRace, M: G.:
Samar eIN TALI AICSIAI DIDIAZIA bee) c cine ay. © nes vices wc.e ow Swvee eres cee sees sens raens
ENE
CHIN, S. C.: |
MEEees One, Fill Flora Of Malaya (TV) ..05 oon. cic ce ce eee ce ev eieiniv ce meesscceeces 31-91
STONE, BENJAMIN C.:
Studies in Malesian Rutaceae. III. Melicope suberosa,
a mew species and new generic record for the Malayan flora ..............0.eeeeeeeee 93-100
STONE, BENJAMIN C.:
Contributions to the Flora of the Solomon Islands.
RU CERIEC COMIDMIALIOUS UNV ALANACCHG oc sarc c cs tcc sc. s scence eccw eee see cnaacate 101-102
WO QE OE OR
t=
KENG, HSUAN:
mmnotated! Mist of.seed Plants of Singapore (VIM) ........6.. 0s cee cette cee tweets 103-124
VELDKAMP, J. F. AND J. C. VAN DER HAVE:
ihe Genus Trisefurn (Gramineae) in Malesia and Taiwan ..............ececeseeeceees 125-135
KONING, R. DE, M. S. M. SOSEF AND J. F. VELDKAMP:
A Revision of Heteropholis and Thaumastochloa (Gramineae) .............6-2.e scenes 137-162
Published by the Botanic Gardens \)
Parks and Recreation Department
Ministry of National Development
Singapore 1025
<e-
Printed by Amsterdam Type Printers, Singapore
Yon YI OK
ZS OOD Rn Ren Rn PR PE PE EB)
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foe GARDENS, BULLETIN
SINGAPORE
VOL. 36 (Part 1)
CONTENTS
LEyarsorvell IN(ojwae Jee So Sse se ee ee eer eee
VINCENT, J.R. AND P. B. TOMLINSON:
Architecture and Phyllotaxis of Anisophyllea disticha (Rhizophoraceae) ....
YAP, S. K..:
Amesiodendron and Litchi (Sapindaceae): new records
Tayr Tie LY ALESIS THT EIS eege On one ORO OOOO COO ene eS arta
PRICE, M..G.:
SMC AA NUSHAlsMidlesian MIPlaziaiis seat. days etnies. kee ecslerdiels clan. ate aroc
GHIN, S:.C.;
The Limestone Hill Flora of Malaya (IV)
STONE, BENJAMIN C.:
Studies in Malesian Rutaceae. III. Melicope suberosa,
a mew species and new generic record for the Malayan flora .............
STONE, BENJAMIN C.:
Contributions to the Flora of the Solomon Islands.
LeBive new combinations: IN. AraliaCcede™ .yse<, «oc conse Stones) otaveatiwie vies wnelele cine
KENG, HSUAN:
Annotated List of Seed Plants of Singapore (VIII) ..................0205.
VELDKAMP, J. F. AND J. C. VAN DER HAVE:
The Genus Trisetum (Gramineae) in Malesia and Taiwan .................
KONING, R. DE, M. S. M. SOSEF AND J. F. VELDKAMP:
A Revision of Heteropholis and Thaumastochloa (Gramineae) .............
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Singapore 1025
Printed by Amsterdam Type Printers, Singapore
Ist June 1983
ee aboot or 19-24
de Met acts ett 25-29
SES a Soe 31-91
te EOC ae 101-102
Reachoman eehes 103-124
SES CRO 125-135
Soren cea 137-162
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Architecture and Phyllotaxis of
Anisophyllea disticha (Rhizophoraceae)
J. R. VINCENT and P. B. TOMLINSON
Harvard Forest, Harvard University, Petersham, U.S.A.
Conspectus
Page
ABSTRACT 3
INTRODUCTION 3
MATERIAL AND METHODS 5
RESULTS 6
Architecture 6
Leaf morphology 8
Leaf anatomy 10
Nodal anatomy 12
Internode length 13
DISCUSSION 14
Phyllotaxy 14
Adaptive considerations 16
BIBLIOGRAPHY 17
Abstract
-
Anisophyllea disticha does not show distichous phyllotaxis. Erect (orthotropic) shoots with + spiral
phyllotaxis and radial symmetry give rise to tiers of sylleptic branches at regular intervals. The branches
are horizontal (plagiotropic) and have marked dorsiventral symmetry. Their phyllotaxis is unique and
consists of four ranks of alternately arranged leaves, two ranks of scale leaves on the dorsal side and two
ranks of foliage leaves on the ventral side, dorsal and ventral leaves of the two ranks alternating regularly
along the stem on opposite sides. Homology between the three kinds of leafy appendage is indicated by
their constant unilacunar node, but dorsal (adaxial) scales on plagiotropic axes do not subtend axillary
buds. The leaf arrangement is assumed to maximize photosynthesis and corresponds closely to ideal
systems established by theoretical considerations.
Introduction
In the present study attention is drawn to some aspects of morphology and
anatomy of a species of Anisophyllea, A. disticha (Jack)Baillon which allows a bet-
ter comparison with its putative relatives. The genus Anisophyllea R. Br. ex Sabine
includes about 25 species of trees and shrubs distributed primarily in tropical Africa,
Ceylon, India and Southeast Asia (Ding Hou, 1958). Its relatively recent discovery
in South America (Sandwith, 1952) where 3 species are presently known (Pires and
Rodrigues, 1971) makes its known range almost pan-tropical. Species vary from
treelets of the lower forest storey to tall, canopy trees.
The taxonomic position of Anisophyllea is somewhat controversial. Most authors
have included it in the Rhizophoraceae, usually within a separate tribe
Plate 1. Anisophyllea disticha
A. Habit of sapling, obliquely from above.
B. Terminal tier of 5 plagiotropic branches from above, apex of orthotropic axis quiescent.
C. Part of plagiotropic branch tier from above, with second-order branches.
D. Detail of plagiotropic branch from above showing pronounced anisophylly.
A and C from Ektachrome transparencies provided by Dr. J. B. Fisher (Kedah collection); B
and D from Singapore collection.
Architecture & phyllotaxis of Anisophyllea disticha 5
Anisophylleae, together with Combretocarpus, Polygonanthus and Poga. These
four genera differ from other members of the family in having alternate, exstipulate
leaves (rather than opposite, stipulate leaves). The other three genera in the tribe
have more limited distribution than Anisophyllea: Combretocarpus (1 sp.) is
restricted to Sumatra and Borneo, Poga (1 sp.) to West Africa and Polygonanthus
(2 spp.) to Amazonia (Pires and Rodrigues, 1971). This tribal aggregation is strongly
supported by evidence from wood anatomy (van Vliet, 1976). A contrasted view is
to segregate a separate family Anisophylleaceae (Schimp.)Ridl. (e.g. Takhtajan,
1969; Airy Shaw, 1973), but van Vliet points out similarities of the Anisophylleae
to other Rhizophoraceae especially in its resemblance to the tribe Gynotrocheae.
It should be useful to establish the comparative phyllotaxis of members of the
family, since the contrast between the tribes in this character is extreme.
Anisophyllea disticha is significant in this respect because of the pronounced
heterophylly of some of its axes which might represent a transition between the
stipulate and exstipulate conditions, especially as the smaller leaves are often
described as ‘‘stipule-like’’.
Anisophyllea disticha, ‘‘leechwood’’, is restricted to the Malay Peninsula,
Sumatra and Kalimantan (Borneo) where it is a characteristic understorey treelet up
to 7.5 m high occurring on a diversity of soils, ranging from swampy areas to drier
granitic sands and ridges (Ding Hou, 1958). According to Hallé et a/ (1978, p. 200)
the architecture of some Anisophyllea species represents an extreme expression of
Massart’s model, which refers to trees with an orthotropic, monopodial trunk which
grows rhythmically and produces tiers of plagiotropic branches at regular intervals.
Anisophyllea disticha in particular, stands apart from most other members of the
genus in the extreme plagiotropy of the horizontal branches which are
anisophyllous, with an apparent distichous series of scale leaves superimposed on
the distichous foliage leaves (cf. Tay, 1977). This condition is occasionally found in
A. scortechinii so that there is a possible phyletic link between Anisophyllea disticha
and the rest of the genus (Ding Hou, 1958).
The present study establishes to what extent the phyllotaxis of the plagiotropic,
anisophyllous branches is primary (i.e. determined by their method of inception at
the shoot apex) or secondary (i.e. modified by later re-orientation through differen-
tial growth or torsions). Previous description has not addressed this problem (e.g.
Ding Hou, 1958) and has not described microscopic details of anatomy of different
leaf types (cf. Geh and Keng, 1974).
Materials and Methods
Two collections of fluid-preserved (FAA) material which included young or-
thotropic and plagiotropic axes were available for this study (J. B. Fisher, 5.vii.77,
Kedah Peak, Malaysia and P. B. Tomlinson, 5.viii.82, Garden’s Jungle, Singapore
Botanic Gardens).
The distal 1 to 2 cm of a number of both types of axes were embedded in
‘*Paraplast’’ and then serially sectioned at 8 » with a rotary microtome. The sections
6 Gard. Bull. Sing. 36(1) (1983)
were mounted on slides and stained in safranin and fast green. In order to facilitate
investigation of leaf development and nodal anatomy the individual sections were
photographed in series through a Wild microscope with a Bolex movie camera, using
the cinematographic drawing method described by Zimmermann and Tomlinson
(1966).
Single transverse and longitudinal sections of mature leaves from these two collec-
tions were also prepared, using a sliding microtome to cut sections of 20-30 ». These
sections were mounted in glycerine and examined unstained. Finally, leaves from
these collections were cleared in 5% alcoholic NaOH and examined unstained.
Diversity in external leaf size, form, and disposition was explored by examining
the collections of A. disticha in the Harvard University Herbaria, which consisted
mainly of plagiotropic axes.
Results
Architecture. Massart’s model is well represented by this species in the orthotropic
(trunk) axis which produces a tier of horizontal plagiotropic axes (branches) at wide
regular, intervals (Plate 1A). There are usually 5 branches in each tier (Plate 1B).
Our material did not include seedlings but the older trunk axis which we studied
seems similar to the seedling axis described by Geh and Keng (1974). Germination
is described as hypogeal and the plumule bears spirally-arranged scale leaves.
Growth of the axis is rhythmic, eventually with the production of a tier of branches
at the end of each of flush. We have had no opportunity to make extensive
phenological studies but the apex of the orthotropic shoot immediately above the
tier undergoes an extended period of rest before growth is renewed and the next ver-
tical increment is made. Individuals within a population seem asynchronous with
regard to flushing since orthotropic shoots at various stages of development can be
found at any one time. Field study of marked individuals is needed, however, to
monitor events precisely.
Leaves on orthotropic shoots are spirally-arranged with = phyllotactic arrange-
ment (Fig. 1A, 2A). At maturity the leaves are separated by extended internodes at
the base of the shoot, but they are crowded towards the region of tier insertion.
Leaves are scale-like, usually appressed to the stem and each subtends a minute
lateral bud which normally undergoes no further development (Fig. 2D). Leaves
towards the end of the flush are somewhat larger than those found at the beginning.
Branches (Fig. 1A, arrows) are produced at the end of each flush by syllepsis (Hallé
et al., 1978), each branch subtended by a scale leaf. Usually five branches are pro-
duced i.e. one for each orthostichy in the phyllotactic spiral (Plate 1B). The subten-
ding leaves apparently represent the last 5 leaves of the series produced by the apex
of the orthotropic shoot before it undergoes rest. Although there is a short
hypopodium, there is something of a transition in leaf size along the axis at its base
(Plate 1B).
Branches extend almost horizontally and are markedly dorsiventral. They branch
infrequently to produce daughter axes of a second and even third order which repeat
Architecture & phyllotaxis of Anisophyllea disticha 7
the plagiotropic organization (Plate 1C). Evidence for rhythmic growth of these
plagiotropic axes is limited since the only articulations are the daughter branches
themselves, which are usually produced in pairs from adjacent internodes (Plate 1A,
C). No discrete terminal resting buds are produced and there is no regular fluctua-
tion in size of the two kinds of leaf. Direct measurement of growth frequency is re-
quired to demonstrate rhythmic growth, if it exists.
Dorsiventrality is determined by phyllotaxis, as shown in serial sections of buds,
with no secondary reorientation of leaves other than their separation by internodal
extension (Fig. 2E-F). Leaves are four-ranked (Fig. 2E) and consist of two ranks of
scale-leaves arranged alternately on the upper side of the branch (dorsal scales) and
two ranks of larger foliage leaves on the lower side of the branch (ventral foliage
leaves). The internodes between them are extended such that in an acropetal direc-
tion the sequence of leaves is - left scale - left foliage - right scale - right foliage
- left scale ... etc. (Plate 1D; Figs. 1B & 5A). In terms of the genetic spiral, the
1mm
Fig. 1. Anisophyllea disticha (Singapore collection). Morphological details.
A. Apex of orthotropic shoot; sylleptic branches forming terminal tier cut off (arrows).
B. Detail of plagiotropic axis from above to show relative position of branch scales and foliage
leaves; LS-left scale; RS-right scale; LF-left foliage leaf; RF-right foliage leaf; branch scale -
stippled (cf. Fig. 4).
C. T.S. trunk axis at level of insertion of trunk scale to show position of multiseriate gland;
vascular tissue-solid black.
D. Detail of multiseriate gland.
8 Gard. Bull. Sing. 36(1) (1983)
following values represent observed angular divergences proceeding acropetally
(n = 11):-
Sequence Angle of Divergence
Left scale - left foliage 46° + 13
Left foliage - right scale 180° +0
Right scale - right foliage 314° + 13
Right foliage - left scale 180° +0
In Figure 5B, these four sequences are indicated by the numbered sequences 4-5,
5-6, 6-7, and 7-8, respectively.
Scale leaves subtend no axillary buds (Fig. 2K) whereas foliage leaves always sub-
tend at least one bud (Fig. 21) which usually develops as an inflorescence, but occas-
sionally as a vegetative branch which grows out by syllepsis and, as stated, repeats
the construction of its parent axis.
Reiteration of the architecture in the sense of Hallé er a/. (1978) is seen in the pro-
duction of additional orthotropic shoots from the trunk axis, presumably from dor-
mant lateral buds. This most typically occurs in damaged stems with the resulting
‘‘repair’’ of the original crown. However, lower buds occasionally grow up without
obvious damage to the plant. Since the architectural crown-form is so precise and
plagiotropic shoots never produce orthotropic axes, the result is always a narrow
cluster of orthotropic axes each with its own set of branch tiers which become
somewhat internested.
Leaf morphology. Three types of leaves can be recognized on the basis of mor-
phology and position. Symmetrical scale leaves are the leaves of orthotropic shoots,
asymmetrical (dorsal) scale leaves and (ventral) foliage leaves are the two kinds on
plagiotropic shoots (Fig. 3A, C and E). These three types will be referred to as trunk
scales, branch scales, and foliage leaves, respectively. Representative dimensions
from the two fluid-preserved collections are included in Table 1 to show that the two
types of scale leaves are about the same in length. Herbarium specimens confirm
these observations in terms of relative size, but several specimens had unusually
large foliage and scale leaves, up to 80 x 35 mm and 10 x 5 mm, respectively. Ding
Hou (1958) notes that the size of leaves (referring only to foliage leaves) is
‘‘variable’’ and comments that ‘‘all specimens from the Malay Peninsula have small
leaves’’. There may be some overlap in size between leaves of different types, with
large distal scales on orthotropic shoots approaching the size of proximal foliage
leaves on plagiotropic shoots. The two contrasted leaf types on plagiotropic shoots
always retain their relative size difference, with transitional forms restricted to the
branch base.
Mean length of foliage leaves in the two collections varied from 16 mm (Kedah
collection) to 25 mm (Singapore collection) and mean length was in both cases two
to three times greater than mean width. Both types of scale leaves in both collections
were about the same mean length, 5 mm, but trunk scales were fifty percent wider
than branch scales (cf. Fig. 3C and EB).
Fig. 2. Anisophyllea disticha (Singapore collection). Phyllotaxis and nodal anatomy.
A-D. Orthotropic shoot.
A. T.S. bud 8 » below shoot apex.
B-D. T.S. three successive levels to show nodal anatomy.
B. Leaf trace with single leaf gap. C. Branch traces from margin of leaf gap. D. Level of
axillary bud.
E-F. Plagiotropic shoot.
E. T.S. bud at level of shoot apex.
F. T.S. bud 64 » below shoot apex.
G-I. T.S. three successive levels to show: G. Departure of foliage leaf trace; H. Departure of
bud trace.
J-K. T.S. insertion of branch scale to show: J. Unilacunar node; K. Absence of bud.
Vascular tissue - solid black; branch scale - stippled; axis - cross-hatched.
10 Gard. Bull. Sing. 36(1) (1983)
Table 1. Comparative leaf dimensions (in mm)
Kedah Foliage 16 + 2.4 x 7.0 + 1.3 (n=42)
Branch scale 4:95 Ol6) x<eIE Ieee 0447)
Trunk scale Sy) ae Its 6 ila7/ se (45) ((N=7/)
Singapore: Foliage Jy ae 747) S< OD) Se IA) (NSS)
Branch scale So O04 x<aoleg = O!37(n—33)
Trunk scale Sys) ae Wet) Se ALS se OO (nS ily
Foliage leaves (Fig. 3A) are rhombic but asymmetrical, with an acute tip and an
acute, unequal base; branch scales (Fig. 3C) are asymmetrical, lanceolate-ovate but
somewhat falcate with an acute tip and an acuminate or rounded but unequal base.
Trunk scales (Fig. 3E) are almost symmetrical, with an acute tip and an attenuate
or rounded, unequal base. The margins of all leaves are entire, those of the foliage
leaves sometimes slightly inrolled abaxially.
The venation of foliage leaves may be described as acrodromous (Fig. 3A) with
a mid-vein which is prominent below (Fig. 3G). Two pronounced secondary (lateral)
veins originate basally from the mid-vein, with a third lateral vein originating
suprabasally from the anodic or acroscopic side of the lamina. All three secondary
veins run almost to the apex and are connected by regularly arranged cross-veins
(tertiaries) which form a scalariform pattern (Fig. 3B). Venation of both types of
scale leaf is much simpler and may be described as camptodromous and
cladodromous (Fig. 3D, F). The single mid-vein gives rise to minor secondaries
which extend towards and sometimes along the margins and interconnect only occa-
sionally to form an open reticulum.
Both leaves and stems are covered with fine, brown, apparently uniseriate hairs.
These differentiate early, when the leaf primordium is at about the fourth to sixth
plastochron. Each originates from a four-celled basal complex of cells which gives
rise to a Single elongated distal cell, which becomes thick-walled in the mature leaf.
The indumentum of the two types of leaf on plagiotropic shoots is somewhat con-
trasted, since hairs are usually absent from the scale leaves except along the leaf
margins whereas the foliage leaves are quite densely hairy, on the surface but
especially along the margins. Trunk scale leaves have frequent abaxial hairs,
especially at the base of the lamina, as well as marginal hairs.
Multicellular clavate glands, in pairs or sometimes fours occur at the base of each
type of leaf in a stipular position (Fig. 1C, D). They are not vasculated, and develop
precociously so that they are conspicuous in the buds.
Leaf anatomy. Leaves of the three types are of comparable thickness, measured
halfway between base and apex and mid-rib and margin, shown in Table 2, but
mesophyll structure varies appreciably.
1mm
eS
© ‘ec
°
©
2
Fig. 3. Anisophyllea disticha (Singapore collection). Leaf anatomy.
A-B; G-H. Foliage leaf.
C-D; I-J. Branch scale.
E-F; K-L. Trunk scale.
A, C, E. Leaf outline showing major veins, from cleared specimens.
B, D, F. Details of venation in same leaves.
G, I, K. Outline of complete leaf (half-leaf in G).
H, J, L. Detail of mesophyll anatomy.
Colourless hypodermis - lumen dotted in H and K.
11
12 Gard. Bull. Sing. 36(1) (1983)
Table 2. Comparative Leaf Thickness (in microns) (n= 10)
Foliage 130 + 11
Branch scale 140 + 18
Trunk scale 140 + 19
All leaves have a thin, smooth cuticle. The epidermal cells have a sinuous outline
in surface view. Stomata occur on all three kinds of leaves and are restricted to the
abaxial surface, except for a few adaxial stomata towards the leaf apex in the trunk
scale leaves. As shown in Table 3, differences in stomatal distributions are greatest
between foliage leaves and branch scales. Stomata are each surrounded by 4-7
epidermal cells with somewhat less sinuous anticlinal walls.
Differences among the three leaf types occur mainly in the mesophyll. Foliage
leaves (Fig. 3G, H) have a single almost continuous, colourless abaxial hypodermal
layer and a well-differentiated mesophyll consisting of a single adaxial palisade layer
and 2-4 layers of spongy mesophyll. Trunk scale leaves (Fig. 3K, L) may also have
a single layer of colourless hypodermal cells below the abaxial surface (Fig. 3L) but
in neither type of scale leaf is the central mesophyll of 2-4 layers differentiated into
palisade and spongy tissue (Fig. 3J, L). Branch scale leaves (Fig. 31, J) are
distinguished by the numerous epidermal tannin cells, staining dark red with
safranin, which differentiate early but become less conspicuous in mature leaves.
Tannin cells are otherwise common in the mesophyll of all three kinds of leaf. The
general conclusion is that the biggest differences in leaf anatomy are between the
two types of leaf on plagiotropic axes.
Table 3. Comparison of stomatal frequency and stomatal index in different
leaf types (n=10 for each leaf type).
Foliage Leaf Branch Scale Trunk Scale
Stomatal frequencies 93 + 10 SSyebel'§ 70 + 18
(stomata/mm?)
Stomatal index .060 + .008 .049 + .012 .061 + .013
(stomata/epidermal cells)
Nodal Anatomy. Ciné-analysis shows that all leaves have a single leaf trace deriv-
ed from a single leaf gap (Fig. 2B-D; G-I; J-K). Geh and Keng (1974) report the
nodal anatomy for the family as ‘‘many-traced, trilacunar’’, and thereby imply that
this also applies to Anisophyllea disticha. This is a curious discrepancy but it is
possible that these authors refer to the two lateral traces from the upper margins
of the leaf gap which are actually traces to the axillary bud and not the leaf (Fig.
AX, IDB Jal, 10).
Architecture & phyllotaxis of Anisophyllea disticha 13
Internode length. Evidence was sought to show that the scale leaves on
plagiotropic shoots had a constant association with a foliage leaf, since this could
relate to their interpretation as stipules. Figure 4 shows that in the Singapore popula-
tion (lower histogram) there was an average greater length of the internode between
a scale leaf and foliage leaf on the same side of the axis than between scale leaf and
foliage leaf on opposite sides of the axis. However, this relationship was reversed
in the Kedah sample (upper histogram). Herbarium specimens provided further ex-
amples of variation in spacing between the two types of appendage. In many ex-
amples, spacing is about equal. This leads to the conclusion that there is no constant
association between the scale leaf and adjacent foliage leaf which might assist in an
interpretation of the distinctive phyllotaxis.
20 KEDAH
15
1.0
SINGAPORE
O
PSaeeratiemako: waikosne | || 6 RE-LS
INTERNODE
Fig. 4. Anisophyllea disticha. Histogram of average internode length along plagiotropic branch in two
collections (Kedah, Singapore); vertical lines above bars represent standard error, numbers within
bars represent sample size. Internodes are those between left scale and right foliage leaf (LS-LF);
left foliage leaf and right scale (LS-RS); right scale and right foliage leaf (RS-RF) and right
foliage leaf and left scale (RF-LS); cf. Fig. 2B.
O
1
LENGTH IN MM
es N
Ol Orel)
O
i)
Ol
14 Gard. Bull. Sing. 36(1) (1983)
Discussion
Phyllotaxy. Evidence from nodal anatomy demonstrates that the 3 kinds of leaves
in Anisophyllea are homologous since they all have the same unilacunar, single-trace
vascular configuration, and therefore cannot be regarded as referring to two
categories, viz. leaf and stipule, even though accounts of the branch scale leaves
have referred to them descriptively as either ‘‘stipule-like’’ (Ding Hou, 1958) or ‘‘ap-
pearing as stipules’’ (Corner, 1952). Nor are they consistently associated with foliage
leaves in a way which might suggest that they are leaf-opposed stipules (Fig. 4). Fur-
ther evidence is provided by the similar anatomy of scales on orthotropic and
plagiotropic shoots (Fig. 3). The homology between trunk scales and leafy appen-
dages is not disputed since these scales are solitary, with a normal spiral phyllotaxis
and subtend axillary buds. Morphological evidence that the branch scales are
stipules therefore comes solely from the absence of axillary buds. The suggestion
that they might be stipular can otherwise be based only on out-group comparison
since stipules are a characteristic feature of other tribes within the Rhizophoraceae
- (e.g. Rhizophoreae, Gynotrocheae). The microscopic glands at the leaf base are not
stipule homologues despite their position (Fig. 1C).
If we rule out the possibility that Anisophyllea possesses stipules, the phyllotaxis
of the different shoot systems becomes easier to interpret. The orthotropic shoots
have a regular spiral phyllotaxis (Fig. 2A) and offer no interpretative problems.
However, the four-ranked leaf arrangement of the plagiotropic shoots (Fig. 2E) is
difficult to interpret as a modification of any more regular or familiar phyllotaxis
without envisaging major rearrangement of appendages. Each of these major ar-
rangements may be considered in turn: -
1. Spiral. To accommodate a genetic spiral of the type found in spiral phyllotaxis,
in which the angular divergence between successive leaves in the spiral is some
regular Fibonacci fraction of the total stem circumference, would require modifica-
tion of each leaf postion. A change from 144° (the Fibonacci angle) for a
phyllotaxis would have to result in different displacements for each successive leaf
(cf. the values on p. 8). Such a modified spiral is a difficult interpretation to accept.
2. Distichous. A distichous leaf arrangement in the plagiotropic shoots could only
exist if it is hypothesized that two independent sets, of distichous leaves are present.
Two such arrangements can be hypothesized. Following one interpretation, one
series would consist of foliage leaves (leaves 1-3-5-7 in Fig. SA, B), and the other
series would consist of scale leaves (leaves 2-4-6-8 in Fig. 5A, B). The orthostichies
of these separate series would be set an angle of approximately 135° to each other
(Fig. 5B). The sequence of initiation of appendages which is continuously acropetal,
without any evidence for the suggested two separate series, does not support this in-
terpretation. Furthermore, this interpretation is purely descriptive, and does not ex-
plain the phyllotaxis as a modification of a more standard arrangement.
Alternatively, the two series could each consist of a scale and a foliage leaf on op-
posite sides of the stem, that is one series of left foliage-right scale (leaves 1,2,5,6
in Fig. SA, B) and the other of right foliage-left scale (leaves 3,4,7,8 in Fig. 5A, B).
The orthostichies of the two series in this case would be set at an angle of approx-
Fig. 5. Anisophyllea disticha (Singapore collection). Leaf arrangement on plagiotropic shoot
(Diagrammatic).
A. Plagiotropic axis from above, with foliar appendages (both foliage leaves and branch scales)
numbered acropetally from 1 (oldest, basal) to 8 (youngest, distal). Odd-numbered appendages
are foliage leaves; even-numbered appendages are branch scales (stippled).
B. Diagram (not to scale) of axis (cross-hatched) shown in A, with the eight numbered appen-
dages projected onto a transverse plane, in their four orthostichies. Angles between orthostichies
are approximate (see text).
imately 45° to each other (Fig. 5B). This interpretation requires that the sequence
of leaves of each series be interrupted alternately by 0,2,0,2, etc., leaves of the con-
trasted series. This is hardly a standard distichous arrangement.
3. Decussate and bijugate. Particularly difficult problems exist if the phyllotaxis
of the plagiotropic shoots is considered to be a modification of a system in which
leaves are opposite with pairs either at right angles (decussate) or some other regular-
ly repeating angle (bijugate). The last arrangement needs to be considered for com-
15
16 Gard. Bull. Sing. 36(1) (1983)
parative purposes because the phyllotaxis of the Rhizophoreae is of this kind
(Tomlinson and Wheat, 1979) and it seems also to occur in at least some of the non-
mangrove genera (e.g. Carallia, Crossotylis, Gynotroches, Pellacalyx). In Figure
5A, B, a bijugate arrangement would be represented by pairing a foliage and a scale
leaf in the fashion 1-2, 3-4, 5-6, 7-8. However, to accommodate such a modification
would require internodes to be separated, the bijugate angle to be changed both in
magnitude and direction and the development of anisophylly between members of
a pair.
Adaptive considerations. The conclusion is therefore that there are constant dif-
ficulties in seeking for a hypothetical ancestral form of shoot with any kind of or-
thodox phyllotaxis from which Anisophyllea disticha might be derived. The present
conclusion must be that the leaf arrangement on the plagiotropic shoots is unique
and represents a departure from those arrangements usual in angiosperms. A careful
examination of architecture in other species of Anisophyllea might throw light on
the subject.
Contrasted leaf arrangements on plagiotropic and orthotropic shoots are not
unusual in tropical trees conforming to Massart’s and Roux’s models (Hallé et ai.,
1978) but do not usually take the extreme form found in Anisophyllea disticha. It
is reasonable to interpret them as mechanisms for maximizing photosynthesis by
maximizing the surface area exposed and minimizing overlap both among leaves within
one branch complex, and, between branch complexes of different tiers. The adap-
tive success is particularly appreciated because Anisophyllea disticha is a treelet of
forest understorey and seemingly able to survive under a closed canopy under
relatively dense shade, At the same time it is, according to Ding Hou (1958), the
most widespread species of Anisophyllea, both geographically and ecologically. Giv-
nish (1979) has hypothesized that optimizing the shape of leaves in planar arrays in
order to minimize overlap will result in a leaf that ‘‘must be modified from a wedge-
shaped form to one in which the leaf margin roughly parallels the midrib over much
of the midleaf and tapers toward either end. The apical section should remain
roughly wedge-shaped’’. His diagram of such a theoretical system (Fig. 6) shows an
uncanny resemblance to the arrangement and shape of foliage leaves in
Anisophyllea disticha (cf. Fig. 1B). Givnish suggests that this modification of leaf
shape from wedge to parallelogram results in a more efficient ‘‘ribbon of photosyn-
thetic tissue along the branch’’. Anisophyllea disticha seems not only to have
brought this theoretical model to life, but to have improved upon it as well. Scale
leaves further increase the surface area of the ‘‘ ribbon of photosynthetic tissue’’
by patching the holes within it i.e. filling the gaps between the bases of the foliage
leaves as well as covering the dorsal side of the branch. This interpretation does not,
of course, rule out the possibility that the scale leaves have some other function not
associated with photosynthesis. For example, they may be involved in close-packing
of appendages within the dorsiventrally flattened shoot apex as a consequence of its
apparently continuous growth.
Although the phyllotaxis of these plagiotropic branches may be unique within the
angiosperms, the same leaf arrangement occurs in other groups. Several writers have
commented upon the considerable similarity between species of Se/aginella (section
Heterophyllum) and Anisophyllea disticha (e.g. Corner, 1952). However, in
Selaginella leaves are in pairs. Dengler (1983a, b) has recently described leaf
Fig. 6. Theoretical optimal leaf shape and display along one side of a horizontally orientated branch seen
from above; leaves shown on one side of the branch only (after Givnish, 1979).
development in plagiotropic shoots of Se/aginella martensii which shows a similar
dorsiventrality determined by primary leaf arrangement. Her studies were under-
taken to show that appendages which are ‘‘homologous’’ diverge considerably from
each other very early in ontogeny and that the development of small dorsal leaves
cannot be regarded as a simple truncating of the developmental process which oc-
curs in larger, ventral leaves.
Our initial observations lead to the same conclusion in Anisophyllea disticha since
dorsal and ventral leaves can be distinguished histologically within three to four
plastochrons of their inception. At their third plastochron primordia which will
develop into foliage leaves have six cell layers (at a point midway between
longitudinal axis and margin) versus only five layers in scale primordia at the same
developmental stage. Foliage primordia also show evidence of an abaxial ridge cor-
responding to the position of the midvein (Fig. 2E), and stain more densely with
safranin than do scale primordia. Dengler’s work (1983a, b) provides a model for
the further investigation in developmental differences between contrasted leaf types
in Anisophyllea disticha. Such an investigation would complement our initial obser-
vations and would contribute considerably to our understanding of the mor-
phological plasticity of the vegetative parts of higher plants in response to limiting
environmental circumstances.
Biblography
Airy Shaw, H. K. (1973). Jn J. C. Willis, A Dictionary of Flowering Plants and
Ferns. Ed. 8. Cambridge University Press, Cambridge.
Corner, E. J. H. (1952). Wayside Trees of Malaya in two volumes. Vol. 1, Second
Edition. Government Printing Office, Singapore.
17
18 Gard. Bull. Sing. 36(1) (1983)
Dengler, N. G. (1983a). The Developmental Basis of Anisophylly in Selaginella
martensii. I. Initiation and morphology of growth. Amer. J. Bot. 70: 181-192.
(1983b). The Developmental Basis of Anisophylly in Selaginella martensii.
II. Histogenesis. Amer. J. Bot. 70: 193-206.
Ding Hou. (1958). Rhizophoraceae. Flora Malesiana. Ser. 1, vol. 5, pp. 429-493.
Noordhoff-Kolff N.V., Djakarta.
Geh, S. Y. and Keng, H. (1974). Morphological Studies on Some Inland
Rhizophoraceae. Gdns’ Bull. Singapore. 27: 183-220.
Givnish, T. J. (1979). On the Adaptive Significance of Leaf Form. pp. 375-407 in:
O. T. Solbrig, S. Jain, G. B. Johnson and P. H. Raven (eds.) Topics in plant
population biology. Columbia University Press, New York.
Hallé, F., Oldeman, R. A. A. and Tomlinson, P. B. (1978). Tropical Trees and
Forests: an architectural analysis. Springer Verlag. Berlin, Heidelberg and New
York.
Pires, J. M. and Rodrigues, W. A. (1971). Notas SObre os Géneros Polygonanthus
e Anisophyllea. Acta Amaz. 1: 7-15.
Sandwith, N. Y. (1952). Contributions to the Flora of Tropical America LV-
Discovery of Anisophyllea in America. Kew Bull.: 303-306.
Takhtajan, A. (1969). Flowering plants, origin and dispersal. Smithsonian Institu-
tion Press, City of Washington.
Tay, Eng Pin. Architecture and Growth Dynamics in Rhizophoraceae. Unpublished
B. Sc. Honours Thesis, 1977, University of Malaya, Kuala Lumpur, W.
Malaysia.
Tomlinson, P. B. and Wheat, D. W. (1979). Bijugate phyllotaxis in Rhizophoreae
(Rhizophoraceae). Bot. J. Linn. Soc. 78: 317-321.
Van Vliet, G. J. C. M. (1976). Wood anatomy of the Rhizophoraceae. Leiden Bot.
Ser. 3: 20-75.
Zimmermann, M. H. and Tomlinson, P. B. (1966). Analysis of Complex Vascular
Systems in Plants: optical shuttle method. Science. 152: 72-73.
AMESIODENDRON AND LITCHI (SAPINDACEAE):
New Records for the Malay Peninsula
S. K. YAP
Forest Research Institute, Kepong, Malaysia
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983
Abstract
Two genera of Sapindaceae, previously not recorded for the Malay Peninsula, have been confirmed
to occur in local forests. They are Amesiodendron and Litchi, each represented by one species.
Introduction
A total of 16 genera of the Sapindaceae were recognised as occurring in the Malay
Peninsula by King (1893). Ridley (1922, 1925) added a further four genera, Tristira
Radlk., Curtisina Ridl. Napeodendron Ridl. and Phoenicimon Ridl., with the last
three described as new and monotypic. However, all these three genera were later
reduced as Curtisina penangensis was proved to be conspecific with Dacryodes
longifolia in Burseraceae (Lam, 1932), Napeodendron altissimum with Walsura
neuroides in Meliaceae (Symington, 1937), and Phoenicimon rubiginosus as a
species of Glycosmis of the Rutaceae (Leenhouts 1967).
In his monograph of the family, Radlkofer (1932) added two more genera for the
Malay Peninsula, Euphoria Commers. and Pseudonephelium Radlk. Subsequently
Leenhouts (1969) reduced Aphania Bl., Erioglossum Bl. and Otophora Bl. to
Lepisanthes Bl. The two genera Euphoria and Pseudonephelium were reduced to
Dimocarpus Lour. (Leenhouts, 1971) while Tristira sensu Ridley was later reduced
to Glenniea Hook. f. (Leenhouts, 1975). A new record, Ganophyllum falcatum BI.
was added by Whitmore (1969) from a specimen collected from south-east Johore.
The present study documents a further two genera that have not been previously
recorded for the Malay Peninsula viz. Amesiodendron Hu and Litchi Sonn. Each
of the two genera is represented by one species in this region.
Amesiodendron Hu
Amesiodendron chinense (Merr.) Hu, Bull. Fan Mem. Inst. Biol. Bot. 7 (1937)
207. Plates 1 & 2, Fig. 1
Basionym: Paranephelium chinense Merr., Lingnan Sci. J. 14 (1935) 30 & fig 10.
Medium tree 20-30 m tall, 30-60 cm diameter; crown dense, spreading; bole fluted;
bark brown to greenish brown, slightly dippled; inner bark red, laminated.
Twigs glabrous, lenticellate; leaves paripinnate, up to 20 cm long, with 4-6 pairs of
alternate leaflets; leaflets 5-10 cm long, 1.5 cm wide, glabrous on both surfaces, with
19
Fig. 1. Flowers of Amesiodendron chinense (Merr.) Hu — a: part of inflorescence with the male flower
at anthesis; b: female flower; c: the bilobed petal of the male flower; d: the bilobed petal of the
female flower; e: stamen of the female flower; f/: stamen of the male flower; g. transverse section
of the ovary.
20
Plate 1. Inflorescences of
Amesiodendron
chinense (Merr.) Hu.
Plate 2. Fruits of Amesiodendron
chinense (Merr.) Hu.
toothed margin, pink when young, dull green when mature. Inflorescences
paniculate, rather erect, terminal and axillary from the distal nodes, with both male
and female flowers on the same panicle. Male flowers: sepals 5, valvate; petals 5,
bilobed, outer lobe of each petal overlapping the feathery inner lobes; stamens 8(-9),
inserted on a raised disc, filaments scantily hairy and exsert; anthers dorsifixed and
extrorse. Female flowers: smaller than the male flowers, sepals 5; petals 5, bilobed
as in the male flowers; stamen short, not exert from corolla; ovary hairy with 3(-4)
locules; styles hairy. Nuts developing from 1 - 3 lobes of the ovary, usually hard,
dark brown when ripe, dehiscing longitudinally into two; seeds smooth, hard and
brown, 2 x 3 cm; germination hypogeal; seedling epicotyl with numerous scale-
leaves; seedling leaves in spiral phylotaxy, each with 2 - 3 pairs of leaflets and a
scale-like extension on the rachis tip.
The genus is distinct from others in the Malayan Sapindaceae by the combination
of paripinnate leaves and serrate leaflet margins.
21
22 Gard. Bull. Sing. 36(1) (1983)
SPECIMENS EXAMINED:
Kedah: Everett FRI 14145, Gunong Bungsu Forest Res. (KEP! SING; SAN; K;L; A); Kochummen
KEP 98760, Jitra, Bt. Bintang (KEP!)
Perak: Whitmore FRJ 15670, Cameron Highlands Road, 18th milestone (KEP; SING; L); Yap FRI
29392, Cameron Highlands Road, 20th milestone (KEP!)
Selangor: Kochummen FRI 16150, Genting-Gombak Road (KEP!; SING; SAR; K; A), Yap FRI 30656,
Gombak Forest Res. (KEP!)
Negri Sembilan: Yap FRI 28488, Kuala Kelawang (KEP!)
Pahang: Burgess FRI 19370, Lipis (KEP!); Kochummen KEP 97776, (KEP!), FRI 2584, Bentong
(KEP! SING; K; L; A); Whitmore FRI 229, Bt. Tinggi (KEP! SING; K; L; A)
This species occurs in valleys of hill forests, at 300 - 400 m elevation. Observation
on two individuals in the Gombak Forest Reserve, Selangor revealed that flowering
and fruit formation extended over a period of about six months. Over 40% of the
fruits were found to be viable.
Geographical distribution: Sumatra, Indo-China, Malay Peninsula and South
China.
Litchi Sonn.
Litchi chinensis Sonn., Voy. Ind. Or. Chine 2 (1782) 230; Radlk., Pfl. R. Helft. 98
(1932) 914-921. Plate 3
Plate 3. Infructescence of
Litchi chinensis Sonn.
L. chinensis subsp. chinensis, Leenhouts (1978), Blumea 23: 395-403. Medium
trees 25-30 m tall, 40-70 cm diameter; crown compact; inner bark orange. Leafy
twigs up to 3 mm diameter. leaves paripinnate, 10-20 cm long, with 2-4 pairs of
leaflets; rachis glabrous; leaflets stiffly held erect, coriaceous, glaucous below, 6-13
cm long, 2-3 cm wide, midveins distinct below and slightly sunken above, secondary
veins inconspicous, petiolules dark brown to black on drying. Infructescences
paniculate, terminal and axillary from distal nodes, rather erect, 8-18 cm long.
Amesiodendron & Litchi in the Malay Peninsula 23
Fruits developing from one (rarely two) lobes of the ovary, globose 2.5 x 3 cm, in-
dehiscent; pericarp leathery and muricate (warts up to 1 mm high); arillodes thin but
succulent and easily detached; seed ovate with glossy black testa.
Germination hypogeal, seedling leaves paripinnate with one pair of leaflets and
scale-like extension of the rachis tip, dull red and limp when young.
The stiffly held coriaceous leaflets that are glaucous underneath separate this
genus from the other Malayan Sapindaceae.
SPECIMENS EXAMINED:
Kedah: Corner SFN 31543, Jitra (SING!; L) cultivated; Taib KEP 105323, Gunong Inas Forest Res.,
Kulim (KEP!); Ghazali KEP 105272, Gunong Inas Forest Res. (KEP!); Othman KEP 74976, Sik,
Enggang Forest Res. (KEP!)
Penang: Yap FRI 30652, FRI 30653, Botanical Garden, Penang (KEP!) cultivated.
Perak: Chan FRI 19124, Gunong Bubu Forest Res. (KEP!); Wyatt-Smith KEP 76728, Dindings, Lumut
Forest Reserve (KEP!)
Trengganu: Mahirden KEP 79825, 37th milestone, Jerangan, Dungun (KEP!)
Negri Sembilan: Loh FRI 17092, Tampin Forest Res. (KEP!; SING; K; L; A) Yap FRI 30451,
Bt. Tampin (KEP!), Yap FRI 30696, Bt. Tampin (KEP!)
Selangor: Motan & Sow KEP 52214, Bt. Lagong (KEP!)
Leenhouts (1978) regarded the two species of Litchi recorded in Radlkofer (1932)
as the same but recognised three subspecies viz. chinensis, javensis Leenh. and
philippinensis (Rad|k.) Leenh. Comparison with non Malayan specimens annotated
by Leenhouts (at the Singapore Herbarium) confirmed that the Malayan specimens
belonged to subsp. chinensis. In contrast to subsp. chinensis, subsp. philippinensis
has fewer leaflets (2 pairs as opposed to 3 pairs) and has more pronounced warts
on the fruit pericarp. The subsp. javensis has inflorescences with a few spike-like
branches thus differing from the subsp. chinensis. In the Malay Peninsula, L.
chinensis subsp. chinensis occurs on ridges of low hills up to 800 m elevation. With
the wild form previously only recorded from Indo-China, the present records
therefore extend the geographical range of this subspecies further south than was
previously known.
Litchi chinensis is grown for its highly favoured fruits (popularly known as
“‘lychee’’). The introduction of Litchi trees to China is recorded as far back as 100
B.C. and extensive selection for the quality of the fruits has resulted in many
cultivars. Three types of flowers are found on the cultivated lychee viz. male, her-
maphrodite but functioning as male, and hermaphrodite but functioning as female.
These flowers appear consecutively on the same panicle with a very gradual transition
from one type to another (Mustard, Liu & Nelson, 1954). The same authors also
indicated that a favourable range of temperature and humidity encourages fruit set.
This may explain the rarity of flowers and fruits on the cultivated lychee tree in the
Malay Peninsula. The discovery of indigenous Litchi chinensis in the forests of the
Malay Peninsula has important implications for the potential cultivation of this
species.
24 Gard. Bull. Sing. 36(1) (1983)
Geographical distribution: Indo China and Malay Peninsula but widely cultivated
in subtropical countries.
Acknowledgement
I am grateful to Mr. Wong Khoon Meng and Dr. P.W. Leenhouts for going
through the manuscript critically and for their valuable suggestions.
References
King, G (1893). Materials for a Flora of the Malay Peninsula. J. Asiat. Soc. Bengal
62: 705-739.
Lam, J.H. (1932). The Burseraceae of the Malay Archipelago and Peninsula. Bull.
Jard. Bot. Btzg 42: 281-559.
Leenhouts, P.W. (1967). Phoenicimon Ridl. (Sapindaceae) is Glycosmis Correa
(Rutaceae). Blumea 15: 452.
(1969). Florae Malesianae Pracecursores L. A revision of Lepisanthes
(Sapindaceae). Blumea 17: 33-91.
(1971). A Revision of Dimocarpus (Sapindaceae) Blumea 19: 113-131.
(1975). Taxonomic Notes on Glenniea (Sapindaceae). Blumea 22: 411-414.
(1978). Systematic Notes on the Sapindaceae-Nephelieae. Blumea 24:
395-403.
Mustard, M.J., Liu S.Y. & Nelson, R.O. (1954). Observations of Floral Biology and
Fruit Setting on the Lychee Varieties. Proc. Florida State Hort. Soc. 66: 212-220.
Radlkofer, L. (1932). Sapindaceae. Jn A. Engler, Das Pflanzenreich 98.
Ridley, H.N. (1922). The Flora of the Malay Peninsula Vol. 1: 487-501. L. Reeve
& Co., Ltd. London.
(1925). The Flora of the Malay Peninsula Vol. 5: 301. L. Reeve & Co.,
Ltd. London.
Symington, C.F. (1937). Imperfectly Known Species Misplaced in the Flora of the
Malay Peninsula. Kew Bull: 318-320.
Whitmore, T.C. (1969). Notable New Tree Species for Malaya, 1966-1968. Malay.
Forester 32: 70-72.
Several Unusual Malesian Diplazia
M. G. PRICE
Herbarium, NUB, University of Michigan
Ann Arbor, MI 48109, U.S.A.
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983
Summary
The affinities of four small and deceptively similar species of Diplazium (Filices: Athyriaceae) are
discussed. Five names are newly reduced to synonymy and two new combinations are made.
In Genera Filicum, Copeland (1947, p. 151) said he was tempted to describe a new
genus for the Malesian diplazia with narrow dark fronds and abundant black paleae,
of which he accepted six species. He reiterated his belief that they constitute a well-
marked natural group in Fern Flora of the Philippines 3 (1961, p. 412). In 1973
(British Fern Gazette 10: 258) I added what I thought was an additional species and
expressed the opinion that my new species provided a link with Diplazium, so that
separating the group from that genus would be unjustified.
I have now examined types of what I believe are all relevant names and have seen
many other specimens from K, L, MICH, NY, P, PNH, UC, and US, finding that
only four species are involved, and that they belong in three different not very close-
ly related lines within the genus Diplazium. However, they agree in the following
characters: rhizome short-erect, with thick wiry black roots; fronds narrowly ellip-
tic, pinnate or subpinnate, apex coadunate, segments numerous, usually dark;
paleae abundant on stipe and rachis, narrow, usually dark and shiny; veins free. The
four species may be distinguished as follows:
Key to the species
1. Fronds pinnate except near apex; rachis blackish beneath paleae
2. _Pinnae lobed or incised; grooves of pinna-costae entering rachis groove; paleae strongly toothed
SR eo et i ee ae eR TS cs Be ah =, cn, = wine pore ape Ze aieie & « 1. D. egenolfioides
2. Pinnae subentire; rachis channel uninterrupted; paleae entire .......... 2. D. fuliginosum
1. Fronds pinnatifid except sometimes at the very base; rachis brownish beneath paleae
3. Paleae blackish, numerous on stipe, rachis, costae, and veins beneath, also present above
where costae meet rachis; frond narrowing gradually towards base, drying dark greenish
SAUNT a ae SOO EE ME ed niin <clevans cuevaya wava/e oa woes om 3. D. lomariaceum
3. Paleae brown, present on stipe, rachis, and costae beneath; frond not or only moderately nar-
rowed downwards, drying dull greyish brown ..................-- 4. D. porphyrorachis
1. Diplazium egenolfioides Price, Brit, Fern Gaz. 10 (1973) 258, t.1.
Type: Philippines, Luzon, Laguna, Mt Makiling, Price 1762, 29 Aug. 1971
(PNH; BM, GE, K, L, OSAK, SING, US).
25
26
Gard. Bull. Sing. 36(1) (1983)
Paleae black, shiny, strongly and regularly toothed, to 8 x 0.8 mm, tapering
evenly, abundant on stipe and rachis, few on costae beneath. Stipes to 10 cm
long. Lamina to 42 x 15 cm, narrowly elliptic, fully pinnate nearly throughout.
Pinnae to 8 x 2 cm, lanceolate, the lowest on stalks to 1 mm, sessile upwards
and truncate at base, apex acute, margins cut up to % of the way to costa into
segments to 6 mm broad. Colour very dark green. Rachis groove above open
to and joining grooves of pinna-costae, the raised cartilaginous sides of grooves
of pinna-costae joining the sides of the rachis groove, not paleate. Indusia pale
brown, margin slightly undulate. Spores pale brown with a broad plane median
wing.
Distribution: Luzon, the type locality (several collections) and Nueva Vizcaya,
Sta. Fe, Mts. N. of Imugan, c. 1200 m. L.L.Co 1460 (MICH, PNH, PUH).
Notes: The prevalent type of axis architecture in Diplazium is confluent raised
cartilaginous ridges bordering the grooves of rachises and costae. Of the four
species treated here, only D. egenolfioides has this axis character, which im-
mediately links it to the bulk of the genus. Of species with which I am familiar,
fairly close to D. egenolfioides are D. banahaoense (Copel.)C.Chr. and
Diplazium symmetricum (Copel.)Price, comb. nov. (basionym: Athyrium sym-
metricum Copel. Philip. J. Sci. 81 (1952) 41. Lectotype: Philippines, Samar, Mt
Cabayanan, Edafio PNH 15148, MICH).
. Diplazium fuliginosum (Hook.)Price, Brit. Fern Gaz. 10 (1973) 260; Asplenium
fuliginosum Hook. Sp. Fil. 3 (1859) 120; C.Chr. Gard. Bull. S.S. 7 (1934) 280;
Athyrium fuliginosum (Hook.)Copel. Philip. J. Sci. 56 (1935) 476; Asplenium
lugubre Hook. Second Cent. Ferns (1861) t.3 (non Liebm. 1849). Type: North
Borneo, Mt Kinabalu, H. Low (K).
Diplazium acrocarpum Rosenst. Repert. Sp. Nov. Fedde 10 (1912) 328;
Asplenium acrocarpum (Rosenst.)Hieron. Hedwigia 61 (1919) 32; Bot. Jahrb.
56 (1920) 148; Athyrium acrocarpum (Rosenst.)Copel. Philip. J. Sci. 78 (1951)
472; Asplenium fuliginosum forma acrocarpum (Rosenst.)C.Chr. Brittonia 2
(1937) 290. Type: North-east New Guinea, Sattelberg, Keysser 27, 1911 (B?
isotypes NY, UC).
Athyrium longissimum Copel. Philip. J. Sci. 38 (1929) 139, syn. nov.; Fern
Fl. Philip. 3 (1961) 411; Diplazium longissimum (Copel.) C.Chr. Ind. Fil.
Suppl. 3 (1934) 74. Type: Philippines, Leyte, Dagami, Ramos BS 15269, Aug.
1912 (MICH).
Paleae blackish, shiny, brittle, entire, to 6 x 1 mm, abundant on stipe,
rachis, costae and veins beneath. Stipes to 16 cm long. Lamina very narrowly
elliptic, to 79 x 13 cm, fully pinnate in at least the lower a Pinnae to 8 x
2 cm, lanceolate, sessile and truncate at base, apex sharply acute, margins
subentire, very thin and translucent. Colour very dark green, when living suc-
culent with a bluish cast. Rachises above concave or channelled but without
raised cartilaginous sides, insertion of pinnae not affecting channel, not paleate.
Indusia blackish with a thin brown outer edge. Spores pale brown with broad
crisped pale anastomosing wings.
Unusual Malesian Diplazia Dil
Distribution: Bismarck Arch. (New Ireland), New Guinea (widespread), North
Borneo (Mt Kinabalu, common), Philippines (Leyte, one collection), occurring
in shaded moist ravines in montane forests, c. 1000-3000 m.
Notes: Diplazium acrocarpum has sori confined to the pinna-apices but this
character is inconstant and unreliable, as both fully fertile and only apically fer-
tile pinnae may occur on a single frond, for example as in Holttum SFN 25529
from Mt Kinabalu (US). Athyrium longissimum has narrow fronds (c. 5 cm
broad) but agrees in all essential characters. Hoogland 9323 (NY) is aberrant in
having shallowly lobed pinnae.
D. fuliginosum is one of the most unusual of all diplazia, and was not
transferred to that genus until recently. It is strikingly peculiar by the smooth
rachis channel without raised sides, uninterrupted by the insertion of pinna-
costae. In small fronds, and distally on large fronds, the rachis is almost flat
above. Other of its features unusual to Dip/azium are sori uniformly extending
from costa to margin, and thin translucent pinna-margins.
Rather than to the other three species treated here, I believe that the closest
relationship of D. fuliginosum is to D. cumingii (Presl)C.Chr., with which it
agrees in dark frond colour and black axes aging to greyish; axes without car-
tilaginous ridges; paleae on stipe abundant, dark, entire; and indusia black with
very narrow pale brown margin, curling back at maturity; even though D. cum-
ingii is very different in its conform frond apex and broadly elliptic pinnae.
However, it should be noted that almost exactly the same frond form (and even
margin structure) as D. fuliginosum was independently evolved in the Central
American D. harrisonii (Bak.)C.Chr., which otherwise differs markedly in
paleae and the architecture of the axes.
3. Diplazium lomariaceum (Christ)Price, comb. nov.; Asplenium lomariaceum
Christ, Verh. Naturf. Ges. Basel 11 (1895) 229. Type: Central Celebes,
Takalekadjo, F. & P. Sarasin 994, 8 Feb. 1895 (P, lectotype).
Diplazium merrillii Copel. Philip. J. Sci. 2C (1907) 128, t.2A, syn. nov.;
Hieron. Bot. Jahrb. 56 (1920) 134; Athyrium merrillii (Copel.)Copel. Philip. J.
Sci. 3C (1908) 300; 56 (1935) 476; Fern Fl. Philip. 3 (1961) 411. Type: Philip-
pines, Mindoro, Mt Halcon, Merrill 5914, Nov. 1906 (Lectotype MICH; US).
Diplazium porphyrolepium v.A.v.R. Bull. Jard. Bot. Buitenz. II, 20 (1915)
11, syn. nov. Type: Celebes, Soemalilah, Capt. van Vuuren’s Explor. Comm.,
Rachmat 481 (BO?; L).
Diplazium porphyrophyllum v.A.v.R. Bull. Jard. Bot. Buitenz. II, 28 (1918)
18, syn. nov. Lectotype: Ceram, Wai Lantabi, L. Rutten’s Explor. Comm.,
Kornassi 1240, 4 May 1918 (L).
Athyrium altum Copel. Philip. J. Sci. 38 (1929) 138, syn. nov.; Fern FI.
Philip. 3 (1961) 411; Diplazium altum (Copel.) C.Chr. Ind. Fil. Suppl. 3 (1934)
72. Type: Philippines, Mindanao, Agusan, Mt Urdaneta, Elmer 14081, Oct.
1912 (MICH; L, NY, US).
28
Gard. Bull. Sing. 36(1) (1983)
Paleae blackish, shiny, entire, to9 x 1 mm, gradually narrowed to a hair-tip,
abundant and persistent on stipe, rachis, costae, and veins beneath. Stipes of
fertile fronds to 15 cm long, of sterile to 9 cm. Lamina narrowly elliptic, to 50
x 9.5 cm, deeply pinnatifid, one pair of reduced basal pinnae sometimes free
and sessile. Lobes to 4.5 x 1 cm, oblong-lanceolate, narrowing towards apex,
subentire, blunt. Colour dark greenish-brown, + shiny below when dry, dark
bluish-green when living. Rachises above with a channel formed by raised car-
tilaginous sides, either continuous or interrupted at each junction with a mid-
vein of a lobe, and paleate at that point whether or not interrupted. Indusia
brown, margin erose. Spores brown, with irregular short wings.
Distribution: Philippines, including Luzon (Ilocos Norte - Price 2932, Rizal,
Quezon), Mindoro, Leyte, Negros, Mindanao, Basilan; Borneo, including
Sabah, Sarawak, and Kalimantan; Celebes, Ceram, New Guinea (collected only
in North-east), in moist montane forest, 400-2000 m.
Notes: Diplazium merrillii was a small plant only partially fertile. The holotype
was destroyed in 1945 at the PNH so I here designate the MICH specimen as
lectotype. D. porphyrolepium and D. porphyrophyllum are not exceptional in
any way. The latter had a syntype purportedly from Sumatra, Brooks 322S
which I have not seen and from where this species is otherwise unknown.
I designate the specimen from Ceram at L as lectotype. Athyrium altum is a
form with narrow fronds not otherwise distinguishable. Three specimens from
eastern Kalimantan (Kostermans 9089, Meijer 577, 872 - all L) differ by having
fewer and brownish paleae but agree in distribution of paleae, and in frond
form and colour. Diplazium lomariaceum is very closely related to D. por-
Dhyrorachis and until now the name seems to have been ignored since Christ
himself reduced /omariaceum to porphyrorachis in Ann. Jard. Bot. Buitenz. 15
(1898, p. 119).
. Diplazium porphyrorachis (Bak.)Diels, Nat. Pflanzenf. I, 4 (1899) 225;
Asplenium porphyrorachis Bak. J. Bot. 17 (1879) 40; Icon. Pl. 17 (1886) t.1650;
C.Chr. Gard. Bull. S.S. 7 (1934) 279, p.p.; Athyrium porphyrorachis (Bak.)
Copel. Philip. J. Sci. 3C (1908) 300; Polypodium subserratum Hook. Sp.
Fil. 4 (1863) 202 (non Diplazium subserratum (Bl.) Moore, 1861). Type:
Borneo, prob. W. Sarawak, A. R. Wallace s.n. 1855 (K).
Paleae dull brown, entire, to5 x 0.5 mm, rapidly narrowed to a hair-tip, pre-
sent on stipe, rachis and costae beneath but relatively inconspicuous. Stipes of
fertile fronds to 37 cm, of sterile to 13 cm long. Lamina lanceolate, to 35 x
9 cm, deeply pinnatifid with up to two free pairs of sessile opposite basal pin-
nae. Lobes to 5.5 x 1 cm, narrowly oblong with mostly parallel sides, apex
usually rounded and denticulate, bluntly acute in the largest fronds. Colour dull
greyish-brown when dry, dark green when living. Rachises above with a channel
formed by raised cartilaginous sides, continuous or interrupted at each junction
with the midvein of a lobe, not paleate. Indusia brown, margin erose. Spores
pale brown with short irregular translucent wings.
Distribution: Borneo, including Sarawak (where often common), Brunei,
29
Unusual Malesian Diplazia
Sabah, and Kalimantan, terrestrial on embankments and petrophytic, in forest
shade, 300-1000 m.
Notes: Christensen gave the unpublished varietal name murudense to a large
specimen narrowed towards the base with long-pointed lobes (Sarawak, Mt
Murud, Sar. Mus. Nat. Coll. 2937, PNH). Similar specimens have been found
on the nearby Gunong Mulu (M. Hotta 15188, L) and at Matang (Sinclair &
Kadim 10346, K).
The two species D. porphyrorachis and D. lomariaceum are very closely
related and some specimens show signs of apparent gene interchange. They were
combined by Christensen (1934) but seem to be amply distinct by the characters
utilized in the key. I believe that the closest relationship of these two with other
species is not at all with the other two treated in this paper, but, of the diplazia
I know, with D. sorzogonense (Presl)Presl. D. sorzogonense has similarities to
especially D. porphyrorachis in paleae, indusia, spores, frond colour and tex-
ture, and the structure of the pinna axes of the former is similar to the rachis
structure of the latter.
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The Limestone Hill Flora of Malaya IV*
S.C. CHIN
Botany Department, University of Malaya
Kuala Lumpur, Malaysia
Angiosperms—Monocotyledons cont.
GRAMINEAE
. Inflorescence a series of successive nodal spikelet-clusters on a leafless axis. Bamboos ......
Inflorescence different, paniculate or spicate. Not bamboos ................0..00eeeeeeee
MRCAVESTSOM-NAILY DEDEALA, Sonic ieee aw riasiaiers SteinvevSevd eieiscecevsisie asers Dendrocalamus elegans
Meavesmousort-nairy beneath «sco en seis Fe to aero ste Odes eleaeles Dendrocalamus dumosus
. Inflorescence from the upper leaf-axis, female spikelets enclosed within a hard bead-like sheath
5.0.0 0-46, O05. COOL CIS OCOEIOTTOR CHICO CIO LA Daan AIH CAE Cie at ema emee Coix lacryma-jobi
. Inflorescence of a single unbranched spike, usually at the end of the stem; or of 2 or more spikes
radiating from or near the end of the stem; or the inflorescence compact and sparsely branched;
all prt eGs aii (etl Aol. Ue ke Oe arts POs Ne 9 ie tse ne a ee een eee
. Inflorescence of a single spike at the end of stem or upper leaf-axils; or compact and sparsely
BME lICA MDL ANGCHES EW RODIER ete ae ieee ee Schone eke ha a tere Salons oe eerie deine
Inflorescence of 2 or more unbranched spikes radiating from or close together at or near the end
DETLE SIG ‘oss Wek bese GUE co me obs toe Sto bon Geto Coen Cc Gore ae ae i item
Inflorescence hairy or bristly, the bristles from the base of spikelets or as awns from the glumes
OP WETNESS * bien d's oR DOD AG OO ORES O SORTS GOERS eREI 5 Chics Pema a eR A Fs AISI re PSR A eer AE
. Inflorescence covered with fluffy hairs ................... Imperata cylindrica var. major
MTL IGRESCET CCRC INL CLT Dern: PHI Te Lecorer rete enero te sere ovchers ashe rene cocoon ctanatane vorctete HOR RTe dette ore nets
MBS DIKELEES PEACHIL WILE DNA WISH Pavey sr evaxrorcestocclcfevs) ener sicterelsnenehore’s axthevetel ste Pogonantherum paniceum
SPIRCICES PCACHEWILE WAWTIESSOMEy WItE OU Gos taps role yore ralalave <tercusseintererayercvoters lo-slevera sete mettre ae eis
Splkcleissin, pairsiyl sessilevand Mepedicelledie a iyirsals 8c ccabottye lec aeeteve Gate aes MR orate ie Sees
APC ICESEITINOTOU Sp Olt EDL CLS tik Noy of stacey Naor 01h 0 sunt oof error Ovwh ov cndy ate) ov ove, Af oKonor A aneet av 8) OBlererore dae
Leaf-base narrowed gradually; lower glume of the pedicelled spikelet with a 3 point-tip ....
RRM Tote ere foc ince atte sic cys Sccravara\otata ciefotayeerenslarcioxs- sete ciavee Snare Dichanthium mucronulatum
Leaves usually less than 0.5 cm wide; spikelets imbricate; nodes of stem glabrous .........
MUM PTV TS Bs cs, «. s\evca. vis e.tue sieved. wiwiate; ave anes Atane stele Ghd, SPS Dichanthium caricosum
Leaves usually more than 0.5 cm wide; spikelets distantly spaced, axis visible; nodes of stem
HETEAY. 25 gua cP) cn O'S or Geeicna OME TERZ a ae AAS OCICS RRC Emer Dichanthium annulatum
Inflorescence a spike-like raceme; 1 spikelet of each group pedicelled, 2 sessile ............
PPE TTT TOTTORI. 50a) ciceve; ot cVavot oy 5 -cic TOVNG Re ae MNO else have Abe, coche Me ialatove Soave ROR YE Polytrias amaura
Inflorescence a small panicle, branches in whorls; 2 spikelets in each group pedicelled, 1 sessile
12
11
13
* This is the final part. Parts I, II and III appeared in the Gardens’ Bulletin xxx (1977) 165-219, xxxii
(1979) 64-203 and xxxv (1983) 137-190, respectively. The author expresses his profound thanks to Dr.
Chang Kiaw Lan for her painstaking assistance.
31
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30.
Gard. Bull. Sing. 36(1) (1983)
Creeping, grasses:sLeaves to sncm 1ongie eeite. ciety erased Chrysopogon aciculatus
Erect grasses: eavestmoresthan) L5icmplonge eer cee et ieee natty reine tener nen ates
Awns 2-3.5 cm long; leaves to 35 cm long; the joint beneath the 3 spikelets swollen .......
ee eine RIOR Oa cco. cto obo omertgoen wndoD.ct 6 Chrysopogon fulvus
Awns 4-5 cm long; leaves to 20 cm long; the joint beneath the 3 spikelets not swollen .....
en a eee na PTC Oo ao cole POS OCC 6 bk OU N Oe UEC Chrysopogon orientalis
Spikelets witht ws) sco. 2:02. cissneiede suanerevaperaeiieuatts couazs' ava) cwueiepel stoi coseteetaMenereles Stone ten ctiotete cotter rr seen en nen
Spikelets without @ws: cassis sccsieie ed yspseht lacie te oils nos iach vwlarelin atene oeueenegete ned oveteeetenetelesomsteke rat area teker tens
Leaves linear-acuminate, to 20 by 0.3 cm; base not cordate. Only from Langkawi .........
sev & a /5igep Sire apejeavonenieripa-eveitE She ations Gus vrareuaila wa tavon seta 80 rene: vbororenece tego Mente ee Mere Eulalia quadrinervis
Leaf bases gradually narrowed to the petiole; inflorescence of 2 often closely-opposed racemes
eocavay aud Svaes beta Ole: oie eNAT <I e SS ole oR) SA erties oe reer oe Ischaemum timorense
Leaf bases cordate; inflorescence of 2-many racemes from the end of a common peduncle
drole: aaa rales nors Wolpe ple eraabe « 7 (ales ovaha) 3 Safe alae eh ete tere ete tetera Arthraxon prionodes
Spikelets fringed with fine hairs (from the upper glume) ........... Paspalum conjugatum
Spikelets not fringed) with hains:.«..,3.sss.cccgscjoe ones orisiero mcs oie) sate rei iees ee ee ROR Ore ae
Spikelets flat, with 4-6 florets arranged alternately ....................... Eleusine indica
Spikelefs not flat, florets 2. <:¢ sitetu.o Sele eicate scott ce eteae, Weetreene stele aaceteue = aerate ee eet ae ee
Inflorescence single, from the topmost leaf axil; lemma of the upper floret thin, margin flat
Oe re eA ere ra noch osaediGacdc os Gach Aco door Digitaria violascens
Inflorescence usually 2, from the topmost leaf axil; lemma of the upper floret thickened, margin
hetge) | (2c ne ir ae REN SoG oo amo oa dD ach. com00 66 Axonopus compressus
Inflorescence with) fineisilky»hairs}from\thelspikelets) ey. eens eileen
Inflorescence! with hairs not silkysifi presenti mcrasckiciie elie raie einer eke ieee
Plant tall and reed-like; inflorescence much branched, 30-50 cm long. Florets 4-8, hairs from
13 so (1011 te ee ee Sener hemes Bite OO BG a HMS Santa cot ac.giae Neyraudia reynaudiana
Plant not reed-like; inflorescence smaller, 10-15 cm long. Florets 2, hairs from all over the
Spikelets; often, pinkish. 5, cac5%c.1rs sores orton are eee raeeS Rhynchelytrum repens
eaves withsfineslongitudinalipleats seem eiieis ciel aes ie Setaria palmifolia
Leaves Mot. Pleated seca iave:ta.é;s,cosonasauossne sorera iene ane wo shysueueres eget yeaeueouaisaelevoreiere COR a cee ae ee
Leaves: broads Siem Or MOre: WIGS, s...6.55<: scaversvopsnendvajlovevov, kecess tonsa.) oiRTeTeRe te Rete: HORE onan aera
Tu@AVES: MALTOWET .ncs 6 pss ssieve so asap esondganw sso ous faye souk soceue SNORT e ONTO tr ne a eee
Leaves elliptic, usually 20-25 by 5-6.5 cm. Lemmas with 5 or more nerves. Plant to 1 m tall
#54 6/6, Uralle carey aus ees ond, wpey atopy apectave tows egal ep el gy Oey See Gere er Oey avers, Seer cog es One cee een Leptaspis urceolata
Leaves lanceolate, usually 40-45 by 5-6.5 cm. Lemmas with 1-3 nerves. Plant to 3 m tall
we ssoarbelin faceenne sie a: tneaeye ie RUCISuc TN RO OPER EL TAL OSES SICeemie euel ere ere arene Thysanolaena maxima
Spikelets, all or some: withawns cyeticctohetcereiias crore tate ciate arenes Penn) cere ean
Spikelets) without awns,ashore bristle presen tlOmnOtmnitcerrieiericieneiieieirsiel icteric rere ieieneteentiaes
Mufted\icoarseigrass-sleaves) to lOOkcmiulongiemee eee ieee Cymbopogon calcicola
Different: leaves: shorter) s.. dcx etcetera athe SOE Seer A Set. Re OE eee
Inflorescence of 3-7 spike-like racemes arranged alternately on the axis. Spikelets in pairs
eC a ero cat CaCO oc LOO ROTO NSO ont ode Oplismenus compositus
Inflorescence different. Spikelets in ultimate groups of threes ............. Apluda mutica
Spikelets with) prominent bristles)or hairs. @uliis) CUnte den a rerererensretereteney ete tetenenet teten-f-teuetenetedeers
Spikelets without prominent bristles or hairs; glabrous or with inconspicuous bristles. Culms
scrambling - i ¢3.ci08.atieniedt dag eee ee ae ae ee Panicum sarmentosum
Leaves more than 1.5 cm wide. Spikelets with stiff, backward pointing hairs on the distal half
of thellemmayoftheluppentlorcteeeeee Ener eccentric Centotheca lappacea
Leaves narrower, 0.2-0.9 cm wide. Spikelets with soft erect hairs all over the outer surface of
the upper glime® <::.5..6. 60 vad sees OO ee Isachne langkawiensis
16
18
17
19
20
22
23
25
26
27
29
28
30
Limestone hill flora of Malaya IV 33
Apluda mutica L., Sp. Pl. (1753) 82; Gillil., Fl. Mal. 3 (1971) 273.
A. varia Hack. var. intermedia Hack., in DC., Monogr. Phan. 6 (1889) 196;
Ridl., Mats. 3 (1907) 164; id., Fl. 5 (1925) 207; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 86.
Slender grass, 1-2 m tall. Leaves pale bluish green, blade to 30 by 1.5 cm. In-
florescence as terminal panicle of numerous racemes with three spikelets each; the
lowermost spikelet with an inflated joint.
Distributed in Ceylon, India and Australia. In Malaya, only from the northern
half, not common but often on limestone.
Arthraxon prionodes (Steud.) Dandy, in Andrews, Fl. Pl. Sudan 3 (1956) 399;
Gillil., Fl. Mal. 3 (1971) 287.
Andropogon prionodes Steud., Syn. Pl. Glum. 1 (1854) 383.
Slender decumbent grass, leafy almost to the inflorescence. Blades 3-6 by 0.8-1.5
cm, lanceolate with cordate base. Inflorescence of several slender racemes to 6 cm
long. Spikelets paired, one sessile and the other pedicelled. Sessile spikelet 5.5 mm
long; the lower glume lanceolate, the upper boat-shaped, both with a wide hyaline
margin. Upper lemma with an awn 4.5 mm long. Stamens 3. Pedicelled spikelet
awnless, with a short 2-lobed lemma.
Distributed in India. Recorded once from Malaya, from limestone crevices on the
summit of Gunong Baling.
Axonopus compressus (Sw.) Beauv., Ess. Agrost. 12 (1812) 154, 167; Ridl., Fl. 5
(1925) 215; Burk., Dict. Econ. Prod. Mal. (1935) 276; Gillil., Fl. Mal. 3 (1971)
187.
Paspalum platycaulon Poir., Ridl., Mats. 3 (1907) 125.
Probably the commonest lawn and wayside grass in Malaya. Recorded from
disturbed localities on limestone.
Centotheca lappacea (L.) Desv. Nouv. Bull. Soc. Phil. Paris 2 (1810) 189; Ridl., Fl.
5 (1925) 253; Burk., Dict. Econ. Prod. Mal. (1935) 508; Gillil., Fl. Mal. 3 (1971)
53.
C. latifolia Trin., in Henders., J. Mal. Br. R. As. Soc. 17 (1939) 86.
Cenchrus lappaceus L., Sp. Pl. ed. 2 (1763) 1488.
All over the secondary and disturbed forests. Recorded several times from
limestone.
Chrysopogon aciculatus (Retz.) Trin., Fund. Agrost. (1820) 188; Ridl., Fl. 5 (1925)
207; Burk., Dict. Econ. Prod. Mal. (1935) 535; Gillil., Fl. Mal. 3 (1971) 236.
Andropogon aciculatus Retz., in Ridl., Mats. 3 (1907) 166.
Common all over Maiaya, in lawns, by wayside and in cultivated area. Recorded
once from the disturbed summit of Bukit Takun.
Chrysopogon fulvus (Spreng.) Chiov., Fl. Somala 1 (1929) 327; Gillil., Fl. Mal. 3
(1971) 237.
C. collinus Ridl., Fl. 5 (1925) 208; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 86.
34 Gard. Bull. Sing. 36(1) (1983)
Recorded twice in Malaya, once from Pulau Burong in Pahang (not limestone)
on dry, sandy or stony soil and the other from limestone on Bukit Wang, Kedah
(Haniff 649).
Chrysopogon orientalis (Desv.) A. Camus, in Lecomte, Fl. Gen. Indochine 7 (1922)
332; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 86; Gillil., Fl. Mal. 3 (1971) 238.
C. wightianus (Nees ex Steud.) Thw., in Ridl., Fl. 5 (1925) 208.
Andropogon wightianus Nees ex Steud., in id., Mats. 3 (1907) 167.
Recorded in Malaya from limestone in Langkawi and from sandy coastal areas
of the east coast. Like C. fulvus, this is a species of dry, well-drained, open
localities.
Coix lacryma-jobi L., Sp. Pl. (1753) 972; Ridl., Fl. 5 (1925) 191; Burk., Dict. Econ.
Prod. Mal. (1935) 629; Gillil., Fl. Mal., 3 (1971) 304.
Common in Malaya, in cultivation or in wasteland near villages. Recorded from
limestone near the railway station at Merapoh in Pahang, probably an escape from
nearby villages.
Cymbopogon calcicola Hubb., Kew Bull. Misc. Inf. (1941) 24; Gillil., Fl. Mal. 3
(1971) 297.
Culms tufted. Blades linear, to 100 by 1.5 cm. Inflorescence a much branched
panicle, characterised by numerous brown spathes each subtending a pair of spike-
like racemes.
Distributed in peninsular Thailand. Recorded in Malaya from Kedah and
Pahang; abundant on the exposed rocky slope of Bukit Chintamani. Restricted to
limestone. Faintly but distinctly scented.
Dendrocalamus dumosus (Ridl.) Holtt., Gard. Bull. S. 11 (1947) 296; id., 16 (1958)
96; Gillil., Fl. Mal. 3 (1971) 30.
Schizostachyum elegans Ridl., J. Str. Br. R. As. Soc. 73 (1916) 146; id., Fl. 5
(1925) 271.
Culms to 5 m long and 2.5 cm thick. Culm-sheaths to 18 cm long; auricles small,
bristly; ligule 1.5 mm tall, toothed. Blades usually 18 by 1.8 cm, sometimes larger.
Spikelets, 1-flowered, tufted at nodes of slender leafless branches. Palea 7 mm long,
7-veined, thin, not keeled. Stamens 3-6; ovary glabrous.
Distributed in the southern Thai islands. In Malaya from Langkawi and mainland
Kedah, probably restricted to limestone. Resembles D. elegans (which see).
Dendrocalamus elegans (Ridl.) Holtt., Gard. Bull. S. 11 (1947) 296; id., 16 (1958)
95; Gillil., Fl. Mal. 3 (1971) 30.
Schizostachyum elegans Ridl., J. Str. Br. R. As. Soc. 73 (1916) 146; id., Fl. 5
(1925) 271.
Culms to 6 m long and 2.5 m diameter. Internodes to 26 cm long; culm-sheaths
not seen. Blades to 12 by 1.2 cm, lower surface densely short hairy. Auricle small;
ligule short. Spikelets in dense tufts at nodes of infloresence branches, about 8 mm
Limestone hill flora of Malaya IV 35
long, glabrous. Florets 1 or 2; lemma of upper floret 7 mm long. Palea of lower
floret 2-keeled, short-hairy on the keels. Upper palea 4-veined. Anthers 6. Ovary
smooth; style slender, stigma plumose.
So far known only from Langkawi and probably restricted to limestone. (The
original plant from which the species was described is cultivated at the Penang
Botanic Gardens, but with the source unknown. Subsequent collections have all
been from limestone).
This species is very much like D. dumosus but differs in the hairy leaves, the 6
instead of 3-6 stamens, the glabrous upper half of the ovary and the frequent
presence of 2 florets with the lower palea keeled and the upper with 4 instead of
about 7 veins. However, in one specimen (Chin 523), the leaves are glabrous, the
stamens number 3-6 and the upper palea has 4-7 veins, yet the upper half of the
ovary is glabrous and there are 1-2 florets.
The position of these two species would be clearer if more specimens were
examined; there is a possibly a gradation of morphological characters from one
species to the other.
Dichanthium annulatum (Forsk.) Stapf., in Prain, Fl. Trop. Af. 9 (1917) 178;
Gillil., Fl. Mal. 3 (1971) 284.
Andropogon annulatus Forsk., Fl. Aegypt. -Arab. (1775) 173.
Slender, branched, erect grass to 1 m tall. Blades to 30 by 0.8 cm. Inflorescence
1-several spike-like racemes from almost every leaf-sheath.
Distributed in Africa, India, Burma and Borneo. In Malaya restricted to
limestone and then only found in Kedah and Selangor; the note in Gilliland, l.c.,
that it has been recorded from Singapore is an error and could have been intended
for D. mucronulatum (which see).
Dichanthium caricosum (L.,)A. Camus, Bull. Mus. Hist. Nat. Paris 27 (1921) 549;
Ridl., Fl. 5 (1925) 210; Gillil., Fl. Mal. 3 (1971) 284.
Andropogon caricosus L., Sp. Pl. ed. 2 (1763) 1480.
Dichanthium mucronulatum Jansen, Act. Bot. Neerl. 1 (1953) 474; Gillil., Fl. Mal.
3 (1971) 283.
Ischaemum beccarii sensu Ridl., Mats. 3 (1907) 160, non Hackel; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 86.
Slender grass to 75 cm tall. Blades narrowly linear to 30 by 0.3 cm, glabrous. In-
florescence a terminal spike-like raceme about 2.5 cm long. Spikelets in pairs, one
sessile, one pedicelled. Sessile spikelet with lower glume 7-nerved and margin inflex-
ed, the upper glume 3-nerved. Pedicelled spikelet with the lower glume tipped with
3 points.
Endemic to limestone in Malaya. Ridley recorded that this species appeared in the
Singapore Botanic Gardens, and, apart from this very unusual occurrence (details
not available) this species has been recorded only from Pahang and Selangor. A
plant of dry rocky and exposed or partially exposed places.
Digitaria violascens Link, Hort. Berol. 1 (1827) 229: Gillil., Fl. Mal. 3 (1971) 191.
D. chinensis sensu Ridley, Fl. 5 (1925) 215, non Hornem.
36 Gard. Bull. Sing. 36(1) (1983)
Pan-tropical. Found all over Malaya from the lowlands and hills to 1600 m.
Recorded once from the disturbed summit of Bukit Takun, and according to Ridley
from Gua Batu.
Eleusine indica (L.) Gaertn., Fruct. 1 (1788) 8; Ridl., Mats. 3 (1907) 174; id., Fl. 5
(1925) 250; Gillil., Fl. Mal. 3 (1971) 78.
Cynosurus indicus L., Sp. Pl. (1753) 72.
A common grass all over Malaya in open places and a weed of cultivation.
Recorded once from the disturbed summit of Bukit Takun.
Eulalia quadrinervis (Hack.) Ktze, Rev. Gen. Pl. 2 (1891) 775; Gillil., Fl. Mal. 3
(1971) 244.
Pollinia quadrinervis Hack., DC., Monog. Phan. 6 (1889) 158.
Culms slender, densely tufted, 50-100 cm tall. Blades linear to 20 by 0.3 cm. In-
florescence of 3-6 racemes arranged subdigitately at the end of a peduncle.
Distributed in India, Burma, Thailand and China. (Restricted to the limestone in
Langkawi, Malaysia.)
Imperata cylindrica (L.) Beauv., Ess. Agrost. (1812) 165; Gillil., Fl. Mal. 3 (1971)
220.
var. major (Nees) Hubb. ex Hubb. et Vaugh., Grass. Maur. (1914) 96; Gillil., op.
Git2205
Widespread everywhere in Malaya, often on disturbed summits of limestone hills.
Isachne langkawiensis Jansen, Reinw. 2 (1953) 284; Gillil., Fl. Mal. 3 (1971) 120.
Tufted, slender grass to 40 cm tall. Blades 2-8 by 0.2-0.9 cm. Inflorescence a ter-
minal panicle 5-12 by 4 cm, with many very fine branches. Spikelets ovate, upper
glume with many rows of hairs with bulbous base.
Endemic to the limestone in Langkawi, often abundant in open places with little
moisture and thin soil cover. Distinguished by the finely hairy leaves and the very
finely bristled upper glume.
Ischaemum indicum (Holtt.) Merr., J. Arn. Arb. 19 (1938) 320; Gillil., Fl. Mal. 3
(1971) 263.
I. aristatum sensu Ridley, Fl. 5 (1925) 203; Henders., J. Mal. Br. R. As. Soc. 17
(1939) 86.
Ischaemum timorense Kunth, Rev. Gram. 1 (1830) 369; Ridl., Fl. 5 (1925) 203;
Gillil., Fl. Mal. 3 (1971) 264.
I. macrurum Stapf. ex Ridl., Fl. 5 (1925) 203.
Leptaspis urceolata (Roxb.) R. Br., in Benn., Pl. Java Rar. (1838) 23; Ridl., Fl. 5
(1925) 255; Gillil., Fl. Mal. 3 (1971) 47.
Pharus urceolatus Roxb., Fl. Ind. 3 (1832) 611.
Distributed in lowland and hill forest, recorded once from limestone (Gunong
Pondok, Perak, Chin 886).
Limestone hill flora of Malaya IV 37
Neyraudid reynaudiana (Kunth) Keng ex Hitch., Amer. J. Bot. 21 (1934) 131; Gillil.,
Fl. Mal. 3 (1971) 61.
Triraphis madagascariensis sensu Ridley, Fl. 5 (1925) 251.
Common in the North as a grass of open wastelands. It has been collected as part
of secondary vegetation on dry summits of limestone. On the top of Gua Musang,
Kelantan, it was common (in 1971) about 1% years after a fire had destroyed the
original vegetation and on the Perak Cave Temple limestone (in 1971) it formed a
prominent part of the vegetation in the disturbed areas. Though frequently recorded
as a grass of damp ground it tolerates the dry conditions on limestone very well,
growing in soil pockets amidst boulders.
Oplismenus compositus (L.) P. Beauv., Ess. Agrost. 54 (1812) 168, 169; Ridl., FI.
5 (1925) 221; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 87, Gillil., Fl. Mal. 3
(1971) 171.
Panicum compositum L., Sp. Pl. (1753) 57.
A true forest grass of lightly shaded places; common in the northern part of
Malaya and recorded from limestone in Langkawi, Kedah mainland and Perlis only.
Panicum sarmentosum Roxb., Fl. Ind. 1 (1820) 311; Ridl., Fl. 5 (1925) 227; Gillil.,
Fl. Mal. 3 (1971) 139.
Paspalum conjugatum Berg., Act. Helv. Phys. Math. 7 (1772) 129; Ridl., Fl. 5
(1925) 218; Gillil., Fl. Mal. 3 (1971) 180.
Widespread in lawns, waysides and cultivated ground. Recorded from disturbed
areas on limestone.
Pogonatherum paniceum (Lamk.) Hack., Alg. Bot. Z. 12 (1906) 178; Gillil., Fl.
Mal. 3 (1971) 251.
Saccharum paniceum Lamk., Encycl. Meth. Bot. 1 (1785) 595.
P. saccharoideum Beauv., in Ridl., Fl. 5 (1925) 195.
Common in Malaya by rocky streams, growing from rock crevices, on steep earth
banks and other rocky places. Recorded several times from limestone.
Polytrias amaura (Buese) Ktze, Rev. Gen. Pl. (1891) 788; Gillil., Fl. Mal. 3 (1971)
244.
Andropogon amaurus Buese, in Miq., Pl. Jungh. (1854) 360.
Eulalia praemorse (Nees) Stapf. ex Ridl., Fl. 5 (1925) 197.
Rhynchelytrum repens (Willd.) Hubb., Kew Bull. Misc. Inf. (1934) 110; Gillil., Fl.
Mal. 3 (1971) 150.
Saccharum repens Willd., Sp. Pl. 1 (1798) 322.
Tricholaena rosea Nees, in Ridl., Fl. 5 (1925) 235; Henders., J. Mal. Br. R. As.
Soc. 17 (1939) 87.
Setaria palmifolia (Koen.) Stapf., J. Linn. Soc. Bot. 42 (1914) 186; Gillil., Fl. Mal.
3 (1971) 157.
S. plicata sensu Ridl., Fl. 5 (1925) 236, non Cooke.
38 Gard. Bull. Sing. 36(1) (1983)
Stenotaphrum helferi Munro ex Hk.f., F.B.I. 7 (1896) 91; Ridl., Fl. 5 (1925) 220;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 87; Gillil., Fl. Mal. 3 (1971) 205.
Thysanolaena maxima (Roxb.) Ktze. Rev. Gen. Pl. 2 (1891) 794; Gillil., Fl. Mal.
3 (1971) 45.
Agrostis maxima Roxb., Fl. Ind. 1 (1820) 319.
T. agrostis Nees, in Ridl., Fl. 5 (1925) 241.
Dubious Record
Eulalia lanipes Ridl., J.F.M.S. Mus. 7 (1916) 56; id., Fl. 5 (1925) 196; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 86; Gillil., Fl. Mal. 3 (1971) 243.
An endemic species and according to both Ridley and Gilliland l.c., recorded
only from Kedah Peak (not limestone). Henderson however, mentions that it is
known from Langkawi limestone. I have not seen any specimens of this plant from
limestone and believe that Henderson was mistaken.
NOTE
HYDROCHARITACEAE
Hyadrilla verticillata (L.f.) Roy.
Widely distributed in the Old World tropics and the sub-tropics, common in most
permanently wet places. Recorded from Tambun, Perak in a stream underlain by
limestone.
HYPOXIDACEAE
Curculigo latifolia Dryand., in Ait., Hort. Kew. 2 (1811) 253; Hk.f., F.B.1. 6 (1894)
280; Ridl., Fl. 5 (1925) 300; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 81.
LILIACEAE
1. Plant scandent, stem long; leaves reduced to scales and with needle-like branches (cladodes) in
1} ed i ee Tm atin Cri arco Some como mo tao Asparagus racemosus
Plant not so, stem short; leaves large, elliptic to lanceolate, cladodes absent .............. 2
2. Leaves slightly plicate with prominent longitudinally parallel veins, elliptic-lanceolate, to about
30) om LOng;, OFFER TESS i: ocicscc:ceca sacs euaite ouestotgenevareueYeusuere tenets Glos asta eae e eR me RSTO CRC Ree 3
Leave not plicate, veins not so, lanceolate, usually about 60 cm long .................... 4
3. Flowers greenish with shades of purple. Fairly common on limestone ... Peliosanthes lurida
Flowers deep purple. Rare on limestone ........+...........-.+...- Peliosanthes violacea
4. Flowers pedicelled, white, perianth lobes linear-acuminate; inflorescence a raceme or panicle.
Leaves about 60 by 5 cm; petioles not distinct ............... Chlorophytum orchidastrum
Flowers sessile, dark purple, perianth lobes broad and short; inflorescence a spike. Leaves about
60 by 10-15 cm; with a long narrowly winged petiole ................... Tupistra grandis
Limestone hill flora of Malaya IV 39
Asparagus racemosus Willd., Sp. Pl. 2 (1799) 152; Ridl., Fl. 4 (1924) 331; Henders.,
J. Mal. Br. R. As. Soc. 17 (1939) 81.
Plant, scandent. Leaves reduced to scales. Racemes solitary or fascicled,
sometimes branched. Flowers small, 0.2-0.3 cm across.
Distributed in Africa, across to India, SE Asia and southwards to Java and
Australia. A very rare plant in Malaya, and so far known only from a single collec-
tion from Langkawi on the limestone. (Curtis 1674).
Chlorophytum orchidastrum Lindl., Trans. Hort. 6 (1825) 79; Ridl., Fl. 4 (1924)
327.
C. malayense Ridl., in Henders., J. Mal. Br. R. As. Soc. 17 (1939) 81.
Peliosanthes lurida Ridl., J. R. As. Soc. S. Br. 31 (1898) 96; id., Fl. 4 (1924) 325;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 81.
Endemic to Malayan, not uncommon in rocky forest and on limestone, usually
in shaded spots.
Peliosanthes violaceae Wall., Cat. (1831) 5084; Bak., J. Linn. Soc. Bot. 17 (1880)
502; Hk.f., F.B.I. 6 (1894) 266; Ridl., Fl. 4 (1924) 324.
Tupistra grandia Ridl., J. Bot. 38 (1900) 73; id., Fl. 4 (1924) 330; Henders., J. Mal.
Br. R. As. Soc. 17 (1939) 81; Molesworth-Allen, M.N.J. 19 (1966) 303.
Endemic to Malaya and apparently quite rare. Molesworth-Allen reports that the
inflorescence smells strongly of male cats! It has been collected twice from limestone
(in Kedah and Kelantan).
LOWIACEAE
Orchidantha longiflora (Scort.) Ridl., Fl. 4 (1924) 292, nomen tantum; Keng, Gard.
Bull. S. 24 (1969) 347.
Lowia longiflora Scort., Nuov. Giorn. Bot. Ital. 18 (1886) 308.
Orchidantha calcarea Hend., Gard. Bull. S.S. 7 (1933) 125; id., J. Mal. Br. R.
As. Soc. 17 (1939) 80.
Endemic and uncommon; probably abundant locally, as in the forest at the
University of Malaya Field Study Centre (270 m) at Gombak, off the 16th mile
Gombak Road, Selangor. (Sporadic flowering late April and early May, 1972).
Recorded once from limestone (Henderson 26023), Lenggong, Upper Perak).
MARANTACEAE
Stemless. Petioles to more than 100 cm; flower pairs without bracteoles ..................
2) Gtet.0) Ba ty eS Bie be eels + 2 beeen es 28 ASTOR Ste eA ed ee ee Stachyphrynium cylindricum
Stems 200-300 cm. Petioles much shorter; flower pairs subtended by small bracteoles ......
PRPS Bate es te pase Sass oh tch coe Peas otays ous cgen Dok oy ancestal yoityege erect ate ays) cer~ Saree 3 Ets Donax grandis
Donax grandis Ridl., J. Str. Br. R. As. Soc. 32 (1899) 176; id., Fl. 4 (1924) 286;
Holtt., Gard. Bull. S. 13 (1951) 268.
40 Gard. Bull. Sing. 36(1) (1983)
Stachyphrynium cylindricum (Ridl.) Schum., Pflanz. Marant. (1902) 49; Ridl., Fl.
4 (1924) 287; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 80; Holtt., Gard. Bull.
S., 13.1951) 278:
Phrynium cylindricum Ridl., J. Str. Br. R. As. Soc. 32 (1899) 178.
Plants tufted; leaves from the base, at ground level; petioles to 120 cm long, blade
to 60 by 20 cm. Inflorescence on an erect scape from the base of plant, spicate with
2 rows of overlapping bracts.
Endemic to limestone in Malaya, rare; recorded from Kelantan, Perak and
Kedah.
MUSACEAE
Musa malaccensis Ridl., Trans. Linn. Soc.(ser. 2) 3 (1893) 383; id., Fl. 4 (1924) 294.
var. minor Ridl., J. Str. Br. R. As. Soc. 59 (1911) 205; id., Fl. 4 (1924) 294;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 80.
This variety is recorded from limestone and is known only from Perlis; it is the
only Musa collected from limestone.
Cheesman in Kew Bull. 3 (1948) 17-28, has reduced M. malaccensis Ridl.,
together with M. truncata Ridl., and M. zebrina Van Houtte ex Planch. to M.
acuminata Colla. Used in this sense M. acuminata is a very widespread species in
Malesia; it is then very variable, and as yet there are no attempts to subdivide it,
apart from the original consideration of three species. M. malaccensis sensu stricto,
on the other hand, occurs only in the forest of Malacca, Selangor and Perak in
Malaya where it is common. The var. minor Ridl. however, could be specifically
distinct, but this needs more study.
M. acuminata sensu lato is, however, not to be found on precipitous cliffs or on
the pinnacled domes of hills though it has been frequently seen on the bases of hills
in partly shaded localities with accumulation of some soil and debris.
ORCHIDACEAE
An introductory key is provided. The numbers on the right hand side refer to the
Ones in the main key with which one should continue.
In the main key at lead 5 a little difficulty may be experienced in deciding whether
to go on to No. 6, ‘‘plant terrestrial or on rocks; not with a climbing stem’’, or to
No. 38, ‘‘plant epiphytic or with a climbing stem ....’’
Plants with a climbing stem pose no problems, however plants without a distinc-
tive climbing stem may prove difficult, as epiphytes have been frequently found
growing on rocks and roots at ground level, this being especially so on hill tops with
a dense herbaceous and mossy ground flora. However, if such plants are also com-
monly found as epiphytes on the limestone hills and tend to be epiphytic outside the
limestone field according to past records, then they are included under lead No. 38.
Limestone hill flora of Malaya IV 41
There are some plants which are generally regarded as epiphytes or rockplants;
however, if on the limestone they are more frequently found on rocks, then they are
included under lead No. 6. Coelogyne asperata and Coelogyne pandurata form very
good examples in this category.
Finally there are those plants usually epiphytic which are very rare on the
limestone, having been collected only once or twice; and if they are recorded from
the limestone only as growing on rocks and roots at ground level, then, of course,
they have been included under lead No. 6.
INTRODUCTORY KEY
Pe Sastre casini ple iicd ieOr leatletS mee Ween. a oo oss cls ese ovine c.sjcines seieen eee sun vases 2
1. Plant different
2. Plant terrestrial or on rocks; not with a climbing stem
4S. LIP WaTED ALLENG EES Oa cs eee oR NO ec EES Re 6
Paphiopedilum niveum
3. Lip not so
4. Plant with a prominent tuber and erect leafy stem .................00ecceececeees 8
Habenaria
4. No tuber; pseudobulbs may be present
5. Pseudobulbs absent; growth not regularly sympodial ...................02-0000- 12
Eria vestita, Vandopsis gigantea, Thunia
alba, Goodyera hispida, Corymborchis,
Malaxis
5. Pseudobulbs present or not; growth sympodial
G@ One leation cach shoot/of, the sympodium 2c 12's ojeceie seis wei ele elere oes owas 23
6. Two or more leaves on each shoots of the sympodium
TPE SHOOLS LOEMIMNG NOME leALY “SLMS! (2 oo isie = Rayesias Sa Sain sn ey cere ie oe are elelclelc,cuieisie ete 26
Tropidia curculigoides, Dendrobium
salaccense
7. Shoots not forming long leafy stems
SeiteavesvabOUtslnCminwiG ern yiyRreisesr=blesetstove a os © wiers o.ci0 + w,srershalaie mrecacavere SySiataw 28
Eulophia keithii, Cymbidium dayanum
Seu lecavessmoOrent any lnCirpWwIde serie) cele cree a/cie a1= o« ciclciels celal. o\swtereieaie ele(evevere 29
Calanthe, Coelogyne, Spathoglottis har-
dingiana, Geodorum citrinum, Cymbidium
finlaysonianum
2. Plant epiphytic or with a climbing stem
9. Plant monopodial
10. Leaves closely 2-ranked with overlapping bases ............. ee eect eee eee eres 39
10. Plant different
11° Stem short, leaves close, laterally flattened .....-.2.....--.. cece e cece ens 41
11. Plant different
12. Flowers lasting 1 day; producing one or a few at atime .................. 44
Pteroceras, Thrixspermum, Ascochilopsis
myosurus
42
Gard. Bull. Sing. 36(1) (1983)
12. Flowers lasting longer; usually many opening together
1°3 “Teeaves) CEretes <2 sceacccra sc corareore eon ovale tepe cs’ obeuoue cs) ec eretsisy opehar sarah one nla) ie carer 50
13. Leaves not terete
145 Upper sepalimorejthan 2icmplong en -eeeeseeoe ea eee 52
Arachnis flos-aeris, Staurochilus fasciatus
14. Upper sepal 2 cm long or less
15. Flowers with lip at the top; on erect inflorescences ............. 54
Acampe longifolia, Camarotis apiculata,
Pomatocalpa
15. Flowers with lip at the bottom if inflorescence erect
16. Inflorescence producing 1-4 flowers .................22005- 61
Adenoncos, Uncifera tenuicaulis,
Trichoglottis
16. Inflorescence producing more than 4 flowers .............. 67
Aerides odoratum, Renanthera, Phalaenop-
sis, Sarcanthus, Schoenorchis micrantha,
Abdominea_ minimiflora, Pomatocalpa
spicatum, Malleola
9. Plant sympodial
17. Pseudobulbs or stems with 1 leaf
18. Inflorescence from the top of pseudobulbs ............... cece eee cee ee eee 82
Ceratostylis pendula, Dendrobium,
Pholidota pallida, Liparis
18. Inflorescence from the side or base of pseudobulbs or from the rhizome ....... 87
Thecostele alata, Bulbophyllum, Thelasis
17. Pseudobulbs with 2 or more leaves
19. Pseudobulbs with 2 leaves
20. Flowers solitary from the pseudobulbs, or opening singly, or 2-3 from an
inflorescence: "WF RTI SRE oe ate Sana. Sots ee RD Oe ah lei eee cara 98
20. Flowers more than 3’ opening together .......................:0-+seeee 102
19. Pseudobulbs with more than 2 leaves
2). ‘Stemvandileaves laterally flattened imei creel rericiene re erecta 104
Hippeophyllum scortechinii, Oberonia,
Dendrobium
21. Stem and leaves not laterally flattened.
Z2> CAVES HOTS t Sse zee Whe re ar erie Rte Tae a ea rats a aS DS aIeE auc tcitc to OSE OTR 117
Dendrobium setifolium
22. Leaves not terete
23. Stems with a swollen base of 2 internodes, the rest slender ......... 118
Dendrobium planibulbe
23. Stems not so
24. Stems with leaves at apex only, lower part long and bare ....... 120
24. Stems not so
25. Stems over 5 cm long
26. Inflorescence a terminal dense head ................... 123
Agrostophyllum majus
26. Inflorescence different
Limestone hill flora of Malaya IV
10.
11.
12.
13.
14.
27. Lip with 1-2 appendages near its base ..............
Podochilus, Appendicula, Dendrobium
27. Lip with no appendages
28. Inflorescence terminal or subterminal ...........
Eria citrina, Agrostophyllum
Paphiopedilum lowii, Polystachya
flavescens, Phreatia secunda, Thelasis
ORCHIDACEAE - MAIN KEY
PlAanEsacitintaisMelCuIcAITOL WITNOUEMCAVES. 30. ok cccsiaieic ao Saieis npoieveen-« npavsiave erdv ve aseeae
SUPA SmDUI tm OLeIE Any Ones Cals mye ciereh ee hrc ciere cis Caio s Diaven cic ehle oak archos aa eee cee bes
Leaf present, single, heart-shaped to 2 cm long .................... Corybas mucronatus
Leaves absent, or reduced to tiny brown scales; stem very short, roots, green, long and
SPLCAMIN OME PID MYLES wae Peston. FA reiernersie roe eae eee helo eae Srcte sete tele oaks See BT
PECAN TNO T ATLAS SCT ects Peay on hoped pacar oFe fara tarayotod velo re¥ops:s eae eI RR cote ot dhs erated tr
Bracts not in 2 ranks, facing all ways, scape to 3 cm long ..... Taeniophyllum culiciferum
Anther with a short beak; lip not lobed ....................... Taeniophyllum obtusum
Anther with a long beak; lip slightly lobed .................... Taeniophyllum filiforme
Plantsiterrestrial or‘on rocks; nof with a climbing stem ...............-.0..-ccceeeccecs
Plants epiphytic or with a climbing stem, sometimes terrestrial ...................-.000-
Lip with a large pouch, lateral sepals united. From the extreme north. A slipper orchid ...
250.9 OA. ten uiolg ocean) Sao CRO ae Sie Oe ee NS te a er ae Paphiopedilum niveum
Plant with a prominent root-tuber, erect leafy stem and terminal inflorescence ...........
No tuber; but may have a prominent pseudobulb (stem structure, with scale leaves or their
GETS). Seon cbnen peop oo TCOOUO NOD CUD OC OS OCT On Cae eet See SSers erICraS Oem Orn aarrer
Spur of lip shorter than sepals and petals. Only from Langkawi .. Habenaria goodyeroides
Spimetsuprlongerst hanssepalseand. petal Se ieee exec ais tere Gye clove, «0 o:eve.o:~ olcicscueislererw. oresa:s atetoras'arave
Side lobes of lip nearly as broad as long. Only from Langkawi ........ Habenaria carnea
SIGEMODESIOLMUPEVERVAILAET OW, precios cris a Ar iavecs i iets sia voueec la Volnis: wforaiatalaversle alm pyeters afe.c sa ace: oveievernss
Upper edge of lateral sepals 6 mm wide. Only from Perak ............. Habenaria kingii
Wpper edge/of lateral sepals'8-10!mmi wide «<< 2.5520.6.- 0-5-0002 +2502- Habenaria reflexa
Pseudobulbs absent; growth not regularly sympodial ................ 00 cece cence eens
Pseudobulbs present, growth sympodial, or plant with underground rhizome bearing a succes-
SOME EB IeCAVESE ANCE OLESCENCES we ats spars ciara citiae_o cians ic eve ainian Suaieue wie) cfe oe leas ane a eels sisi eoke
LETTE GTP Oo) 2 etek S Se eb COORG ae Be Oe Be Boe a0 6 GOED ES SEO TSO CUTE ee cee arcs
Stems very fleshy, leaves strap-shaped or stems close together, to 60 cm tall, with many thin
OES: AD YT SS Gis Sepa an Obs 50 OEEO DOO E COS UBUCOSCGaS COCR E OO Ce CeO UC eee aOretnet
Stems not fleshy, if fleshy not tall and erect, not with many thin leaves .................
Leaves very fleshy, 35 by6cm. Stem very stout, erect, short or to 30 cm; sepals and petals 2.5-3
cm, fleshy. On rocks from Langkawi .............. Seco ooo pEEOoOr Vandopsis gigantea
Leaves thin 20 by 5 cm, stem fleshy, erect to 60 cm tall; flowers at the end of stems. Only from
DCL ANC DEMMINSH ArH UTALLANG tara ciate essere sro, has sera: «a0 ey nv ovate. cnn aire. ovaj & saoheretolrete Thunia alba
38
1}
10
12
22
13
14
15
44
5)
19.
20.
De
22.
23.
24.
25.
26.
Dili
28.
29.
30.
SHilE
323
33%
34.
Gard. Bull. Sing. 36(1) (1983)
Flowers: with liptatsthe bottom yeti cs « mike eth tereke enckotes oysretere sy ess wiciet crete enalsyoneiciets or otaleuavetete
Flowers*with hip atthe itop ee sersee te oniee segs oeveyaiel «cram teyatevenete aleve ecile Ve eleioney de elercteetenetree
Herbaceous, less than 20 cm tall. Leaves small, flushed with pink ...... Goodyera hispida
Woody; more: thanys0lemitallsWeavesilargerssereemime street tsrerrystelelsrenhrelet-taekeleystetieretetetete
Hepeneitee mire aenynp ye, Uae Mer tes Aer eo ate oR IB EES OC MOOS Oo Corymborchis rhytidocarpa
Sepals and petals 3-5 cm long; column about 3-4 cm long ...... Corymborchis veratrifolia
Lip ‘without: aliricles cast once ne oe eee OEE cee Malaxis latifolia
Lip with auricles: «00:5 oz sgsvojsjgusccre ts, sys eee sesnapoyeaegensls be eked voter eekegeteners ches a1 reuet ve deren
Apex of lip not toothed. Endemic and very rare, only from Perlis ....... Malaxis reniloba
Apex Of lip toothed oe sis cis cccs sie cieiwseteserayerece (olsrevete re fouls epeclorel aise tars, spe eneenaeetie ae Ren ere an
Feeths Only 2) oic.e oats soca wus eras Seats eine wleee Plein eke ee ee Malaxis calophylla
Teeth; 4-6: Once fromiPahane: .22..0eee eee coer een ene Malaxis micrantha
One: leaf onkeach shoot ofiithe sympodiumimn creer crete ian eee eer eee
Leaves) 2) or more) on!each shoot of, the’sympodiumiys:- +e eee eee ene
Bracts. in: 2:ClOS€ Opposite LAMKS) _ya.6:c6,<jcus,eceusus,sveyecereneseseneseveuousloregersisisieleseiekes Cea cee oe Re
Bractsinot in) 2)closejopposite ranks) eee ee eee eee eer eee Liparis caespitosa
Leaf to 35 by 3 cm; lip 1 cm wide. Doubtfully from Kelantan ......... Liparis compressa
Leaf to 20 by 0.8 cm; lip 3.5 mm wide. Common in Pahang and Kelantan ..............
Shoots: forming: long: leafy stemsaraapeeeeenoe asec ane eee Geen Geter eee
Shoots not forming long leafy, stems! tj..-rmctes eters cere ieee ee leeeroienere eerie eee
Inflorescence of more than 2 flowers; leaves 14-22 by 2.5-5 cm .... Tropidia curculigoides
Inflorescence of 2 flowers; leaves narrower; plant grass-like ....... Dendrobium salaccense
Leaves about 1 cm wide. Only from the extreme north .......................40+-ssees
Leaves more: than: 1 ice Wide Seve. pete roe etree ace rete heute tes eee) On ea era ere
Pseudobulbs prominent, about 10 cm long, ovoid, green and shining ..... Eulophia keithii
Pseudobulbs) notobvious | anaeienie rooereeteere orictetten: etcetera Cymbidium dayanum
Lip with a prominent-spur’ Ve Miccek cs cee oe oteiele soe e, aye ee ishe encase te erecta isecne ete ol her one reer
Lip without a prominent spur; spur sometimes not developed at all .....................
Plant flowering. when leafless: <.0.<:..55<acesse-ecavecevn wo onontie pete tener ere eee oTs GRAS ST oT EN ee rat el eet
Plant ‘never Jeafless, ...s2 sce cs cette eae Mar ae ae re oO ae tate Te ere
Flowers white, midlobe of lip as broad as the width of the side lobes ..... Calanthe vestita
Flowers pink, midlobe of lip much narrower than the width of the side lobes ............
Pee One TI Kes tad Noite Lotito D a toncdocto om0oo 3.0ti0 c Calanthe rubens
Spur less than 3 cm long; leaf blade to 50 cm long. The most common species of Calanthe on
limestone... 5: 5:5:0.<40% are NA INR es a ee are Calanthe triplicata
Spur 3-5 cm long; leaf blade about 30 cm long. On limestone, only from Pahang .......
ee Seer ned acc irc.n On nb Oloicohc eo-cci ot Logo POO cern Emcor ee OTL Calanthe ceciliae
Leaves 2. to: each pseudobul be caciracicvorteverieleee tere ercmereke IPO VOR Itech teri rcioveree tore ietae
Reayes' more than) 2itoleach) pseudobul bin aerriei eerie ereie nine arta ete serene aerate eee
Leaves less than 20 cm long. Once from limestone in Selangor ......... Coelogyne pallens
Leaves more than 20 cm long, usually more than 40 cm long .................00eeueeee
17
18
20
21
23
25
24
26
27
28
29
30
33
31
32
34
36
Limestone hill flora of Malaya IV 45
35. Flowers creamy white, with brown markings. From limestone in Kelantan ...............
2 6 Sip Cong AIG ale D.ORNe Otc ae CERTO TO OIEE.O 6 OO CPN cco DLAI ae nea EES ae Coelogyne asperata
Flowers greenish with black markings ....................eeeeeeee Coelogyne pandurata
36. Leaves to about 15 cm long, pseudobulbs, small, rounded ..... Spathogolottis hardingiana
Leaves larger, to 30 cm long and 7-10 cm wide, or longer and narrower ................. 37
37. Leaves to 30 cm long and 7-10 cm wide; pseudobulbs rounded, subterranean. On limestone only
TOM ANC KAWIME Re ie eek Seis aciobtr ie ails wile g alciels Ausiaa.« ssicisinekwleies Geodorum citrinum
Leaves more than 50 cm long, about 4 cm wide; thick and fleshy; pseudobulbs long and com-
pletely hidden by the sheathing leaf bases ................... Cymbidium finlaysonianum
38. Plants monopodial, apical growth of stem continuing indefinitely ....................... 39
Plants sympodial, apical growth of stem not continuing indefinitely, ending in a pseudobulb
MtMnCALCSHOMMIGWELSONIOOLE ee cnn ns ncaa otis oe necro onees asisias tby ties bscee as bine 80
39. Leaves to 25 by 2 cm, closely 2-ranked, with overlapping bases, blades curved outwards; sepals
and petals with crimson blotches on the outside. Once from limestone, Pahang ..........
TI eee Paynes HAA TS Ot er ave loielele arg, wlerars-onpovotas o 8 sje ave, aelayeo4 Dipodium pictum
PIA GRC Hood onaSeie Ss Naa er Gunes DO S6 COST D OCT OREO Oren ce eet: 40
40. Stem short, leaves close, fleshy and laterally flattened; flowers small, in pairs ............ 41
ELEGY DOVES) cingic'cln6 Bab 08 on OO OG Oa BC 8 OB or ID OS Cart Oa eae eee rer ae 43
41. Stem with leaves about 1 cm across; from Selangor and Pahang .. Microsaccus brevifolius
SCITIMMIENEAVES Hl es) CI OFsMOLesACLOSS ia (enyefeas «fo sees a overseer eys SIO «aie sce a aes woes o epernse 42
42. Lip much narrower than spur, petals wider than sepals; on limestone from Pahang .......
DoD OC an oO Bg OBOL DADC 6 OCRICOICTE CO OTTCTICCeInC Ciao ae ier same Microsaccus ampullaceus
Lip not much narrower than spur, petals about as wide as sepals; common on Bukit Takun,
SQELEO? Cg ode douse Sop omos DOR On OMe 26 be Sereno a Rar eta Microsaccus javensis
43. Flowers short-lived, lasting one day, produced one or few at a time at the end of the in-
EIQECSCELICC METRE PI rhe ter ole iene tie, cloleret eset iis cieim savers! > alaa e stelarsrereus, eigioweonmiliyedeeraye: arse ve cians 44
Flowers lasting longer, usually many opening together ..................0202 000 eee eeee 49
44. Lip loosely hinged, movable, distinct from the column foot .................00e ee ee eee 45
Lip not so; immovable, not distinct from the column foot ...............-.-eeecereeees 47
45. Inflorescence-scape glabrous, to 10 cm. On limestone in Kelantan and Pahang ...........
9 6 COCAB OG HS SOD COSC ACS tar TLCS S DTERCOTS IC ITER ee ei nme Pteroceras hirsutum
Mutlorescence-scape minutely prickly, 2-4 cm long ... 2... 22. nce ere cile one cles wis wisi ee 46
46. Sepals and petals spotted crimson, sepals prominently hairy ......... Pteroceras hirsutum
Sepalsrandupetalsinot spotted, not hairy .........---meei dei e+ rvs! pie Pteroceras ciliatum
47. Bracts 2-ranked, alternating; rachis to 10 cm. On limestone only from Perlis .............
S'S ASIGUG 0 Cress CRO CRC GIDC CO EL EITO Se OCI OTIS CROCE DECODER CEE RTCaE Thrixspermum amplexicaule
SLAGISMUO bye -Faniked rachis SHOILET, eon ..cees.c pore acs G ae visre ¢ slnaic 6) e eee, sveeiaia stems aise sine ajelehels 48
48. Scapes to 15 cm long, bracts 3-4 mm long, acute ................. Thrixspermum album
Scapes 2-5 cm long, bracts minute, on a thick rachis bearing very small flowers ..........
3 dd A oe Re Te Oe ee Teena Fete otro ates eA SCOCHIODSISHINYOSULUS
49 MeIEAVESICILCHIA GIN AtrANSVELSE.SECEION pie lasik eich «eb tllor« Clave fale ctavaveleve susle0ot alatciaqorat aleTatene ailslla site. 50
eaves not circularinitransVverse SCION sOOL tETeLe 1s syos)< sie sice aiel ste vlc chelele w) aleiclsio eel alele ete 51
50. Leaves grooved; midlobe of lip 6 mm wide at the base ........... Sarcanthus machadonis
Leaves not grooved; midlobe of lip narrower. Only from the extreme north ..............
SLOTS SO SOC eee OE ot 0 nic, Gon teprmacen rae! el Chace Sarcanthus sacculatus
51. Upper sepal more than 2 cm long; plant with a long climbing stem ...................-. 52
Wines agua om lGne Oven addeacco spn codcdospseopntne Ged cUS Uso OU ton coo ae Goma 53
52. Lip loosely hinged, freely movable; leaves to 17 by 5 cm. The scorpion orchid ...........
Meets APTS A LAST IEE s SENG kesS leo VOR LaU SEE aN nga lnies She. gh sm nejayaesina card QUA ee altuna Arachnis flos-aeris
Lip not so movable; leaves to 12 by 2.5 cm. Only from the extreme north ...............
JSGb0db BESO TOU EOS von DE ObE DOD ORD OEND oUt oe OCDE eC eee Tats Staurochilus fasciatus
46
53.
54.
55:
56.
Sil
58.
2)
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
Tle
2:
Gard. Bull. Sing. 36(1) (1983)
Flowers with the lip at the top, on erect inflorescences ..................e eee eeeeeeeee
Flowers with the lip at the bottom, if inflorescence is erect .................000-es eee
Stem stout, often branching, leaves thick and stiffly ascending to 20 by 3.5 cm. Inflorescence-
scape stout. Lip of flower not spurred. Only from Langkawi .......... Acampe longifolia
Plant different; lip) deeplyzsaccatevorsspurredi mec cre cere terete eee tee eee
Spur with a 2-lobed callus attached below the midlobe; rostellum with a long beak; leaves to
CO Fal oh ia) IS o1 Winch Per, Sirens one renee neem ner eeeee eae eR ROTO cae OOM ree PO ic Camarotis apiculata
Spur without calluses; back wall of spur with an erect tongue ....................00000-
Stem short;internodesiunderslaicmblongs rian ease eee roe een neta
Stem long-climbing,) internodes!2-sacmplongeer pacts eit etter reer tee eae
Flowers pinkish, inflorescence minutely hairy .................... Pamatocalpa kunstleri
Plowersiyellowishspinflorescence: notnalnyaeceicieeeieieieireterietter etait eee
Sepalsiandiipetalsawithsaredimarcingee er ree rere err Pomatocalpa latifolium
Sepals and petals without a red margin. From limestone of the extreme north, in peninsular
WALANG: ioe. sers odin eye xe ens axa wteceuer oye d nse uence re eae eee Pomatocalpa setulense
Inflorescence: producing I=4° flowers: c.ctrcs co oicpore cr tones erie oe eke eae ea
Inflorescence producing more! than’ 4 flowers. sees ace ae etter aeteeie sree ate sere eerie ene
Lip shallowly saccate at the base, papillose or with a callus .................0.00e0e0e0
Lip deeply saccate or spurred, entrance to the spur often hairy ...................00000-
Rachis of inflorescence zigzag, to 8 mm long, bearing 3-4 flowers .. Adenoncos sumatrana
Rachis of inflorescence not so, very short, bearing 1-2 flowers ...................0.008-
Leaves to 7 by 1 cm; slightly constricted near the acute tip; upper sepal to 5 mm long
GOO GIG ORSTOCOOOHEKTOOISDOMEDO OD GOOFS THHOOCUDNO es OSSAHDOOOOONOES Adenoncos major
Leaves and flowers smaller. Only from Selangor, at Gua Batu ...... Adenoncos parviflora
Lip, upper half of midlobe turned up, the tip with 2 small lobes...... Uncifera tenuicaulis
Kip; midlobelsimpleror si lobed: mever2-1o bed ir rererseraeeeierersreieiensi iste ricte celts events
Midlobe:of lip3-lobedta az .ossccncciaacnamec cee tee ei lee Trichogolottis winkleri
Midlobe-of lip’ simples 4.082.< occas ceegs te een See ee AOR eee
Midlobeihainy.s9)mmiplones Commoners scence arena Trichoglottis retusa
Midlobe not hairy, 5 mm long. Only in the very North ............. Trichoglottis misera
Spur turned up in front like a horn, flowers with purple markings, upper sepal about 1.2 cm
LON: ev. d esos. Eis. nravelai ih doneseranieas te vers Neca Mee tei ene Poe OR ee Se ee Aerides odoratum
Spursifepresentnotsso;susuallyssaccatemen eee teen ceit cine reiterate enn
Flowers redvor yellowiswithoredispotaes a mrten torre. ae ate eee ere nee ey
Flowers*notisOuGolOured! F536. coze8 occa apo aac tetas roe wae ean OR oars sree
Column slender, curved; leaves fleshy, stiff and sharply pointed ... Renanthera histrionica
Column short; leaves bilobed at the tip. Only once from limestone, in Kedah ............
a Oe Ror LO TEs OD OO Goo oO O OO re ooo MoDUoL ae Renanthera elongata
Lip not spurred, sometimes slightly saccate, with a forked appendage ...................
Lip)spurrediior saceate, with nosforked| appendages: jase cei areata aeiaeerenieiens
Flowers alternate, 2 ranked; rachis of inflorescence flattened; upper sepal 1.6-2 cm long. .
5snBe dicen (allave. 5 caa/folin 95hcRw) wile oy enops Cay Son een ee ene CARRE REECE PETE CORRE IG Renn ee Phalaenopsis cornu-cervi
Flowers not so, facing all ways; rachis not flattened; upper sepal 6.5-8.5 mm long ........
= seal ops semetion)ee\ cio rua sifev' (ors [stosan's: 0h seh cava for G1 oN eT Toe Roe Pe RETR ET Re RERERReR eoae Phalaenopsis decumbens
Callus present, closing the entrance to the spur of lip. ...............eeeeceeeeeeeeueees
No such callus
55
56
57
59
58
63
65
66
68
69
70
71
72
73
76
Limestone hill flora of Malaya IV 47
11h
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Stemmtos0;cm On more-long: leaves wellspaced!*. 20. fn ncwaeee aa vale s teeslee. ces cee cei. 74
Sie CHASHOLtensleAVeSiClOSEstOGEENEN vayer a: cxaycterereiaratarateren chet afeict anarai state svepsierate: -aierw: sl atsi ere alie'oy' 75
Leaves to 30 by 1.5 cm, fleshy, constricted at about 2.5 cm from the tip ................
5 OAS S03 OOS AE ci be Ate nts Bree bee a kek tod Bid tS ee Sarcanthus subulatus
Inflorescence pendulous, to 25 cm long; stem to 6 cm long, stout. Only from Perak ......
ea TE ey atlas ate aCarale ler atetaPeve sca resaveliave tc Weliefers 'e"sle win Yela'a'ele la ele e's eels ele Sarcanthus termissus
Inflorescence short to 1.5 cm long; stem to 15 cm long, slender. Only from Kelantan .....
| ooo en eos GO OB BR UIO DR CSREES OIE DOT OR OEE ORCC Cece tea onan Sarcanthus rugulosus
Hiei OLESCERCEFELECHAOL, HOLIZONLALL sep-tav-otetct fetal cyoreiersietetersioistaleletels)claielcletvisiete a.aluineine ene ees nate Wy
ONES cence GLOODINELOTEPENGU IOUS we cry seteaeyeiclete|escveuscicyoscs<lonsaci=ke oh ovaisiene eysssi cle: <tetersasys sueieccis) Ss 78
Leaves 5 mm wide or less. Stern to 15 cm long, much branched, tufted. Only from Gua Batu
CoO COO OCS ODOT OIOO TIO CIID CE IOI CORTES cca ena eae Schoenorchis micrantha
Leaves more than 1 cm wide. Stem very short .................. Abdominea minimiflora
Stem very short, leaves leathery to 18 by 4 cm; a tongue present on the back wall of the spur
PT CUTIER: SINE TS CTSNET a5 c5 cr gah ore) oyet schol cia) at sieve dheyat ovary abevat'e eq pleleinlone atats Pomatocapla spicatum
Stem long, climbing, 10-25 cm; no such tongue in the spur .................2 eee eeeee 79
Stem and leaf sheaths flattened, leaves green. Only from Perak ........ Malleola undulata
Stem and leaf sheaths not flattened; leaves usually flushed with purple ..................
BACHAO DU DSSOTEStCINS iWILMOlUleatians eevee tte ets alc Perens cle chslaers mnsitale eelcle ish Mae edaeleatniets 81
SENG DSEOLISLEMIS AW IEG 2TICAV ES) fire coued caste cal ois ta oue%ersqnyore'usyenyavey senses oe beya/oFoyssereuene cep creyere pou 97
SENG ODI OSEOleSeEMSiwithemOLe tare? slEAVOS a cteyaictereiclepekckeveisiese feeiaieiei-ieierelsiey evekeloneyesebocT> 103
rin OLescenceMLGMutnentOplOfnPSEUGODUI Die tetas a crsyers ss ys.2ysysienee las cicsatescho sae isisnbasls ans leis susya,e/sterctc 82
Inflorescence from the base or side of pseudobulb, or from the rhizome between pseudobulbs 87
Blawersml -severalcOn SHOT NNGIVIGUALIStAIKS oo 4. oees cposeos0rs oyn{eycio/ein © oysteies sie) sye aie cel sicle of 83
Flowers 1-numerous on an inflorescence with a distinct peduncle ....................0.. 85
Stems with pseudobulbs distinctly fleshy, not entirely covered by leaf sheaths ............ 84
Stems slender not bulbous, covered by leaf sheaths ................. Ceratostylis pendula
Blades of midlobe abruptly widened, whole lip purple-spotted ..... Dendrobium luxurians
Blade of midlobe gradually widened, side lobes and base of midlobe with purple spots ....
3 RPO OO bo Con O ) GOCE IG Oe a OOO oD CCI CN: ae Sener eet et einer Dendrobium plicatile
Bracts in 2 close opposite rows; inflorescence with 1-3 flowers opening together .......... 86
Bracts different; inflorescence with numerous flowers opening together .. Pholidota pallida
Leaf to 35 by 3 cm; lip 1 cm wide. Once doubtfully recorded from limestone ............
PORN Mette eee Fon isi Te dhs toh 201 soos: ov sb oss) so) wich, oles Susisi eleveses a. eusiase iecsle Liparis compressa
MeAtstOPZOLD YAO. SiCme lips -SeMM WIGE) eeye om gree 2) + aperb poco epsceicasyw chorus ofeas Liparis gibbosa
Lip joined at its base to an outgrowth from the column to the column-foot; inflorescence pen-
dulous with»many flowers, Leaf stalk grooved. 2.55.6... 00. saear once Thecostele alata
MNP ATI OE SO) reve A cayeccrn nyt ieee ce MRS Tere RTA Te rere are CR Le cava rel or SUONST OTE ar cia inicogutaraga voy cheievale: al onnlenere oe 88
ENO WCLSESOLIPAGN Ai Parare tte e Oise t Fore ba vey cia vi cov Sue icvael fore, v's e esaefovaioieioile 0 et S cure a1Suayay sieve) aneve veya ops a sreyegsy ate 89
BIO WEESHALNeAS Lean DEI IIIT OTESCEMCE vee reya.cicyeiere ci visisieke.slejeveletovere averers) ete epevevelajars aisle reyes svsya © «se 91
RAIZOMeRS ION Ge aMUCHHDranCHedimee aya cits, male oho ecaiere alice vaisioyorevote ya aise’ a ave) cers serene sisreisve iets 90
Rhizome different, pseudobulbs to 2 cm apart, 1.5 cm long; leaf to 16 by 2.5cm ........
mid ber Oigliaoe ShOCIDO EPEC DIDAGICEOIDINTTI TE EOI TIT ERCICE CIO EIR OI OTe Bulbophyllum pulchellum
Leaves very fleshy, usually 2 by 0.7 cm, pseudobulbs 3 mm long, appressed to rhizome ...
BYP ROPSPCN TNR R TER str t ctietek taVotict ctotelrod otct ciclo "afatar tatalis Sete) otattehatas els tateletere’ ohateratshenateere Bulbophyllum sessile
Leaves thin, usually 3.5 by 2 cm, pseudobulbs 1 cm long .. Bulbophyllum membranaceum
48
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110.
Gard. Bull. Sing. 36(1) (1983)
Pseudobulbs hardly noticeable, leaves to 23 by 3.5 cm with a 6-cm stalk ................
COUT OL OC OCS COD Coan OT Ob ORoC oD DECC. oHobcocoaaae Bulbophyllum apodum
Pseudobulbs'distinct; smalliomlargema-ss sacri oe eee EOL 92
Pseudobulbs large to 6 cm long, ovoid, fleshy; inflorescence to 15 cm long ..............
a emo ot b.cn coo On un atr sca coomocoomenCedGon Bulbophyllum lilacinum
Pseudobulbsisinallene........jerecsaecso esas eee oe eee Eee 93
Leaves almost terete, to 15 cm long and 1 cm wide, rarely with a second shorter leaf .....
OO era cee COCO TORRID Coe dod T soon te rao oe Thelasis succosa
Leaves: different). c 4..0d2%.scalepentoncecrtsintts, 0 Som ecieer sees Semele, 3 \oe ee eerie: 94
Pseudobulbs broader than tall, oblong, with a terminal leaf ............. Thelasis triptera
Pseudobulbs not SO. ss... é sjecece seca ce vasare oye. cereus 020,018) 000 000g SORTER lo aa eee 95
Leaves small, about 4 by 1.3 cm, narrowed abruptly at the base and apex; rhizome slender and
long creeping 2.24%: <astectAckefnrbetarcte chSeise naka elke ex Pee Bulbophyllum concinnum
Leaves: larger,over 10cm long. t5..3 ce ssiaciccss ne eters eterna reel oe eT eee 96
Scape to 2.5 cm long, covered by sheaths, whole inflorescence 5-12 cm long .............
PT oa ora tec dan BO OSOn NOG ae ac ob Ao OAtS. Ob c Bulbophyllum flammuliferum
Scapeitorl2iemylongy dullipurples .5-4.4- see ceioeee eee Bulbophyllum fenestratum
Flowers solitary, from the pseudobulbs, or opening singly on a slowly elongating inflorescence,
or 2-3,.on'a single inflorescence» 3 «cn1.01./22eieh o(snivicl ate: cents e nel ae else he eee eee 98
Flowers more than 3 opening together on the same inflorescence .................-0005: 102
Flowers solitary from the stem below the leaves ................... Dendrobium spurium
Flowers or inflorescence from the apex of pseudobulbs ................000.0 cece ce eueee 99
Inflorescence slowly elongating, one flower opening at a time; short hairy .. Eria pulchella
Inflorescence: not ‘slowly elongating 2.7.45 ccs cteemiaer cc tenie tors arerarecn eve orn clerer sista lekev ati eneeere 100
Inflorescence with no distinct peduncle, flowers solitary ........... Dendrobium pumilum
Inflorescence with a distinct peduncle, woolly hairy, with 1-3 flowers ................... 101
Leaves fleshy, terete, slightly flat on one side; young shoots white-hairy ..... Eria pannea
Leaves elliptic-oblong; young shoots brown-hairy ..................-008- Eria leiophylla
Pseudobulbs to 12 cm long, ribbed; leaves to 30 by 4.5cm ..... Coelogyne foerstermannii
Pseudobulbs narrowly cylindrical, to 20 cm long, ribbed; leaves to 30 by ll cm ..........
a leiavpre, ene stayareihere tepals 0 oan Metabo Ghee pale eit re etaittor. aire Re toke see eters Coelogyne rochussenii
. ‘Stems’and leaves: laterally"flattened™ <icoccmcevaec sss ceveiccaieicaseas Grinch erro ene 104
Stems and leaves not laterally flattened; leaves sometimes circular in transverse section .... 117
. Inflorescence elongate, of many small flowers many opening together. ................+-- 105
Inflorescence'short, bearing 1-2" flowers at aitime ss eee ce ote nie ence cee aici ieee eee 112
Stems about 4 cm apart, leaves to 20 cm long and! ] cm wide ~- 2... 0 <n e....- sees
aa. 6: ahd’ bs wins Herbie ce roy choca te tepene UGS ane AVENEL ST oh sysitece je, aoe ahe overs) cusiezeeesancuey exe Hippeophyllum scortechinii
Stemsiclose together, forming tufts) <.ccececeie cere error cre ietete = raieret berths teiencie ere kisiee ner 106
. Leaves jointed ‘atithe base. <2 sesecsccusis ere susie ove o.s olehegeietayede rieseust overenes otorevefeasyeisKeaeie Tee rere 107
Léaves ‘not Jointed’ at: the base. s...0.. 0. scott Sacaenewasrosjetns cites caste ieTeeteie citer costo acl Oe eT eee 110
Rms 10 0) 9 ee ee Son er eR SEO ods OOO LOD LOO OF OTE COGS Oe oS 108
Stem 2:5) cmior more: LOMR? Siero. /ote re erent orese oye.creseie tw eves ogetey 1 eeeve topsloncPOL RS Se RCPS CREE RCI 109
Leaves to6cm long: ‘Only from! Mangkawili i -retteieiier tetetieletatsisieyareiei Oberonia calcicola
eaves) to Sicmior morevlongarcrem cece rene nee iste tr rier Oberonia dissitiflora
Midlobe. of lipsbilobed.ws:<)..c4 osha eect ees seer eae Oberonia flava
Midlobe of lop transversely oblong 35.22. .5...2260-0 4++ eee Oberonia transversiloba
Stem: 10 \emor MOTE ONG, s5:6ye:0;5 015-01 ereeseceterey see penederorerey suelo elkeLeVdechevsis eastexe rere Loses a evan 111
Stem! 3=5cmilonginflorescence’5 cmulong, = miseries sci arene telat Oberonia caudata
Limestone hill flora of Malaya IV 49
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Flowers on distinct pedicels; not completely covering rachis ......... Oberonia spathulata
Flowers lying on rachis, completely covering it ....................000. Oberonia anceps
Leaves very close together, bases overlapping. Once from limestone, Perlis ..............
I ers ee ee Settee ate Pods Mea laeaeals Sees bb heves Dendrobium excavatum
Leaves distinctly spaced; only sheathing parts overlapping .....................2..000-. 113
Meaves atiease SuInmMEapatt Stemithin, TIEXUOUS) <2. .cf2 2 sence cece cece occieccveteenset s = 114
MecAvespalraneeaicloser LOLEWIET; SEEMS |SEMTLCL fetes creiseccherte selec saree.6 ols wists wie aoe quale cetjenore 115
Leaves 3-5 cm long; flowers about 1.2 cm long .................. Dendrobium acerosum
Leaves under 2 cm long; flowers about 8 mm long .............. Dendrobium subulatum
Stem with the apical part devoid of normal leaves, floriferous .... Dendrobium aloifolium
PCIE AVAIL OUPTIOU Darr tiats, ctetere ac fepa occ te feasts .e.oc, yay s/o eae, 5 ahs w. ens wo alee wianeiarsvaie'si sidiclars ss 116
Stem with leaves to 2 cm wide; a large bilobed callus present at the base of the lip .......
PER ree act Ste ay ssc} cvs eiegsiels ats ahaha ee Bi cia s'tise wuapshays.ay2,aye's, Cae Dendrobium indivisum
Stem with leaves to 3.5 cm wide; a small unlobed callus present on the lip ...............
26 o bbe CORSE SSB OECD EGER AGNES RiGe TOOLS tte iE ern ee Dendrobium leonis
Leaves round in transverse section; stem to 40 cm long, very slender; flowers white .......
pong aca 20 ACOSO Mot D6 BODIE EGE Beene ae ee eee Dendrobium setifolium
IRCAL ESENGLALOUNGEANESECLION Marte aa ore oe ee arte Peete tec olga oa Dene 6 abe wlndlowae decals 118
Stem with a swollen base of 2 internodes, the rest slender ........ Dendrobium planibulbe
EC HINMO NSO MN sean ante a Aen a Ata rs 3 cn Sas se ste we See he ee fees Drees 119
Stem elongate, with a few leaves near the apex only (old stems may have lost all their leaves),
PandispaLmlOnecrathall the eaLyiPalts bares jercte a. oo cte eeidie o, dione nea he eles Sates Oe wee wees ae et 120
BETH OLESO MP Ey Toe a eroeiaa ay aaa av ave Ad vac ale els aware Rattan tee eee os is 122
Stem covered by red brown sheaths; flowers solitary ...................00-- Eria nutans
BrcinmMOmsorcOvered: flowers in! inflorescences... > .(2) 22 co he Secs he dee te acces tenes 121
Pseudobulbs strongly 4-angled in the upper part; inflorescence drooping to 20 cm long ....
PNM ON Far 57 fo¥.8 reve Yotetars ie tare tons a fa teviotove tole to so 10% 'nrevaie'a-erd oeetotate Dendrobium farmeri
Pseudobulbs not angled; inflorescence shorter ................... Dendrobium callibotrys
SCIMSLOVEL- 5 CIP lONP sa With SEVeLal COMMAMNYICAVES) «.<.c50.c css <seee wie aciew viciie nd oases qevac 123
Stems under 5 cm long, with a few leaves, the bases of which often cover the stem completely 138
Inflorescence a terminal dense head, 2.5-3 cm in diameter, hardly stalked ...............
2.5.0 10:0 6 BIS COREA ORO Oto Bae in Ori Sne Orin Seta Agrostophyllum majus
MLAMOENEST CS GTM gta s wi ONS B55 ODS SEI Ero 8 CHA TIE SOGo TESS O et OC OrrSe aera 124
IDEN LIB -2ea DPenGAages) MeATaILS DAS apie te reels erence ayorerel Siare isinia less ial eioiis «mee ae. saiciave 125
MIPRWItHENOrapPEHGAGES\e ere er. She Maes Peta d PRT a oe eee Reel Cowen sueeesaeees 133
BIO WersrsmidllemunGehal, Cir AGkOSS seine Aes) <P ween nic Deh Se kiew este da Seat clon ehlarheleels'er 126
FR OMCESBIELE CopILLOLG aM MINCMLACKOSS) stecssssave oisicicievorcyec Vac i) sresaysge Ovens eels: yioveveve « Syar aye aleve whys 132
Meavesmlessmnanwled Cm slONe =p POM imas Alc -.- seye: oterrs Roe os oys,ov cued Sieyd = tefere ia leat sevouer oimantels, c/Syete cloves ars 127
ILGAVES THORS TEMES ETI OTES fo ITE iio? i SSB Be a paes decon comer cero nn acer nemn dene 129
MeAVeSMOB Tn Widessacutel Seay ok cc focrreee i. «oes hes Nek «6 Seie%is 2 Cees Seale cit BINS 128
REAM ESEWIGET a CNUSELOUNGEG! ceicktsisic cisieiciciereicie.c # a-06 2% Gis s srarcievoiiatea Podochilus lucescens
Leaves about 6 by 1 mm, at a very acute angle to the stem; lip with 2 narrow appendages
AO yO NO NTE OC aC ee Reh PROCITE CECI COOTER RCE teen ane Podochilus tenuis
Leaves about 8 by 2 mm, spreading; lip with a simple broad appendage .................
= pelo 2 NS ES Oo DO Gy 5 oR © CITGO cRE RIS EARNERS O PRCIS Ie oer ieee cena Podochilus microphyllus
Floral bracts leaf like, closely overlapping, the inflorescence terminal ... Appendicula torta
BlOralbractseGinerentme irre. A. a.c tee See ea ccintale clavate titel sleyate/eioabe Moses cue lslelestele cues Oe tee 130
Stemandjleafssheaths) strongly flattened::.; i< sac. soja. foci) fue «fe Appendicula anceps
Srtembanlcsleatesheatnsenoteilatteme cn. titnce ea Secs ates, spasvenentper spore ovens) cbepncaxctamn yd voretetane ska 131
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141.
Gard. Bull. Sing. 36(1) (1983)
Inflorescence lateral, slender, the scape as long or longer than the leaves ................
asta ery ofaNaSareYSNGLEL GW asi idiaqevayeel Se (a. @yRlicteage Mums Us we SHOE STSREI TET Hie ee ees Appendicula undulata
Inflorescence usually terminal, shorter than the leaves .............. Appendicula cornuta
Flowers numerous and densely arranged on one side of the inflorescence. Stems, 50 cm or more
LONG. 220.6 Few cee a cawlonn Gerke ees Se ee ee ee eee Dendrobium secundum
Flowers different, apparently solitary. Stems to 30 cm long. Only from Langkawi ........
a iat amataciaxhianie spa ondcve she aveoueneraherseatcter ore eu ate tanoreretaiezsversie tetera tere retens Dendrobium langkawiensis
Inflorescence}terminallionsub-terminallasce eee Cee eee reece eee EEE rer arncrn 134
Inflorescence ‘lateral = airocsssa ssccevory onties Cees Vetere, areies roy ToLet TORTOER eh Te Reena 136
Inflorescence!short;s1-2) cm withicloser2 ranked: bractSmr eee eee errr een ener 135
Inflorescence to 20 cm long. Only from Kelantan .......................-5- Eria citrina
Leaf sheaths with slender appendages on either side, at the base of the blade ............
To ae roe OT Eas IO OS Gals ABS OS CUO GOO dine Agrostophyllum bicuspidatum
Leaf sheaths without such appendages .....................-.. Agrostophyllum hasseltii
Inflorescence slender, 1 cm long, with 3-4 flowers, each less than 5 mm across ...........
a saan tneil' an anche evarat san edaee ou ateyetscays ener eT: Caen Tete Ic eRe one Pe eke Poaephyllum pauciflorum
Inflorescence! different;) flowers) larger, about. les! Cmracross) ei eesti eee 137
Stem erect, leaves thin, to 20 by 2 cm; inflorescence of 4-6 flowers ....... Eria leptocarpa
Stem pendulous, leaves thick, to 12 by 1 cm; inflorescence usually of 1 flower Eria rigida
Flowers' small; sepalsitoyabout;4ommi longi cere ciseireeiernisictercnsie cis cictcreiaicrie arene nee 139
Flowers much larger, lip) pouch=shaped) j.5- eee cniieeieeie eis ae Paphiopedilum lowii
Column-foot- well developed) 22.c.45.5 sins seen eels oe SOE IAS ae eee eee 140
Column-foot not developed: ..5:5 0.5 sysi0:6,0 ssjparnaiesoiery ney ues epee cxrareia accent eater en nee 141
Leaves 3-4; to 20 by 3 cm or smaller; scapes about 5 cm long or more Polystachya flavescens
Leaves to 9; to 5 cm by 4 mm; scapes to 2 cm long ................... Phreatia secunda
Rachis of inflorescence to 15 cm long. Stem tufted bearing about 5 leaves, the lower leaves
muchishorterithanithewuppern ....1ac. cee eerie eek Thelasis carinata
Rachis of inflorescence about 1.5 cm long; plant like the above but smaller ..............
se ays ce atte es aca cw Sates Moparre ageNa Pate tam odiane ote oMey orale Toren iohete aktene cP ohcic eee eRe renee Thelasis micrantha
Abdominea minimiflora (Hk.f.) J.J.S., Bull. Btzg (ser. 2) 25 (1917) 98. Carr, Gard.
Bull. S.S. 5 (1929) 20. Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75. Holtt.,
Fl. Mal. 1 (1964) 640.
Saccolabium minimiflorum Hk.f., in Ridl., Fl. 4 (1924) 166.
A small plant, leaves to 5.5 by 1.7 cm, elliptic. Flowers reddish brown.
This us a rare plant in Malaya, which is also found in Java; it has been collected
mainly from the limestone hills usually as epiphytes but also on rocks.
Acampe longifolia Lindl., Foi. Orch., Acampe (1853) 1; Holtt., Fl. Mal. 1 (1964)
625.
Saccolabium longifolium Hk.f., F.B.I. 6 (1980) 62.
Acampe penangiana Ridl., Fl. 4 (1924) 155.
Adenoncos major Ridl., J. Linn. Soc. 32 (1896) 350; id., Fl. 4 (1924) 154. Henders.,
J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 597.
A species found in many lowland localities. It has been collected a number of
times over limestone as epiphytes, and possibly once on rocks.
Limestone hill flora of Malaya IV 51
Adenoncos parviflora Rild., J. Linn. Soc. 32 (1896) 350; id., Fl. 4 (1924) 154;
Holtt., Fl. Mal. 1 (1964) 598.
Recorded in Malaya from Perak and Selangor, uncommon. On limestone in
Selangor and also from Telok Forest Reserve (freshwater swamp).
Adenoncos sumatrana J.J.S., Bull. Dep. Ag. 22 (1909) 44; Holtt., Fl. Mal. 1 (1964)
596.
A. virens Bl., in Ridl., Fl. 4 (1924) 154; Henders., J. Mal. Br. R. As. Soc. 17
(1939) 75.
In Malaya, found in many localities from the lowlands up to 1300 m. It is a com-
mon epiphyte on the dry summit of Bukit Takun in Selangor.
Aerides odoratum Lour., Fl. Cochinch. 2 (1790) 525; Ridl., Fl. 4 (1924) 182; Carr,
J. Mal. Br. R. As. Soc. 6 (1928) 65; Holtt., Fl. Mal. 1 (1964) 698.
Agrostophyllum bicuspidatum J.J.S., Ic. Bog. 2 (1903) 55; Ridl., Fl. 4 (1924) 108;
Holtt., Fl. Mal. 1 (1964) 487.
Agrostophyllum hasseltii (B1.) J.J.S., Ic. Bog. 2 (1903) 55. Ridl., Fl. 4 (1924) 108;
Holtt., Fl. Mal. 1 (1964) 488.
Agrostophyllum majus Hk.f., F.B.I. 5 (1890) 824; Ridl., Fl. 4 (1924) 107; Holtt.,
Fl. Mal. 1 (1964) 489.
Appendicula anceps Bl., Bijdr. (1825) 299; Ridl., Fl. 4 (1924) 195; Holtt., Fl. Mal.
1 (1964) 50S.
Appendicula cornuta Bl., Bijdr. (1825) 302; Ridl., Fl. 4 (1924) 196; Holtt., Fl. Mal.
(1964) 505.
Common in Malaya in the lowlands. However, it has been collected from the
limestone only from Bukit Takun in Selangor, usually as epiphytes but also on
rocks.
Appendicula torta Bl., Bijdr. (1825) 303; Ridl., Fl. 4 (1924) 198; Holtt., Fl. Mal.
1 (1964) 50S.
Stems to 25 cm long, leaves to 1.6 by 0.5 cm; nearly at right angles to the stem,
oblong-elliptic, tips rounded. Inflorescence terminal, bracts leaf-like and closely
overlapping.
Distributed in Java, Sumatra and Borneo. In Malaya, chiefly found as epiphytes
on limestone hills; also on Taiping Hills (not limestone).
Appendicula undulata Bl., Bijdr. (1825) 301; Holtt., Fl. Mal. 1 (1964) 505.
A. purpurascens De Vr., in Ridl., Fl. 4 (1924) 195; Henders., J. Mal. Br. R.
As. Soc. 17 (1939) 75.
52 Gard. Bull. Sing. 36(1) (1983)
Arachnis flos-aeris (L.) Rchb.f., Bot. Cent. 28 (1886) 343; Henders., J. Mal. Br. R.
As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 616.
A. moschifera Bl., in Ridl., Fl. 4 (1924) 159.
Stems long and stout, climbing; leaves to 17 by 5 cm; apex bilobed. Inflorescence
large, over 100 cm long; flowers about 10 cm across; with broad dark purple brown
streaks.
This scorpion orchid is distributed in Java, Sumatra and Borneo. In Malaya, it
is chiefly found on the limestone, growing from organic debris and climbing over
rocks and on trees stems.
Ascochilopsis myosurus (Ridl.) Carr, Gard. Bull. S.S. 5 (1929) 21; Holtt., Fl. Mal.
1 (1964) 707.
Saccolobium myosurus Ridl., J. Str. Br. R. As. Soc. 39 (1903) 84; id., Fl. 4 (1924)
M73:
Bulbophyllum apodum Hk.f., F.B.I. 5 (1890) 766; Ridl., Fl. 4 (1924) 73; Holtt., Fl.
Mal. 1 (1964) 466.
Bulbophyllum concinnum Hk.f., F.B.1. 6 (1890) 189; Ridl., Fl. 4 (1924) 70; Holtt.,
Fl. Mal. 1 (1964) 454.
In Malaya, apart from the limestone field, found only in Johore and Singapore.
Collected from the limestone in Selangor and Kelantan from rather dry situations.
In Singapore and Johore found as epiphytes on old mangrove trees and by rivers.
Bulbophyllum fenestratum J.J.S., Bull. Dep. Ag. 13 (1907) 48; Carr, Gard. Bull.
S.S. 7 (1932) 33; Holtt., Fl. Mal. 1 (1964) 411.
B. punctatissimum Ridl., Fl. 4 (1924) 77.
B. rupicolum Ridl., |.c.; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75.
Rhizome creeping, pseudobulbs to 3.5 cm long and 1.5 cm wide, 4-angled. Leaves
fleshy 18 by 3 cm. Scape to 12 cm, dull purple. Sepals with purple spots.
Distributed in Java, Borneo and Sumatra. In Malaya, found mainly on the
limestone as epiphytes and also on rocks.
Bulbophyllum flammuliferum Ridl., J. Bot. 36 (1898) 211; id., Fl. 4 (1924) 74;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 451.
Rhizome creeping, stout. Pseudobulbs 3-6 cm or more apart, to 4.5 cm high.
Leaves about 12 by 2.8 cm, or more. Flowers yellow with orange tips.
Endemic and rare, chiefly from limestone; recorded from Kota Glanggi, Pahang,
and Gua Batu and Bukit Takun, Selangor.
Bulbophyllum lilacinum Ridl., J. Linn. Soc. 32 (1896) 276; id., Fl. 4 (1924) 68;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 460.
Rhizome creeping, pseudobulbs to 6 cm long, narrowly ovoid and slightly
4-angled. Leaves fleshy, to 27 by 3.5 cm. Flowers pale mauve or nearly white.
Limestone hill flora of Malaya IV 53
Distributed in peninsular Thailand and northern Malaya. In Malaya, chiefly on
limestone as epiphytes and occasionally on rocks.
Bulbophyllum membranaceum T. et B., Nat. Tijdschr. Ned. Ind. 3 (1885) 397;
Ridl., Fl. 4 (1924) 64; Holtt., Fl. Mal. 1 (1964) 430.
Bulbophyllum pulchellum Ridl., Mats. 1 (1907) 83; id., Fl. 4 (1924) 80; Holtt., Fl.
Mal. 1 (1964) 415.
Bulbophyllum sessile (Koen.) J.J.S., Fl. Btzg 6 (1905) 448; Holtt., Fl. Mal. 1 (1964)
451.
Epidendrum sessile Koen., Retz. Obs. 6 (1791) 60.
B. clandestinum Lind; in Ridl., Fl. 4 (1924) 69.
Calanthe ceciliae Rchb.f., Gard. Chron. 1 (1833) 432; Ridl., Fl. 4 (1924) 119;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 153.
Calanthe rubens Ridl., Gard. Chron. 1 (1890) 576; Holtt., Fl. Mal. 1 (1964) 151.
Preptanthe rubens Ridl., Fl. 4 (1924) 123.
Pseudobulbs ovoid and angled, about 8 cm long. Leaf blades to 40 by 15 cm.
Scape hairy, to 50 cm long; rachis many flowered. Flowers pink with crimson.
Distributed in Thailand and northern Malaya, terrestrial and restricted to
limestone.
Calanthe triplicata (Will.) Awes, Philip. J. Sc. Bot. 2 (1907) 326; Holtt., Fl. Mal.
1 (1964) 154.
C. veratrifolia R. Br. in Ridl., Fl. 4 (1924) 119.
Orchis triplicata Will., Usteri Ann. Bot. 6 (1796) 52.
Found in the lowlands, especially in freshwater swamp-forest, and extending on
to the hills to almost 1600 m. It is also the commonest species of Calanthe on
limestone; being frequently found in partly to completely shaded areas with deep or
shallow soils, especially where there is a slight accumulation of decaying litter.
Calanthe vestita Lindl., Gen. et Sp. Orch. (1833) 250; Holtt., Fl. Mal. 1 (1964) 151.
Preptanthe vestita Rchb.f, in Ridl., Fl. 4 (1924) 123.
Pseudobulbs large, ovoid and angled to 10 cm long; leaves to about 45 by 12 cm.
Scape to 70 cm long hairy; flowers, well-spaced; bracts broad, 1.5 cm long.
Distributed in Thailand, Borneo and Celebes; in Malaya, collected only once.
This specimen was growing as an epiphyte on the top of Gua Batu in Selangor
(Ridley s.n. Dec. 1896).
Camarotis apiculata Rchb.f., Bonpl. 5 (1857) 39; Carr, Gard. Bull. S.S. 7 (1932) 54;
Holtt., Fl. Mal. 1 (1964) 633.
Saccolabium saxicolum Ridl., Trans. Linn. Soc. 3 (1893) 374; id., Fl. 4 (1924)
172; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 78.
54 Gard. Bull. Sing. 36(1) (1983)
Ceratostylis pendula Hk.f., F.B.1. 5 (1890) 826; Ridl., Fl. 4 (1924) 111; Holtt., Fl.
Mal. 1 (1964) 496.
Endemic to Malaya, widely distributed in the lowlands but not very common, col-
lected only once over limestone (UNESCO 455 from Gua Musang, Kelantan).
Coelogyne asperata Lindl., J. Hort. Soc. 4 (1849) 221; Ridl., Fl. 4 (1924) 131;
Holtt., Fl. Mal. 1 (1964) 253.
Coelogyne pallens Ridl., J. Str. Br. R. As. Soc. 39 (1903) 81; id., Fl. 4 (1924) 135;
Holtt., Fl. Mal. 1 (1964) 248.
This species is known from Kedah Peak, Taiping Hills and one limstone locality
(Bukit Takun in Selangor).
Coelogyne pandurata Lindl., Fol. Orch. Coelogyne (1854) 7; Ridl., Fl. 4 (1924) 133;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 254.
Coelogyne rochussenii de Vr., III. Orch. (1954) t. 2; Ridl., Fl. 4 (1924) 131;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 257.
Corbyas mucronatus (Bl.) Schltr, Fed. Rep. 19 (1923) 20; Holtt., Fl. Mal. 1 (1964)
93.
Corysanthes mucronata Bl., in Ridl., Fl. 4 (1924) 205;; Henders., J. Mal. Br. R.
As. Soc. 17 (1939) 75.
Corymborchis brevistylis (Hk.f.) Holtt., Fl. Mal. 1 (1953) 144, op. cit. (3rd ed.)
(1964) 146.
Corymbis brevistylis Hk.f., F.B.I. 6 (1890) 91.
Stem to 60 cm tall. Leaves 12-15 by 5-7.5 cm. Inflorescence with few flowers, col-
umn 8 mm long.
Endemic and known from a single record made on limestone at Kuala Dipang,
Perak.
Corymborchis rhytidocarpa (Hk.f.) Holtt., Fl. Mal. 1 (1953) 144, op. cit. (3rd ed.)
(1964) 146.
Corymbis longiflora Hk.f., in Ridl., 4 (1924) 208.
C. rhytidocarpa Hk.f., F.B.I. 6 (1890) 92.
This is a lowland plant. It has been recorded only once from limestone, growing
from rock crevices. (Molesworth-Allen 4435 from Gunong Tempurong, Perak)
Corymborchis veratrifolia Bl., Fl. Jav. N.S. (1858) 105; Holtt., Fl. Mal. 1 (1964)
144.
Corymbis longiflora Hk.f., F.B.I. 6 (1890) 92; Ridl., Fl. 4 (1924) 208; Henders.,
J. Mal. Br. R. R. As. Spc. 17 (1939) 75.
Cymbidium dayanum Rchb.f., Gard. Chron. (1869) 710; Ridl., Fl. 4 (1924) 146;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 75; Holtt., Fl. Mal. 1 (1964) 519.
C. acutum Ridl., J. Linn. Soc. 32 (1896) 334.
Limestone hill flora of Malaya IV 55
Cymbidium finlaysonianum Lindl., Gen. et Sp. Orch. (1833) 164. Ridl., Fl. 4 (1924)
145; Holtt., Fl. Mal. 1 (1964) 520.
Dendrobium acerosum Lind., Bot. Reg. 30 (1841) 86; Ridl., Fl. 4 (1924) 38; Holtt.,
Fl. Mal. 1 (1964) 338.
Dendrobium aloifolium (Bl.) Rchb.f., Walp. Ann. 6 (1861) 279; Holtt., Fl. Mal. 1
(1964) 330.
D. serra Lindl., in Ridl., Fl. 4 (1924) 35.
Dendrobium excavatum (Bl.) Miaq., Fi. Ind. Bat. 3 (1859) 644; Holtt., Fl. Mal. 1
(1964) 336.
D. atropurpureum quoad Ridl., Fl. 4 (1924) 39.
Dendrobium farmeri Paxt., Mag. 15 (1849) 241; Holtt., Fl. Mal. 1 (1964) 282.
Dendrobium indivisum (Bl.) Miq., Fl. Ind. Bat. 3 (1859) 630; Holtt., Fl. Mal. 1
(1964) 322.
D. eulophotum Lindl., in Ridl., Fl. 4 (1924) 35; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 76.
Dendrobium langkawiense Ridl., J. Str. Br. R. As. Soc. 54 (1909) 49; Holtt., Fi.
Mal. 1 (1964) 320.
Stem to 30 cm long; leaves about 5 by 0.6 cm. Flowers apparently solitary.
Endemic and known only from a single collection from Langkawi. A second
specimen collected by Corner in 1941 from Pulau Chupak, Langkawi is doubtfully
this species.
Dendrobium leonis (Lind.) Rchb.f., Walp. Ann. 6 (1861) 280; Ridl., Fl. 4 (1924) 37;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 334.
Dendrobium planibulbe Lindl., Bot. Reg. (1843) 54: Ridl., Fl. 4 (1924) 41: Holtt.,
Fl. Mal.1 (1964) 328.
Dendrobium pumilum Roxb., Hort. Beng. (1814) 61: Ridl., Fl. 4 (1924) 34; Holtt.,
Fl. Mal. 1 (1964) 275.
Dendrobium salaccense (Bl.) Lindl., Gen. et Sp. Orch. (1830) 86; Henders., J. Mal.
Br. R. As. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 341.
D. gemellum Lindl., in Ridl., Fl. 4 ( 1924) 40.
Stem slender to 100 cm long. Leaves to 13 by 1.7 cm. Inflorescence of 2 flowers
at the nodes.
Distributed in Java. An epiphyte but more frequently found on rocks and roots
on shaded to semi-exposed summits on limestone, fairly common, but unknown
outside the limestone field.
56 Gard. Bull. Sing. 36(1) (1983)
Dendrobium secundum (Bl.) Lind., Gen. et Sp. Orch. (1830) 81. Ridl., Fl. 4 (1924)
46; Holtt., Fl. Mal. 1 (1964) 316.
Pedilonum secundum BI., Bijdr. (1825) 322.
Dendrobium setifolium Ridl., J. Linn. Soc. 31 (1896) 270; Carr, Gard. Bull. S. 5
(1929) 6; Holtt., Fl. Mal. 1 (1964) 326.
D. gracile Lindl., in Ridl., Fl. 4 (1929) 41.
Dendrobium spurium (Bl.) J.J.S., Fl. Btzg 6 (1905) 343; Holtt., Fl. Mal. 1 (1964)
Did
D. euphlebium Rchb.f. ex Lindl., in Ridl., Fl. 4 (1924) 44; Henders., J. Mal. Br.
R. as. Soc. 17 (1939) 76.
Dendrobium subulatum (B\.) Lindl., Gen. et Sp. Orch. (1830) 71; Ridl., Fl. 4 (1924)
39; Holtt., Fl. Mal. 1 (1964) 339.
Dipodium pictum (Lindl.) Rchb.f., Xen. Orch. 2 (1862) 15; Ridl., Fl. 4 (1924) 147;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 513; Holtt., Fl. Mal. 1 (1964) 513.
Ephemerantha fimbriata (Bl.) P.F. Hunt et Summ., Taxon 10 (1916) 103.
Desmotrichum fimbriatum Bl., Bijdr. (1824) 329; Ridl., Fl. 4 (1924) 30;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76.
Dendrobium plicatile Lindl., Bot. Reg. (1840) 10: Holtt., Fl. Mal. 1 (1964) 267.
Ephemerantha luxurians (J.J.S.) Hunt et Summ., Taxon 10 (1961) 105.
Dendrobium luxurians J.J.S., Bull Btzg Ser. 3 (1921) 288; Holtt., Fl. Mal. 1
(1964) 267.
Desmotrichum luxurians Carr, Gard. Bull. S.S. 7 (1932) 9.
Eria citrina Ridl., Kew Bull., (1924) 204; id., Fl. 4 (1924) 89; Holtt., Fl. Mal. 1
(1964) 366.
Eria leiophylla Lindl., J. Linn. Soc. 3 (1859) 57; Hkf., F.B.I. 5 (1890) 809; Ridl.,
Fl. 4 (1924) 99; Holtt., Fl. Mal. 1 (1964) 379.
Eria leptocarpa Hk.f., F.B.I. 5 (1890) 805; Ridl., Fl. 4 (1924) 88; Holtt., Fl. Mal.
1 (1964) 370.
Eria nutans Lindl., Bot. Reg. 26 (1840) 83; Ridl., Fl. 4 (1924) 94; Holtt., Fl. Mal.
1 (1964) 368.
Eria pannea Lindl., Bot. Reg. 28 (1842) 64; Ridl., Fl. 4 (1924) 99; Henders., J. Mal.
Br. R. as. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 380.
Eria pulchella Lindl., Bot. Reg. 27 (1841) 52; Holtt., Fl. Mal. 1 (1964) 372.
E. pendula Ridl., Fl. 4 (1924) 89; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76.
E. rigida Rchb.f, in Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76.
Tylostylis rigida Bl., in Ridl., op. cit. 104.
Limestone hill flora of Malaya IV 3y7/
Eria vestita Lindl., Bot. Reg. 30 (1844) 76; Holtt., Fl. Mal. 1 (1964) 359.
Trichotosia vestita Krzl., in Ridl., Fl. 4 (1924) 100.
Eulophia keithii Ridl., J. Linn. Soc. 32 (1896) 333; id., Fl. 4 (1924) 142; Holtt., Fl
Mal. 1 (1964) 536.
Pseudobulbs about 10 cm long, with about 5 leaves. Leaves to 30 by 1 cm, tapered
at both ends. Inflorescence 30-40 cm long; flowers greenish.
According to Holttum found only in Langkawi and mainland Kedah; but ap-
parently also found in peninsular Thailand (Ridley). Probably restricted to
limestone but the records are not conclusive.
Geodorum Citrinum Jacks., Andr. Bot. Rep (1810) t. 626; Ridl., Fl. 4 (1924) 143;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 539.
Goodyera hispida Lindl., J. Linn. Soc. 1 (1857) 183; Holtt., Fl. Mal. 1 (1964) 123.
Plant to 20 cm tall, with 4-6 leaves. Leaves 2.5-7 by 1.2-2.7 cm, base rounded,
apex acute, flushed pink with whitish or pinkish vein-network. Inflorescence erect,
to 12 cm; flowers close together and subtended by prominent bracts.
Distributed in the southern Himalayas. In Malaya, restricted to limestone; first
found on Gua Ninek, Kelantan. Subsequently also from Selangor. Grows on
sheltered localities in small soil pockets amongst boulders.
Habenaria carnea N.E. Br., Gard. Chron. 2 (1891) 729; Ridl., Fl. 4 (1924) 227;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 84.
Tuberous; stem erect, leaves from near the base, to 10 by 4 cm; olive green with
pale spots. Inflorescence terminal; upper sepal and petal forming a hood, lip to 3
cm long, 3-lobed.
This species is restricted to limestone in Langkawi and peninsular Thailand at low
elevation.
Habenaria kingii Hk.f., F.B.1. 6 (1890) 144; Ridl., Fl. 4 (1924) 299; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 76; Holtt., Fl. Mal. 1 (1964) 87.
Plant 30-50 cm; leaves 4-6, near the base; blade to 16 by 5 cm. Inflorescence to
30 cm long, flowers pale green, upper sepal about 6 mm long.
Endemic and restricted to the limestone in Perak and once from Kedah.
Habenaria reflexa BI., Bijdr. (1825) 403; Holtt., Fl. Mal. 1 (1964) 85.
H. kingii Hk.f., quoad Ridl., Fl. 4 (1924) 228.
Plant like H. kingii Hk.f., but inflorescence to 10 cm long; upper sepal and petal
forming a hood 4 mm long.
Distributed in Java and Sumatra. In Malaya usually from limestone in Pahang,
Perak and Kedah.
58 Gard. Bull. Sing. 36(1) (1983)
Hippeophyllum scortechinii (Hk.f.) Schltr, Engl. Jhrb. (1911) 45; Ridl., Fl. 4 (1924)
18; Holtt., Fl. Mal. 1 (1964) 255.
Liparis caespitosa (Thou.) Lindl., Bot. Reg. 11 (1825) 882; Holtt., Fl. Mal. 1 (1964)
205.
L. comosa Ridl., J. Linn. Soc. 32 (1896) 229; id., Fl. 4 (1924) 23; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 77.
Liparis compressa (Bl.) Lindl., Gen. et Sp. Orch. (1830) 32; Ridl., Fl. 4 (1924) 24;
Holtt., Fl. Mal. 1 (1964) 209.
Malaxis compressa Bl., Bijdr. (1925) 390.
Distributed from Sumatra to the Philippines. Found in mountain forests in
Malaya at 1300-1600 m. Recorded once on limestone at low elevation (UNESCO 55
from Bertam, Kelantan, growing on rocks).
Liparis gibbosa Finet, Bull. Soc. Bot. Fr. 55 (1908) 342; Holtt., Fl. Mal. 1 (1964)
209.
L. disticha Lindl., in Ridl., Fl. 4 (1924) 24; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 77.
Malaxis calophylla (Rchb.f.) Ktze, Rev. Gen. Pl. (1891) 673; Holtt., Fl. Mal. 1
(1964) 197.
Microstylis calophylla Rchb.f., in Ridl., Fl. 4 (1924) 11.
Distributed in peninsular Thailand. Recorded in Malaya from Penang Hill and
on limestone at Baling, Kedah; now also known to be found on the limestone at Gua
Musang, Kelantan. This Kelantan record is the southernmost location for this
species.
Malaxis latifolia Sm., Rees Cycl. (1819) 22; Holtt., Fl. Mal. 1 (1964) 195.
Microstylis congesta Rchb.f., in Ridl., Fl. 4 (1924) 12; Henders., J. Mal. Br. R.
As. Soc. 17 (1939) 77.
Malaxis micrantha (H.k.f.) Ktze, Rev. Gen. Pl. (1891) 673; Holtt., Fl. Mal. 1 (1964)
197.
Microstylis flavoviridis Ridl., Fl. 4 (1924) 12.
Microstylis micrantha Hk.f., in Ridl., l.c.
Malaxis reniloba (Carr) Holtt., Gard. Bull. S. S. 11 (1947) 263; Holtt., Fl. Mal. 1
(1964) 195.
Microstylis reniloba Carr, Gard. Bull. S.S. 7 (1932) 5; Henders., J. Mal. Br. R.
As. Soc. 17 (1939) 77.
Restricted to limestone in Perlis and peninsular Thailand.
Malleola dentifera J.J.S., Bull. Btzg (ser. 3) 9 (1927) 171; Holtt., Fl. Mal. 1 (1964)
02.
Saccolabium undulatum quoad Carr, J. Mal. Br. R. As. Soc. 6 (1928) 64.
Limestone hill flora of Malaya IV 59
Malleola undulata (Ridl.) J.J.S. et Schltr, Orch. Dtsch. Neu-Guin. (1914) 981;
Holtt., Fl. Mal. 1 (1964) 711.
Saccolabium undulatum Ridl., in Fl. 4 (1924) 172.
S. sylvestre Ridl., op. cit. 168.
Stem to 15 cm or more long. Leaves to 14 by 2 cm, apex unequal. Inflorescence
pendulous to 16 cm long. Flowers many, yellow.
Endemic. Recorded only twice, from Perak, as epiphytes on limestone hills.
Miscrosaccus ampullaceus J.J.S., Bull. Btzg (ser. 3) 5 (1922) 99; Carr, Gard. Bull.
S.S. 7 (1922) 52; Holtt., Fl. Mal. 1 (1964) 594.
Microsaccus brevifolius J.J.S., Ic. Bog. 3 (1906) 63; Carr, Gard. Bull. S.S. 7 (1922)
52; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 77; Holtt., Fl. Mal. 1 (1964) 594.
Microsaccus javensis Bl., Bijdr. (1825) 368; Ridl., Fl. 4 (1924) 175; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 77; Holtt., Fl. Mal. 1 (1964) 595.
Distributed in Java. In Malaya, recorded from the southern half. Collected from
limestone in Selangor and Kelantan. (The identification of Kelantan specimens
UNESCO 300 and 472 is doubtful). This species is common on Bukit Takun,
Selangor.
Oberonia anceps Lindl., Sert. Orch. (1838) t. 8; Ridl., Fl. 4 (1924) 15; Holtt., Fl.
Mal. 1 (1964) 219.
Oberonia calcicola Holtt., Gard. Bull. S.S. 11 (1947) 248; id., Fl. Mal. 1 (1964) 217.
Stems very stout. Leaves about 6, spreading out evenly like a fan, to 5.5 by 0.9
cm. Inflorescence slender, to 10 cm long. Flowers in whorls of about 6.
Endemic. Known in Malaya from a single record (Henderson 21398) from the
limestone at Langkawi as an epiphyte. Also from peninsular Thailand
Oberonia caudata King et Prantl., J.R. As. Soc. Beng. 66 (1897) 581; Ridl., Fl. 4
(1924) 18; Seid. et Smit. Orch. Thai. (1959) 157; Holtt., Fl. Mal. 1 (1964) 221.
Stem 3-5 cm long with 4-5 leaves. Leaves at 45 degrees to the stem, tips incurved,
5 by 0.4 cm. Inflorescence about 5 cm long, decurved.
Stem 3-5 cm long with 4-5 leaves. Leaves at 45 degrees to the stem, tips incurved,
5 by 0.4 cm. Inflorescence about 5 cm long, decurved.
Recorded once from Thailand. In Malaya, known from a collection made by
Scortechini in Perak. More recently recorded on limestone from Kelantan
(UNESCO 337, from Gua Serai).
Oberonia dissitiflora Ridl., J. Linn. Soc. 32 (1896) 218; Holtt., Fl. Mal. 1 (1964)
218.
O. lunata Lindl., in Ridl., Fl. 4 (1924) 14.
60 Gard. Bull. Sing. 36(1) (1983)
Oberonia flava Ridl., J. F.M.S. Mus. 4 (1909) 64; id., Fl. 4 (1924) 17; Holtt., Fl.
Mal. 1 (1964) 214.
This very rare endemic, until 1962, was known only from a single collection from
Telom, to the East of Cameron Highlands in Pahang. The second known collection
(UNESCO 309) is from over limestone in Kelantan.
Oberonia spathulata Lindl., Gen. et Sp. Orch. (1830) 16; Ridl., Fl. 4 (1924) 17;
Holtt., Fl. Mal. 1 (1964) 220.
Oberonia transversiloba Holtt., Gard. Bull. S.S. 11 (1947) 285; Henders., J. Mal.
Br. R. As. Soc. 17 (1939) 77; Holtt., Fl. Mal. 1 (1964) 214.
Stems 3-5 cm long with about 6 leaves. Leaves at a very acute angle to the stem,
to 9.5 by 0.5 cm. Inflorescence erect, to 16 cm long. Flowers in whorls, yellow
orange.
Endemic to limestone in Malaya. Until 1962 known from Gua Tipus in north
Pahang (Henderson 19448). Now recorded from Kelantan. This species is probably
more common than the records suggest.
Paphiopedilum lowii (Lindl.) Pfitz., in Engl., Bot. Jahrb. 19 (1894) 42; Ridl., Fl.
4 (1924) 232; Holtt., Fl. Mal. 1 (1964) 73.
Cypripedium lowii Lindl., Gard. Chron. (1847) 765.
Paphiopedilum niveum (Rchb.f.) Pfitz., Engl. Bot. Jahrb. 19 (1894) 40; Ridl., Fl.
4 (1924) 231; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 77; Seid. et Smit., Orch.
Thai. (1958) 10; Holtt., Fl. Mal. 1 (1964) 69.
Plant fleshy. Leaves almost horizontal, surface with pale purple mottling on a
dark green background; undersurface purple. Inflorescence about 12 cm long with
1-2 large flowers which are 5-6 cm across.
Distributed in peninsular Thailand. In Malaya, only from Langkawi where it is
restricted to limestone and often locally abundant in partly shaded situations; grow-
ing on soil and from rock crevices.
Phalaenopsis cornu-cervi (Breda) Bl. et Rchb.f., Hamb. Gartenz. 16 (1860) 116;
Ridl., Fl. 4 (1924) 156; Holtt., Fl. Mal. 1 (1964) 667.
Phalaenopsis decumbens (Griff.) Holtt., Gard. Bull. S.S. 11 (1947) 286; id., Fl.
Mal. 1 (1964) 669.
Aerides decumbens Griff., Notul. 3 (1851) 365.
Kingiella decumbens Rolfe., in Ridl., Fl. 4 (1924) 158.
Pholidota pallida Lindl., Bot. Reg. (1836) 1777; Holtt., Fl. Mal. 1 (1964) 237.
P. imbricata sensu Lindl., in Ridl., Fl. 4 (1924) 139; Henders., J. Mal. Br. R. As.
Soc. 17 (1939) 77.
Pseudobulbs conical, 3-6 cm long. Leaves fleshy to 30 by 6 cm; plicate. In-
florescence to 30 cm long, with numerous flowers and persistent semi-circular
bracts.
Limestone hill flora of Malaya IV 61
Distributed in Burma and southern China and southwards to Australia. Common
on the limestone in Malaya, but also found in lowland forest. This species grows
in partly shaded areas but will tolerate extreme exposure.
Phreatia secunda (Bl.) Lindl., Gen. et Sp. Orch. (1830) 64; Holtt., Fl. Mal. 1 (1964)
eee
P. microtidis Lindl., in Ridl., Fl. 4 (1924) 106.
P. minutiflora Lindl., in Ridl., op. cit. 105; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 77.
Poaephyllum pauciflorum (Hk.f.) Ridl., Mats. 1 (107) 109; id., Fl. 4 (1924) 109;
Holtt., Fl. Mal. 1 (1964) 511.
Agrostophyllum pauciflorum Hk.f., F.B.I. 5 (1890) 824.
Eria minutiflora Ridl., J. Linn. Soc. 32 (1896) 297.
Podochilus lucescens Bl., Bijdr. (1825) 295; Ridl., Fl. 4 (1924) 193; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 77; Holtt., Fl. Mal. 1 (1964) 502.
Podochilus microphyllus Lindl., Gen. et Sp. Orch. (1825) 234; Ridl., Fl. 4 (1924)
193; Holtt., Fl. Mal. 1 (1964) 500.
Podochilus tenuis (Bl.) Lindl., Gen. et Sp. Orch. (1833) 235; Ridl., Fl. 4 (1924) 193;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 77; Holtt., Fl. Mal. 1 (1964) 500.
Polystachya flavescens (Bl.) J.J.S., Fl. Btzg 6 (1905) 284; Holtt., Fl. Mal. (1964)
500.
P. penangensis Ridl., Fl. 4 (1924) 152.
P. siamensis Ridl., op. cit. 152; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 77.
Pomatocalpa kunstleri (Hk.f.) J.J.S., Nat. Tijdschr. Ned. Ind. 72 (1912) 104.
Saccolabium pubescens Ridl., J. Linn. Soc. 31 (1896) 295; id., Fl. 4 (1924) 174.
Cleisostoma kunstleri Hk.f., Ic. Pl. (1895) t. 2335.
Pomatocalpa latifolium (Lindl.) J.J.S., Nat. Tijds. Ned. Ind. 72 (1912) 105; Holtt.,
Fl. Mal. 1 (1964) 628.
Cleisostoma latifolium Lindl., Bot. Reg. 26 (1840) 60.
Saccolabium hortense Ridl., Fl. 4 (1924) 167.
S. latifolium var parviflorum Ridl., op. cit. 166.
Pomatocalpa naevatum J.J.S., Bull. Btzg (ser. 2) 9 (1913) 106; Seid et Smit., Orch.
Thai (1964) 653; Holtt., Fl. Mal. 1 (1964) 630.
Saccolobium latifolium var strictum Ridl., Fl. 4 (1924) 166.
Stem climbing. Leaves 10-18 by 1.5-2.4 cm. Inflorescence 15-40 cm, branched,
flowers greenish yellow with red brown spots.
Distributed in Thailand and Java. Restricted to limestone in Malaya. Known
from Gua Batu, Selangor and Gua Musang, Kelantan.
62 Gard. Bull. Sing. 36(1) (1983)
Pomatocalpa spicatum Breda, Orch. Kuhl et Hass. (1827) t. 15; Holtt., Fl. Mal. 1
(1964) 628.
Saccolabium hobsonii Ridl., Fl. 4 (1924) 167.
S. uteriferum Ridl., op. cit. 168.
Pteroceras ciliatum (Ridl.) Holtt., Kew Bull. 14 (1960) 269; Seid. et Smit., Orch.
Thai. (1964) 820.
Dendrocolla ciliata Ridl., Fl. 4 (1924) 188.
Sarcochilus ciliatus (Ridl.) J.J.S., Bull. Btzg (ser. 3) 8 (1926) 63; Holtt., Fl. Mal.
1 (1964) 690.
Pteroceras hirsutum (Hk.f.) Holtt., Kew Bull. 14 (1960) 270.
Ascochilus hirsutus Ridl., Fl. 4 (1924) 182.
Sarcochilus hirsutus Hk.f., F.B.I. 6 (1890) 38; Holtt., Fl. Mal. 1 (1964) 691.
Pteroceras tanyphyllum (Ridl.) Holtt., Kew Bull. 14 (1960) 271.
Sarcochilus tanyphyllus Ridl., Fl. 4 (1924) 179; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 78; Holtt., F. Mal. 1 (1964) 686.
Stems to 5 cm long. Leaves many, to 40 by 4 cm or usually less, with an unequally
bilobed apex. Inflorescence to 13 cm long; bracts spreading. Flowers pale yellow
with purple spots at the base of lateral sepals; about 3-4 cm across.
Endemic to limestone, previously known only from Kota Glanggi in Pahang.
Now known from Kelantan as well, where it is not uncommon.
Renanthera elongata Lindl., Gen. et Sp. Orch. (1833) 218; Ridl., Fl. 4 (1924) 160;
Holtt., Fl. Mal. 1 (1964) 634.
Renanthera histrionica Rchb.f., Gard. Chron. (1878) 74; Seid. et Smit., Orch. Thai
(1962) 591; Holtt., Fl. Mal. 1 (1964) 634.
Renantherella histrionica Ridl., J. Linn. Soc. 32 (1896) 355; id., Fl. 4 (1924) 161.
Sarcanthus machadonis (Ridl.) J.J.S., Nat. Tijdschr. Ned. Ind. 72 (1912) 89; Holtt.,
Fl. Mal. 1 (1964) 652.
Saccolabium machadonis Ridl., Fl. 4 (1924) 168.
Sarcanthus rugulosus (Ridl.) Holtt., Gard. Bull. S.S. 11 (1947) 288, id., Fl. Mal. 1
(1964) 659.
Saccolabium rugulosum Ridl., Fl. 4 (1924) 168.
Stem 15 cm. Leaves spreading to 13 by 1.3 cm, slightly constricted at 0.5-1.5 cm
from the apex. Infloresence 1.5 cm long with a few flowers. Flowers yellow, spotted
red.
Endemic to Malaya; formerly known only from a single collection from Kedah
Peak (not limestone). Now known from two other numbers from the Kelantan
limestone (UNESCO 38, 358).
Limestone hill flora of Malaya IV 63
Sarcanthus sacculatus Ridl., J. Linn. Soc. 32 (1896) 368; Holtt., Fl. Mal. 1 (1964)
653. Seid. et Smit., Orch. Thai. (1964) 695.
Saccolabium sacculatum Ridl., Fl. 4 (1924) 172; Henders., J. Mal. Br. R. As.
Soc. 17 (1939) 78.
Sarcanthus scortechinii Hk.f., F.B.1. 6 (1890) 68; Holtt., Fl. Mal. 1 (1964) 655; Seid.
et Smit., Orch. Thai (1964) 675.
Saccolabium scortechinii Ridl., Fl. 4 (1974) 169.
Sarcanthus subulatus (B\.) Rchb.f., Bonpl. 5 (1857) 41; Holtt., Fl. Mal. 1 (1964)
654.
Saccolabium secundum Ridl., Fl. 4 (1924) 169; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 78.
Sarcanthus termissus Rchb.f., Hamb. Gartenz. 16 (1860) 15; J.J.S., Bull. Btzg (ser.
3) 8 (1926) 367; Holtt., Fl. Mal. 1 (1964) 657.
Stem to 6 cm long, stout, with about 10 leaves. Leaves to 23 by 1.8 cm. In-
florescence pendulous, to 25 cm, unbranched or with a few branches. Flowers pale
green; sepals and petals with dull purple-red median bands.
Distributed in Java and Sumatra. In Malaya, known only from Langkawi, and
Lenggong in in Perak; the Lenggong collection is from limestone and the Langkawi
ones are probably also.
Schoenorchis micrantha Bl., Bijdr. (1825) 362. Seid. et Smit., Orch. Thai. (1962)
612; Holtt., Fl. Mal. 1 (1964) 663.
Saccolabium perpusillum Hk.f., in Ridl., Fl. 4 (1924) 171.
Spathoglottis hardingiana Par. et Rchb.f., Otia. Bot. Hamb. 1 (1878) 45; Ridl., FI.
4 (1924) 118; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 78; Seid. et Smit., Orch.
Thai. (1961) 341; Holtt., Fl. Mal. 1 (1964) 167.
Plant with small round pseudobulbs. Leaves to about 15 cm long; Scape to:20
cm long, rachis elongated; flowers about 2.5 cm wide; crimson to pale pinkish-
purple.
Distributed in Lower Burma and Thailand. In Malaya, it is restricted to the
limestone on Langkawi. This is a terrestrial species, growing from soil and rock
crevices in partly shaded localities, often close to the sea.
Staurochilus fasciatus (Rchb.f.) Ridl., J. Linn. Soc. 32 (1896) 351, id., Fl. 4 (1924)
157; Seid. et Smit., Orch. Thai. (1962) 600.
Trichoglottis fasciata Rchb.f., Gard. Chron. (1872) 699; Holtt., Fl. Mal. 1 (1964)
644.
Taeniophyllum culiciferum Ridl., Fl. 4 (1924) 176; Carr, Gard. Bull. S.S. 7 (1932)
73; Holtt., Fl. Mal. 1 (1964) 592; Seid. et Smit., Orch. Thai (1964) 718.
Taeniophyllum filiforme J.J.S., Bull. Inst. Bot. Btzg 7 (1900) 4; Carr, Gard. Bull.
64 Gard. Bull. Sing. 36(1) (1983)
S.S. 7 (1932) 80; Holtt., Fl. Mal. 1 (1964) 593; Seid et Smit., Orch. Thai. (1964)
720.
T. macrorhizum Ridl., Fl. 4 (1924) 176.
Taeniophyllum obtusum Bl., Bijdr. (1825) 35; Holtt., Fl. Mal. 1 (1964) 592; Seid.
et Smit., Orch. Thai. (1964) 722.
T. serrula Hk.f., in Ridl., Fl. 4 (1924) 176.
Thecostele alata (Roxb.) Par. et Rchb., T. Linn. Soc. 30 (1874) 135; Holtt., Fl. Mal.
1 (1964) 558; Seid. et Smit., Orch. Thai. (1961) 514.
T. maculosa Ridl., Fl. 4 (1924) 191; Henders., J. Mal. Br. R. As. Soc. 17 (1939)
78.
T. zollingeri Rchb.f., in Ridl. op. cit. 192.
Thelasis carinata Bl., Bijdr. (1825) 385; Ridl., Fl. 4 (1924) 200; Henders., J. Mal.
Br. R. As. Soc. 17 (1939) 78; Holtt., Fl. Mal. 1 (1964) 550.
Thelasis micrantha (Brongn.) J.J.S., Fl. Btzg 6 (1905) 495; Seid. et Smit., Orch.
Thai. (1961) 461; Holtt., Fl. Mal. 1 (1964) 551.
T. decurva Hk.f., in Ridl., Fl. 4 (1924) 200; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 78.
Thelasis succosa Carr, Gard. Bull. S.S. 7 (1932) 35; Holtt., Fl. Mal. 1 (1964) 550.
Pseudobulbs lying on rhizome, about 1 cm diameter and 7 mm tall. Leaves fleshy,
almost terete, to 15 cm long and 1 cm wide. Inflorescence from between
pseudobulbs, scape to 15 cm long, rachis gradually elongating to about 4 cm; bracts
persistent and deflexed, overlapping. Flowers whitish.
Endemic and restricted to limestone. Previously known only from Kota Glanggi
in Pahang. Now also known from Gua Musang, Kelantan and Bukit Chintamani,
Pahang. Epiphytic.
Thelasis triptera Rchb.f., Bonpl. 3 (1855) 219; Seid. et Smit., Orch. Thai. (1961)
458; Holtt., Fl. Mal. 1 (1964) 550.
T. elongata Bl., in Ridl., Fl. 4 (1924) 199; Henders., J. Mal. Br. R. As. Soc. 17
(1939) 78.
Thrixspermum album (Ridl.) Schltr, Orchis 5 (1911) 2;; Holtt., Fl. Mal. 1 (1964)
613.
Dendrocolla alba Ridl., Fl. 4 (1924) 188.
Thrixspermum amplexicaula (Bl.) Rchb.f., Xen. Orch. 2 (1867) 121; Holtt., Fl. Mal.
i (1964) 602.
T. lilacinum Rchb.f., in Ridl., Fl. 4 (1924) 184.
Trichoglottis misera (Ridl.) Holtt., Gard. Bull. S.S. 11 (1947) 292; Seid. et Smit.,
Orch. Thai. (1962) 604; Holtt., Fl. Mal. 1 (1964) 643.
Saccolabium miserum Ridl., J. Linn. Soc. 32 (1896) 359; id., Fl. 4 (1924) 171.
Limestone hill flora of Malaya IV 65
Trichoglottis retusa Bl., Bijdr. (1825) 360; Ridl., Fl. 4 (1924) 162; Henders., J. Mal.
Br. R. As. Soc. 17 (1939) 78; Holtt., Fl. Mal. 1 (1964) 642.
Stem long-climbing. Leaves 7-12 by 1.5-2 cm; ends unequally bilobed. In-
florescence 1-flowered, 1-3 at a node. Flowers greenish yellow with red-brown spots,
2-3 cm across.
Distributed in Java, Sumatra and Borneo. In Malaya, found chiefly on limestone,
in Kelantan, Pahang and Selangor.
Trichoglottis winkleri J.J.S., Engl. Bot. Jahrb. 48 (1912) 105; Holtt., Fl. Mal. 1
(1964) 641.
var. minor J.J.S., Bull. Btzg (ser. 2) 26 (1918) 102; Holtt., op. cit. 642.
Stem pendulous, about 30 cm long. Leaves about 5 by 1.3 cm. Inflorescence
1-flowered, 1-3 at a node. Flowers pale yellow with brown spots and streaks.
The typical variety is distributed in Borneo and Sumatra; known in Malaya only
from Port Swettenham and Cameron Highlands. Var. minor is distributed in Java
and recorded from Malaya once only (Kota Glanggi, Pahang).
Tropidia curculigoides Lindl., Gen. et Sp. Orch. (1840) 497; Ridl., Fl. 4 (1924) 209;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 79; Holtt., Fl. Mal. 1 (1964) 143.
Uncifera tenuicaulis (Hk.f.) Holtt., Gard. Bull. S.S. 11 (1947) 292; Seid. et Smit.,
Orch. Thai. (1962) 642; Holtt., Fl. Mal. 1 (1964) 665.
Saccolabium tenuicaule Hk.f., in Ridl., Fl. 4 (1924) 171.
Stem to 30 cm long, more or less pendulous. Leaves well spaced, to 12 by 0.6 cm.
Inflorescence very short, from nodes, each with 1-3 flowers. Midlobe as long as the
forward curving spur, with 2 small diverging curved horns at the very tip.
Distributed in peninsular Thailand. In Malaya, found chiefly on limestone in
Pahang, Selangor, Perak and Langkawi.
Vandopsis gigantea (Lindl.) Pfitz., Nat. Pflanz. 6 (1889) 210; Seid. et Smit., Orch.
Thai (1962) 586; Holtt., Fl. Mal. 1 (1964) 647.
Vanda gigantea Lindl., Gen. et Sp. Orch. (1833) 215.
Stauropsis gigantea Benth., in Ridl., Fl. 4 (1924) 155.
Excluded Species
Tainia plicata Ridl. ex Henders., J. Mal. Br. R. As. Soc. 17 (1939) 78.
According to Henderson this species is known only from Gua Tipus limestone in
Pahang. I have not been able to trace this in any literature, nor have I seen any
specimens.
Notes
Dendrobium tetrodon Rchb.f. ex Lind., J. Linn. Soc. 3 (1859) 10; Ridl., Fl. 4 (1924)
45; Holtt., Fl. Mal. 1 (1964) 293.
66 Gard. Bull. Sing. 36(1) (1983)
Endemic to Malaya, not uncommon in open places in the lowlands. Two numbers
from limestone are doubtfully identified as this species, UNESCO 444 & 456 from
Gua Musang, Kelantan.
Paphiopedilum barbatum (Lindl.) Pfitz., Engl. Bot. Jahrb. 19 (1894) 41; Holtt., Fl.
Mal. 1 (1964) 77.
Locally common, recorded on Gunong Blumut and G. Ledang in Johore, Penang
Hill, Kedah Peak and several other isolated small hills in westcoast. Strangely not
recorded on the main range.
Usually in shaded places with peaty or mossy substrate. What appears to be this
species has been seen on Gunong Tempurong, Perak. (I am grateful to Mr. Lee Toh
Ming for his information.)
Pomatocalpa setulense (Ridl.) Holtt., Gard. Bull. S.S. 11 (1947) 287; Seid. et Smit.,
Orch. Thai. (1964) 652; Holtt., Fl. Mal. 1 (1964) 628.
Saccolabium setulense Ridl., Fl. 4 (1924) 167; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 78.
Stem to 5 cm long. Leaves: fleshy, at the end of stems, to 19 by 2.5 cm. In-
florescence to 15 cm with few branches. Flowers pale yellow, lateral sepals
sometimes with reddish spots at the base.
A rare plant which according to records is restricted to limestone in peninsular
Thailand and Perlis, Malaysia. The specimen thought to be from Malaya (Ridley
15226) was collected from Bukit Rajah Wang, Setul. All previous authors have im-
plicitly or explicitly expressed Setul to be within the Malayan boundary. According
to present maps, however, Setul is within Thai territory. But this plant is very likely
to be found in Malaya proper.
Thunia alba Rchb.f., Bot. Zeitschr. (1852) 764; Holtt., Fl. Mal. 1 (1964) 185.
This species has also been recorded from limestone in Setul.
PALMAE*
1. Leaves. palmate: .. .. 0.455. 2a oe Bee oo es See oes es ners. Sener ocier eee 2
Léaves-pinnaté or bipinnate .j3.3054 seca es Sea see Oe CE aerate 6
2. Major divisions of leaf reaching the insertion on the petiole; leaflets wedge-shaped ........
one Mir cnn Grn oo dao Good CO ABO ds 6 DO QADOL Oe OOO Good Licuala modesta
Major divisions of leaf notreachingsthespetioleseeeere reece oeeeriieeteiertiante 3
3. Petioles armed with stout spines; fruits 1.5-2 cm diameter, blue green .... Livistona saribus
Petioles tinarmed! '07s<5:...2 55, Hie ocak are Sire One oA ee ete mT reve ear arnt eray oreteyarer nee ere Taner 4
4. Leaves massive, petiole channelled, with very sharp edges; leaflets compound .............
sriguailayave'S wxocdlshsd « ehage'h -«: aim yehd Bye Saeko epee ees Deer eee ee eee Borassodendron machadonis
Keaves:small> petiolenotchannelledae- eee eee rere reer eer eterna 5
5. Leafsheaths formed of broad laminate fibres. Only from Langkawi ... Maxburretia gracilis
Leafsheaths formed of fine dark fibres. Only from Selangor ........ Maxburretia rupicola
* The generous help of Dr. J. Dransfield (KEW) with this family is gratefully acknowledged.
Limestone hill flora of Malaya IV
6.
11.
a2
13:
14.
ME AVCSESIIT DMMP ITITiAl Caper Sete RNP Sorcte a oe re toes. ssl atelo cPeeie siete he Sie. wig Sh hae ee ete as pe oROE EE
CARES EDIDINNALC ML riot ae Sirsa choy sista lars ape, © ios alee fee Nees «ha de e Lela’ Caryota mitis
. SITS PEUTIC eo eee Soe so SES BOSE OHSS Ge FO Uo SCE Brg O CCE Cet i aaa
PAH SENV | ALIN SERENE eee PCE Cari e So a AER ie a1) Pelein, «1 ts od eevee auaia. as vt d we aos So vee He de Ree
. Massive clustering palm with crownshaft ........................ Onchosperma horridum
Slender palms without crownshaft (rattans, 1 species acaulescent) .....................00-
. Leaflets with white indumentum below; whole leaf ending in a barbed whip (cirrus) .......
cn. SSSA 00 SQOS08 BODES p SURES O Oe COR DOCS OC Oe Inet iit Plectocomia griffithii
Leaflets concolorous; leaf without cirrus; barbed whip (flagellum) borne on leafsheath .....
meenealemeatictsels-30/Dygc4-1 SiCMs ne ce se cic ee cece gece ssc bes ac ueeihe dope bile se faae
Ge PEAS GAS TID ENSUE ee Ne SE ee Oe en Se ee ee ae
Leaflets equidistant. Leafsheaths densely covered with 2-5 cm-long spines ................
lier acted eng eee arts Bicey GR OOURE Gee eae Calamus balingensis
Leaflets grouped. Leafsheaths with spines to 2 cm long .. Calamus siamensis var malaianus
Middle leaflets 30-60 by 3-3.5 cm. Fruits 0.8-1 cm long ............... Calamus concinnus
Middle leaflets 60-80 by 5-8 cm. Fruits 3-3.5 cm long .................. Calamus ornatus
GIP GENTS GTin@ APs Behctatiige alist or gio OF c GE OCS MES CTO rere tones Areca triandra
PAMES-EIPISISCLE ALCS (PLACIN GESC) Pat tsts <b te 152s ciao < BS Oye «aye oR S SWISS a hs Te a slesios a\eetee Se
Massive palms, with, leaves; to, 10)m.Or More Ong «opera, s:chcin.,diepo¥s « oie ere 6,018 046.8 02'S © Sisie aici e ee «
Slender undergrowth palms; leaves rarely exceeding 2m ..................00ceeeeeeeeeee
. Stem solitary, monocarpic with basipetal inflorescence production; trunk covered with black
TES - ns gediat ce dhGend bg de Dene a to SOPs Gone fe AO nee eee ae Arenga pinnata
Stem clustered, polycarpic with acropetal inflorescence production; trunk eventually bare
SR Oe ne to MSc Joie aca Siojsuc tele lois eneveis''a Gia a Sieve adtares Gla aes Arenga westerhoutii
BERCERST AME ALIEESDICSEMUAT tet - everson ties oe abuses’ ova gale bladed vadiec Arenga hookeriana
“ERRATTEN TEER CECI SUTTER Se ede US GLE ORCS OG cL Gene OPEC eM Germ ne ae ee
. Inflorescence unbranched; fruits elongate with a pronounced hook at the tip .............
+ aor Sty SoBe oeneigh fe Se ikoc 0 IOS OCR CIE ats, Ceo a eee ie ee Iguanura corniculata
Inflorescence branched (rarely unbranched); fruits curved, with no pronounced hook at the tip
ee Te TN. IL AP yos AMES SS ft Nek aide wisiel slaves Bre FS Sue 0 a's Iguanura polymorpha
14
15
16
17
Areca triandra Roxb., (Hort. Beng. (1814) 68, nom. nud.), Fl. Indica 3 (1832) 617;
Ridl., Fl. 5 (1925) 4; Henders., Mal. Br. R. As. Soc. 17 (1939) 85; Whitmore,
Palms Mal. (1973) 35.
Common palm at 700-1300 m, uncommon in the lowlands. Recorded from
limestone in Langkawi at near sea-level.
Arenga hookeriana (Becc.) Whitmore, Principes 14 (1970) 124; id., Palms Mal.
(1973) 37.
Didymosperma hookerianum Becc., Malesia 3 (1889) 186; Ridl., Fl. 5 (1925) 20;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 85.
South Thailand. In Malaya only from the east coast; not uncommon in lowland
forest, recorded several times from limestone.
Arenga pinnata (Wurmb.) Merr., Int. Herb. Amb. (1917) 119; Whitmore, Palms
Mal. (1973) 37.
68 Gard. Bull. Sing. 36(1) (1983)
Recorded once from limestone in Malaya (Whitmore FRI 4267 from Gua
Musang, Kelantan).
Arenga westerhoutii Griff., Calc. J. Nat. Hist. 5 (1845) 475; Ridl., Fl. 5 (1925) 19;
Whitmore, Palms Mal. (1973) 38.
Locally gregarious in lowland forest. Not uncommon on the lower slopes and
gullies of limestone hills.
Borassodendron machadonis (Ridl.) Becc., Webbia 4 (1914) 361; Whitmore, Palms
Mal. (1973) 39.
Borassus machadonis Ridl., Fl. 5 (1925) 71; Molesworth-Allen, M.N.J. 18 (1964)
168.
Peninsular Thailand. Recorded from limestone at Gua Musang, Kelantan; con-
spicuous on Gunong Datok, Perak. Rare in the wild.
Calamus balingensis Furt., Gard. Bull. S. 15 (1956) 240.
Calamus sp. in Henders., J. Mal. Br. R. As. Soc. 17 (1939) 85.
Solitary. Leaf-sheaths densely covered by 2-5 cm long spines. Leaves 1.6-2 m
long. Male spadix slightly longer than the leaves, with about 6 branches. Female
inflorescence unknown.
Endemic and known from only one collection (Furtado 33073, Gunong Baling,
Kedah).
Calamus concinnus Mart., Hist. Nat. Palm. 3 (1838) 208; Ridl., Fl. 5 (1925) 50;
Furt., Gard. Bull. S. 15 (1956) 211.
Plectocomiopsis ferox Ridl., op. cit. 66.
Peninsular Thailand. Rare; in Malaya known from only three collections, one of
which is from limestone (Haniff & Nur 7087, Telok Apau, Langkawi).
Calamus ornatus Bl. var horridus Becc. in Hk.f., Fl. Br. Ind. 6 (1893) 460; Furt.,
Gard. Bull. S. 15 (1956) 202.
C. ornatus Bl. sensu Griff., Calc. J. Nat. Hist. 5 (1844) 37; Ridl., Fl. 5 (1925) 54.
Endemic; in lowland forest. Recorded on limestone from Gua Musang and from
the small outcrops in Johore.
(Dransfield, J. (pers. com.) does not regard C. ornatus var. horridus as distinct from
the typical variety C. ornatus var. ornatus.
Calamus siamensis Becc. var. malaianus Furt., Gard. Bull. S. 15 (1956) 215.
C. densiflorus Becc., in Ridl., Fl. 5 (1925) 53.
C. siamensis Becc., in Ridl. op. cit. 59.
The species is described from Thailand and the variety is found only in Kedah
and Perlis; once from limestone (Henderson 23028, Tebing Tinggi, Perlis).
Caryota mitis Lour., Fl. Cochinch. (1790) 569; Ridl., Fl. 5 (1925) 20; Whitmore,
Palms Mal. (1973) 44.
Limestone hill flora of Malaya IV 69
Iguanura corniculata Becc., Malesia 3 (1886) 187; Ridl., Fl. 5 (1925) 15; Whitmore,
Palms Mal. (1973) 63; Kiew, Gard. Bull. S. 28 (1976) 205.
Plant similar to J. polymorpha. Rare; known from two collections only, one from
limestone (Henderson 25059, Bukit Serdam, Pahang).
Iguanura polymorpha Becc., Malesia 3 (1886) 189; Ridl., Fl. 5 (1925) 15; Whitmore,
Palms Mal. (1973) 64; Kiew, Gard. Bull. S. 28 (1976) 213.
Not uncommon on limestone.
Licuala modesta Becc., Malesia 3 (1886) 195; Ridl., Fl. 5 (1925) 28; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 85; Furt., Gard. Bull. S.S. 11 (1940) 60.
L. wrayi Becc. ex Ridl., Fl. 5 (1925) 28.
Endemic, from the lowlands to 1600 m. Recorded from the limestone at Gunong
Pondok, Perak.
Livistona saribus (Lour.) ex Chev., Bull. Econ. Indochine (ser 2) 22 (1919) 501;
Whitmore, Palms Mal. (1973) 72.
L. cochinchinensis Mart., Hist, Nat. Palm. 3 (1838) 242; Henders., J. Mal. Br.
R. As. Soc. 17 (1939) 85.
Indochina, Java, Sumatra, Borneo and the Philippines. Widely distributed in
Malaya, except in the South. Recorded once on limestone (Henderson s.n. 7th June
1930, from Gunong Pondok, Perak).
Maxburretia gracilis (Burr.) Dransfield, Gentes Herb. 11 (1978) 194.
Liberbaileya gracilis (Burr.) Burr. & Potztal, Willdenowia 1 (1956) 530; Whit-
more, Principes 14 (1970) 97; id., Palms Mal. (1973) 67.
L. langkawiensis Furt., Gard. Bull. S. S. 11 (1940) 238.
Symphyogyne gracilis Burr., Notizbl. Bot. Gart. Berl. 15 (1941) 317.
Endemic and known only from the limestone at Dayang Bunting, Langkawi.
Very likely to be found on the limestone islands of southern Thailand too.
Maxburretia rupicola (Ridl.) Furt., Gard. Bull. S. 11 (1940) 240; Whitmore, Palms
Mal. (1973) 76.
Livistona rupicola Ridl., Fl. 5 (1925) 23; Henders., J. Mal. Br. R. As. Soc. 17
(1939) 85.
Stem to Im. Petiole 50-90 cm, lamina 50-60 cm across with 30-35 segments. In-
florescence to 60 cm with a strong musty-sweet fragrance. Fruits ovoid, 0.8-0.9 cm
long; ripening black.
Endemic and restricted to the limestone in Selangor.
Onchosperma horridum (Griff.) Scheff., Tijdschr. Ned. Ind. 32 (1871) 191; Ridl.,
Fl. 5 (1925) 16; Whitmore, Palms Mal. (1973) 82.
Areca horrida Griff., Calcutta J. Nat. Hist. 5 (1845) 465.
70 Gard. Bull. Sing. 36(1) (1983)
There is a clump of this palm on the top of the Gua Batu limestone growing in
a broad gully amongst tangled shrubby vegetation.
Plectocomia griffithii Becc. ex Hk.f., F.B.I. 6 (1893) 478; Ridl., Fl. 5 (1925) 70;
Furt., Gard. Bull. S. 13 (1951) 346; Whitmore, Palms Mal. (1973) 94.
Thailand, Peninsular Malaya. Common especially in mountain forests and
coastal hills. Once from limestone (Chin 1273, Gua Batu, Selangor).
PANDANACEAE
1. Plant of sandy or rocky shores, spines on leaves white ........... Pandanus odoratissimus
Inland'plant; 1f by shoresispines. not white! ee eeEeeeGe crac cer eee 2
2. Leaf apex very abruptly acuminate-caudate, boat-shaped, when older, splitting and appearing
bilobed «.:'e isto Peta tases ake ae eter cna aia eee serene enor eae isteneeer serene Pandanus irregularis
Leaf apex Ot SO. wis c cnalaccicheececeie sie engid ovenese gueveter ale areyereieresecelaiele Gal oer aera ree ere 3
3. Leaf less than 2.5 cm wide, usually less than 150 cm long; stem slender, less than 5 cm in
Giameter® «2 o..460 teed enw eae hole ae S Se SO OG Pee race a EE TOT ee oe ere eee 4
Leaf more than 2.5 cm wide, usually more than 4 cm wide, and often more than 200 cm long;
stem\stout,, more than) 5) cm diameter aac eee er ecreeecricr eae Cie ieee ene 5
4. Plant much branched; stems 1-5 m tall, thorny; older parts devoid of leaf bases; very rare on
limestone,a fOreSt SPeCleS’ ssclscuac ite recreeisie icin cise eee Pandanus recurvatus
Plant seldom branched; often forming dense clumps; stems about 30 cm or more long; all
covered by/persistentaleaf bases! ia -rciertaecraee eieits siesta ae Pandanus alticola
5. Drupes all simple, one-celled. Plant large; leaves to 4 m long and 9-12 cm wide (on younger
plants with stem less than 1 m tall); older plants with leaves smaller, about 3 m long; tip of apical
leaves filiform and 10-15 cm long. Common on Gua Batu, Selangor ... Pandanus calcicola
Drupes connate, 1-4 celled. Plant large; leaves smaller, to 4-7 cm wide; tip 4-5 cm long. So far
onlysknown) from) the» eulamareay herd kerries ieee Pandanus piniformis
Pandanus alticola Holtt. & St. John, Pac. Sci. 16 (1962) 218; Stone, M.N.J. 21
(1968) 136.
A fairly common endemic epiphyte in Malaya. Recorded from limestone; seen on
Bukit Anak Takun, Bukit Takun, and Gua Batu in Selangor, and Gua Musang in
Kelantan. They are always fround growing on rocks and from rock crevices, often
forming large dense clumps. Common on Bukit Takun, in very exposed to partly
shaded situations. Distinguished by its linear, narrow leaves and its often clump-
forming habit.
Pandanus calcicola Holtt. & St. John, Pac. Sci. 16 (1962) 230; Stone, M.N.J. 21
(1962) 128.
Plants to 5 m tall, 2-3 times branched when old. Stem 10-20 cm in diameter at
the basal part. Leaves about 3 m long and 9 m wide; often larger in young plants
when stems are as yet unbranched. Fruits terminal, either a single syncarp to 20 cm
by 10 cm, or with 2-3 laterals which are smaller, about 12 by 9 cm each. (Dimensions
all refer to mature fresh fruits.)
Endemic to limestone in Malaya. Recently found to be common on the slopes of
Gua Batu in Selangor, on rocky localities with soil pockets. Originally described
from a Perlis specimen.
Limstone hill flora of Malaya IV 71
Pandanus irregularis Ridl., Fl. 5 (1925) 76; Henders., J. Mal. Br. R. As. Soc. 17
(1939) 82; St. John, Pac. Sci. 17 (1963) 344; Stone, M.N.J. 21 (1968) 6.
Erect plant, when old to 5 m tall, 2-3 times branched, lower half of stem 5-10
cm in diameter. Leaves 100-150 by 6-10 cm, tip abruptly acuminate-caudate, boat-
shaped, but this is lost in older leaves which are split at the tip and apparently bilob-
ed. Peduncle to 30 cm long, bearing 3-7 syncarps each of which globose to sub-
globose, about 6 cm in diameter when mature.
Endemic to limestone in Malaya. Common on some of the limestone hills around
Gua Musang in Kelantan; often a conspicuous feature of the precipitous, pinnacled
ridges.
Pandanus odoratissimus L.f., Suppl. (1781) 424; Stone, M.N.J. 21 (1968) 3.
P. fascicularis Lam., in Ridl. Fl. 5 (1925) 72; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 82.
P. tectorius Sol. ex Balf.f., J. Linn. Soc. 17 (1878) 63.
Pandanus piniformis Holtt. & St. John, Pac. Sci. 16 (1962) 224; Stone M.N.J. 21
(1968) 130.
Plant forming clumps, stem 3-4 m tall. Leaves about 3 m long and 4-7 cm wide;
glaucous beneath. Penduncle about 30 cm long, bearing up to 7 heads of syncarps,
each ovoid and 6-11 cm long.
Endemic to limestone in Malaya, and so far known only from limestone in the
Pulai area in Perak.
Pandanus recurvatus St. John, Pac. Sci. 19 (1965) 227; Stone, M.N.J. 19 (1965) 210;
id., 21 (1968) 136.
STEMONACEAE
Stemona tuberosa Lour., Fl. Cochinch. (1790) 404; Ridl., Fl. 4 (1924) 320;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 82; Hutch., Fam. Fl. Pl. 2 (1959) 656.
S. curtisii Hk.f., F.B.I. 6 (1892) 298.
TACCACEAE
Tacca leontopetaloides (L.) Ktze, Rev. Gen. Pl. 2 (1891) 704.
T. pinnatifida Forst., Char. Gen (1776) 70; Ridl., Fl. 5 (1925) 309; Henders., J.
Mal. Br. R. As. Soc. 17 (1939) 81.
TRIURIDACEAE
Sciaphila asterias Ridl., J.F.M.S. Mus. 6 (1915) 188; id., Fl. 4 (1924) 365; Henders.,
J. Mal. Br. R. As. Soc. 17 (1939) 82.
Endemic to Malaya, uncommon; often on limestone but also on the mountains.
Recorded from Langkawi, Gua Ninek, and Batu Bayan in Kelantan, and on Gua
Tipus in Pahang.
72
Gard. Bull. Sing. 36(1) (1983)
NOTE
TYPHACEAE
Typha angustifolia L.
A very widely distributed plant found throughout the world between the arctic cir-
cle and lat. 35 degrees South; in marshy places; not uncommon in Malaya in such
places. Recorded from swampy ground at the base of limestone cliffs near Ipoh,
Perak.
ZINGIBERACEAE
Ii; WLeaves spirally arranged; sheathsitubularn) “epee eerie eee eile ete eee
Leaves arranged in 2 opposite vertical rows; sheaths open on the side opposite the blade ...
2 sinflorescencerat theyapexcols leatyaSDOOtS meee ene eer ere renee nee Costus speciosus
Inflorescence on short leafless shoots at ground level .................... Costus globosus
3. Staminodes (infertile stamens) on either side of the lip petaloid, conspicuous; free or adnate to
tHE: LP) oss cisterns S) pirates wrsitns eb ae Sue ole. DE Slate eteae eS TOOT oe no eT
Staminodes (infertile stamens) never petaloid, usually reduced to short, linear appendages or
small teeth ‘atithe baseiof the lips aaa ene oe or enor nOr cee er iene
4. Lip and filament joined for some distance above the insertion of petals and staminodes; filament
10.
IE
12.
13)
much longer than lip’ s.. 2.6 sc5 een coetns clei cee ovoleere o cucene ere oetsenete Tciche anise erent eens
Lip and filament not so joined, filament usually shorter than lip .....................05-
I eee Mee Farrel erecta Glc Hiarol OA CCIE iO) Read rc CHGS SORENSON che cho EOI Globba patens
Anthers with 2 appendages; leaves usually more than 7, narrowly elliptic, 19-29 by 3-5.5 cm
. Appendages of anther attached to the base of anther, their bases extending up along the side
(0) i 04 (=) Galen eee eee ae chs one Oe A Cola CO Ae OO. cThOD Of Dodo omo ote Globba fasciata
Appendages of anther spreading from about the middle of each side of the anther; their bases
spreading along the whole length of each side of the anther .............. Globa albiflora
= (Leafy stemsito:2smi tallies. ...a,.cck eae earn eee eine Zingiber spectabile
Leafy, ‘stems. short, less) than 15cm) s.-5-..:cr prcite ution ae en eta Tae een
. Bracts in 2 opposite rows, alternate and overlapping ................ Boesenbergia curtisii
Bracts not in 2:OppOSitE TOWS. %ccc.cc.c/scccmtesstopsiale a = atevertorenetay ater etate ee teha eed ereue key cele evel Si eROrey Nene
. Petiole and sheath to 6 cm long, scape enclosed by leaf-sheaths; anther-crest narrow, spathulate
ee ae eee ee ee nent Oo Fact bond TAdot. 7 oeae mo uRdaueaD Kaempferia pulchra
Petiole and sheath longer, scape exerted beyond leaf-sheaths; anther-crest as wide as long, or
Nearly SOnsjascsa acc ee eee Ee ae OSI ort oon Ob ao b Kaempferia elegans
Inflorescence)terminallion! ai leafysstem=s sae eee eee iene ree Catimbium speciosum
Inflorescence not terminal but on a leafless peduncle from the rhizome or from the base of a
C0) 1010) ee ee nnn on ano as loyeoen oddncod doocosdabe ands veoLdac
Inflorescence surrounded at the base or covered all over by large overlapping sterile bracts; the
floral bracts ‘smaller: «0... 6.0 53 we. nende ret eae aE tere are iotessitiesy Ae meer ic heereen toe
Inflorescence on a long erect peduncle 40-80 cm, a common forest species, very rare on limestone
er cnr a nar Go cnr On oan'S OF nocd 66.00 bamela Soo Phaeomeria maingayi
Inflorescence on a submerged short peduncle; inflorescence at ground level ...............
Bracts: large, tov8:by S.cmy ac ccw.ciics caicn ee ee eee oO eee Achasma triorgyale
Bractsrsmallers usually,less: thant 2.oicimewides-wereme creel teicheriieierereicieie erties reentrant
10
11
15
Limestone hill flora of Malaya IV 73
14. Petals longer than sepals, dorsal petal about 1 cm wide ........... Achasma macrocheilos
Petals as long as sepals, dorsal petal about 0.5 cm wide. ......... Achasma megalocheilos
15. Inflorescence with only 2 to 3 flowers, short. Fairly common in forest and waste grounds but
MeTMaFALCTOMPUIMIESEONG Fis fe > cise. -eisteid- seiacieie = ode ciein« Bldem tans wie's Amomum biflorum
Inflorescence with numerous flowers, elongate, narrowly cylindric when old, with persistent,
buff-coloured, papery bracts. Not uncommon on northern limestone .. Amomum testaceum
Achasma macrocheilos Griff., Notul. 3 (1851) 429; Holtt., Gard. Bull. S. 13 (1950)
188.
Hornstedtia macrocheilos Ridl., Fl. 4 (1924) 271.
H. metriocheilos Ridl., op. cit. 271; Henders., J. Mal. Br. R. As. Soc. 17 (1939)
80.
A common species in lowland forest. According to Henderson l.c., recorded from
limestone in the Ipoh area, Perak.
Achasma megalocheilos Griff., Notul. 3 (1851) 426; Holtt., Gard. Bull. S. 13 (1950)
191.
Hornstedtia megalocheilos Ridl., Fl. 4 (1924) 270.
Achasma triorgyale (Bak.) Holtt., Gard. Bull. S. 13 (1950) 186.
Amomum triorgyale Bak., F.B.I. 6 (1892) 237.
Hornstedtia triorgyalis Ridl., Fl. 4 (1924) 269.
Endemic to Malaya, not common; recorded from limestone in Perak.
Amomum biflorum Jack, Mal. Misc. 1 (1820) 2; Bak., F.B.I. 6 (1892) 240; Holtt.
Gard. Bull. S. 13 (1950) 199.
Elettariopsis pubescens Ridl., Fl. 4 (1924) 275.
Endemic and fairly common in Malaya in lowland forest and waste grounds.
Once recorded from limestone.
Amomum testaceum Ridl., J. Str. Br. R. As. Soc. 32 (1899) 135; id., Fl. 4 (1924)
266; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 79; Holtt., Gard. Bull. S. 13
(1950) 205.
Leafy shoots 2-4 m tall. Leaves to 60 by 10 cm, base narrowly cuneate, sessile. In-
florescence elongating, narrow cylindrical, bracts persistent, papery, and buff-
coloured.
Distributed in Borneo; in Malaya common especially in the North, very often on
or near limestone, but not confined to it.
Boesenbergia curtisii (Bak.) Schltr. Fed. Rep. 12 (1913) 316; Holtt., Gard. Bull. S.
13 (1950) 112.
Gastrochilus curtisii Bak., Bot. Mag. (1894) t. 7363; Ridl., Fl. 4 (1924) 249;
Henders., J. Mal. Br. R. As. Soc. 17 (1939) 79.
Leafy stems short, to about 5 cm tall. Leaf to 40 by 12 cm, slightly asymmetric,
elliptic; base cuneate. Bracts numerous, 4-5 cm long, narrow.
74 Gard. Bull. Sing. 36(1) (1983)
Endemic to limestone in Malaya; common in the North, growing on soil and from
rock crevices in part shade.
Catimbium speciosum (Wendl.) Holtt., Gard. Bull. S. 13 (1950) 152.
Alpinia nutans Andr., in Ridl., Fl. 4 (1924) 277.
Costus globosus Bl., Enum. Pl. Jav. (1827) 62; Ridl., Fl. 4 (1924) 256; Holtt., Gard.
Bull. 13 (1950) 243.
C. kingii Bak., in Ridl., sp. cit. 257.
C. velutinue Ridl., op. cit. 257.
Costus speciosus (Koen.) Sm., Trans. Linn. Soc. 1 (1800) 249; Ridl., Fl. 4 (1924)
256; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 79.
Globba albiflora Ridl., J. Str. Br. R. As. Soc. 32 (1899) 96; id., Fl. 4 (1924) 237.
Holtt., Gard. Bull. S. 13 (1950) 31; Lim, Notes R. Bot. Gard. Edin. 31 (1972)
263.
Plant to 80 cm tall. Leaves 10-16, narrow, to 24 cm by 3-3.5 cm. Inflorescence
20-30 cm long; flowers with white corolla.
Endemic and found only on Penang Hill.
var aurea Holtt., l.c.
This differs from the typical form by having orange flowers and the longer lip
1 cm instead of about 0.6 cm long. Recorded once from Gua Lambok limestone,
Kelantan (Henderson 29115).
Globba fasciata Ridl., J. Str. Br. R. As. Soc. 57 (1910) 101; id., Fl. 4 (1924) 136;
Holtt., Gard. Bull. S.\13 (1950) 28; Lim, Notes R. Bot. Gard. Edin. 31 (1972)
263.
Plant 30-50 cm tall. Leaves 5-8; 6-21 by 1.5-6 cm. Inflorescence 20-60 cm long.
Flowers with orange corolla.
Endemic. Until recent collections from limestone, this species was known only
from the type collection, Ridley 14415, from ‘banks of woods by the Temengoh
river’. The correct spelling of Ridley’s ‘Temengoh’ is —Temengor’. This is in Upper
Perak and as far as I know is not a limestone locality.
Recently (1970 & 1971) this species was collected from Gua Musang and Batu
Neng (about 2 miles to the south west of Gua Musang), both limestone hills in Ulu
Kelantan. The Gua Musang collections (Chin 1420 & Stone 9524) were from a small
population of 6-10 scattered plants at and around the opening of the large cave half-
way up the hill, on the side facing Gua Musang town. They were growing in small
pockets of accumulated soil and humus and from crevices in limestone ledges in this
fairly sheltered locality which is at about 100 m elevation. The Batu Neng specimen
(Chin 1543) was solitary and growing in a similar sheltered situation in a small
shallow gully on the lower half of the outcrop.
It may be interesting to note that this new locality for the species is about 50 miles
to the south-east of the type locality in Upper Perak and separated from it by the
Limstone hill flora of Malaya IV 75
main range of mountains which are all well above 1500 m. This species is probably
more common than originally thought.
Globba patens Miq., Fl. Ind. Bat. Suppl. (1860) 613; Lim, Notes R. Bot. Gard.
Edin. 30 (1972) 259.
G. aurantiaca Miq., in Ridl., Fl. 4 (1924) 239; Henders., J. Mal. Br. R. As. Soc.
17 (1939) 80; Holtt., Gard. Bull. S. 13 (1950) 36.
Plant to 60 cm tall. Leaves 3-6, broadly elliptic, 19-24 by 6.5-10 cm. Inflorescence
15-30 cm long; flowers with orange corolla.
Distributed in Sumatra; not uncommon in lowland and mountain forest and with
only one record from limestone.
Kaempferia elegans Wall., Cat. (1832) 6593; Hk.f., F.B.I. 6 (1890) 222; Ridl., Fl.
4 (1924) 245; Henders., J. Mal. Br. R. As. Soc. (1939) 80.
Kaempferia pulchra Ridl., J. Str. Br. R. As. Soc. 32 (1899) 107; id., Fl. 4 (1924)
245; Henders., J. Mal. Br. R. As. Soc. 17 (1939) 80.
Stem short, leaves 2-3, 8-4 cm, asymmetric, elliptic. Scape of inflorescnce enclos-
ed by leaf-sheaths.
Distributed in peninsular Thailand; in Malaya apparently restricted to limestone
in the extreme North. Very similar to K. elegans, but slightly smaller in size. There
is supposedly some floral differences but this is referred to with some doubt. More
specimens are needed for verification.
Phaeomeria maingayi (Bak.) K. Schum., Pflzr. Zingib. (1904) 266; Ridl., Fl. 4
(1924) 272; Holtt., Gard. Bull. S. 13 (1950) 180.
Amomum maingayi Bak., F.B.I. 6 (1892) 180.
Zingiber spectabile Griff., Notul. 3 (1853) 414; Ridl., Fl. 4 (1924) 258; Henders.,
J. Mal. Br. R. As. Soc. 17 (1939) 80; Holtt., Gard. Bull. S. 13 (1950) 56.
Endemic to Malaya, found throughout in lowland forest, recorded once from
limestone.
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78 Gard. Bull. Sing. 36(1) (1983)
Index to Specimens
The Index to Specimens is a list of all specimens examined. Species names are arranged in the
alphabetical order of the genus and then of their epithets. The order of citation is: species, collector(s)’
name(s) and collection number(s), all printed in italics. A full list of collectors in this index is found in
Part I of this paper (Gard. Bull. vol. xxx (1977) 168-170). Where a species is represented by more than
one specimen and collected by different persons, the collectors’ names are arranged alphabetically. Only
collectors’ surnames or, in their absence, their principal names are cited. A name, if followed by ‘er a/.’,
indicates that the specimen has been collected by more than two individuals. When a collecting number
is absent, the date of the collection, which is printed in roman is cited instead. Months are indicated in
Roman numerals; Twentieth Century years are abbreviated to, e.g. ‘41 (= 1941) but Nineteenth Century
ones are in full. The few specimens with neither number nor date are listed as ‘s.n. & s.a.’ (since numero
& sine annum), followed (if available) by an accession number prefixed by the code of the Herbarium
which houses them. KEP and FRI are herbarium codes for the Forest Research Institute, Kepong, KLU,
for the University of Malaya, and SING, for Singapore.
Abacopteris urophylla (Wall.)Ching: Chin 588,710. Abdominea minimiflora (Hk.f.)J.J.S.: Carr 9;
Ridley xii 1896. Abutilon indicum (L.)Sweet: Henderson 22994. Acalypha lanceolata Willd.:
Molesworth-Allen 4847. Acampe longifolia Lindl.: Corner 19 xi 41. Achasma megalocheilos Griff.:
Burkill 6292. A. triorgyale (Bak.)Holtt.: Curtis 3317. Acrotrema costatum Jack.: UNESCO 202.
Actephila excelsa (Dalz.)M.A.: Henderson 19521; Ng FRI 1799; Whitmore FRI 4239. A. ovalis
(Ridl.)Gage: Best 21204; Corner 28 xi 41; Henderson 23005, 29111; Kiah 35247; Stone 6937. Adenia
nicobarica (Kurz)King: Boey 541; Chin 525, 1745; Stone 6991. Adenoncos major Ridl.: Henderson
22257, 22570; UNESCO 291, 433. A. parviflora Ridl.: Chin 418 A; Kelsall i 1891. A. sumatrana J.J.S.:
Chin 418 B; Dransfield 914; Henderson 22447, 22576; Nur 34401; Ridley 1891. Adenosma capitatum
Benth. ex Hance.: Corner 37825; Henderson 29103. Adiantum malesianum Ghatak: Burkill 13933; Chin
84; 319; Corner & Henderson 23009; Curtis xii 1895; Evans 266; Haniff 4048; Henderson vii 29; Johnson
s.n. KLU; Kiah 35240; King’s coll. 8351; Merton 4102; Samat 128; Shimizu & Fukuoko M14134; Sinclair
40058; Stone 5912, 6572, 7299, 8979. A. soboliferum Wall. apud HK.: Haniff & Nur 7523; Holttum
15145; Kiah 35241. A. stenochlamys Bak. Corner 21 xi ‘41, 22 xi ‘41; Henderson 29706. A. tenerum
Sw.: Chin 357, 812. A. zollingeri Mett. ex Kuhn.: Corner & Henderson 22818. Aerides odoratum Lour.:
Chin 1274A; Gimlette 12 viii ‘17. Aeschynanthus parvifolia R.Br.: Chin 1441, 1812; Stone 5889;
UNESCO 239, 440. A. radicans Jack: Nur 34400; UNESCO 440A. Agathis dammara
(Lambert)L.G.Rich.: Loh FRI 17201. Aglaia argentea Bl\.: Stone 9139; UNESCO 503. A. odoratissima
Bl.: Henderson 19553; King’s coll. 10710; UNESCO 64. Aglaonema costatum N.E.Br.: Henderson 21
xi 734. A. oblongifolium Schott.: Burkill 6301; Chin 1767; Henderson 22879, 29145; UNESCO 510.
Agrostistachys gaudichaudii M.A.: Chin 895, 897; Ogata 110194; Samsuri & Mahmud 612.
Agrostophyllum bicuspidatum J.J.S.: Chin 90, 453, 1142; UNESCO 327. A. hasseltii (Bl.) J.J.S.: Chin
1160. Alchornea rugosa (Lour.)M.A.: Stone 6945. Aleurites moluccana (L.) Willd.: Chin 560. Allophylus
cobbe (L.)Raeusch. var. glaber Corner: Chin 109, 1398A, 1590; King’s coll. 8206. A. cobbe (L.)Raeusch.
var. villosus Corner: Chew 189; Kiah 35316. Alocasia denudata Engl.: Chin 694; Henderson 23836;
UNESCO 481. A. lowii Hk.f. Boey 530; Chin 485, 834; Curtis x 1894; Henderson 22851, 23024; Ridley
1897; Sinclair 40071. Alstonia scholaris (L.)R.Br.: Chin 977. Alyxia anqustifolia Ridl.: Chin 385, 571.
A. pumila Hk.f.: Chin 1616; Henderson 19466; UNESCO 305. A. selangorica K. & G.: Ridley 8558.
Amaracarpus saxicola Ridl.: Chin 1035, 1180, 1181, 1457; Ridley 11884. Amomum biflorum Jack.: Chin
730. A. testaceum Ridl.: Kiah 35236, 7 v ’38; Ridley 13122. Amorphophallus carnosus Ridl.: Curtis s.n.
& s.a. (SING); Ridley 15200. A. haematospadix Hk.f.: Corner 19 xi ’41; Holttum 23 viii ’25. A. prainii
Hk.f.: Chin 22; Henderson 19467; UNESCO 438. A. variabilis Bl.: Curtis 2815. Amydrium humile
Schott: Chin 395, 1143; Corner & Nauen 25 xi ’41; Furtado B. 4 vi ’37; Henderson 22995; Nur 3 xi ’37;
UNESCO 360, 686. Anadendrum latifolium Hk.f.: Henderson 19600. A. marginatum Schott: Henderson
25013; Ridley 20 vi 1889, 23 vi 1889. A. montanum Schott: Chin 727; Ridley 8172, 23 vi 1889, 13 xii ’20.
Anaxagorea javanica Bl.: Cockburn FRI 10563; UNESCO 42. Andrographis tenuiflora T. Anders.: Cor-
ner 19 xi ’41; Corner & Nauen 25 xi ’41; Curtis 2578; Henderson 21370; Holttum 15120. Aneilema
nudiflorum Br.: Samat 3. Antidesma japonicum Sieb. et Zucc.: Chin 1509 A; Corner 30 xi ’41; Kiah
35233; Sinclair 9865. A. montanum Bl.: Stone 6944. A. tomentosum BI.: King’s coll. 4808; Loh FRI
17195. Antrophyum callifolium Bl.: Chin 1613; Henderson 22466; UNESCO 247, 579. A. parvulum Bl.
Burkill 13932; Chin 143, 688, 358, 1116, 1182, 1475; Henderson 19699, 22248, 22260, 23890, 25024,
Limstone hill flora of Malaya IV 79
29681; Holttum 8924; Molesworth-Allen 1504; Nauen 38013; Samat 522; Shimizu & Stone M 13724;
Stone 6571, 7456, 8822A; Whitmore FRI 4041. A. semicostatum Bl.: UNESCO 634. Apluda mutica L.:
Chin 522; Corner & Nauen 37842; Henderson 28938. Aporuellia sumatrensis C.B. Clke var. ridleyi C.B.
Clke: Allen 4339; Burkill 6370; Chin 479, 1250; Curtis 2364; Ridley 8213, 23 vi 1889, xii ’20; Stone &
Wycherley 8986. Aporusa stellifera Hk.f.: King’s coll. 7102. Appendicula anceps Bl|.: Ridley 8130. A.
cornuta Bl.: Chin 361; Stone 5919. A. torta Bl.: Chin 549, 1566; Kadim & Mrs. Molesworth-Allen K506;
Stone 9514; UNESCO 36. A. undulata Bl.: Henderson 22457. Arachnis flos-aeris (L.)Rchb.f.: Hender-
son 22452; UNESCO 160. Aralidium pinnatifidum Miq.: UNESCO 79. Arcypteris irregularis (Pr.)Holtt.:
Chin 589. Ardisia andamanica Kurz: Chin 763; Samsuri 547. A. biflora K.et G.: Ridley 11933, ii ’04.
A. colorata Roxb.: Chin 477. A. crenata Sims: Chin 532. A. fulva K.et G.: Nauen 38119. A. kunstleri
K.et G.: Corner 38145. A. lanceolata Roxb.: Burkill 6352; UNESCO 665. A. oxyphylla Wall.: Henderson
29712; Kiah 35420; UNESCO 411, 508, 590. A. pendula Mez: Ridley 14935. A. platyclada K.et G.: King’s
coll. 8136. A. solanacea Roxb.: Henderson 21394. A. tahanica K.et G.: Henderson 19692. A. vaughani
Ridl.: Best 21226; Kiah 35383, 2 v ’38. Areca triandra Roxb.: Henderson 29081. Arenga hookeriana
(Becc.) Whitm.: Chin 1547, 1600, 1601; Cockburn FRI 10580; Henderson 29673; Whitmore FRI 4032.
A. pinnata (Wurmb.)Merr.: Whitmore FRI 4267. A. westerhoutii Griff.: Chin 1463; Stone 8825.
Argostemma diversifolium Ridl.: Ridley 14479. A. inaequilaterum Benn.: Chin 2108; Ridley 8233; Stone
8949. A. pictum Wall.: Holttum 17402. Argyeia maingayi (C.B.Clke)Hoogl.: Chin 1544; Stone 7313;
UNESCO 28, 496. A. mollis (Burm.f.)Choisy: Chin 1735, Henderson 29108, Nauen 37953. Arisaema
fimbriatum Mast.: Corner 13 xi ’41, 19 xi 41, 20 xi ’41; Ng FRI 1605; Nur 25160, 34376; Stone 8936;
UNESCO 427. A. roxburghii Kunth: Chin 394, 1614; Henderson 25214; Nur 34394. Artabotrys gran-
difolius King: King’s coll. 7222. Arthraxon prionodes (Steud.)Dandy: Corner & Nauen 25 xi ’41.
Ascochilopsis myosurus (Ridl.)Carr: Henderson 25246. Asparagus racemosus Willd.: Curtis 1674.
Asplenium adiantoides (L.)C. Chr.: Chin 108, 1451, 1607, 1768; Corner & Henderson 22842, 23089;
Henderson 22266; Molesworth-Allen 4429; Ridley 14756; Sinclair 9873; UNESCO 246, 261. A.
macrophyllum Sw.: Chin 4, 320; Henderson 6 x ’31; Mahmud 29 v ’70; Poore 347; Ridley xii 1896; Samat
523; Samsuri & Mahmud 580; Shimizu & Stone M 13719, T 14376; Stone 7294, 8833. A. pellucidum
Lamk.: Chin 544, 1075, 1450; Corner & Nauen 25 xi ’41; Henderson 19529, 19693, 25236. A. phyllitidis
Don: UNESCO 244, 471, 656. A. salignum Bl.: Chin 3; Henderson 19455, 25207; King’s coll. 8130, 8424;
Ng FRI 1612; Samat 521; Stone 8831; UNESCO 256, 526. A. squamulatum Bl.: Chin 8, 137, 321, 1415;
Henderson 25076; Ridley xii 1896; Shimizu & Stone T 14354; Stone 7296, 8843. A. unilaterale Lamk.:
King’s coll. 7188; Molesworth-Allen 1505A; Ridley vii 1897. Atalantia monophylla DC.: Chin 516; Cor-
ner 37816; Henderson 29161; Kiah 35570; Stone 9085. A. roxburghiana Hk.f.: Chin 535, 1680, 1807;
Chin & Mahmud 1308; Loh FRI 17197; Stone 8796, 9388; Symington KEP 37404. Athyrium cordifolium
(Bl.)Copel.: Chin 682. A. esculentum (Retz.)Copel.: Chin 529. A. montanum (v.A.v.R.)Holtt.: Chin
607. A. pinnatum (Blanco)Copel.: Henderson 22422. Athyrium prescottianum (Wall.)Holtt.: UNESCO
517. Axonopus compressus (Sw.)Beauv.: Chin 106.
Baccaurea lanceolata (Miq.) M.A.: Henderson 23767. Balanophora fungosa Forst.: Henderson 22810.
Barleria siamensis Craib.: Henderson 29150. Barringtonia asiatica (L.)Kurz: Henderson 29149. B.
fusiformis King: Henderson 23796; Ridley 8284. Bauhinia acuminata L.: King’s coll. 8288. B. pottsit
G.Don: UNESCO 139, 653. Becheria parviflora Ridl.: Ridley 8250. Begonia foxworthyi Burk.: Hender-
son 25249, 29672; UNESCO 29, 234, 365. B. guttata Wall.: Holttum 15127. B. kingiana Irmsch.: Burkill
6300; Burkill & Haniff 13922; Chin 845, 1958; Curtis s.n. & s.a. (SING); Henderson 29705; UNESCO
395. B. nurii Irmsch.: Chin 153, 1121, 1406. B. phoeniogramma Ridl.: Burkill 6362, 6366, 6367; Chin
1668, Kho | vii ’74; Ridley 2882, 8281; Stone 8971; UNESCO 26. Beilschmiedia lumutensis Gamble:
UNESCO 504. Berrya cordifolia (Willd.)Burret: Henderson 29148. Bidens pilosa L.: Chin 1093.
Biophytum adiantoides Wight ex Edgew. et Hk.f.; Kiah 35246. Blechnum finlaysonianum Hk. et Grev.:
Chin 672. Boea acutifolia Ridl.: Chin 1769; Henderson 23014; Stone 6916. B. brachycarpa Ridl.: Hender-
son 19668; UNESCO 303, 560, 593. B. caerulescens Ridl.: Chin 441, 1066; Henderson 25048; King’s coll.
7062, 7175, 8276; Mills & Henderson 15062, 15068a, Ng FRI 1610; Stone 7449; UNESCO 154, 264, 339,
429; Whitmore FRI 4253. B. divaricata Ridl.: Corner 20 xi ’41; Stone 6915. B. lanata Ridl.: Alphonso
& Samsuri 89; Chin 1727; Henderson 21400; Holttum 15139; Nauen 37894. B. minutiflora Ridl.: Hender-
son 22392. B. paniculata Ridl.: Alphonso & Samsuri 88; Chin 327, 393, 1251; King’s coll. 7026, 8271;
Mills & Henderson 15068; Sinclair 9872, 40072. B. parviflora Ridl.: King’s coll 7108..B. suffrutocosa
Ridl.: Corner 37811. B. treubii Forbes: Henderson 19459, B. verticillata Ridl.: Best 21280; Chin 33, 326,
411, 1252; Henderson 19459, 2 vi ’30; Kiah 9 v ’38; Nauen 38021; Ng FRI 1628; Nur 34366; Stone 8935;
Stone & Mahmud 9399. Boehmeria nivea Hk. & Arn.: Kiah 35427. Boerhaavia chinensis (L.)Aschrs. &
80 Gard. Bull. Sing. 36(1) (1983)
Schweinf.: Henderson 23115, Ridley 15153. Boesenbergia curtisii (Bak.)Schl.: Best 21259; Chew 133;
Chin 512; Corner 13 xi ’41; Curtis 2678; Haniff & Nur 7494; Henderson 22874, 25015, 25086, xi ’29.
Bombax anceps Pierre: Corner 19 xi ’41; Henderson 29098. Borassodendron machadonis (Ridl.)Becc.;
Whitmore FRI 4031. Brassaiopsis polycantha (Wall.)R.N. Ban.: Henderson 25212. Breynia vitis -idaea
(Burm.f.)Fisher: Chin 1385; Stone 7464. Bridelia ovata Decne: Henderson 21395; Stone 9079. B.
stipularis (L.)Bl.: Corner 38131. B. tomentosa Bl.: Best 21273; Corner & Nauen 25 xi ’41; Stone &
Wycherley 8978. Buchanania sessilifolia Bl.: UNESCO 201. Bulbophyllum apodum Hk.f.: Chin 121,
458; UNESCO 212. B. concinnum Hk.f.: Allen xii 56; UNESCO 297. B. fenestratum J.J.S.: Carr 263;
UNESCO 133, 362. B. flammuliferum Ridl.: Allen xii ’56. B. lilacinum Ridl.: Corner 38133; Henderson
22804, 29088. B. membranaceum T.et B.: Chin 548, 1115. B. pulchellum Ridl.: Stone 7455. Burmannia
championii Thw.: Chin 1829; UNESCO 50. B. lutescens Becc.: Henderson 21383. Buxus holttumiana
Hatusima: Chew 181; Kiah 35239. B. malayana Ridl.: Allen 5 ii °57; Chin 54, 376, 1162; Loh FRI 17226;
Mills & Henderson 15078; Nur 34382; Stone 5906, 8944; UNESCO 12 & 180. B. rupicola Ridl.: Curtis
2662.
Caesalpinia crista L.: Henderson 29110. Calamus balingensis Furt.: Furtado 33073. C. concinnus
Mart.: Haniff & Nur 7087. C. ornatus Bl. var. horridus Becc.: Chin 605. C. siamensis Becc. var.
malaianus Furt.: Henderson 23028. Calanthe ceciliae Rchb.f.: Carr 14; Henderson 22354, 25002. C.
rubens Ridl.: Corner 20 xi ’41; Curtis 2182, 1890; Ridley 14976. C. triplicata (Will.)Ames.: Chin 893,
1033; Henderson 22275, 23761; UNESCO 227. C. vestita Lindl.: Ridley xii 1896. Callicarpa angustifolia
K. & G. Chin 341, 443, 1067; Corner 38127; Henderson 19597, 22265, 23757, 23805, 25049, 29152; Ng
FRI 1637, 1798; Nur 34369; Ridley 8207; Stone 5896, 6926; UNESCO 146, 605. C. lanata L.: Chin 809;
Henderson 23754; UNESCO 63. Camarotis apiculata Rchb.f.: Haniff 10331; Henderson 22449; Ridley
1891. Cananga odorata (Lamk.) Hk.f. et Th.: King’s coll. vii 1883. Canarium perlisanum Leenh.: Kiah
35311. C. pilosum Benn.: Loh FRI 17208. C. pseudodecumanum Hochr.: Loh FRI 17257. Canscora pen-
tanthera C.B. Clke: Burkill 2554; Chin 340, 437, 803, 1534; Hardial & Sidek 476; Henderson 19458,
21380, 23118; Kiah 35371; Nauen 38026; Ng FRI 1632; Nur 34396; Sinclair 9868, 40065; Spare 36318,
Stone 7460; UNESCO 285, 446. Canthium aciculatum Ridl.: UNESCO 388. C. didymum Roxb.: Chin
27, 1377. Stone 6923. Capparis diffusa Ridl.: Chin 870, 1069, 1104; Ridley 15174; Stone 6988. C.
pubiflora DC.: Henderson 22319. Capsicum frutescens L.: Chin 939, 1100. Carallia brachiata
(Lour.)Merr.: Chin 980; Kiah 35377; Soepadmo ix 68. Cardiopteris javanica Bl.: Henderson 22820.
Carex breviscapa C.B. Clke: UNESCO 56. C. malaccensis C.B. Clke: Chin 1712, 1724. C. perakensis
C.B. Clke: UNESCO 277. C. speciosa Kunth: Chin 436, 1378; UNESCO 314, 568. Caryota mitis Lour.:
Chin 553; Chin & Badaruddin 1691; Henderson 22486, 29068; Ridley 13452. Casearia capitellata Bl.:
Chin 755; Samsuri 543. C. grewiaefolia Vent.: UNESCO 538. Cassia timoriensis DC.: Best 21276. Catim-
bium speciosum (Wendl.)Holtt.: Ridley 14776. Cayratia japonica (Thunb.)Gagn.: Nur 3 xi 737. C.
mollissima(Wall.)Gagn.: Chin 1428, 1513; Henderson 23751. Celtis philippensis Blanco: Chin 1758,
1783; Corner 17 xi ’41; Kiah 35413; Loh FRI 17176; Stone 6982; UNESCO 214. Centotheca lappacea
(L.)Desv.: Chin 58, 105, 1380; Corner 22 xi’41; Henderson 29189. Centranthera hispida R.Br.: Hender-
son 29057. Ceratostylis pendula Hk.f.: UNESCO 455. Cheilanthes farinosa (Forsk.) Hk.f.: Chin 433;
Nauen 38027. Chionanthes calcicolus (Kerr)Kiew: UNESCO 346. Chirita caliginosa C.B. Clke: Burkill
2253, 2258; Chin 331, 390; Corner 37832; Curtis 3109; Hardial & Sidek 477; Henderson 22419, 25033,
25223; King’s coll. 7028; Ng FRI 1629; Nur 34389; Samsuri & Mahmud 560; Sinclair 10732, 40066; Smith
435; Stone 5897, 6570, 8934; Wycherley & Stone 8988. C. hamosa R.Br.: Henderson 29185. C. rupestris
Ridl.: Corner & Nauen 25 xi’41; Henderson 22816; Nauen 38120A. C. sericea Ridl.: Burkill 2552, 6284;
Corner & Nauen 25 xi ’41; Curtis 3131; Sinclair 9844. C. viola Ridl.: Chin 505; Corner 13 xi ’41; Curtis
2570; Henderson 28931, 29185; Holttum 17433; Keng et al. 6142 & 6239; Molesworth-Allen 15 xii ’50;
Nauen 38120B; UNESCO 268. Chloranthus elatior R. Br. ex Link: Chin 585, 676, 1666. Chlorophytum
orchidastrum Lindl.: Burkill & Haniff 13910; Chew 177; Curtis 3211. Chrysopogon aciculatus
(Retz.)Trin.: Chin 377. C. fulvus (Spreng.)Chiov.: Haniff 649. C. orientalis (Desv.)A. Camus: Corner
& Nauen 37840; Henderson 29058. Cinnamomum iners Reinw. ex Bl.: Corner & Nauen 25 xi ’41;
UNESCO 487. Cissus discolor Bl.: Chin 1755; Corner 38138; Corner & Nauen 25 xi ’41; Henderson
21472. C. glaberrima (Wall.)Steud.: Henderson 29067; Ng FRI 1631; Nur 34399. C. hastata Miq.: Chin
110, 404; Henderson 22256, 23807, 25050; Nur 8472; Stone 5909, 7306, 9493; UNESCO 141, 236, 558.
C. novemfolia (Wall.)Planch.: Burkill & Haniff 13938. C. pyrrhodasys Miq.: Chin 490; Corner 20 xi ’41;
UNESCO 242. C. repens Lamk.: Chew 188; Chin 1741; Corner 37804; Henderson xi ’34; Holttum 22
viii ’25. C. rostrata (Miq.)Planch.: Chin 398, 1384; Corner 38139; Stone 8836. Citrus macroptera
Montr.: Loh FRI 17183. C. medica L.: UNESCO 1017. Cladogynos orientalis Zipp. ex Span.: Chew 132;
Limestone hill flora of Malaya IV 81
Chin 907, 1487, 1561; Henderson 29678; Loh FRI 17163; Whitmore FRI 4236. Claoxylon longifolium
(Bl.)Endl. ex Hassk.: Henderson 21391; Kerr 21700. Clausena excavata Burm.f.: Chin 345, 438, 818,
1086; Corner & Nauen 25 xi 41. C. harmandiana (Pierre)Guill.: Henderson 23820. Cleidion javanicum
BI.: Chin 57; Mahmud 9 v ’70; Whitmore FRI 4240. Cleistanthus decurrens Hk.f.: King’s coll. 11285;
Loh FRI 17162. C. gracilis Hk.f.: Chew 186; Chin 103, 826, 898, 1057; Corner 20 xi ’41; Henderson
23830; Holttum 15119; Loh FRI 17165; Molesworth-Allen 4292. C. hirsutulus Hk.f.: King’s coll. 5870.
C. kingii Jabl.: Burkill 6270; Henderson 23789, 23801. C. macrophyllus Hk.f.: Chin 859; Samsuri &
Mahmud 603. C. polyphyllus Wall.: Chin 25. Clerodendron paniculatum L.: Best 21281; Corner &
Nauen 25 xi ’41; Nauen viii ’41. C. penduliflorum Wall.: Chin & Mahmud 1278; Henderson 19445. C.
serratum Spr.: Chin 29, 796; Chin & Badaruddin 1709; Samsuri & Mahmud 569; UNESCO 330.
Clidemia hirta (L.)Don.: UNESCO 600. Cnesmone javanica B\.: Whitmore FRI 751. C. subpeltata Ridl.:
Chin 330, 1247. Coelogyne asperata Lindl.: Chin 138, 1440; UNESCO 48, 218. C. pallens Ridl.: Allen
27 i ’57. C. pandurata Lindl.: Chin 552; King’s coll. 7176. Coffea canephora Pierre ex Froehner: Chin
914; Samsuri & Mahmud 617. C. malayana Ridl.: Henderson 23808. Coix lacryma-jobi L.: Chin 1520.
Colocasia esculenta (L.)Schott: Mahmud 9 iii’71. C. gigantea (Bl.)Hk.f.: Burkill 2266, 6275; Chin 831A;
Henderson 22331, 22359; Stone 9495. Colona merguensis (Planch. ex Mast.)Burr.: Boey 521; Chin 1763;
Corner 38137; Henderson 21387, 23026; Soepadmo & Mahmud 1216; Whitmore FRI 15065. Colubrina
asiatica (L.)Brongn.: UNESCO 570. Combretum latifolium Bl\.: Chin 832; Samsuri & Mahmud 583. C.
porterianum (C.B. Clke)Wall ex Craib: Burkill 6288. Connarus sp.: Chin 1424; Everett FRI 14391;
Whitmore FRI 4268. Congea vestita Griff.: UNESCO 606. Cordia griffithii C.B. Clke: Henderson 29155.
C. obliqua Willd.: Henderson 29155; Kiah 35418. Corybas mucronatus (Bl.)Schltr: Allen s.n. & S.a.
(SING); Chin 94, 355. Corymborchis rh ytidocarpa (Hk.f.)Holtt.: Molesworth-Allen 4435. C. veratrifolia
BI.: Burkill 6345; Chin 1588. Costus globosus Bl\.: Chin 134, 606, 709. C. speciosus (Koen.)Sm.: Chin
1581, 1754. Cotylelobium malayanum V.S\.: Chin 774. Cratoxylum maingayi Dyer: Chin 113, 1398,
1459; Stone 9496; UNESCO 321; Whitmore 4271. Crinum defixum Kerr.: Henderson 26 xi ’34. Croton
cascarilloides Raeusch.: Chin 150, 815, 1050; Corner 37802; Henderson 23012, 28930; Kiah 35242; King’s
coll. 8419; Ng FRI 5536. Samsuri & Mahmud 576; Stone 6936, 7450. C. laevifolius Bl.: Chin 926; Samsuri
& Mahmud 618. Crypsinus enervis (Cav.)Copel.: UNESCO 103. Cryptocarya griffithiana Wight: Chin
1246. Cryptocoryne affinis N.E.Br.: Chin 1480. C. minima Ridl.: UNESCO 279. C. purpurea Ridl.:
UNESCO 607. Ctenopteris alata (Bl.)Holtt.: Chin 147; UNESCO 59, 361. C. moultoni (Copel.)C.Chr.
et Tard.: Henderson 22253. Curanga amara JSuss.: Burkill 13925. Curculigo latifolia Dryand.: Mill &
Henderson 15074. Cyathula prostrata (L.)Bl.: Chin 869, Stone 8818. Cycas rumphii Miq.: Chin 799,
1148; Stone 9531-A; Whitmore FRI 4232. C. siamensis Miq.: Chin 1788; Henderson 23826; Nauen 38032;
Ng FRI 1606; Ridley iii ’10 (= SING 503403). Cyclea laxiflora Miers: Chin 985; Stone 7451; Whitmore
759. Cyclopeltis crenata (Fee)C.Chr.: Chin 478, 876, 1416; Corner & Nauen 25 xi 41; Holttum 15141;
King’s coll. 8282; Nur 3 xi ’37; Samat 415; Shimizu & Stone T 14391. Cyclosorus extensus (Bl.)Ching:
Sinclair 9869. C. interruptus (Willd.)Ching: Kiah 35250. C. megaphyllus (Mett.)Ching: Phang 15 xi 68.
C. unitus (L.)Ching: UNESCO 262. Cymaria dichotoma Benth.: Fox 10686; UNESCO 555. Cymbidium
dayanum Rchb.f.: Kiah 13 iv ’38. C. finlaysonianum Lindl.: Chin 1814; Corner 19 xi’41. Cymbopogon
calcicola Hubb.: Chin 434, 559, 1106; Henderson 21906, 25224; Kiah 13 v ’38; Mat 36256; Nauen 38031:
Cynoctonum mitreola (L.)Britt.: Curtis 2114; Henderson 28932; Nauen 37954. Cynometra malaccensis
Meeuwen: Chin 835; Loh FRI 17234. Cyperus kyllingia Endl.: Molesworth-Allen 4271. C. trialatus
(Boeck.)Kern.: UNESCO 190. Cyrtandra cupulata Ridl.: Loh FRI 17261. C. lanceolata Ridl.: UNESCO
547. Cyrtosperma lasioides Griff.: Henderson 22506.
Dacryodes kingii (Eng|.) Kalkman ex Leenh.: Henderson 23800. Dalbergia phyllanthoides Bl\.: Hender-
son 25054. D. scortechinii Prain: Henderson 29694; UNESCO 322. Davallia denticulata (Burm.)Mett.:
Chin 518, 1076; UNESCO 15, 627. D. solida (Frost) Sw.: Chin 1603. Debregeasia squamata King ex
Hk.f.: Molesworth-Allen 4880; Ridley viii 1897; Samsuri & Mahmud 594. Decaspermum fruticosum J.R.
et G. Forst.: Allen xii ’56; Chin 55, 124, 374, 1164, 1255; Henderson 22272, 29695; Nur 34373; Ridley
Vii 1893; Stone 5911 & 8943; Whitmore FRI 4242, 4271. Deeringia polysperma (Roxb.)Miq.: Chin 442;
Henderson 19520; Kiah 35362; Molesworth-Allen 4286; UNESCO 271, 597. Dehaasia curtisii Gamble:
Chin 155, 328, 375, 912, 1267, 1389, 1458, 1548; Corner 34389; Stone 5901, 8940, 9515; UNESCO 124;
Whitmore FRI 753. D. longipedicellata (Ridl.)Kosterm.: UNESCO 80. Dendrobium acerosum Lindl.:
Chin 1425; UNESCO 290. D. aloifolium (Bl.)Rchb.f.: Chin 120, 1567; Stone 9507; UNESCO 375. D.
excavatum (Bl.) Miq.: Henderson 23017. D. farmeri Paxt.: UNESCO 315. D. indivisum (Bl.) Miq.: Cor-
ner 17 xi’41, 22 xi’41; Henderson 29089; Stone 6977. D. langkawiense Ridl.: Corner 19'xi ’41. D. leonis
(Lindl.)Rchb.f.: Henderson 22283. D. pumilum Roxb.: UNESCO 414. D. salaccense (Bl.)Lindl.: Chin
82 Gard. Bull. Sing. 36(1) (1983)
129, 1084; Chin & Badaruddin 1705; Henderson 23140, 25074; Nur 25156. D. setifolium Ridl.: Chin
1560. D. spurium (B1.)J.J.S.: Chin 420; Henderson 22282, 25233; Nur 34397, 14x31; Stone & Anderson
8841; UNESCO 372, 445. D. subulatum (Bl.)Lindl.: Dransfield 913. D. tetrodon Rchb.f. ex Lindl.:
UNESCO 444 & 456. Dendrocalamus dumosus (Ridl.)Holtt.: Corner 37839; Corner & Nauen 25 xi ’41;
Kiah 35409; Nauen 38017. D. elegans (Ridl.)Holtt.: Burkill 785; Chin 523; Henderson 29097. Dendroc-
nide sinuata (Bl.)Chew: Molesworth-Allen 4667, 4846, 4879 & 4887. D. stimulans (L.f.)Chew:
Molesworth-Allen 4493, 4718, 4878, 4880. Derris thyrsiflora Benth.: Corner 22 xi ’41. D. uliginosa
(Roxb.)Benth.: Chin 609; Stone & Wycherley 8980. Desmodium rugosum Prain: Corner 37833; Hender-
son 28935. D. umbellatum DC.: Henderson 21 xi ’34. Desmos cochinchinensis Lour.: Henderson 23762;
Molesworth-Allen 4288. D. dasymaschalus (B1.)Saff. var wallichii (Hk.f. et Th.)Ridl.: King’s coll. 4564;
Stone 7487. D. dunalii (Hk.f. et Th.)Saff.: King’s coll. 4483. Dichanthium annulatum (Forsk.)Stapf.:
Corner & Nauen 37859, 25 xi ’41; Nur 34367; Reid 29 x ’50. D. caricosum (L.)A.Camus: Henderson
22270, 6 x ’31. D. mucronulatum Jansen: Chin 379, 460, 1060, 1158, 1271; Ridley 8129, Stone 8938.
Dichiloboea speciosa (Ridl.)Stapf.: Chin 499; Corner 38134; Henderson 29186; Nauen 37956. Dicliptera
rosea Ridl.: Ridley 15050. Digitaria violascens Link: Chin 575. Dillenia indica L.: Henderson 29560;
UNESCO 209. Dimocarpus longan Lour. var. longan Leenh.: Henderson 23812. Dioscorea bulbifera L.:
Chin 488; Stone 8847. D. calcicola Prain et Burk.: Corner & Nauen 25 xi ’41; Stone 7462. D. esculenta
(Lour.)Burk.: Haniff 641; Kiah 35416. D. filiformis Bl.: Burkill 2269, Chew 40996, Henderson 21389,
22882, 23004. D. glabra Roxb.: Corner 19 xi ’41; Henderson 23098, 28939. D. hispida Dennst.: Hender-
son 21 xi ’34. D. polyclades Hk.f.: Burkill & Haniff 13942. D. prainiana Kunth: Burkill 6308. D. prazeri
Prain et Burk.: Henderson 22884. D. pyrifolia Kunth: Burkill 2265; Chin 930; Samsuri & Mahmud 619.
Diospyros adenophora Bakh.: Chin 1154, 1167, 1168. D. buxifolia (Bl.)Hiern: UNESCO 320. D.
cauliflora Bl.: Chin 775, 1551. D. ferrea (Willd.)Bakh.: Chin 126, 1390, 1555, 1592; Corner & Nauen
25 xi ’41. D. frutescens Bl.: Chin 662. D. transitoria Bakh.: Chin 1833. D. undulata Wall. ex G. Don.:
Stone 6935. Diploclisia glaucescens (B1.)Biels: Stone 9398. Dipodium pictum (Lindl.)Rchb.f.: Henderson
22222. Dipterocarpus oblongifolius Bl.: UNESCO 343. Dischidia benghalensis Colebr.: Chin 1438;
UNESCO 144. D. hirsuta Decne: Chew 146; Chin 132, 1037; Stone 7459; UNESCO 131, 233, 489. D.
rafflesiana Wall.: Stone 6983. D. scortechinii K.et. F.: UNESCO 681. D. tomentella Ridl.: Henderson
22802. Distyliopsis dunnii (F.H. Hemsl.)Endress: Chin 109, 1072, 1081; Stone 9492. Distylium stellare
Ktze: Spare 36326. Donax grandis Ridl.: Chin 716, 1545. Doryopteris allenae Tryon: Chin 1270; Ridley
8135. D. ludens (Wall.)J. Sm.: Chin 1726; Corner 17 xi ’41, 19 xi ’41; Corner & Henderson 22815;
Henderson 29066. Dracaena angustifolia Roxb.: Chin 324; Kassim 17 viii ’62; Ng FRI 1593; Stone 7297.
D. congesta Ridl.: Chin 526; Corner 13 xi ’41, 17 xi ’41; Nur 34395. D. curtisii Ridl.: Haniff ix 1900;
Wooldridge vi 1896. D. graminifolia Wall.: Chin 764; Samsuri 548. D. nutans Ridl.: UNESCO 178. D.
porteri Bak.: Chin 1584. D. yuccaefolia Ridl.: Chin 514. Drynaria bonii Christ.: Chin 1764. D.
quercifolia (L.)J. Sm.: Chin 519; Corner 17 xi’41, 22 xi’41; UNESCO 474. D. rigidula (Sw.)Bedd.: Chin
406; Corner 19 xi ’41; Henderson 29069, 29168; Nur 34368; Stone 8925. D. sparsisora (Desv.)Moore:
Chin 1452. Dryobalanops aromatica Gaertn.f.: Chin 660. D. oblongifolia Dyer: Chin 620. Drypetes ner-
vosa (Hk.f.)P. et H.: Scortechini s.n. & s.a. (SING). D. oxyodonta Airy Shaw: Chin 1169, 1387, 1453,
Loh FRI 17174. Dysoxylum arborescens Miq.: Chin 954; Samsuri & Mahmud 633.
Echinodorus ridleyi Steen.: Ridley 8464. Ehretia timorensis D.C.: Chin 336, 777, 1248; Henderson
29115. Elaeocarpus pedunculatus Wall.: Chin 1262. Elastostema curtisii (Ridl.)H. Schrot.: Molesworth-
Allen 4652. E. latifolium (B\.)H. Schrot.: Burkill 13580; Chin 540, 597, 1611; Henderson 19503, 22329,
29083; Sinclair 9857. Eleusine indica (L.)Gaertn.: Chin 573. Endospermum diadenum (Miq.)Airy Shaw:
Henderson 7 vi ’30. Enicosanthum congregatum (King)Airy-Shaw: Ridley 24 vi 1889 (= SING 1367),
xii 1896 (= SING 1368). Ephemerantha fimbriata (Bl.)P.F. Hunt et Summ.: Kiah 35365. E. luxurians
(J.J.Sm.)Hunt et Summ.: Chin 47, 1101. Epipremnum giganteum Schott: Chin 971, 1681; Corner 20
xi ’41; Henderson 23131. Epithema saxatile B\.: Burkill 4223, 6274; Chin 392, 1404, 1759; Corner 20 xi
?41; Corner & Nauen 25 xi’41; Henderson 19406, 22382, 22580, 22823, 22875, 25010, 25205, 29079; Holt-
tum 15125; Keng et al. 6140; King’s coll. 5872, 7046; Molesworth-Allen 4272, 4654; Ng FRI 1795; Sinclair
9846; Stone 8830; Stone & Mahmud 8400; UNESCO 147. Erechtites valerianifolia (Wolf)DC.: Chin 572.
Eria citrina Ridl.: UNESCO 359. E. leiophylla Lindl.: UNESCO 301, 325. E. nutans Lindl.: UNESCO
453. E. pannea Lindl.: Henderson 22225. E. pulchella Lindl.: Chin 164, 421, 1144; Henderson 19465;
Stone 7480; UNESCO 217, 302. E. vestita Lindl.: Chin 1065. Eriobotrya bengalensis Hk.f.: Chin 410,
1152, 1550; UNESCO 308, 466; Whitmore 19 iv ’57. Erismanthus obliquus Wall. ex M.A.: Chin 1061
& 1090. Ervatamia peduncularis K. et. G.: Stone 6574. Erythroxylum cuneatum (Miq.)Kurz: Chin 1048,
1268. Eugenia chlorantha Duthie: UNESCO 157. E. claviflora Roxb.: UNESCO 204. E. pendens Duthie:
Limstone hill flora of Malaya IV 83
Chin 902; UNESCO 66, 71, 507. E. porphyranthera Ridl.: Chin 1031. E. spicata Lamk.: King’s coll.
4674. Eulalia quadrinervis (Hack.)Ktze: Corner & Nauen 37841; Henderson 29051. Eulophia keithii
Ridl.: Curtis 2 collections, s.n., 1892; Ridley 14973, iii ’10; Stone 6928. Euonymus cochinchinensis
Pierre: Chin 1068, 1070, 1082, 1146; UNESCO 347. E. javanicus Bl.: UNESCO 150. Eupatorium
odoratum L.: Chin 550. Euphorbia antiquorum L.: Chin 1749; Henderson 23025, 29167. E. hirta L.:
Henderson 25052. Eurycles amboinensis (L.)Loud.: Ridley 14790. Excoecaria oppositifolia Griff.:
Henderson 23794, 29164, 29675.
Fagraea auriculata Jack: Stone 5915. F. blumei G. Don: Chin 569. F. calcarea Henders.: Henderson
25036. F. carnosa Jack: Chin 367, 566, 1151; Henderson 6 x’31. F. curtisii K. et G.: Chin 910, 967, 1524;
Samsuri & Mahmud 628; Whitmore FRI 4243. Ficus annulata Bl.: Corner 37851. F. binnendykii Miq.:
Corner 31678; Henderson 29697. F. botryocarpa Miq.: Henderson 23771. F. calcicola Corner: Burkill
6283; Chin 32, 370; Chin & Badaruddin 1701; Nur 34388; Symington all KEP 37426, 37451 & 39590;
Whitmore FRI 12162. F. curtipes Corner: Chew 200; Chin 503; Corner 38143; Henderson 29073. F.
deltoidea Jack: Best 21235; Chin 423, 972, 986, 1381, 1589; Corner & Nauen 25 xi’41; Henderson 19701,
29689, 35379; Nauen 38028; Nur 25161; Stone 7465; Whitmore FRI 4258. F. elastica Roxb. ex Hornem.:
Curtis 3305. F. hispida L.f.: Corner 37899; Henderson 23132. F. microcarpa L.f.: Chew 205; Chin 123,
1102, 1149; Corner 38142; Henderson 21384. F. montana Burm.f.: Chew 125; Henderson 22441.
F.oligodon Miq.: Best 21227. F. parietalis Bl.: Henderson 10 x ’31. F. racemosa L.f.: Chew 197. Ficus
sagittata Vahl.: Burkill & Haniff 13945; Henderson 23753. F. scortechinii King: Best 21286. F. semicor-
date B.Ham. ex J.E. Smith: Turnau 1199. F. stricta Miq.: Ng FRI 5873. F. subulata Bl.: Chin 568, 646,
703, 712. F. sundaica Bl\.: Chin 1272, 1810 & 1827. F. superba Miq.: Chew 206; Chin 491. F. tinctoria
Forst.f.: Chew 208; Kiah 35422. F. trichocarpa Bl.: Nur 34387. F. virens Ait.: Henderson 23752, 25752.
Fimbristylis calcicola Kern: Corner 19 xi ’41. F. fusca (Nees)C.B. Clke: Corner & Nauen 37848; Hender-
son 29683; Nur 34378. F. malayana Ohwi: Corner & Nauen 37849; Henderson 29052. F. trichophylla
Ridl.: Corner 19 xi ’41; Corner & Nauen 37847; Henderson 29062, 29683; Nur 34378. Flacourtia
jangomas (Lour.)Raeusch.: Kiah 35255. Forrestia monosperma C.B. Clke: Chin & Mahmud 1275A;
UNESCO 428, 534.
Garcinia cowa Roxb.: Chin 768; Ogata FRI 110202; Samsuri 551. G. eugeniaefolia Wall. ex Anders.:
Kiah 35368; UNESCO 354. G. minutiflora Ridl.: Chin 1418, 1596, 1722; Corner 13 xi ’41, 37855; Curtis
2802; Henderson 22 xi ’34; Kiah 35319. G. montana Ridl.: Ng FRI 1600. G. murdochii Ridl.: Chin 1147,
1253; Loh FRI 17175. G. nigrolineata Planch. ex. T. Anders.: 556; UNESCO 612. G. opaca King: Chin
645; Cockburn FRI 10559; Henderson 19654. Garuga floribunda Decne: Whitmore FRI 4247. Geophila
repens (L.)Johnston: Chin 154, 1549; UNESCO 689. Globba albiflora Ridl. var aurea Holtt.: Henderson
29715. G. fasciata Ridl.: Chin 1420, 1543; Stone 9524. G. patens Miq.: Henderson 25015. Glochidion
obscurum (Roxb. ex Willd.)Bl.: Chin 472; King’s coll. 4630. G. perakense Hk.f.: Chin 835A; Samsuri
& Mahmud 587. G. rubrum Bl.: Chin 350, 1256, 1682; Nur 34377; Reid 51679; Symington KEP 37413,
KEP 37453. Glossocarya mollis Wall.: Henderson 23 xi ’29. Glycosmis calcicola Stone: Chin 342, 373,
502, 959; Henderson & Symington KEP 43289; Samsuri & Mahmud 634; Stone 5907, 8789, 8846, 8937,
8977, 11004, 11078; Symington KEP 37420; Whitmore FRI 4040. G. clacicola Stone, var. kelantanica
Stone: Stone 7479, 9521, 12611; Whitmore FRI 4255. G. chlorosperma (Bl.)Spreng.: UNESCO 506, 577.
G. rupestris Ridl.: Chin 816, 904, 1491; Kiah 35229; Samsuri & Mahmud 578; Stone & Mahmud 12615;
UNESCO 120. G. sapindoides Lindl. ex Wall.: Stone 6942. Glyptopetalum quadrangulare Prain ex King:
Chin 334, 473, 1261; Kiah 35405; Ridley s.n. & s.a. (SING). Gmelina asiatica L.: Chin 937; Keng 6113.
G. villosa Roxb.: Chin 793, 927. Gnetum cuspidatum Bl.: Chin 530; Chin & Badaruddin 1703; Hender-
son 19560. G. gnemon L.: Chin 652. Gomphandra quadrifida (B\.)Sleum.: Kiah 35256. Gomphia serrata
(Gaertn.)Kanis: Chin 555, 1074; Corner 37815, 37854. Gomphostemma crinitum Wall.: Burkill 6298;
Sinclair 9848. G. curtisii Prain: UNESCO 54. G. javanicum (B1.)Benth.: Corner & Nauen 25 xi 41; Kiah
35381; UNESCO 355. G. microcalyx Prain: Chin 1558; Poore 16 vii ’61. Gongylosperma lanuginosum
Ridl.: Corner 37836; Curtis 2663; Henderson 29141; Stone 11024; Symington KEP 46742.
Goniothalamus fulvus Hk.f. et Th.: UNESCO 394 & 532. G. scortechinii King: UNESCO 90. G.
subevenius King: King’s coll. 8260. G. unvarioides King: Loh FRI 17192. Goodyera hispida Lind.: Allen
xii 56; Chin 1425A; Chin & Badaruddin 1702; Stone 5925. Grewia acuminata Juss.: Stone 6930, 9175.
G. paniculata Roxb. ex DC.: UNESCO 196, 521. G. viminea Wall. ex Burr.: Chin 1765, 1832; Corner
19 xi °41; Haniff & Nur 7487; Henderson 21377, 22814; Soepadmo & Mahmud 1219; Stone 11009. Guet-
tarda speciosa L.: Chin 524, 1789; Holttum 15133; Stone 9141. Gymnanthera insularum K. et G.: Chin
506, 1743; Corner 19 xi ’41. Gymnopetalum cochinchinense Kurz.: Burkill 13937. Gymnostachyum
84 Gard. Bull. Sing. 36(1) (1983)
decurrens Stapf: Boey 267; Chin 91, 922, 1123, 1456, 1609; Corner 20 xi ’41; Spare 36329; Stone 7458.
G. diversifolium C.B. Clke: Henderson 29094; Holttum 15128.
Habernaria carnea N.E.Br.: Corner 37861, 20 xi ’41; Curtis 2104. H. kingii Hk.f.: Best 337; Curtis
3342, xii 1895; vii 1896; (Anon.) xii 1895 (SING). H reflexa Bl.: Carr 315; Corner & Nauen 25 ix ’41.
Hanguana malayana (Jack)Merr.: Chin 680; UNESCO 86, 86A. Hapaline brownii Hk.f.: Henderson
19510; Ridley 14784. Hedyotis congesta R.Br. ex Don: UNESCO 65. H. coronaria (Kurz)Craib: Hender-
son 22809; 29181; Sinclair 9858; UNESCO 155, 270, 490. H. tenelliflora Bl.: Chin 1744; Corner 13 xi
’41; Henderson 28944, 29105; UNESCO 618. Helicteres angustifolia L.: Corner 37826, 37830; Holttum
17430. H. hirsuta Lour.: Chin 1711; Whitmore FRI 12995. Helixanthera coccinea (Jack)Dans.: Stone
7474. H. pulchra (DC.)Dans.: UNESCO 170, 329. Hemigraphis ridleyi C.B. Clke: Henderson 28933. He-
mionitis arifolia (Burm.)Moore: Curtis xi 1890; King’s coll. 4174. Heritiera littoralis Ait.: Chin 1786;
Henderson 29169. H. pterospermoides Kosterm. Chin 824, 1183; Ng FRI 1790, 1792. Heterogonium
alderwereltii Holtt.: King’s coll. 465; Ridley 864; Samat 599-607; Scortechini s.n. & s.a. (SING); Smith
1966. H. pinnatum (Copel.) Holtt.: Chin 426, 584, 594, 665; Evans 265; Henderson 19400, 22303, 22376,
22420, 25004; Keng 282; King’s coll. 5871; Molesworth-Allen & Kadim 489; Ridley 13434, xii 1896;
Samat 468, 472, 525, 593-598, 608-611; Sinclair 40062; Stone 7299B, 8822. Heterostemma piperifolium
K.et G.: Burkill 2556, 6265; Burkill & Haniff 13931; Chin 28, 387, 413, 1508. Hippeophyllum scortechinii
(Hk.f.)Schltr: Henderson 25090. Holarrhena curtisii K. et G.: Holttum 17428. Homalanthus populneus
(Geisel.)Pax: Chin 781; Henderson 22301; Samsuri & Mahmud 558. Homalium dasyanthum
(Turcz.)Warb.: Chin 497; Henderson 29116; Holttum 15096; Whitmore FRI 15101. H. foetidum
(Roxb.)Benth.: Whitmore FRI 8518. H. kunstleri King: King’s coll. 7109. H. undulatum King: Corner
38132; King’s coll. 8184. Homalomena argentea Ridl.: Chin 648. H. deltoidea Hk.f.: Chin 593, 728,
1585. H. griffithii Hk.f.: UNESCO 280. H. humilis (Jack)Hk.f. var. pumila (Hk.f.) Furt.: Burkill 2559,
6304. H. rubra Hassk.: UNESCO 101. Homonoia riparia Lour.: Loh FRI 17237. Hopea ferrea Lianess.:
Corner 38149; Henderson 21 xi ’34, 29157. Horsfieldia tomentosa Warb.: UNESCO 159. Hoya cor-
onaria Bl.: Burkill 6297; Chin 107, 797; Stone 5918. H latifolia Don.: Chin 42; Stone & Mahmud 29 v
°70. H. maingayi Hk.f.: UNESCO 125. H. occlusa Ridl.: Ridley xii 1890. Hoya parviflora Wight: Corner
26 xi °41. H. revoluta Wight ex Decne: Chin 95. H. ridleyi K. et. G.: UNESCO 34, 371. Humata
heterophylla (Sm.)Desv.: Chin 1437, 1606; UNESCO 230, 670. H. pectinata (Sm.)Desv.: Chin 545, 1605;
Henderson 19538, 25247; UNESCO 657, 672. Hunteria zeylanica (Retz.)Gard. ex Thw.: Henderson
23850. Hydnocarpus castanea Hk.f. & Th.: UNESCO 6. H. ilicifolia King: Chew 194; Chin & Badarud-
din 1693; Kiah 35272; Loh FRI 17174; UNESCO 541. H. woodii Merr.: Chin & Badaruddin 1693. H
wrayi King.: Loh FRI 17191. Hydnophytum formicarum Jack: UNESCO 638. Hygrophila angustifolia
R. Br.: Samsuri & Mahmud 589. Hypserpa cuspidata Miers: Chew 190. Hyptis rhomboidea Mart. & Gal.:
UNESCO 614. H. suaveolens (L.)Point.: Chin 837.
Iguanura polymorpha Becc.: Burkill 6299; Chin 871; Henderson 22685, 23792, 25059; Samsuri &
Mahmud 604. Ilex maingayi Hk.f.: King’s coll. 8177. Illigera pulchra Bl.: Corner & Nauen 25 xi ’41 (=
SING 645109); Henderson 23015, 29174. Impatiens cryptoneura Hk.f.: Curtis 3172. I. mirabilis Hk.f.:
Chin 1756; Enoch 8; Henderson 23023. I. opinata Craib: Chin 1312, 1495, 1610; Henderson 19509,
19558, 22374, 25087, 29677; UNESCO 5, 491, 535, 580, 602. I. ridleyi Hk.f.: Burkill 2260; Chin 332 &
1275; Henderson 22324; Sinclair 40064. I. scortechinii Hk.f.: Curtis 3115; Haniff 13904; Henderson
23839. I. tipusensis Henders.: Henderson 19339. Imperata cylindrica (L.)Beauv. var. major (Nees)Hubb.
ex Hubb. et Vaugh.: UNESCO 616. Iodes cirrhosa Turcz.: Chin 792, 953, 994; Samsuri & Mahmud 565,
653. I. ovalis Bl.: Mills & Henderson 15067; Sinclair 9875. Isachne langkawiensis Jansen: Chin 1787; Cor-
ner 37843; Corner & Nauen 37845; 37959; Henderson 28945 & 29054; Holttum 17423. Ischaemum
timorense Kunth: Corner 37844. Isonandra perakensis K. & G. var. kelantanensis Ng: Loh FRI 17179;
17212; UNESCO 654. I. perakensis K. & G. var. perakensis Ng: Ng FRI 1598, 1599, 1906, 1908, 1914,
1915, 1916, 5846. Ixora brunonis Wall. ex Don: Best 21274. I. clerodendron Ridl.: Henderson 25245;
30 vii 29, 6 viii ’29. I. congesta Roxb.: Kiah 35366; UNESCO 383. I. grandifolia Zoll. & Mor.: UNESCO
379. I. lobbii K. & G.: UNESCO 208. I. nigricans Wight & Arn.: UNESCO 40. I. nigricans Wight &
Arn. var. ovalis Pitard: Chin 1402, 1594; Stone 9179. I. pendula Jack: Chin 161; Samsuri & Mahmud
615; Stone 6575; UNESCO 96, 381. I. scortechinii K. & G.: Henderson 19518. I. umbellata Koord. et.
Valet.: Kiah 7 v ’38: Stone 9137.
Jacquemontia paniculata (Burm.f.)Hall.f.: Henderson 28941; Ridley 14900a. Jasminum
adenophyllum Wail.: Kiah 35424. J. bifarium Wall.: UNESCO 661. J. cordatum Ridl.: Chin 366; Mills
Limestone hill flora of Malaya IV 85
& Henderson 15064; Molesworth-Allen 4289; Stone 5893. J. curtisii K. & G.: Henderson 23010. J. in-
sularum Kerr: Chin 454. J. wrayi K. & G.: King’s coll. iv 1884; UNESCO 19. Justicia henicophylla C.B.
Clke: Henderson 22312. J. hirticarpa J.B. Imlay: Henderson 29176. J. pectinella Ridl.: Chin 915; Kiah
35359. J. ptychostoma Nees: Henderson 29136; Nauen 38123; Nur 34391. J. robinsonii Ridl.: Henderson
29133. J. subcymosa C.B. Clke: Sinclair 9889. J. uber C.B. Clke: Chin 397; Henderson 19508, 22399;
Stone 5921. J. vasculosa Wall.: Burkill 2553; Samsuri & Mahmud 582.
Kaempferia elegans Wall.: Curtis 2637. K. pulchra Ridl.: Henderson xi ’29. Kibara chartacea Bl.:
Henderson 25070. Knema cinerea (Poir.)Warb. var. patentinervia (Sinclair) Sinclair: Loh FRI 17243. K.
cinerea (Poir.)Warb. var. rubens (Sinclair) Sinclair: UNESCO 200. K. globularia(Lamk.)Warb.: Corner
22 xi °41; Haniff & Nur 7569; Henderson 22978. K. hookeriana (Hk.f. et Th.)Warb.: Loh FRI 17188.
K. laurina (Bl.)Warb.: Kiah 35398. Knoxia corymbosa Willd.: Burkill 1392. Koilodepas longifolium
Hk.f.: Stone 6946. Kopsia macrophylla Hk.f.: Henderson 25044.
Lagerstroemia langkawienis Furt. et Mont.: Chin 486; Corner 19 xi ’41. Lantana camara L. var.
aculeata (L.)Moldenke: Chin 804. Laportea interrupta (L.)Chew: Molesworth-Allen 4643. Lasia aculeata
Lour.: Chin 836A; Henderson 22487; Kiah 35429; Ridley 23 vi 1889. Lasianthus stipularis Bl. var. hirtus
Ridl.: Ridley 8575. Lasiobema curtisii (Prain)De Wit: Corner 13 xi ’41; Kiah 35415. L. flavum De Wit:
Henderson 29146. L. strychnoideum (Prain)De Wit: Chin 840; Samsuri & Mahmud 590; Stone 9530. Leea
aequata L.: Kiah 35234; Samsuri & Mahmud 566. L. rubra B\.: Kiah 35421. L. sambucina Willd.: Chin
127, 476, 779, 1110; Samsuri & Mahmud 556; Stone 6940; Whitmore 4246. L. saxatilis Ridl.: Merton
4098; Poore 800. Lemmaphyllum accedens (B\.)Donk: UNESCO 601. Lepisorus longifolius (Bl.)Holtt.:
Chin 1414. Lepistemon binectariferum (Wall.)Ktze: Stone & Wycherley 8976. Leptaspis urceolata
(Roxb.)R.Br.: Chin 886. Leptochilus decurrens Bl.: UNESCO 27. Leptonychia glabra Turcz.: Chin 887
& 921; Henderson 23760; Molesworth-Allen 4270. Leptopus australis (Zoll. et Mor.)Pojark.: Best 21233;
Chin 879; Corner 37802, 13 xi’41; Henderson 19513, 22839; Kelsall s.n. 1891; Kiah 35249; Nauen 38012;
Ridley 8203; Samsuri & Mahmud 638. Leucaena leucocephala (Lamk.)De Wit: Chin 979. Leucas
mollissima Wall. ex Benth.: Chin 1776; Kiah 35381. L. zeylanica R. Br.: Chin 846. Licuala modesta
Becc.: Chin 884; Henderson 23804. Ligustrum confusum Decne: Chin 400, 414, 440, 574; Henderson
22865, 22941; Stone 8942. Liparis caespitosa (Thou.)Lindl.: Chin 4178; Nur 14 x31; Stone 5926. L. gib-
bosa Finet: Carr 162; Chin 163, 456, 1176; Henderson 19456, 22242, 25046; UNESCO 37, 250, 366.
Lithocarpus elegans (Bl\.)Hatus. ex Soepadmo: Henderson 23806. L. urceolaris (Jack)Merr.: UNESCO
344. Litsea glutinosa (Lour.)C.B. Rob.: Chin 31; Chin & Badaruddin 1689; Corner & Nauen 25 xi ’41;
Kiah 35372; Stone & Wycherley 8987. L. noronhae Bl.: Burkill 6289; Henderson 23810. Livistona saribus
(Lour.)Merr. ex Chev.: Henderson 7 vi ’30 (= SING 39682). Loeseneriella pauciflora (DC.)A.C. Smith:
Henderson 29688. Loxogramme avenia (Bl.)Presl: Chin 1447; King’s coll. 8280. L. scolopendrina
(Bory)Presl: Chin 26, 139; Henderson 19449; Kiah 12 v ’38; Molesworth-Allen 18 i ’56; Shimizu & Stone
T 14396; Stone 7311, 8828, 8842; Stone & Mahmud 9397; UNESCO 245, 435. Ludwigia hyssopifolia (G.
Don)Exell.: Chin 838; Samsuri & Mahmud 588. Luvunga eleutherandra Dalz.: Keng 6174. Lycopersicon
esculentum Mill.: Chin 863. Lygodium flexuosum (L.)Sw.: UNESCO 188. L. polystyachum Wall. ex
Moore: Chin 904; Henderson 7 vi ’30; UNESCO 83. Lysimachia peduncularis Wall. ex Kurz: Henderson
21371; Keng et al.: 6211.
Macaranga tanarius (L.)M.A.: Chin 444, 577 & 1804; Samsuri & Mahmud 597. Macrosolen cochin-
chinensis van Tiegh.: Chin 398 & 447. Madhuca ridleyi Lam: Chin 1071; Henderson 25055; Stone 6924;
UNESCO 113. Maesa pahangiana K. & G.: UNESCO 410, 530. M. striata Mez, Chin 1580. Malaisia
scandens (Lour.)Planch.: Chin 857, 928; Samsuri & Mahmud 599. Malaxis calophylla (Rchb. f.)Ktze.:
Kiah 13 v ’38; UNESCO 357. M. latifolia Sm.: Henderson xi 34. M. micrantha (Hk.f.)Ktze: Henderson
22332. M. reniloba (Carr)Holtt.: Henderson xi ’29. Malleola dentifera J.J.S.: UNESCO 35, 240, 1014.
Mallotus brevipetiolatus Gage: Chew 209; Chin 753, 819, 850, 1488; Molesworth-Allen 4644; Ng FRI
1601; Ogata KEP 110159; Samsuri 542; Samsuri & Mahmud 595; Sinclair 9853, 9856. M. cuneatus Ridl.:
Chew 198; Henderson 23020; Kiah 35238. M. dispar (Bl.)M.A.: Chew 199; Chin 83, 314, 917, 1488A;
Corner 13 xi ’41 (= SING 28327); Henderson 29143; Kiah 35375; Poore 352; Samsuri & Mahmud 620;
Stone 6947, 7478, 9178; Stone & Wycherley 8974. M. eriocarpus (Thw.)M.A.: Henderson 25211. M. grif-
_fithianus (M.A.)Hk.f.: King’s coll. 4602. M. miquelianus (Scheff.)Boerl.: Chin 1552, 1597; Henderson
19543. M. oblongifolius (Miq.)M.A.: Molesworth-Allen 4269. M. peltatus (Geisel.)M.A.: Sinclair 9859.
M. philippensis (Lamk.) M.A.: Chin 830; Kiah 35423. M. repandus (Willd.)M.A.: Stone 8849. M. wrayi
King ex Hk.f.: Henderson 23803. Mammea brevipes (Craib)Kosterm.: Chin 1729; Henderson 29163.
86 Gard. Bull. Sing. 36(1) (1983)
Mangifera sp.: Chin 329, 911; Ng FRI 5577; UNESCO 399; Whitmore FRI 15626. Marsdenia tinctoria
Br.: Burkill 13928; Chin 77; Kiah 35245. Maxburretia gracilis (Burr.)Dranfield: Curtis 2661; Fox xii
04; Stone 9147; Stone & Mahmud 6951; Whitmore FRI 15053. M. rupicola (Ridl.)Furt.: Chin 362, 363
& 364; Durant 56257; Nur 34370; Ridley 8285; Stone 8920; Symington KEP 39585; Whitmore FRI 699,
12167, 15173, 15174, 15560 & 15636; Wong FRI 99501. Maytenus curtisii (King) Ding Hou: Chin 1462,
1725, 1835; UNESCO 143; Whitmore FRI 4261. Medinilla crassifolia (Reinw. ex Bl.)Bl.: Chin 528. M
scortechinii King: Chin 1595; UNESCO 629. Meiogyne virgata (Bl.)Miq.: Chin & Badaruddin 1692;
Henderson 19609; UNESCO 1007. Melaleuca cajuputi Powell: ’Students’ 45. Melanolepis multiglan-
dulosa (Reinw. ex Bl.)Rchb.f. et Zoll.: Chin 855, 951, 952. Melastoma polyanthum Bl.: UNESCO 619.
Melochia umbellata (Houtt.)Stapf: Chin 982, 989; Samsuri & Mahmud 648; Stone 9176. Melodinus
orientalis Bl.: King’s coll. 4530. Melothria affinis King: Curtis 3784; Ridley xii 1902. Memecylon
acuminatum Sm.: Corner 13 xi ’41 (= SING 54941). M. dichotomum C.B. Clke. ex King: Corner 17
xi ’41; Henderson 22 xi 34, 25064. M. edule Roxb.: Chew 207; Chin 313; Stone & Wycherley 8972. M.
floribundum Bl.: Henderson 22881; Kiah 35280. M. laevigatum Bl.: Chin 112, 371, 372, 452, 1161; Kiah
35281; Molesworth-Allen & Kadim 494. M. pauciflorum Bl.: Chin 823; Henderson 22 xi ’34; Nur
34371; Samsuri & Mahmud 579; Stone 6934. M. wallichii Ridl.: UNESCO 95, 609. Mesua ferrea L.:
UNESCO 76. Microdesmis casearifolia Planch.: Chin 346, 879; Kiah 35231; Samsuri & Mahmud 607.
Microlepia speluncae (L.)Moore: Shimizu & Stone M 13726, M 13728, T 14417. M. speluncae (L.)Moore
var. villosissima C. Chr.: Chin 427; Henderson 25210; Ridley 8641; Stone 7483. Micromelum minutum
(Forst.)W. & Arn.: Whitmore: FRI 4260. Microrphium pubescens C.B. Clke: Corner 37857; Henderson
29184; ‘Students’ 148. Microsaccus ampullaceus J.J.S.: Carr 69; Nur 25153. M. brevifolius J.J.S.: Chin
417A; Henderson 22261.M. javensis B\.: Chin 396; Nur 34402; Stone 5927; UNESCO 300. Microsorium
musifolium (Bl.)Ching: Henderson 19704; UNESCO 635. M. punctatum (L.)Copel.: Chin 786. Mikania
cordata (Burm.f.)B.L. Robins.: Henderson 23781. Miliusa amplexicaulis Ridl.: Chin 900; Kiah 35363;
Samsuri & Mahmud 614; UNESCO 516. M. longipes King: Henderson 25038. M. parviflora Ridl.: Cor-
ner 19 xi’41 (= SING 4818); Henderson 28949; Ridley 15249 & 15340. Millettia hemsleyana Prain: Loh
FRI 17238; UNESCO 195. M. pterocarpa Dunn: Fox 10788. M. sericea (Vent.)W.et Arm.: UNESCO
122. Mimosa pudica L.: UNESCO 620. Mimusops elengi L.: Chin 1737, 1782; Corner 38141; Henderson
21363, 21369; Stone 11002. Mitrephora maingayi Hk.f. et Th.: King’s coll. 4701. Moghania strobilifera
(L.)St. Hill. ex. Jacks.: Stone 9078. Momordica subangulata Bl.: Curtis 3329. Monophyllaea glabra
Ridl.: Henderson 29072. M. hirticalyx Franch.: Allen v ’46; Burkill 4222; Chin 773, 2107; Curtis 3136;
Henderson 19380, 19404, 22578, 29790; King’s coll. 7052; Spare 36324; Tomlinson ix ’55. M. horsfieldii
R.Br.: Chin 391, 546, 881, 1539; Henderson 25067, 29714; Kassim 375; Keng & Keng 70; Ng FRI 1594;
Sinclair 9848; Stone 7298; UNESCO 10, 21, 396, 533, 1001; Wray 597. Morinda elliptica Ridl.: Chin 805,
828; Chin & Mahmud 1313; Kiah 35408; Stone 6985; UNESCO 8. M. umbellata L.: Burkill 2555; Chin
53, 96, 388; Stone 5904, 7305A, 7477, 8929, 9501. Mucuna biplicata Teysm. et Binn.: Samat 417; Start
23 i ’72. Muntingia calabura L.: Chin 445. Murraya koenigii (L.)Spreng.: Henderson 28947. M.
paniculata (L.)Jack: Chin 756, 794, 822, 991, 1509; Molesworth-Allen 4647; Samsuri & Mahmud 571;
UNESCO 177, 536. Musa malaccensis Ridl. var. minor Ridl.: Ridley 15202. Mycetia malayana Craib:
Burkill 13571; Chin 760, 783; Kiah 7 v ’38; Mills & Henderson 15069; Poore 16 vii ’61; Stone 8786;
UNESCO 493, 525. Myrsine porteriana Wall.: Chin 415, 547, 1565; Corner & Nauen 25 xi ’41; Kiah
35373; Stone 5892; Whitmore FRI 12161.
Naravelia dasyoneura Korth.: UNESCO 219. N. laurifolia Wall.: Chin 128. Nauclea junghuhnii Merr.:
King’s coil. 8179. Neesia synandra Mast.: UNESCO 589. Neolitsea zeylanica Merr.: Chin 145; Stone
7474; UNESCO 284, 645. Neonauclea calycina Merr.: Chin 1481; UNESCO 206. Nephrolepis biserrata
(Sw.)Schott: Chin 135. N. dicksonioides Chr.: Allen xii ’56; Best 21236, 21284; Chin 405, 1602; Evans
202; Henderson 19528, 22263; Kiah 35407; Nauen 38023; Stone 5895. N. falcata (Cav.)C.Chr.: Chin 527,
570, 813; Henderson 19405, 22304; Poore 348; UNESCO 198, 376. N. hirsutula (Forst.)Pr.: Chin 561.
Neyraudia reynaudiana (Kunth)Keng ex Hitch.: Chin 974, 1421; Samsuri & Mahmud 641.
Oberonia dissitiflora Ridl.: Corner 20 xi ’41. O. flava Ridl.: UNESCO 309. O. spathulata Lindl.:
UNESCO 420, 421. O. transversiloba Holtt.: UNESCO 248, 422. Oldenlandia rosettifolia Geddes: Chin
1721; Henderson 28929; Holttum 15138. Oleandra undulata (Willd.)Ching: Kadim K491. Ophiorrhiza
communis Ridl.: Kiah 25376. O. discolor R. Br. ex Don: Chin 335, 537, 890; Chin & Badaruddin 1688;
Stone 8975. O. fruticosa Ridl.: Ng FRI 1639; Nur 34374; Ridley 8237. O. hispidula Wall. ex Don.:
UNESCO 393, 1006. O. kunstleri King: Chin 1394, 1732; Corner 38147; Henderson 21382, 29711; Kiah
35251; UNESCO 53: O. longerepens Ridl.: Chin 85, 1395, 1454, 1563; Henderson 19524; Stone 7457 &
Limstone hill flora of Malaya IV 87
9519. O. pallidula Rid\.: Henderson 29677; Nur 34374; Spare 36322. O. remotiflora Ridl.: Chin 1612.
Oplismenus compositus (L.)P. Beauv.: Corner & Nauen 25 xi’41; Henderson 23130, 28934. Ornithobosa
flexuosa (Ridl.) Burtt: Chew 203; Haniff 640; Kiah 35419, 18 v ’38. Orophea cuneiformis King; Haniff
10335. O. enterocarpa Maingay ex Hk.f. et. Th.: Henderson 25040; Stone 7496. O. hirsuta King: Hender-
son 35402; Kiah 35276; King’s coll. 4283; UNESCO 43, 412, 595. O. maculata King: Chin 769, 780; Chin
& Badaruddin 1687; Henderson 43281; Md. Shah & Md. Ali 2999; Samsuri 552; Samsuri & Mahmud 559.
O. polycarpa A. DC.: Chin 149; Henderson 22443; Ng FRI 1595, 5836; Ogata KEP 110158 & 110160;
Sinclair 9892; UNESCO 136. Orthosiphon aristatus (Bl.)Miq.: Chin 1715; Henderson 29092. Oxymitra
biglandulosa (B1.)Scheff.: Chin 1040.
Pachycentria constricta (B\.)Bl.: Henderson 25242. Paedaria tomentosa Bl.: Henderson 29147. Palaquium
obovatum (Griff.)Engl.: Loh FRI 17207. Pandanus alticola Holtt. & St. John: Chin 1684A; Stone 5885,
5886. P. calcicola Holtt. & St. John: Chin 321A, 1274, 1663, 1664; Kiah 35285; Stone 8969; Whitmore
FRI 15625. P. irregularis Ridl.: Chin 1529, 1530, 1531, 1532, 1617; Henderson 19468; Stone & Mahmud
7448. P. ordoratissimus L.f.: Henderson 29153. P. piniformis Holtt. & St. John: Henderson 23759;
Stone & Mahmud 9182. P. recurvatus St. John: UNESCO 47. Panicum sarmentosum Roxb.: Chin 938.
Paphiopedilum lowii (Lindl.)Pfitz.: UNESCO 443. P. niveum (Rchb. f.)Pfitz.: Chin 1723; Curtis ’20’s;
Stone 6979. Paraboea bakeri Henders.: Henderson, 25094. P. bettiana Henders.: Henderson 25250. P.
capitata Ridl.: Burkill 6276, 13557; Chin 831; Curtis xii 1895; Henderson 22252; King’s coll. 4325; Sam-
suri & Mahmud 581; Spare 36327; UNESCO 409. P. ferruginea Ridl.: Curtis 2566. P. laxa Ridl.: Alphon-
so & Samsuri 122; Chin 1731, 1785; Corner 13 xi ’41, 17 xi ’41; Henderson 21366, 29187. Parameria
polyneura Hk.f.: Henderson 23758. Paramignya scandens (Griff.)Craib subsp. ridleyi (Burkill)Swing.:
Stone 8981. Paranephelium macrophyllum King: Chin 841, 856; Samsuri & Mahmud 598. Parashorea
lucida (Miq.)Kurz: UNESCO 377. Paspalum conjugatum Berg.: Chin 378. Passiflora foetida L.: Chin
976. Pavetta naucleiflora R.Br. ex G.Don: Henderson 21374. P. pauciflora Ridl.: Ridley 13 xii ’20, xii
’20, xii ’27. Peliosanthes lurida Ridl.: Chin 543, 564; Corner 17 xi ’41; Henderson 29180; Kiah 35426.
P. violaccea Wall.: Nur 31 xi ’37. Peltophorum pterocarpum (DC.)Back. ex Heyne: Stone 10994. Pen-
tacme siamensis (Miq.)Kurz: Chin 510; Henderson 29102; Stone 6990; Whitmore 10 xii ’69, FRI 12969,
15066, 15104. Peperomia dindigulensis Miq.: Chin 1172; Chin & Mahmud 1310; Henderson 22560,
25237. P. kotana C. DC.: Corner & Nauen 25 xi ’41; Stone 7454. Peperomia sp. A (= P. maxwellana
C.DC.): Chin 539, 1695; Stone 14086. Petunga hirta Ridl.: Mills & Henderson 15070. Phaeomeria ma-
ingayi (Bak.)K. Schum.: Chin 1583. Phalaenopsis cornu-cervi (Breda)Bl. et Rchb.f.: Carr 289; Hender-
son 22274, 25073; UNESCO 221, 417. P. decumbens (Griff.)Holtt.: Henderson 22235, 25244; UNESCO
413. Phanera integrifolia (Roxb.)Benth.: Chin 541. Phaseolus mungo L.: Chin 483. Pheobe lanceolata
(Wall. ex Nees)Nees: UNESCO 511. Pholidota pallida Lindl.: Best 21279; Chin 419, 459; Corner 19 xi
41; Corner & Nauen 25 xi ’41; Henderson 25241; Holttum 15136; Nur 34386; Ridley xii 1896; Stone
7312, 7481, 7484; UNESCO 418. Photinopteris speciosa (Bl.)Presl.: Best 21237; Chin 1163, 1559;
UNESCO 266. Phreatia secunda (Bl\.)Lindl.: Carr 250; Chin 322, 416; Henderson 25235; Nur 34406;
UNESCO 288, 416. Phyllanthus columnaris M.A.: Chin 1710; Corner 38135; Henderson 22840, 29056.
P. filicifolius Gage: Henderson 29053. P. oxyphyllus Miq.: Chin 316, 348, 761; Henderson 22271; Ridley
8509; Samsuri 546; Stone 8966. P. pulcher Wall. ex M.A.: Henderson 29690; Loh FRI 17216. P.
ridleyanus Airy Shaw: Chin 114, 790, 966, 1059, 1386; Henderson 23764; King’s coll. 8219; Ogata KEP
11061; Samsuri & Mahmud 564, 627; Stone 9494; Whitmore FRI 4273. P. sikkimensis M.A.: Henderson
22872. Phymatodes nigrescens (B\.)J. Sm.: Burkill 13930; Chin 43, 88; Corner & Nauen 25pxin Ale
Henderson 22397; Nur 34393; Samat 511; Shimizu & Stone M 13721; Stone 8824. P. scolopendria
(Burm.)Ching: Chin 1150, 1604; Corner & Henderson 22801; UNESCO 434. Physalis minima L.: Sam-
suri & Mahmud 592. Pilea fruticosa Hk.f.: Burkill 2560, 6264; Burkill & Haniff 13927; Chin 24, 765,
1405; Corner & Nauen 25 xi’41; Henderson 19385, 22375, 22436, 25008, 29710; Molesworth-Allen 4015,
4432; Ng FRI 1609; Samsuri 549; Sinclair & Forman 9851; Spare 36325; Stone 7461, 8797, 8835. P.
microphylla (L.)Lieb.: Chin 975; Samsuri & Mahmud 641. Piper boehmeriaefolium Wall.: Henderson
23756. P. caninum Bl.: Kiah 35396. P. mucronatum C. DC.: Poore 801. P. nigrum L.: Ridley 8180. P.
porphyrophyllum N.E.Br.: Chin 611. P. umbellatum L.: Burkill & Haniff 13923. Pisonia aculeata L.:
Henderson 23121, 29175. P. umbellifera (Forst.)Seem.: Burkill 6291; Chin 860; Henderson 25011; Ridley
9681; Samsuri & Mahmud 600. Pistacia malayana Henders.: Chin 401, 1064, 1808. Pittosporum fer-
rugineum Ait.: Ng FRI 5572. Pityrogramma calomelanos (L.)Link: UNESCO 259. Planchonella obovata
(R. Br.)Pierre: Allen 5 ii ’57; Chin 81, 339, 365, 1182; Ng FRI 1623, 1625, 1796, 1802; Stone 5898, 8937;
UNESCO 312, 369; Whitmore FRI 4248. Plectocomia griffithii Becc.ex Hk.f.: Chin 1273. Plectranthus
kunstleri Prain: Chin 1760. Plethiandra sessiliflora Ridl.: Chin 565; Stone 11081. Poaephyllum
88 Gard. Bull. Sing. 36(1) (1983)
pauciflorum (Hk. f.)Ridl.: Spare 36323. Podocarpus neriifolius D. Don: Ng FRI 1634. P. polystachyus
R.Br. ex Mirb.: Chin 157, 368, 369, 538, 1811, 1260; Loh FRI 17214; Ng FRI 1634; Stone 5905.
Podochilus lucescens Bl.: Corner 20 xi ’41; Henderson 29075. P. microphyllus Lindl.: Chin 466, 1401;
UNESCO 373. P. tenuis (Bl.)Lindl.: Symington 10 x 34. Pogonanthera pulverulenta (Jack)BI.: Chin 30.
Pogonatherum paniceum (Lamk.)Hack.: Chin 435, 1521; Chin & Mahmud 1306A. Poikilospermum
suaveolens (Bl.)Merr.: Chew 126, 195, 210; Chin 788; Henderson 22453; Loh FRI 17199; Samsuri &
Mahmud 570. Pollia sorzogonensis Endl.: Henderson 22703, 14 vii ’35. P. sumatrana Hassk.: Burkill
& Haniff 13907. P. thyrsiflora Endl.: Chin 360. Polyalthia brunneifolia Sinclair: Chin 141, 149, 312,
1062, 1094, 1170, 1407, 1507; Ng FRI 5548; Whitmore FRI 757; Stone & Wycherley 8982. P. cauliflora
Hk.f. et Th. var. beccarii (King)Sinclair: Loh FRI 17239. P. cinnamomea Hk.f. et Th.: Whitmore FRI
468. P. hypogaea King: Mills & Henderson 15075. P. lateritia Sinclair; Chin 877; Samsuri & Mahmud
608. P. rumphii (Bl.)Merr.: Stone & Wycherley 8983. P. stenopetala (Hk.f. et. Th.)Ridl.: Loh FRI 17252.
Polygala cardiocarpa Kurz: Corner xi ’41; Henderson 29104. P. malesiana Adema: Allen 27 1’57; King’s
coll. 4814. P. triphylla Buch. et Ham. ex Don: Henderson 22254. Polygonum chinense L.: Ng FRI 1918.
Polypodium papilosum Bl.: King’s coll. 7206. Polystachya flavenscens (B1.)J.J.S.: Chin 422; Curtis ix
1890; Henderson 25075, xi’34; Ng FRI 1922. Polystichum lindsaeifolium Ridl.: Henderson 19651, 22321;
Molesworth-Allen 4105. Polytrema cupreum Ridl.: Burkill 6285. P. vulgare C.B. Clke: Burkill 6307;
Henderson 25001. Pomatocalpa kunstleri (Hk. f.)J.J.S.: UNESCO 228. P. /atifolium (Lindl.)J.J.S.:
UNESCO 215. P. naevatum J.J.S.: Kelsall i 1891; UNESCO 275. P. setulense (Ridl.)Holtt.: Ridley
15226. P. spicatum Breda: Boey 301; Chin 1499; Henderson 23825; Nur 14 x ’31; UNESCO 225A, 374.
Popowia velutina King: King’s coll. 4418. Pothos latifolius Hk.f.: Furtado 4 vi ’37. P. macrocephalus
Scort. ex Hk.f.: Chin 999; Henderson 19557, 23835; Kiah 35282; Molesworth-Allen 4116; Samsuri &
Mahmud 638; Wary 4254. P. scandens L.: Curtis 2394; Kiah 35428; UNESCO 2394. Premna pyramidata
Wall.: Henderson 23766. P. rubens Ridl. UNESCO 407. Prismatomeris malayana Ridl.: Chew 187; Kiah
35271. Procris pendunculata (J.R. & G. Forst.)Wedd.: Chin 16; Stone 8834. Pseuderanthemum
crenulatum Radlk.: Burkill 6273; Chin 667; Corner 13 xi ’41; Henderson 21 xi ’34, 23091; Stone 8821.
P. graciliflorum (Nees ex Wall.)Ridl.: Henderson 21396; Stone 6941, 9080. Pseuduvaria macrophylla
(Oliv.)Merr.: Best 21262; UNESCO 69. P. setosa (King)Sinclair: Chin 955; Henderson 29713; Ogata KEP
110156. Psychotria angulata Korth.: Nur 34372; UNESCO 323. P. cantleyi Ridl.: Burkill 6363. P. mon-
tana Bl.: Burkill 6286; Henderson 23774. P. rhinocerotis Reinw. ex Bl.: Burkill 6310; Chin 811, 1099,
1439; Stone 6939; UNESCO 163, 225. P. rostrata B\l.: Henderson 22400. P. sarmentosa Bl.: Stone 6938.
P. viridiflora Reinw. ex Bl.: Henderson 25218. Pteridium aquilinum (L.)Kuhn var. wightianum
(Ag.)Tryon: Chin 99. P. caudatum (1.) Maxon var. yarrabense Domin.: Chin 383; Corner & Nauen 25
xi ’41. Pteridrys syrmatica (Willd.)C.Chr. et Ching: Best 21225; Chin 782, 849; Corner & Nauen 25 xi
’41; Henderson 22320; King’s coll. 8178. Pteris ensiformis Burm.: Chin 1174, 480; Johnson s.n. & S.a.
(KLU); King’s coll. 8334; Samat 524; Shimizu & Stone T 14389, T 14409; Stone & Wycherley 8990;
UNESCO 553. P. longipinnula Wall.: Burkill 13929; Chin 79, 428, 534; Henderson 19505; Samat 469;
Shimizu & Stone M 13722; Stone 8823. P. mertensioides Willd.: Ridley 8640. P. scabripes Wall. apud
Hk.: ‘Students’ 19. P. tripartita Sw.: Henderson 29581; Holttum 8975. P. vittata L.: Chin 100, 381, 432,
1423; Henderson 19517; Nur 3 xi’37; Samat 590; Shimizu & Stone M 13729. Pterisanthes coriacea Korth.
ex Miq.: UNESCO 87. Pteroceras ciliatum (Ridl.)Holtt.: UNESCO 298 & 415. P. hirsutum
(Hk.f.)Holtt.: Chin 1533; Stone 9509. P. tanyphyllum (Ridl.)Holtt.: Carr 142; Chin 93; Ridley 1891;
UNESCO 349, 363, 380. Pterolobium densiflorum Prain: Chin 352. Pterospermum jackianum Wall. ex
Masters: Chin 839; UNESCO 448. P. pectiniforme Kost.: Stone 9134, 10993. Pterygota alata
Roxb.)R.Br.: Chin 942. Pyrrosia adnascens (Sw.)Ching: Chin 424, 448, 495, 1108; Corner 17 xi ’41, 19
xi 41, 22 xi’41; Corner & Henderson 23105; Henderson 21378; Nur 34408; UNESCO 137. P. penangiana
(Hk.)Holtt.: Best 21230; Chin 1446; Henderson 19447, 22238, 25239; King’s coll. 7083; Nauen 38035;
Shimizu & Stone T 14353; Stone 7442; UNESCO 216, 243, 639. P. stigmosa (Sw.)Ching: Chin 431, 1080,
1449; Corner & Henderson 22811; Henderson 19453, 22572; King’s coll. 8361; Stone 6573. P. varia
(Kaulf.)Farwell: Best 21231; Chin 101, 430, 1053, 1109, 1436; Henderson 19450, 22575, 25154, 29076;
Stone 9508; UNESCO 161, 628, 660.
Quisqualis parvifolia (Rid\.)Exell. Stone 9082.
Radermachera lobbii (T.&B.)Miq.: Chin 116, 1400; Ng FRI 5570; Stone 9506; UNESCO 482, 651.
Randia densiflora (Wall.)Benth.: Burkill 6290; Chin 23, 337, 489, 851, 973; Henderson 29156;
Molesworth-Allen 4656; Samsuri & Mahmud 596; Stone 6996, 8790, 8837; UNESCO 3, 515; Whitmore
FRI 12159. Raphidophora beccarii Engl.: UNESCO 281. R. korthalsii H. Schott: Chin 997. R. kunstleri
Limestone hill flora of Malaya IV 89
Hk.f.: Chin 754; Samsuri & Mahmud 573; Sinclair 9852; Stone 9174. R. maingayi Hk.f.: Chin 592; Nur
8965; Ridley xii 1896. R. silvestris (Bl.)Engl.: Nur 8965; Ridley xii 1896. Rauwolfia reflexa Teys. et Binn.:
Burkill 6280. Renanthera elongata Lindl.: Best 21232. R. histrionica Rchb.f.: Carr 133; Chin 1244; Poore
12 ix 60. Rhododendron longiflorum Lindl.: Stone 5887. Rhoeo spathacea (Sw.)Stearn: Chin 806; Sam-
suri & Mahmud 557. Rhynchelytrum repens (Willd.)Hubb.: Burkill 2273; Chin 848; Samsuri & Mahmud
593. Rhynchoglossum obliquum Bl.: Henderson 19402. Richeriella malayana Henders.: Burkill 6281;
Henderson 23790. Rinorea anguifera (Lour.)Ktze: UNESCO 385, 1005. R. bengalensis (Wall.)Ktze:
Burkill 6278; Chew 201; Chin 82, 861, 885, 920, 1492; Ng FRI 1804; Samsuri & Mahmud 601, 610 &
630; Stone 5923, 6933, 8848; UNESCO 610; Whitmore FRI 15078, 15167. R. horneri (Korth.)Ktze: Chin
338, 475, 758, 1030, 1096; Cockburn FRI 10569; Henderson 19495, 22277; Stone & Wycherley 8967;
UNESCO 1, 108, 596. Rostellaria procumbens Nees: Corner 37829; Henderson 28942; Holttum 17425.
Ruellia repens L.: Chin 500. Rungia minutiflora C.B. Clke: Chin 1557; Henderson 21 xi 734.
Sageraea elliptica (A. DC.)Hk.f. et Th.: Chin 469, 1266. Salacia grandiflora Kurz: Henderson 23811;
Wyatt-Smith KEP 79149. S. korthalsiana Miq.: Henderson 22565. S. macrophylla B\.: Chin 771, 874;
Henderson 19602, 22708; Samsuri 553, 605. Salomonia ciliata D.C.: Corner 19 xi’41; Henderson 29059;
Holttum 17426. Saraca delinata (Jack)Miq.: Chin 776, 872; Ogata KEP 110197. S. indica L.: Henderson
19562. S. thaipingensis Cantley ex Prain: Chin 1568; Cockburn FRI 10565; Henderson 22700; Loh FRI
17193. Sarcanthus machadonis (Ridl.)J.J.S.: Stone 9510; UNESCO 391. S. rugulosus (Ridl.)Holtt.:
UNESCO 38, 358. S. sacculatus Ridl.: Chin 1777; Henderson 22805, 29129. S. scortechinii Hk.f.: Chin
1141, 1442; UNESCO 213. S. subulatus (B}.) Rchb.f.: Chin 1505; Corner 20 xi ’41; Henderson 29170;
UNESCO 340. S. termissus Rchb.f.: Henderson 23821. Sauropus brevipes M.A.: Henderson 23022. S.
calcareus Henders.: Henderson 22316. S. suberosus Airy Shaw: Chin 827, 1522; Henderson ii vi ’30;
Molesworth-Allen 4651. S. villosus (Blanco)Merr.: Henderson 22846, 25202; Nauen 38118. Schefflera
elegans (Ridl.)Ridl.: UNESCO 432. S. junghuhniana (Miq.)Harms.: Chin 882, 970; Samsuri 609; Sam-
suri & Mahmud 610, 631. S. musangensis Henders.: Chin 1828. S. subulata (Seem.)Vig.: Chin 1257,
1258, 1259, 1518; Poore 16 vii ’61; Stone 8973; UNESCO 121. S. tomentosa (Hassk.)Vig.: Chin 757;
Sinclair 9843; Stone 13 ii’66; UNESCO 666. S. venolusa (Seem.)Harms.: Chin 7. Schismatoglottis calyp-
trata Zoll. & Morr.: Chin 595, 697; UNESCO 528. S. mutata Hk.f.: Henderson 22427. Schizaea in-
opinata Selling: Chin 455, 1378A; Henderson 19460, 22246, 25157; Nur 34379; Ridley xii 1896; Shimizu
& Stone T 14419; Stone 5908, 7310, 7463; UNESCO 451. Schoenorchis micrantha Bl.: Ridley xii 1896.
Sciaphila asterias Ridl.: Holttum 15113; UNESCO 51. Scindapsus hederaceus Schott: Kiah 35230; Ridley
1897. S. perakensis Hk.f.: Burkill & Haniff 13563; Molesworth-Allen 4273; Ridley 23 vi 1889; Sinclair
9845. S. scortechinii Hk.f.: UNESCO 437. Scleria lithosperma Swartz: Chin 1105; Corner 19 xi ’41;
Henderson 21364, 22467, 25047; UNESCO 313. S. purpurascens Steud.: Chin 969. Sclerophyrum
wallichianum Arn.: Henderson 22278; Kiah 35374. Scolopia spinosa (Roxb.) Warb.: Chin 354, 603;
Stone 8838; Stone & Wycherley 8985. S. steenisiana Sleum.: Chin 399, 2104; Ng FRI 5580; UNESCO
127, 127A. Scoparia dulcis L.: Turnau 743. Scurrula ferruginea Dans.: Henderson 21386. Scutellaria
discolor Wall. ex Benth.: Chin 905; Samsuri & Mahmud 616. Secamone micrantha Decne: Chin 351, 987;
Samsuri & Mahmud 647; Stone & Mahmud 12602; UNESCO 162. Setaria palmifolia (Koen.) Stapf.: Chin
& Badaruddin 1700. Shorea guiso (Blanco)Bl.: UNESCO 497. S. leprosula Miq.: UNESCO 498A. S.
ovalis (Korth.)Bl.: UNESCO 497A. Sida javensis Cav.: Henderson 23117. Solanum biflorum Lour.:
Henderson 22709. S. ferox L.: Stone 9497. S. nigrum L.: Burkill 13939. Sonerila tenera Royle: Hender-
son 21375, 28928. Spathogolottis hardingiana Par. et Rchb.f.: Corner 19 xi’41; Curtis 2150, xi’01. Spon-
dias dulcis Forst.f.: Loh FRI 17184. Stachyphrynium cylindricum (Ridl.)Schum.: Kiah 35410. Stauran-
thera grandifolia Benth.: Henderson 19395, 10 viii 29; Nur & Foxworthy 11813; UNESCO 44, 387. S.
umbrosa C.B. Clke: King’s coll. 7144. Staurochilus fasciatus (Rchb.f.)Ridl.: Stone 6980. Stelechocarpus
cauliflorus (Scheff.)R.E.Fr.: Henderson 23802; Loh FRI 17194. Stemona tuberosa Lour.: Ridley 14780.
Stenotaphrum helferi Munro ex Hk.f.: Kiah 35360. Stenothyrsus ridleyi CB.Clke: Curtis 3149; Hender-
son 23765. Stephania venosa (B\.)Spreng.: Chin 511, 932; Symington 46790. Sterculia lancaviensis Ridl.:
Corner 17 xi’41 (2, from different localities); Stone 7004. S. rubigniosa Vent.: Chin 798, 894; Henderson
23837, 29162; Loh 17242, 17250; Ng FRI 5858; Samsuri & Mahmud 567. Streblus asper Lour.: Chin 144;
Kiah 20 vy ’38. S. ilicifolius (Vidal) Corner: Chew 131, 41021; Chin 151, 785, 832A, 833A; Holttum
Holttum 15118; Keng et al. 6198; Loh FRI 17166; Molesworth-Allen 4789; Ng FRI 5537; Samsuri &
Mahmud 586; Stone 6931; UNESCO 13. S. laxiflorus (Hutch.)Corner: Chew 202. S. taxoides
(Heyne)Kurz: Chin 940, 962; Corner 17 xi ’41; Henderson xi ’34; Samsuri & Mahmud 629; Stone 6929.
Strobilanthes leucopogon Ridl.: Henderson 29140. Strychnos axillaris Colebr.: Chin 984; Corner 38136;
Henderson 21379, 21 xi ’34. S. ignatii Berg.: UNESCO 311. Sumbaviopsis albicans (B1.)J.J.Sm.: Chin
1088.
90 Gard. Bull. Sing. 36(1) (1983)
Taeniophyllum culiciferum Ridl.: Henderson 22240. T. filiforme J.J.S.: Ridley xii 1896; UNESCO
183. T. obtusum Bl.: UNESCO 334. Taenitis blechnoides (Willd.)Sw.: UNESCO 91. Tamarindus indica
L.: Stone 6950. Tarenna appressa (King)Corner: Stone 6932. T. calcarea Ridl.: Burkill 13944; Ridley i
21. T. curtisii (King)Ridl.: Allen 5 ii ’57; Chin 111, 347, 386, 403, 507, 762, 1058, 1698; Corner 37892,
38140; Henderson 29692, 22568; Holttum 15124; Kerr 21734; Kiah 35310; Molesworth-Allen 4293; Ng
FRI 1597, 5576; Nur 34383; Ogata KEP 110162, KEP 110171; Ridley 8241; Stone 5902, 5903, 7476, 8927,
8928, 9522; UNESCO 164, 331. T. pulchra Ridl.: UNESCO 16, 272, 545. T. ridleyi (Pears.)Ridl.: Chin
451; Henderson 23827. Tectaria amplifolia (v.A.v.R.)C.Chr.: Burkill 13940; Chin 2, 770; Chin &
Mahmud 1309; Curtis 3374; Henderson 19394, 19511, 22276, 22379, 23755, 29708; King’s coll. 4713,
5908; Nur 3 xi ’37; Sinclair 9867; UNESCO 551. T. barberi (Hk.)Copel.: Chin 650. T. devexa
(Ktze)Copel.: Chin & Mahmud 1307; Curtis 3375; Henderson 22432; Keng 281; Shimizu & Stone T
14377, 14382, 14383; UNESCO 253, 260. T. griffithii (Bak.)C.Chr.: Chin 425; Shimizu & Stone M 13730.
T. macrodonta (Fee)C.Chr.: Curtis 3376. T. variolosa (Wall.)C.Chr.: Chin 517, 1739; Corner 19 xi ’41;
Corner & Henderson 22807; Holttum 15123; Kiah 35315. Terminalia calamansanai (Blanco)Rolfe: Cor-
ner 29399; Henderson 28946. T. triptera Stapf.: Chin 515; Corner 38128; Keng 6214; Kiah 35414.
Tetracera scandens (L.)Merr.: UNESCO 194. Tetrastigma kunstleri (King)Craib: Chin 494, 787, 833,
1254; Henderson 25201; Stone 8827. T. lanceolarium (Roxb.)Planch.: Kiah 35243. T. peduncularis
(Wall.)Planch.: Chin 789, 810; Samsuri & Mahmud 562. T. scortechinii (King)Gagn.: Henderson 23793.
T. wrayi (King)Craib: Corner & Nauen 25 xi ’41; Ridley 27 xii ’20. Thecostele alata (Roxb.) Par. et
Rchb.: Henderson 22251; UNESCO 169. Thelasis carinata Bl.: Henderson 22255. T. micrantha
(Brongn.)J.J.S.: Chin 162, 457, 1399; Henderson 22460, 25248, 29684; Ng FRI 1911; UNESCO 67, 158.
T. succosa Carr: Chin 429, 1403. T. triptera Rchb.f.: Chin 467; Dransfield 912; Henderson 22249. 22461;
Nur 25155; UNESCO 145, 294, 295. Thelypteris immersa (Bl.)Ching: Carr 22382, 22384; Chin 80, 1805;
Corner & Nauen 25 xi ’41; Henderson 22357; Phang 15 xi ’68. Thespesia populnea (L.)Sol. ex Corr.:
Chin 520. Thrixspermum album (Ridl.)Schltr: Stone 6989. T. amplexicaula (Bl.)Rchb.f.: Henderson
22944. Thunbergia fragrans Roxb.: Chin 1740. Timonius atropurpureus Craib: Chin 1834; Corner 37852;
Henderson 29101; Holttum 15129; Spare 36328; Stone 6993, 9143. Tinomiscium petiolare Miers: ex Hk.f.
et Thoms.: Whitmore FRI 4241. Tinospora crispa (L.)Miers ex Hk.f. et Thoms.: Chin 960; Samsuri &
Mahmud 636. Toxocarpus curtisii K.et G.: Chin 402; Stone 6925; UNESCO 237; Whitmore FRI 12163.
T. pauciflorus Henders.: Chin 353, 791; Stone & Mahmud 12620; UNESCO 184. Trema orientalis
(L.)BI.: Chin 382. T. tomentosa (Roxb.)Hara: Chin 1379. Trichoglottis misera (Ridl.)Holtt.: Henderson
21 xi ’34. T. retusa Bl.: Carr 87; Chin 115, 1253, 1540, 1599; Henderson 22243, 22573, 29685; Kelsall
i 1891; UNESCO 168, 292. T. winkleri J.J.S. var. minor J.J.S.: Carr 423. Trichomanes bipunctatum
Poir.: Chin 89, 554, 693, 1044, 1124, 1465; Henderson 19512; Ridley 8143; UNESCO 81, 186, 591. T.
christii Copel.: UNESCO 39. T. humile Forst.: Chin 687. T. motleyi Bosch: Chin 695. Trichosanthes
tricuspidata Lour.: Corner 20 xi ’41; Henderson 25225; Stone 9077. Tridax procumbens L.: Chin 978.
Trigonostemon aurantiacus (Kurz ex Teijsm. et Binn.) Boerl.: Burkill 6302, 6311; Chew 40995; Hender-
son 29080; Nauen 38124. T. villosus Hk.f.: Whitmore FRI 752. T. viridissimus (Kurz)Airy Shaw: Chin
1034, 1097, 1179; Strugnell KEP 20275. Tristania merguensis Griff.: Nur 25100. T. subauriculata King:
King’s coll. 8253. Trivalvaria macrophylla (B\.)Miq.: Burkill 6342; Chin 470, 531, 998; Chin & Badarud-
din 1697, Sinclair 40060. Tropidia curculigoides Lindl.: Kiah 35400. Tupistra grandis Ridl.: Henderson
29179; UNESCO 565. Turpinia ovalifolia Elmer: UNESCO 683. Tylophora calcicola Henders.: Hender-
son 19583, 25204. T. perakensis K.et G.: Haniff 635. T. tenuis Bl.: Burkill 6351; Henderson 22916.
Typhonium filiforme Ridl.: Henderson 22824; Ridley 9620. T. fultum Ridl.: Henderson 25041; Ridley
8165.
Uncifera tenuicaulis (Hk.f.)Holtt.: Carr 233; Chin 1665; Henderson 21399, 22236. Urena lobata L.:
Chin 579. Urophyllum corymbosum Korth.: UNESCO 546. U. glabrum Wall.: Stone 7495; UNESCO
94. Utricularia minutissima Vahl: Holttum 17427. Uvaria javana Dunal: King’s coll. 5945; Whitmore
FRI 700.
Vaccinium littoreum Miq.: Chin 56; UNESCO 644A. Vandopsis gigantea (Lindl.)Pfitz.: Ridley ii 1897;
Stone 6981. Vatica cinerea King: Kiah 35257, UNESCO 174, 307. Ventilago gladiata Pierre: Chin 981;
UNESCO 138. V. oblongifolia Bl.: Chin 533; Ridley 13349. Vernonia cinerea (L.)Less.: Samat 5. V. cur-
tisii Craib & Hutch.: Chin 508. V. rupicola Ridl.: Corner 37828; Henderson 28927; Ridley 15863. Vibur-
num sambucinum Bl.: UNESCO 117. Villebrunea sylvatica (Miq.)Bl.: Burkill 4419; Chin 78; Stone 8816;
Whitmore FRI 12155. Viscum orientale Willd.: Chin 1747. Vitex pubescens Vahl: Henderson 28943. V.
siamica Willd.: Best 21283; Chew 191; Chin 46, 1388; Corner 38148, 13 xi’41; Henderson 22269, 22464,
Limstone hill flora of Malaya IV 91
25053, 29696; Holttum 15095; Keng et al. 6223; Kiah 35309, 35380; Nur 25151, 34381; Ogata KEP
110180; Ridley 8535; Smith FRI 72558; Stone 5894, 6922, 6994, 8931; Turnau 773, UNESCO 173, 319;
Whitmore FRI 12158. Vitis martinelli Kew ex Planch: Kiah 35392. Vittaria angustifolia Bl.: Corner &
Henderson 22813, 23131; Henderson 29070. V. elongata Sw.: Chin 1705, 1762A.
Wikstroemia androsaemifolia Decne: Chin 146, 412, 1159; Stone 5888, 7308; Whitmore FRI 12164.
W. indica (L.)C.A. Mey.: Chin 521, 1748, 1837; Corner 37837, 37856; Hashim Jaafar 238; Henderson
25093, 29055; Ridley xii 1896; Symington FMS 46733; Whitmore FRI 15063. W. polyantha Merr.:
UNESCO 338. Wrightia dubia (Sims)Spreng.: Kiah 11 iv ’38. W. laevis Hk.f.: Henderson 19599; Kiah
35275.
Xylocarpus granatum Koen.: Henderson 29159.
Zingiber spectabile Griff.: Henderson 23818. Zippelia begoniaefolia Bl.: Henderson 19556. Zizyphus
oenoplia Mill.: Allen 3 iii °57; Henderson 28940, 7 vi ’30; Kerr 21742; Molesworth-Allen 4291; Samsuri
& Mahmud 591; Stone 7471, 9527; Stone & Lewis 9574; UNESCO 468. Z. pernettyoides Ridl.: Chin 343.
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STUDIES IN MALESIAN RUTACEAE
III*. Melicope suberosa, a new species and new generic
record for the Malayan flora
BENJAMIN C. STONE
Department of Botany, University of Malaya, Kuala Lumpur, Malaysia
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983
Abstract
The genus Melicope, previously believed not to be represented in the flora of Peninsular Malaysia, has
been found, in the form of a new species, occurring in montane rainforest on the upper slopes of Gunung
Ulu Kali, Genting Highlands, Pahang, in Malaysia. The only known locality, a densely wooded gully at
1550 m alt., is in the midst of a zone rapidly being cleared by bulldozing and two of the only five trees
found have been felled. The new species, M. suberosa Stone, is apparently related to Bornean species of
Melicope, particularly an as yet undescribed unifoliolate species known from Mt. Kinabalu in Sabah. The
addition of Melicope to the Malayan flora brings the number of Rutaceous genera known to occur in
Peninsular Malaysia up to seventeen.
Introduction
The family Rutaceae as represented in Peninsular Malaysia was last revised in
1972 (Stone, 1972) with sixteen genera and 58 species known to occur in the area.
Since that time, further work on the family by T.G. Hartley (1974, 1979, 1982) and
myself (Stone, Lowry, Scora, and Jong 1973; Stone 1978a, 1978b) improved our
understanding of the Malayan members of the family and resulted in several changes
of name and descriptions of new taxa. Continuing field work in the Peninsula in
recent years has widened our knowledge of the distribution of various taxa. The ex-
ploration of the summit areas of the Genting Highlands, along and just past the
Selangor border in Pahang, with particular concentration on the flora of Gunung
Ulu Kali, the fourth highest peak in Peninsular Malaysia (almost 1800 m) was
recently summarized in a checklist of the flora (Stone, 1981). This flora was already
known to include several endemic taxa (including another species, Maclurodendron
magnificum Hartley, of the Rutaceae, which was only discovered a few years ago),
and all indications were that further exploration would likely result in an increase
in the number of species including novelties in various genera. This conclusion is
demonstrated by the discovery documented here, which not only proves to be an
undescribed species but, in addition, to be a representative of a genus hitherto
unreported from Peninsular Malaysia. This genus, Melicope J.R. & G. Forst., has
long ago been recorded from India (M. indica R. Wight), China, Taiwan, Borneo,
Philippines, Okinawa, New Guinea, Australia, and various Pacific Islands.
As the genus Melicope is being revised currently by Hartley, along with the closely
related genera such as Euodia, specimens of the new species described below were
* Parts I and II of this series appeared in Federation Museums’ Journal (Kuala Lumpur), N.S., 23:
75-116, 1978.
93
94 Gard. Bull. Sing. 36(1) (1983)
sent to Canberra for his examination. He has confirmed (in /itt.) that the species
belongs to Melicope sensu stricto, in the original sense of the Forsters and therefore
also to the more restricted diagnosis which will be using in his restructuring of the
generic division of the tribe Zanthoxyleae. Dr. Hartley has also indicated some rela-
tionships of the new species particularly to Bornean plants, as detailed below.
With the addition of Melicope, the Malayan flora now is known to possess
members of 17 genera of Rutaceae.
RUTACEAE — Zanthoxyleae — Euodiinae
Melicope suberosa B.C. Stone, sp. nov. Plate 1; Figures 1, 2.
Arbor parva etecta usque ad 10 m alta, ramis majoribus fragilibus oppositis, trunco usque ad 15 cm
diametro, cortice crassiter rugososuberoso, partibus glabris floribus et innovationibus primo
ephemeraliter minute sparseque puberulentis exceptis; foliis simplicibus oppositis integris ellipticis,
petiolis ad 55 mm longis, laminis vulgo 10-15 cm longis, 4-8 cm latis (maxime ad 22 x 11 cm), tenue
subcoriaceis, minute glanduloso-punctatis non-aromaticis, costa infra elevato, nervis 7-12-paribus; in-
florescentiis axillaribus, 15-30 mm longis, pedunculo 10-20 mm longo, cymulis 3-4 congestibus
trifloriferis; floribus pallide albo-viridibus, tetrameris; sepalis c. 1.3 mm longis deltoideis scarioso-
marginatis, extus minute puberulentis; petalis imbricatis galndulosis, 4.3 mm longis, anguste ovatis, apice
cucullato, extus subglabris vel minute sparseque puberulentis; staminibus 8 (in specimine viso infertilibus)
inaequalibus albis glabris, filamentis 1-1.5 mm longis, antheris 0.75 mm longis connectivo dorso minute
glanduloso, apice apiculato; disco albo, 1 mm alto; ovario 4-lobato, viridi, 1.4 mm lato, minute
puberulento; stylo 1.25 mm longo, in parte dimidio inferiori minute puberulento; stigmate obscure
4-lobato- capitato; fructibus 4-lobatis, stellato-radiatis, ad 20 mm latis, folliculis ad 9 mm longis, 8 mm
latis, 9 mm altis, uniseminatis, dehiscentibus; endocarpio cartilagineo glabro; seminibus ovoideis c. 7 mm
longis, nitide nigris, endospermio albo.
Type: Stone & Lowry 15338 (KLU, holo: isotypes A, BISH, BO, K, L, SING,
etc.), from Ulu Kali, Pahang, Malaysia.
A small glabrous tree up to 10 m tall, the trunk straight, to c. 15 cm diameter,
branched laxly above with opposite, elongated branches; innovations at first very
minutely sparsely puberulent (hairs simple, clear, 0.15-0.1 mm long, densest on leaf
margins) but quickly glabrate; bark of trunk and older branches becoming thickly
rugose-corky, 2-8 mm thick (or more), spongy, light brown; bark of youngest bran-
chlets 0.5 mm thick, pale tan, sparsely and coarsely fissured; inner bark green; only
the youngest 1 or 2 internodes green and as yet not corky; wood white, very brittle;
leafy branchlets mostly 3-9 mm diameter. Leaves opposite, simple, elliptic, the
petioles to 5.5 cm long, but usually 1.5-3.5 cm long, 3.5 mm diameter, thickened
at base and apex and there later becoming pale corky-barked, the rest remaining
green, subterete, distally shallow-channelled above, not articulated, abscissing at the
base; blade to about 22 x 11 cm but commonly smaller, usually 10-15 x 4-8 cm,
acute or obtuse at apex and at base, thin and rather delicately coriaceous, rather
rapidly wilting; margins entire to somewhat sinuous or shallowly irregularly crenate
distally; upper surfaces medium-dark glossy green, lower surfaces paler apple-green,
somewhat duller; glands pellucid or green but minute, numerous but not evident
without a lens; midrib slightly raised or flat above, well raised beneath; main secon-
dary nerves about 7 to 12 pairs, looped well within (3-5 mm) the margin; tertiary
BCS 15338
Fig. 1. Melicope suberosa B.C. Stone, sp. nov. Details of holotype — a: flowering branch; b: older
branch showing rugose corky bark; c: functionally pistillate flower, side view; d: the same, with
petals removed, stamens sterile; e: capsule in top view; f. capsule in side view; g- capsule in top
view, enlarged; A: one ripe carpel with extruded seed. (From Stone 15338; b,e-f, to same scale
as @.).
95
Fig. 2. Melicope suberosa B.C. Stone, sp. nov. Some details of the seedlings — a: simple leaf; b:
bifoliolate leaf; c: trifoliolate leaf; d: petiole apex from the trifoliolate leaf; e: leaflet, margin
showing glands and crenulation. (From Stone 15349).
nerves and reticulations rather obscure in life, fine but evident in sicco; fresh blades
very slightly scented on bruising.
Inflorescences axillary, pedunculate, up to 15-30 mm long, the peduncle somewhat
compressed, 10-20 mm long, to 2.5 mm wide when fresh, channelled and somewhat
shrunken when dry, subglabrous, but at first with minute and rather scattered con-
ical 1-celled whitish hairs c. 0.1 mm long; bracts and bracteoles early caducous, c.
1 mm long, deltoid, scurfy; cymes congested, usually 3-flowered, or up to 3 or 4 par-
tial cymes crowded near apex of inflorescence, with very short pedicels c. 1 mm
long. Flowers greenish-white, the calyx about 2 mm high, the 4 deltoid lobes c. 1.3
mm long, green with pale or white scarious margins, minutely puberulent externally;
petals 4, imbricate, folded, white tinged green, glandular, c. 4.3 mm long, narrowly
ovate, the margins pale, the apex cucullate, glabrous within and subglabrous to
minutely sparsely puberulent externally; staminodes in pistillate flower 8, glabrous,
white, small, thin, with 4 slightly longer, anthers devoid of pollen, filaments 1-1.5
mm long, tapered, anthers 0.75 mm long, the connective dorsally with several small
pale glands, the apex minutely apiculate; disk white, 1 mm high, slightly wider than
the ovary; ovary 4-lobed, green, to 1.4 mm wide, minutely puberulent; style 1.25
mm long, pale green, puberulent in lower half; stigma pale, capitate but 4-lobed,
minutely papillose, glabrous. Pedicels in fruit pale, corky, 3-4 mm long, 2.5 mm
diameter. Fruits 4-lobed, or by abortion 3- to 1-lobed, dark green, radiate, dehis-
cent and turning brown, to 20 mm wide, each lobe ovoid, to 9 mm long, 8 mm wide,
9 mm tall, glabrate, glandular, 1-seeded. Endocarp thin, horny, glabrous,
detaching. Seed ovoid, c. 7 mm long, testa thin, glossy black; endosperm white.
96
Melicope, new for Malaya 97
MALAYSIA: Pahang: Genting Highlands, Gunung Ulu Kali, 1550 m alt., below hotel complex in rem-
nant patch of montane forest; small tree to 10 m tall, trunk and all branches with thick, rugose, softly
corky bark and brittle wood; leaves all simple, opposite, as are the branches; flowering and fruiting
simultaneously; 17 November 1982, B.C. Stone & J.B. Lowry 15338 (KLU and to be distributed).
A very characteristic species. The most remarkable features are the thick, deeply
corrugated but soft and corky bark, which is a light or medium brown and covers
all the trunk and main branches and even twigs, except the youngest; and the
uniformly simple, non-articulate leaves, which are very slightly undulate-subcrenate
toward the tips, glossy on both sides, somewhat paler below, with rather con-
spicuous nerves. The leaves when crushed are only very slightly scented, and the
fruit pericarp is also but faintly odorous. The capsular follicles dehisce, the seed is
extruded but remains attached on its placenta, and the pericarp quickly withers and
turns brown. Fruits with 1-4 ripening lobes are found, but the lobes seem to be
uniformly 1-seeded. Only pistillate flowers were seen, and the number of staminodes
was uniformly 8, with no fertile anthers even in the youngest buds examined. The
minute hairs on the ovary are rather ephemeral and the fruits appear quite glabrous
when ripe.
Ecology: Five or six apparently full-grown individuals were found, all close
together in a moist, originally well-shaped gully with dense vegetation and granite
boulders, at 1550 m altitude on the uppermost slopes of G. Ulu Kali (the location
is about 50 m higher than the Sri Genting Villa, and is on the SE. side of the road
leading up to the Genting Hotels complex). The gully is rich in bryophytes but lacks
Sphagnum. Currently it is being cut, the upper end opened and the understorey now
largely exposed to much more insolation than originally must have been the case.
So far, this species has not been found elsewhere; this small gully population is clear-
ly threatened. (Seeds, seedlings and cuttings have been taken for cultivation, and it
is hoped to distribute this species widely). Fruiting was rather abundant in the
population and ripe seeds were visible on most of the trees at the time of collection
(November). In addition, there were numerous seedlings underneath, especially on
the rich organic matter.
All the individuals found were pistillate; the flowers, though possessing stamens,
do not produce pollen, the anthers being somewhat rudimentary. It seems possible
that the seeds found, and perhaps the seedlings as well, are the result of a non-sexual
process of reproduction. Further exploration is required in the hope of finding addi-
tional individuals and particularly to find staminate (or perhaps bisexual)
individuals.
The canopy trees in the area include Eugenia, Garcinia, Symingtonia, Lithocar-
pus, Litsea, and various other genera (see Stone, 1981), forming a canopy about
18-26 m tall, i.e. about twice as tall as the Melicope trunks. It is clear that this species
is essentially an understorey species of the primary montane forest, and seems not
to be a pioneer or colonizer species like the Euodias. The very brittle, easily broken
wood, and quick wilting character of the Melicope is very noticeable and somewhat
unusual in this forest. Other members of the substorey stage which reach about the
same dimensions as the Melicope are Polyosma parviflora King, Prunus arborea
(Bl.) Kalkm., Casearia capitellata Bl., and somewhat smaller shrubs include
98 Gard. Bull. Sing. 36(1) (1983)
Chasalia minor Ridl., Pinanga polymorpha Becc., and Arthrophyllum stonei Lim.
Relationships: The most similar species is one found on Mt. Kinabalu in Sabah,
represented by J. & M.S. Clemens 51077, 51184, G. Mikil SAN 46579, Chew, Corner
& Stainton 197, and RSNB 4547 and 4558. This species, as yet undescribed, has
unifoliolate leaves. It differs from M. suberosa in having smaller flowers (with
petals about 2.5 mm long), a glabrous perianth, and a glabrous gynoecium. There
are also on Mt. Kinabalu plants which are very similar, grading to a taxon with
trifoliolate leaves (e.g. J. & M.S. Clemens 29810, 31048, 32562, 51052, Carr 27490,
and Stone 11398). The latter taxon with trifoliolate leaves (represented by J. & M.S.
Clemens 29477, 33623, Carr 27663, and Ding Hou 232) is in turn highly similar and
may grade into a Sarawak taxon also with trifoliolate leaves (represented by Sibat
ak Luang S. 22129 and Banying ak Nyudong S. 19031), which has smaller leaves,
more delicate and longer inflorescences, and smaller fruits.
Amongst the already described species of Melicope, there do not appear to be any
strikingly close to the new species. Some, such as M. mindanaensis Elmer, have
unifoliolate leaves, but differ in important floral features. M. indica Wight differs
in its more oboval leaves, pubescent stigma, and blunt anthers. M. monophylla
Merr. differs in its hirsutulous branches, villous ovary, very short style and rugose
cocci. M. curranii Merr. differs in its smaller flowers, oblong-obovate to oblong-
lanceolate leaves, and very long petioles. M. unifoliolata Merr. and M. helferi
Hook. fil., of Borneo and Burma respectively, are not true Melicopes and have been
reduced to Maclurodendron by Hartley (1982). The others mostly have trifoliolate
leaves, or differ in other important respects.
In Engler’s treatment (1931) Melicope was subdivided into four sections, of which
nowadays the last, Tetractomia, is best regarded as a well-defined genus of its own,
as J.D. Hooker had originally ranked it. The other three are Entoganum (Banks ex
Gaertn.) Engl., to which most of the species pertain; Astorganthus (Endl.) Engl.,
including only M. simplex A. Cunn. of New Zealand; and Brombya (F.v.Muell.)
Engl. with only M. platynema (F.v.Muell.) Engl. of Queensland. The latter two are
remote from the present new species.
Thus the relationships of M. suberosa appear so far to be with a complex of taxa,
mostly undescribed, which occur in Borneo, especially on Mt. Kinabalu.
Seedlings: The seedlings of Melicope suberosa are noteworthy in that they occa-
sionally produce some trifoliolate, or intermediate (bifoliolate) leaves. These seem
always to be among the first four or five true leaves produced by the seedling. Later
stages appear to have only simple leaves. These, as in the adult plants, are not ar-
ticulated and can scarcely be called unifoliolate. Details of these seedling leaves are
shown (Fig. 2).
Plate 1. Melicope suberosa B.C. Stone, sp. nov. a: leafy shoot with fruits. b: older stem, showing corky
rugose bark. c: leafy stem with inflorescences; flowers at anthesis. d: ripe capsules. (All from
the holotype; photographs by the author).
99
100 Gard. Bull. Sing. 36(1) (1983)
Acknowledgements
Thanks are expressed to Dr T.G. Hartley (CSIRO, Herbarium Australiense,
Canberra) for his very helpful and encouraging remarks and his indication of the
specific relations of M. suberosa. I am also grateful to Dr Brian Lowry for compa-
nionship in the field and for the good luck he always seems to bring.
References
Engler, A. (1931). Rutaceae. Jn Engler, A., and H. Harms, Die Natiirlichen
Pflanzenfamilien. Bd. 19a: 187-357. Leipzig: W. Engelmann.
Hartley, T.G. (1974). A Revision of the Genus Acronychia (Rutaceae). J. of the Ar-
nold Arb. 55: 469-523, 525-567.
(1979). A Revision of the Genus Tetractomia (Rutaceae). J. of the Arnold
Arb. 60: 127-153.
(1982). Maclurodendron: a new genus of Rutaceae from Southeast Asia.
Gard. Bull. Sing. 35: 1-19.
Stone, B.C. (1972). Rutaceae. Jn Whitmore, T.C. (ed.) Tree Flora of Malaya. Forest
Department, Malaysia. Vol. 1: 367-387. Kuala Lumpur: Longmans.
(1978a). Studies in Malesian Rutaceae, I: Notes towards a revison of the
genus Glycosmis Correa. Fed. Mus. J., new ser., 23: 75-110.
(1978b). Studies in Malesian Rutaceae, II: New records and new taxa of
Aurantioideae from Borneo. Fed. Mus. J., new series., 23: 111-116.
Stone, B.C., J.B. Lowry, R.W. Scora, and K. Jong. (1973). Citrus halimii: a new
species from Malaya and peninsular Thailand. Biotropica 5(2): 102-110.
CONTRIBUTIONS TO THE FLORA OF
THE SOLOMON ISLANDS
II. Five new combinations in Araliaceae
BENJAMIN C. STONE
Department of Botany, University of Malaya
Kuala Lumpur, Malaysia
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983
Abstract
Five species of Boerlagiodendron (Araliaceae), including four from the Solomon Islands and one from
the New Hebrides, are transferred, according to nomenclatural necessity, to Osmmoxylon.
Some years ago I revised briefly the East Melanesian species of Boerlagiodendron
Harms, describing two new species and demonstrating that four species could be
found in the Solomon Islands and another in the New Hebrides. Since then it has
been shown that the generic name Osmoxylon Miq. must be used for this very
distinctive group of plants. As the Araliaceae, apart from Schefflera, has now been
published in the Flora Malesiana (Philipson, 1979), it is appropriate to prepare a
correct nomenclatural list of the East Melanesian species to supplement my original
paper (Stone, 1962). This list follows.
Osmoxylon Miq.
Ann. Mus. Bot. Ludg.-Bat. 1 (1863) 5. Philipson, Blumea 23 (1976) 99; Fl. Males.
ser. I, 9(1) (1979) 31-53. -- Eschweileria Zipp. ex Boerl. Ann. Jard. Bot. Btzg 6
(1887) 112, non Eschweilera Mart. 1828. -- Pseudo-sandalum O.K. Rev. Gen. PI.
1 (1891) 271, as ‘‘Pseudosantalum’’, nom. illegit. -- Boerlagiodendron Harms, E.
& P. Nat. Pfl. Fam. 3, 8 (1894) 31; Hutchinson, Gen. Fl. Pl. 2 (1967) 72.
1. Osmoxylon puniceopolleniferum (Stone) Stone comb. nov.
Boerlagiodendron puniceopolleniferum Stone, Proc. Biol. Soc. Wash. DC. 75
(1962)"27, f. 1.
SOLOMON ISLANDS: Malaita, Stone 2450 (BISH holo, A, K, US iso).
2. Osmoxylon tetrandrum (C.T. White) Stone comb. nov.
Boerlagiodendron tetrandrum C.T. White, J. Arn. ARB. 31 (1950) 102;
Walker, For. Brit. Sol. Is. Prot. (1948) 98.
SOLOMON ISLANDs: San Cristoval, Wa/ker BSIP 260 (K).
3. Osmoxylon orientale (Guill.) Stone comb. nov.
Boerlagiodendron orientale Guillaumin, J. Linn. Soc. Bot. 51 (1938) 554.
NEW HEBRIDES: Espiritu Santo, J. & Z. Baker 258 (K).
101
102 Gard. Bull. Sing. 36(1) (1983)
4. Osmoxylon reburrum (Stone) Stone comb. nov.
Boerlagiodendron reburrum Stone, Proc. Biol. Soc. Wash. DC. 75 (1962) 30,
fees
SOLOMON ISLANDS: Malaita, Stone 2397 (BISH holo, US iso).
5. Osmoxylon russellense (Philipson) Stone comb. nov.
Boerlagiodendron russellensis Philipson, Bull. Brit. Mus. (Nat. Hist.) Bot. 1(1)
(1951) 12.
SOLOMON ISLANDS: Russel Islands, R.7. Brice 18 (A holo).
References
Philipson, W. R.(1979). Araliaceae, part 1. Flora Malesiana ser. I, 9 (1): 1-105.
Stone, B. C.(1962). Boerlagiodendron (Araliaceae) in Eastern Melanesia. Proc.
Biol. Wash. DC. 75: 25-32.
rr
Annotated List of Seed Plants of Singapore (VIII)*
HSUAN KENG
Department of Botany, National University of Singapore
Index to Families
Page Page
Apocynaceae 112 Myrsinaceae 117
Asclepiadaceae 123 Oleaceae 121
Ebenaceae 106 Plantaginaceae 111
Epacridaceae 103 Plumbaginaceae 110
Ericaceae 103 Sapotaceae 103
Gentianaceae 112 Styracaceae 115
Loganiaceae 111 Symplocaceae 116
II. Angiospermae-Dicotyledons (cont’d)
111. Ericaceae
Rhododendron indicum Sweet
A densely branched shrub; leaves narrowly lanceolate to oblanceolate, 1.8-3.5
cm long, red-brown hairy. Flowers rose-red to scarlet, with round lobes, borne
in short clusters at branch-tips. A pot plant introduced from Japan, has never
set fruit here.
Rhod. longiflorum Lindl.
An epiphytic shrub with bright red flowers, formerly collected from lofty
Hopea trees near the top of Bukit Timah, Ridley 2030 in 1891, now extinct.
112. Epacridaceae
Styphelia malayana Spr. (= Leucopogon malayanus Jack)
A shrub; leaves narrow and stiff with a sharp-pointed tip. Flowers white, in ax-
illary clusters. Fruit red, baccate, Formerly found on the sea-shore near Changi,
Tanah Merah, Ridley 1832, now extinct.
113. Sapotaceae
Key to the genera
APR SEDAIRTS WINE WORWROLIS: Bitet:. Sate vce sac oareis a octhee Sales Soe ae oboe Ganua, Madhuca, Payena
1. Sepals 5, 6 or 8, in one or two whorls
* continued from Gdns’ Bull. Sing. 35: 103, 1982.
103
104 Gard. Bull. Sing. 36(1) (1983)
2. Sepals 5, in one whorl
3. SStaminodesiS: 3. .005 setorsce rele toraist ee eres tavedeee ey terreleitere mieieene Planchonella, Pouteria
3. Staminodes absent: aacncwc ssc este eee ea Oe ROO ace eee Chrysophyllum
2. Sepals 6 or 8, in two whorls
4. Sepals 6
§ Stamens:6- ccc arouses ete rautavalecars evstere er ekeue ate nieneereurte eletelorertere rarer orers ree onenetate Manilkara
§:Stamens’ about HQ) 2505065: afs ois savaievare siesta, acuta aber ereravale tate el @kayoiaie: a/aiere sie tarelevsieasiats Palaquium
As Sepals:8 ..cicrsicsiwiw stevens drones We en Sees Sete: ee tint. ee aera Mimusops
Chrysophyllum cainito L.
A tree with elliptic, drooping leaves. Leaf-blades coppery-silvery beneath.
Corolla purplish white. Fruit large, broadly ellipsoid to spherical, 5-7 cm across,
white or purple, with a star-like core in transverse section (hence Star-apple).
Native of tropical America, occasionally planted.
Chrys. lanceolatum DC. (= C. roxburghii G. Don)
Tree; leaves dark green, with numerous pairs (12-35) of parallel secondary
veins. Corolla yellow, 5-merous; staminodes absent. Fruit globose, 5-lobed.
Tanglin, Bukit Timah (Sinclair SF 40200).
Ganua kingiana (Brace) v.d. Assem (= Madhura kingiana (Brace) Lam)
Large tree; leaves closely clustered. Flowers white, 1-6 per cluster, above the leaf
scars. Bukit Timah, Ridley 6294, type of Bassia kingiana.
Ganua motleyana Pierre ex Dubard
Tree; leaves spiral, glabrous. In freshwater and peat swamps, Seletar, Jurong,
Kranji, Bukit Mandai, Changi (Ridley 5645).
Ganua sessilis (K. & G.) Lam (= Payena sessilis K. & G.)
Leaves obovate, the base gradually tapered downward, hence appearing sessile.
Tuas, Goodenough 5076, type.
Madhuca malaccensis (Clarke) Lam
Small or medium tree; leaves loosely clustered. Flowers white or yellow, many
in axillary clusters. Gardens’ Jungle, Ridley 6521.
Madh. sericea (Mig.) Lam
Medium tree; leaves yellowish brown to silvery, velvety below. MacRitchie
Reservoir, Sinclair SF 39656.
Manilkara zapota (L.) van Royen (= Achras zapota L.)
Small tree; leaves glabrous, dark green below. Fruit roundish ovoid, up to 9 x
7 cm, the rind thin, brown; flesh pinkish, juicy, in which are embedded about
10 black seeds. The milky latex tapped from the trunk is known as Chicle or
Chicle gum, the main ingredient of chewing gum. A native of tropical America,
locally grown for its edible fruit, Chicku. Aux
Mimusops elengi L.
Small or medium tree; leaves alternate or laxly spirally arranged, margin wavy
and curled; flowers white, with 8 flat and hairy staminodes. Native to India and
Thailand, often cultivated as an ornamental and roadside tree.
Seed plants of Singapore VIII 105
Palaquium gutta (Hook. f.) Baill.
Medium-sized tree, buttressed; leaves obovate to oblong-elliptic, spirally ar-
ranged, golden brown velvety below. Flowers pale green, in small axillary
clusters. Fruit ovoid, 2.5 cm across. This species furnishes the top grade of Gut-
ta Percha which was an important commodity for insulation material during the
last century. Probably introduced from the Malay Peninsula, formerly widely
planted; a small patch can still be found at the foot of Bukit Timah Nature
Reserve.
Pal. hexandrum (Griff.) Baill.
Large tree; leaves obovate or oblong-elliptic, to 20 cm long; petioles glabrous,
narrowly grooved above. Bukit Timah, Ridley 11373.
Pal. microphyllum K. & G.
Large tree; leaf-blades spathulate, less than 11 cm long; petioles finely hairy.
Bukit Timah, Sinclair SF 38245.
Pal. obovatum (Griff.) Engl.
Medium-sized or large tree; leaf-blades obovate, leathery, to 35 cm long;
flowers green, crowded along the twigs behind the leaves. Tanglin, Changi,
Pulau Senang (Burkill & Kiah HMB 503). Vern. Nyatoh.
Pal. oxleyanum Pierre
Small to medium-sized tree; leaf-blades obovate, to 28 cm long; petioles strong-
ly grooved above; very similar to P. gutta. Pulau Dammar Darat, Goodenough
in 1892.
Pal ridleyi K. & G.
Large tree; leaf-blades obovate to elliptic, to 12 cm long. In swamps; Kranji,
Ridley 4796.
Pal. rostratum (Mig.) Burck (= P. bancanum Burck)
Large tree; leaf-blades obovate to spathulate, to 80 cm long. Gardens’ Jungle,
Bukit Timah (Mat 6/34).
Pal. xanthochymum Pierre
Large tree; leaves obovate to spathulate, to 15 cm long. In freshwater swamps,
Changi, Ridley 3639a.
Payena lucida (G. Don) DC.
Small or medium tree; leaves oblong, to 25 cm long; flowers in small axillary
clusters; sepals 4, in two pairs; corolla white, mostly 8-lobed; stamens 16; fruit
ovoid to ellipsoid, 2 cm across. Seletar, Ridley 5644, type.
Pay. maingayi Clarke
Medium-sized or large tree; leaves oblong, to 30 cm long, velvety beneath.
Flowers white or pale yellow. Chua Chu Kang, Ridley 6696, Mandai Road.
Pay. obscura Burck (= P. havilandii K. & G.)
Leaves elliptic, to 15 cm long, glabrous beneath. Bukit Timah, Ngadiman SF
35800.
106 Gard. Bull. Sing. 36(1) (1983)
Planchonella maingayi (Clarke) Van Royen (= Sideroxylon maingayi Clarke)
Tree; leaves obovate. Like the next species, but leaves glabrous below, tertiary
veins transverse. Less common; Gardens’ Jungle, Bukit Timah (Liew 36465).
Planch. obovata (R.Br.) Pierre (= Sideroxylon ferrugineum Hook. & Arn.)
Small or medium-sized tree; leaves obovate, usually velvety below, tertiary veins
reticulate; flowers small, 5-merous; staminodes tongue-shaped, alternate with
corolla-lobes; fruit obovoid, small. In mangroves and on the sea-coast; Pulau
Dammar Darat, Goodenough 2763.
Pouteria malaccensis (Clarke) Baehn. (= Lucuma malaccensis Dubard)
Large or medium-sized tree; flowers small, in leaf-axils, 5-merous; staminodes
5, alternate with corolla-lobes. Seletar, Mat 6499.
114. Ebenaceae
Diospyros argentea Griff.
Small tree with dense golden-yellow hairs on various parts; leaves oblong, 15-32
cm long; flowers 4-5 merous, in axillary cymes, usually dioecious; fruit round,
about 5 cm across, covered with red hairs. In dense jungle, Bukit Timah,
Changi (Hullett 410), Chua Chu Kang.
Diosp. buxifolia (Bl.) Hiern. (= D. microphylla Bedd.)
Tree; leaves small, ovate, 1.2-5 cm long. Fruit ellipsoid, 1.1 cm across, 1-seeded.
Producing black heartwood, the Malacca Ebony. Bukit Timah, MacRitchie
(Sinclair SF 39482).
Diosp. clavigera Clarke
Large tree; leaves obovate, 5-13 cm long. Fruit round, 1.2 cm across. Sungei
Morai, Bukit Timah (Ngadiman SF 36396).
Diosp. diepenhorstii Mig. (= D. pyrifera Ridl.)
Leaves oblong, 15-30 cm long. Fruit obovoid or oblong, 8 cm across. Bukit
Timah, Ridley 8101.
Diosp. discolor Willd.
Small, densely crowned tree; leaves lanceolate or oblong-elliptic, 8-30 cm long,
densely hairy below; fruit depressed globose, 5-12 cm across, densely hairy.
Native to the Philippines, occasionally planted for the edible fruits (Buah
Mentega). #4%
Diosp. ferrea (Willd.) Bakh. (= Maba buxifolia Pers.)
Leaves small, elliptic or obovate, 1.6-6 cm long. Fruit globose, 0.8 cm across.
Often near the sea, Tempinis, Kranji (Ridley 6703).
Diosp. kaki Thunb.
Fresh fruits of the Chinese persimmon ( ## ) are imported from China, Japan
and Java usually during the end of the year. Compressed dried ones are sold
in stores almost throughout the year.
Seed plants of Singapore VIII 107
Diosp. lanceifolia Roxb. (= D. lucida Wall.)
Leaves lanceolate to oblong-elliptic, 4.5-15 cm long. Fruit globose, about 2 cm
across, beaked. In the forest, sometimes near the sea, Bukit Timah, Changi,
Loyang, Tempinis.
Diosp. maingayi (Hiern.) Bakh. (= D. bilocularis Oliv.)
Leaves elliptic, 6-17 cm long. Fruit oblong, 3.5 cm across. Gardens’ Jungle,
Changi (Goodenough s.n. in 1890).
Diosp. pilosauthera Blanco var oblonga (Wal. ex G. Don) Ng
Leaves oblong, 15-33 cm long. Fruit ovoid, 2.5 cm across, Bukit Timah, Tan-
jong Gul, Gardens’ Jungle (Abu Kassim 267).
Diosp. styraciformis K. & G.
Leaves lanceolate to oblong, 5-13.5 cm long. Fruit globose, to 2.5 cm across,
pointed. MacRitchie, Samsuri 1463.
Diosp. venosa Wall. ex A. DC. (= Maba venosa (Wall.) K. & G.
Leaves lanceolate to elliptic. Fruit globose to oblong, 1.3-1.7 cm across.
Changi, Ridley 4667.
115. Styracaceae
Styrax benzoin Dryand
Medium-sized tree; buttressed. Leaves ovate to elliptic, 6-20 cm long, glaucous
beneath; corolla white, fragrant; fruit depressed-globose to globose, 2-3.8 cm
across. Gum benzoin is a fragrant resin, tapped from deep incisions into the
bark. It is used as incense and in medicine. Bukit Timah, Seletar (Ridley 2650),
Chua Chu Kang, Gardens’ Jungle. Vern. Kemenyan. £8.&
Styr. crotonoides Clarke
Leaves elliptic to broadly ovate, 8.5-20 cm long, densely rust-hairy below; fruit
ovoid to round, to 3 cm across. Recorded in Singapore by a single collection,
Wallich 7848.
116. Symplocaceae
Symplocos adenophylla Wall, ex D. Don
Shrub or small tree; young twigs and undersurface of leaves covered with red
brown hairs. Leaves ovate-lanceolate, 4-14 cm long; flowers white; fruits ellip-
soid, to 0.8 cm long. The fruits of Symplocos are drupes, usually bluish when
ripe, crowned with a persistent calyx, and have a ribbed stone. Seletar (Ridley
2752b), Tuas.
Sympl. barringtonifolia Brand (= S. rigida Clarke)
Small tree; leaves oblong or elliptic, 26-31 cm long; flowers white; fruit oblong,
pointed, to 3 cm long. Changi, Ridley 5960.
108 Gard. Bull. Sing. 36(1) (1983)
Sympl. celastrifolia Griff. ex Clarke
Leaves elliptic to ovate, 5.5-12 cm long; fruit ovoid, to 1 cm long. Jurong,
Ridley 8423.
Sympl. fasciculata Zoll.
Leaves oblong-lanceolate, 5-11 cm long; fruit ovoid, with a short beak, blue,
6 mm long. In the forest and belukar, Bukit Timah, Chua Chu Kang, Tanglin,
Bukit Mandai (Ridley 3626a).
Sympl. odoratissima (Bl.) Choisy ex Zoll.
Leaves narrowly elliptic to obovate, 7.5-17 cm long; fruit ovoid, 0.8-1.5 cm
long. Bukit Timah, Ridley 4428.
Sympl. rubiginosa Wall. ex DC.
Leaves narrowly elliptic to obovate, 15-35 cm long; fruit ellipsoid, 0.8-1 cm
long. Chua Chu Kang, Bukit Timah (Ngadiman SF 36363).
117. Myrsinaceae
Key to the genera
1. Woody climbers
2. Ovary and fruitssuperior, seedisolitanys elOOOSe mere aeianeeieieiee rere eters Embelia
2. Ovary and fruit half-inferior, seeds numerous, angular ...................-.0eeeeeee Maesa
1. Erect subshrubs, shrubs or small trees; fruit 1-seeded
3. Herbaceous subshrubs, rhizomatous; flowers in small fascicles along the rachis of a racemose in-
5100) ¢ Cielo | 6 a ee Io i hi a ani aiden ctheitow toi Ga cic Labisia
3. Shrubs or small trees, not rhizomatous; flowers in umbellate or other inflorescences, but not as
above
4. Fruit cylindric; curved; acute; pericarpidry. leathenys .-r1e ele terete Aegiceras
4. Fruit globose, pericarp fleshy
5. Corolla-lobes contorted in bud, flowers perfect, style and stigma tapering to a point
ee ete ner te i Oe ono SOO Aa. oe ROU aD Anns GOOD Elo ot O06 Ardisia
5. Corolla-lobes imbricate or valvate, stigma tongue-shaped ..................... Rafania
Aegiceras corniculatum (L.) Blanco
Shrub; leaves thick-leathery, obovate, 4-8 cm long; flowers pale pinkish, 10-20
in terminal and axillary umbels. Fruit cylindric, acute, curved, pericarp
leathery. In mangroves, Tuas, Changi (Ridley s. n. in 1890), Lim Chu Kang,
Pulau Ubin.
Ardisia corolata Roxb. (= Ard. stylosa Miq.)
Shrub or small tree; leaves papery, oblong-lanceolate, 12-25 cm long, variable,
reddish-brown beneath. Bukit Timah, Gardens’ Jungle, Changi Road, Jurong
Road (Ridley 11342).
Ard. crassa Clarke
Shrub or small tree; leaves leathery, narrowly oblong; panicles rusty pubescent,
flowers pink. Rare, Seletar, Chua Chu Kang (Ridley 3844).
Seed plants of Singapore VIII 109
Ard. crispa DC. (= A. crenata Roxb.)
Glabrous shrub; leaves thinly leathery, lanceolate, with a crenate margin, 5-10
cm long; flowers pink, in short umbellate racemes. In open places, Alexandra
Road, Changi, Sentosa Island (Samsuri 1347).
Ard. lanceolata Roxb.
Shrub or small tree; leaves leathery, lanceolate or elliptic-oblong, 15-28 cm
long, petioles winged; panicles terminal. Seletar, Bukit Timah (Ridley 6436).
Ard. littoralis Andr. (= A. elliptica)
Large shrub; leaves leathery, obovate, 8-12 cm long, nerves nearly invisible;
flowers rose pink, 6-8 in an umbel, corolla gland-dotted. Drupe flattened,
globose, red then black. In tidal swamps and muddy river banks. Changi,
Ridley 20115.
Ard. ridleyi K. & G.
Slender spreading shrub; leaves membranous, oblong-lanceolate, 10-20 cm
long. Sole record in Singapore: Ang Mo Kio. No specimens available.
Ard. singaporensis Rid.
Small tree, young parts densely red-scurfy; leaves thinly leathery, elliptic-
lanceolate, gland-dotted, 10-18 cm long. Rare, Pulau Ubin (Ridley 2816, type),
and Changi Road; probably extinct.
Ard. teysmanniana Scheff. (= Pimelandra wallichii DC.)
Shrub or small tree; branches rusty tomentose; leaves papery, oblong or
oblanceolate, 30-45 cm long, margin entire; flowers in branched short racemes.
Gardens’ Jungle, Bukit Timah (Corner s.n. in 1940), Changi, Nee Soon.
Ard. tuberculata Wall.
Glabrous shrub; leaves leathery, elliptic, 8-15 cm long, coppery scaly beneath;
flowers pink, in a lax terminal panicle. In forests, Bukit Timah, Seletar (Ridley
432), Jurong, Gardens’ Jungle.
Ard. villosa Roxb.
Small shrub; leaves papery thin, oblong-lanceolate, villose beneath, 10-20 cm
long. In dense forests, Pulau Ubin (Ridley 2809).
Embelia amentacea Clarke
Slender climber, branches brown velvety; leaves papery, oblong, 3-8 cm long,
pubescent beneath; flowers small, in panicles of slender hanging racemes. In
secondary forests and their edge; Bukit Timah Road, Changi Road.
Emb. canescens Jack (= E. garcinifolia)
Slender woody climber; inflorescences terminal, red-hairy; fruit small, black.
Catchment forests (Goodenough 1822), Changi (Ridley 5927).
Emb. coriacea DC.
Large liana; leaves stiff, coriaceous, oblong, 16-25 cm long; panicles very large,
pendent, corolla white; fruit black. In secondary forests, Seletar, Bukit Mandai,
Chua Chu Kang (Ridley 2812).
110 Gard. Bull. Sing. 36(1) (1983)
Emb. lampani Scheff.
Woody climber; leaves oblong, 4-8 cm long. In forest edge, Changi, Bukit
Timah, Gardens’ Jungle, Catchment forests (Ridley s.n. in 1906).
Emb. ribes Burm.
Slender woody climber; in forest edge, Changi, Bukit Timah (Ridley s.n. in
1889), Gardens’ Jungle.
Emb. ridleyi K. & G.
Large climber; leaves thinly papery, oblong-acute, 10-18 cm long; panicles ax-
illary. Forest edge; Chua Chu Kang, Bukit Mandai. Probably extinct.
Labisia pumila (Bl.) Benth. & Hook. f. (= L. pothoina Lindl.)
A shrublet; leaves variable, usually lanceolate-elliptic, crenulate, petioles short
or long, sometimes winged (var. alata). Corolla pink, flowers in small clusters
on a spike-like inflorescence; fruit globose, bright red, 3-4 mm across. In dense
forests; Sungei Bajau, Bukit Timah, Kranji (Ridley 1799a), Changi. Vern. Akar
Fatimah.
Maesa ramentacea Wall.
Scandent shrub or small tree; branches often bearing masses of abortive branch-
lets; leaves membranous, ovate-lanceolate, 8-20 cm long; flowers with a white
corolla, very small, in axillary panicles. In secondary forests, Catchment
forests, Bukit Timah (Ridley 2064).
Rafania avensis (Bl.) Mez (= Myrsine avensis DC.)
Small tree; leaves oblong-lanceolate, 3-6 cm long, nerves obscure; flowers in
compact, few-flowered umbels. Drupe globose, 3-4 mm across. In mangrove
forests, Bajau, Tanjong Gul (Sinclair SFN 39637).
Raf. umbellulata (Wall.) Mez (Myrs. umbellulata A. DC.)
Shrub or small tree; leaves oblanceolate, 4-8 cm long, nerves visible; flowers 3
or more in an umbel; drupe globose, 7-8 mm across. Near the sea, Changi, Chua
Chu Kang (Ridley 6827).
118. Plumbaginaceae
Plumbago auriculata Lamk. (= P. capensis Thunb.)
A dwarf shrub, much branched; flowers in umbel-like clusters, corolla tubular,
light blue. Native of S. Africa, planted as an ornamental or hedge. # it ©
Plumb. indica Linn. (= P. rosea L.)
Corolla rose-pink or red. Native of India.
Plumb. zeylanica Linn.
Corolla white. Native of tropical Asia; cultivated or as garden escape on road-
side, waste ground or near villages.
Seed plants of Singapore VIII 111
119. Plantaginaceae
Plantago major L. (or as P. asiatica Michx)
A herb with thin or thick rhizomes; leaves lanceolate to ovate, in a radical
rosette; flowers small, white, in a spike-like inflorescence. A cosmopolitan
weed, used in herbal medicine. 227% -
120. Loganiaceae
Key to the genera
1. Leaves with one main vein from the base; tendrils often present ............-....... Strychnos
1. Leaves with 3-5 basal veins; no tendrils
2. Goerolia-lobes imbricate or contorted; fruit baccate ........2..-....2..-0.022..0005- Fagraea
Za eormmlalopes valvate m bud; Capsules 2-valved .. 2. 2-222. .c. oes cece ccc ese scces Norrisia
Fagraea auriculata Jack
Epiphytic or climbing shrub; leaves stalked, with auricles present at the base of
stalk; corolla tubular, large, to 30 cm long. Pulau Pawai (Sinclair SF 38900).
Fagr. crenulata Maingay ex Clarke
Large tree, trunk and branches often thorny; leaves obovate, 18-35 cm long,
sessile, like those of the cabbage plant (hence Cabbage Tree). Native to the
swamp-forests in S. Malaya, cultivated in large gardens or along roadsides.
Superficially it resembles the Ketapang tree (Terminalia catappa), but the leaves
are green and larger. Vern. Birah.
Fagr. fragrans Roxb.
Medium-sized tree; leaves elliptic, 5-13 cm long; flowers cream-white, becoming
yellow, fragrant, 2 cm across; berry orange, round, 8 mm across. Usually found
in open places and belukar, also widely planted. A form with a wavy leaf-
margin and reddish brown bark, found in the primary forest, is sometimes con-
sidered as a separate species, namely Fagr. gigantea Ridl. Vern. Tembusu.
Fagr. racemosa Jack ex Wall.
Shrub or small tree, rarely climbing; leaves elliptic, 15-30 cm long, without
distinct auricles, but stipules united around the twig into a short ocrea. In secon-
dary forests, Changi (Goodenough 2783), Kranji.
Fagr. ridleyi K. & G.
Climbing or straggling shrub or small tree; leaves obovate, 15-25 cm long,
secondary veins prominent below. Very rare, Bukit Timah (Ridley 11363),
Sungei Karang.
Norrisia major Solered. (= N. malaccensis Gardn.)
Small tree; leaves oblong to elliptic,6-10 cm long; flowers cream-yellow, in ter-
minal corymbs, branched; capsules opening by parting of 2 valves. Mandai
Road (Corner s.n. in 1937).
112 Gard. Bull. Sing. 36(1) (1983)
Strychnos axillaris Colebr. (= S. malaccensis Benth. & S. pubescens Clarke)
A liana, sometimes a shrub or small tree; twigs pubescent; leaves lanceolate to
suborbicular, 9-12 cm long, 3-5 nerved from the base; flowers less than 0.5 cm
long, corolla-tube nearly as long the limb; fruit ovoid, oblique, 1-1.5 cm across.
Bukit Timah (Ridley 6317), Gardens’ Jungle.
Strychn. ignatii Berg. (= S. ovalifolia Wall, ex G. Don)
Large liana; leaves ovate to elliptic, 8-12 cm long; flowers in axillary cymes,
salver-shaped, 1-1.5 cm long, the corolla-tube longer than the limb; fruit
globose, 5-8 cm across, on thickened branches. The seeds contain strychnine
(Saint Ignatius’ Bean); the roots are used for poison arrows and as fish poison.
Changi, Gardens’ Jungle, MacRitchie (Whitmore 68).
Strychn. ridleyi K. & G.
Climbing shrub; leaves oblong, 8-10 cm long; flowers 0.5 cm long, corolla-tube
longer than the limb. Rare, once collected at Tuas, Ridley 6313 (type).
121. Gentianaceae
Nymphoides indica (L.) O. Kuntze (= Limnanthemum indicum Griseb).
Aquatic plant; leaves broadly ellipsoid or rounded, solitary at the top of stem
or branches, floating; lowermost leaves submerged; flowers in fascicles, at the
top of the petiole-like stem; corolla white, the lobes densely covered with long,
white hairs. In shallow stagnant water at the margin of the Catchment Area,
Haviland & Lim 5741.
122. Apocynaceae
Synoptic key to the genera
I) Herbs: leaves opposite (introduced! and maturalised)ie cre rise cre sere ctersiereieieneeteeneisters Catharanthus
1. Trees, shrubs or woody climbers
2. Erect trees or shrubs
3. Leaves alternate and spiral
4. Native plants; both) wildvandiplanted ses een eis eects Cerbera
4. Introduced: plants) ...cnin26 casas en Oar Ce Thevetia, Plumeria, Adenium
3. Leaves opposite or in whorls
5. Leaves opposite
6:Native plants= 7). 3.0.2. eee eee eae Kibatalia, Kopsia, Tabernaemontana
6= Introducedvandicultivatediplantsmem eee ee Carissa, Tabernaemontana
5. Leaves usually in whorls of 3-8
fie Native plants* smalll toillargeritreesy sere eerie eee eee Alstonia, Dyera
7. Introduced cultivated plants, small trees or shrubs .. Nerium, Ochrosia, Strophanthus
2. Climbers or scramblers,
8. Leaves opposite
9. Native plants Anodendron, Leuconotis, Melodius, Parameria, Strophanthus, Willughbeia
9. Introduced cultivated plants ...... Allamanda, Beaumontia, Chonemorpha, Odontadenia
8. Leaves in whorls of 3-4
10. Native: plants: ocis5.05,.:2tigetye oc sare a io eae ene ee Alyxia
LOS Introducedicultivatediplantss...-5..c cee eee eee eee een Allamanda
Seed plants of Singapore VIII 113
Adenium obesum Balf. (= A. coetaneum Stapf)
A bushy shrub; 1-1.5 m tall; stem and branches thick, succulent; leaves ovate,
5-10 cm long, spirally arranged in tufts. Flowers purple or pink, funnel-shaped,
4-5 cm long, in small clusters. Native to E. Africa. A pot plant.
Allamanda cathartica L.
Climbing shrub; leaves opposite or in whorls of 3-4, oval-shaped, tapering at
both ends, 10-15 cm long. Flowers broad-tubular, 4-5 cm long and across, in
small clusters at branch-tips. A large flowered form, sometimes called A.
hendersoni Rafill., native to Brazil, is often planted in gardens. *x##3% -
Allamanda violacea G. & E.
A climbing shrub also from Brazil. Differs from the above species in having
flowers 5-7 cm across and a reddish-purple corolla. Less commonly planted.
Alstonia angustifolia Wall. ex A. DC.
Small tree, to 10 cm tall; leaves in whorls of 3, oblanceolate, 6-14 cm long. Fruit
of 2 long, slender, woody follicles. Seeds small, numerous, flat, with a tuft of
silky hairs at both ends. In the forest, Gardens’ Jungle, Bukit Timah, Jurong
(Corner s.n. in 1933), Pasir Panjang.
Alst. angustiloba Miq.
Big tree, buttressed; branches in distinct whorls; leaves in whorls of 5-8, oblong-
elliptic, 8-16 cm long. In the forest; Gardens’ Jungle (Ridley s.n in 1890), Sen-
tosa Island. Vern. Pulai.
Alst. scholaris (L.) R. Br.
Small or big tree; leaves in whorls of 5-8, elliptic, 7-17 cm long. Native to the
Malay Peninsula, sometimes planted.
Alst. spathulata BI.
Small to big tree; leaves in whorls of 4-6, obovate, 7-10 cm long. In wet forests
or swamps; Cluny road, Bukit Mandai, Changi (Sinclair SF 40541).
Alyxia reinwardti Bl. var. lucida (Wall.) Markgr. (= A. /ucida Wall.)
Woody twiner; leaves thin-leathery, elliptic or obovate, 4-8 cm long, in whorls
of 3-4. Flowers white, with an orange throat, fragrant. Drupe ellipsoid, pulpy,
black. On the sea-coast; Kranji, Sungei Morai (Ridley 4427 ).
Anodendron candolleanum Wight
Big climber; leaves opposite, leathery, elliptic or ovate, 12-20 cm long. Flowers
cylindric, yellow, very small, in panicles. Follicles dagger-shaped, 15-18 cm
long. In swampy forests. No specimens available.
Beaumontia multifora T. & B.
Woody climber, leaves opposite, oblong, 10-25 cm long, wavy-margined.
Flowers funnel-shaped, 5-7 cm long and 8-10 cm across, white, fragrant,
clustered at the end of branches. Native to SE. Asia; cultivated.
114 Gard. Bull. Sing, 36(1) (1983)
Carissa carandas L.
A much branched shrub; small branches zigzag and with opposite, sharp, bifid
spines to 3 cm long. Leaves opposite, oblong, 3-7 cm long. Flowers in terminal
cymes, solitary, salver-shaped, white, 3-4 cm across. Fruit a black berry. Native
to S. Asia, sometimes grown as a hedge plant or for its edible fruit.
Catharanthus roseus G. Don (= Lochnera rosea Rchb. ex Steud.)
A perennial herb, 30-60 cm tall, ever blooming; leaves opposite, oblong, 3-8 cm
across, rose-purple or white (var. a/bus Sweet) with a slender tube 2-3cm long.
Native to tropical America; cultivated as an ornamental, often wild. K#it ©
Cerbera manghas L.
Small tree; leaves spirally arranged, obovate, 12-30 cm long. Flowers large,
fragrant, white, with a pink eye in the throat of the corolla tube )2.5-4 cm long).
Drupes solitary or in pairs, oblong, slightly compressed, 5-7 cm long. Rare, at
Pasir Panjang and Katong. Vern. Pong Pong. Pinked-eyed Cerbera. i#+¢ 4 »
Cerb. odollam Gaertn.
Like the above species but the corolla-tube is shorter (1.5-2 cm long), the eye
yellow, and the drupe is roundish, 5-10 cm across. Common on sea-shores and
mangroves, and planted along roadsides. Vern. Pong Pong. Yellow-eyed
Cerbera.
Chonemorpha fragrans Alst. (= Ch. macrophylla G. Don)
Large liana with milky sap; leaves opposite, elliptic with a heart-shaped base,
pubescent, 10-15 cm long. Flowers in cymes, bell-shaped, white, fragrant, 3-4
cm long. Native to the Malay Islands, sometimes found in gardens.
Dyera costulata (Miqg.) Hook.f.
Huge tree, to 60 m tall, the bark dark grey, not buttressed; branches in whorls;
leaves in whorls of 6-8, ovate-oblong, 7-18 cm long. Flowers in branched cymes.
Fruit a pair of massive woody follicles, 20-30 cm long, a commercial timber
tree, the latex used to be tapped for chewing gum. Vern. Je/utong.
Kibatalia maingayi (Hook.f.) Woodson (= Vallaris maingayi Hook.f.)
Medium tree, to 20 m tall; leaves decussate, elliptic, 7-10 cm long; flowers
white, in axillary cymes or fascicles. Follicles oblong, slender, to 15 cm long.
Seeds plumed. In the forest, Tanglin, Gardens’ Jungle (Hassan SF 36262).
Kopsia fruticosa (Ker) DC.
Shrub or small tree; leaves oblong-lanceolate, 16-23 x 5-9 cm; flowers pink with
a dark eye, grouped in a much-branched sessile cyme. Native to Burma,
sometimes planted.
Kops. singaporensis Rid.
Shrub or:small tree; leaves oblong-ovate, 14-26 x 7-11 cm; flowers in large ter-
minal dense cymes; corolla white, with a crimson eye. In forests, Chua Chu
Kang, Seletar (Md Noor 17).
Seed plants of Singapore VIII mS
Leuconotis griffithii Hook.f.
Climber; leaves leathery, elliptic-oblong, 8-10 cm long; corolla yellow, 0.75 cm
long; calyx thin, dry. In the forest, Serangoon Road (Ridley 9166), Seletar,
Bukit Mandai, Jurong.
Leuc. maingayi Dyer
Leaves leathery, often in whorls of 3; corolla smaller and shorter (0.5 cm long)
than the above species, calyx fleshy. In forest edge, Bukit Timah, Changi,
Jurong (Ridley 3868).
Meiodinus micranthus Hook.f.
Climber; leaves leathery, oblong-lanceolate, with 10-15 slender parallel nerves;
flowers in axillary cymose panicles; corolla-tube slender, with bifid scales in the
mouth. In the forest, Jalan Bray, Mat 5999.
Nerium indicum Mill.
Erect shrub; leaves leathery, narrowly lanceolate, 10-30 cm long, usually 3 in
a whorl; flowers red, pink or white (often double), fragrant, 4-5.5 cm across.
This is a native to South Asia, easily differentiated from the European Oleander
(N. oleander L). by its scented flowers. #*+#& <
Ochrosia borbonica Gmel. (= Calpicarpum oppositifolium Boireau)
Small tree with stout branches; leaves fleshy, 3-4 in a whorl, obovate, 10-25 cm
long; flowers white, in short cymes; drupe ovoid, oblique. Serimbum, Ridley
s.n. in 1894.
Odontadenia macrantha Markgr. (= O. speciosa Benth.)
A woody climber; leaves opposite, oblong, 10-25 cm long; flowers yellow or
orange, many in terminal or upper axillary cymes; corolla salver-shaped, 6-10
across, the tube 3-4.5 cm long, 5-lobed. Native to tropical America.
Parameria glandulifera Benth.
Large climber; leaves opposite or 3 in a whorl, elliptic to obovate, 8-12 cm long;
flowers white, bell-shaped, fragrant, very small (0.3 cm long), in terminal
corymbose panicles, glabrous; follicles slender, 25-30 cm long. Locality
unstated in Cantley’s collection.
Param. polyneura Hook.f.
Like the above, but flowers pink, and, follicles much longer, 50-60 cm.
Garden’s Jungle, Changi. No specimens available.
Parsonia helicandra Hook. & Arn. (= P. spiralis Wall.)
Slender twiner, glabrous; leaves membranaceous, ovate-oblong, 7-15 cm long;
flowers pinkish yellow, with a pink eye, very small. Follicles cylindric, linear-
lanceolate, 8-15 cm long. Climbing up bushes, usually near the sea. Seletar,
Tuas, Pulau Brani, Tempinis (Rid/ey s.n. in 1891).
Plumeria acuminata Ait.f. (= P. acutifolia Poir.)
A broad-crowned tree; branches thick, succulent; leaves oblong, pointed, 20-40
cm long, alternate and spiral, congested at branch ends; flowers funnel-shaped,
116 Gard. Bull. Sing. 36(1) (1983)
4-5 cm across, waxy, white with a yellow centre. The Frangipani is a native of
tropical America. ##it °
Plum. alba L.
Like the above, but the leaf-tips blunt, and the white flowers much longer (the
spread corolla 8-15 cm across). Other species and hybrids are also planted, one
of them, P. rubra or the Red Frangipani with leaves having a pointed tip and
flowers pink to red, also from tropical America.
Strophanthus brevicaudatus Wight (= ? S. singaporianus Gilg)
Erect shrub; leaves opposite, membranaceous, oblong-elliptic, 5-10 cm long;
flowers small, purple, corolla-lobes not tailed; terminal cymes much branched;
follicles cylindric, to 20 cm long. In damp, open places; Holland Road, Jurong,
Balestier Plain (Ridley 9149), probably extinct.
Stroph. dichotomus DC. (= ? S. caudatus Kurz)
Stout shrub with long, slender trailing shoots; leaves obovate-oblong, 7-12 cm
long; flowers white, corolla-lobes ovate, with long, slender tails, 7-10 cm long,
purple; the follicles cylindric, twinned, stout, divaricate, each 20 cm long and
5 cm across. In open places and edge of forests; Changi (Ridley s.n. in 1890)
Seletar. Vern. Sembrong, Seram.
Tabernaemontana corymbosa Roxb. (= Ervatamia corymbosa K. & G.)
Shrub or small tree; leaves leathery, elliptic, 15-40 cm long; petiole-base enlarg-
ed, clasping stem; corymbs in upper axils, much branched; follicles often twinn-
ed, ovoid, curved, beaked, 3.5-4 cm long. Mandai Road, Corner 30668.
Tabern. divaricata Roxb. (= T. coronaria Willd., Ervatamia divaricata Burke)
Cultivated shrub, less than 60 cm tall; corolla white, often in double form;
flowers 2-3 in a cyme. Probably native to N. India. 4 it °
Tabern. malaccensis Hook.f. (= Ervatamia malaccensis K. & G.)
Slender shrub; leaves membranaceous, lanceolate, 4-10 cm long; cymes terminal
and axillary; corolla white with a yellow throat; follicles oblong, falcate, beak-
ed, orange, 2.5-3 cm long. Gardens’ Jungle (Hullet 521), Changi.
Thevetia peruviana K. Schum.(= T. neriifolia Juss.)
Much branched shrub; leaves leathery, alternate and spirally arranged, narrow-
ly lanceolate, 8-14 cm long; flowers, many together in cymes, 1-2 blossoming
at a time; corolla bright yellow, 6-7 cm long; drupe compressed-globose, 4-5.5
cm across. Native of Central America. Yellow oleander. 3% ii R41
Urceola brachysepala Hook.f.
Large climber; leaves papery, elliptic, 10-14 cm long; panicles terminal and ax-
illary; flowers white, small, corolla globose, short; follicles cylindric, slender,
15-18 cm long. Jurong, Bukit Timah (Ridley 8397), Bukit Mandai.
Ure. lucida Benth.
Stout liana; leaves papery, elliptic, 7-12 cm long; follicles dagger-shaped below,
recurved. Jurong, Changi, P. Ubin (Ridley s.n. in 1894).
Seed plants of Singapore VIII LY)
Ure. maingayi Hook.f.
Large climber; leaves leathery, elliptic, 6-8 cm long; follicles terete, 15 cm long,
the tip hooked. In the forest, Kranji, Goodenough 2717.
Ure. malaccensis Hook.f.
Climbing shrub; leaves leathery, ovate, 5-12 cm long; follicles terete, recurved,
20 cm long. Gardens’ Jungle, Changi (Hullet 398), Bukit Timah, Bedok.
Ure. torulosa Hook.f.
Climber; leaves leathery, oblong-lanceolate, 10-17 cm long; follicles very
slender, constricted at intervals. Gardens’ Jungle, Tempinis (Ridley 3602a),
Seletar, Nee Soon.
Willughbeia apiculata Miq. (= W. flavescens Dyer)
Differs from W. coriacea in the flowers being in slender-peduncled, lax cymes
and in the smaller, spherical fruit (2-5 cm across). Chua Chu Kang (Ridley
6143), Changi.
Will. coriacea Wall. (= W. firma BI.)
Large woody climber with tendrils (modified peduncles); leaves leathery, op-
posite, distichous, oblong-elliptic, 8-14 cm long; cymes sessile, axillary; flowers
white; berry pear-shaped, orange to dull red, to 10 cm across. In the forest,
common; Bukit Timah, Gardens’ Jungle, P. Ubin, Tanjong Gul (Ridley 3594a).
Will. tenuiflora Dyer
Like W. coriacea, but leaves thin, leathery, oblong, 10-12 cm long; fruit larger,
to 14 cm across. In the forest; Bukit Timah, Catchment Reserve forest (Corner
S.n. in 1937).
Wrightia religiosa (T. & B.) Hook.f.
Shrub, less than 1 m tall, with many short and slender branchlets; leaves op-
posite, oblong-lanceolate, 2-5 cm long; flowers in terminal cymes; corolla white,
salver-shaped, less than 0.5 cm long and 2 cm across. native to Thailand, a
popular bush planted for training as bonsai or miniature plant. #4 ©
123. Asclepiadaceae
Synoptic key to the genera
1. Erect herbs, shrubs or small trees
At. TSIRETID GES 65 6 ak ale SO OEE Oe OES DOR GEO oe ae Teac aoe ee Stapelia, Asclepias
PPE SHEM ES SHOMESIN ALLELE COSI Lire tere ie eke late ver vate cenek ss cs yas ers) 67 6) Sle 0 6S eysieustes oy Steifore, che euers.cusysrepesdjave eva evs Calotropis
1. Climbers
2. Corolla-tube very short or almost none, or corolla wheel-shaped .... Cynanchum, Finlaysonia,
Genianthus, Hoya, Sarcolobus, Tylophora
2. Corolla with a distinct tube
2) COPGIE TES ees apo. on ed adoo CeO GaES Oe SEO DE CONG OA TAO MRS NOOSE ORS Dischidia
Sy (Cla TEMS IEG Te oes lenge Gta Oaes OO Tere enn be teaches o Reece me Physostelma, Telosma
3. Corolla salver-shaped or funnel-shaped .............. Ceropegia, Stephanotis, Toxocarpus
118 Gard. Bull. Sing. 36(1) (1983)
Asclepias curassavica L.
Perennial herb, 50-60 cm tall; leaves opposite, narrowly oblong, both ends
pointed, 5-12 cm long; flowers umbellate; corolla red; corona orange, pro-
truding; fruit a pair of follicles, erect; seeds with a tuft of silky hairs. The
milkweed is a native to tropical America, occasionally planted as an
ornamental. #44: it «
Calotropis gigantea (Willd.) Dryand ex Ait.f.
Bushy shrub or sometimes a small tree, to 5 m tall; young parts woolly-greyish;
leaves elliptic, 10-35 cm long, fleshy; flowers in umbellate clusters; corolla white
to pale lilac, 2.5-3.5 cm across; corona prominent. Native to India and the
Malay Islands.
Ceropegia woodii Schlecht.
Slender tuberous vine; leaves heart-shaped, mottled with dark green; flowers 2-3
together, pinkish; corolla tube 1.5-2 cm long, with swollen base. Native to S.
Africa, occasionally planted in hanging baskets.
Cynanchium ovalifolium Wight
A slender twiner; leaves thin, oval or oblong, 6-10 (-15) cm long, tip pointed;
flowers 5-15 in a bundle; corolla 5-6 mm across, yellowish, tipped red; follicles
narrow oval, 8-10 cm long, pointed. Cluny Road, Balestier Plain, Bukit Timah
Road, Holland Road (Ridley 11456).
Dischidia albiflora Griff. (= D. collyris Wall.)
Slender creeper, epiphytic; leaves rounded or kidney-shaped, buff-yellow
above, purple beneath, lying flat on the trees; flowers white. Ants often nesting
between the leaves of the epiphyte and the branches of the host tree. Tanglin,
Chua Chu Kang (Ridley 2928), Bukit Timah, Kranji.
Disch. bengalensis Colebr.
Large creeping epiphyte; stem glaucous green, long and nearly leafless, or with
a few narrow oblong leaves, 2-3 cm long; flowers creamy white. In the forest
near the sea; Bajau, Kranji, Ridley 2729.
Disch. cochleata Bl. (= D. coccinea Griff.)
Stem slender; leaves fleshy, rounded (1.5-2 cm across), crowded together, the
edges pressed close to the bark of the host tree, harbouring ants and roots;
flowers scarlet, tipped blue. Usually on lofty trees; Bukit Timah, (Ridley s.n.
in 1896), Seletar, Chua Chu Kang.
Disch. hirsuta Decne
Stem slender; leaves small (1-2.5 cm long), purple green, hairy; flowers red. In
the forest; Seletar, Chua Chu Kang (Goodenough s.n. in 1889), Kranji.
Disch. major Merr. (= D. rafflesiana Wall.)
Epiphytic twiner; leaves opposite, of two forms: the smaller ones rounded (2-2.5
cm across), the pitcher leaves oblong, pouch-like (7-12 cm long), outside
yellowish green, inside purple, with holes at the base for ants and aerial roots.
Flowers golden yellow, in small umbels. Common on trees all over the Island
especially near the sea. Berlayer (/.H. Burkill 239).
Seed plants of Singapore VIII 119
Disch. nummularia R. Br. (= D. gaudichaudii Decne)
Creeping epiphyte; stem very slender; leaves small (5-6 mm long), thick and
fleshy; flowers pale yellow or white, 2-3 in leaf-axils. In open places often in
masses closely covering the tree trunk and limbs and causing damages to the
host trees, throughout the Island. Tyersall Road Ridley 5746.
Disch. singaporensis Rid.
Leaves ovate, flat, 1.5-2.5 cm long. Collected only once at Changi by Ridley
(Ridley s.n. in 1908, type).
Finlaysonia obovata Wall. (= F. maritima Backer)
Woody climber; leaves fleshy, green, broadly obovate, 5-10 cm long; flowers in
branched axillary cymes; corolla 1 cm across, yellowish green, purplish in the
centre; follicles pear-shaped, oblique, 3-4 ribbed, 8-10 cm long. In mangroves
and on tidal river banks, Kranji, Geylang (Ridley s.n. in 1893).
Genianthus maingayi Hook.f.
Slender climber, with a purply pubescence; leaves leathery, oblancecolate to
obovate, 6-9 cm long; flowers reddish white, very small (2-3 mm long), in long
spikes. Bukit Timah, Cantley s.n. (not seen).
Hoya coriacea Bl.
Epiphytic climber; leaves thin, ovate-lanceolate, 12-15 cm long; flowers
yellowish white, tipped purplish. Collected once at Tempinis (D’A/meida s.n.
in 1893).
Hoya coronaria BI.
Densely hirsute; stem stout; leaves coriaceous, elliptic, 8-10 cm long. Flowers
waxy, 2-3.5 cm across, the largest of all the species; when opened, white, with
a yellow tint, then becoming pink. Common near the sea, Serangoon, Changi,
Kranji, P. Tekong, Ulu Berih U.H. Burkill 4336).
Hoya diversifolia Bl.
Leaves fleshy, elliptic-obovate with narrowed base, 10-13 cm long; umbels 1-20
flowers; flowers pink. Often densely covering trees; Jurong, Chua Chu Kang,
Kranji, Serangoon (Ridley 2733).
Hoya finlaysonii Wight
Leaves coriaceous, ovate-oblong. Collected once from Singapore (Wallich
2724).
Hoya lacunosa BI.
Creeping epiphyte; leaves thick-fleshy, lanceolate to ovate, 2-4 cm long; corolla
white. On trees; Tanglin, Seletar, Bukit Mandai, Chua Chu Kang, Bukit Timah
(Ridley s.n. in 1892).
Hoya latifolia G. Don
Stem deep red young; leaves fleshy, ovate with a heart-shaped base, up to 25
cm long; flowers small, pink. On trees in dense jungles; Kranji, Seletar, Pong-
gol, Changi, P. Tekong, Changi U/.H. Burkill 10013).
120 Gard. Bull. Sing. 36(1) (1983)
Hoya obtusifolia Wight
Stout climber; leaves oblong thick, 12-15 cm long; umbels 1-20 flowers; flowers
yellow, with a pinkish centre. Serangoon (Ridley 8932), Changi, Bukit Timah.
Hoya parasitica Wall (= H. ridleyi K. & G.)
Leaves fleshy, elliptic with cuneate bases, 8-10 cm long; umbels 1-40 flowered;
flowers pink or white. Often near the sea, Changi, Chua Chu Kang, Tuas, Kran-
ji, Pulau Tekong (Ridley 1796).
Physostelma wallichii Wight
Slender climber; leaves oblong to elliptic, abruptly pointed, 5-12 cm long;
flowers large, in axillary bundles; corolla bell-shaped, 2-2.5 cm across, thin,
cream-white, with a purple centre; fruits long and narrowly cylindric, 15-20 cm
long. Tempinis River (Ridley s.n. in 1894), Kranji, Tuas.
Sarcolobus globosus Wall.
Small twiner; leaves fleshy, oblong, pointed, 7-10 cm long; flowers small in
umbels, light purple with brown streaks; follicles nearly globose, ridged on one
side, greyish brown; seeds not plumed. On edges of mangroves and river banks.
Changi, Seletar, Serangoon (Ridley 11640). Pulau Ayer Chawan.
Stapelia nobilis N.E. Br.
Small leafless herb; stem cactus-like, succulent, bluntly 4-angled, 5-20 cm long,
with soft spines; flowers star-shaped, dark purple, 3-5 cm across, hairy. Native
to Africa, sometimes planted.
Stephanotis floribunda Brongn.
Climber; leaves oval, leathery, shiny, 5-10 cm long; flowers in axillary clusters,
tubular, waxy, white, 2.5-5 cm long, fragrant. Native to Malagasy
(Madagascar).
Steph. maingayi Hook.f.
Climber; flowers white, bigger than the above species. Once collected at Changi
(Aullett 147).
Telosma cordatum Merr. (= Pergularia minor Andr.)
Slender vine; branches downy; leaves cordate, 4-11 cm long, pubescent below;
flowers yellowish green, bell-shaped, about 1 cm long, in axillary clusters. Pro-
bably a native to E. Asia, occasionally cultivated. 4°
Toxocarpus griffithii Decne
Slender twiner; leaves papery, elliptic-oblong, 5-10 cm long; flowers in small
clusters; corolla-lobes pinkish-crimson outside, yellow inside; follicles cylindric,
25-30 cm long. Jurong, Corner s.n. in 1933.
Tylophora asthmatica W. & A.
Slender twiner; leaves oblong or ovate-lanceolate, 4-10 cm long; flowers small
in compound umbels, yellow or pink; follicles narrowly lanceolate, 8-10 cm
long. In sandy places by the river or sea; Changi, Ridley s.n. in 1891.
Seed plants of Singapore VIII 121
Tyl. squarrosa Ridl.
Slender twiner; leaves ovate, 5-8 cm long; flowers in racemes. Tanah Merah
(Ridley 2737). Probably synonymous with T. wallichii.
Tyl. tenuis Bl.
Slender climber; leaves small, lanceolate, tip pointed, 2-6 cm long; flowers
small, purple, in axillary, spreading inflorescence. In open places or on banks
of tidal rivers; Alexandra Road. (Ridley 5746), Sentosa, Tanjong Rhu.
124. Oleaceae
Key to the genera
Ie Scranipline scandent Or Climbing ‘SHTUDS 2.1.2.6 6cra 0:00 1s. + oy. cres ayes wie e oinyeresiasea vein ee as Jasminum
1. Erect shrubs or trees
2. Flowers 4-merous
Seebreesscorolla-lobes nearly free tonthe base )...e5 50) lal ee cleave eee bes Linociera
Ba SHENDS-ACOrolla-lobes contates below . cas. <tc iesie oie, > « isis sis so isle SSMS s Sees chee ore Olea
PERO WEES?S-INCEOUS:) CUEIVALCE SHTUDS) e:0../:-0 oe cccc cae dae cece sevens eee esate eens Nyctanthes
Jasminum bifarium Wall.
Scrambling or climbing shrub; leaves simple, opposite, ovate-lanceolate, 4-8 cm
long; flowers 3-5 in axillary clusters higher up; calyx 5-7 lobed; corolla-tube
1.5-2.5 cm long, 5-7 lobed, the pointed; drupe black. In open country and
hedges; Tanglin, Balestier Plain, Mandai Road (Sinclair SF 40007), Changi,
Vern. Melor Hutan.
Jasm. griffithii Clarke
Scandent or climbing shrub; hairy; leaves ovate or elliptic, 10-15 cm long; cymes
densely hairy; corolla-tube 1.5-2 cm long, drupe white. In forests, Gardens’
Jungle, Chua Chu Kang (Goodenough s.n. in 1893).
Jasm. longipetalum K. & G.
Slender climber; leaves elliptic to lanceolate, 5-15 cm long, 3-nerved from the
base; flowers in short cymes; corolla-tube 2 cm long, 9-lobed, the lobes linear-
oblong, acute, 2-2.5 cm long; fruit glabrous, black. In forests, Mandai, Ridley
10937.
Jasm. rex Dunn
Climbing shrub, glabrous; leaves simple, 3-5 nerved at the base; flowers white,
often tinted violet, odourless; corolla-tube 2.5-3 cm long, 5-11 lobes, the lobes
oblong-obovate. The Royal Jasmine is native to Thailand, often cultivated.
Jasm. sambac Ait.
Scandent or climbing shrub; leaves oval or rounded, 2.5-5 cm long; flowers
white, often violet outside, fragrant; corolla-tube 0.7-1.5 cm long, 8- or more
lobed, the lobes oblong to oval, blunt, or often the corolla double. Native to
India, often planted in gardens. #€# #
122 Gard. Bull. Sing. 36(1) (1983)
Linociera ramiflora (Roxb.) Wall (= L. pauciflora Clarke)
Small tree; leaves coriaceous, elliptic oblong, 10-20 cm long; flowers in panicles;
petals oblong. In wet forests, Jurong, Bukit Timah, Bukit Mandai
(Goodenough 5079), Seletar.
Nyctanthes arbor-tristis L.
Shrub, much-branched, the branches 4-angled, hairy; leaves ovate, 4-11 cm
long, flowers 3-7 in an axillary head; corolla-tube orange, 1-2 cm long, the lobes
white; capsule dark brown, 2-seeded. Night blooming, strongly fragrant; native
to India, one of the sacred plants of that country; occasionally cultivated. (The
genus Nyctanthes is sometimes classified under Verbenaceae). Vern. Seri
Gading.
Olea brachiata Merr. (= O. maritima Wall.)
Much-branched shrub; young parts pubescent; leaves leathery, elliptic or ovate-
lanceolate, 7-10 cm long; flowers in cymose panicles; corolla globose or cam-
panulate, yellow, very small; drupe pear-shaped, 7-8 mm long, black. In open
places near the sea, Changi (Sinclair SF 40017), P. Ubin.
ADDENDA, CORRIGENDA ET EMENDANDA
A) MALPIGHIACEAE
Through an oversight the family Malpighiaceae was left out in the previous treat-
ment. The following is to be read after 85. Erythroxylaceae and before 86. Ox-
alidaceae on page 341, in Gdns’ Bull. Sing. 33(1980).
85 A. Malpighiaceae
Key to the genera
1. Erect shrubs; fruit smooth, neither winged nor horned
2. Leaves membranous; entire; with’ 2)elanasrat tie) bDaScu a eecirerersielieiieireierenarerarate Galphimia
2; eaves leathery,-usually with’spiny teeth. elandless: s-eisa-jpieiecrsiersie sie crerererserereierent Malpighia
1. Scandent or twining shrubs; fruits winged or horned
3. Petals yellow
4. Flowers)in) racemes; petals) = equals, Keeledis sermon ieee Tristellateia
4. Flowers in umbellate clusters; petals unequal, not keeled .................. Stigmaphyllon
3. Petals white or pink
5. The central mericarp surrounded by a large oblong wing, and the 2 lateral by smaller and narrow
WINGS ohh a oar ctate e eeatlatecera dl a. 24re Koel eeelere tenets RRL eM estas te cntec ay tte is tee lererees cece oe eee Hiptage
5. Mericarps surrounded by broad, oblong to orbicular wings ................. Aspidoptreys
Aspidoptreys concava (Wall.) Juss.
Liana to: 20 m tall; young parts dark-red hairy; leaves ovate-elliptic, opposite;
panicles axillary, flowers white, ovary glabrous, with 3 winged sides; samara of
3 membranous, ovate to oribular, winged mericarps. In forests, Bukit Mandai,
Seletar, Chua Chu Kang, Bukit Timah (Ridley 3583a), Nee Soon.
Seed plants of Singapore VIII 123
Galphimia glauca Bartl.
Shrub, to 2 m high; young parts reddish hairy, older twigs brown, glabrous;
leaves oblong, 2.5-5 cm long, with 2 small glands at the base of the blade;
flowers yellow, in terminal racemes. Native to Central America, cultivated in
gardens.
Hiptage sericea Hook.f.
Climber; young twigs densely brown hairy; leaves oblong, 4-6 cm long; flowers
pink and white, in axillary racemes; samara 3-winged, the central wing 3-4 cm
long, and 2 lateral wings 1.5-2 cm long. In open places, Changi (Ridley 3989),
Bukit Mandai.
Malpighia coccigera L.
Small shrubs, 1-2 m high; leaves leathery, elliptic, 1-2 cm long, entire or with
a few spiny teeth; flowers axillary, single or in pairs; petals pinkish, fringed,
abruptly narrowed at the base. From the West Indies, planted as hedges or in
pots as a miniature plant or bonsai.
Stigmaphyllon ciliatum (Lam.) Juss.
A slender woody vine; leaves heart-shaped, 2.5-6 cm long, with hairy edge;
flowers 3-7 in a cluster; petals yellow, wavy. Native from the West Indies to
Brazil, occasionally planted.
Tristellateia australasiae A. Rich.
Liana to 10 m tall; twigs purple, lenticellate; leaves oblong-ovate, 7-12 cm long;
petioles with 2 small stipules at the base; racemes terminal; flowers bright
yellow; samara of 3 mericarps, the lateral wings of the mericarp 4-10 lobed or
horned, expanding in one plane, star-shaped. In mangroves, tidal swamps or in
forests; often cultivated in gardens. Jurong, Ridley s.n. in 1894.
B) VITACEAE (Gdns’ Bull. Sing. 35: 86-88, 1982)
Dr. Abdul Latiff Mohamed of Universiti Kebangsaan Malaysia, in a letter dated
19th January 1983, kindly informed that the following 8 species should be included
in this family.
1. Pterisanthes cissoides Bl.
Nee Soon, Enoch s.n. in 1955.
2. Pter. eriopoda (Miq.) Planch.
Chua Chu Kang, Ridley 425.
3. Ampelocissus ascendiflora Latiff
Catchment Reserves, Bukit Timah (Ridley 5846).
4. Ampel. polystachya (Wall.) Planch.
Bukit Timah, Sinclair SF 39649.
5. Ampel. thrysiflora (Bl.) Planch.
Changi, Ridley s.n. in 1893.
124
Gard. Bull. Sing. 36(1) (1983)
Tetrastigma lanceolarium (Roxb.) Planch.
Bukit Timah, Ridley s.n. in 1894.
Tetr. curtisii (Ridl.) Suesseng.
Bukit Timah, Ridley s.n. in 1894.
Cissus quadrangularis L. (alt. names: Vitis quadrangularis Wall.)
Native to Bengal, India; sometimes cultivated as a hedge plant, also used
in local herbal medicine
Dr. Latiff also mentioned that he has elevated Vitis macrostachya Miq. or
Ampelocissus spicifera Planch. to a generic status, namely Nothocissus, recent-
ly published in Mus. J. (N.S.), 27, 1983.
The Genus Trisetum (Gramineae) In Malesia And Taiwan
J. F. VELDKAMP & J. C. van der HAVE
Rijksherbarium, Leiden, the Netherlands
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983
Summary
In Malesia and Taiwan there are three taxa of Trisetum Pers. (Gramineae): T. bifidum (Thunb.) Ohwi
in Taiwan and New Guinea, 7. spicatum (Linné) Richt. ssp. kinabaluense Chrtek in Sabah and the ssp.
formosanum (Honda) Veldk., comb. nov., in Taiwan. Trisetum flavescens (Linné) Beauv. must be the
conserved type of the generic name. Some subspecific epithets proposed by Hultén (1959) under T.
spicatum are validated.
Introduction
The genus Trisetum Pers. belongs to the Aveneae and is characterised by the lem-
ma, which is awned in the distal half and has a bifid to bisetose apex. It is very close
to Koeleria Pers., in which the lemma has a (sub-) apical awn and an entire apex. Some
species are intermediary in these characters, as could be expected, but since they are
annuals, the life-cycle has been used to place them conveniently in separate satellite
genera. Thus we have Parvotrisetum Chrtek and Trisetaria Forssk. next to Trisetum
and what is generally called Lophochloa Reichenb. next to Koeleria. Life form
seems a very weak character, especially when it is the only one to base genera on,
and it is then not surprising that some authors, e.g. Paunero (1950), Jacques-Félix
(1962) have united Trisetum with Trisetaria, the latter name being the oldest. Jonsell
(1978, 1980) has kept them separate, but retained some annuals in Trisetum. Unfor-
tunately he has not yet explained the exact reasons for doing so. As we intended tp
study only the Malesian and Taiwanese taxa, we did not want to get involved in the
problems of generic delimitation and have here used the more usual generic name
Trisetum.
There is yet another problem about the application of the name T7risetum,
however. The generally accepted lectotype of this is 7. pratense Pers., a superfluous
name for T. flavescens (Linné) Beauv. (cf. Hitchcock, 1920; Kerguélen, 1975); the
Index Nominum Genericorum erroneously followed Britton & Brown’s (1913) ar-
bitrary typification, which must be rejected. They appointed 7. striatum (Lamk.)
Pers., but that is a taxonomic synonym of Helictotrichon sempervirens (Vill.) Pilg.,
the lectotype of Helictotrichon Besser! (See Sevenster & Veldkamp, 1983). It has not
been realised, however, that as early as 1827, Besser (between July and December)
and Link (in October or November) restricted Trisetum to the annual species, and
by retaining 7. parviflorum (Desf.) Pers. as the only species in it, Link made that
the obligatory type! Jonsell has kept that species in Trisetum but Holub (1974) has
included it in Rostraria Trin., which he claimed is the correct name for Lophochioa,
but which then rightfully would be the ‘true’ Trisetum. What everyone else has call-
ed Trisetum must then be named Acrospelion under which name Besser placed the
perennial species including 7. flavescens. This is of course most undesirable and
125
126 Gard. Bull. Sing. 36(1) (1983)
therefore Veldkamp (1983) has proposed to make T. flavescens the conserved type
of both Trisetum and Acrospelion, whereby the latter name becomes superfluous.
Various attempts have been made to distinguish infra-generic entities in Trisetum.
For America, Louis-Marie (1928) has proposed a system which was based on mor-
phological characters; for Europe, Chrtek (1965) made another using the anatomy
of the leaves and roots. The two systems are incompatible: Louis-Marie’s subgenus
Heterolytrum (= Trisetum) includes all of Chrtek’s subgenera, while sections of the
latter were not recognised by Louis-Marie at all.
The species under study here belong to the subgenus Trisetum, with T. bifidum
in the sect. Sibirica Chrtek (1968), reduced to a subsection of sect. Trisetum by Pro-
batova (1978), and 7. spicatum in sect. Trisetaera Aschers. & Graebn.
Ridley (1916) described 7. /atifolium from New Guinea and mentioned the
presence of 7. spicatum [as T. subspicatum (Linné) Beauv.] there. The type of the
first name belongs to Arundinella furva Chase, the material of the second to An-
thoxanthum angustum (Hitche.) Ohwi.
Some 18th century collections labeled ‘Timor’ represent T. antarcticum (Forst. f.)
Trin. This locality is no doubt erroneous, for this species is otherwise only known
from New Zealand and some Antarctic islands. If it really occurred in Timor, it
could only have been found in the mountains, but the collectors at that time had
not yet penetrated so far inland. This species is certainly not the same as T. spicatum
spp. australiense Hultén, although the name is cited in the latter’s synonymy by
Hultén (1959). It is a ‘good’ Trisetum and certainly not a Hierochloé as stated by
the International Code of Botanical Nomenclature (1975)! Here its basionym, Aira
antarctica, is cited as the basionym of Disarrenum antarcticum Labill., the type of
Disarrenum Labill. a name rejected against Hierochloé. De Labillardiére in fact did
mention Forster’s name as possibly belonging to what he described and depicted,
but his doubts were soon validated when Forster’s material turned out to belong to
Trisetum and his own, to what now is known as Anthoxanthum redolens (Vahl)
Royen. The reference to Forster must therefore be deleted from his protologue and
the Code emended. If it is maintained that Aira antarctica must nevertheless be
regarded as the basionym, the whole entry of Disarrenum must be deleted, as that
name then be regarded as a later, heterotypic synonym of Trisetum!
Acknowledgements
This study was initiated by the second author during a course in advanced
Angiosperm taxonomy at the Rijksherbarium and finished by the first author. The
descriptions are mainly based on material present in L, specimens of the 7.
spicatum-complex seen by Hultén were kindly sent on loan from S by Dr. K.
Bremer, making possible the validation of some of Hultén’s combinations, while
Veldkamp was able to annotate specimens in BO and SING during a visit to these
Institutes. The latter also made the drawings, which were carefully inked by Mr. J.
van Os, Leiden.
Trisetum in Malesia & Taiwan Wil
Literature
Besser, W.S. J. T. von (1827). In: J. A. Schultes & J. H. Schultes, Mantissa 3: 526
(‘326’). Stuttgart.
Britton, N. L. & A. Brown. (1913). An illustrated flora of the northern United
States, Canada and the British possessions, ed. 2, 1: 216. New York.
Chrtek, J. (1965). Bemerkungen zur Gliederung der Gattung Trisetum Pers. Bot.
Not. 118: 210-224.
(1968). Bemerkungen zu einigen Arten der Gattung Trisetum aus dem
Sikkim-Gebiet. Acta Univ. Carol., Biol. 1967: 103-107.
Hitchcock, A. Si: (1920). The genera of grasses of the United States. U. S. Dept.
Agric. Bull. 772: 107-109.
Holub, J. (1974). New names in Phanerogamae 3. Folia Geobot. & Phytotax. 9:
271.
Hultén, E. (1959). The Trisetum spicatum complex. Svensk Bot. Tidskr. 53:
203-228.
Jacques-Félix, H. (1962). Les Graminées d’Afrique tropicale 1: 187. Paris.
Jonsell, B, (1978). Nomenclatural and Taxonomic Notes on Trisetum Pers. and
Lophochloa Reichenb. (Gramineae). J. Linn. Soc. Bot. 76: 320-322.
(1980). Lophochloa, Trisetum, Trisetaria, in: T. G. Tutin et al/., Fl. Eur.
5: 121-123, 220-224. Cambridge.
Kerguélen, M. (1975). Les Gramineae (Poaceae) de la Flore Frangaise. Essai de mise
au point taxonomique et nomenclaturale. Lejeunia n.s. 75: 275.
de Labillardiere, J. J. H. (1806). Novae Hollandiae Plantarum Specimen 2: 82.
Paris.
Link, H. F. (1827). Hortus Regius botanicus Berolinensis 1: 120. Berlin.
Louis-Marie. (1928). The Genus Trisetum in America. Rhodora 30: 209-223,
237-245.
Paunero, E. (1950). Las Especies espafiolas del Género Trisetaria Forsk. Anal. Jard.
Bot. Madrid 9: 503-582.
Probatova, N. S. (1978). De Generum Trisetum Pers. et Koeleria Pers. Speciebus
Caucasicis Notulae Systematicae. Nov. Syst. Pl. Vasc. 15: 17-22.
Ridley, H. N. (1916). Report on the Botany of the Wollaston Expedition fo Dutch
New Guinea, 1912-13. Trans. Linn. Soc., London (Bot.) 9, 1: 250.
Sevenster, J. G. & J. F. Veldkamp. (1983). A Revision of Helictotrichon
(Gramineae) in Malesia. Blumea 28: 329-342.
Veldkamp, J. F. (1983). Proposal to Conserve Trisetum (Gramineae) and its Type
Species. Taxon 32: 487-488.
128 Gard. Bull. Sing. 36(1) (1983)
Trisetum
Trisetum Pers., Syn. 1 (1805) 97; Louis-Marie, Rhodora 30 (1928) 209, 237; C. E.
Hubb., Fl. Trop. E. Afr. 10 (1937) 99; Hitchc. & Chase, Man. Grasses U. S.,
ed. 2 (1951) 287; Bor, Grasses (1960) 447; Ohwi, Fl. Jap. (1965) 148; Chrtek, Bot.
Not. 118 (1965) 210; Hsu, Fl. Taiw. 5 (1978) 422; Nicora, Fl. Patag. 3 (1978) 238;
Jonsell, Fl. Eur. 5 (1980) 220. — Acrospelion Bess. in Schult. & Schult. f., Mant.
3 (1827) 526 (‘326’), nom. superfl. — Lectotype: Avena flavescens Linné =
Trisetum pratense Pers., nom. .superfl., = Trisetum flavescens (Linné) Beauv.,
typ. cons. prop. (see introduction).
Graphephorum Desv., Nouv. Bull. Soc. Philom. Paris 2 (1810) 189. — Lectotype:
Graphephorum melicoideum (Michx.) Desv. [= Trisetum melicoides (Michx.)
Scribn.].
Trichaeta Beauv., Agrost. (1812) 86, 179, t. 17, f. 8. — Type: Trichaeta ovata (Pers)
Beauv. [= Trisetum ovatum Pers.; the basionym of this is Bromus ovatus Cav.,
1801, non Gaertn., 1770, so Persoon is the validating author].
Rupestrina Provancher, Fl. Canad. (1862) 689. — Type: Rupestrina pubescens Pro-
vancher, non. superfl. [= Trisetum spicatum (Linné) Richt.].
Perennials. Innovations folded or involute. Ligule membranous. Panicle lax to
densely contracted. Spikelets laterally compressed, (1-) several-flowered, disar-
ticulating above the glumes and between the lemmas (rarely below the glumes in
some N. Am. spp.), all florets bisexual except for the variously reduced uppermost
one(s). Glumes in situ shorter to longer than the lemmas, of + the same consistency
or thinner, unawned, keeled; lower glume usually 1-nerved; upper glume usually
3-nerved, equal to or longer and wider than the lower one. Rachilla prolonged,
nodes puberulous to pubescent. Lemmas usually awned above the middle, apex
usually bifid and bi-aristate, 3-5-nerved; callus usually shortly hairy. Palea
2-nerved, not enclosed by the lemma in fruit. Lodicules 2, membranous, bilobed,
margin puberulous to glabrous. Anthers 3. Ovary apically glabrous or with only a
few hairs; styles free at base, protruding laterally. Caryopsis fusiform, + terete, not
furrowed; hilum subbasal, punctiform to elliptic, inconspicuous; embryo small; en-
dosperm + liquid.
Distribution. Temperate regions, in the tropics in the high mountains. Depending
on the classification, 50-75 species, 2 in Malesia and Taiwan.
Key
1. Peduncle densely pubescent below the contracted panicle. Axis densely pubescent. Lower glume
0.6-1 times as long as the first lemma, Apical teeth of the first lemma 0.4-0.7 mm long ... 2
1. Peduncle and axis glabrous, panicle lax, nodding. Lower glume 0.5-0.6 times as long as the first
lemma. Apical teeth of the first lemma 1.5-2 mm long.—Lower glume 2-5 mm long. Upper
glume 4.5-6.5 mm long. First lemma 5-7.5 mm long. Anthers 0.9-1.3 mm long. Taiwan, New
Guinea: 2,25. cee Sie sis BORIS coe haa ree eee ee Bee 1. T. bifidum
2. Lower glume 3.5-5 mm long, 0.6-0.8 times as long as the first lemma. Upper glume 5-7 mm
long. First lemma 4.5-6 mm long, apical teeth + 0.4 mm long. Taiwan ..................
siete: ei6ye) ayerleves soueive Ings] eceeneue cei ehel one Lene isocvaKe ORs rece PekereR rR Kote RRC 2a. T. spicatum spp. formosanum
2. Lower glume 6-7 mm long, + as long as the first lemma. Upper glume + 8 mm long. First lem-
ma 6-6.5 mm long, apical teeth + 0.7 mm long. Sabah.. 2b. T. spicatum ssp. kinabaluense
Trisetum in Malesia & Taiwan 129
1. Trisetum bifidum (Thunb.) Ohwi
Trisetum bifidum (Thunb.) Ohwi, Bot. Mag. Tokyo 45 (1931) 191; Acta Phytotax.
& Geobot. 3 (1934) 81; J. Jap. Bot. 17 (1941) 444; Fl. Jap. (1965) 149; Chung,
Korean Gr. (1965) 89; Lee, Man. Korean Gr. (1966) 121, f. 90; Hsu, Taiwania
16 (1971) 235; Taiwania 17 (1972) 54, f. 62; Anon., Icon. Corm. Sin. 5 (1974)
91, 845, f. 7012; Hsu, Taiwan Gr. (1975) 285, fig.; Tateoka, Bull. Nat. Sc. Mus.
Tokyo B 4 (1978) 1; Hsu, Fl. Taiwan 5 (1978) 422, f. 1384. — Bromus bifidus
Thunb., Fl. Jap. (1784) 53. — Trisetum flavescens (Linné) Beauv. var. bifidum
Makino, Bot. Mag. Tokyo 26 (1912) 215. — Type: Hb. Thunberg 2576 (UPS,
holo, n.v., IDC 1036 seen; L), Japan, Nagasaki, 1774.
Bromus avenaeformis Steud., Syn. 1 (1854) 326. — Lectotype: Burger s.n. (L, holo,
908.98-239, here proposed), Japan.
Trisetum flavescens (Linné) Beauv. var. papillosum Hack., Bull. Hb. Boiss. 7 (1899)
702; Honda, J. Fac. Sc. Univ. Tokyo III, 3 (1930) 126; Chase, J. Arn. Arb. 24
(1943) 83; Henty, Bot. Bull., Lae 1 (1969) 189; Royen, Alp. Fl. N. G. 2 (1980)
1158, f. 375. — Trisetum sibiricum Rupr. ssp. papillosum Roshev., Bull. Jard.
Bot. Russ. 21 (1922) 2. — Trisetum bifidum (Thunb.) Ohwi var. papillosum
Ohwi, J. Jap. Bot. 17 (1941) 445. — Lectotype: Faurie 2359 (G, holo; P, n.v.)
(cf. Van Royen, 1980), Japan, Tokyo.
Trisetum flavescens (Linné) Beauv. var. macranthum Hack., Bull. Hb. Boiss. 7
(1899) 703; Honda, J. Fac. Sc. Imp. Univ. Tokyo III, 3 (1930) 126. — Trisetum
bifidum (Thunb.) Ohwi var. macranthum Ohwi, J. Jap. Bot. 17 (1941) 445. —
Type: Faurie 7171 (G, holo, n.v.), Japan, Prov. Ishikari, Sapporo.
Trisetum taquetii Hack. in Fedde, Repert. 12 (1913) 386; Honda, J. Fac. Sc. Imp.
Univ. Tokyo III, 3 (1930) 128. — Type: Taquet 3403 (W, holo; US, n.v.), Korea,
Quelpaert, Hallaisan, July 1909.
Trisetum biaristatum Nakai, Bot. Mag. Tokyo 35 (1921) 150; Honda, J. Fac. Sc.
Imp. Univ. Tokyo III, 3 (1930) 125. — Trisetum bifidum (Thunb.) Ohwi var.
biaristatum Honda, fide Ohwi, J. Jap. Bot. 17 (1941) 445. — Type: Matsuzaki
s.n. (n.v.). Japan, Prov. Izo, Aogashima, 1920.
Trisetum bifidum (Thunb.) Ohwi forma contractum Ohwi, Bot. Mag. Tokyo 45
(1931) 192. — Types: Faurie 883, 2288 (n.v.), Korea.
Trisetum bifidum (Thunb.) Ohwi var. oshimense Honda, Bot Mag. Tokyo 49 (1935)
697. — Type: Jotani s.n. (TI, holo, n.v.), Japan, Prov. Izu, Oshima, Sashikiji,
1933.
Trisetum bifidum (Thunb.) Ohwi var. viride Honda, Bot. Mag. Tokyo 49 (1935)
697. — Type: Jotanis.n. (TI, holo, n.v.), Japan. Prov. Izu, Oshima, Motomura,
1932.
130 Gard. Bull. Sing. 36(1) (1983)
Tussocky perennial, up to 90 cm high, with a shortly creeping rhizome. Culms
densely pubescent at base, becoming glabrous upwards. Ligules collar-shaped, 0.5-2
mm long, margin usually glabrous. Blades flat, up to 20 cm by 5 mm, + appressed
pubescent to glabrous above, glabrous beneath. Peduncle glabrous. Panicle + lax,
nodding, 10-20 by 2-5 cm across, axis glabrous, lower branches 2 or 3 together,
longest one up to 5.5 cm long with 20-25 spikelets, scabrous. Spikelets 5-9 mm long,
2-4 flowered, at least the lower two florets bisexual. Glumes acuminate, scabrid on
the nerves; lower glume 2-5 mm long, 0.5-0.6 times as long as the first lemma,
l-nerved; upper glume 4.5-6.5 mm long, 0.6-0.75 times as long as the second lem-
ma, 3-S-nerved. Rachilla nodes 1.5-2 mm long, process 2-2.7 mm long, hairs
0.2-0.5 mm long. First lemma lanceolate, 5-7.5 mm long, scabrid-punctate,
somewhat shiny at base, 3-nerved, callus hairs + 0.3 mm long, apical teeth 1.5-2
mm long; awn L-shaped, inserted in the apical quarter, usually horizontally patent,
column 2-4 mm long, arista 5-7 mm long. First palea 3-5 mm long, keels ciliate,
apex bifid, brown-suffused. Lodicules 0.8-1 mm long, oblong, bidentate. Anthers
0.9-1.3 mm long. Caryopsis fusiform, + 2.75 mm long.
Distribution. China (E. of Szechuan-Kwantung), Korea, Japan, Taiwan,
Malesia: New Guinea (Snow Mts., Lake Habbema, Brass 9118; W. Sepik, Star Mts.,
Tel Besin, Veldkamp 6248; Central, Iwan Swamp, Van Royen 10867).
Ecology. Banks of rivulets, well-drained slopes, on limestone, old native camps,
2660-3225 m in New Guinea, at around 2000 m in Taiwan (Hsu, 1978).
Collectors’ notes. Tussocks to 30 cm across, culms divergent, to 60 cm long.
Leaves bright green. Inflorescence pendulous, spikelets green to silvery purple, awns
divergent in fruit. Anthers and stigmas white.
Chromosome numbers. 2n = 28 (Lee, 1966; Hsu, 1971; Tateoka, 1978), 42 (Hsu,
1972).
Notes. This species is not as closely related to Trisetum flavescens as is sometimes
thought. In Probatova’s system it belongs to subsection Sibirica (Chrtek) Pro-
batova, while the other species, being the generic type, belongs to subsect. Trisetum.
As the latter is a good forage grass, it has been introduced in many parts of the
world, and may perhaps sometime appear in Malesia and Taiwan also. It differs
from 7. bifidum by having at most only a slightly scabridulous lemma with an S-
shaped awn, white to silvery paleas, and anthers 1.5-3 mm long (for the con-
noisseurs: xX = 6, not 7).
Thunberg’s specimen is filed under Avena antarctica in UPS. It is annotated
Bromus bifidus and Aira antaractica and under that latter name there is a collection
by Forster f. marked Bromus bifidus, but with the ‘bifidus’ partly crossed out.
Schweickerdt (Bothalia 3, 1937, 198) who personally studied these specimens noted
that the last one matches another in K, which, we may add, is 7risetum antarcticum
(Forst. f.) Trin. In L there is a specimen given by Thunberg to D. van Royen, which
we assume is an isotype of Bromus bifidus.
The Code errs where it cites, under the nomen conservandum 206 Hierochloé,
Trisetum in Malesia & Taiwan 131
Aira antarctica Forst. f. as the basionym of Disarrenum antarcticum Labill. De
Labillardiére only doubtfully included the reference to Forster’s plant, which is
Trisetum antaracticum (Forst. f.) Trin., while his own is Anthoxanthum redolens
(Vahl) Royen. (See introduction here and R. Brown, Prodr., 1810, 209).
This species has a very curious distribution, ‘jumping’ from Continental Asia and
Taiwan to New Guinea, while it is not found anywhere else in Malesia. Comparable
patterns are known for only a few other species: Carex bilateralis Hayata, C.
brachyathera Ohwi, C. curta Gooden., C. duriuscula C. A. Mey., C. finitima Boott
(also in Sumatra), C. michauxiana Boeck, Eleocharis attenuata (Franch. & Sav.)
Palla, Fimbristylis dictyocolea S. T. Blake (Cyperaceae), Drosera rotundifolia Linné
ssp. bracteata Kern & Steen. (Droseraceae), Germainia capitata Balansa &
Poitrasson, Hemarthria pratensis (Balansa) Clayton (Gramineae), Myriophyllum
propinquun Cunn. or M. verticillatum Linné (Haloragaceae) to which a sterile col-
lection from the Wissel Lakes is thought to belong; Hypericum gramineum Forst.
(Hypericaceae), Utricularia minor (Lentibulariaceae). It is remarkable that, but for
the savannah grasses, all these species are montane to subalpine and rare to very rare
(Germainia ranges from 0-2000 m). The cause of this phenomenon can only be
speculated on. Some also occur in Australia.
2. Trisetum spicatum (Linné) Richt.
Trisetum spicatum (Linné) Richt., Fl. Eur. 1 (1890) 59; Hultén, Svensk Bot. Tidskr.
53 (1959) 203; Bor, Grasses (1960) 448, 701; Chrtek & Jirasek, Webbia 17 (1963)
569; Chrtek, Acta Univ. Carol., Biol. 1967 (1968) 95; C. L. Hitchc. et a/., Vasc.
Pl. Pacif. N. W. (1969) 721; Chrtek, Folia Geobot. & Phytotax. 5 (1970) 447;
Tzvel., Nov. Syst. Pl. Vasc. 7 (1970) 59; Kerguélen, Lejeunia n.s. 75 (1975) 277;
Jonsell et al., Svensk Bot. Tidskr. 69 (1975) 113; Petrovsky, Nov. Syst. Pl. Vasc.
15 (1979) 22; Jonsell, Fl. Eur. 5 (1980) 222. — Aira spicata Linne, Sp. Pl. 1 (1753)
64, non Linné, l.c., p. 63. — Aira subspicata Linné, Syst. Nat., ed. 10, 2 (1759)
873, nom. superfl. — Trisetum subspicatum Beauv., Agrost. (1812) 88, 149, 180,
nom. superfl. — Type: Linné s.n. (LINN, holo, no. 85-7, seen on IDC micro-
fiche; S, iso, n.v., depicted by Hultén, 1959, f. 1), Sweden, Lapland, 1732.
For further synonymy see Hultén (1959) and Kerguélen (1975).
This species has one of the widest natural distributions of flowering plants known
(Hultén, 1959): Eurasia, North and South America, New Zealand and Australia. It
is no wonder that it is very variable and so Hultén thought he could distinguish at
least 14 subspecies in it. Chrtek (1968, 1970) added two more and Petrovsky (1979)
another. Unfortunately Hultén gave no key, while his diagnoses are often incom-
patible and vaguely worded; several of his names are moreover invalid as he did not
indicate types. Opinions on the acceptance of these subspecies differ widely. Hit-
chcock ef a/. (1969) remark that at least for their area ‘well marked geographical
races are not delimitable ... (and) serve(s) no useful purpose’, while Bor (1960)
remarked that for the taxa in the Himalaya ‘it is by no means possible to make any
clear-cut division ... (It) is a polyploid complex which cannot be worked out by the
examination of even a large number of dried specimens.’ Chrtek (1970), however,
‘is inclined to accept’ them. Undoubtedly the best guide for identification at present
132 Gard. Bull. Sing. 36(1) (1983)
is provided by the provenance, but at least the plants from Sabah and Taiwan ap-
peared to be really distinct from each other and from material identified by Hultén
as ssp. alaskanum in S, although that does not seem to be homogenous and can
possibly be split up further. Therefore, although a much clearer account of the com-
plex is necessary to clear up the difficulties, we have decided to follow Hultén and
Chrtek. The plants from Taiwan must therefore be regarded as a subspecies to avoid
the suggestion that they might represent a taxon subjugated to one of Hultén’s, e.g.
ssp. alaskanum.
2a. Ssp. formosanum (Honda) Veldk., comb. & stat. nov. — Fig. 1b.
Trisetum formosanum Honda, Bot. Mag. Tokyo 41 (1927) 636; J. Fac. Sc. Imp.
Univ. Tokyo III, 3 (1930) 128. — Trisetum spicatum (Linné) Richt. var. for-
mosanum Ohwi, J. Jap. Bot. 17 (1941) 443; Chen & Hsu, J. Jap. Bot. 37 (1962)
301; Hsu, Taiwania 17 (1972) 54, f. 63; Taiwan Gr. (1975) 287, f. 38; Fl. Taiwan
5 (1978) 424, f. 1384. — Type: Matsuda 29 (TI, holo, n.v.), Taiwan, Mt.
Nokozan, 1919.
Trisetum spicatum auct. non Richt., e.g. Anon., Icon. Corm. Sin. 5 (1976) 92, 845,
pro specim. taiw.
Trisetum spicatum (Linné) Richt. ssp. alaskanum auct. non Hultén, e.g. Hultén,
Svensk Bot. Tidskr. 53 (1959) 212, pro specim. taiw.
Trisetum subspicatum auct. non Beauv., e.g. Honda, J. Fac. Sc. Imp. Univ. III, 3
(1930) 129, pro specim. taiw.
Tussocky perennial, up to 50 cm high. Culms glabrous at base, becoming pubes-
cent upwards to villous under the panicle. Ligules lingular, 1.5-3 mm long, margin
sometimes erose, glabrous. Blades flat, up to 15 cm by 5 mm, glabrous,
scabridulous. Peduncle villous. Panicle contracted, erect, 8-15 by 0.5-1.5 cm
across, axis villous. Spikelets 5-7 mm long, 3-flowered, the uppermost florets
sometimes variously reduced. Glumes acute, scabrid on the nerves, greenish, rarely
purple; lower glume 3.5-5 mm long, 0.6-0.9 times as long as the first lemma,
1-nerved; upper glume 5-7 mm long, + 0.9-1 (or more) time as long as the second
lemma, 3-nerved. Rachilla nodes + 1.2 mm long, process + 1.5 mm long, hairs
0.8-1.1 mm long. First lemma 4.5-6 mm long, scabridulous, 3—-5-nerved, callus hairs
0.2-0.5 mm long, apical teeth + 0.4 mm long; awn nearly straight to S-shaped, in-
serted in the upper>-=th, column 1-2 mm long, arista + 4 mm long. First palea
3-5 mm long, keels scabridulous, apex bidentate. Lodicules + 0.5 mm long, biden-
tate. Anthers + 1.75 mm long. Caryopsis not seen.
Distribution. Endemic to Taiwan [Van Steenis 20698, 20937 (L), Tamura et al.
20823, 22135 (S) }.
Ecology. Subalpine grasslands, roadsides in Pinus forest, 3000-3950 m.
Chromosome number. N = 14 (Chen & Hsu, 1962).
Note. The ecological data have been taken from the numbers cited above as other
records are scanty, at most stating ‘alpine grass’.
Fig. 1. Spikelets of Trisetum spicatum (Linné) Richt. — a. subsp. kinabaluense Chrtek, J.M.B. Smith
512; b. subsp. formosanum (Honda) Veldk., Tamura et al. 20823; all x 6.
2b. Ssp. kinabaluense Chrtek — Fig. la.
Trisetum spicatum (Linné) Richt. ssp. kinabaluense Chrtek, Folia Geobot. &
Phytotax. 5 (1970) 447. — Type: Clemens 51668 (BM, holo, n.v.; L), Sabah, Mt.
Kinabalu, Donkey’s Ears and Giant Saddle near Low’s Peak, over 3960 m,
January 1934.
Deschampsia sp.: Steen., Bull. Jard. Bot. Btzg. III, 13 (1934) 214.
Trisetum spicatum auct. non Richt.: J. B. M. Smith, New Phytol. 84 (1980) 563,
565, 566.
Trisetum spicatum (Linné) Richt. ssp. alaskanum auct. non Hultén: Hultén, Svensk
Bot. Tidskr. 53 (1959) 212, pro specim. born.
Tussocky perennial, up to 40 cm high. Culms pubescent at base, becoming villous
upwards. Ligules lingular, 2-3 mm long, margin sometimes incised, ciliolate. Blades
flat, up to 15 cm by 5 mn, slightly to moderately pubescent. Peduncle villous. Pani-
cle contracted, 8-10 by 1-2.5 cm across, axis villous. Spikelets 6-9 mm long,
3-flowered, the uppermost floret variously reduced. Glumes acute, slightly scabrid,
with distinct brown, green and purple areas, sometimes green all over; lower glumes
6-7 mm long, about as long as the first lemma, 1(-3)-nerved; upper glume + 8 mm
long, + 0.8 times as long as the second lemma, 3-nerved. Rachilla nodes 1.5-2 mm
long, process + 1.2 mm long, hairs + 0.7 mm long. First lemma 6-6.5 mm long,
scabridulous, 5-nerved, callus hairs 0.1-0.3 mm long, apical teeth + 0.7 mm long;
awn L-shaped, horizontally patent, inserted in upper th, column 2-3 mm long,
arista + 4 mm long. First palea + 5 mm long, keels scabridulous, apex bidentate.
Lodicules + 0.3 mm long, bifid. Anthers + 1.5 mm long, yellow. Caryopsis not
seen.
133
134 Gard. Bull. Sing. 36(1) (1983)
Distribution. Endemic to Mt. Kinabalu, Sabah. (Clemens 30261, 51668, SAN
29281 Meijer, 82997 Cockburn & Aban, Rao et al. 56, Sinclair et al. 9185, J. M.
B. Smith 459, 512, B. C. Stone 11342, Wong 30).
Ecology. Seepage fens, disturbed places, sometimes weedy, 2650-3760 m alt.
Excluded Taxa
1. Trisetum antarcticum (Forst. f.) Trin. var. densum Ridl. in Forb., A naturalist’s
wanderings (1885) 522, nomen. — Lectotype: Wiles & Smith s.n. (BM, holo;
L), Timor, Kupang, October 1792.
Although this combination was not validly published, it represents the pur-
ported occurrence of the species for Malesia. The plants on which it was based
belong to a form of 7. antarcticum. At BM there are two collections from
Timor, one by Nelson, who accompanied Capt. Bligh on the expedition of the
Bounty, and one by Wiles & Smith, who joined the Captain on his voyage with
the Providence. Trisetum antarcticum, however, Occurs only in New Zealand
and some Antarctic Islands. It is, moreover, doubtful whether Nelson collected
any plants in Timor, as Bligh (Voyage to the South Sea, 1792, 240, fide Britton,
J. Bot. 54, 1916, 352) reported that he was too ill to move around. If the species
indeed did occur in Timor (it has not been collected since), it could only have
grown in the mountains, where none of these collectors could have gone. It is
obvious that mislabeling must have taken place.
2. Trisetum latifolium Ridl., Trans. Linn. Soc., London, Bot. 9, 1 (1916) 250. —
Lectotype: Kloss s.n. (BM, holo; L), New Guinea, Mt. Carstensz, Camp 1x-x,
1710-1950 m, 26 January 1913. = Arundinella furva Chase, non A. latifolia
Fourn. (1886).
3. Trisetum subspicatum auct. non Beauv.: Ridl., J. Linn. Soc., London, Bot. 9,
1 (1916) 250; Steen., Bull. Jard. Bot. Btzg. III, 13 (1934) 217. — Trisetum
spicatum auct. non Richt.: Henty, Bot. Bull. Lae 1 (1969) 189. — Anthoxan-
thum horsfieldii auct. non Mez: Royen, Alp. Fl. N. G. 2 (1980) 1188. = An-
thoxanthum angustum (Hitche.) Ohwi.
Ridley’s original speciments (Kloss s.n., Mt. Carstensz, camps xiii-xiv) were
seen at BM. Van Royen equated it with Anthoxanthum horsfieldii, but that is
an endemic Javanese species. Henty, who had no access to the specimens, mere-
ly corrected the name used by Ridley.
Appendix
(by J. F. Veldkamp)
Hultén did not validly publish all his new taxa because he forgot to indicate their
types. Now that we have seen some of his specimens in S which were mentioned by
him and sometimes even depicted, these names can be validated:
Trisetum in Malesia & Taiwan 135
1. Trisetum spicatum (Linné) Richt. ssp. australiense Hultén, Svensk Bot. Tidskr.
53 (1959) 220, f. 3-e, 4-1, 10, excl. syn., quae ad T. antarcticum (Forst. f.) Trin.,
speciem diversam pertinent. — Lectotype: S. Helms (not ‘Helmes’) 847 (S,
holo; C, n.v.), Australia, N. S. Wales, Mt. Kosciusko, 2135 m, 15 April 1922.
2. Trisetum spicatum (Linné) Richt. ssp. himalaicum Hultén, Svensk Bot. Tidskr.
53 (1959) 213, f. 2-e, 4-1, 7, 9. — Lectotype: Polunin 1175 (S, holo; BM, n.v.),
Central Nepal, Chilime Kharka, 4570 m, July 1949.
3. Trisetum spicatum (Linné) Richt. ssp. mongolicum Hultén, Svensk Bot. Tidskr.
53 (1959) 214, f. 6, 9. — Lectotype: Roborovsky s.n. (S, holo; LE, n.v.), Tibet,
Kuen-Lun, Ulan Bulak, 5 July 1894.
4. Trisetum spicatum (Linné) Richt. ssp. tol/uccense (Kunth) Hultén var. bar-
batipaleum Hultén, Svensk Bot. Tidskr. 53 (1959) 223. — Lectotype: Pringle
10032 (S, holo; L; C, n.v.), Mexico, Hidalgo State, Trinidad Iron Works, 1770
m, 21 August 1905.
Trisetum spicatum (Linné) Richt. ssp. ovatipaniculatum Hultén was validated by
Jonsell, Svensk Bot. Tidskr. 69 (1975) 132 with its lectotype represented by Hoppe
s.n. (S, holo, n.v.: L), Austria, Karinthia, summit of Mt. Glockner, mistakenly cited
by Hultén (f. 2b) ‘Tirol, Treffer’. Jonsell suggested that Hultén had indicated types
on the sheets, but the ones seen by us were not so designated. Some had remnants
of a red label which had been torn up.
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77
A Revision of Heteropholis and Thaumastochloa (Gramineae)
R. de KONING, M. S. M. SOSEF & J. F. VELDKAMP
Rijksherbarium, Leiden, the Netherlands
EFFECTIVE-PUBLICATION DATE: 14TH OCT. 1983.
ABSTRACT
Heteropholis C. E. Hubb. (Gramineae, Rottboelliinae) has 4 allopatric species in Central Africa,
Madagascar, Sri Lanka and from India to central Malesia. For Taiwan H. cochinchinensis (Lour.)
Clayton var. chenii (Hsu) Sosef & Koning is here distinguished. The related genus Thaumastochloa C.
E. Hubb. has 7 species mainly in Australia of which 3 are newly described here: T. monilifera Sosef &
Koning, T. rubra Sosef & Koning, T. striata Sosef & Koning. T. major S. T. Blake also occurs in the
Aru Islands off the south-west of New Guinea, and T. rariflora (F. M. Bailey) C. E. Hubb. in Papua
New Guinea. A suspected hybrid between T. major and T. pubescens (Benth.) C. E. Hubb. is reported.
A cladistic study indicates that the two genera may be paraphyletic.
INTRODUCTION
HISTORICAL BACKGROUND
When C. E. Hubbard (1936) described Thaumastochloa (Gramineae, Rott-
boelliinae) he included T. cochinchinensis (Lour.) C. E. Hubb. from SE. Asia and
for Australia T. brassii C. E. Hubb., 7. pubescens (Benth.) C. E. Hubb., T.
rariflora (F. M. Bailey) C. E. Hubb. and an unnamed, insufficiently known species.
Ohwi & Odashima (1937) included 7. shimadana (Ohwi & Odashima) Ohwi &
Odashima from Taiwan, which was soon reduced to a form of 7. cochinchinensis
by Ohwi (1942). In 1941 S. T. Blake added T. major S. T. Blake and T. constricta
S. T. Blake from Australia, while Hsu (1971) proposed 7. chenii Hsu for Taiwan.
Heteropholis as described by C. E. Hubbard (1956) originally included the
African H. sulcata (Stapf) C. E. Hubb., the Ceylonese H. nigrescens (Thw.) C. E.
Hubb. and an unnamed species from Madagascar, which he graciously left to Miss
Camus to describe. She described it in 1956 as H. benoistii.
Clayton (1981) transferred 7. cochinchinensis to Heteropholis and remarked that
as a result Thaumastochloa would be restricted to Asutralia. This aroused our
curiosity as Jansen in his unpublished treatment of the grasses of Malesia, Chase
(1939), Reeder (1948), Henty (1969), and Lazarides (1980) had recorded at least two
other species for Malesia. As the identification of these collections had turned out
to be difficult, a full survey of all material became necessary, and as we were not
quite satisfied about the generic delimitation of the two genera, Heteropholis was
also completely revised.
RELATIONSHIPS OF THE GENERA
Both genera can be placed without any problem in the Roftboelliastrae sensu
Clayton (1973), but the generic delimitation within this group and its distinction
137
138 Gard. Bull. Sing. 36(1) (1983)
from related genera is far from clear, as was also indicated by Clayton. The use of
fused versus free stipes of the pedicelled spikelets to distinguish the Rottboelliastrae
from the Coelorachidastrae, respectively, makes the species with partly fused ones
difficult to place, e.g. Rottboellia coelorachis Forst.f., Robynsiochloa purpurascens
(Robyns) J.-Fél. of the Rottboelliastrae and Coelorachis striata (Steud.) Camus, oc-
casionally of the Coelorachidastrae.
The presence of two sessile and one pedicelled spikelet per joint is supposed to
be the character that distinguishes Mnesithea Kunth from Coelorachis Brongn. The
expression of this feature, however, is variable even within a single inflorescence of
M. laevis (Retz.) Kunth, the type, while on account of the presence of stipes that
are at least apically fused to the rachis, the species should not have been included
in the Coelorachidastrae. Other species sometimes regarded as belonging to
Mnesithea often have only a few triads at the base of the inflorescence and with their
free stipes are better accommodated in Coelorachis (Heidweiller & Van der
Klaauw, MS). One collection of 7. major (q.v.) also had such triads and fused
stipes.
Several species of Heteropholis and Thaumastochloa have been included in
Ophiuros Gaertn., which differs mainly by having the sessile spikelets in two oppos-
ing rows while the lower floret of the sessile spikelet is usually paleate and male, but
may also become epaleate and sterile or paleate and female or bisexual with lodicules
present whenever the sexual organs are developed (Avé, MS). In other Rott-
boelliinae the pedicelled spikelets alternate from the right hand side to the left, joint
for joint, throughout the raceme.
Close to Heteropholis seems to be Robynsiochloa purpurascens of which we have
seen no material. Here the lower floret of the sessile spikelet is paleate and male,
while the pedicelled spikelet is well-developed, 1- or 2-flowered with the lower floret
male and the upper male or sterile (apparently without lodicules). Although this
species resembles especially H. su/cata, it is an annual and thus does not fit the trans-
formation series thought to be represented in Heteropholis.
The distinction between Heteropholis and Thaumastochloa is at first rather ob-
vious: Heteropholis comprises perennial plants with a persistent peduncle and com-
pletely fragmenting spikes or racemes, the species allopatrically distributed in the
Afro-asian tropics, while Thaumastochioa contains annual plants in which the
peduncle breaks off at its base and remains attached to the lowermost joint of the
fragmenting spike (of course not fragmenting when only one joint is developed), the
species being found more or less sympatrically in tropical Australia.
On the other hand there appears to be a gradual transition in the reduction of the
various parts from West to East and we have tried to test the hypothesis that the
present situation would be the result of a West-East migration with an ultimate cen-
tre of speciation in Australia from an ancestor which had become annual and had
evolved this peculiar diaspore.
Heteropholis & Thaumastochloa 139
DIFFERENTIATING CHARACTERS FOR GENERA
When looking at the differentiating characters singly they seem not to be too im-
pressive at the generic level.
Life form has occasionally been used to distinguish between genera, e.g. in Euro-
pean literature, but even for Europe this is not very satisfactory: Avena L. would
include annual species only, but Baum (1977) grudgingly had to include one peren-
nial, which renders a distinction against Helictotrichon Besser and Avenula (Dum.)
Dum. very vague (see also Sevenster & Veldkamp, 1983). A similar situation where
life form is apparently the single distinctive and thus an unsatisfactory feature is pre-
sent in TJrisetum Pers., Koeleria Pers. (perennials), and Rostraria Trin., and
Trisetaria Forsst. (annuals) (see Veldkamp & Van der Have, 1983). On the other
hand many genera are generally accepted to contain both annual and perennial
species, a distinction which in the humid tropics becomes difficult to maintain
anyhow. The species of Thaumastochloa are apparently annual, but a few collec-
tions of 7. major have been annotated ‘biennial’ and ‘short-lived perennial’, so
whatever the latter may be, doubt creeps in whether under favourable situations an
extended longevity would not be possible in at least that species.
Is mode of dispersal a ‘dependable’ generic character? Apart from the fact that
in taxonomy no a priori value can be given to any feature, although some act as if
it were otherwise, and that evaluation depends on the situation and the careful con-
sideration of the revisor, it is frequently employed in this alliance, Clayton (1981),
for instance, distinguished Manisuris L. from Glyphochloa Clayton on the mode of
disarticulation of the raceme. In many cases, however, the form of the diaspore
depends more on the degree of modifications of structures already present in the
related taxa than on the presence of really different ones. In Heteropholis the lowest
point of breakage in the inflorescence is below the lowest joint of the rachis, and
in Thaumastochloa this is at the base of the peduncle, but it seems illogical to confer
a separate generic status to 7. rariflora where the articulation often takes place at
the one but last vegetative node, so that the diaspore is composed of the uppermost
internode and its leaf, the peduncle and the single or double joints of the rachis.
It should be obvious that distribution by itself cannot be considered as a generic
character, and even its employment at the infra-specific level may not be absolute.
Theoretically, disjunctions and isolation may cause distinct forms to evolve into
distinct species and in the end, perhaps, genera, but in the present case the ‘unit’
Thaumastochloa fits in very well with the allopatric pattern shown by the species
of Heteropholis. If there was but one species in Australia, how great would then be
the inclination to distinguish it at generic level? The fact that there are several in-
stead should not be an inducement to do so, tempting as it may be. Distribution
should be correlated with other characters and can then at most be regarded as a
confirmation of a conclusion already reached.
The correlation between life form, diaspore and distribution shown by the present
twe genera is of course interesting, but do three rather weak characters constitute
proof when found together? At most they are very suggestive, but the alternative
possibility that only a single genus with radiate speciation in Australia is involved
could not be overlooked.
1 10
4 1
5 4
6 5
14 6
10 10
11 14
12 11
14 12
7 7
9 9
8 8
12 12
13 13
2 2
3 3
1 1
2 P 2 P
3 3
4 4
: : \
6 6
7 © P 7 @) P
8 8
9 9
10 10
11 P 11 P
12 P 12 P
13 P 13 P
14 (P) 14
H1 H2 H3 H4 F172 1374 157617 H1 H2 H3H4 11 T2 73 14 15 T6T7
Fig. 1. Alternative models of phylogenetic relations in Heteropholis and Thaumastochloa.— @ © a:
different apomorphous character-states. — p: parallel apomorphous states. — 1-10: character-
states according to Table 1. — H1-4, T1-7: species of Heteropholis and Thaumastochloa. For
further explanation see text.
Table 1. Survey of characters
1. Plants perennial yes / no
2. Sheaths longer than 0.5 times the internodes yes / no
3. Peduncle falling of with the uppermost
internode and its leaf no / yes
4. Peduncle articulating at base no / yes
5. Peduncle adnate to the lower-most joint of the spike no / yes
6. Peduncle smooth yes / no
7. Number of spikelets per spike 11--more / 4--10 / 1--3
8. Spikes homomorphous yes / no
9. Articulation of joints straight yes / no
10. Lower glume distinctly winged yes / narrowly / no
11. Margin of lower glume puberulous at base yes / no
12. Nerves of lower glume UH SY Ml
13. Anthers (length in mm) 2--more / 1--2 / up to 1
14. Pedicelled spikelet male / 2 glumes only / | glume / 0
It is presumed that the character state on the left hand side is plesiomorphous, except for 2, /2 and /3,
where the sequence is arbitrary.
140
Heteropholis & Thaumastochloa 14]
We disagree with the views of Roberty (1960), who regarded all species of
Thaumastochloa as subvarieties of Rottboellia corymbosa L.f. (now Ophiuros ex-
altatus (L.) O. Ktze) and those of Heteropholis as varieties of Rottboellia myuros
(L.) Benth. (now Manisuris myuros L.).
A CLADISTICAL APPROACH
The use of cladistics causes one to look very carefully at the characters employed
in a taxonomic study and may reveal certain erroneous preconceptions. We have
employed this method in an attempt to shed some light on the present situation.
After extensive descriptions of all the taxa were made, at least fourteen characters
seemed to be of some significance (Table 1) from which two alternative models
could be deduced, having as few parallel apomorphies as possible (Fig. 1). In order
to estimate the probable polarity of a character-state, the account of Clayton (1973)
was used where numerical taxonomy was applied to the awnless genera of the An-
dropogoneae. The Rottboelliastrae as distinguished by him, excluding our genera,
has been used as the out-group.
Characters 4, 5 and 6 should be regarded as belonging to the complex that makes
an effective pseudo-awn possible and are probably best considered as a single
character; they have therefore been enclosed by a broken line.
To characters 12 and 13 no polarity could be attributed, but the arbitrary choice
made here does not seem to have much effect on the models.
For T. major, character 7 has been placed between brackets in the lower part of
fig. 1 (summation of features) because of the heteromorphous spikes. Characters 3
and 8 might have been omitted because of their uniqueness for which reason the
creeping habit of H. nigrescens was not mentioned, although these features set the
various species apart.
In model A one can decide that a single genus is represented, if no generic value
is accredited to the set of characters offered by 1, 4, 5, 6, 10 and 14. These divide
the taxa into two groups which correspond with the two original genera. When,
however, the correlation of these, perhaps minor characters is considered as
evidence for a generic status, the old situation is maintained.
In model B a single genus can also be accepted, the extent of which remains uncer-
tain; it is more easy here to envisage further steps ‘upwards’, whereby more taxa
would be included. No infra-generic groups can formally be distinguished, however,
because of the paraphyly involved, although the Thaumastochloa-alliance remains
distinct. To do so would require the creation of three or four distinct taxa, which
seems highly unsatisfactory. Nevertheless, because of one lesser partial parallelism
between Heteropholis and Thaumastochioa, this model has a slight advantage. This
model corresponds with a clinal variation coupled with the reduction of the pedicell-
ed spikelet terminating in a secondary radiative speciation in Australia, which was
the alternative explanation outlined above before the cladistics were done. It should
142 Gard. Bull. Sing. 36(1) (1983)
be noted that in neither model there is a complete correlation with the geography:
the pedicelled spikelet is more reduced in H. benoistii from Madagascar than in H.
nigrescens from Sri Lanka.
To make an educated guess as to which model should have preference, the out-
group must be known in more detail. Only then a good insight can be obtained of
the evolutionary tendencies within it. At present it seems therefore irresponsable to
choose in favour of either model with any certainty. Hence, for the present, matters
are better left as they were, and so we have kept the genera distinct.
ACKNOWLEDGEMENTS
This survey was started during a course in advanced Angiosperm taxonomy by the
first two authors based on the material present in L. Subsequently other material
was kindly sent on loan from A, BM, BO, BRI, K, KYO, P, PERTH, SING, TAI,
TI, and W, to the Keepers of which we are very grateful. The last author guided
the process, edited and translated the final report. The Director and Staff of the
Rijksherbarium are much thanked for the opportunity and assistance, especially Mr.
J. H. van Os who prepared the fine drawings and advised on the graphical presenta-
tion. Advice rendered by Dr. M. Zandee, Laboratory of Experimental Plant
Systematics, Leiden, and Dr. R. van der Meijden (L) on the intricacies of cladistics
are much appreciated. Finally, the authors wish to thank Mr. Wong Khoon Meng,
Forest Botanist, Forest Research Institute in Kepong, for his critical comments on
the orginal manuscript.
LITERATURE CITED
Avé, W. Ophiuros exaltatus (Linné) O. Ktze (Gramineae). Manuscript. Rijksher-
barium, Leiden.
Baum, B. R. (1977). Oats: wild and cultivated: 25. Ottawa.
Camus, A. (1956). Une nouvelle espéce du genre Heteropholis Hubbard
(Graminées). Bull. Soc. Bot., France 103: 476.
Chase, A. (1939). Papuan grasses collected by L. J. Brass. II. J. Arn. Arb. 20:
314-315.
Clayton, W. D. (1973). The awnless genera of Andropogoneae. Studies in the
Gramineae: XXXIII. Kew Bull. 28: 49-57.
(1981). Notes on the tribe Andropogoneae (Gramineae). Kew Bull. 35:
813-818.
Heidweiller, J. & M. A. F. van der Klaauw. Differences between Mnesithea and
Coelorachis (Gramineae) in South East Asia. Manuscript. Rijksherbarium,
Leiden.
Henty, E. E. (1969). A Manual of the Grasses of New Guinea. Bot. Bull., Lae 1:
181.
Hsu, C. C. (1971). A guide to the Taiwan grasses, with keys to sub-families, tribes,
genera and species. Taiwania 16: 216-218, 335, f. 2, 2a.
Heteropholis & Thaumastochloa 143
Hubbard, C. E. (1936). Thaumastochloa C. E. Hubbard, in W. J. Hooker, cones
plantarum 34: t. 3313, 3314.
—— (1956). Heteropholis sulcata (Stapf) C. E. Hubbard, in W. J. Hooker,
Icones plantarum 36: t. 3548.
Jansen, P. Gramineae of Malesia. Manuscript. Rijksherbarium, Leiden.
Lazarides, M. (1980). The Tropical Grasses of Southeast Asia: 74-75. Vaduz.
Ohwi, J. (1942). Gramina japonica. IV. Acta Phytotax. & Geobot. 11: 178.
— &K. Odashima. (1937). In J. Ohwi, Symbolae ad Floram Asiae orientalis.
15. Acta Phytotax. & Geobot. 6: 151.
Reeder, J. R. (1948). The Gramineae-Panicoideae of New Guinea. J. Arn. Arb. 29:
354-355.
Roberty, G. (1960). Monographie systématique des Andropogonées du Globe.
Boissiera 9: 65-66, 81-82.
Sevenster, J. G. & J. F. Veldkamp. (1983). A Revision of Helictotrichon (Grami-
neae) in Malesia. Blumea 28: 329-342.
Veldkamp, J. F. & J. C. van der Have. (1983). The Genus Trisetum (Gramineae)
in Malesia and Taiwan. Gard. Bull. Sing 36: 125-135.
KEY TO THE GENERA
1. Perennials. Peduncle smooth, not articulating at base. All joints of the spike articulating in fruit.
Central Africa to Micronesia, not in New Guinea. .................-0e0 000s Heteropholis
1. Annuals. Peduncle scabrous, articulating at base or with the uppermost internode and its leaf.
Lowermost (or the only) joint of the spike persistent in fruit. Australia, New Guinea. .....
25 66 050 (ACRE BOOT 6 GUIS BEUISE RTO IRR Ce eI ee eee ean ieee ioe Thaumastochloa
HETEROPHOLIS
HETEROPHOLIS C. E. Hubb. in Hook., Ic. Pl. 36 (1956) t. 3548; Camus, Bull. Soc.
Bot., France 103 (1956) 475; Clayton, Kew Bull. 35 (1981) 816. — Type: H.
sulcata (Stapf) C. E. Hubb.
Perennials, branching intra-and/or extra-vaginally at base. Ligules collar-shaped,
membranous, margin ciliolate. Peduncle glabrous, smooth, not articulating at base.
Spikes + terete, dehiscing transversally into swollen joints, which bear a sessile
spikelet and a more or less reduced pedicelled one, with a ‘knob’ at base (remnants
of the vasculature). Spikelets unilaterally in 1 or 2 slightly alternating rows, un-
awned. Sessile spikelets sunken into the joints, with 2 florets, the lower one reduced
to a lemma, the upper one bisexual. Callus short, glabrous. Lower glume slightly
convex, indurated, 5-11-nerved, smooth or with longitudinal rows of small pits be-
tween the nerved or rugose, 2-keeled, glabrous or margins at base puberulous, mar-
gins narrow, stiff, infolded, apex more or less winged, acutish. Upper glume boat-
shaped, slightly keeled, 3-7-nerved, scarious to membranous. Lemmas and paleas
obtuse, scarious to slightly membranous. First lemma 2-nerved. Second lemma
0-3-nerved. First palea absent, second palea present, 0- or 2-nerved. Lodicules pre-
Fig. 2. Sessile spikelets of Heteropholis. — a. H. sulcata (Quarré 137); b. H. benoistii (Bosser 15241);
c. H. nigrescens (Davidse & Sumithraarachchi 7961). x 15.
Map 1. Solid stars: Heferopholis sulcata; diamond: H. benoistii; open star: H. nigrescens; circles: H.
cochinchinensis, s.l.; triangles: Thaumastochloa spp.
144
Heteropholis & Thaumastochloa 145
sent in the upper floret, broad- to oblong-cuneate, truncate, glabrous. Anthers 3,
purple. Styles 2, free; stigmas purple. Caryopsis ovoid to broadly ellipsoid, dorsally
flattened; hilum (sub-)basal, punctiform; embryo 0.5-0.8 times as long as the
caryopsis. Stipe of the pedicelled spikelet partly to entirely adnate to the joint.
Pedicelled spikelet much reduced to absent, 1-flowered, male or neuter. Glumes,
when well-developed, herbaceous to indurated, the lower one keeled, 5-9-nerved,
not sculptured, the upper one boat-shaped, sometimes with an infolded margin,
3-5-nerved. Lemma and palea much reduced. Lodicules present or not. Anthers 0
or 3.
Distribution. Four species allopatric in E. Africa, Madagascar, Sri Lanka, or
from SE. Asia to S. China, E. Malesia and Micronesia.
Ecology. Savannahs, dry grasslands, forest edges, up to 1800 m alt.
Chromosome numbers. N = 18 [H. cochinchinensis (Lour.) Clayton var. cochin-
chinensis, H. nigrescens (Thw.) C. E. Hubb.].
Note. Hubbard described the sessile spikelets as 2-flowered with the lower floret
being male or sterile and with a palea equal to the lemma or shorter to absent. Hsu
(1971) also described and depicted a first palea for H. cochinchinensis (Lour.)
Clayton (q.v.). However, in all specimens seen by us this floret was always reduced
to an empty lemma.
MELCICEHEGESDI KCletS7ADSETIT AS Ee ASIAL Root ec rvel a: als Spee aie SSS Stee a0 1S a) e Visieicw)siercie Ve eiyeyere oS. 2
1. Pedicelled spikelets more or less developed. Africa, Madagascar, Sri Lanka. .............. 3
2. Sessile spikelets 2.7-4 mm long. Lower glume narrowly winged. Anthers 1.5-1.7 mm long. .
» os tho can thOOOthe DU ORO Ce OCD Tae eC aera 4a. H. cochinchinensis var. cochinchinensis
2. Sessile spikelets 4-4.5 mm long. Lower glume distinctly winged at the apex. Anthers + 2.8 mm
[One "TERE -Aseslou oe cc SC BS OCU ett ane ee 4b. H. cochinchinensis var. chenii
3. Blades 4-10 mm wide, at least the margins pubescent. Pedicelled spikelets 4-5 mm long. ... 4
Blades 1.5-2 mm wide, glabrous. Pedicelled spikelets 2-3 mm long. Madagascar. .........
Deak e oo SLOG Ho OAD oer OEE nec ee aaa ee ae eee Eb adeeb dat ena, = Eds }DENOISHE
4. Blades up to 12 cm long. Spike 6-10-jointed. Lower glume of the sessile spikelet with small pits.
Rediceledispikelet neuter: pon Malka pa fey aye tant cor oayey ets tS eae seks Stelahe’s lekead nish ls 3. H. nigrescens
4. Blades up to 60 cm long. Spike 20-30-jointed. Lower glume of the sessile spikelet areolate.
Pesiccledispikelen males Central Bo AUmICA. Sieh © naseceus 65) of < Gye sole jopsyere ee ae 1. H. sulcata
1. Heteropholis sulcata (Stapf) C. E. Hubb. — Map 1, fig. 2a
Heteropholis sulcata (Stapf) C. E. Hubb. in Hook., Ic. Pl. 36 (1956) t. 3548, p. 2;
J.-Félix, Gram. Afr. Trop. 1 (1962) 282, f. 218; Clayton & Renvoize, Fl. Trop.
E. Afr., Gram. 3 (1982) 849. — Peltophorus sulcatus Stapf, Fl. Trop. Afr. 9
(1917) 59, comb. ill.; Robijns, Fl. Agrost. Congo Belge 1 (1929) 63. — Manisurus
sulcata Dandy, J. Bot. 69 (1931) 54. — Rottboellia myuros (L.) Benth. var.
sulcata Roberty, Mon. Androp. (1960) 82. — Type: Homblé 56 (K, holo; BR),
Zaire, Katanga, Lubumbashi (Elisabethville), February 1912.
146 Gard. Bull. Sing. 36(1) (1983)
Culms 60-150 cm high. Nodes 3-5, glabrous, rarely hairy. Sheaths 8-12 cm long,
0.4-0.75 times as long as the internodes, subglabrous to hirsute. Blades linear, flat
to involute, up to 60 cm by 4-10 mm, subglabrous to hirsute. Peduncle 12-25 cm
long. Joints 20-30. Sessile spikelets slightly alternating, 3.5-5 mm long. Lower glume
5-9-nerved, areolate because of longitudinal and transverse ribs, margins
puberulous at base, apex winged, acuminate. Anthers 2.4-2.5 mm long. Pedicelled
spikelet more or less well-developed; stipe adnate in the lower half to completely so
with the joint and approximately as long as the joint; spikelet 4-5 mm long, male,
lodicules present.
Distribution. Zaire (Katanga: Bianos, Kalule, Kienge, Kivilolo, Kundelungu
Plateau, Lubumbashi, Manika Plateau, Musoka-Tanda, Tumbwe), Zambia (Aber-
corn, Chishinga, Lundazi, Ndola, Solwezi), Malawi (Mzimba, Mzuzi), Tanzania
(Songea).
Ecology. Open woodland dominated by Brachystegia (e.g. B. boehmii, B.
floribun—., B. microphylla) and with Andropogon schirenzis, Isoberlinia sp.,
900-1830 m alt.
Collector’s notes. Loosely tufted, to 1.5 m high. Rhizome creeping, roots brown,
wiry. Lower sheaths reddish purple. Culms bright green. Leaves erect, apex + pen-
dulous. Sessile spikelets yellowish green. Pedicelled spikelets pale green with dark
green nerves. Anthers, stigmas dark purple. Entire plant turning black. (Mainly after
Milne-Redhead & Taylor 9058, BR).
Vernacular name. Kipolo (Katanga: Kibemba).
Note. The plants are nigrescent with a greyish hue when dried, as was also noticed
in the field.
2. Heteropholis benoistii Camus — Map 1, fig. 2b
Heteropholis benoistii Camus, Bull. Soc. Bot., France 103 (1956) 476. — Type:
Benoist 1639 (P, holo), Madagascar, Domaine Central, Manjakatompo,
Ankaratra, 1700 m alt., 20 December 1951.
Culms 35-50 cm high. Nodes 6-more, glabrous. Sheaths up to 7 cm long, shorter
than the internodes, margins distally pilose, otherwise glabrous. Blades linear, flat
to involute, up to 25 cm by 1.5-2 mm. Peduncle up to 13 cm long. Joints 12-more.
Sessile spikelets slightly alternating, 3-4.35 mm long. Lower glume 5-7-nerved, with
rows of small pits, margins glabrous at base, apex minutely winged, acuminate. An-
thers + 2.1 mm long. Pedicelled spikelets reduced, to the lower glume; stipe adnate
with and + 0.8 times as long as the joint; spikelet 2-3 mm long.
Distribution. Madagascar (Domaine Central).
Ecology. Savannahs, up to 1700 m alt.
Note. Only two collections seen (Benoist 1639, Bosser 15241, P).
Heteropholis & Thaumastochloa 147
3. Heteropholis nigrescens (Thw.) C. E. Hubb. — Map 1, fig. 2c
Heteropholis nigrescens (Thw.) C. E. Hubb. in Hook., Ic. Pl. 36 (1956) t. 3548, p.
4; Bor, Grasses (1960) 162. — Rottboellia nigrescens Thw., Enum. Pl. Zeyl.
(1864) 364; Hack., Mon. Androp. (1889) 296; Hook. f., Fl. Br. Ind. 7 (1896) 157;
Hook. in Trimen, Handb. Fl. Ceyl. 5 (1900) 207; Senaratna, Gr. Ceylon (1956)
180; Gould & Soderstrom, Can. J. Bot. 52 (1974) 1085, 1088, f. 71. — Manisurus
nigrescens O. Ktze, Rev. Gen. PI. 2 (1891) 780. — Rottboellia myuros (L.) Benth.
var. nigrescens Roberty, Mon. Androp. (1960) 81. — Type: Thwaites CP 867
(PDA, holo, K. n.v.; BM, BO, SING), Ceylon, Central Prov., Bagavantalawa,
1220 m alt.
Culms up to 60 cm high. Nodes 10-15, glabrous. Sheaths 3-6 cm long, 0.45-0.75
times as long as the internodes, pilose along the margins, otherwise subglabrous.
Blades linear, flat, up to 12 cm by 6-11 mm, margins hairy at base, otherwise
glabrous. Peduncle up to 5 cm long. Joints 6-10. Sessile spikelets + in 1 row, +
4.5 mm long. Lower glume 11-nerved, with rows of small pits, margins puberulous
at base, apex narrowly winged, acute. Anthers + 2 mm long. Pedicelled spikelet
reduced to the glumes; stipe entirely adnate to and + 0.67 times as long as the joint;
spikelet up to 4 mm long.
Distribution. Sri Lanka: Central and Trincomalee Prov.
Ecology. Cracks of cliffs, forest edges, 1200-1560 m alt.
Chromosome number. 2n = 36 (Gould & Soderstrom, 1974; ‘60’ sphalm. p.
1088).
Collector’s notes. Plants scandent, base long-decumbent, rooting at the nodes.
Note. Plants nigrescent.
4. Heteropholis cochinchinensis (Lour.) Clayton — Map 1.
a. var. cochinchinensis. — Fig. 3a
Heteropholis cochinchinensis (Lour.) Clayton, Kew Bull. 35 (1981) 816. — Phleum
cochinchinense Lour., Fl. Cochinch. (1790) 48. — Paspalum ? cochinchinense R.
& S., Syst. Veg. 2 (1817) 317. — Ophiuros cochinchinensis Merr., Trans. Am.
Philos. Soc., n.s. 24, 2 (1935) 72. — Thaumastochloa cochinchinensis C. E.
Hubb. in Hook., Ic. Pl. 34 (1936) t. 3313, 3314, p. 2; Ohwi, Bot. Mag., Tokyo
55 (1941) 551; Raizada et al., Ind. For. Rec., n.s. 4 (1957) 211; Bor, Grasses
(1960) 247; Dansk Bot. Ark. 23 (1965) 165; Koyama in Walker, Fl. Okinawa
(1976) 238; Anon., Ic. Corm. Sin. 5 (1976) 195, t. 7219. — Rottboellia corymbosa
L.f. subvar. cochinchinensis Roberty, Mon. Androp. (1960) 65. — Thaumasto-
chloa cochinchinensis forma cochinchinensis: Hsu, Taiwania 16 (1971) 335, f. 2a;
Fl. Taiwan 5 (1978) 701, f. 1481. — Type: Hb. Loureiro (BM, holo, n.v.),
Cochinchina.
148 Gard. Bull. Sing. 36(1) (1983)
Ophiuros monostachyus Presl, Rel. Haenk. 1 (1830) 330; Miq., Fl. Ind. Bat. 3
(1857) 405; Hack., Mon. Androp. (1889) 318; Camus, Fl. Gén. I.-C. 7 (1922) 373,
f. 38; Merr., Enum. Philip. Fl. Pl. 1 (1925) 41. — Rottboellia monostachya
Schmid, l’Agron. Trop. 13 (1958) 193, nom. inval. — Type: Haenke s.n. (PR,
holo, n.v.), Philippines, Luzon.
Ophiuros undatus Nees in Hook., J. Bot. Kew 2 (1850) 100; Steud., Syn. 1 (1854)
360 (‘undulatus’); Miq., Fl. Ind. Bat. 3 (1857) 405 (id.); Benth., J. Linn. Soc.,
London, Bot. 19 (1881) 69 (id.). — Type: Cuming 1339 (BM, L, P, W; K, n.v.).
Ophiuros shimadanus Ohwi & Odashima, Acta Phytotax. & Geobot. 5 (1936) 185.
— Thaumastochloa shimadana Ohwi & Odashima, Acta Phytotax. & Geobot. 6
(1937) 151. — Thaumastochloa cochinchinensis C. E. Hubb. forma shimadana
Ohwi, Acta Phytotax. & Geobot. 11 (1942) 178; Hsu, Taiwania 16 (1971) 335;
Taiwan Gr. (1975) 779, f. 285; Fl. Taiwan 5 (1978) 701. — Type: Shimada 4852
(KYO, holo, n.v.), Taiwan, Hsinchu.
Thaumastochloa chenii auct. non Hsu: Hsu, Taiwania 16 (1971) 335, pro Chen 123.
Thaumastochloa cochinchinensis auct. non C. E. Hubb.: Chen & Hsu, J. Jap. Bot.
37 (1962) 306, 313, f. 28, 29, pro Chen 123.
Culms up to 60 cm high. Nodes 3-4, glabrous. Sheaths 3-6 cm long, 0.25-0.75
time the length of the internodes, margins at the top with the long hairs, otherwise
glabrous. Blades linear, flat to involute, up to 45 cm by 2-5 mm, margins at base
with long hairs, otherwise (sub-)glabrous. Peduncle up to 10 cm long. Joints
12-more. Sessile spikelets + in 1 row, 2.7-4(-4.5) mm long. Lower glume 7-nerved,
smooth to minutely pitted in longitudinal rows, margins glabrous at base, apex nar-
rowly winged, obtuse to acute. Anthers 1.5-1.75 mm long. Pedicelled spikelet re-
duced to a small scale; stipe entirely adnate to and + as long as the joint.
Distribution. India (e.g. Uttar Pradesh, Bihar, no doubt elsewhere), Thailand
(Northern: Sukhotai, Nakhon Sawan; Southwestern: Ratchaburi), Vietnam (Tu
Phap, Hué, Hanoi, Phocam), China (Canton, Hainan, Hong Kong, Putien),
Taiwan, Ryu Kyu Isl. (Iriomote, Okinawa), Malesia: Java (Banjumas), Lesser Sun-
da Isl. (Bali, fide Jansen MS), Philippines (Luzon, Mindanao), Moluccas (Buru),
Carolines (Palau, Yap), Marianes (fide C. E. Hubbard).
Ecology. Savannahs, disturbed places, e.g. roadsides, bunds of rice fields, in rice
fields, grass fields, up to 600 m alt.
Collectors’ notes. Very few: caespitose.
Chromosome numbers. N = 18 (Chen & Hsu, 1962, at least pro Chen 123, TAI).
Notes. The only record for Java (not mentioned in the Flora of Java!, Kievits
1543, L) probably constitutes an introduction. It has relatively large lower glumes
and anthers. Jansen (MS) mentioned an unnamed collection present in BO, purported
from Bali but Veldkamp could not find any trace of it there. The record for the
Fig. 3. Sessile spikelets of Heteropholis cont. — a. H. cochinchinensis var. cochinchinensis (Merrill
9875); b. H. cochinchinensis var. chenii (Hsu 511, type). x 15.
Marianas could not be verified with the loan from K.
Although the types of ‘cochinchinensis’, ‘monostachyus’ and ‘shimadanus’ have
not been seen, there is enough evidence from the descriptions, later remarks and
subsequent material for the authors to be certain of their identities. The last epithet
was employed by several authors for specimens with pitted lower glumes, but some
collections (e.g. Balansa 503, L; Chuang 1560, Hsu & Kuoh 13658A, Suzuki s.n.,
Tanaka & Shimada, TAI, etc.) were seen to have both pitted and smooth lower
glumes even within the same inflorescence, from which we conclude that this
character has no great taxonomic significance. However, specimens with both pitted
lower glumes and distinctly larger spikelets and anthers have been tentatively kept
distinct by us as the next variety.
Hsu (1971) described, compared and even depicted lower paleas for both varieties,
but in all the materal seen by us, also that from TAI, we have not even encountered
a trace of them.
b. var. chenii (Hsu) Sosef & de Koning, stat. & comb. nov. — Fig. 3b
Thaumastochloa chenii Hsu, Taiwania 16 (1971) 216, 335, f. 2; Taiwan Gr. (1975)
777, f. 284; Fl. Taiwan 5 (1978) 700. — Type: Hsu 511] (TAI, holo; TI), Taiwan,
Pingtung Co., O-luan-pi, + at sea level, 21 September 1959.
Differs from var. cochinchinensis by the lower glumes of sessile spikelets being
(3.5-)4-4.5 mm long, apex somewhat more broadly winged, always with pits in
longitudinal rows, and especially by the + 2.8-mm long anthers.
149
150 Gard. Bull. Sing. 36(1) (1983)
Distribution. Taiwan (Pingtung Co.).
Ecology. Littoral grasslands.
Chromosome number. Chen 123 on which a count of n = 18 was based and which
was included in this taxon, has + 3.8-mm long sessile spikelets and + 1.7 mm-long
anthers and obviously belongs to the typical variety (q.v.).
THAUMASTOCHLOA — Map 1
THAUMASTOCHLOA C. E. Hubb. in Hook., Ic. Pl. 34 (1936) t. 3313, 3314; Simon,
Techn. Bull. Bot. Br. Dept. Prim. Industr., Brisbane 3 (1978) 13. — Type:
Thaumastochloa pubescens (Benth.) C. E. Hubb.
Annuals (or short-living perennials?), branching intra-vaginally at base. Ligules
collar-shaped, membranous, margin ciliolate. Peduncles scaberulous, articulating at
base or breaking off with the uppermost internode and its leaf, gradually narrowing
to base, at least in the lower third. Spikes + terete, the lower joint (or the only one)
+ persistently adnate to the peduncle, the others (if any) dehiscing transversely to
obliquely into swollen to inflated joints which bear a sessile spikelet and the remnant
of a pedicelled one, with a ‘knob’ at base (remnants of the vascularity). Spikelets
unilaterally in 1 or 2 somewhat alternating rows, unawned. Sessile spikelets sunken
into the joints, with 2 florets, the lower one reduced to a lemma, the upper one
bisexual. Callus short, glabrous. Lower glume convex to slightly concave, in-
durated, 7-9-nerved, smooth or variously longitudinally and/or transversely
sculptured, 2-keeled, margins at base puberulous, otherwise usually glabrous,
margins narrow, stiff infolded, apex somewhat acute, unwinged. Upper glume con-
vex, slightly keeled, 3-5-nerved, scarious to slightly indurated. Lemmas and paleas
obtuse, scarious. First lemma 2-nerved, second one 0-3-nerved. First palea absent,
second one present, 0-nerved. Lodicules present in the upper floret, cuneate, trun-
cate, glabrous. Anthers 3, purple. Styles 2, free; stigmas purple. Caryopsis broadly
ovoid, dorsally flattened; hilum sub-basal, punctiform; embryo 0.5-0.67 times as
long as the caryopsis. Stipe of the pedicelled spikelet adnate to the joint; spikelet
absent or reduced to a miniscule scale.
Distribution. Seven species (and a possible hybrid) more or less sympatric in Nor-
thern Australia (W. Australia, N. Territory, Queensland) and Malesia (Aru Isls,
New Guinea, each with 1 sp.).
Ecology. Savannahs, sandy places, up to 1000 m alt.
Notes. The spikes of most species are homomorphous, but in 7. major S. T. Blake
two different types may be present even on the same plant: long ones with 4-10
relatively broad joints, and short ones with 1-3(-4) relatively narrow joints.
As these plants grow in often nearly inaccessible places and may not be evident
because of their annual life cycle, a wider distribution, certainly in Malesia, may be
expected with increase of the variability as more material is acquired, while
Heteropholis & Thaumastochloa 151
undescribed species may well turn up. Compare for instance the supposedly mono-
typic genus Micraira F.v.M., which now has at least 8 species (Lazarides, Brunonia
2, 1979, 67-84)!
KEY
Peete CASEESOMECISPIKES WITHA —INOLE YOUNES co ace wee ws 0 woe cis 5 nie cero ayer ayeie sieeve e geisie weee ee e,s 2.
REE MBS PIR CS avi tlle —3 OMNES ere eerie acc ecs creer eie 2 oc eyes So) 6ib esi aoleree cave « Geidee srees o < 5
2. Ali joints 1-1.8 mm across — Lower glumes areolate to weakly rugulose, at least in some spikes
209504 CC ORAORB AS SIIORRS oniieeiaies Scioto ect ACIS Re eens a gO a 3
2. At least some joints 2-3.5 mm across — Lower glumes usually smooth to at most weakly
PLORIUTS? = snot cossddcessct cad Outed OOO CED COM TOES SO OeR Oe Og aCe as eT 4
3. Spikes homomorphous, joints 4-9. Lower glumes all weakly rugulose to areolate, but the basal
sometimes smooth. Anthers 0.75-1 mm long ....................2-2-05- 1. T. pubescens
3. Spikes heteromorphous, some 4-9-jointed and lower glumes areolate, others 1-3-jointed and
lower glumes smooth. Anthers + 1.1 mmlong ................. ? T. major x pubescens
4. Culms up to 70 cm high. Spikes homomorphous. Joints inflated. Anthers 0.5-1 mm long
bout cor oar eoatey So FeO CeO BOO E GEL oo 8 oo SOO TGC 2 eee nae ea ae 2. T. monilifera
4. Culms up to 35 cm long. Spikes homo- or heteromorphous. Joints cylindric. Anthers 1.4-1.7
TH OE oaceeonbdodc eee eaauneeocen 4hct Sp SS 3. T. major
SD. DG? GUMS COIN weed aones oapnde 5.505. cso et oe cee eee eae ee ee 6
5. Lower glume longitudinally grooved or transversely rugose ...................-2-.000-- 7
6. Well-developed sheaths 0.17-0.33 time length of internode. Peduncle 0.5-3 cm long, usually
falling off with the uppermost internode. Joints 2.5-3 mm long ........... 7. T. rariflora
6. Well-developed sheaths 0.5->length of internodes. Peduncle 1.5-18 cm long, articulating at
AS. OS SG WIA Gaon os On OB an OR ee se eae eee eee 3. T. major
7. Culm-nodes 4-8. Lower glume longitudinally grooved. Anthers 1.1-1.7 mm long ......... 8
7. Culm-nodes 10-15. Lower glume transversely rugose. Anthers 0.6-0.7 mm long 6. T. brassii
8. Well-developed sheaths 0.17-0.33 time length of internode. Peduncle 0.5-2 cm long, glabrous
cant todas SORE AAO Por Be OO SOOO. TT ote Mec Cee ee eee 5. T. rubra
8. Well-developed sheaths 0.5-same length as internode. Peduncle 4-18 cm long, with rows of
THT TES CAPD in ie Tip oS Ol UE ee ee ee ee 4. T. striata
1. Thaumastochloa pubescens (Benth.) C. E. Hubb. — Map 2, fig. 4
Thaumastochloa pubescens (Benth.) C. E. Hubb. in Hook., Ic. Pl. 34 (1936) t. 3313,
p. 2; Chippendale, Proc. Linn. Soc. N. S. W. 96 (1972) 223; Simon, Techn. Bull.
Bot. Br. Dept. Prim. Industr., Brisbane 3 (1978) 13; ibid. 4 (1980) 84. — Ophiuros
corymbosus (L.f.) Gaertn. var. ? pubescens Benth., Fl. Austr. 7 (1878) 512;
Hack., Mon. Androp. (1889) 318. — Ophiuros pubescens Dom., Bibl. Bot. 85
(1915) 262. — Ophiuros pollockii Marquand, Kew. Bull. (1925) 284, nom.
superfl. — Rottboellia corymbosa L.f. subvar. pubescens Roberty, Mon. An-
drop. (1960) 66, excl. Mnesithea pubescens Ridl. — Type: F. v. Mueller s.n. (K,
holo, n.v.), Australia, N. Territory, sources of Hooker’s Cr., + 18° S, 130° E.
Thaumastochloa constricta S. T. Blake, Papers Dept. Biol. Univ. Queensl. 1, 18
(1941) 19; Simon, Techn. Bull. Bot. Br. Dept. Prim. Industr., Brisbane 3 (1978);
ibid. 4 (1980) 84. — Type: S. T. Blake 12460 (BRI, holo, n.v.; L), Australia,
Queensland, Burke Distr., Croydon, 18° 12’ S, 142° 15’ E, 110 m alt., 8 August
1936.
152 Gard. Bull. Sing. 36(1) (1983)
Ophiuros corymbosus auct. non Gaertn.: R. Br., Prodr. (1810) 207.
Culms 12-30 cm high, rarely more. Nodes 5-9. Sheaths 1-3 cm long, 0.5-same
length as internode, pilose. Blades linear, flat, 2-9 cm by 1.5-4 mm, pilose. Pedun-
cle erect, + straight, gradually narrowed towards the base where it articulates, up
to 10 cm long, glabrous. Spikes homomorphous, cylindric to constricted, 1.5-3.5
cm by 1-1.8 mm across. Joints 4-9, articulation straight to oblique, 3-4 mm long,
glabrous, rarely puberulous. Spikelets somewhat alternatingly in 2 rows, 2-3 mm long.
Lower glume ovate-oblong, flat to convex, the lowermost usually smooth, the others
weakly transversely rugulose to areolate by longitudinal and transverse ribs, 7-9-
nerved. Upper lemma 2-nerved. Anthers 0.75-1 mm long.
Fig. 4. Sessile spikelets of Thaumastochloa — T. pubescens (McKee 9183). x 15.
Map 2. Stars: Thaumastochloa pubescens; circles: T. major x T. pubescens.
Heteropholis & Thaumastochloa 153
Distribution. Malesia: New Guinea (Western Distr.: Arufi, Rouku); Australia:
W. Australia (N. of Broome, Karunjie St., Kimberley Res. St.), N. Territory (Hoo-
ker’s Cr., Katherine, Mittibah St.), Queensland (Atherton, Batavia, Burra, Cam-
paspe R., Cheltenham, Conjuboy St., Cooktown, Croydon, Doomagee, Endeavour
R., W. of Ingham, Jericho, Lizard Isl., Mungana, Mareeba, Normaton, Pentland,
Poison Cr., Port Douglas, Torrens Cr., Townsville, Yarromeere).
Ecology. Low lying, damp sandy places, gullies, exposed coastal sand dunes,
weedy in fields and disturbed places, open savannah forests with e.g. Acacia cow-
leana, Erythrophloeum sp., Eucalyptus calleni, E. dichromophloia, E. miniata, E.
polycarpa, Grevillea glauca, Petalostigma sp., Spinifex sp., Triodia pungens, up to
840 m alt.
Collector’s notes. Tufted, culms prostrate to erect, up to 30 cm high or radiating
to 1.5 m in diam., subglaucous, dull green, brown, or reddish. Racemes green,
spikelets paler to purplish.
Notes. Although Hubbard and many others cited Domin as the author of the ba-
sionym, it should be Bentham. The presence of a question mark does not invalidate
the latter’s publication (Art. 34.2).
Roberty included Mnesithea pubescens Ridl. in his synonymy, but that species has
little to do with the present genus; the correct name for it is Coelorachis mollicoma
(Hance) Bor (Heidweiler & Van der Klaauw, MS).
S. T. Blake distinguished 7. constricta because of its puberulous and constricted
joints. As several intermediate collections were available these distinctions could not
be upheld.
Four collections (S. T. Blake 13604, 13738, Clarkson 3146, Symon 4883 from
Queensland, BRI) possess heteromorphic spikes, with one type similar to those of
T. pubescens and the other to the small form found in 7. major. The anthers are
1.1 mm long. These collections may be of hybrid origin (7. major x T. pubescens),
or represent an infraspecific taxon (but of which species?), or belong to a distinct
species altogether (see Map 2). At present we cannot offer a satisfactory solution.
On the other hand there are mixed collections of both species, which may confuse
those unaware of the heteromorphy of the spikes of 7. major, e.g. Bailey s.n. (K)
from Thursday Isl., which led Hubbard to observe that it might represent ‘probably
a new species related to 7. pubescens.’.
2. Thaumastochloa monilifera Sosef & Koning, sp. nov. — Map 4, fig. Sa
Thaumastochloa sp.: Simon, Techn. Bull. Bot. Br. Dept. Prim. Industr., Brisbane
3 (1978) 13; ibid. 4 (1980) 84, pro Symon 4795S.
Culmi ad 70 cm alti. Spicae homomorphicae, articulis 4-8 inflatis 2-3.5 mm in diam. Glumae inferiores
modice rugulosae. Antherae 0.5-1 mm longae. — Type: Brass 19712 (L, holo; A), Australia, Queensland,
Cook Distr., Wenlock R., 12° 11’ S, 141° 53’ E, 150 m alt., 27 July 1948.
ae b
Fig. 5. Sessile spikelets of Thaumastochloa cont. — a. T. monilifera (Latz 1509); b. T. major (Collins
87) xaos
Culms up to 70 cm high. Nodes 5-9. Sheaths 2.5-4 cm long, 0.33-0.68 times as
long as the internodes, pilose. Blades linear, flat, 4-11 cm by 2-5 mm, pilose.
Peduncle + erect, straight, gradually narrowed towards the base, where it ar-
ticulates, 1-8 cm long, glabrous. Spikes homomorphous, moniliform, 1-1.5 cm by
2-3.5 mm across. Joints 4-8, articulation oblique, 1.7-5 mm long, glabrous.
Spikelets alternatingly in 2 rows, 1.6-3.5 mm long. Lower glume triangular-ovate,
slightly convex, weakly rugulose, 7-nerved. Upper lemma not nerved. Anthers 0.5-1
mm long.
Distribution. Australia: Queensland (Batavia, Bing Bong, Koolburra, Laura, S.
of N. Kennedy R., Wenlock R.).
Ecology. Dry sandy banks of gullies, road-sides, savannah forest with e.g. Acacia
sp., Eucalyptus tetrodonta, Melaleuca sp., Callitris intratropica-heath.
Collector’s notes. Plants semi-prostrate, culms weak, 40-60 cm long.
Note. Named for the bead-like joints of the spike.
3. Thaumastochloa major S. T. Blake — Map 3, fig. Sb
Thaumastochloa major 8. T. Blake, Papers Dept. Biol. Univ. Queensl. 1, 18 (1941)
20; Chippendale, Proc. Linn. Soc. N. S. W. 96 (1972) 223; Simon, Techn. Bull.
Bot. Br. Dept. Prim. Industr., Brisbane 3 (1978) 13; ibid. 4 (1980) 84. — Type:
S. T. Blake 13360 (BRI, holo, n.v.; L), Australia, Queensland, Cook Distr.,
Cairns, 16° 55’ S, 145° 46’ E, + 1.5 m alt., 24 March 1938.
Thaumastochloa pubescens auct. non C. E. Hubb. and Th. rariflora auct. non C.
E. Hubb.: Gardn., Fl. W. Austr. 1, 1 (1952) 309, 310, f. 91 A-E.
154
Map 3. Thaumastochloa major.
Culms up to 35 cm high. Nodes 6-10. Sheaths 1-4 cm long, (0.33-)0.5-> length
of internode, pilose at least along the margins. Blades linear, flat to involute, 2-12
cm by 1.5-5.5 mm, pilose, rarely glabrous. Peduncle erect to recurved, usually
gradually narrowed towards the base, where it articulates, 1.5-18 cm long, glabrous.
Spikes hetero- or homomorphous: some 1.5-3 cm long, rarely with a small secon-
dary branch at base, joints 4-10, relatively broad, 4-5.5 by 2-3.5 mm across, ar-
ticulation straight to oblique, spikelets alternating in 2 very distinct rows, but still
secund, others 0.35-1.6 cm long, joints 1 or 2 (or 3), relatively narrow, 3.5-10 by
1-1.5 mm across, articulation straight, spikelets + in 1 row. Joints cylindric,
glabrous. Spikelets 2.3-4 mm long. Lower glume triangular to ovate-oblong, convex
to flat, rarely concave, smooth, rarely rugulose or pustulate and with some hairs,
9-nerved. Upper lemma 2-nerved. Anthers (1.1-)1.4-1.7 mm long.
Distribution. Malesia: New Guinea (Aru Islands, P. Trangan); Australia: W.
Australia (Camden Sound, Cape Domett, Cockburn Ra., Elcho Isl., Giddy R.,
Glenelg R., Groote Eylandt, Kalumbara, Kimberley Res. St., Oobagooma, Prince
Regent R. Res., NE. of Tableland St.), N. Territory (Batchelor, Borroloola, Coastal
Plains Res. St., Cooinda, Kapalga Ref., Katherine Gorge Nat. P., Maranboy Pol.
St., McMillan’s Rd., Mudginbarry, Mummalary St., Nourlangie Safari Camp, Port
Bradshaw, Twinn Falls, Wessel Isl., Yirrkala), Queensland (Ayr/Bowen Rd.,
Cairns, S. of Cape York, Chillagoe, Coalbrook, S. of Coen, Cooktown, Croydon,
Einasleigh, Endeavour R., Glenore, Jardine R., Kennedy Rd., Mapoon Res., Nor-
manby R., St. George R., Thursday Isl., Wrotham Park).
155
156 Gard. Bull. Sing. 36(1) (1983)
Ecology. Low-lying, damp, sandy places, coastal dunes, open woodland with e.g.
Allosyncarpia ternata, Aristida hygrometrica, Erythrophloeum sp., Eucalyptus
crebra, E. dichromophloia, E. ferruginea, E. latifolia, E. miniata, E. papuana, E.
polycarpa, E. tetrodonta, Melaleuca viridiflora, Perotis rara, Petalostigma banksii,
Plectrachne pungens, Sorghum sp., locally gregarious, up to 600 m alt.
Collector’s notes. Solitary or tufted annual, biennial (!), or short-lived perennial
(!), suberect to diffuse and spreading, bent over as if to bury the rigid tips into the
soil, culms up to 45 cm high, tufts up to 45 cm in diam., rather pale to bright green
to pale or reddish brown, but spikes green, spikelets paler to whitish.
Notes. Most remarkable for its heteromorphic spikes which, if not found on the
same plant, would never have prompted the suggestion that one species was involv-
ed. It is no wonder that Gardner was led astray, thinking that Holmes ex Black
5000.033 (PERTH) was a mixture of 7. pubescens and T. rariflora.
Occasionally a secondary basal branch is present in the spike, but although we
have seen the material from PERTH where Gardner worked, we have not seen this
as well-developed as depicted by him. It is more likely that two spikes were placed
one above the other; the heteromorphy is then evident.
As stated in the introduction, some spikes of Brass 18880 (A, L) have some lower
joints with 2 sessile spikelets and the stipe of a pedicelled one, reminiscent of the
apparently single feature that characterises the genus Mnesithea. One of these
spikelets may be regarded as a very short lateral branch as described above.
The confusion that results when a mixture is made in collecting this species with
another is mentioned under 7. pubescens.
4. Thaumastochloa striata Sosef & Koning, sp. nov. — Map 5, fig. 6a
Thaumastochloa sp.: Simon, Techn. Bull. Dept. Prim. Industr., Brisbane 3 (1978)
13.
Nodi culmi 5-8. Vaginae bene evolutae internodiis 0.5-1-plo breviores. Pedunculi 4-18 cm longi,
minute puberuli (20x!) in parte tertia superiore. Internodia (1 vel) 2 vel 3. Gluma inferior longitudinaliter
inter nervos 7 sulcata. Antherae 1.1-1.7 mm longae. — Type: Lazarides & Adams 10 (L, holo; BRI,
CANB, DNA, E, K, NSW, NT, US, 7.v.), Australia, N. Territory, 25 m N.N.E. of Maranboy Pol. St.,
14° 40’ S, 132° 39’ E, 3 March 1965S.
Culms 15-25 cm high. Nodes 5-8. Sheaths 1.2-2.5 cm long, 0.5-1 imes as long
as the internodes, shortly soft-hairy to nearly glabrous with densely hairy margins.
Blades linear, usually somewhat involute, 2-6 cm by 1.5-3.5 mm across, hairy
behind the ligule and along the margins, otherwise glabrous. Peduncle erect,
straight, narrowed in the lower half towards the base, where it articulates, 4-18 cm
long, upper third with rows of minute hairs (20x!). Spikes homomorphous, (0.45-)
1.1-2 cm by 0.6-1.5 mm across. Joints (1 or) 2 or 3, articulation straight, glabrous.
Spikelets slightly alternating, 3-4 mm long. Lower glume triangular-oblong, slightly
convex, grooved.between the 7 nerves. Upper lemma 2-nerved. Anthers 1.1-1.7 mm
long.
Heteropholis & Thaumastochloa I Si7/
Distribution. Australia: N. Territory (Maranboy Pol. St., Elcho Isl.).
Perri tere noes Coad
Fig. 6. Sessile spikelets of Thaumastochloa cont. — a. T. striata (Lazarides & Adams 10, type); b. T.
rubra (S.T. Blake 17636, type). x 15.
Ecology. Sandy soil in open Eucalyptus tetrodonta forest with Plectrachne
pungens, alt. low.
Collector’s notes. Tufted, erect annual, culms spreading from base, inflorescence
erect.
Note. Named for the conspicuously striate joints and lower glumes.
5. Thaumastochloa rubra Sosef & Koning, sp. nov. — Map 5, fig. 6b
Thaumastochloa striata similis, sed in vaginis internodiis 0.17-0.33-plo brevioribus, pedunculis 0.5-2
cm longis glabris differt. — Type: S. T. Blake 17636 (BRI, holo; L), Australia, N. Territory, 16° 28’ S,
134° 59° E, 198 m alt., 4 May 1947.
Culms up to 45 cm high. Nodes 4-6. Sheaths 0.8-1.6 cm long, 0.17-0.33 time the
length of the internode, pilose. Blades linear, flat to involute, 1.5-5 cm by 1-2 mm,
pilose. Peduncle + erect, + straight, gradually narrowed towards the base, where
it articulates, 0.5-2 cm long, glabrous. Spikes homomorphous, 0.9-2.7 by + 0.7
mm across. Joints (1 or) 2, articulation straight, 4-5 mm long, glabrous. Spikelets
(see note) 3-4 mm long. Lower glume triangular-oblong, flattened to convex,
7-nerved, grooved between the nerves. Upper lemma 2-nerved. Anthers + 1.1 mm
long.
158 Gard. Bull. Sing. 36(1) (1983)
Distribution. Only known from the type.
Ecology. On flagging sandstone ridges with Eucalyptus dichromorphloia, 198 m
alt.
Collector’s notes. Green to reddish, + erect tufts to 45 cm.
Notes. The exact position of the spikelets could not be ascertained, because all
spikes were either 1-jointed or the second joints had dropped off; a unilateral posi-
tion seems most likely. The joints are + reddish-brown, hence the epithet.
6. Thaumastochloa brassii C. E. Hubb. — Map 4, fig. 7a
Thaumastochloa brassii C. E. Hubb. in Hook., Ic. Pl. 34 (1936) t. 3314, p. 3;
Simon, Bull. Bot. Br. Prim. Industr., Brisbane 3 (1978) 13; ibid. 4 (1980) 84 —
Rottboellia corymbosa L.f. subvar. brassii Roberty, Mon. Androp. (1960) 65. —
Type: Brass 370 (K, holo, n.v.; BRI), Australia, Queensland, Burke Distr., Set-
tlement €r.;-18°%S; 138° 77 E, Junesi9237
Culms 7-35 cm high. Nodes 10-15. Sheaths 0.5-1.5 cm long, 0.25-1 times as long
as the internodes, sparsely pilose to subglabrous. Blades linear, flat to involute, 1-3
cm by + 3 mm, sparsely pilose to subglabrous. Peduncle slightly curved, gradually
narrowed towards the base, where it articulates, 0.8-6 cm long, glabrous. Spikelets
erect to erecto-patent, homomorphous, 0.3-0.8 cm by 0.8-1 mm across. Joints
1(-3), articulation straight, 3-4 mm long, glabrous. Spikelets in + 1 row, 2-2.7 mm
long. Lower glume oblong, flat, transversely rugose, 7-nerved. Upper lemma
2-nerved at base. Anthers 0.6-0.7 mm long.
a ‘ote
Fig. 7. Sessile spikelets of Thaumastochloa cont. — a. T. brassii (Craven 4103); b. T. rariflora (Brass
18956). X 15.
Distribution. Australia: N. Territory (Bing Bong, Cooinda, Maria Isl., McArthur
R., Katherine), Queensland (N. of Chillagoe, near Croydon, Dunbar, Esmeralda,
Guthalungra, Musgrave, Settlement Cr., Wenlock R.).
Map 4. _ Triangles: Thaumastochloa brassii; circles: T. monilifera.
Map 5. Squares: Thaumastochloa striata; circles: T. rariflora; triangle: T. rubra.
159
160 Gard. Bull. Sing. 36(1) (1983)
Ecology. Wet sandy soil, Callitris intratropica-heath, open savannah forest with
e.g. Eucalyptus sp., Melaleuca viridiflora, Petalostigma sp., Pheidochloa gracilis,
road-sides, up to 300 m alt.
Collector’s notes. Solitary or tufted, pale green, reddish, brownish, or purplish,
to 35 cm diam. Culms sprawling to erect.
Note. Hubbard described the upper lemma as 1-nerved, we saw it consistently
with 2 short nerves at the base only.
7. Thaumastochloa rariflora (F. M. Bailey) C. E. Hubb. — Map 5, fig. 7b
Thaumastochloa rariflora (F. M. Bailey) C. E. Hubb. in Hook., Ic. Pl. 34 (1936)
t. 3313, p. 4; Chase, J. Arn. Arb. 20 (1939) 314; Reeder, J. Arn. Arb. 29 (1948)
355; Henty, Bot. Bull., Lae 1 (1969) 181; Simon, Techn. Bull. Bot. Br. Dept.
Prim. Industr., Brisbane 3 (1978) 13; ibid. (1980) 84. — Rottboellia rariflora F.
M. Bailey, Queensl. Dept. Agric. Bull. 8 (1893) 86; Rept. Austr. Ass. Adv. Sc.
7 (1898) 446; Compreh. Cat. Queens]. Pl. (1912) 617. — Rottboellia corymbosa
L.f. subvar. rariflora Roberty, Mon. Androp. (1960) 66. — Type: F. M. Bailey
15 (BRI, holo, n.v., see note, K, n.v.), Australia, Queensland, Cook Distr.,
Somerset, 10° 45’ S, 142° 35’ E, June 1897.
Ophiuros pubescens auct. non Dom.: Hitchce., Brittonia 2 (1936) 128.
Culms 10-60 cm high. Nodes (4-)8-12. Sheaths 0.8-2.2 cm long, 0.17-0.33(—more
than 1 in apparently juvenile plants) times as long as the internodes, pilose, rarely
subglabrous. Blades linear, flat to involute, 2-10 cm by 2-4 mm, pilose, rarely sub-
glabrous. Peduncle erect, straight to curved, gradually narrowed towards the base,
usually falling off with the uppermost internode and its leaf, 0.5-3 cm long,
glabrous. Spikes homomorphous, cylindric, 0.35-0.9 cm by 0.7-1 mm across. Joints
1 (or 2, then the lower one usually without a spikelet), articulation + straight, 2.5-3
mm long, glabrous. Spikelets slightly alternating in 1 row, 2-3 mm long. Lower
glume ovate-oblong, somewhat concave, smooth, 9-nerved. Upper lemma
0-3-nerved. Anthers 0.5-0.8 mm long.
Distribution. Malesia: New Guinea (Western Distr.: near Morehead Patrol Post,
Mabaduan, Wuroi); Australia: Queensland (Andoom, Batavia, Cooktown, Locker-
bie, Mareeba, Musgrave, Mt. Mulligan, Portland Roads, Somerset, Thursday Isl.,
Townsville, Vallack point, Weipa).
Ecology. Sandy patches in savannah woodlands, e.g. with Eucalyptus, Melaleuca
cunninghamii, Themeda sp., clearings, sometimes gregarious, up to 1000 m alt.
Collector’s notes. Erect to spreading, tufted or not, sometimes densely mat-
forming, up to 30 cm high, green to rather dull green, culms oblique to erect, red-
dish. A troublesome spear-grass.
Notes. The type, presumed to be in BRI, was not among their material although
all was sent. The specimen in K cited by Hubbard was not lent to the authors; it
Heteropholis & Thaumastochloa 161
may be the holotype.
Brass 18554 (A, L) has glabrous blades and pedicelled spikelets reduced to a
minute scale, otherwise it seems to belong here.
INDEX OF COLLECTORS
Only numbered collections have been included. The ‘H-’ and ‘T-’ -numbers refer to the sequence of
the taxa as employed above. Specimens cited in literature but not seen have been included with their iden-
tifications between brackets when these seemed acceptable, otherwise they have been deleted.
Alcorn 8160: T7, Alston 1046: (H3), Aplin 5136: T3, Astle(?) 1354: H1.
Bailey 15: (17), Balansa 503: H4a, 1776: H4a, 1777: H4a, 1778: H4a, Beeston 15: T3, 77: T3, Benoist
1639: H2, Bequaert 282: H1, Bishop 266: T3, S. T. Blake 8571: T1, 9540-A: T1, 11678: T1, 12459: T3,
12460: T1, 12461: T1, 12612: T6, 13360: T3, 13386: T7, 13387: T1, 13591: T6, 13604: T1x3, 13605: T3,
13689: T1, 13716: T3, 13738: T1x3, 13739: T1, 17474: T6, 17480, p.p.: T6, 17636: TS, 18600: T3, 18624:
T6, 19520: T3, 19637: T6, 21803: T1, 21804: T3, 21812: T1, 21866: T7, 23217: T3, S. T. Blake & Webb
15720: T1, 15767: T7, 15768: T1, Bosser 15241: H2, Brass 370: T6, 1904: (T1), 6014: T7, 6554: T7, 18829:
T3, 18554: T7, 18803: T7, 18880: T3, 18956: T7, 19712: T2, 19713: T1, Bredo 2342: H1, 4155: H1,
Brynaert 648: H1, BS 19978 (McGregor): (H4a), 32781 (Ramos): (H4a), 37450 (Ramos & Edano): (H4a),
39203 (id.): H4a, 85252 (id.): H4a, Burbidge 5712: (T1), 5719: T3, But 51: H4a, Buwalda 5519: T3, 5554:
T3, Byrnes 693: T3.
Chang 2130: (H4a), Chatterjee 551: H3, Chen 87: (H4b), 123: H4a, Chippendale 7737: T3, Chuang
1560: H4a, C. B. Clarke 33807: (H4a), Clarkson 3100: T2, 3146: T1x3, 3146-A: T2, 3176: T3, 3198: T3,
- 3285: T1, Clemens 156: (H4a), 17093: H4a, 18224: H4a, 21234: H4a, Collins 377: T3, Craven 4103: T6,
Cuming 1339: H4a.
Davidse & Sumithraarachchi 7961: H3, 8698: H3, Distr. Off. Atherton 19: T1, Dujardin 342: H1,
Dunlop 2882: T6, 3787: T6.
Edwards 25: H1, Evrard 7027: H1.
Fanshawe 3073: H1, Flecker 477: (17), Flecker Qid. Nat. Club 13189: T6, 13190: T2, Fosberg 55015:
ive
Gardner 9651: T3, Gathy 289: H1, George 12559: T3, Gould & Cooray 13740: H3.
Hance 1248: H4a, Hartley 14632: T3, 14757: T3, Henry 1201: H4a, Holmes ex Black 5000.033: T3,
Homblé 56: H1, Hosokawa 8868: H4a, Hsu 287: H4a, 511: H4b, 1170: H4a, 5300-1: H4a, 12101: H4a,
13555: H4a, Hsu & Kuoh 13658-A: H4a, Hu 10365: H4a.
Jackson 444: (H1), 731: H1, Jacobsen 21: (17).
Kanehira & Hatusima 4425: H4a, 4428: (H4a), 4688: (H4a), Kenneally 5969: T1, Kievits 1543: H4a,
Kneucker 789 (Merrill & Robinson): H4a.
Larsen 8113: (H4a), Latz 1461: T3, 1507: T6, 1509: T2, 3539: T3?, 3736: T3, 6071-A: T3, 6071-B: T4,
7761: T3, Lazarides 3663: T1, 3804: T1, 3929: T6, 4503: T3, 4564: T1, 4746: T6, 4766: T3, 6404: T3,
6823: T3, 7564: T3, 7801: T3, 8850: T6, 8851: T3, 8981: T3, Lazarides & Adams 10: T4, 11: T3, 76: T3,
295: T3, Lin Pin 6226: H4a, Lisowski 208: H1, 539: H1, 540: H1, 1120: H1, 2661: H1, 8892: H1, Lynes
645-A: H1.
Maconochie 2594: T1, Malaisse 17538: H1, McCallum Webster T 204: (H1), McDonald 2628: T1,
McKee 9183: T1, 9505: T7, Melville et al. 3731-C: T1, Merrill 236: H4a, 9875: H4a, Merrill Philip. Pl.
162 Gard. Bull. Sing. 36(1) (1983)
124: (H4a), Milne-Redhead & Taylor 9058: H1, Morain 307: T1, Morton 601: T7, 1707: T3, Murata et
al. T-16580: H4a, 16988: H4a.
NGF 38699 (Henty & Katik): T1, 49393 (Henty & Forman): T7, 49686 (Henty & Carr): T1.
Odashima 129: (H4a), 562: H4a, Oersipuny 18: H4a.
Perry 1974: T1?, Pedley 2637: T3, 2648: T6, 2716: T3, 2745: T2, Peterham 389: T3, Pollock 32: T1,
Pullen 7088: T7, 7173: T7.
Quarré 137: H1.
Rust 33: (T1).
Salesiéns 832: H1, Santos 4055: H4a, 4986: H4a, Schmitz 1009: H1, Schrooten 1091: H1 (wrong label),
Scribner 3: (H4a), Shantz 543: (H1), Shimada 4852: (H4a), Simon & Williamson 1872: H1, L. S. Smith
04364: T1, Specht 93: T3, 190: T3, 683: T3, 731: T3, Squires 165: H4a, Staples 010572/12: T1, Suzuki
20066: H4a, Symoens 3341: H1, 8267: H1, 12237: H1, 12285: H1, Symon 4785: T2, 4845: T3, 4883:
M1x354926:5655022- Tie 76le nS:
Thwaites CP 867: H3, Trapnell 1522: H1, 1706: (H1).
Vanoverbergh 2802: H4a, Vesey-Fitzgerald 1628: H1, 2975: H1.
Wang 12768: H4a, C. T. White 8674: T1, Wiehe N. 182: (H1).
Yamamoto et al. 11: H4a, Yamamoto & Suzuki 579: H4a.
Zerny 479: H1.
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IE IYER IE EERE EE EEE EEE}
THE GARDENS’ BULLETIN
eZ
a=
‘a SINGAPORE -
y
; VOL. 36 (Part 2) lst December 1983
i) | CONTENTS aX
A
4 i
PAGES >
JARZEN, DAVID M.: \
Meese Polliensmecord of the Pandanaceaé .........52... 008s cesses ee eeeeetens 163-175
NG, F. S. P. AND M. JACOBS: :
A Guide to King’s ‘‘Materials for a Flora of the Malayan Peninsula” ................ 177-185 >
KOCHUMMEN, K. M.:
_ Notes on the Systematy of Malayan Phanerogams. XXX. Anacardiaceae ............. 187-196
KURATA, SHIGEO:
A new Species of Nepenthes from Sulawesi, Indonesia ..................00.0-00eeees 197-200
WONG, KHOON MENG AND AH LAN LIM: ()
On the Nature of leaf-opposed Inflorescences in Aidia cochinchinensis ..............- 201-204 X
STONE, BENJAMIN C.:
Some new and critical Pandanus Species. I. ()
EMMI SLOMRCVIRIONEANGANACCATUIN 5 <<<: 210s 022 een an ce eee eee nese eeeeeseess 205-212 ‘
LATIFF, A.: ()
q Studies in Malesian Vitaceae. VII. The genus Tetrastigma \
gl ERTL OM 8 ie i et 213-228 e
" X
-MAXWELL, J. F:: ‘)
New and interesting Plant Records for Singapore, II ...........-2-.eeeeeeeccceeeees 229-232 \
~ GRUEZO, WILLIAM SM.: ¢
_ Lobaria clemensiae Vain. (Lobariaceae, Lichenes) \
MTC HITE ESI ATCT SIMGGHESIR Vile ction sc nce sonic cle ee cle cote a ae nicce ene ewcedccecess 233-236 ()
j yy
BHATTACHARIEE, S. K.:
Influence of growth-regulating Chemicals on Hippeastrum hybridum hort. ..........-- 237-242
Published by the Botanic Gardens
Parks and Recreation Department
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THE GARDENS’ BULLETIN
SINGAPORE
VOL. 36 (Part 2)
lst December 1983
CONTENTS
JARZEN, DAVID M.:
iunesbossileollen Record of the Pandanaceae: 22 ..02 0502 sce. oe ene see cs
NG; F.S. P. AND-M. JACOBS:
A Guide to King’s ‘‘Materials for a Flora of the Malayan Peninsula”’ ...
KOCHUMMEN, K. M.:
Notes on the Systematy of Malayan Phanerogams. XXX. Anacardiaceae
KURATA, SHIGEO:
A new Species of Nepenthes from Sulawesi, Indonesia .................
WONG, KHOON MENG AND AH LAN LIM:
On the Nature of leaf-opposed Inflorescences in Aidia cochinchinensis_ ..
STONE, BENJAMIN C.:
Some new and critical Pandanus Species. |.
SepmementtanRevisio PandanacearuMm Aca <sccuccee seve soe e ds eelda eee
RATIFF, A.:
Studies in Malesian Vitaceae. VII. The genus Tetrastigma
MMHCRVIAIAVRRCHINSU LAs nt torrerte rte eels crates Sock MascnieG singe sae ee
MAXWELL, J. F.:
New and interesting Plant Records for Singapore, II ................-.
GRUEZO, WILLIAM SM.:
Lobaria clemensiae Vain. (Lobariaceae, Lichenes)
LEAMIIA CH As SIANG CINIGONESIAn. 5 35 cc ele cs: 6s ele die he nae nprutatie n EE
BHATTACHARJEE, S. K.:
Influence of growth-regulating Chemicals on Hippeastrum hybridum hort.
Published by the Botanic Gardens
Parks and Recreation Department
Ministry of National Development
Cluny Road, Singapore 1025.
Printed by Amsterdam Type Printers, Singapore
PAGES
163-175
177-185
187-196
197-200
201-204
229-232
233-236
INSTRUCTIONS TO AUTHORS
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to the surname of the (first) author and in the following form:
Singh, H. (1967). Sclereids in Fagraea. Gard. Bull. Sing. 22, 193-212.
Abbreviations of titles of journals should be those of the World List of Scientific Periodicals
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Ridley, H. N. (1930). The Dispersal of Plants Throughout the World, L. Reeve; Ashford,
Kent; 242-255.
For literature citations in taxonomic papers the following style is required:
Medinilla alternifolia B\l., Mus. Bot. Lugd.-Bat. I:2 (1849) 19.
Sterculia acuminatissima Merr., Philip. J. Sci. 21 (1922) $24.
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The Fossil Pollen Record of the Pandanaceae
DAVID M. JARZEN
Paleobiology Division
National Museum of Natural Sciences
National Museums of Canada
Ottawa, Canada
Abstract
The fossil record of pollen comparable to the family Pandanaceae and sometimes directly comparable
with the extant genus Pandanus extends back to the latest Upper Cretaceous. The family which once had
a wide geographic distribution on all continents except Australia, has, since the mid-Tertiary, become
restricted to the Old World tropics and subtropics.
Introduction
The monocotyledon genus Pandanus Rumph. ex L. comprises about 600 species
of trees, shrubs and less frequently subshrubs. Several, such as P. epiphyticus
Martelli and P. alticola Holt. and St. John from Malaya and Borneo are truly
epiphytic though facultatively so, also occurring on boulders of limestone or sand-
stone (Stone, 1978). The tree habit is the most common, and such pandans form a
conspicuous part of the vegetation of many tropical shorelines. The pandans (or
screw-pines) are distributed throughout the palaeotropics, with species occurring on
nearly all tropical and marginally subtropical islands of the Pacific, the northern
tropical regions of Australia, tropical Southeast Asia, Indonesia, the Philippines,
southern India and islands of the Indian Ocean, the Malagasy Republic and East
and West Africa (Stone, 1976). The genus does not occur naturally in the neotropics.
Map | illustrates the overall distribution of the genus, as well as the known fossil
pollen reports of Pandanaceae.
The pandans are dioecious, with the staminate plants, particularly of the forest
species, being less frequently collected than the pistillate plants, due to the brief,
ephemeral staminate anthesis. The leaves, usually long, linear, spirally arranged and
congested along branch tips, the characteristic stout aerial prop roots, and the often
(but not always) large pendulous female cephalia (syncarps) allow recognition of the
genus even at some distance.
The pulpy fruits (drupes or polydrupes) are often a bright-coloured red, orange
or yellow and are dispersed not only by birds and fruitbats, but within several
sections of the genus by ocean or freshwater currents (Stone, 1976). The fruits of
Pandanus helicopus Kurz which occur today in Bangka Island, Malaya, Borneo and
Sumatra, are eaten and thus dispersed by fish and turtles (Ridley, 1930; Stone,
1976). Turtles no doubt also perform a dispersal role for Pandanus aquaticus F.
Muell. in Australia (Stone pers. comm., 1978). The small, and very light fruits of
Pandanus basedowii C.H. Wright, native in the Northern Territory of Australia
may be distributed by wind (Stone, 1974).
163
164 Gard. Bull. Sing. 36(2) (1983)
Pollen Morphology
The pollen of Pandanus (29 species examined) is free, radiosymmetrical, spherical
to ovoid (often irregular) monoporate (rarely pseudocolpate). The ornamentation is
typically echinate but some species are psilate, finely granulate or finely rugulate
(e.g. granulate in Pandanus dorystigma Mart.; rugulate in P. matthewsii Merr.;
psilate in P. jiulianettii Mart. and P. elostigma Mart.). The echinae are conical,
blunt tipped or capitate, and range in length from <1.0 » to 4-5 pw. They are more
or less evenly distributed over the surface of the grain. The pore (often difficult to
observe) is circular and situated at one end of the long axis, with either an entire
or diffuse margin, annulate or non-annulate, diameter 2-4 yw (rarely larger). The
exine is two-layered. In some species the sexine is thicker than the nexine, while in
other species the opposite is true. Wall thickness ranges from <1.0 » to ca. 1.5 p.
Dimensions for 29 species examined are 16 (22.2) 37 w x 13 (17.1) 25 un.
Studies on the pollen morphology of Pandanaceae include: Erdtman (1972),
Huang (1970), Huynh (1980), Selling (1947), Sharma (1968), and Sowunmi (1974).
Comparisons with other extant Pollen Forms
Several angiosperm families have genera with certain pollen-morphological
similarities to Pandanus. Sharma (1968) placed significance on the monocolpate
condition of P. odoratissimus L.f. as providing evidence of palynological affinity
of Pandanus with Hypoxis (Hypoxidaceae) and Smilax (Liliaceae). Hypoxis pollen
(12 species examined) is monosulcate, often foveolate, has a rather thick exine and
the grains measure about 50 » in diameter. Smilax is non-aperturate, or monosulcate,
but never monoporate; the exine is tenuiexinous (nexine thinner than sexine). Pan-
danus pollen may be quite easily differentiated from either of these two genera.
Pandanus pollen is much smaller than that of Hypoxis, and is never foveolate; the
presence of a pore (if discernable) will exclude Smilax, and in spinate forms of
Pandanus the nexine is thicker than the sexine.
In the family Araceae, the genus Remusatia has pollen which superficially
resembles that of Pandanus, but upon closer examination can be easily distinguished.
Although the sculpturing elements in Remusatia are echinate, interestingly the
spines are of two distinct sizes and shapes. The larger spines, as wide as they are
high are interspersed with much smaller spines somewhat longer than wide. This
distinctive feature, not observed in Pandanus, as well as the non-aperturate condition
of Remusatia allows easy recognition of its pollen.
The generally larger diameter and extremely small scrobiculi (very small, more or
less circular pits) differentiate Joinvillea (Flagellariaceae) pollen from that of
Pandanus.
Although a number of palm genera (e.g. Arenga, Manicaria, Maurita, Socratia
et al.) have spherical, echinate pollen, none are monoporate.
As far as is known, only three dicotyledon genera have pollen which might be
confused with that of Pandanus.
Fossil Pollen Record, Pandanaceae 165
The pollen of Peumus (monospecific genus of the Monimiaceae) much re-
sembles that of Pandanus, but the grains are inaperturate, and the bases of the
spines do not appear sunken below the surface, a feature common to Pandanus
pollen. The size of Peumus pollen ranges 38-41 », somewhat larger than that of
Pandanus.
Hernandia (Hernandiaceae) pollen although echinate can not be confused with
that of Pandanus, because of its larger size (generally up to 90 »), the equidimen-
sional spines, and granulate surface between spines.
Stachyanthus (Icacinaceae) has inaperturate, spinose pollen. The characteristic,
long (up to 4 »), recurved spines however allow recognition of this pollen as distinct
from Pandanus.
The Fossil Occurrences
Forty-five references to fossil pandanaceous pollen are to be found in the published
literature table 1). Thirty-five of these are Tertiary or younger occurrence, eight of
which occur within the present distribution of the genus (the paper by Leopold,
1969, contains four localities, thus there are 11 fossil occurrences shown on map 1
within the present range of Pandanus). The combined stratigraphic ranges of fossil
Pandanaceae pollen from major geographic regions of the world are illustrated in
graph 1.
The stratigraphically oldest reports of pollen referable to the family of Pan-
danaceae, are from the Maestrichtian of the western interior of North America.
Oltz (1969) reported its occurrence from the Hell Creek and Tullock Formations
(Maestrichtian-Palaeocene) of central Montana; Norton and Hall (1969) described
Spinamonoporites typicus Norton from the Maestrichtian and Palaeocene of the
type Hell Creek Formation, Montana, which they compared with question to Pan-
danus, and Leffingwell (1971) recovered Pandaniidites radicus Leffingwell from the
Lance and Fort Union Formations (Maestrichtian-Palaeocene) of Wyoming.
Within the Canadian western interior, pollen referable to the family Pan-
danaceae occurs in the Palaeocene of Alberta (Snead, 1969) and in the Maestrichtian
of the Morgan Creek badlands of southern Saskatchewan (Jarzen, 1978). This latter
occurrence is accepted as the oldest record of Pandanus pollen (Muller, 1981).
Archangelsky (1973; 1976) described but did not name a species of Pandaniidites
from the Salamanca Formation (Palaeocene) in the Chubut Province of Argentina,
which he compared directly to the genus Pandanus. The reports of Archangelsky are
the only known occurrences of fossil Pandanus pollen from South America. Reports
of pollen referable to Pandanus from European sediments are those of Raukopf
(1959), Kedves (1963), Kedves and Endredi (1963; 1965), Martin (1976), Zaklinskaya
(1967) and Simpson (1961) who described ?Pandanus shiabensis from the inter-
basaltic lignites of Shiaba, Island of Mull, Scotland. Simpson had earlier (Simpson,
1936) assigned this pollen form to the extant genus Smilax but after a re-
examination, felt it more closely resembled Pandanus.
166 Gard. Bull. Sing. 36(2) (1983)
Table 1. Published reports of fossil pollen referable/comparable with the family Pandanaceae. (numbers
in parenthesis refer to numbers on map 1).
AUTHOR(S) LOCALITY AGE
Zaklinskaya, 1978a (1) $262 DSPP, Timor Pliocene-Pleistocene
Shatilova, 1964, 1967 (2) Georgian, USSR Pliocene-Pleistocene
Anderson and Muller, 1975 (3) Northwest, Borneo Holocene and Miocene
Mchedlishvili 1963 (4) USSR Upper-lower Pliocene
Simpson, 1961 (5) Shiba, Mull Scotland Miocene or lower Pliocene
Leopold, 1969 (6) Eniwetok & Bikini Atolls, Miocene
Guam & Palau Islands
Muller, 1964 (7) Northwest Borneo Upper Miocene
Rao & Ramanujam, 1975, 1976 (8) Kerala, India Lower to Middle Miocene
Sah, 1967 (9) Burundi Upper Neogene
Raukopf, 1959 (10) Eastern Germany Oligocene-Miocene
Li, ef al., 1967 (11) Trans-Urals, USSR Oligocene, Miocene
Sedova, 1956 (12) Primorsk Kray, USSR Oligocene
Tarakanova, 1965 (13) Siberia, USSR Eocene-Oligocene
Aristova, 1973 (14) Siberia, USSR Eocene-Oligocene
Potonié, 1960 (15) Kalewa, Burma Eocene
Fisk, 1976 (16) Wyoming, USA Eocene
Tschudy, 1973a (17) Mississippi embayment, USA Eocene
Love, et al., 1978 (18) Wyoming, USA Middle Eocene
Zaklinskaya & Prokof’yev, 1971 (19) Guinea Lower to Middle Eocene
Kedves, 1963 (20) Hungary Lower Eocene
Kedves and Endredi, 1963, 1965 (21) Hungary Lower Eocene
Elsik, 1967 (22) Texas, USA Lower Eocene
Martin, 1976 (23) England Palaeocene-Eocene
Bebout & Traverse, 1978 (24) North Dakota, USA Palaeocene-Eocene
Fairchild & Elsik, 1969 (25) Gulf Coast, USA Palaeocene-Eocene
Jarzen, 1982 (26) Saskatchewan, Canada Palaeocene
Archangelsky, 1973, 1976 (27) Chubut, Argentina Palaeocene
Petriella & Archangelsky, 1975 (28) Chubut, Argentina Palaeocene
Elsik, 1968 (29) Texas, USA Palaeocene
Snead, 1969 (30) Alberta, Canada Palaeocene
Zaklinskaya, 1967 (31) Europe Palaeocene
Zaklinskaya, 1978b (32) Cape Verde Rise Cretaceous-Palaeogene
Sedova, 1960 (33) Far East, USSR Upper Cretaceous-Danian
Khlonova, 1971 (34) USSR Maestrichtian-Danian
Oltz, 1969, 1971 (35) Montana, USA Maestrichtian-Palaeocene
Norton & Hall, 1969 (36) Montana, USA Maestrichtian-Palaeocene
Leffingwell, 1971 (37) Wyoming, USA Maestrichtian-Palaeocene
Tschudy, 1973b (38) New Mexico, USA Maestrichtian-Palaeocene
Leffingwell, ef a/., 1970 (39) Wyoming, USA Maestrichtian
Jarzen, 1978 (40) Saskatchewan, Canada Maestrichtian
The first report of fossil pollen comparable to Pandanus (or inferred to be Pan-
danus by reference to the family Pandanaceae since the other members of the family,
Freycinetia and Sararanga, are not usually echinate except two Freycinetia species
which have echinate pollen) is that of Sedova (1956) who designated Pandanus sp.
from the Oligocene of Primorsk Kray, U.S.S.R.
The geologically most recent occurrences of fossil Pandanaceae pollen are of
Pliocene-Pleistocene age deposits of the U.S.S.R. (Shatilova, 1964; 1967) and from
al
31qe], Ul SJOQUINU 0} JOJaI SIOQUINAY *(S}OP YOR]Q) oRODBURpUR A]IWIRY dy) O} 9[QeJOJOI UD|IOd [ISSOJ
Jo syodas poystjqnd umouy ‘(pojddns) sevsovuepued oy) Jo UONNQSIp s1ydes1d093 Aep-juasoig “| dey
167
168 Gard. Bull. Sing. 36(2) (1983)
core material of Site 262 drilled by the Glomar Challenger in the Timor Trough near
the Island of Timor (Zaklinskaya, 1978a). Zaklinskaya refers only to the family
Pandanaceae, and does not illustrate the pollen thus assigned. It is probable that the
pollen recovered by Zaklinskaya represents Pandanus inasmuch as the other two
genera within the family usually have psilate exines and would be rather difficult to
assign to the family Pandanaceae.
Of the numerous reports of pandanaceous megafossils, nearly all have subse-
quently been determined to be Nypa (Tralau, 1964), and in fact Stone (1976)
considers the report of Pandaniites rhenanus Krausel and Weyland (1950) from the
Oligocene of Germany as a true pandan. The megafossil record of Pandanaceae is
thus meager and provides little information on the geologic history of the family.
Discussion
The fossil pollen grains illustrated in plate 1, figures 1-5, were recovered from the
uppermost Cretaceous (Maestrichtian) Frenchman Formation of southwestern
Saskatchewan, Canada (Jarzen, 1978). These grains were selected to illustrate
several features which are diagnostic of extant Pandanaceae pollen. Figure | illustrates
the general shape (ovoid), circular pore with entire margin and the conical isometric
echinae (spines) typical of many species of Pandanus. These features are comparable
with those of P. /ongicaudatus Holtt. and St. John as illustrated in figure 11.
The fossil grain in figure 2 was photographed in optical section to illustrate the
nature of the spine bases which are depressed into the interior of the pollen wall (see
also Erdtman, 1972, p. 308, fig. 179A). This feature of ‘‘sunken bases’’ is clearly
seen in the phase contrast photomicrograph (fig. 7) of Pandanus tectorius Sol. and
as faint protuberances in the electron micrograph (fig. 13) of P. vandra St. John.*
Figure 3 also illustrates the ‘‘sunken bases’’ of the spines but additionally shows the
less frequent annulate pore condition of some of the fossil grains. P. odoratissimus
L.f. (fig. 8-9) also displays an annulate pore. Pandaniidites radicus Leffingwell
(1971), recovered from the Maestrichtian of Wyoming, and P. texus Elsik (1968) are
clearly annulate.
The fossil grain type illustrated in figure 4 is rarely encountered in Saskatchewan
sediments. The unique elongated, clavate spines were initially considered unlike the
usual conical spines of Pandanus. However, as illustrated in figure 12 the pollen of
P. mozambicus St. John bears identical spines. Figure 5 and figure 10 illustrate the
rather rare monocolpate condition in the fossil specimens (fig. 5) and extant Pan-
danus tectorius Sol. (fig. 10). Figure 14 illustrates spine shape in modern Pandanus,
with a bulbous enlarged base. This feature may or may not occur at random on
certain fossil specimens. Some of the spines on the fossil grain in figure 2 approach
this condition.
There is little doubt as to the validity of reports of fossil pollen referable to the
family Pandanaceae from the Tertiary period (see map 1) and especially so for
* P. vandra St. John is regarded by Stone (pers. comm., 1983) as an undoubted synonym of P. tectorius Sol.
Fossil Pollen Record, Pandanaceae 169
those occurrences within the present range of Pandanus. Less confidence is held for
the uppermost Cretaceous occurrences, since all of these reports occur outside the
present geographical range of the family.
Stone (1976) and Muller (1981), accept palynological reports of Maestrichtian
Pandanaceae pollen. From the descriptions and illustrations of the six published
reports of ‘‘pandanaceous’’ pollen from the Maestrichtian/Palaeocene interval of
the western interior of North America it appears that at least two species of the
family Pandanaceae once occurred in North America 65-63 million years ago. Since
the Cretaceous, continental movements, withdrawal of the eperic seaways, mountain
building, and changes in climate have dramatically reduced suitable conditions for
the survival of the Pandanaceae in North America. Today no wild plants of the
family occur in North or South America, Europe, or temperate Asia where mega-
and micro- fossils have been recovered (Stone, 1976). Speculation as to the origin
of the family Pandanaceae should accept as baseline data the reports of fossil Pan-
danaceae from North and South America, which must be considered valid.
Jarzen (1978) has suggested a modern analogous environment for the fossil flora
recovered from the Maestrichtian of the Morgan Creek badlands of Saskatchewan
as being in the Southeast Asia-Indomalaysian area based on the occurrence of several
= me GEOGRAPHIC DISTRIBUTION
Te ea ©
ti Ja! NORTH | SOUTH INDONESIA
2 100) america [AMERICA [EURASIA | AFRICA | cic
TERTIARY
fess eo [ ee
CRETACEOU
Graph 1. Generalized stratigraphic and geographic distribution of reports of fossil Pandanaceae pollen.
170 Gard. Bull. Sing. 36(2) (1983)
diagnostic pollen taxa, including that of Pandanaceae. Other angiosperm pollen
recovered from the Morgan Creek sediments could be compared with extant families
including the Juglandaceae, Myrtaceae, Buxaceae, Cercidiphyllaceae, Gunneraceae,
and all found together today only in Southeast Asia-Indomalaysia.
Acknowledgements
Appreciation is expressed to Mr. Gregory Whalen who processed the samples and
assisted in photography.
Pollen material of Pandanaceae was collected from herbarium specimens at United
States National Herbarium and Herbarium, Missouri Botanical Garden. The assistance
and courtesy of the staff of these herbaria is gratefully acknowledged. Additional
fresh pollen of Pandanus and Freycinetia was received through the courtesies of
Dr. B.C. Stone (Kuala Lumpur, Malaysia), and Dr. K.-L. Huynh, (Institut de
Botanique de |’Université, Neuchatel, Suisse).
Thought provoking and useful discussion concerning the occurrence of Pan-
danaceae in the North American Cretaceous was provided by Drs. Arthur Cronquist,
Harry Leffingwell, Gordon McPherson and Robert Tschudy.
Drs. A.K. Graham, P.H. Stauffer and B.C. Stone kindly read the manuscript and
provided many useful suggestions and criticisms for its improvement.
Special appreciation is expressed to Dr. Benjamin Stone, without whose help and
encouragement this project would not have been completed.
opposite page
Plate 1. Morphological variation in extant and fossil Pandanaceae pollen. Fig. /. Fossil grain illustrating
the ‘‘normal’’ conical shaped spines (11/03 —86.3 x 36.9). Fig. 2. Fossil grain showing, in
optical section, the bases of spines sunken into the interior of the grain (11/07—81.3 x 37.3).
Fig. 3. Fossil grain with annulate pore (07/04 —95.4 x 36.8). Fig. 4. Fossil grain with club-like
ornamentation (11/04 — 89.0 x 35.6). Fig. 5. Fossil grain in colpate condition (11/03 — 82.9 x 35.5).
Fig. 6. Pandanus tectorius Sol., bearing the ‘‘normal’’ conical spines (10513 — 88.2 x 41.6). Fig. 7.
P. tectorius Sol., photographed in phase contrast to illustrate sunken bases of spines into interior
of grain (10513—88.2 x 41.6). Fig. 8. P. odoratissimus L.f., with annulate pore as observed
in optical section (228—93.1 x 40.6). Fig. 9. P. odoratissimus L.f. annulate pore observed in
surface view (228 — 93.2 x 39.9). Fig. 10. P. tectorius Sol., in colpate condition (227 —95.3 x 31.5).
Fig. 11. P. longicaudatus Holtt. & St. John, illustrating the conical shaped spines (7371-SEM
012/7). Fig. 12. P. mozambicus St. John, with club-like ornamentation similar to fossil grain
in Fig. 4. (870-SEM 003/10). Fig. /3. P. vandra St John, split open to show sunken bases of
spines into interior of grain (872-SEM 001/1). Fig. /4. P. furcatus Roxb. detail of surface showing
spines with enlarged bases (7369-SEM 011/7). Figs. /-/0: X 1,000; Figs. //, 12: X 2,500; Fig.
13: X 2,200; Fig. 14: X 3,400.
Plate 1
Morphological Variation in Extant
and Fossil Pandanaceae Pollen
171
172 Gard. Bull. Sing. 36(2) (1983)
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Erdtman, G. (1972). Pollen Morphology and Plant Taxonomy - Angiosperms.
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Fairchild, W.W. and W.C. Elsik (1969). Characteristic palynomorphs of the Lower
Tertiary in the Gulf Coast. Palaeontographica 128B: 81-89.
Fisk, L.H. (1976). Palynology and Palaeoecology of the Eocene and fossil forests
of Yellowstone National Park, Wyoming, USA. (Abstract). JV Internat. Palyn.
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Huang, T.-C. (1970). Pollen grains of Formosan plants (6). Taiwania 15(1): 73-179.
Huynh, Kim-Lang (1980). La morphologie du pollen de Pandanus subg. Vinsonia
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Jarzen, D.M. (1978). Some Maestrichtian palynomorphs and their phytogeographical
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——— . (1982). Angiosperm pollen from the Ravenscrag Formation
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Kedves, M. (1963). Contribution a la flore Eocene inferieure de la Hongrie sur la
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—— . and L. Endredi (1963). Investigation of the bitumen and humic acid
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Fossil Pollen Record, Pandanaceae 173
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Khlonova, A.F. (1971). Palynological characteristics of Cretaceous deposits of
Siberia and the Far East. Akademiya Nauk S.S.S.R. Sibirskoe Otdelenie. Institut.
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Leffingwell, H.A. (1971). Palynology of the Lance (Late Cretaceous) and Fort
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A Guide To King’s ‘‘Materials for a Flora of The
Malayan Peninsula”’
Pass P2 NG"'& TM. JACOBS?
‘Forest Research Institute, Kepong, Malaysia
*Rijksherbarium Leiden, Holland
Abstract
The first publication on the flora of Malaya was Sir George King’s Materials for a Flora of the
Malayan Peninsula. This appeared in 26 instalments over a period of 47 years (1889-1936). The manner
of publication was somewhat complicated. This paper explains the various complications and gives the
dates of issue of the various instalments.
Introduction
Although the best known Flora of the Malay Peninsula is that published by
H.N. Ridley in five volumes between 1922 and 1925, it is a matter of general
knowledge among Malesian botanists that much of Ridley’s work was based on an
earlier work done at Calcutta by Sir George King and his various associates, entitled
Materials for a Flora of the Malayan Peninsula, published in 25 instalments between
1889 and 1915, and with a 26th instalment in 1936. The geographical area of the
Materials covers the Malay Peninsula, and the Andaman and Nicobar Islands. In
terms of quality, the Materials is superior to Ridley’s Flora because whereas the
former was based on original and careful study, the latter appears, in many places,
to have been merely a hastily and inaccurately rewritten copy of the former work.
Unfortunately, the Materials can be quite complicated for a beginner to use,
because, instead of being published in a few self-contained volumes with tables of
contents, regular pagination and indices, the 26 instalments were published as
separate papers in a periodical, the Journal of the Asiatic Society of Bengal, over
the rather long period of 47 years. In fact it was never completed. The Materials in
the Journal of the Asiatic Society of Bengal deal only with Dicotyledons and even
so, the Urticales viz. Cannabinaceae, Moraceae, Ulmaceae, Urticaceae, and most of
the Euphorbiaceae never appeared in print. As for Ficus, this is hardly a problem
since King gave an extensive account of the genus in the Annals of the Royal Botanic
Garden Calcutta Vol. 1, (1887-1888).
The Monocotyledons were all written up by H.N. Ridley and published in-
dependently in 3 volumes (called Parts) under the title Materials for a Flora of the
Malayan Peninsula, Monocotyledons in 1907-1908, at Singapore.
King apparently realised that the interrupted manner of publication in the Journal
would make the Materials difficult to use. He tried to overcome this by having extra
copies of the first 21 instalments printed and assembled into 4 volumes wiz. Vol. 1
Thalamiflorae, Vol. 2 Disciflorae, Vol. 3 Calyciflorae, Vol. 4 Gamopetalae, each
volume with continuous pagination from 1 onwards, except Vol. 2 which continued
ig)
178 Gard. Bull. Sing. 36(2) (1983)
the pagination from Vol. 1. Instalments 22-26 were published in sequence hence the
problem of repagination did not arise when they were assembled into Vol. 5, In-
completae. Botanists who use the Materials will almost certainly come across the
5-volume ‘“‘reprint’’, found in most botanical libraries, rather than the original
Journal which seems to be quite scarce.
However there are a number of complications in the use of the ‘‘reprinted’’
volumes. Firstly, each page of volumes 1-4 bears two page-numbers. The number
series on the top of the page is the original Journal pagination, and is not continuous.
The number series at the bottom is continuous. King had an index compiled for each
volume, using the continuous pagination, but for purposes of citation, it is necessary
to use the original Journal pagination.
Another complication is that tables of contents were not provided. This would not
be a problem to anyone familiar with the system of Bentham and Hooker, which
King tried to follow except for the fact that King was obliged now and again to
postpone the treatment of difficult families. Consequently the family sequence is
disrupted in several places.
A minor complication also arose when the Journal of the Asiatic Society fell
behind schedule in publication e.g., when a number officially dated 1903 actually
appeared in 1904. In such cases, both the official and actual dates of publication
were given on the original Journal covers, to which it is necessary to refer whenever
there is a need to establish priority. King himself cited dates of ‘‘publication’’ in
short prefaces to the first four ‘‘reprinted’’ volumes but these dates sometimes refer
to the dates on which the instalments were read at meetings of the Asiatic Society
rather than to the dates on which they were actually published in printed form.
In the guide below, we have listed the instalments, families, dates of publication,
and other details that may help users of the Materials to find their way. The dates
of publication were obtained from the covers of an unbound set of numbers of the
original Journal in the Singapore Botanic Gardens. For ordinary purposes of
citation, we would recommend that the official year of publication be used.
Instalments 1-21 appeared in Series 2 (called ‘‘Part 2’’) of the Journal, (there were
3 parallel series) and we would recommend that the series be cited before the
volume, for instance:
Diospyros wrayi King & Gamble, J. Asiat. Soc. Bengal II, 74 extra no. (1905)
209.
Capparis cucurbitina King, J. Asiat. Soc. Bengal II, 58 (1889) 395.
Instalments 22-26 appeared in Volume 75 of the Journal without any series number.
Ridley’s account of the Monocotyledons appeared independently and should be
cited as a book, e.g.,
Daemonorops calospatha Ridley. Mat. Fl. Mal. Pen. Monocots 2 (1907) 179.
Volume 1. Thalamiflorae
All families by G. King
(Column A = instalment; Column B = journal pages; Column C = renumbered
pages; Column D = date of publication.)
Family Ser. Vol. No. Year B (6 D
1. Ranunculaceae Il 58 4 1889 359-361 3-5 30.XI11.1889
2. Dilleniaceae 361-368 5-12
3. Magnoliaceae 368-376 12-20
4. (see Instalment 4)
5. Menispermaceae 377-388 21-32
6. Nymphaeaceae 388-391 32-35
7. Capparideae 391-399 35-43
8. Violaceae 399-408 43-52
9. Bixineae II 59 2 1890 113-128 53-68 16.1X.1890
10. Pittosporeae 128-129 68-69
11. Polygaleae 129-144 69-84
12. Portulaceae 144-145 84-85
13. Hypericineae 145-147 85-87
14. Guttiferae 147-185 87-125
1S. Ternstroemiaceae 185-206 125-146
16. (see Instalment 5)
17. Malvaceae Il 60 1 1891 38-57 147-166 19.V.1891
18. Sterculiaceae 57-95 166-204
19. Tiliaceae 95-140 204-249
4. Anonaceae Il 61 1 1892 1-130 250-379 13.VI.1892
16. Dipterocarpeae Il 62 2 1893 87-137 379-429 12. VIII.1893
Volume 2. Disciflorae
All families by G. King
Family Ser. Vol. No. Year B te: D
20. Lineae II 62 4 1893 189-192 431-434 6.111.1894
21. Malpighiaceae 192-197 434-439
22. Geraniaceae 197-204 439-446
23. Rutaceae 205-226 447-468
24. Simarubeae 226-230 468-472
25. Ochnaceae 231-235 473-477
26. Burseraceae 235-262 477-504
27. Meliaceae Il 64 1 1895 16-90 504-578 15.1V.1895
28. Chailletiaceae 90-94 578-582
(also Instalment 8)
29. Olacineae 94-133 582-621
30. Ilicineae 133-137 621-625
179
Volume 2. Disciflorae Continued
A Family Ser. Vol. No. Year B Cc D
8 31. Celastrineae Il 65 3 1896 339-370 625-656 1.X.1896
32. Rhamneae 370-384 656-670
33. Ampelideae 384-419 670-705
34. Sapindaceae 419-453 705-737
35. Sabiaceae 453-459 739-745
36. Anacardiaceae 459-514 745-800
Addition to Chailletiaceae 514-516 800-802
Volume 3. Calyciflorae
All families by G. King except Leguminosae by D. Prain and Sonerila
(Melastomaceae) by O. Stapf. Droseraceae was independently described twice, in
Instalments 9 and 13, without explanation, and the second account made no
reference to the first.
A Family Ser. Vol. No. Year B (G:
9 37. Connaraceae Il 66 1 1897 1-21 1-21 8.VI.1897
38. Leguminosae 21-275 21-275
39. Rosaceae 275-297 275-297
40. Saxifragaceae 297-305 297-305
41. Droseraceae 305-306 305-306
(also Instalment 13)
42. Hamamelideae 306-309 306-309
43. Halorageae 309-311 309-311
44. Rhizophoreae 311-326 311-326
45. Combretaceae 326-345 326-345
46. (see Instalment 12)
47. (see Instalment 11)
10 48. Lythraceae WE af 1 1898 1-11 345-355 5§.VIII.1898
49. Onagraceae 11-14 355-358
50. Samydaceae 14-24 358-368
51. (see Instalment 13)
52. Cucurbitaceae 24-42 368-386
53. (see Instalment 13)
54. (see Instalment 13)
55. (see Instalment 13)
56. Araliaceae 42-63 386-407
11 47. Melastomaceae Il 69 1 1900 1-87 409-495 9. VIII.1900
12 46. Myrtaceae II 70 1 1901 66-142 496-572 14.VIII.1901
13 Datiscaceae NC 7A 1 1902 46-47 574-575 8.V.1902
(wrongly numbered 67)
Droseraceae 47-48 575-576
180
(wrongly numbered 54;
also see Instalment 9)
Volume 3. Calyciflorae Continued
A Family Ser. Vol. No. Year B Cc D
13 51. Passifloreae WS. i 1 1902 49-56 577-584
53. Begoniaceae 56-68 584-596
(wrongly numbered 52)
54. Ficoideae 68-69 596-597
55. Umbelliferae 69-72 597-600
57. Cornaceae 72-80 600-608
Volume 4. Gamopetalae
The various families were written by G. King (K.), J.S. Gamble (G.), D. Prain, C.B.
Clarke (Cl.) and H.N. Ridley (Ridl.), with King as editor.
A Family (author) Ser. Vol. No. Year B Cc D
14 58. Caprifoliaceae (G.) me 72 4 1903 112-115 2-5 6.11.1904
59. Rubiaceae (K. & G.) 115-229 §-119
15 Rubiaceae cont. Il 73 3 1904 47-135 121-209 26.X.1904
16 Rubiaceae cont. II 74 Extra No. 1905 1-20 211-230 4.1.1906
60. Valerianeae (K.) 21=22 231-232
61. Compositae (K.) 22-48 232-258
62. Stylidieae (G.) 48-49 258-259
63. Goodenovieae (G.) 49-51 259-261
64. Campanulaceae (G.) 51-58 261-268
65. Vacciniaceae (K.) 58-67 268-277
66. Ericaceae (K.) 68-80 278-290
67. Monotropeae (Prain) 81-82 291-292
68. Epacrideae (G.) 82-84 292-294
69. Plumbagineae (G.) 84-86 294-296
78. Gentianaceae (Cl.) 86-91 296-301
17 70. Myrsinaceae (K. & G.) Il 74 Extra No. 1905 93-157 303-367 841.1906
71. Sapotaceae (K. & G.) 157-201 367-411
72. Ebenaceae (K. & G.) 202-230 412-440
73. Styraceae (K. & G.) 230-254 440-464
74. Oleaceae (K. & G.) 254-273 464-483
75. (see Instalment 19)
76. (see Instalment 19)
77. (see Instalment 19)
78. (see Instalment 16)
18 79. Hydrophyllaceae (G.) II 74 Extra No. 1905 275-277 485-487 4.1.1906
80. Boragineae (K.) 277-284 487-494
81. Convolvulaceae (Prain) 284-327 494-537
82. Solanaceae (Prain) 327-340 537-550
83. Scrophulariaceae (Prain) 341-364 551-574
84. Orobanchaceae (Prain) 364-366 574-576
85. Lentibulariaceae (Prain) 366-374 §76-584
86. (see Instalment 21)
181
Volume 4. Gamopetalae Continued
A Family (author) Ser. Vol. No. Year B € D
18 87. Bignoniaceae (G.) I] 74 Extra No. 1905 375-385 585-595
88. Pedalineae (Prain) 385-386 595-596
19 75. Apocynaceae (G.) II 74 Extra No. 1907 387-505 597-715 20.11.1908
76. Asclepiadaceae (G.) 505-597 715-807
77. Loganiaceae (K.) 598-625 808-835
20 89. Acanthaceae (Cl.) II 74 Extra No. 1907 628-698 838-908 27.11.1908
90. (see Instalment 21)
91. Labiatae (Prain) 699-727 909-937
92. Plantaginaceae (Prain) 727-728 937-938
21 86. Gesneraceae (Ridl.) II 74 Extra No. 1908 730-793 940-1003 6.V.1909
90. Verbenaceae (G.) 794-869 1004-1079
Volume 5. Incompletae
King died after completion of Vol. 4 and editorship of Vol. 5 passed over to J.S.
Gamble. The various families were written by Gamble (G.), J.M. Macfarlane
(Macf.), C. de Candolle (C. DC.), H.N. Ridley (Ridl.) and A.T. Gage. Family
numbers 96 and 105 were allocated twice, the second one marked ‘bis’. The account
of the Euphorbiaceae gave only a key to the genera and accounts of the first 6
genera. It was explained in a note in Vol. 75 of the Journal by the Acting Secretary
General of Asiatic Society of Bengal that the complete manuscript for Euphorbiaceae
was awaiting publication when Ridley’s Flora of the Malay Peninsula (1922-1925)
appeared. The Society then felt that publication of the complete manuscript could
not be justified.
A Family (author) Ser. Vol. No. Year B (e D
ph ky. Nyctaginaceae (G.) = 75 1 1912 2-3 - ?
94. Amarantaceae (G.) 3-17
95s Polygonaceae (Gage) 18-23
96. (see Instalment 24)
97. Aristolochiaceae (G.) 23-33
98. (see Instalment 24)
99. Chloranthaceae (G.) 33-34
100. (see Instalment 23)
101. (see Instalment 23)
102. Lauraceae (G.) 35-202
103. Hernandiaceae (G.) 203-204
23 100. Myristicaceae (G.) — 75 2 1912 205-249 ~ ?
101. Monimiaceae (G.) 250-256
102. (see Instalment 22)
103. (see Instalment 22)
104. (see Instalment 24)
105. Thymelaeaceae (G.) 256-265
105 bis. Gonystylaceae (G.) 266-267
106. Elaeagnaceae (G.) 267-269
107. (see Instalment 24)
182
Volume 5. Incompletae Continued
A Family (author) Ser. Vol. No. Year B Cc D
108. Santalaceae (G.) SPW 2 1912 269-278
24 96. Nepenthaceae (Macf.) - 75 3 1914 279-288 4.V1.1914
98. Piperaceae (C. DC.) 288-339
104. Proteaceae (G.) 339-349
107. Loranthaceae (G.) 349-39]
25 96 bis. Cytinaceae (Ridl.) =a 4 1915 393-395 1.VI.1915
109. Balanophoraceae (Ridl.) 395-399
110. (see Instalment 26)
111. Urticaceae, unpublished
112. Juglandaceae (G.) 399-403
113. Myricaceae (G.) 403-405
114. Casuarinaceae (G.) 405-406
115. Fagaceae (G.) 406-466
116. Salicaceae (G.) 466-468
26 110. Euphorbiaceae (Gage) - 75 5 1936 469-527 IX.1936
Monocotyledons
The monocotyledons were written up by H.N. Ridley and published separately in
Singapore in 1907, in three parts.
Family Part Pages
R1 Hydrocharideae 1 3-6
R2 Orchideae 7-231
R3 Apostasiaceae 231-233
R4 Zingiberaceae 2 1-64
RS Amaryllideae 64-69
R6 Burmanniaceae 69-75
R7 Taccaceae 75-78
R8 Dioscoreaceae 78-85
R9 Roxburghiaceae 85-86
R10 Liliaceae 86-108
R11 Pontederiaceae 108-110
R12 Philydraceae 110
R13 Xyrideae 111-113
R14 Commelinaceae 113-125
R15 Triurideae 125-126
R16 Alismaceae 126-127
R17 Naiadaceae 128-129
R18 Flagellariae 129-133
R19 Palmae 133-221
R20 Pandanae 221-235
R21 Araceae 3 1-52
R22 Lemnaceae $2-53
R23 Eriocauloneae $3-55
R24 Cyperaceae $5-118
R25 Gramineae 118-197
183
184
Index
Gard. Bull. Sing. 36(2) (1983)
With this index, a family in the above lists can be located. The families are taken in their
modern sense, and referred to their sequence number in the lists. Their spelling is the officially
conserved one, as given in the International Code of Botanical Nomenclature, so our
Styracaceae are referred to 73 Styraceae. Their circumscription follows the (non-official) list
by Dr. C.G.G.J. van Steenis, Nomina Generum Malesianorum (Leiden 1972), hence our
Symplocaceae are referred to that same number, 73, because King included them in his
Styraceae. Monocotyledons are referred to the work by Ridley, which has a number sequence
of its own, here denoted with ‘‘R’’. If families were later combined, reference is made to both
numbers, so our Orchidaceae which includes Apostasiaeae are referred to R2 and R3 because
Ridley recognised them as separate families.
Acanthaceae
Actinidiaceae
Aizoaceae
Alangiaceae
Alismataceae
Amarantaceae
Amaryllidaceae
Anacardiaceae
Ancistrocladaceae
Annonaceae
Apocynaceae
Apostasiaceae
Aquifoliaceae
Araceae
Araliaceae
Aristolochiaceae
Asclepiadaceae
Balanophoraceae
Balsaminaceae
Begoniaceae
Bignoniaceae
Bixaceae
Bombacaceae
Boraginaceae
Buddleiaceae
Burmanniaceae
Burseraceae
Campanulaceae
Capparaceae
Caprifoliaceae
Casuarinaceae
Celastraceae
Chloranthaceae
Combretaceae
Commelinaceae
Compositae
Connaraceae
Convolvulaceae
Cornaceae
Crypteroniaceae
Cucurbitaceae
Cunnoniaceae
Cyperaceae
R16
Instalment 13
28 and
Datiscaceae
Dichapetalaceae
Instalment 8
Dilleniaceae 2
Dioscoreaceae R8
Dipterocarpaceae 16
Droseraceae ; 41 and
Instalment 13
Ebenaceae 72
Elaeagnaceae 106
Elaeocarpaceae 19
Epacridaceae 68
Ericaceae 65, 66, 67
Eriocaulaceae R23
Erythroxylaceae 20
Euphorbiaceae 110
Fagaceae 115
Flacourtiaceae 9, 50
Flagellariaceae R18
Gentianaceae 78
Gesneriaceae 86
Gonystylaceae 105 bis
Goodeniaceae 63
Gramineae R25
Guttiferae 14
Haloragaceae 43
Hamamelidaceae 42
Hernandiaceae 45, 103
Hydrocharitaceae Rl
Hydrophyllaceae 79
Hypericaceae 13
Icacinaceae 29
Illiciaceae 3
Juglandaceae 112
Labiatae 91
Lauraceae 102
Lecythidaceae 46
Lemnaceae R22
Leguminosae 38
Lentibulariaceae 85
Liliaceae R10
Linaceae 20, 29
Loganiaceae 77
Loranthaceae 107
Lowiaceae R4
Lythraceae 48
Magnoliaceae 3
Malpighiaceae 21
Malvaceae 17
Marantaceae R4
Melastomataceae 47
Meliaceae 27
Menispermaceae 5
Monimiaceae 101
Musaceae R4
Myricaceae 113
Myristicaceae 100
Myrsinaceae 70
Myrtaceae 46
Najadaceae R17
Nepenthaceae 96
Nyctaginaceae 93
Nymphaeaceae 6
Nyssaceae $7
Ochnaceae 75)
Olacaceae 29
Oleaceae 74
Onagraceae 49
Opiliaceae 29
Orchidaceae R2, R3
Orobanchaceae 84
Oxalidaceae 22
Palmae R19
Pandanaceae R20
Passifloraceae 51
Pedaliaceae 88
Piperaceae 98
Pittosporaceae 10
Plantaginaceae 92
Plumbaginaceae 69
Polygalaceae 1]
Polygonaceae 95
Portulacaceae 12
Pontederiaceae R11
Proteaceae 104
Rafflesiaceae 96 bis
=
Guide to King’s ‘‘Materials’’
Ranunculaceae
Rhamnaceae
Rhizophoraceae
Rosaceae
Rubiaceae
Rutaceae
Sabiaceae
Salicaceae
Santalaceae
Sapindaceae
Sapotaceae
Sarcospermataceae
Saurauiaceae
116
108
Saxifragaceae
Scrophulariaceae
Simaroubaceae
Solanaceae
Sonneratiaceae
Staphyleaceae
Stemonaceae
Sterculiaceae
Stylidiaceae
Styracaceae
Symplocaceae
Taccaceae
Tetrameristaceae
40
83
24
82
48
34
R9
18
62
73
73
R7
25
Theaceae
Thymelaeaceae
Tiliaceae
Trigoniaceae
Triuridaceae
Umbelliferae
Valerianaceae
Verbenaceae
Violaceae
Vitaceae
Xyridaceae
Zingiberaceae
185
15
105, 105 bis
19
11
R15
55
NOTES ON THE SYSTEMATY OF MALAYAN PHANEROGAMS
XXX. Anacardiaceae
K. M. KOCHUMMEN
Forest Research Institute, Kepong, Malaysia
EFFECTIVE-PUBLICATION DATE: 7TH MAR. 1984
Abstract
The purpose of these notes is to formalise several taxonomic changes in the family Anacardiaceae
which has just been revised for the Tree Flora of Malaya. The changes are as follows:-
Mangifera
M. indica L.: reduction of M. longipes to M. indica and resultant range extension
of M. indica.
M. magnifica Kochummen, sp. nov.
M. quadrifida Jack var. longipetiolata (King) Kochummen, stat. nov.
M. whitmorei Kochummen, sp. nov.
Melanochyla
M. caesia (Bl.) Ding Hou: reduction of M. kunstleri to M. caesia.
M. fasciculiflora Kochummen, sp. nov.
M. longipetiolata Kochummen, sp. nov.
Rhus
R. succedanea L.: new record.
Semecarpus
S. rufovelutinus Ridl.: new record
S. trengganuensis Kochummen, sp. nov.
Swintonia
S. floribunda Griff. var. penangensis (King) Kochummen, stat. nov.
Introduction
The Anacardiaceae for Malesia was revised by Ding Hou of Flora Malesiana
Foundation. This was published in Flora Malesiana Vol. 8 (1978).
While I was writing up the Anacardiaceae for the 4th volume of Tree Flora of
Malaya, based on Hou’s revision, I discovered some new species as well as new
records. Critical study of the family required reduction of certain taxa as well as
establishment of new status for some taxa.
MANGIFERA
Mangifera indica L.
Mangifera longipes Griff., Notul. 4 (1854) 419; type: Griffith 1096 (K)
syn. nov.
According to Hou, the main differentiating character between M. indica and
187
KEP 20562
SFN 34970
Fig. 1. Mangifera magnifica Kochumm. sp. noy.
Leafy twig, with longitudinal section of a fruit (right),
c: sepal, inside view; d: petal, inside view; e: disc, stamen and pistil. (a, x 4; DEG:
and a seed (/eft). Inset - a & b: flowers;
al, 2, Aull S< 3)
188
Systematy of Malayan Anacardiaceae 189
M. longipes is the degree of hairiness of the calyx, i.e., densely puberulous on both
surfaces especially on the outside in M. indica, but sparsely pubescent to glabrous
in M. longipes. However I have come across several collections of the cultivated
mango (M. indica) with glabrous calyx and so this character does not hold good.
The degree of hairiness varies among the various cultivars. Consequently I have
decided to merge M. longipes Griff. with M. indica L. I have not seen the type of
M. longipes, but there is one sheet at Kepong, KEP 3637 on which is written in
C. F. Symington’s hand: ‘‘matches with the type, Griffith 1096’’. Many of the
collections formerly distinguished as M. /ongipes are undoubtedly wild forest trees
in Malesia and hence the natural range of M. indica must now be considered to
extend into the Malesian region.
Mangifera magnifica Kochummen, sp. nov. Fig. 1.
Arbo excelsa usque at 54 m alta et 76 cm diametro; caulis sine anteridum; cortex extus cinereo-brunneus,
laevis vel fissuratis et squamatis; intus luteolus; latex albus macilentus.
Ramunculi c. 7 mm crassi, glabri, subangulati. Folia spiraliter dispositi, petiolis crassis, 2 - 4.5 cm
longis, basi gibbosis. Lamina crassa elliptica vel oblonga, variabilissima, 6 - 26.5 x 5 - 12 cm, apice
obtuso vel rotundato, basi rotundato vel cuneato, raro asymmetrica, marginibus reflexibus; nervis
secundariis 11-20-paribus, nervis intermediis paucibus, nervis tertiariis reticulationes formatibus obsoletis
raro visibilis. Inflorescentia paniculata terminalis axillaris glabra 15 cm longa breviter ramosa, axibus
complanatis; floribus brevipedicellatis c. 1.5 mm longis. Sepala 4 triangularia, 1.5 mm lata, acuminata,
extus papillosa. Petala 4 oblonga glabra, c. 4 x 1.5 mm, extus intusque papillosa, costata, costa
e dimidio petali 3-4-diviso glanduloso. Discus extrastaminalis. Stamen fertilium unicum. Ovarium
glabrum sulco distincto, stylo laterale. Drupa oblonga 11 x 7 cm (in sicco), mesocarpio 1 cm crasso,
endocarpio longitudinaliter striato; semen 10 x 6 cm, testa papyraceo.
Typus: KEP 20562, Gunung Jerai, Kedah (Holotypus KEP, isotypus SING).
Distribution: common, lowlands and hill forests throughout Malaya except
Perlis, Penang and Malacca.
Other collections seen: KEDAH: KEP 11251, 20562, 105245; PERAK: KEP 8025,
9534, 11612, 40613, 63259, 65917, 69063, FRI 13941; KELANTAN: KEP 68315;
TRENGGANU: FRI 16869; PAHANG: KEP 17288, 69602; SELANGOR: KEP
84613; NEGRI SEMBILAN: KEP 18422, 64749, 93265, 93797, 104762; JOHORE:
KEP 69918, 70058, 70085, 70221, 84698, 92204, 93555, 104734, 105241, 116033,
SFN 34970.
This is the thickest-leafed Mangifera with strong recurved margins and having
floral parts similar to M. quadrifida.
Mangifera quadrifida Jack var. longipetiolata (King) Kochummen, stat. nov.
Basionym: M. longipetiolata King, J. As. Soc. Beng. 65 (1896) 470; type: King’s
Collector No. 7266 (K).
Ding Hou reduced M. longipetiolata to M. quadrifida Jack. After examining a
photograph of the type of M. Jongipetiolata and the herbarium collections at
190 Gard. Bull. Sing. 36(2) (1983)
Kepong, I have come to the conclusion that vegetative characters like the very large
leaves, often exceeding 24 cm long and 8 cm wide, the leaf stalks 4 - 10 cm long,
prominently swollen at their base, and the raised tertiary nerves and lax reticulations
on the upper leaf surface are sufficient to merit varietal status.
Mangifera whitmorei Kochummen, sp. nov. Fig; 2:
Arbor usque ad 30 m alta, corona subrotunda coronellis parvioribus numerosis. Cortex tenuiter
fissuratus. Cortex interioris subroseus et rubrostriosus; latex albus guttulosis. Lignum succosum album.
Ramunculi rubro-brunnei glabri inconspicue striati. Petiolus tenuis 4 - 4.5 cm longis basi tumido
rugoso. Lamina tenue coriacea anguste oblonga 11 - 13 x 3 - 3.8 cm, acuminata, basi cuneato, nervis
secundariis 15-17-paribus, (nervis intermediis paucibus) improminentibus, supra elevatis; nervis tertiariis
reticulationes formantibus, infra visibilis, supra inconspicuis; costa media supra elevata. Inflorescentia
axillaris paniculata glabra usque ad 23 x 10 cm, pauciramosa. Flores bisexuales cremei pentameri,
pedicelli 6 mm longi, infra apice 1 mm articulati. Sepala 5 glabra deltoidea 2 x 1.5 mm, obscuriter
6-lineata. Petala 5 oblonga glabra 4 x 1.5 mm pentacostulata (costuli quam petali “3s breviori), in parte
basilari tertiali confluenta. Stamina 5 fertilia, 4 mm longa, filamentis separatis, antheris globosis;
staminodiis 5 valde parvioribus oppositepetalis. Ovarium subglobosum 2 mm latus glabrum, glandulosum,
glandis papilliformibus dispersis. Stylus lateralis 2.5 mm longus. Cetera ignota.
Typus: FRI 15820 (Whitmore), Upper Perak, 150 m alt. (Holotypus KEP, isotypi
K, L, A, SING).
Known only by the type, however the collector stated that it is common on the
hill slopes in the area.
The only other species with 3 - 5 fertile stamens is M. pentandra from which it
differs by the glabrous inflorescences and long-pedicelled flowers.
MELANOCHYLA
Melanochyla caesia (Bl.) Ding Hou
Melanochyla kunstleri King, J. As. Soc. Beng. 65 (1896) 504; type: King’s
Collector No. 6810 (SING); syn. nov.
When King described M. kunstleri he commented that it is very close to M. maingayi
Hook.f. and the only difference he could discover was the smaller, more shiny leaves
in M. kunstleri. Ding Hou later reduced M. maingayi to M. caesia. After examining
the type of M. kunstleri and all collections under M. caesia and M. kunstleri at
Kepong and Singapore herbaria I could find no floral or vegetative differences
between the two.
Melanochyla fasciculiflora Kochummen, sp. nov. Fig. 3.
Frutex c. 5 m altus, ramunculis pallidis, gemma terminale ferrugineo hirsuto. Folia tenuicoriaceae,
laminis 10.5 - 16.5 x 2.5-5 cm, acuminatis (acumen ad 1 cm longis), basi cuneato, pagina supra glabro,
infra uniformiter papilloso et ad costam venamque sparsiter hirtello; costa infra canaliculato; nervis
secundariis 14-18-paribus curvato-adscendentibus juxta marginem transientibus; nervis intermidiis
cm 10 + 4 in.
4 mm
KEP 15820
Fig. 2. Mangifera whitmorei Kochumm. sp. nov.
Leafy twig with inflorescence. Inset - a: flower; b: sepal, inside view; c:
d: disc, stamens and pistil.
petal, inside view;
19]
192 Gard. Bull. Sing. 36(2) (1983)
brevibus; nervis tertiariis scalariformi reticulatis infra valde elevatis, supra inconspicuis; petiolo 0.7 - 2.2
cm longo. Flores ignoti. Infrustescentia axillaris, fructibus aggregatis, maturitate flavidis, oblongis,
c. 1.5 x 1 cm, hispidis; acuminoides.
Typus: FRI 16366 (Kochummen), Johore, Rengam V.J.R. (Holotypus KEP,
isotypi K, L, A, SING).
FRI 16374, from the same locality, is another collection belonging to this species.
M. fasciculiflora is very distinct from all other Melanochyla species because of the
clustered axillary fruits.
Melanochyla longipetiolata Kochummen, sp. nov. Fig. 4.
Arbor usque ad 30 m alta, 100 cm, diametro; cortex cinereus, laevis, rimosus.
Ramunculi pallide brunnei, angulati. Folia crassiter coriacea, petiolo 1 - 4.5 cm longo, basi incrassato;
lamina elliptico vel obovato 16 - 26 x 4 - 9.5 cm, acuminato, basi cuneato, infra glauco papillosoque;
nervis secundariis 14-16-paribus, tertiariis scalariformibus reticulatis tenuibus. Inflorescentia terminalis
paniculata 22 cm longa, floribus incognitis. Drupa ovoidea 4 x 2 cm, acuminata (in sicco), adpressiter
pilosa, rugosa.
Typus: FRI 8478 (Cockburn), Trengganu, Kg. Petang, Ulu Trengganu (Holotypus
KEP, isotypi K, L, A, SING).
Distribution ; very rare, lowlands to mountain forests.
Other collections seen: TRENGGANU: FRI 20341; PAHANG: KEP 26123.
RHUS
Rhus succedanea Linn.
This species is a new record for Malaya. Ding Hou (in Jitt.) confirmed the
identification as I was not able to see the type. It is a common tree along the valleys
of mountain forests 1400 m at Fraser’s Hill and also at Cameron Highlands in
Pahang. It can grow up to 40 m tall and 200 cm girth.
Collections seen: PAHANG: KEP 27166, 27171, 31031, 44864, 45411; FRI 23106,
29379, 29425, 29467, 32538.
SEMECARPUS
Semecarpus rufovelutinus Ridl.
This species is a new record for Malaya. It is acommon small tree in the lowland
forests of east Johore and Pahang. I have not seen the type, but the material match
Bornean specimens SAN 39887 & 88330 which have been identified by Ding Hou.
4 in.
KEP 16366
Fig. 3. Melanochyla fasciculiflora Kochumm. sp. nov.
Leafy twig with fruit. Inset - part of the underside of a leaf, enlarged to show the channelled
midrib.
193
ee
we, cm! 1O=4sine
FRI 8478
Fig. 4 Melanochyla longipetiolata Kochumm. sp. nov.
Leafy twig and fruits.
194
>
ere
KEIN VAYE AOR PEER ILS
saws ee cant
eet
FRI 28300
a We FRI 16872
Fig. 5. Semecarpus trengganuensis Kochumm. sp. nov.
Leafy twig with inflorescence. Inset - : a: flower bud; 5: petal, inside view; c: petal, outside
view; d: longitudinal section of flower. (a & b, x 12; b, c, & e, x 20).
195
196 Gard. Bull. Sing. 36(2) (1983)
Collections seen: PAHANG: Samsuri Ahmad 397; JOHORE: KEP 97982, FRI
13802, Hardial Singh 1064, Samsuri Ahmad 506.
Semecarpus trengganuensis Kochummen, sp. nov. Fig. 5.
Frutex usque ad 3 m altus, ramunculis griseo-brunneis striatis, glabris. Folia alternata sed verticillata,
petiolo c. 2-4 cm longo, supra canaliculato; lamina obovato vel anguste elliptico 16.5 - 37 x 6-15 cm,
acuminato, basi cuneato, margine reflexo sinuato albido; nervis secundariis 13-16-paribus, juxta
marginem recurvatis; tertiariis reticulationibus distinctis formantibus in paginis ambis. Inflorescentia
paniculata terminalis et axillaris usque ad 18 cm longa pauciramosa hirtellae. Flores masculas (in
alabastro) 1.5 mm longi, pedicello 0.2 mm longo hirtello, bracteis deltoideis 1 mm longis hirtellis, calyci
lobis 5 obtusis 0.5 mm longis marginibus albidis; petalis 5 oblongis 1 x 0.5 mm intus glabris extus
hirtellis prominente 4-venatis; disco intrastaminale apice longe hirsuto; staminis 5, ad 0.5 mm longis, an-
theris oblongis basifixis. Flores foemineas non vidi. Fructus subglobsus 3 mm latus sparsiter hirtellus,
obscure canaliculatus, hypocarpio obconico hirtello; pedicello hirtello 3 mm longo.
Typus: FRI 16872 (Chan), TRENGGANU, Bukit Bauk F.R. (Holotypus KEP
isotypi K, L, A, SING).
Other collections seen: TRENGGANU: FRI 3940, 25163, 28300; PAHANG: KEP
28446, SFN 10688.
This species is close to Semecarpus curtisii but differs in the short few branched
inflorescences, in the pale wavy-margined leaves and in the obconic hypocarp.
SWINTONIA
Swintonia floribunda Griff. var. penangiana (King) Kochummen, stat. nov.
Basionym: Swintonia penangiana King, J. As. Soc. Beng. 65 (1896) 490; type:
Curtis 1579 (SING).
Hou reduced Swintonia penangiana King to S. floribunda Griff. After examining
the type and studying the collections at Kepong and Singapore herbaria I find that
S. penangiana King differs from S. floribunda Griff. in having non-glaucous leaves
with very faint to inconspicuous veins, abruptly acuminate apex, and a characteristic
brownish colour when dried. On the basis of these features I am raising it to varietal
status.
A new Species of Nepenthes from Sulawesi, Indonesia
SHIGEO KURATA
Insectivorous Plants Society of Japan, Department of Biology, Nippon Dental College
Fujimi, Chiyoda-Ku, Tokyo, Japan
EFFECTIVE-PUBLICATION DATE: 7TH MAR. 1984
Introduction
During my stay at the Herbarium Bogoriense in 1972, for the study of their
Nepenthes collection, I was able to examine much undetermined material from
several Indonesian islands. While going through those collected by P. J. Eyma in
Sulawesi, I came across a very interesting Nepenthes. After subsequent study, I am
now able to conclude that it should be described as a new species.
Nepenthes dentata Kurata, sp. nov. Plate 1 & Fig. 1
Nepenthes dentata Kurata, nomen nudum, in Nepenthes of Mount Kinabalu,
(1976) 11.
Planta tenuis, alte scandens. Caulis cylindricus vel obtuse trigonus, 4-5 mm crassus, glabra. Folia
inferiora ignota. Folia superiora coriacea, sessilia, lanceolata vel elliptica, glabra, 10-12 cm longa, 3-4 cm
lata, apice obtusa, basi rotundata vel leviter cordata, caulim = amplectens, sine vagina, utrinque glabra;
nervi longitudinales utrinque 3-4, nervi transversales obscure, oblique ascendentes; cirrhus tenuis,
cylindricus, 10-12 cm longus, glaber. Ascidia inferiora ignota. Ascidia superiora coriacea 18-22 cm
longa, 3.5-4.5 cm lata, parte inferiori anguste ovata, parte superiori cylindrica vel leviter infundibulifor-
mia, intus parte - inferiori glandulosa, costis 2 prominentibus; os ovatum, obliquum; peristomium
pectinatum, dentibus magnis, lamellaribus, falciformibus vel lunaribus, 12-16 mm longis, 2-3 mm latis,
5-6 mm distantibus; operculum elliptico-ovatum, 4.5-5.5 cm longum, 4-5 cm latum, subtus planum;
calcar ca. 6 mm longum, filiforme. Inflorescentia mascula dense racemosa, ca. 15 cm longa; pedunculus
ca. 10 cm longus, glaber; pedicelli insigniter tenues, 10-15 mm longi, 0.1-0.2 mm crassi, uniflori, glabri;
sepala 4, elliptica, ca. 1.5 mm longa, ca. 1 mm lata, extus glabra, intus glandulosa; columna staminea
2-2.5 mm longa; antherae 6-8, uniseriatae. Inflorescentia feminea ignota.
SULAWESI. Central Sulawesi; G. Lumut, between bivouac II and III on the
north spur, 3 Sept. 1938, P. J. Eyma 3572 (holotype & isotype, BO).
Stem slender, climbing high, the part with adult leaves 4-5 mm thick, cylindrical
or obtusely trigonous, glabrous. Lower leaves unknown. Upper leaves scattered,
coriaceous, sessile, lanceolate or elliptic, 10-12 cm long, 3-4 cm broad, the apex
obtuse, the base rounded or slightly cordate, obliquely clasping the stem for @,
longitudinal nerves 3-4 on both sides, pinnate nerves running obliquely towards the
margin, irregularly reticulate; tendrils slender, 10-12 cm long, cylindrical, glabrous.
Lower pitchers unknown. Upper pitchers coriaceous, 18-22 cm high, 3.5-4.5 cm
broad, narrowly ovate in the lower part, cylindrical or slightly infundibuliform in
the upper part, with 2 prominent ribs; mouth oblique, ovate; peristome pectinate,
the teeth large, lamellate, falcate or crescent-shaped, 12-16 mm long, 2-3 mm
broad, arranged on the narrow cylindrical base at intervals of 5-6 mm; inner surface
of the pitcher glandular in the lower ovate part, the glands rounded and exposed
LOT,
5 Cm a
'
(Eyma 3572).
Plate 1. Holotype of Nepenthes dentata Kurata sp. nov.
198
Nepenthes dentata Kurata sp. nov. 199
(not overarched); lid elliptic/ovate, 4.5-5.5 cm long, 4-5 cm broad, the lower
surface without appendage, with small round glands on the marginal part; spur
filiform, about 6 mm long. Male inflorescence densely racemose, about 15 cm long,
the peduncle about 10 cm long, glabrous; pedicels extremely thin, 0.1-0.2 mm thick,
10-15 mm long, 1-flowered; sepals 4, elliptic, about 1.5 mm long, 1 mm broad, the
outer surface glabrous, the inner surface glandular; staminal column 2-2.5 mm
long, the anthers 6-8, uniseriate. Female inflorescence unknown. Colour of the
B
Y
AONE Sop ia)
xO £y
Q@pDOLP™
Fig. 1. Nepenthes dentata Kurata sp. nov. A: leaf & pitchers; B: male flower; C: side view of a tooth.
200 Gard. Bull. Sing. 36(2) (1983)
pitcher light-green with dark-red blotches or stripes, tooth yellow to light-green.
This new species is closely related to N. tentaculata Hook.f. by its sessile leaves,
exposed (not overarched) pitcher glands and purplish brown colour of its dried
specimen but differs clearly from it by the presence of its unique and remarkably
large teeth at the mouth of the pitcher (hence the epithet ‘‘dentata’’). The flower
also characterizes this species by its small dimensions, which is considered to be the
smallest in the genus. The pedicel, extremely slender in relation to its length — at
most 0.2 mm in thickness — distinguishes this species too.
The teeth, roughly arranged along the slender peristome with an interval of about
5 mm, have the shape of a thin blade of a sickle or crescent and are about 15 mm
long. This shape reminds us of two Bornean species (viz. N. villosa Hook.f. and N.
edwardsiana Hook.f.) but the present species is not to be confused with them by its
superficial semblance as these Bornean species are, all parts considered, quite
robust.
The distribution of this species seems limited to a particular area on the mountains
of Central Sulawesi.
The author wishes to commemorate P. J. Eyma for his courage and achievement,
in bravely stepping into the unknown world of Gunung Lumut forty-five years
ago. I also wish to express my thanks to Professor A. J. H. Kostermans of
Herbarium Bogoriense for his valuable suggestions in this study and to Dr. R. D.
Hoogland who corrected my Latin diagnosis.
Bibliography
Danser, B. H. (1928). The Nepenthaceae of the Netherlands Indies. Bull. Jard. Bot.
Buitenz., 9 (ser. ili).
. (1935). Note on few Nepenthes. Bull. Jard. Buitenz., 13 (ser. ii).
Harms, H. (1936). Nepenthaceae. In A. Engler & K. Prantl, Die Naturlichen
Pflanzenfamilien, 2nd ed. 17b.
Hooker, J. D. (1873). Nepenthaceae. In De Candolle, Prodromus systematis univer-
salis regni vegetabilis 17: 90-105.
Kurata, S. (1976). Nepenthes of Mount Kinabalu. Sabah National Parks Trust.
. (1973). Nepenthes from Borneo, Singapore and Sumatra. Gard. Bull.
Sing 26: 227-232.
Macfarlane, J. M. (1908). Nepenthaceae. In A. Engler, Das Pflanzenreich, 36 (ser.
ili): 53-54.
On the Nature of Leaf-opposed Inflorescences in
Aidia cochinchinensis (Rubiaceae)
WONG KHOON MENG! & LIM AH LAN:
‘Forest Research Institute, Kepong, Malaysia
?University of Malaya, Kuala Lumpur, Malaysia
Abstract
Leaf-opposed inflorescences in Aidia cochinchinensis, previously ascribed a terminal origin, is here
shown to be axillary instead. Hence, the branch system in this species is not sympodial as believed, but
a true monopodium along which inflorescences develop from the axils of reduced leaves at alternate
nodes, exemplifying a specialised control of floral development.
Introduction
Aidia cochinchinensis Lour., a small tree-species distributed from the north-
eastern part of the Indian subcontinent to Malesia, exhibits a specialised morphology
in its branches. The flowering cymes develop only at alternate nodes along each
branch, on its upper side. Along the branches, alternate leaf-pairs have laterally
disposed leaves and the intervening (flowering) nodes have a fully developed leaf
from the lower side of the branch, the leaf on the upper side being reduced to a tiny
scale which subtends the inflorescence. As such, each inflorescence appears leaf-
opposed.
Previous Hypothesis for the Origin of Leaf-opposed Inflorescences
Fagerlind (1943) had characterised A. cochinchinensis Lour. (therein as Randia
cochinchinensis (Lour.) Merr.) as having inflorescences terminating the lateral
shoots from which subsequent branch extension is formed by an axillary shoot from
the most distal leaf-pair, so that a series of such shoots forms a sympodial branch
system (Fig. 1A). It was explained that this inflorescence would subsequently be
displaced to the upper side of the branch sympodium by the developing shoot from
the final node preceding the inflorescence. The displaced inflorescence thus appears
lateral and leaf-opposed because the leaf on the upper side of this final node fails
to develop normally. Subsequently, Tirvengadum (1978) has also adopted this
explanation for the leaf-opposed inflorescences in this species.
Hallé (1967) has recorded a similar mechanism of branch development in
Schumanniophyton problematicum (Chev.) Aubr. and Massularia acuminata
(Benth.) Bullock ex Hoyle; in the former he noted a series of three buds in the leaf
axil on the lower side of the branch at the final node preceding the inflorescence.
The most distal of these buds then develops to continue sympodial extension of the
branch, and Hallé designated the period between two successive flowering nodes as
equivalent to one year (Fig. 1B). In other words, the branch system is a sympodium
of hapaxanthic modules (Hallé ef a/. 1978).
201
202 Gard. Bull. Sing. 36(2) (1983)
Evidence for an Axillary Origin of the Inflorescences in Aidia
In A. cochinchinensis, the origin of the inflorescence is in fact axillary. Several
morphological observations lend evidence to this conclusion:-
(a) The branch system is remarkably straight, and during its growth may extend by
several internodes and nodes which remain vegetative. In the vegetative state, the
current season’s branch-extension would be equivalent to a series of successive
internodes, and to accept that this is in fact a series of two-internode segments
forming a sympodium requires that a whole series of buds terminal to the
sympodial segments remain dormant for some time.
(b) Often, an axillary shoot develops on the lower side of the node of leaf reduction,
simultaneous with (or even earlier than) the development of the bud from the
upper side of the node (which becomes the inflorescence).
An axillary origin of the inflorescence is confirmed by a longitudinal section
through the apex of a branch shoot terminated by a node of leaf reduction (Fig. 2).
Here, it can be seen that the primordia of the leaf axillary buds (the one subtended
by the reduced leaf being destined to form an inflorescence) are already present at
an early stage of development, and are distinct from the central terminal bud which
develops to continue shoot extension.
At a later stage of development, the central terminal bud extends its growth to
form the next node of two normally developed leaves (Fig. 3), while the axillary buds
are still dormant. There are no serial buds in leaf axils, as confirmed by sections
through nodes of vertical axes and the branches; each leaf axil produces one
axillary bud.
On older parts of the branches which have not yet produced inflorescences,
longitudinal sections through the potentially flowering (one-leaf) nodes indicate that
while there is already a clear and straight continuation of the main vascular cylinder
across into the next internode, the inflorescence bud can still remain in a dormant
state, not supplied by any clear vascular connection.
The Flowering Habit in Aidia cochinchinensis
The branch system in A. cochinchinensis is thus a monopodium along which there
is leaf reduction at alternate nodes. The dormant bud from the axil of the (reduced)
scale leaf, on the upper side of the branch, is destined to develop into an in-
florescence. Two modes of flowering behaviour may be seen: either several successive
potentially flowering nodes can initially remain vegetative and subsequently flower
together, or such nodes can develop inflorescences progressively as branch extension
continues. These two modes of behaviour may occur on the same plant, their
prevalence seems to depend on the number of internodes added through branch
extension during each growing season.
Fig. 1. A. Sympodial branch development explained for Randia sect. Gynopachys, from Fagerlind
(1943); he included R. cochinchinensis (a synonym of Aidia cochinchinensis) in that section.
B. Sympodial branch development explained for Schumanniophyton problematicum, from Hallé
(1967); note serial buds in the leaf axils.
Fig. 2. Longitudinal section through the apex of a branch shoot in Aidia cochinchinensis.
To the left of the section is the lower side of the branch, where the petiole of the developed leaf
(pd) is found; to the right is the petiole of the reduced leaf (pr), and the stipules corresponding
to these leaves are designated by S/; ad: axillary bud of developed leaf; ar: axillary bud of
reduced leaf, which is equivalent to the inflorescence bud; S2: stipules corresponding to the next
pair of leaves (at right angles to the plane of the section) and which protect the central terminal
bud; gf: glandular trichomes or colleters found at the inner basal parts of the stipules.
203
204 Gard. Bull. Sing. 36(2) (1983)
=
Co
i=
oO
=
oOo
_
o
upper side
of branch
@
a
o
=
=
Fig. 3. Longitudinal section through the apex of a branch shoot in Aidia cochinchinensis, at the stage
of development subsequent to that in Fig. 2. pd: petiole of developed leaf; pr: petiole of reduced
leaf; S/: stipules corresponding to pd and pr; ad: axillary bud of developed leaf; ar: axillary bud
of reduced leaf (which develops to form the inflorescence); L: one leaf from the most distal pair
of differentiated leaves; $2: stipules corresponding to the most distal pair of differentiated leaves;
t: terminal bud; v- vascular tissue.
Acknowledgements
Drs. Diane M. Bridson, F.S.P. Ng, Ong Siew Ling and B.C. Stone kindly
commented on a preliminary manuscript. We are grateful to Dr. Ong for providing
extensive translations from German.
Literature Cited
Fagerlind, F. (1943). Die Sprossfolge in der Gattung Randia und ihre Bedeutung fiir
die Revision der Gattung. Arkiv for Botanik 30A (7): 1-57.
Halle, F. (1967). Etude biologique et morphologique de la tribu des Gardenieées
(Rubiacées). Mem. ORSTOM, no. 22.
Hallé, F., R.A.A. Oldeman & P.B. Tomlinson. (1978). Tropical Trees and Forests:
an architectural analysis. Springer-Verlag, New York; xvii+441 pp.
Tirvengadum, D.D. (1978). A Synopsis of the Rubiaceae — Gardenieae of Ceylon
(Sri Lanka). Bull. Mus. natn. Hist. nat. Paris, 3° ser. no. 521 Botanique 35:
3-33.
SOME NEW AND CRITICAL PANDANUS SPECIES
OF SUBGENUS ACROSTIGMA
I*. Supplement to Revisio Pandanacearum
BENJAMIN C. STONE
Department of Botany, University of Malaya, Kuala Lumpur, Malaysia
EFFECTIVE-PUBLICATION DATE: 7TH MAR. 1984
Abstract
Eight species of Pandanus subgen. Acrostigma are discussed in this supplement. Seven of them are
additions:— four have recently been published, two more are proposed as new herein and one has been
omitted. P. ellipsoideus was poorly known and is here given a fuller description based on a recent
collection. Both new species pertain to sect. Acrostigma subsect. Dimissistyli; one (P. gladiator) is from
Sulawesi, the other (P. jacobsii) is from Papua New Guinea. As is usually the case, most of these species
are as yet not represented by staminate specimens (known only in P. merrillii). Only one, P. microglottis,
presents features which suggest a possible addition to the infrageneric scheme, but in the absence of
sufficient specimens no change is suggested for the present.
Introduction
A synopsis, based on a critical revision, of two of the eight subgenera of Pandanus,
subgen. Acrostigma and subgen. Coronata, was issued recently (Stone, 1978). That
study, forming a component of the preliminary work leading to the revision of the
family and to a family treatment in the Flora Malesiana, recognized three species
in subgen. Coronata and 66 species in subgen. Acrostigma. Since then, a few
additional species have been recognized, either as new or newly understood taxa
from among the synonyms or nomina dubia. In this paper, these additions are listed
in the same order of sections, subsections, and taxonomic order as was followed
in the publication cited.
PANDANUS subgen. ACROSTIGMA (Kurz) Stone Sect. Acrostigma Kurz
SUBSECT. Acrostigma
Pandanus ellipsoideus Warburg, Pflanzenr. 3 (1900) 81. Martelli, Webbia 4 (1913)
13, 4 (1914) t. 31, f. 19-20. Stone, Fedn. Mus. J. n.s. 23 (1978) 15. _—‘Fig. 1.
SULAWESI: Sopu valley, west slope, Roroka Timbu, ridge forest at 1200 m. alt.,
clay soil; short-stemmed stooling herb, inflorescence erect, fruit green, leaves used
for making mats (tikar), 4 May 1979, M. J. van Balgooy 3147 (L).
Extended description: short-stemmed, stooling herb. Leaves linear, c. 120 cm long
* This paper is dedicated to the late Marius Jacobs of the Rijksherbarium, Leiden, in memory of his
contributions to Malesian botany and his ungrudging help and assistance.
205
2
>
°
8
SS
S
N
yep soe
Below left: Fig. 1. Pandanus ellipsoideus Warb. — A: cephalium. 1.s.; B & C: drupe, B: profile and
C: 1.s., note the chamber above the endocarp in C; D: leaf apex, adaxial surface. All from Balgooy 3147
(L).
Above left: Fig. 2. Pandanus merrillii Martelli — Cephalium (upper) + x 1; and drupes (/Jower) + x
1; from the type collection, Merrill 840 (FI). (Courtesy of Dr. C. Steinberg).
Right: Fig. 3. Pandanus merrillii — Infructescence with cephalia, + x 2/3; from Bermejos BS 232
(FI). (Courtesy of Dr. C. Steinberg).
206
Supplement to Revisio Pandanacearum 207
(or more), 3.8 cm wide, the apex gradually acuminate; lower surface glaucous;
upper surface finely striolate, indistinctly cross-nerved toward apex; lower surface
with fine, close longitudinal nerves (c. 84-86 per leaf), very slightly indistinctly
raised, intervein areas densely stomatophorous. Leaf margins near base with spreading
slender prickles c. 2 mm long, usually 3-5 mm apart; near middle, the prickles
shorter, more appressed, c. 1 mm long, 3-6 mm apart; near apex, the prickles still
smaller, c. 0.5 mm long, 1-2 mm apart. Beneath, midrib near base with short erect
or retrorse prickles at least 1 mm long (or reduced and nut-like on scale leaves and
lower bracts), often 3-4 mm apart but rather irregularly spaced; near middle, the
midrib carinate, with antrorse prickles c. 1 mm long, 4-10 mm apart; near apex, with
prickles similar in size and spacing to those on adjacent margins. Apical ventral pleats
prickly along distal 30-33 cm with numerous small antrorse prickles c. 0.5-0.7 mm
long, nearer the apex mostly 1.5-4 mm apart. /nflorescence erect, the pistillate
peduncle to 50 cm long, 8 mm wide, triquetrous, glabrous, bracteate, distally
zig-zag-helical, bearing 5 to 7 smallish ellipsoid cephalia c. 6 cm long, 5 cm wide,
composed of numerous drupes, these with projecting, sharp, straight or slightly
curved styles. Drupes 15-20 mm long (including style), c. 5 mm wide, the corpus
clavoid, mostly 11-13 mm long, the pileus 5—6-angled, steeply pyramidal, gradually
and rather indistinguishably produced upward as the sharp spiniform style, this
erect or only slightly antrorsely curved, sharp, to 9 mm long; stigma linear, grooved,
papillose, almost reaching the style tip. Apical mesocarp chamber large, rounded,
c. 4mm long; lower mesocarp very short fibrous; endocarp pale, c. 6 mm long, apex
concave, base tapered, walls c. 0.5-0.6 mm thick.
In its inflorescences, leaves, and fruits, this species very much resembles P. affinis
Kurz, a freshwater swamp lowland species common in Western Malesia, but in habit
(as noted above) and certainly in habitat, this species appears quite different. The
drupes of the cited specimens are an excellent match for those illustrated by Martelli.
Pandanus ketele B. C. Stone, J. Arn. Arb. 64 (1983) 312.
A slender upland swamp species of Papua New Guinea, belonging to the group
of species including P. adinobotrys Merr. & Perry.
PAPUA NEW GUINEA: Southern Highlands Prov., Tari valley, Itipu village,
5600 ft. alt., 6 Sept. 1981, B. C. Stone & C. Rose 15005 (holotype, KLU; isotypes,
BISH, LAE).
A shrub similar to P. adinobotrys but more slender and smaller in all respects
except the longer pistillate peduncles.
Pandanus lustrorum B. C. Stone, J. Arn. Arb. 64: (1983) 313.
This recently described Papuan species is an addition to the small cluster of
species centering around P. adinobotrys Merr. & Perry.
PAPUA NEW GUINEA: West Sepik Distr., Telefomin, alt. 1800 m, wet swamp,
208 Gard. Bull. Sing. 36(2) (1983)
erect pandan to 3 m tall, with many short side branches, leaves grey, fruit pale
orange-brown solitary, Mar. 1975, Womersley & Waikabu NGF 48718 (holotype,
KLU: isotypes, LAE, L, BRI).
Pandanus merrillii Warburg, in Perkins, Fragm. Pl. Philipp. 50. 1904. Martelli,
Philipp. J. Sci. Bot. 3 (1908) 71. Martelli, Webbia 4 (1913) 23, 4 (1914) t. 35-37.
Merrill, Enum. Philipp. Fl. Pl. 1 (1925) 19. Figse 2.056
PHILIPPINES: Palawan; Paragua Island, San Antonio Bay, alt. 1500 ft., on dry
slopes, Feb. 1903, E. D. Merrill 840 (isotypes FI, NY).
Additional collections, all from Palawan; Merrill 7250, 9274 (staminate), 9375,
and Bermejos B.S. 232 (FI).
This species is so similar to P. affinis Kurz that, on the basis of herbarium
material alone, discrimination is virtually impossible. The sole difference worthy of
note seems to be the presence, albeit slight, of minute bristles on the pistillate
peduncle, noticeable in the type collection. The slightly hispidulous peduncle is not,
so far, known in P. affinis, but is found in P. monticola F.v.M. of Queensland,
another (but quite different) species of sect. Acrostigma. Following are some
supplementary descriptive notes on P. merrilli drawn from the isotypes.
Leaves linear attenuate acute, the apex somewhat prolonged, subflagellate, to
162 cm long (and over), 2.8 cm wide; base with sheath about 3.5 cm long; margins
near base with spreading prickles with slightly recurved tips 2-2.8 mm long,
1.5-5 mm apart; near the middle, the marginal teeth slender antrorse, nearly 1 mm
long, 3-6 mm apart; near apex, the teeth antrorse, 0.25 mm long, 1 mm apart.
Midrib near base with stout broad-based retrorse prickles to nearly 3 mm long,
1.5-11 mm apart; farther out along basal fifth of leaf, the prickles recurved,
2 mm long, more remote, to 25 mm apart; near the middle, the teeth scarcely 1 mm
long, antrorse, 10 mm apart; near apex, the teeth similar in size and spacing to those
of the adjacent margin. Apical ventral pleats prickly with antrorse prickles scarcely
0.5-0.9 mm long, irregularly spaced, sometimes only | mm apart, sometimes several
cm apart, extending along the most distal 30 cm of the leaf. Undersurface pale
glaucous. Longitudinal nerves about 67-69 per leaf, dorsally overlain by the minute
punctiform stomatal complexes visible as 3-5 parallel rows of dots. Infructescence
spicate, cephalia 5 (or more?) together, each c. 5.5 cm diameter, the drupes crowded.
Penduncle slightly hispidulous toward apex, or not (?). Drupes c. 27 mm long
(including styles), the corpus 15 mm long, 4-5 mm wide; pileus 3-4 mm long,
produced into a style c. 6 mm long. Staminate inflorescence spicate; spike bearing
apparently solitary stamens, each with a short filament 1 mm long and a long
anther 8 mm long, the apiculus 0.5 mm long. (Staminate description based on
Merrill 9274).
Pandanus microglottis B. C. Stone, Bot. J. Linn. Soc. 85 (1982) 34, f. 7.
This Sarawak species is noteworthy for its polycephalic infructescence and the
Supplement to Revisio Pandanacearum 209
short, Ovate-acute stigmas on similarly short, broad, non-spiniform styles. The
latter feature suggests that the species has an isolated position in the section and may
even deserve a subsectional rank on its own. However, pending the discovery of the
staminate plants, this remains a potential status rather than a clearly deserved one,
and for the time being, it is merely ranked as a species of subsection Acrostigma.
SARAWAK: Mulu National Park (4th Division), Sungei Melinau Paku drainage
area along rocky riverbanks beside Transit Camp 2 at c. 150 m alt., 2 Apr. 1978,
Stone 13654 (holotype, KLU; isotypes, K, SAR).
SUBSECT. Dimissistyli Stone
Pandanus gladiator B. C. Stone, sp. nov. Fig. 4.
Frutex arborescens usque ad 5 m altus, erectus, sparsiter ramosus, strictus; solitarius; corona foliorum
densa; foliis lineari-attenuatis coriaceis munitis, usque ad 300 cm longis, et 8 cm latis, apice acutis breviter
subcaudatis; marginibus in basi dentatis, dentibus rigidis deltoideis patentibus c. 2.5 mm longis et 2-5 mm
sese separatis; in medio, dentibus subappressiter curvatis antrorsis c. 2 mm longis, (3—) 5-9 (-12) mm sese
separatis; apicem versus, denticulis parvioribus, subappressiter antrorse curvatis, klydoniformibus,
densis, c. 1-1.4 mm longis et | mm sese separatis; costa media dorso in basi hamato-aculeato dentibus
reflexis 3-3.5 mm longis, 3-12 mm sese separatis; in medio, denticulis subappressiter antrorsis parvioribus
c. 1.4-1.8 mm longis, (2-) 4-8 (-11) mm sese separatis; apicem versus, denticulis antrorsis gracilibus c.
1-1.2 mm longis et 2-3 mm sese separatis; in cauda, denticulis ad eis in margine adjacenti simillimis;
pagina infra ut videtur pallido et glauco, densiter et obscuriter venosostriatulo; plicibus apicalibus
foliorum ventraliter denticulatis. nflorescentia axillaris, spicata, robusta, bracteata (bracteis delapsis),
polycephala (cephaliis c. 5-7); cephaliis aggregatis, oblato-rotundis, lateraliter subcompressis, c. 9-10.5
cm longis, 10-11 cm latis, receptaculo c. 3-4 x 2.5 cm, drupis numerosis ferenti; pedunculo triquetro
c. 40 cm longo, 2.5 cm crasso, robusto. Drupa (ex loco mediani) c. 40 mm longa; pileo (corpore
styliforme) 25-30 mm longo (stylo incluso); pyrena basali anguste obovoidea 8.9-9.5 mm longa et 3 mm
lata, vel (mesocarpio basali incluso) c. 11 mm longa; stylo (sensu stricto) spiniforme, antrorse curvato
vel recto, 4-6 mm longo, acutissimo; stigmate anguste canaliculati nigro, per totam longitudinem styli;
endocarpio pallido pariete c. 0.5 mm crasso. Cetera ignota.
Type: SULAWESI (Celebes), Mt. Roroka Timbu, west slope, c. 80 km SSE. of
Palu, disturbed lower montane forest c. 35 m high, with little undergrowth, on steep
terrain with shallow clayey soil; solitary ‘*schopfbaum’’ 5 m tall, fruit bright red;
leaf c. 3 m long; trunk hardly branched; 11 May 1979; E. F. de Vogel 5287 holotype,
BO!; isotypes, L! KLU!).
This species fits readily into subsect. Dimissistyli, but is the first species in that
group to present a spicate, polycephalic infructescence, and this seems its principal
distinctive character. However, drupe size is smaller than in P. danckelmannianus
K. Schum. and larger than in P. misimaensis St. John ex Stone; but about the same
as in P. erinaceus Stone and P. lictor Stone. The long pilei of the drupe are relatively
smooth, unlike the coarsely verruculose pilei of P. verruculosus Backer ex Stone.
The species is known so far only from the type collection.
The specific epithet is well known and means ‘‘sword-fighter’’ in allusion to the
sword-like leaves; but also more broadly appropriate because of the various weapons
which the Roman gladiators used, including the mace, to which the fruit-heads bear
210 Gard. Bull. Sing. 36(2) (1983)
a certain resemblance. The species is also a companion of P. lictor, the specific
epthet of which means ‘‘mace-bearer.’’
Pandanus gladiator is the western-most of subsect. Dimissistyli, and probably
indicates the absolute western boundary of this subsection. There is one other
Moluccan species (P. verruculosus), from Halmaheira, but probably a few more
remain to be found. Most of the species are in New Guinea, the Bismarck Archipelago,
and the Solomon Islands.
Pandanus jacobsii B. C. Stone, sp. nov. Fig 5.
Frutex usque ad 2-3 m altus, stipite ad 4 cm crasso, simplex; radicibus gralliformibus nullis. Folia
linearia ad 220 cm longa, 5 cm lata, infra glauca, apicem versus acutata; marginibus in basi dentatis,
dentibus c. 2 mm longis, 1-5 mm sese separatis; in medio, dentibus antrorsis, c. 1 mm longis, 1-5-4 mm
sese separatis; apicem versus, denticulis subappressiter antrorsis c. 0.5-1 mm longis, c. 1.5 mm sese
separatis. Costa media dorso in basi per spatio 15-18 cm inerme, deinde dentibus deltoideis patento-
retrorsis ad 2.5 mm longis, 6-14 mm sese separatis; in medio, costa carinato, denticulis antrorsis 0.7-0.9
mm longis, 1-3 mm sese separatis; apicem versus, denticulis antrorsis 0.5-0.9 mm longis, 1-3 mm sese
separatis. Plicae foliorum apicali-ventraliter armatae, denticulis antrorsis 0.75 mm longis, irregulariter
separatis, sparsis. Inflorescentia axillaris, pedunculo c. 10 cm longo, 14 mm crasso (apicem versus);
cephalio solitario oblato-globoso c. 10 x 11 cm, receptaculo c. 4 cm longo, drupis numerosis composito.
Drupa c. 35 mm longa, pileo c. 20 x 4 mm anguloso, verruculoso-tuberculato, in stylo 6-7 mm longo
antrorse curvato producto, stigmate lineare subaequilongo; pyreno 12-13 mm longo, 4-5 mm lato,
endocarpio 10 mm longo, pariete pallido c. 0.14 mm crasso. Semen c. 8 mm longum. Cetera ignota.
Type: PAPUA NEW GUINEA: Southern Highlands Prov., limestone country
near Waro airstrip 20 km SSW of Kutubu, 6° 31'S, 143° 10’E, at 500-600 m alt.,
single-stemmed pandan without proproots, 2-3 m tall, leaves glaucous underneath,
fruit light red, 15 Oct. 1973, M. Jacobs 9281 (L holotype, 2 sheets!).
Like most species of subsect. Dimissistyli, P. jacobsii has the erect, not or little-
branched trunk, with no or few basal proproots, an axillary inflorescence, the
pistillate peduncle bearing a solitary oblate globose cephalium, and the drupes
ripening red. In P. jacobsii, the drupes are quite small, approaching P. misimaensis
St. John ex Stone, in their dimensions. However, the additional length and the more
coarsely verruculose-tuberculate pilei distinguish P. jacobsii. The marginal prickles
of the leaf apex are moreover smaller in P. misimaensis. Pandanus verruculosus
Backer ex Stone has also the tuberculate pilei intimated by its name, but in this
Halmaheiran species the cephalia are larger (12-15 cm diameter) and leaves much
larger (to 500 x 14cm). Perhaps the most similar species to P. jacobsii is P. erinaceus
Stone, which however is a taller plant (to 5 m), with basal proproots to 1 m long,
and the drupes have a virtually smooth pileus.
The new species is named after the late Dr. M. Jacobs of the Rijksherbarium,
Leiden, in recognition not only of his merit in obtaining the type collection but for
his botanical contributions to the Flora Malesiana and special assistance to the
author.
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9°] Sumtpeydos iy Soovyans yeixege ‘seq Jeo] :q ‘ovjans [erxeqe “uoNtod
ajpprur ‘yea, 39 fsopyoud jeurdsuew jo jiejap “wWopl sg soovjins [eIXepe
‘xode joy 1p — ‘Aaou “ds oU0Ig “DO “qf “OIDIPH/S SNUDpUD, *p BL “1/27
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211
212 Gard. Bull. Sing. 36(2) (1983)
SUBSECT. Alticolae Stone
Pandanus lepatophilus B. C. Stone, Bot. J. Linn. Soc. 85 (1982) 31, f. 6.
A remarkable, small, sprawling decumbent plant, probably a facultative epiphyte,
known so far only from the massive limestone peak Gunung Api in Mulu National
Park, Sarawak. The solitary cephalium is only 15 mm long (sometimes two occur
together), but the size of the fruit at maturity is not known. At the unripe stage the
drupes are only about | cm long.
SARAWAK: Mulu National Part (4th Division), Gunung Api limestone ridge at
c. 1500 m alt., 14 Apr. 1978, Argent & Jermy 973 (holotype, E; isotypes, K, KLU,
SAR).
It should be mentioned that the type species of this subsection, P. a/ticola Holtt.
& St. John, is also known from Sarawak and has in fact also been collected in Mulu
National Park (/van Nielsen 662, in AAU).
Acknowledgement
My thanks are due to E. F. de Vogel for his kindness in showing me his collection of
Pandanus gladiator before final processing at Leiden in 1981, and to the Rijksherbarium
for making a duplicate available for study at KLU in 1983.
Literature Cited
Stone, B. C. (1978) Revisio Pandanacearum, Part I. Pandanus subgenera Coronata
Martelli and Acrostigma (Kurz) Stone. Flora Malesiana, Precursores. Fedn. Mus.
J. n.s. 23: 1-74, Plates I-VI.
STUDIES IN MALESIAN VITACEAE
VII. The genus Tetrastigma in the Malay Peninsula
Aq LATIF
Botany Department, Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia
EFFECTIVE-PUBLICATION DATE: 7TH MAR. 1984
Abstract
A revision of the genus Jefrastigma (Miq.) Planch. in the Malay Peninsula is presented. Twelve species
are here redescribed and assigned to section Tetrastigma and section Carinata sect. nov. They are:
T. pedunculare (Wall. ex Laws.) Planch., T. papillosum (Bl.) Planch., T. pyriforme Gagnep., T. scortechinii
(King) Gagnep., T. dubium (Law.) Planch., T. cruciatum Craib & Gagnep., T. /Jawsoni (King) Burkill,
T. dichotomum (B\.) Planch., T. /anceolarium (Roxb.) Planch., T. hookeri (Laws.) Planch., T. curtisii
(Ridl.) Suesseng., and 7. godefroyanum Planch. Six species, namely, T. papillosum, T. pyriforme,
T. cruciatum, T. dichotomum, T. hookeri, and T. godefroyanum, represent new additions to the flora
of the Malay Peninsula, and two established species names are reduced, namely, 7. kunstleri (King) Craib
(to T. lanceolarium) and T. wrayi (King) Craib syn. nov. (to 7. dubium). A general discussion on the
growth habits and morphology of stem, inflorescence, flowers, fruits and seeds is also given.
Introduction
The largest genus of the Malayan Vitaceae is Tetrastigma (Miq.) Planch., which
comprises 12 species. The genus is characterized by its 4-lobed or 4-cleft stigma in
the pistillate flowers. In the other genera, the stigma is subcapitate (as in Ampelo-
cissus, Cissus, and Cayratia) or entire (as in Pferisanthes). The value of the stigmatic
characteristic of Tetrastigma was first recognised by Miquel (1861) when he included
all the then known species of Vitis (sensu lato) which have a 4-segmented stigma
under section Tetrastigma Miq. Later Planchon (1887) raised Miquel’s section to
generic status, a taxonomic decision which received mixed support from later
taxonomists. Gilg (1896), Gangepain (1910) and Suessenguth (1953), for instance,
recognised Tefrastigma as a genus whereas King (1896) and Ridley (1922) did not.
Ridley (1922) recognised 3 genera of Malay Peninsula Vitaceae viz. Vitis L.,
Pterisanthes B\. and Leea L. ex Van Royen, and treated Tetrastigma as a section
of Vitis, along with Ampelocissus, Cissus and Ampelopsis.
The first complete account of the Malayan Terrastigma was given by King (1896),
an account later adopted by Ridley (1922). Seven species, viz., Vitis pedunculare,
V. scortechinii, V. wrayi, V. lawsoni, V. kunstleri, V. lanceolarium and V. curtisii
were adequately described by Ridley (1922). In a recent study, the author (Latiff,
1978) recognised only six of the species described; the description of V. wrayi King
was found to be based on two species, that of the vegetative parts agreeing with
Cayratia wrayi (King) Craib and that of the reproductive parts with T dubium
which, as a name, has priority over 7. wrayi (King) Craib.
HABIT AND MORPHOLOGY
Growth habits. All members of the Vitaceae except Leea climb by means of ten-
drils. The tendrils are borne opposite the leaves on the long vegetative shoots.
213
214 Gard. Bull. Sing. 36(2) (1983)
Tetrastigma species may be described as large vines; their stems are comparatively
thick, woody and long. Their trailing stems may be found draping trees and shrubs
along roads, river banks and in forest edges, especially at higher altitudes.
Stem. This description of stem morphology is based largely on the study of
herbarium specimens, supported by limited field observations. The stems are striate
and terete when young and later they either remain terete as in 7. pedunculare and
T. pyriforme or become flattened as in 7. /awsoni and T. cruciatum. Stem lenticels
are very prominent; they appear normal as in 7. pedunculare or protrude externally
as corky excrescences, a character diagnostic for T. papillosum.
Leaves. Only one species, i.e., 7. scortechinii has simple leaves, its diagnostic
character. Other species have compound leaves with leaflets ranging from 3 to 7.
However, leaves of 7. cruciatum, T. dubium, and T. dichotomum are predominantly
trifoliolate while on the same stem some are simple. The simple leaves in these are
comparatively larger than any single leaflet of the compound leaf. A few species,
for instance, 7. pedunculare, T. papillosum, and T. curtisii have only trifoliolate
leaves.
In Tetrastigma the leaf-outline is found to be diagnostic for some species. T.
godefroyanum and T. curtisii have obovate terminal leaflets. Specimens of 7.
hookeri, which have widely elliptical terminal leaflets, are distinguishable from
those of 7. lanceolarium which have lanceolate ones.
Inflorescences. The inflorescence in all species is pedunculate. In most of them,
the peduncles are easily observed but in 7. cruciatum and T. lawsoni they are very
short, the inflorescence being almost sessile. The peduncie-length differs in in-
florescences of different sexes. In 7. dichotomum, T. curtisii, and T. lanceolarium
the male inflorescences have a longer peduncle than those of the female. This has
made identification of species based on general morphology difficult. The :in-
florescence in Tetrastigma species is a cyme which is of the following kinds: corym-
bose in 7. papillosum, umbellate in T. scortechinii, T. cruciatum and T. godefro-
yanum, and dichotomous in others.
Flowers. Tetrastigma is dioecious, having pistillate and staminate flowers in
different individuals while all the other genera of Vitaceae have bisexual flowers.
The flowers of Tetrastigma are largely 4-merous but 5-merous ones are occasionally
found. The calyx is small and subcupuliform. The petals are membranous, on the
outside glabrous or slightly pubescent and are strongly reflexed in the bud.
The stigma which is either 4-cleft or 4-lobed in the pistillate flowers is an important
single character for recognizing the genus. It is entire in the staminate flowers. The
four narrow segments of the stigma are orientated in different ways: vertically, as
in T. pedunculare or horizontally as in 7. dubium. In other species, the four
segments are not very conspicuous because the stigmatic surface is densely covered
with simple multicellular hairs or cilia. The ciliate stigma is perhaps an adaptive
feature to ensure the better deposition of pollen grains in dioecious plants like
Tetrastigma. Within the Vitaceae, it is only in 7efrastigma that staminodes are
present in the pistillate flowers. In shape and position, these are suggestive of
Tetrastigma in the Malay Peninsula 215
reduced stamens.
Fruits. The fruits of Vitaceae are berries. In Tetrastigma the degree of pulpiness
varies from species to species. For example, the berries of T. hookeri are very pulpy,
moderately so in 7. scortechinii and T. dubium while those of T. godefroyanum
contain hardly any pulp.
In section TJefrastigma the variation in size, shape and surface features of the
berries is quite useful for species identification. For example, 7. hookeri has the
largest berries (c. 3.5 cm in diameter) and 7. dubium has the smallest (c. 0.9 cm
in diameter) in that section. The dried berries of 7. godefroyanum are smooth and
hard, whilst those of 7. curtisii are wrinkled and soft. Generally, 7. /awsoni and
T. dichotomum are similar but their berries differ in number and shape. The former
has globose berries, usually 1-3 per fructification while the latter has ellipsoid ones,
usually more than 3 per fructification. Within section Carinata the difference in
fruit size is also useful for separating the three Malayan species. The fruits are c.
0.7 cm long in 7. pedunculare, c. 1.4 cm in T. pyriforme and c. 0.9 cm in T.
papillosum.
Seeds. The difference in fruit and seed morphology form the basis for creating
two sections in this genus as described in the latter part of the paper (under tax-
onomic treatment).
Size in general is not useful for distinguishing species even though seeds in T.
pedunculare and T. papillosum are relatively small (0.5 x 0.4 cm). Shape has been
found to be diagnostic for the two sections. In section Tetrastigma the shape is
plano-convex, globose or oblong while in section Carinata it is convex-carinate.
The two sections in 7etrastigma may also be recognised by the differences in their
testa sculpturing or ornamentation. Observations made on all the Malay Peninsular
species show differences in their testa pattern which suggest a grouping into the
following types (Table 1).
Table 1.
Testa types in Tetrastigma.
Type Sect. Tetrastigma Sect. Carinata
1. Ridged T. pedunculare
T. papillosum
2. Tuberculate T. pyriforme
3. Glabrous T. lawsoni
T. dubium
4. Rugulose T. lanceolarium
T. hookeri
T. curtisii
§. Furrowed T. dichotomum
T. cruciatum
T. scortechinii
6. Reticulate T. godefroyanum
216 Gard. Bull. Sing. 36(2) (1983)
Another character which distinguishes the two sections is the position of the
chalazal knot on the dorsal surface of the seeds. In section Tetrastigma the chalazal
knot extends 34- or full-way the dorsal surface while in section Carinata it extends
'4-way the surface. On the ventral surface the infolds are deep and converging in
section Carinata but shallow and parallel in section Tetrastigma.
Host-parasite relationships. Rafflesia arnoldi R. Br., with flowers 1 m across, was
the first parasite reported on Vitaceae. Brown (1842) collected it in Sumatra, growing
on the roots of (what he reported) a species of Cissus. As Tetrastigma is now known
to be the only host of the parasitic Rafflesia, it is very likely that Brown misidentified
the host. In the State of Perak in the Malay Peninsula, 7. scortechinii is said to be
the host of R. hassel/tii Suring. (B. C. Stone, 1978, pers. comm.).
Stirling (1939) reported the occurrence of Sapria himalayana Griff. as a parasite
on the roots of 7. cruciatum. The seeds of this parasitic member of the Rafflesiaceae
are said to be deposited (and to germinate) in cracks in the roots or stems of the
host. Species such as 7. cruciatum, T. scortechinii, T. lawsoni and T. lanceolarium
have characteristic straplike stems which are easily broken or cracked, thereby
providing a suitable niche for the seeds of the parasite. The physiological relation-
ships between these parasites and their hosts have been the subject of a few in-
vestigations (Stirling, 1939; Brown, 1912). Recently, a Rhizanthes species has been
noted to be parasitic on stems of 7. dubium in the State of Pahang (Francis S. P.
Ng, 1982, pers. comm.). Speculation on the evolutionary relationships between
these plants is currently given attention to by W. Meijer (1982, pers. comm.).
Tetrastigma (Miq.) Planch.
Tetrastigma (Miq.) Planchon in DC., Monog. Phan. 5 (1887) 423; Suessenguth in
Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 318; Backer & Bakhuizen v. d. Brink,
Fl. Java 2 (1965) 88.
Vitis sect. Tetrastigma Miquel, Ann. Mus. Bot. Lugd.-Bat. 1 (1862) 72; King, J. As.
Soc. Beng. 65, 2 (1896) 386; Ridley, Fl. Mal. Penin. 1 (1922) 470.
Type species: T. lanceolarium (Roxb.) Planch.
Large, woody, dioecious vines. Stem terete or flattened, conspicuously lenticellate.
Leaves simple or/and compound. Tendrils simple, leaf-opposed, not ending with an
adhesive disc. Inflorescence axillary and pedunculate, a cyme, the cyme umbellate,
corymbose or dichotomous. Flowers 4(5)-merous; unisexual, pedicellate. Calyx sub-
cupuliform to disciform. Petals reflexed in mature flowers. Disc adnate to ovary in
pistillate flowers, 4-lobed, free in staminate flowers. Filaments filiform. Anthers
orbicular to ovate-oblong. Staminodes filiform. Style terete, short. Stigma minute
and entire in staminate flowers, large, 4-lobed, and ciliolate in pistillate flowers. Ovary
undeveloped in staminate flowers, 2-loculate, the locules 2-ovulate each in pistillate
flowers. Berries pyriform to globose. Seeds with dorsal side convex and ridged or
furrowed, ventral side convex or carinate; endosperm T- *, M- 2, m , or ™ -
shaped in cross-section.
Tetrastigma in the Malay Peninsula PA
Distribution. About 95 species, in tropical and subtropical Asia, in Malesia(c. 57),
Taiwan(4), India(12), Thailand(16), and Indochina(22).
Ecology. Occurs in all kinds of habitat from sea-level to high mountains.
Key to the Species
1. Berries pyriform when dry, 3- to 4-seeded; seeds convex-carinate, the chalaza extending /2-way across
the surface, the ventral infolds converging; endosperm 7*-shaped in cross-section
2. Leaves 5-foliolate; berries 0.8-1.0 cm long; seeds 0.7 cm long, the beak inconspicuous ........
eS Stl sd. 5 ave eet SE ULE acc Me SRS Meee ae wees 12. T. pyriforme
2. Leaves 3-foliolate; berries 0.5-0.7 cm long; seeds 0.5 cm long, the beak prominent
3. Terminal leaflet broadly elliptical to rhombic; lenticels of old stems not modified; inflorescence
dichotomous; leaflets with fine glaucous indumentum beneath; staminodes as long as gynoecium
ete thee ae oN stwislcios oe ricclsa ets ce ceca sea cece gcacs sce 10. T. pedunculare
3. Terminal leaflet narrowly elliptical; lenticels of old stems modified to spinelike corky ex-
crescences; inflorescence corymbose; leaflets glabrous beneath; staminodes 2 the length of
(OT ESAT scale aed Bi pe anaes AP ok eae Ie a 11. T. papillosum
1. Berries globose or ellipsoid when dry, 1- to 2-seeded; seeds globose or plano-convex, the chalaza
extending %4-way or more across the surface, the ventral infolds parallel; endosperm M- to 7 -shaped
in cross-section :
4. Leaves simple, pubescent beneath, leaf-base semi-cordate; anthers kidney-shaped .............
Lis: 4e5 55S ee ee NRE epee a etter FL, SCORLCCHITIE
4. Leaves compound (simple leaves sometimes present on the same individual), glabrous beneath; leaf-
base acute or obtuse; anthers orbicular or ovate-oblong
5. Flowers glabrous
6. Leaves 5- to 7-foliolate; seed testa reticulate; endosperm 7% -shaped in cross-section; bark
usually shredding; inflorescence usually borne on lateral branches .................-..---
6. Leaves simple to 3-foliolate; seed testa rugulose; endosperm M-shaped in cross-section; bark
usually glossy; inflorescence on the main stem
7. Stigma-lobe pointed; staminodes '2 length of the gynoecium; seeds 0.6 cm long, the chalazal
knot clavate and extending %4-way across the surface; leaves chartaceous 2. 7. dubium
7. Stigma-lobe rounded; staminodes as long as gynoecium; seeds 1.2 cm long, the chalazal
knot elongate and extending completely across the surface; leaves coriaceous ..........
epee nae Se Care aM Peete Gene Pech esac scare asfals ee 5. T. dichotomum
5. Flowers pubescent
8. Stigma-lobe pointed, glabrous; flowers flat-topped; leaflets with decurrent bases 4. T. /awsoni
8. Stigma-lobe rounded, ciliolate; flowers conically topped; leaflets with acute base
9. Venation between the secondary veins of leaves very prominent; seed testa reticulate;
EuGOSDEEM eA -SAPeCiM CLOSS-SCCLIOM) p< 2 <15)- = 2 is a es ie nln al 8. T. curtisit
9. Venation obscure; seed testa rugulose or furrowed; endosperm yrt-shaped in cross-section
. 10. Leaves membranous, simple to 3-foliolate; seeds obovoid; inflorescence very condensed,
ie remot Bets ge ee OAs Bocas Reais Ge ao A an pe 3. T. cruciatum
218 Gard. Bull. Sing. 36(2) (1983)
10. Leaves coriaceous, 3- to 7-foliolate; seeds ellipsoid; inflorescence lax, 5-13 cm in
diameter
11. Leaflets lanceolate to narrowly elliptical, the margin obscurely dentate; berries 1.5-2.0
cmunkdiameterscedspl2-0 >< O:6nciin ee eee eee eee 6. T. lanceolarium
11. Leaflets very broadly elliptical, the margin dentate; berries 3.2-3.5 cm in diameter;
Seedsils8\-<O/8 Cri Aa a: Goya Cake ea oe eee ee 7. T. hookeri
I. Sect. Tetrastigma
Baccae globose vel ellipsoidae. Semina 1-2, globose vel plano-convexa, postice sulcis rectis parallelis
2 leviter exarata. chalaza per %4 vel totum longitudinis seminis deorsum extensa, antice sulcis; endosper-
mium in sectione transversali M-forme ad 7*-forme.
Type: T. lanceolarium (Roxb.) Planch.
1. Tetrastigma scortechinii (King) Gagnep.
T. scortechinii (King) Gagnepain, Not. Syst. 1 (1911) 376; Suessenguth in Engler &
Prantl, Nat. Pfl. Fam. 20d (1953) 321.
Lectotype: King’s Coll. 2897, Larut, Perak (SING/, isolectotype K/).
Vitis scortechinii King, J. As. Soc. Beng. 65, 2 (1896) 392; Ridley, Fl. Penin. 1
(1922) 474.
Stem 0.9-1.2 cm in diameter (young), flattened, 2.7-3.1 cm wide (old). Leaves
simple 15.4-19.2 x 8.5-10.3 cm, ovate-oblong, acuminate, semicordate at base,
sinuate-bristled, subcoriaceous, glabrous above, pubescent beneath, the petiole
3.4-4.6 cm. Peduncle 2.9-3.1 cm; cyme umbellate. Staminate flowers c. 5 mm long,
oblong, the stigma-lobe rounded. Pistillate flowers c. 6 mm long, oblong, the
stigma-lobe rounded. Berries 0.4-0.6 cm in diameter globose, usually 2-seeded; seed
0.4-0.5 cm in diameter, globose, the endosperm M-shaped in cross-section.
Ecology. On the margin of limestone vegetation and lowland dipterocarp-forest.
This species is believed to be the host for Rafflesia hasseltii Suring. (B. C. Stone
1978, pers. comm.)
Distribution. Known only from collections in Gopeng, Larut and G. Pondok
(Perak), and Sungai Jelai (Pahang).
Observation. This is the only simple-leafed species in the Peninsula, recognised
by its ovate-oblong, pubescent leaves and compact inflorescence.
Specimens examined. PERAK, Goping, April 1884, Wray Jr. 5998 (K. BM);
Larut, March 1882, King’s coll. 2897 (lectotype, SING); G. Pondok, 7.6.1930,
Henderson 23793 (SING). PAHANG, Sg. Jelai, July 1903, Machado s.n. (SING).
Tetrastigma in the Malay Peninsula 219
2. Tetrastigma dubium (Laws.) Planch.
T. dubium (Laws.) Planchon in DC. Monog. Phan. 5 (1887) 437; Suessenguth in
Engler & Prantl., Nat. Pfl. Fam. 20d (1953) 324.
Type: Hooker f. s.n., Sikkim, India (Isotype, K/)
Vitis dubia Lawson in Hooker f. Fl. Brit. Ind. 1 (1875) 661.
Vitis wrayi King, J. As. Soc. Beng. 65, 2 (1896), in part; Ridley, Fl. Mal. Penin.
1 (1922) 475 in part.
T. wrayi (King) Craib, Fl. Siam. Enum. 1 (1926) 314; Suessenguth in Engler &
Prantl, Nat. Pfl. Fam. 20d (1953) 323.
Stem 1.8-2.4 cm in diameter. Leaves simple or digitately 3-foliolate, chartaceous,
glabrous; simple leaves 10.2-12.8 x 4.6-5.2 cm, ovate-oblong, acuminate, rounded
at base, entire, the petiole 7.4-9.8 cm; compound leaves with terminal leaflets
9.8-19.2 x 4.3-10.6 cm, elliptical, accuminate, entire, acute at base; the petiolule
1.6-4.5 cm; lateral leaflets 5.7-17.4 x 3.4-8.2 cm, oblong, oblique, the apex and
margin as in terminal leaflets, the petiolule 0.3-1.4 cm. Peduncle 1.7-2.2 cm; cyme
dichotomous. Staminate flowers c. 3 mm, pubescent. Pistillate flowers c. 3 mm
long, pubescent, the staminode c. 1 mm, the stigma-lobe terete. Berries 0.5-0.8 cm
in diameter, globose, usually 2-seeded; seeds 0.6 x 0.4 cm, globose, the endosperm
M-shaped in cross-section.
Ecology. On the margin of the lowland dipterocarp-forest, and submontane
forest.
Distribution. India, Burma, Thailand and the Malay Peninsula.
Observation. The specimens of the Malay Peninsula have two forms of leaves,
simple and 3-foliolate, but in India variants with (a) 3- and 5-foliolate leaves (on the
Same specimen) and (b) with S-foliolate ones only, are frequently seen.
Specimens examined. PERAK, Larut, in April 1890, Wray Jr. 3945 (K, SING);
Taiping,, 26.1.1923, Henderson 10328 (SING). SELANGOR, Dusun Tua, May
1896, Ridley s.n. (SING); Gombak, 20 m.s., 25.9.1979, Latiff 61 (UKMB).
KELANTAN, Kg Parit, 2.2.1923, Haniff & Nur 10359 (SING). JOHORE, G. Panti,
Dec. 1899, Ridley s.n. (SING). KEDAH, G. Baling, 25 Nov. 1941, Corner & Nauen
s.n. (SING). PENANG, Govt. Hill, 5.4.1968, Sidek 236 (K, SING).
3. Tetrastigma cruciatum Craib & Gagnep.
T. cruciatum Craib & Gagnepain, Kew Bull. (1911) 31; Craib, Fl. Siam Enum.
1 (1926) 312; Suessenguth in Engler & Prantl. Nat. Pfl. Fam. 20d (1953) 320.
Type: Kerr 599, Chieng Mai, Thailand (isotype, K/, BM’).
220 Gard. Bull. Sing. 36(2) (1983)
Old stem flattened, 0.9-1.2 cm thick, 3 cm wide. Leaves simple or 3-foliolate,
membranous, glabrous, the petiole 0.8-2.4 cm; simple leaves 9.5-11.2 x 3.6-5.2 cm,
oblong, acuminate, rounded at base, subentire; compound leaves with terminal
leaflets 7.9-9.2 x 3.7-4.8 cm, elliptical, the apex, base and margin as in simple
leaves, the petiolule 0.3-0.5 cm; lateral leaflets 7.5-8.9 x 3.6-4.3 cm, elliptical,
oblique, decurrent, the petiolule 0.2-0.4 cm. Peduncle 0.1-0.3 in diameter; cyme
umbellate. Pistillate flowers c. 5 mm long, pubescent, the staminode c. 0.8 mm, the
stigma-lobe rounded and ciliolate. Staminate flowers not observed. Berries 0.9 x
0.6 cm, subglobose; seeds 0.7 x 0.5 cm, the testa furrowed, the endosperm M-shaped
in cross-section.
Ecology. On the margin of lowland dipterocarp-forest and mostly on limestone
hills (e.g. Pulau Langkawi). :
Distribution. Thailand and the Malay Peninsula.
Observation. This species is recognized by its very condensed inflorescence,
flattened stem and membranous leaves. The first collection, of Griffith, in Malacca,
was misidentified by Ridley as 7. /Janceolarium. The other collections are from Perlis.
This species constitutes another addition to the flora of the Malay Peninsula.
Specimens examined. MALACCA, sine loc., Griffith 1323 (K). PERLIS, Kaki
Bukit, 11.6.1938, Kiah 35243 (L, SING). KEDAH, P. Langkawi, 11.11.1980, Latiff
290 (UKMB).
4. T. lawsoni (King) Burkill
T. lawsoni (King) Burkill, Dict. Econ. Prod. Mal. Penin. 2 (1935) 2286; Suessenguth
in Engler & Prantl, Nat. Pfl. Fam 20d (1953) 325.
Lectotype: King’s Coll. 6287, Larut, Perak (K/, isolectotypes BM!, SING!).
Vitis lawsoni King, J. As. Soc. Beng. 65, 2 (1896) 394; Ridley, Fl. Mal. Penin. 1
(1922) 474.
Stem flattened, 2.2-2.8 cm wide, 1 cm thick, tuberculate. Leaves 3-foliolate,
coriaceous, glabrous, the petiole 2.2-4.3 cm; terminal leaflets 10.7-13.8 x 4.8-5.8 cm,
elliptical to lanceolate, caudate, decurrent at base, obscurely bristled, the petiolule
1.4-1.8 cm; lateral leaflets 9.4-10.6 x 2.7-3.8 cm, elliptical to oblong, leaf-base
decurrent, the apex and margins as with terminal leaflets. Peduncle 0.4-0.5 cm,
cyme dichotomous. Staminate flowers c. 3 mm long. Pistillate flowers c. 4 mm long,
ovoid, the staminode c. 0.2 mm, the stigma-lobe terete. Berries 1.7-2.0 cm in
diameter globose, 1-3 per fructification; seeds 1.2 x 0.6 cm, oblong, the endosperm
M-shaped in cross-section.
Ecology. On the margin of lowland dipterocarp-forests of the West Coast of the
Malay Peninsula.
Tetrastigma in the Malay Peninsula 221
Distribution. Malay Peninsula.
Observation. This species is superficially quite similar to the 3-foliolate form of
T. lanceolarium but differs in fruit and seed morphology. The fructification usually
has 3 berries.
Specimens examined. SELANGOR, Ulu Gombak, 20.9.1926, Strugnell 12459
(SING); Bangi, 8.12.1981, Latiff PUS 184 (UKMB). PERAK, Larut, May 1884,
King’s coll. 6134 (SING). PENANG, Gardens’ Waterfalls, 5.2.1916, Burkill 1218
(K, SING). KEDAH, Relau, 1.8.1938, Baba 21486 (K, SING), Singapore, sine /oc.,
sine die, Ridley 4649 (K).
5. Tetrastigma dichotomum (BI.) Planch.
T. dichotomum (Bl.) Planchon, in DC., Monog. Phan. 5 (1887) 441; Suessenguth
in Engler & Prantl, Nat. Pfl. Fam. 20 d (1953) 323; Backer & Bakhuizen v.d.
Brink, Fl. Java 2 (1965) 89.
Lectotype: Blume s.n. (L!).
Cissus dichotoma Blume, Bi jdr. 1 (1825) 186.
Stem 0.8-1.2 cm in diameter, lenticellate. Leaves simple or 3-foliolate, coriaceous,
glabrous; simple leaves 8.7-10.2 x 3.4-4.8 cm, elliptical, acuminate, obtuse at base,
serrate, the petiole 2.7-6.1 cm; compound leaves with terminal leaflets 5.7-9.4 x
2.5-4.4 cm, elliptical, acuminate, decurrent at base, serrate, the petiolule 1.5-2.1
cm; lateral leaflets 4.8-9.7 x 2.1-5.5 cm, the outline, apex, base and margin as in
terminal leaflets, the petiolule 0.8-1.0 cm. Peduncle 4.7-5.4 cm; cyme dichotomous.
Staminate flowers obovoid. Pistillate flowers oblong, the staminode c. 0.6 mm, the
stigma-lobe rounded. Berries 1.2-2.2 x 0.6-1.0 cm ellipsoid, usually 1-seeded; seed
1.2 x 0.5 cm, oblong, the endosperm " -shaped in cross-section.
Ecology. On the fringe of hill dipterocarp-forests and submontane forests.
Distribution. Java, Sumatra and the Malay Peninsula.
Observation. This species is recognized by its decurrent and glossy leaflets, and
ellipsoid berries. Its leaves are mostly 3-foliolate but occasionally larger, simple
leaves have been observed on the same plant. In the Malay Peninsula it is only
recorded from Maxwell Hill, Cameron Highlands, Gunung Berumbun, and these
represent additional records for the flora of the Malay Peninsula.
Specimens examined. PERAK, Maxwell Hill, 4.12.1965, Md. Shah & Sidek 1081
(SING, K). PAHANG, Gunung Berumbun, 21 Nov. 1903, Ridley 13554 (SING);
Cameron Highlands, 15.5.1936, Holttum 31262 (L, K, SING). Cameron Highlands,
21.4.1930, Henderson 23627 (K, SING). Genting Highlands, 19.10.1977, Latiff &
Mohd. Kassim s.n. (UKMB).
222, Gard. Bull. Sing. 36(2) (1983)
6. T. lanceolarium (Roxb.) Planch.
Tetrastigma lanceolarium (Roxb.) Planchon in DC. Monog. Phan. 5 (1887) 423;
Suessenguth in Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 323; Backer & Bakhuizen
v.d. Brink, Fl. Java 2 (1965) 89.
Type: Roxburgh 2429 (CAL, water-colour painting at K/)
Cissus lanceolarium Roxburgh FI. Ind. 1 (1820) 430.
Vitis lanceolarium (Roxb.) Wallich in Wight & Arnott, Prod. Fl. Ind. Orient. 1
(1834) 128; King, J. As. Soc. Beng. 65, 2 (1896) 395; Ridley, Fl. Mal. Penin. 1
(1922) 475.
Vitis kunstleri King, J. As. Soc. Beng 65, 2 (1896) 396; Ridley, Fl. Mal. Penin. 1
(1922) 475
Tetrastigma kunstleri (King) Craib, Fl. Siam Enum. 1 (1926) 313; Suessenguth in
Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 325.
Young stem terete, 2.6-4.1 cm in diameter, old stem flattened, 6.0-11 cm across,
tuberculate. Leaves 3-foliolate to pedately S- to 7-foliolate, coriaceous, glabrous,
the petiole 6.7-11.6 cm; terminal leaflets 13.6-20.5 x 4.2-9.4 cm, lanceolate,
acuminate, acute at base, obscurely dentate, the petiolule 0.6-3.3 cm; lateral leaflets
5.8-13.4 x 2.6-6.7 cm, lanceolate, rounded at base, the apex and margin as in
terminal leaflets, the petiolule 0.4-2.5 cm. Peduncle 0.5-2.0 cm; cyme dichotomous.
Staminate flowers c. 2 mm long ovoid. Pistillate flowers c. 5 mm long, oblong, the
staminode c. 0.5 mm, the stigma-lobe rounded, ciliolate. Berries 1.5-2.0 cm in
diameter, globose, 1-2-seeded; seed 1.2 x 0.6 cm, oblong, the testa rugulose, the
endosperm -shaped in cross-section.
Ecology. Mostly on the margin of lowland dipterocarp-forests, frequently on the
margin of limestone vegetation and hill dipterocarp-forests. According to Willem
Meijer (pers. comm.) this is the commonest host of Rafflesia hasseltii.
Distribution. India, Burma, Thailand, Vietnam, Laos, Kampuchea and Malesia.
Observation. In the Malay Peninsula it is the most widely distributed species,
being recorded from various habitats throughout the Peninsula.
Specimens examined. PAHANG, Kuala Lipis, 22.11.1924, Burkill & Haniff
15765 (SING). KEDAH, P. Langkawi, Bt Batu, 5.11.1968, Keng et al. 72 (SING),
Country Club, 14.11.1980, Latiff 279 (UKMB). NEGERI SEMBILAN, Bt Klanera.
7.3.1885, Alvins 935 (SING). JOHORE. Endau, 24.7.1977, Maxwell 77-362
(SING); P. Pemanggil, 21.1.1980, Latiff 122 (UKMB). PERAK, Bota Kiri,
14.3.1958, Md. Shah & Kadim 350 (K, KEP, SING). SELANGOR, Lake Garden,
31.7.1951, Woods 150 (SING); Gombak 13 m.s., 25.9.1979. Latiff 51 (UKMB).
TRENGGANU, Kemaman, 12.11.1935, Corner s.n. (SING). PENANG, sine loc.,
Wallich Cat 6013 (K). SINGAPORE, Bukit Timah, in 1894, Ridley s.n. (SING).
KELANTAN, Bertam Lama, 19.1.1981, Kamaruddin Mat Salleh 104 (UKMB).
Tetrastigma in the Malay Peninsula 223
7. Tetrastigma hookeri (Laws.) Planch.
T. hookeri (Laws.) Planchon in DC., Monog. Phan. 5 (1887) 428; Suessenguth in
Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 325; Backer & Bakhuizen v.d. Brink,
Fl. Java 2 (1965) 89.
Lectotype: Hooker f. 162, Sikkim (K/).
Vitis hookeri Lawson in Hooker f., Fl. Brit. Ind. 1 (1875) 661.
Stem 2.2-2.8 cm in diameter, tuberculate. Leaves pedately 5-foliolate, coriaceous,
glabrous, the petiole 10.7-16.2 cm; terminal leaflets 15.1-25.6 x 7.0-13.4 cm,
broadly elliptical, caudate, acute at base, dentate, the petiolule 2.6-3.7 cm; lateral
leaflets 12.4-22.8 x 5.1-9.4 cm, elliptical, becoming rounded, the apex and margin
as with terminal leaflets, the petiolule 2.2-3.1 cm. Peduncle 2.3-2.5 cm; cyme
dichotomous. Pistillate flowers c. 5 cm long, the staminode c. 1 mm long, the
stigma-lobe rounded, ciliolate. Staminate flowers not observed. Berries 3.2-3.5 cm
in diameter, globose, usually 1-seeded; seed 1.8 x 0.8 cm, ellipsoid, the testa
rugulose, the endosperm »-shaped in cross-section.
Ecology. On the margin of hill dipterocarp-forests.
Distribution. India, Burma, Thailand, Java, Sumatra, Borneo and the Malay
Peninsula.
Observation. The earliest record of this species is a collection of Ridley from
Penang Hill, which was identified as Vitis lanceolarium (synonym of T. /anceolarium).
It is recognised by its largely broadly elliptical and dentate leaflets and large berries.
It is closely related to T. /anceolarium but differs in the character of the leaves, ber-
ries and seeds. This species is another new record for the Malay Peninsula.
Specimens examined. SELANGOR, Ulu Gombak, 21.7.1968, Stone 8320 (K, KLU).
PENANG, Government Hill, March 1915, Ridley s.n. (K). KEDAH, P. Langkawi,
Selat Pancur, 11.11.1980, Latiff 180 (UKMB).
8. Tetrastigma curtisii (Ridl.) Suesseng.
ti Pei (Ridley) Suessenguth in Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 325.
Vitis curtisii Ridley, J. Roy. Soc. Str. Br. 75 (1917) 24; Fl. Mal. Penin. 1 (1922) 474.
Lectotype: Curtis 3363, Penang (K! isolectotype S/NG).
Stem 1.7-2.5 cm in diameter. Leaves 3-foliolate, coriaceous, glabrous, the petiole
3.4-8.6 cm; terminal leaflets 9.1-12.3 x 4.6-6.2 cm, obovate, caudate, acute at
base, subentire, the petiolule 2.4—4.7 cm; lateral leaflets 7.2-9.4 x 3.7-4.5 cm,
oblong becoming obtuse, the apex and margin as with terminal leaflets, the petiolule
1.4-2.2 cm. Peduncle 0.4-0.6 cm, .cyme dichotomous. Staminate flowers c. 4 mm
long, oblong. Pistillate flowers c. 4 mm long, the staminode c. 0.5 mm, the stigma-lobe
224 Gard. Bull. Sing. 36(2) (1983)
rounded and ciliolate, Berries 1.3-1.7 x 0.7-1.1 cm, ellipsoid; seeds 1.2-1.6 x
0.7-0.9 cm, broadly ellipsoid, the testa reticulate, the endosperm * -shaped in cross-
section.
Ecology. On the margin of the hill and lowland dipterocarp-forests.
Distribution. Malay Peninsula.
Observation. This species is recognized by its diffuse venation between the secondary
veins, particularly conspicuous on the upper surface of the leaves.
Specimens examined. PENANG, West Hill, Oct. 1886, Curtis 1051 (K, SING),
Government Hill, March 1915, Ridley s.n. (K). SELANGOR, Ulu Langat, 15.11.1958,
Gadoh 1175 (SING). PAHANG, Raub, 15.4.1970, Soepadmo 629 (KLU). KEDAH,
Bt Perak, 27.11.1969, Chan 13207 (KEP). NEGERI SEMBILAN, Ulu Bendul,
22.12.1979, Latiff 92 (UKMB).
9. Tetrastigma godefroyanum Planch.
T. godefroyanum Planchon in DC., Monog, Phan. 5 (1887) 436.
Type: Godefroy 663, Kampuchea (Isotype, K/).
Stem 1.5-1.8 cm in diameter, the bark glaucous and usually shredding. Leaves
pedately 5— to 7-foliolate, coriaceous, glabrous, the petiole 5.1-8.3 cm; terminal
leaflets 5.2-8.1 x 3.1-4.8 cm, obovate to obovate-oblong, acuminate, obtuse at
base, crenate, the petiolule 2.5-3.2 cm; lateral leaflets 4.8-7.9 x 3.0-4.5 cm, the
outline, apex, base and margin as in terminal leaflets, the petiolule 1.7-2.7 cm.
Peduncle 2.8-3.3 cm; cyme umbellate, usually borne on lateral branches. Staminate
flowers not seen. Pistillate flowers c. 4 mm long, oblong, the staminode c. 0.6 mm,
the stigma-lobe rounded. Berries 1.2-1.5 x 0.9-1.1 cm, ellipsoid, usually 1-seeded;
seed 0.9 x 0.6 cm, the beak prominent, the testa reticulate, the endosperm 7*-shaped
in cross-section.
Ecology. On the margin of lowland dipterocarp-forests.
Distribution. Vietnam, Laos, Kampuchea and the Malay Peninsula.
Observation. This species is recognized by its thinly pulped berries, hard seed and
obovate leaflets which have abruptly caudate apices. This species is a new addition
to the flora of the Malay Peninsula.
Specimens examined. SELANGOR, Ulu Langat, 1.5.1960, Gadoh 2156 (KEP).
KEDAH, P. Langkawi, Selat Pancor, 21.11.1934, Henderson 28948 (K, SING);
P. Dayang Bunting, 27.11.1934, Henderson 18948 (K, SING); Ayer Hangat,
12.10.1970, Chin 494 (KLU), P. Kedrah, 17.11.1941, Corner s.n. (SING), Dalam
Ru, 6.3.1983 Latiff & Rahim s.n. (UKMB); G. Senyum, 30.7.1929, Henderson
22372 (SING).
Tetrastigma in the Malay Peninsula 225
II. Sect. Carinata Latiff sect. nov.
Baccae pyriformes. Semina 3-4, postice carinata et tuberculata, chalaza per 2 longitudinis seminis
deorsum extensa, antice concavitibus divergentibus 2; endospermium in sectione transversali T-forme.
Type species: T. pedunculare (Wall. ex Laws.) Planch.
10. Tetrastigma pedunculare (Wall. ex Laws.) Planch.
T. pedunculare (Wall. ex Laws.) Planchon in DC., Monog Phan. 5 (1887) 438;
Suessenguth in Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 319.
Holotype: Wallich Cat. 6024, Penang (K/, isotype SING!).
Vitis peduncularis Wall ex Lawson in Hooker f., Fl. Br. Ind. 1 (1875) 655; King,
J. As. Soc. Beng. 65, 2 (1896) 393; Ridley, Fl. Mal. Penin. 1 (1922) 474.
Stem 1.8-2.6 cm in diameter, lenticels conspicuous. Leaves digitately 3-foliolate,
coriaceous, glabrous above, densely covered with glaucous fine indumentum
beneath, the petiole 10.1-13.3 cm; terminal leaflets 16.8-22.2 x 9.8-11.4 cm broadly
elliptical to rhombic, acuminate, acute at base, serrate, the petiolule 2.4-3.2 cm;
lateral leaflets 12.0-16.8 x 8.9-11.8 cm, asymetrically oblong, oblique, the apex
and margin as in terminal leaflets, the petiolule 1.8-2.3 cm. Peduncle 1.2-1.8 cm.
Cyme dichotomous, 10-12 cm in diameter often borne on older branches or stem.
Staminate flowers oblong; pistillate flowers oblong, puberulose, the staminodes
c. 1 mm long, the stigma-lobe terete. Berries 0.5-0.7 x 0.4-0.5 cm; seeds 0.5 x 0.4
cm, the testa ridged.
Ecology. On the margin of the hill and lowland dipterocarp-forest, sometimes on
limestone hills.
Distribution. Borneo, Sumatra and the Malay Peninsula.
Observation. This species is recognized by its large dichotomously branched
inflorescence which is often borne on older stems and the broadly elliptical to
rhombic terminal leaflets. In general appearance, it is similar to 7. Jauterbachianum
Gilg of New Guinea but differs in the characters of the inflorescence and the leaflets.
Holotype: Wallich Cat. 6024, Penang (K!, isotype SING/).
Specimens examined. PAHANG, G. Benum, 23.3.1967, Whitmore 3395 (KEP,
SING); Jerantut, 3.3.1967, Whitmore 3144 (KEP); Maran-Jerantut Road,
30.5.1979, Latiff & Md Kassim 1650a (UKMB). KELANTAN, Kuala Rek, 27.1.1923,
Haniff & Nur 10176 (K, SING). JOHORE, P. Pemanggil, 29.1.1980, Latiff 140
(UKMB). SELANGOR, Ulu Gombak, 1.4.1921, Hume 9398 (SING); Genting
Highlands, 10.9.1979, Latiff 19 (UKMB). PERAK, Larut, Dec. 1883, King’s Coll.
5364 (K, SING). MALACCA, Bt Tampin, May 1896, Goodenough 1953 (SING);
NEGERI SEMBILAN, G. Angsi, Feb. 1904, Ridley s.n. (SING); Ulu Bendul,
226 Gard. Bull. Sing. 36(2) (1983)
22.12.1979, Latiff 90 (UKMB). KEDAH, Koh Mai, 7.4.1938, Kiah 35214 (K,
SING).
11. Tetrastigma papillosum (Bl.) Planch.
T. papillosum (Bl\.) Planchon in DC., Monog. Phan. 5 (1887) 429; Suessenguth in
Engler & Prantl, Nat. Pfl. Fam. 20d (1953) 320; Backer and Bakhuizen v.d. Brink, Fl.
Java 2 (1965) 88.
Lectotype: Blume s.n., Java (L! isolectotypes K! BM!)
Cissus papillosa Blume Bijdr. 1 (1825) 183.
Stem 1.4-1.8 cm in diameter, with spinelike corky excrescences. Leaves digitately
3-foliolate, coriaceous, glabrous, the petiole 3.3-7.5 cm; terminal leaflets 8.2-9.7 x
3.6-4.8 cm, elliptical, acuminate, obtuse at base, coarsely crenate, the petiolule
2.7-3.3 cm; lateral leaflets 7.6-8.9 x 3.1-3.6 cm, elliptical, oblique, the apex and
the margin as in terminal leaflets, the petiolule 1.0-1.4 cm. Inflorescence a corymbose
cyme. Staminate flowers pubescent; pistillate flowers c. 2 mm long, pubescent, the
staminode c. 0.5 mm, the stigma-lobe terete. Berries 0.6 x 0.7 cm. Seed 0.5 x 4.0
cm, the testa ridged.
Ecology. On the margin of hill and lowland dipterocarp-forests, rarely on limestone
hills.
Distribution. Thailand, Borneo, Philippines, Celebes, New Guinea, Java, Sumatra
and the Malay Peninsula.
Observation. The epithet papillosum refers to the papillate structures on the old
stem. Strictly speaking these are not papillae but rather, spiny excrescences formed
by the changes in the lenticels. Although Ridley himself collected the species in
Johore, he did not distinguish it from 7. pedunculare. Recently the author
collected some specimens of the species in Frasers’ Hill, Gombak and Genting
Highlands. These constitute additional records to the flora of the Malay Peninsula.
Specimens examined. PAHANG, Cameron Highlands, 10.4.1934, Symington
36201 (KEP); Fraser’s Hill, 25.10.1979, Latiff 77 (UKMB). JOHORE, Sg. Tebrau,
March 1908, Ridley s.n. (SING); G. Panti, in 1892, Ridley 4180 (K, SING); G.
Sumalayang, 19.2.1971, Chin 715 (SING, KLU). SELANGOR, Gap 24.8.1959,
Burkill 1989 (K, SING); Genting Highlands, 25.10.1977, Latiff 36 (UKMB); Genting
Highlands, 10.9.1979, Latiff 18 (UKMB); Ulu Gombak, 20 m.s, 25.9.1979, Latiff
59 (UKMB).
12. Tetrastigma pyriforme Gagnep.
T. pyriforme Gagnepain, Bull. Soc. Bot. Fr. 91 (1944) 35: Suessenguth in Engler &
Prantl, Nat. Pfl. Fam. 22d (1953) 327.
Type: Poilane 25625, Tonkin. Vietnam (isotype, K/).
. Tetrastigma in the Malay Peninsula 225]
Stem 2.0-2.1 cm in diameter, smooth. Leaves pedately 5-foliolate, coriaceous,
glabrous, the petiole 6.5-11.2 cm; terminal leaflets 9-11.2 x 4.6-5.2 cm, elliptical,
acuminate, acute at base, serrate, the petiolule 2.1-2.3 cm; lateral leaflets 7.9-9.3
x 3.8-4.4 cm, asymetrically oblong, oblique, the apex and margin as in terminal
leaflets, petiolule 2.0-2.1 cm. Peduncle 4.1-4.3 cm; cyme dichotomous. Flowers not
observed. Berries 0.8-1.0 cm in length. Seeds 0.7 x 0.5 cm, the testa tuberculate.
Ecology. Known only from the hill forests in Cameron Highlands, Bukit Kemaman
(Trengganu) and Maran-Jerantut Road.
Distribution. Vietnam, Laos and the Malay Peninsula.
Observation. This is the only species in the Malay Peninsula of sect. Carinata with
5-foliolate leaves. This species could not be identified with the literature of King
(1896) and Ridley (1922). It was finally established by comparative studies with type
material of Laos and Vietnam. The species is a new record for the Peninsula.
Specimens examined. PAHANG, Cameron Highlands, 2.4.1937, Md. Nur 32827
(K, SING). TRENGGANU, Bukit Kemaman, 26.11.1935, Corner s.n. (SING).
PAHANG, Maran-Jerantut Rd, 30.5.1979, Latiff & Md Kassim 1650b (UKMB).
Acknowledgements
I am indebted to Prof. D. M. Moore (Reading University) and Dr. B.C. Stone
(Universiti Malaya) for their advice. The data reported here have been partly
extracted from my PhD thesis, accepted by the University of Reading, England, and
partly obtained while holding Universiti Kebangsaan Malaysia Research Grant No.
34/79.
Literature Cited
Brown, R. (1842). An account of a new genus of plants, named Rafflesia. Trans.
Linn. Soc. 13: 201-234.
Brown, W. H. (1912). The relation of Rafflesia manillana to its host. Philip. J. Sci.
(Bot.) 7: 209-224.
Gagnepain, F. (1910). Essai de classification du genre Tetrastigma. Not. Syst.
1: 306-326.
Gilg, E. (1896). Vitaceae. Jn: Engler, A. & K. Prantl. Die naturlichen Pflanzenfamilien
5(3), 427-454. W. Engelmann, Leipzig.
King, G. (1896). Ampelidaceae. Materials for a flora of the Malayan Peninsula.
Disciflorae, 384-419, Royal Botanic Garden, Calcutta.
Latiff, A. (1978). A systematic study of Malesian Vitaceae. PhD thesis, University
ef Reading, England (Unpublished).
228 Gard. Bull. Sing. 36(2) (1983)
Miquel, F. A. G. 1861. Ampelideae. Flora van Nederlandsch Indie. Suppl. 1: 201-202
and 514-518; Suppl. 2: 600-608. Van der Post, Utrecht.
Planchon, J. E. (1887). Monographie des Ampelidées vraires. In: de Candolle, A.
Monographiae Phanerogamarum 5(2), 306—654. Sumptibus G. Mason, Paris.
Ridley, H. N. (1922). Ampelidaceae. Flora of the Malay Peninsula 1: 469-482.
Reeves & Co., London.
Stirling, J. (1939). Notes on the structure of the female flower of Sapria himalayana
Griff. parasitic on the roots of Tetrastigma cruciatum Craib & Gagnep. Publ.
Hartley Bot. Lab., University Press Liverpool.
Suessenguth, C. (1953). Vitaceae. Jn: Engler A. & K. Prantl, Die naturlichen
Pflanzenfamilien 20d, 235-329, Dunker & Humblot, Berlin.
New and Interesting Plant Records for Singapore, II*
J. F. MAXWELL
Parks & Recreation Department, Singapore
Abstract
Plant collecting in Singapore in 1983 has resulted in 3 new records and other species that have not been
recorded here in many years. Brief notes on these plants, with information on their collection localities
and general distribution, are included.
Introduction
Continued floristic work in Singapore during 1983 has resulted in the following
finds: 3 new records and 3 other rare and interesting plants which have not been
collected recently. It is hoped that other botanists will take an active interest in
Singapore’s native flora and continue the work.
Annonaceae
1. Goniothalamus macrophyllus (Bl.) Hk.f. & Thoms.
Fruiting material has been collected from a small, cauliflorous understorey
treelet at Ngadiman Bridge in the Bukit Timah Nature Reserve in a shaded un-
disturbed area. This species is found throughout most of the Malay Peninsula, in
Thailand, Borneo, Java, and Sumatra. In Singapore, previous collections date
1890’s and early 1900’s from the following localities: Tanglin, Serangoon Road,
Sungei Tengah, Tuas, Bukit Mandai, Ang Mo Kio, and Bukit Timah.
Voucher specimen: Maxwell 83-86, 6 Feb. 1983.
Elaeocarpaceae
2. Elaeocarpus acmosepalus Stapf ex Ridley New Record
Ridley’s descriptions (1917, 1922) of this species do not match well with actual
specimens in our Herbarium, collected so far by several persons, as pointed out by
Corner (1939) and do not match my collection. The problem, as Corner (/oc. cit.)
notes, is that the three type collections are not in the Singapore Herbarium. These
are presumably at Kew. The name used with some reserve by Corner for those
specimens in the Herbarium, which match mine is E. floribundus Bl. (1939, 1978).
The specimens are:— Corner 32290 (18 Feb. 1937), C.F. 4120 (Mohd. Yusop, 8
March 1920), and SFN 33969 (Mohd. Nur, | Oct. 1937). R. Weibel (of the Conser-
vatoire et Jardin Botaniques de Genéve), who has recently published descriptions of
* continued from Gard. Bull. Sing. 35: 198, 1982
229
Fig. 1. Prerocymbium tubulatum (Mast.) Pierre. A: branchlet with leaves (Goodenough 1428, Malacca,
Aug. 1893); B: mature fruits (Mohd. Noor 789, Singapore 8 Sept. 1983). Drawn by Mr. Wan Ah
Keong.
Plate 1. Neptunia plena (L.) Bth. Mature pods and seeds; insert: mature inflorescence with staminodes,
and staminate and bisexual flowers. Photograph by Mr. Wan Ah Keong.
230
Plant Records for Singapore, II 231
new species of the same genus for the Malesia Region, has annotated these collec-
tions as E. acmosepalus. The peculiar setae on the tip of the awns of the anthers
are, by the author’s description, quite distinctive a feature for E. acmosepalus. The
anthers in E. floribundus differ in that the tips have no setae.
A fully mature tree 12 m tall was found along the pipeline at Nee Soon on the
partly open margin of the swamp-forest. This species is, apparently, rather rare as
is shown by the meager collections in our herbarium. These are from Selangor,
Pahang and Johore. It is also known from Sarawak.
Voucher specimen: Maxwell 83-25, 13 March 1983.
Gramineae
3. Eriachne triseta Nees ex Steud. New Record
This interesting grass, which is readily recognised by having 2-awned paleas, was
referred by Gilliland (1971) to Massia triseta (Nees ex Steud.) Balansa in his book
on the Grasses of Malaya. There are only a few collections of this species from
Malacca, Pahang, Trengganu, and several specimens from SE. Thailand, Philippines,
Sabah, and northern Queensland in the herbarium. It is also known from Sri Lanka
and Burma.
A very localized group of this grass was found near the Pasir Panjang Nursery on
a partly shaded, lateritic, periodically mown slope.
Voucher specimen: Maxwell 83-27, 22 March 1983.
Leguminosae
4. Derris amoena Bth. var. maingayana (Baker) Prain
Previous collections are:— without locality (Hullet, 1885), the Economic Garden
of our Botanic Gardens (Ridley, 1898) and Siglap (Ridley, 1899). My collection was
from the open margin of the swamp-forest in Nee Soon. The same variety has been
collected in Malacca and Johore. The flowers on my Singapore plant produced
sparse fruit.
Voucher specimen: Maxwell 83-11, 14 Feb. 1983.
5. Neptunia plena (L.) Bth. Plate 1 New Record
I found several individuals of this hardy, decumbent shrub with spreading
branches and bright yellow inflorescences along West Coast Road in an open waste
232 Gard. Bull. Sing. 36(2) (1983)
area. Aside from having two types of flowers, this species lacks thorns and spines.
It, therefore, would be a very desirable species to plant on Singapore’s sandy
reclaimed land, where it will surely thrive. This species, which is native to tropical
America and the West Indies, produces viable seeds here which have been success-
fully propagated at the Pasir Panjang Nursery.
Voucher specimen: Maxwell 83-2, 7 Jan. 1983.
6. Pterocymbium tubulatum (Mast.) Pierre Fig. 1
Winged fallen fruits of this species, a canopy tree of primary forests, was collected
by Mohd. Noor from one which is in the Fern Valley in Bukit Timah Nature Reserve.
It is the second ever collected in Singapore, Ridley 6435 in 1894, from the same
Reserve, being the first and it bears flowers. This species is apparently rare and is
known from Malaya (Kelantan, Malacca, and Johore), Singapore, Sumatra, Sabah,
Sarawak, and Kalimantan.
Voucher specimen: Mohd. Noor 789, 8 Sept. 1983.
Literature Cited
Corner, E.J.H. (1939). Notes on the Systematy and Distribution of Malayan
Phanerogams, III. Gard. Bull. Str. Settl. 10: 317.
. (1978). The Freshwater Swamp-forest of South Johore and Singapore.
Garda Bull Supp aS
Gilliland, H.B. (1971). Flora of Malaya III, Grasses of Malaya. Government
Printing Office, Singapore.
Ridley, H.N. (1917). New and Rare Malayan Plants. J. Str. Br. Roy. As. Soc. 75: 8.
. (1922). Flora of the Malay Peninsula 1: 314. L. Reeve, London.
Lobaria clemensiae Vain. (Lobariaceae, Lichenes)
on Halmaheira Island, Indonesia
WILLIAM SM. GRUEZO
Museum of Natural History and Department of Life Sciences,
College of Arts and Sciences, University of the Philippines
at Los Banos, College, Laguna, Philippines.
Abstract
Lobaria clemensiae Vain., a lichen species, originally described from the Philippines, is reported from
the Halmahera Island, Indonesia.
During my work on the Philippine material of the genus Lobaria (Schreb.)
Hoffm., it has become necessary to study Lobaria specimens from the neighbouring
countries particularly those collected from within the natural phytogeographical
unit known as Malesia. And, from the abundant specimens obtained on loan from
the Herbarium Bogoriense (BO), Indonesia, one collection represents an interesting
phytogeographical record for Lobaria clemensiae Vain.
Lobaria clemensiae Vain.
Lobaria clemensiae Vain., Philipp. J. Sci. 8C: 136. 1913, ‘‘clemensae’’; Yoshimura,
J. Hattori Bot. Lab. No. 34: 262. 1971, fig. 15q-w, pl. 7a-e. Lectotype (designated
by Yoshimura, 1971): Philippines, Mindanao, Lake Lanao, Camp Keithley, June 1907,
Mary Strong Clemens 1313 [TUR-V 10688, part of the lectotype in hb. Gruezo 00-7
ex TUR-V; US (not seen, fide Yoshimura 1971)].
Thallus adpressed, small to medium-sized, 2-4 (-10) cm wide, strongly lacinulate;
lobes 2-10 mm broad, their margin crenulate, often distinctly fringed with small
simple or forked lobules; dorsal surface of thallus smooth, partly canaliculate,
yellowish grass-green to yellowish brown, without soredia or isidia, instead with
lobules formed from cracked cortex; ventral surface of thallus pale to yellowish
brown, thickly tomentose and sparsely rhizinate at midportion of lobes; tomentum
dark chocolate brown to nearly black, crowded on older lobes, pale brown, scattered
near apex or margins; rhizines simple, pale brown, 2 mm long. Apothecia absent.
Pycnidia not seen. Plate 1, A & B; Fig. 2.
Dorsal cortex paraplectenchymatous, c.30 y» thick; medulla 170-220 yp thick,
medullary hyphae 3 » wide; ventral cortex paraplectenchymatous, pale brown, 15
p thick, composed of 2-3 cell-layers; tomentum elongate, branched, loosely inter-
woven, 5-7 » wide, up to 300 p» long. Inner cephalodia often present.
Chemical reactions: thallus K-, C-, KC-; medulla K-, C-, KC+ rose-red.
Chemical substance: gyrophoric acid.
Specimen examined: HALMAHERA ISLAND. Mt Sembilan (Siu), Bivak Ake
233
234 Gard. Bull. Sing. 36(2) (1983)
Biaur, alt. c.550 m, on tree, 10 October 1951, P. Groenhart 8336 (BO 7307).
Habitat: L. clemensiae is usually found tightly adnate on bark of trees from
200-1300 m altitude in lowland to midmontane virgin or mixed dipterocarp forests,
and up to the mossy forest zone.
Distribution range: In the Philippines, L. clemensiae has been collected from: 1,
Tandul Mati, Naujan, Mindoro Oriental Province; 2, Sitio Manlangco, Sibulan,
Negros Oriental Province; 3, Mt Kampalili, Davao Province; 4, Tungao and
Florida, Butuan City, Agusan Province; and 5, Camp Keithley, Lake Lanao, Lanao
Province, the type locality (Gruezo, 1979). (Fig. 1).
Outside of the Philippines, L. clemensiae is known only from the following
collecting stations: 1, near Mt Silam and Kundasan, Sabah (Borneo) and 2, Mt
Gede, Tjibodas and Mt Tantyjar, all in Java (Yoshimura, 1971). Its discovery in the
Halmaheira Island (Mt Sembilan) extends the distribution range further southeast
(Fig. 1). Very likely, this species is expected to be found in New Guinea and its near-
by islands.
HALMAHEIRA
w *
aS
ee oS a
800 KM |
Fig. 1. Geographical range of Lobaria clemensiae Vain.
Mn a
A
ung
Plate 1. Lobaria clemensiae Vain. A. Dorsal surface of thallus. B. Ventral surface of thallus. (scale:
small divisions in mm). (Groenhart 8336).
i cm.
Fig. 2. Lobaria clemensiae Vain. Thallus showing position of lobules. (Groenhart 8336).
235
236 Gard. Bull. Sing. 36(2) (1983)
The diagnostic features of L. clemensiae are: the presence of very fragile lobules
(phyllidia) along the lobe margins and rims of cracked cortex of the thallus (fig. 2);
the more or less veined type of tomentum on the lower surface of the thallus
(plate 1, B); the presence of gyrophoric acid; and the thallus being comparatively
small, thin and fragile, and usually tightly adnate to the substratum.
Incidentally, Yoshimura (1971) described as new Lobaria clemensiae Vain. var.
crassa based from a single collection (S. Kurokawa 6280, TNS; isotype, NICH) from
the Western Highland district of New Guinea. However, comparison of plate 7 a-e
(L. clemensiae Vain. var. clemensiae) and plate 7f (L. clemensiae Vain. var. crassa
Yoshim.)(Yoshimura 1971, p.343) suggests that when additional materials of var.
crassa are obtained and studied, the latter taxon might prove to be a distinct species.
For the time being, var. crassa is distinguished from the typical variety by its thicker
dorsal cortex (50 » vs. 30 ») and the K+ thalline reaction (Yoshimura 1971).
Acknowledgements
I thank Dr. E. Kuswata Kartawinata, Keeper of the Herbarium Bogoriense (BO),
Indonesia for the loan of Lobaria specimens; Professor Juan V. Pancho, former
Director, Museum of Natural History, University of the Philippines at Los Banos
(UPLB) for facilities placed at my disposal; and Dr. Ricardo M. Lantican, Director,
Institute of Plant Breeding, UPLB College of Agriculture for the Institute’s
Fellowship during which tenure this study was undertaken. I also thank Mr. Carlito
S. Sanchez for drawing figures 1 and 2.
For the genuine concern in the plight of Philippine lichenology and his generous
assistance on many occasions, I specially thank Dr. Reino Alava, Curator Emeritus,
The Herbarium, Institute of Biology, University of Turku, Finland.
Literature Cited
Gruezo, W. Sm. (1979). The Lichen Genus Lobaria (Schreb.) Hoffm. in the
Philippines. Unpublished M.Sc. Thesis, Univerisity of the Philippines at Los
Banos, 214 p., 80 figures.
Yoshimura, I. (1971). The Genus Lobaria of Eastern Asia. J. Hattori Bot. Lab. No.
34: 231-364. 27 figures, + 28 plates.
Influence of Growth-Regulating Chemicals on
Hippeastrum hybridum hott.
S. K. BHATTACHARJEE
Indian Institute of Horticultural Research, Bangalore-560 080, Karnataka, India
Contribution No. 178/82 of I.I-H.R.
Abstract
The effect of bulb-dip application of each of the following - chlormequat chloride (CCC), daminozide
(B-Nine), etephon (Ethrel), maleic hydrazide (MH), triiodobenzoic acid (TIBA), indole-acetic acid
(IAA), gibberellic acid (GA;) and naphthylacetic acid (NAA) - was studied on growth, flowering and
bulb formation of Hippeastrum hybridum. CCC and B-Nine induced early sprouting of bulbs, delayed
flowering, increased the size and longevity of flowers, accelerated bulb production and improved the
quality of bulbs in size and weight. Longer flower spikes were obtained with B-Nine-treated bulbs. Ethrel
and MH delayed sprouting, but induced early flower-bud appearance and increased longevity of flowers.
Beneficial effects on bulb formation with Ethrel was also recorded, while MH, NAA and TIBA were
ineffective. IAA and GA; promoted vegetative growth, induced early flowering, stimulated flower size
and stalk length, yielded a greater number of flowers per stalk, extended longevity of flowers, increased
bulb production, and improved the size and weight of bulbs.
Introduction
Hippeastrum hybridum is one of the well known bulbous ornamental plants,
which produces attractive lily-like flowers on long stalks. This most popular spring-
and summer-flowering plant is commercially grown in the Netherlands and exported
to different countries. Synthetic growth-regulating chemicals are being increasingly
used to manipulate growth and flowering of bulbous plants. These chemicals offer
a broad range of effects, both morphological and physiological. Stem elongation of
Lilium cultivars has been effectively reduced by soil drench of CCC (Pearse, 1972),
ancymidol (Dicks and Rees, 1973), Ethrel (Wade, 1972) and Phosphon (chlor-
phonium chloride) (Stuart ef a/., 1961). Bhattacharjee et a/. (1976) reported that in
dahlia, treatment with GA; increased the number of flowers, and with IAA and
GA;, promotion of shoot length. Application of IAA increased the corm weight of
gladiolus, whereas GA; treatments increased the weight of cormels (Winkler, 1969).
In the present investigation an attempt has been made to evaluate the effectiveness
of different growth-regulating chemicals on growth, flowering and bulb formation
of Hippeastrum hybridum.
Materials and Methods
Experiments were conducted at the Institute in Hessarghatta, Bangalore during
1981-1982. Hippeastrum hybridum bulbs of uniform size and weight were soaked
for five hours in different growth regulating chemicals prior to planting in
September 1981. The untreated bulbs of the control were soaked in distilled water.
The chemicals used were CCC and B-Nine at 1000, 2500 and 5000 ppm of each;
Ethre!, MH and TIBA at 500, 1000 and 2000 ppm of each; and IAA, GA; and NAA
at 10, 100 and 1000 ppm of each. The soaked bulbs were then dried in the shade
237
238 Gard. Bull. Sing. 36(2) (1983)
in open air for eighteen hours. Thereafter the bulbs were planted in 25-cm earthen-
ware pots, one in each, in a pot mixture of one part each of sand, soil, leaf mould
and farm yard manure. A mixed fertilizer at the rate of 25 gms per pot was also
added, irrespective of the treatment. The experiment was laid out in a completely
randomised design, involving twenty-five treatments. Five pots were kept for each
treatment. Observations were taken on vegetative growth, flowering and bulb
formation. Data on bulb formation was collected by carefully removing the bulb
and bulblets at the cessation of vegetative growth, twelve months after planting.
Results and Discussion
Sprouting of bulbs. In both the control and the treated ones there was 100%
sprouting with all the treatments including the untreated control. Days taken for
sprouting of the treated bulbs varied with the chemicals and their concentrations.
At all concentrations, B-Nine and CCC significantly induced advanced sprouting by
12.2 to 20.0 days over the control. The lowest concentration of Ethrel, GA; and
NAA also exhibited earlier sprouting of bulbs by 3.4 to 5.2 days. Treatments with
MH and TIBA, and NAA at 1000 ppm delayed sprouting to a great extent. For
inhibition of sprouting of storage tubers of potato and bulbs of onion, MH is largely
used (Skoog, 1980). Junges (1964) and Rudinicki ef a/ (1976) reported early sprouting
with the application of GA; in Freesia hybrida and tulips respectively (Table 1).
Vegetative growth. It was observed that the highest concentration of Ethrel and
NAA, and treatments with MH and TIBA markedly retarded leaf length. Maximum
retardation was obtained with TIBA at 2000 ppm, and the percentage of retardation
was 37.09 over the control. The lowest concentration of CCC, B-Nine and NAA,
all concentrations of GA; and IAA at 100 and 1000 ppm promoted leaf length
appreciably. Promotive effect was greater with GA;, and the highest accleration,
exceeding 27.33 per cent over the control, resulted with GA; at 1000 ppm. Applica-
tion of MH inhibited growth of Croft Easter lilies (Struckmeyer and Beck, 1952).
Dicks and Rees (1973) observed height reduction in Lilium with Ethrel at 1000 ppm.
Promotion of plant height with GA; in Mid-Century Hybrid Lily was reported by
Dicks et al (1974). The number of leaves per plant increased significantly with CCC
at 1000 ppm, GA; and IAA at 100 ppm of each. An increase in the number of leaves
of Hippeastrum hybridum with the application of IAA and GA; was also reported by
Bose et a/. (1980). Though the number of side shoots per plant due to the treatments
with growth-regulating chemicals had not affected significantly, treatments with
CCC at 1000 ppm, and GA; and IAA at 100 ppm of each considerably stimulated
the production of a greater number of the side shoots (Table 1).
Bulb yield. The application of chemicals increased bulb yield, in general, except
with NAA at 100 ppm, IAA at 1000 ppm, TIBA at 500 and 2000 ppm. A significant
rise in the production of bulbs was recorded with all concentrations of CCC,
B-Nine, GA;, IAA and MH. The two higher concentrations of Ethrel, two lower
concentrations of IAA, and NAA at 10 and 1000 ppm, also showed appreciable
improvement in bulb production (Table 1).
Growth-regulating chemicals on Hippeastrum hybridum 239
Bulb weight and diameter. The total weight of bulbs varied with the chemicals and
concentrations. Among different chemicals only the two higher concentrations of
TIBA, and NAA at 1000 ppm significantly reduced the bulb weight. Increase in the
total weight of bulbs varied from 2.69 to 72.31 per cent. over control with the treat-
ments of CCC, B-Nine, Ethrel, GA;, NAA, IAA and MH. Maximum beneficial
effect was seen with the two lower concentrations of CCC, B-Nine and GA;. Diameter
and weight of the largest bulb markedly increased with all concentrations of CCC
Table 1
Effect of Growth Regulating Chemicals on Vegetative Growth and Bulb Formation of Hippeastrum Hybridum hort.
Control :
CEC 1000 53.1 ( 15.18)
cc 2500 49.4 ( 7.16)
CEC 5000 47.4 ( 2.82)
B-Nine 1000 52.3 ( 13.45)
B-Nine 2500 48.3 ( 4.77)
B-Nine 5000 49.3 ( 6.94)
Ethrel 500 44.2 ( —4.12)
Ethrel 1000 42.4 ( —8.03)
Ethrel 2000 39.4 (—14.53)
MH 500 41.4 (—10.19)
MH 1000 36.0 (—21.90)
MH 2000 35.0 (—24.08)
TIBA 500 36.2 (—21.48)
TIBA 1000 32.0 (-—30.59)
TIBA 2000 29.0 (—37.09)
IAA 10 49.3 ( 6.94)
IAA 100 A3%4. | 652:5; (13:45)
IAA 1000 48.8 53:3:.\( 15:62)
GA, 10 40.0 5522 ( 19:73)
GA, 100 42.0 STS (. 247)
GA, 1000 46.2 S8ih ( 2733)
NAA 10 41.8 SLAG 173)
NAA 100 50.0 47.4 (2.82)
NAA 1000 77.8 35.4 (—23.21)
‘F-test -* ~* ** N.S -_* ** ** **
S.Em 1.97 1.91 0.43 | 0.34} 0.29 4.94 0.16 4.65
L.S.D. at 5% 5.54 5.39 1.19 - 0.81 13.90 0.44 13.07
* * — highly significant: N.S. — non significant.
Figures in the parenthesis indicate the percentage of increase or (—) decrease over control.
240 Gard. Bull. Sing. 36(2) (1983)
and B-Nine, GA, at 10 and 100 ppm. Halevy and Shilo (1970) reported that Ethrel
on gladioli increased the yield of cormels. Application of GA; in Begonia increased
the tuber weight (van Onsem and Haegeman, 1961). In tulips, Mohamed and
Fawzi (1980) recorded an increase in the weight of mother and daughter bulbs.
Flowering. Advanced flower-bud appearance, by 22.0 to 79.8 days, was induced
by treating with Ethrel, GA;, IAA and MH. Application of CCC, B-Nine and NAA
Table 2
Effect of Growth Regulating Chemicals on Flowering of Hippeastrum Hybridum hort.
&
NE oi
Control
CEE 6.95)
Gee 13.58)
CEE 14.57)
B-Nine 50.00)
B-Nine 28.81)
B-Nine 39.75)
Ethrel i 5 2 ( —9.32)
Ethrel (— 20.19)
Ethrel (— 16.52)
MH ( = 592)
MH (— 17.55)
MH (—23.50)
TIBA (— 32.78)
TIBA =
TIBA -
IAA ( — 3.64)
IAA ( 19.21)
IAA ( 33.44)
GA, ( 15.56)
GA, ( 29.80)
GA, p : 9 ( 25.49)
NAA ( —8.61)
NAA (— 16.89)
NAA (— 16.89)
‘F’-test
S.Em ; 1.26
L.S.D. at 5% : SSW)
* * — highly significant.
Figures in the parenthesis indicate the percentage of promotion or (—) retardation over control.
Growth-regulating chemicals on Hippeastrum hybridum 241
delayed fiowering by 40.4 to 68.0 days. Treatment with TIBA at 1000 and 2000 ppm
completely suppressed flowering. Flower-bud opening was also affected by the
chemical treatments. All concentrations of GA; and NAA, and the two lower
concentrations of IAA resulted in the early opening of flower buds. Flower-bud
opening was delayed in MH and TIBA treatments (Table 2). Jansen (1960) reported
early flowering in cyclamen with GA;. Will (1977) observed that B-Nine delayed
flowering in gloxinia. It was observed that treatment with CCC at 2500 and 5000
ppm, B-Nine and GA; at all concentrations, and IAA at 100 and 1000 ppm
accelerated the flower-stalk length significantly, and the resulting acceleration from
these treatments was 13.58 to 50.0 per cent. Higher concentrations of Ethrel and
MH, and TIBA at 500 ppm reduced flower-stalk extension. Bose ef al (1980)
reported increase in flower-stalk length of Hippeastrum with IAA, GA; and CCC
at 10 and 1000 ppm. Application of CCC, B-Nine, GA; and IAA, in general, and
MH at 500 and 1000 ppm appreciably increased the length and diameter of flowers.
The highest number of flower spikes per plant resulted with treatment with GA; at
100 ppm. The number of flowers per stalk also varied with the treatments, and the
application of GA; and IAA, in general, showed the most beneficial effect.
Treatments with the higher concentrations of CCC and all concentrations of B-Nine
also induced a greater number of flowers per spike. The longevity of the first-opened
flower was optimal with MH treatments. Treatments with CCC, B-Nine, Ethrel,
GA, and IAA also extended the life of the first-opened flowers to a significant extent
(Table 2). Sheehan and Joiner (1964-1965) in Lilium longiflorum reported an
increase in the number of flowers with the application of CCC. Increased flower
number and size in Hippeastrum with GA;, IAA and CCC was also observed by
Bose et al., (1980).
Acknowledgement
The author is grateful to Dr. K. L. Chadha, Director, Indian Institute of
Horticultural Research, Bangalore for providing the necessary facilities to carry out
the investigation.
Literature Cited
Bhattacharjee, S. K., T. P. Mukhopadhyay and T. K. Bose (1976). Interaction of
auxin and gibberellin with growth retardants on growth and flowering of Dahlia
variabilis and Malvaviscus conzattii. Indian Agriculturist 20: 193-199.
Bose, T. K., B. K. Jana and T. P. Mukhopadhyay (1980). Effect of growth
regulators on growth and flowering in Hippeastrum hybridum hort. Scientia
Horticulturae 12: 195-200.
Dicks, J. W. and A. R. Rees (1973). Effects of growth regulating chemicals on two
cultivars of Mid-Century Hybrid Lily. Scientia Horticulturae 1: 133-142.
Dicks, J. W., J. McD. Gilford and A. R. Rees (1974). The influence of timing of
application and gibberellic acid on the effects of ancymidol on growth and
flowering of Mid-Century Hybrid Lily cv. ‘Enchantment.’ Scientia Horticulturae
2: 153-163.
242 Gard. Bull. Sing. 36(2) (1983)
Halevy, A. H. and R. Shilo (1970). Promotion of growth and flowering and
increase in contents of endogenous gibberellin in gladiolus plants treated with
growth retardant, CCC. Physiol. Plant. 23: 820-827.
Jansen, H. (1960). The application of gibberellins to cyclamen. Gartenwelt 60:
231-232.
Junges, W. (1964). Effect of rindite and gibberellin on dormancy in corms, cormels,
bulbils and seeds of Freesia hybrida. Arch. Gartenb. 12: 295-310.
Mohamed, B. R. and A. F. A. Fawzi (1980). Tulip production as affected by
chlormequat chloride (CCC) treatments. Annals of Agricultural Science,
Moshtohor. 12: 243-247.
Onsem, J. van, and J. Haegeman (1961). Trials on the effect of gibberellins on
growth and tuber weight in Begonia hybrida. Rev. Agric. Brux. 14: 263-272.
Pearse, H. L. (1972). Beautiful as pot plants - dwarfed lilies. Farming in South
Africa 48: 75-76.
Rudinicki, R. M., J. Nowak and M. Saniewski (1976). The effect of gibberellic acid
on sprouting and flowering of some tulip cultivars. Scientia Horticulturae 4:
387-397.
Sheehan, T. J. and J. N. Joiner (1964-1965). Effects of watering frequencies and
dwarfing chemicals on growth, flowering and chemical composition of Lilium
longiflorum ‘Georgia’. Proc. Flo. St. hort. Soc. 77: 520-523.
Skoog. F. (1980). Plant growth substances. Proc. of the 10th International Con-
ference on plant growth substances. Madison, Wisconsin, July 22-26, 1979:
398-399.
Struckmeyer, B. E. and G. E. Beck (1952). The effect of MH on growth and flower-
ing of Croft Easter lilies. Proc. Amer. Soc. hort. Sci. 59: 542-548.
Stuart, N. W., D. L. Gill and M. G. Hickman (1961). Height control of forced
Georgia Easter lilies. The Exchange 136: 22-23, 42.
Wade, D. R. (1972). Ancymidol drench gives good control of lily height and shape
in Lee Valley EHS pot plant trials. Grower 77: 1383-1384.
Will, H. (1977). Gloxinia, the use of growth regulators. Gb + Gw 77: 709.
Winkler, G. (1969). Investigations on the effect of growth substances on the corm
yleld of gladioli. Arch Gartenb. 17: 325-340.
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INDEX
Volume 36
Basionyms and synonyms appear in italics. Page numbers in italics indicate the presence of illustra-
tions. For the index of pages 177-183 please see pages 184 and 185.
Acampe, longifolia 50
penangiana 50
Achasma, macrocheilos 73
megalocheilos 73
Achras zapota 104
Acrospelion 125, 126, 128
Adenoncos, major 50
parviflora 51
sumatrana 51
virens 51
Adenum, coetaneum 113
obesum 113
Aegiceras corniculatum 108
Aerides, decumbens 60
odoratum 51
Agrostis maxima 38
Agrostophyllum, bicuspidatum 51
hasseltii 51
majus 51
pauciflorum 61
Aidia cochinchinensis,
leaf-opposed inflor. 201-204
axillary in origin 202, 203
development 204
flowering habit 202
a monopodium 202
Aira, antarctica 126, 130, 131
spicata 131
subspicata 131
Allamanda, cathartica 113
violacea 113
Alpinia nutans 74
Alstonia, angustifolia 113
angustiloba 113
lucida 113
scholaris 113
spathulata 113
Alyxia, lucida 113
reinwardti var. lucida 113
Amesiodendron 19
chinense 19, 20, 2/
Amomun, biflorum 73
maingayi 75
testaceum 73
triorgyale 73
Ampelocissus, ascendiflora 123
polystachya 123
spicifera 124
thyrsiflora 123
Anacardiaceae 187
Andropogon, aciculatus 33
amaurus 37
annulatus 35
caricosus 35
prionodes 33
wightianus 34
Andropogonae 141
Anisophyllea, architecture and
phyllotaxis 3
Gisticha 3, 4; 5:6, 7, 9
adaptive success 16
leaf anatomy 10, //
leaf morphology 8
nodal anatomy 12
optimal leaf 17
average internode length 13
phyllotaxy 14
as compared with Selaginella
martensii 17
scortechinii 5
Anodendron candolleanum 113
Anthoxanthum, angustum 126, 134
horsfieldii auct. non Mez 134
redolens 126, 131
Aphania 5
mutica 33
Apluda, mutica 33
varia var. intermedia 33
Appendicula, anceps 51
cornuta 51
majus 51
purpurascens 51
torta 52
undulata 51
Arachnis, flos-aeris 52
moschifera 52
Ardisia, corolata 108
crassa 108
crenata 109
crispa 109
elliptica 109
lanceolata 109
littoralis 109
ridleyi 109
singaporensis 109
stylosa 108
teysmanniana 109
tuberculata 109
villosa 109
Areca, horrida 69
triandra 67
Arenga, hookeriana 67
243
244
Arenga cont.
pinnata 67
westerhoutii 68
Arthraxon prionodes 33
Arundinella furva 126, 134 (4)
Aschochilus hirsutus 62
Asclepias curassavica 118
Ascochilopsis myosurus 52
Asparagus racemosus 39
Aspidoptreys concava 122
Asplenium, acrocarpum 26
fuliginosum 26
forma acrocarpum 26
lomariaceum 27
lugubre 26
porphyrorachis 28
Athyrium, acrocarpum 26
fuliginosum 26
longissimum syn. noy. 26, 27
lugubre 26
merrillti 27
porphyrorachis 28
symmetricum 26
Avena 139
flavescens 128
Avenula 139
Axonopus compressus 33
Bassia kingiana 104
Beaumontia multiflora 113
Boerlagiodendron 101
orientale 101
puniceopolleniferum 101
reburrum 102
russullensis 102
tetrandrum 101
Borassodendron machadonis 68
Borassus machadonis 68
Bromus, avenaeformis 129
bifidus 129, 130
ovatus Cav. 128
Bulbophyllum, apodum 52
clandestinum 53
concinnum 52
fenestratum 52
flammuliferum 52
lilacinum 52
membranaceum 53
pulchellum 53
punctatissimum 52
rupicolum 52
sessile 53
Calamus, balingensis 68
concinnus 68
densiflorus 68
ornatus, var. ornatus 68
var. horridus 68
ornatus sensu Griff. 68
siamensis 68
siamensis var. malaianus 68
Gard. Bull. Sing. 36 (1983)
Calotropis gigantea 118
Calpicarpum oppositifolium 115
Carex, bilateralis 131
brachyathera 131
curta 131
duriuscula 131
finitima 131
michauxiana 131
Carissa carandas 114
Caryota mitis 68
Catharanthus roseus 114
Catimbium speciosum 74
Cayratia wrayi 213
Ceratostylis pendula 54
Cerbera, manghas 114
odollam 114
Ceropegia woodii 118
Chlorophytum, malayense 39
orchidastrum 39
Chonemorpha, fragrans 114
macrophylla 114
Chrysophyllum, cainito 104
lanceolatum 104
roxburghii 104
Chrysopogon, aciculatus 33
collinus 33
fulvus 33, 34
orientalis 34
wightianus 34
Cissus, dichotoma 221
lanceolarium 222
quadrangularis 124
Cleisostoma, kunstleri 61 |
latifolium 61
Coelogyne, asperata 54
pallens 54
pandurata 54 |
rochussenii 54 |
Coelorhachidastrae 138
Coelorachis 138
mollicoma 153
Coix lacryma-jobi 34
Combretocarpus 5
Corbyas mucronatus 54
Corymbis, brevistyles 54
longiflora 54
rhytidocarpa 54
Corymborchis, brevistylis 54
rhytidocarpa 54
veratrifolia 54
Corysanthes mucronata 54
Costus, globosus 74
kingii 74
speciosus 74
velutinus 74
Crossotylis 16
Curculigo latifolia 38
Index
Curtisina 19
penangensis 19
Cymbidium, acutum 54
dayanum 54
finlaysonianum 55
Cymbopogon calicicola 40
Cynanchium ovalifolium 118
Cynosurus indicus 36
Cypripedium /owi 60
Dacryodes longifolia 19
Dendrobium, acerosum 55
aloifolium 55
atropurpureum 55
eulophotum 55
euphlebium 56
excavatum 55
farmeri 55
gemellum 55
gracile 56
indivisum 55
langkawiense 55
leonis 55
planibulbe 55
pumilum 55
salaccense 55
secundum 56
serra 55
setifolium 56
spurium 56
subulatum 56
tetrodon 65
Dendrocalamus dumosus 34, 35
Dendrocolla alba 64
Derris amoena var. maingayana 231
Deschampsia sp. 133
Desmotrichum, fimbriatum 56
luxurians 56
Dichanthium, annulatum 35
caricosum 35
mucronulatum 35
Digitaria, chinensis sensu Ridley 35
violascens 35
Dimocarpus 19
Diospyros, argentea 106
bilocularis 107
buxifolia 106
clavigera 106
diepenhorstii 106
discolor 106
ferrea 106
kaki 106
maingayi 107
microphylla 106
lanceifolia 107
lucida 107
pilosauthera var. oblonga 107
pyrifera 106
styraciformis 107
venosa 107
Diplazium 25, 26, 17
Diplazium cont.
acrocarpum 26, 27
altum syn. nov. 27, 28
banahaoense 26
cumingii 27
egenolfioides 25, 26
fuliginosum 26, 27
harrisonii 27
lomariaceum comb. nov. 27, 28, 29
longissimum 26
merrillii syn. noy. 27, 28
plicatile 56
porphyrolepium syn. nov. 27, 28
porphyrophyllum syn. nov. 27
porphyrorachis 28, 29
var. murudense 29
sorzogonense 29
symmetricum comb. nov. 26
Dipodium pictum 56
Dischidia, albiflora 118
bengalensis 118
coccinea 118
cochleata 118
collyris 118
gaudichaudii 119
hirsuta 118
major 118
nummularia 119
rafflesiana 118
singaporensis 119
Disarrenum 126
antarcticum 126, 131
Donax grandis 39
Drosera rotundifolia subsp. bracteata 131
Dyera costulata 114
Elaeocarpus, acmosepalus 229, 231
floribundus 229, 231
Eleocharis attenuata 131
Eleusine indica 36
Embelia, amentacea 109
canescens 109
coriacea 109
garcinifolia 109
lampani 110
ribes 110
ridleyi 109
Ephemerantha, fimbriata 56
luxurians 56
Epidendrum sessile 53
Erica, citrina 56
leiophylla 56
leptocarpa 56
minutiflora 61
nutans 56
pannea 56
pendula S6
pulchella 56
rigida 56
vestita 57
Eriachne triseta 231
Erioglossum 19
245
246
Ervatamia, corymbosa 116
divaricata 116
malaccensis 116
Eschweileri Zipp. ex Boerl. 101
Eulalia, lanipes 38
praemorse 337
quadrinervis 36
Eulophia keithii 57
Euphoria 19
Fagraea, auriculata 111
crenulata 111
fragrans 111
gigantea 111
racemosa 111
ridleyi 111
Fimbristylis dictyocolea 131
Finlaysonia, maritima 119
obovata 119
Freesia hybrida, early sprouting 238
Freycinetia, pollen 166
Galphimia glauca 123
Ganophyllum falcatum 19
Ganua, kingiana 104
motleyana 104
sessilis 104
Gastrochilus curtisii 73
Genianthus maingayi 119
Geodorum citrinum 57
Germainia capitata 131
Glenniea 19
Globba
albiflora 74
var. aurea 74
aurantiaca 75
fasciata 74
patens 75
Glycosmis 19
Glyphochloa 139
Goniothalamus macrophyllus 229
Goodyera hispida 57
Graphephorum 128
melicoideum 128
Gynotroches 16
Habenaria carnea 57
Helictotrichon 125, 139
sempervirens 125
Hemarthria pratensis 131
Hernandia, pollen 165
Heterolytrum 126
Heteropholis 143
benostii 144, 146
cochinchinensis /44, 147
var. cochinchinensis 147, 149
var. chenii stat. & comb. nov. 149, 1/49
nigrescens 144, 147
sulcata 144, 145
Heterophyllum 16
Gard. Bull. Sing. 36 (1983)
Hierochloé 126, 130 (16)
Hippeophyllum scortechinii 58
Hiptage sericea 123
Hornstedtia, macrocheilos 73
megalocheilos 73
metriocheilos 73
triorgyalis 73
Hoya, coriacea 119
coronaria 119
diversifolia 119
finlaysonii 119
lacunosa 119
latifolia 119
obtusifolia 120
parasitica 120
ridleyi 120
Hydrilla verticillata 38
Hypericum gramineum 131
Hypoxis, pollen 164
Iguanura, corniculata 69
polymorpha 69
Imperata cylindrica var. major 36
Isachne langkawiensis 36
Ischaemum, aristatum sensu Ridley 36
beccari sensu Ridley 35
indicum 36
macrarum 36
timorense 36
Jasminum, bifarium 121
griffithii 121
longipetalum 121
rex 121
sambac 121
Joinvillea, pollen 164
Kaempferia elegans 75
Kibatalia maingayi 114
Kingiella, decumbens 60
kingli 57
kingii quoad Ridley 57
Kopsia, fruticosa 114
singaporensis 114
Labisia, pothoina 110
pumila 110
Lepisanthes 19
Leptaspis urceolata 36
Leuconotis griffithii 115
maingayi 115
Leucopogon malayanus 103
Liberbaileya, gracilis 69
langkawiensis 69
Licuala, modesta 69
wrayi 69
Limnanthemum indicum 112
Linociera, ramiflora 122
pauciflora 122
Liparis, caespitosa 58
comosa 58
compressa 58
Index
Liparis cont.
disticha 58
gibbosa 58
Litchi 19
chinensis 22, 22, 23
subsp. chinensis 22, 23
subsp, javensis 23
subsp. philippinensis 23
Livistona, cochinchinensis 69
rupicola 69
saribus 69
Lochnera rosea 114
Lowia longiflora 39
Lucuma malaccensis 106
Maba, buxifolia 106
venosa 107
Maclurodendron magnificum 93
Madhuca, kingiana 104
malaccensis 104
sericea 104
Malaxis, calophylla 58
compressa 58
latifolia 58
micrantha 58
reniloba 58
Malleola, dentifera 58
undulata 59
Malpighia coccigera 123
Malpighiaceae, emendated 122
Manilkara zapota 104
Manisuris 139
myuros 141
nigrescens 147
sulcata 145
Maxburretia, gracilis 69
rupicola 69
Melicope 93
Melicope sensu stricto 94
Melicope species:
curranii 98
helferi 98
indica 98
mindanaensis 98
monophylla 98
platynema 98
simplex 98
suberosa sp. nov. 93-100, 95, 96, 99
unifoliolata 98
Melodinus micranthus 115
Microsaccus, ampullaceus 59
brevifolius 59
javensis 59
Microstylis, calophylla 58
congesta 58
flavovirides 58
micrantha 58
reniloba 58
Mimusops elengi 104
Mnesithea 138
laevis 138
pubescens 151, 153
Musa, acuminata 40
malaccensis var. minor 40
truncata 40
zebrina 40
Myriophyllum, propinquum 131
verticillatum 131
Myrsine, avensis 110
umbellutata 110
Napeodendron altissimum 19
Nepenthes
dentata sp. nov. 197-200; 198, 199
tentaculata 200
villosa 200
Neptunia plena 230, 231
Nerium indicum 115
Neyraudia reynaudiana 37
Norrisia, major 111
malaccensis 111
Nothocissus 124
Nyctanthes arbor-tristis 122
Nymphoides indica 112
Nypa, pollen 168
Oberonia, anceps 59
calcicola 59
caudata 59
dissitiflora 59
flava 60
lunata 59
spathulata 60
transversiloba 60
Ochrosia borbonica 115
Odontadenia, macrantha 115
speciosa 115
Olea, brachiata 122
maritima 122
Oncosperma horridum 69
Ophiuros 138
cochinchinensis 147
corymbosus auct. non Gaertn. 152
var. pubescens 151
exaltatus 141
monostachyus 148
pollockii 151
pubescens (Benth.) Dom. 151
pubescens auct. non. Dom. 160
shimadanus 148
undatus 148
undulatus 148
Oplismenus compositus 37
Orchidaceae, key to limestone spp. 43
Orchidantha, ca/carea 39
longiflora 39
Orchis triplicata 53
Osmoxylon 101
247
248
Osmoxylon cont.
orientale comb. nov. 101
puniceopolleniferum comb. nov. 101
reburrum comb. nov. 102
russellense comb. nov. 101
tetrandrum comb. nov. 101
Otophora 19
Palaquium, bancanum 105
gutta 105
hexandrum 105
microphyllym 105
obovatum 105
oxleyanum 105
ridleyi 105
rostratum 105
xanthochymum 105
Palm genera, pollen 164
Palms, key to limestone, 66
Pandanaceae
pollen record, 163, 1/69
distribution map 167
reports of fossil pollen 169
Pandaniidites, pollen
radicus 165, 168
rhenanus 168
texus 168
Pandanus 163-169 passim, 205
subgen. Acrostigma
sect. Acrostigma 205
subsect. Acrostigma 205-209
subsect. Alticolae 212
subsect. Dimissistyli 209-211
subgen. Coronata 205
Pandanus species:
adinobotrys 207
affinis 208
alticola 70
alticola, pollen 163
aquaticus, pollen 163
basedowii, pollen 163
calcicola 70
danckelmannianus 209
doraistigma, pollen 164
ellipsoideus, 205, 206
epiphyticus, pollen 163
erinaceus 209, 210
fascicularis 71
furcatus, pollen /70
gladiator sp. noy. 209
helicopus, pollen 163
irregularis 71
jacobsii sp. nov. 210, 2//
jiulianettii, pollen 164
ketele 207
lepatophilus 212
lictor 209
longicaudatus, pollen 168, /7/
lustrorum 207
matthewsii, pollen 164
merrillii 206, 208
microglottis 208
misimaensis 209, 210
Gard. Bull. Sing. 36 (1983)
Pandanus cont.
monticola 208
mozambicus, pollen 168, /7/
odoratissimus 71
pollen 164, 168, /7/
piniformis 71
recurvatus 71
?shiabensis, pollen 165
tectorius 71
pollen 168, /7/
vandra, pollen 168, /7/
verruculosus 209, 210
xanthochymum 105
Panicum, compositum 37
sarmentosum 37
Paphiopedilum, barbatum 66
lowii 60
niveum 60
Parameria, glandulifera 115
polyneura 115
Paranephelium chinense \9
Parsonia helicandra 115
spiralis 115
Parvotrisetum 125
Paspalum, cochinchinense 147
conjugatum 37
platycaulon 33
Payena, havilandii 105
lucida 105
maingayi 105
obscura 105
sessilis 104
Pedilonum secundum 56
Peliosanthes violacea 39
Pellacalyx 16
Pergularia minor 120
Peumus, pollen 165
Phaeomeria maingayi 75
Phalaenopsis, cornu-cervi 60
decumbens 60
Pharus urceolatus 36
Phoenicimon 19
rubiginosus 19
Pholidota imbricata sensu Lindl.
Phreatia, microtidis 61
minutiflora 61
Phrynium cylindricium 40
Physostelma wallichii 120
Pimelandra wallichii 109
Planchonella, maingayi 106
obovata 106
Plantago, asiatica 111
major 111
Plectocomia griffithii 70
Plectocomiopsis ferox 68
Plumbago, auriculata 110
capensis 110
indica 110
Index
Plumbago cont.
rosea 110
zeylanica 110
Plumeria, acuminata 115
acutifolia 115
alba 116
Poaephyllum pauciflorum 61
Podochilus, lucescens 61
microphyllus 61
tenuis 61
Poga 5
Pogonatherum, paniceum 37
sacharoideum 37
Pollinia quadrinervis 36
Polygonanthus 5
Polypodium subserratum Hook. 28
Polystachya, flavescens 61
penangensis 61
siamensis 61
Polytrias amaura 37
Pomatocalpa, kunstleri 61
latifolium 61
naevatum 61
setulense 66
spicatum 62
Pouteria malaccensis 106
Preptante, rubens 53
vestita 53
Pterisanthes 213
cissoides 123
eriopoda 123
Pteroceras, ciliatum 62
hirsutum 62
tanyphyllum 62
Pterocymbium tubulatum 230-236
Rafania, avensis 110
umbellulata 110
Remusatia, pollen 164
Renanthera, elongata 62
histrionica 62
Renantherella histrionica 62
Rhododendron, indicum 102
longiflorum 103
Robynsiochloa 138
Rostraria 125, 139
Rottboellia 138
corymbosa 141
subvar.
brassii 158
cochinchinensis 147
pubescens 151
rariflora 160
Rottboelliastrae 137
Rupestrina 128
pubescens 128
Rutaceae 93
sect. Astorganthus 98
sect. Brombya 98
sect. Entoganum 98
Rutaceae cont.
sect. Tetractomia 98
tribe Tetractomia 98
Rynchelytrium repens 37
Saccharum, paniceum 37
repens 37
Saccolabium, hobsonii 62
hortense 61
latifolium, var. parviflorum 61
var. strictum 61
longifolium 50
machadonis 62
minimiflorum 50
miserum 64
myosurus 52
perpusillum 63
pubescens 61
rugulosum 62
sacculatum 63
saxicolum 53
scortechinii 63
secundum 63
setulense 66
sylvestre 59
tenuicaule 65
undulatum Ridl. 59
undulatum quoad Carr 58
uteriferum 168
Sapria himalayana 216
Saranga, pollen 166
Sarcanthus, machadonis 62
rugulosus 62
sacculatus 63
scortechinii 63
subulatus 63
termissus 63
Sarcochilus, ciliatus 62
hirsutus 62
tanyphyllus 62
Sarcolobus globosus 120
Schizostachyum elegans 34
Schoenorchis micrantha 63
Schumanniophyton problematicum,
sympodial inflor. 201, 203
Sciaphila asterias 71
Selaginella 16
martensii 16, 17
Semecarpus, curtisii 192, 196
rufovelutinus 192
trengganuensis sp. nov. 192, 195, 196
Setaria, palmifolia 37
plicata sensu Ridley 37
Sideroxylon, ferrugineum 105
maingayi 106
Smilax, pollen 164
Spathoglottis hardingiana 63
Spinamonoporites typicus, pollen 165
Stachynanthus, pollen 165
Stachyphrynium cylindricum 40
249
250
Stapelia nobilis 120
Staurochilus fasciatus 63
Stauropsis gigantea 65
Stemona, curtisii 71
tuberosa 71
Stenotaphrum helferi 38
Stephanotis, floribunda 120
maingayi 120
Stigmaphyllon ciliatum 123
Strophanthus, brevicaudatus 116
?caudatus 116
dichotomus 116
?singaporianus 116
Strychnos axillaris 112
ignatii 112
malaccensis 112
ovalifolia 112
pubescens 112
ridleyi 112
Styphelia malayana 103
Styrax, benzoin 107
crotonoides 107
Swintonia, floribunda 196
var. penangiana stat. nov. 196
penangiana 196
Symphyogyne gracilis 69
Symplocos, adenophylla 107
barringtonifolia 107
celastrifolia 108
fasciculata 108
odoratissima 108
rigida 107
rubiginosa 108
Tabernaemontana, corymbosa 116
divaricata 116
malaccensis 116
Tacca, leontopetaloides 71
pinnatifida 71
Taeniophyllum, culiciferum 63
filiforme 63
macrorhizum 64
obtusum 64
serrula 64
Tainia plicata 65
Telosma cordatum 120
Terminalia catappa 111
Tetractomia 98
Tetrastigma 213
Malayan species 213-227
sect. Carinata sect. nov. 215, 216, 225, 227
sect. Tetrastigma 213-218
Tetrastigma species:
cruciatum 214, 216, 219
curtisii 124, 214, 215, 223
dichotomum 214, 215, 221
dubium 213, 214, 215, 216, 219
godefroyanum 214, 215, 224
hookeri 215, 223
kunstleri 222
Gard. Bull. Sing. 36 (1983)
Tetrastigma species cont.
lanceolarium 124, 216, 220, 221, 222
lauterbachianum 225
lawsoni 214, 216, 220
papillosum 214, 215
pedunculare 214, 215, 225, 226
pyriforme 214, 215, 226
scortechinii 214, 215, 216, 218
wrayi 213, 219
Thaumastochloa 144, 150
brassii 158, 758, 159
chenii 149
cochinchinensis 147
forma cochinchinensis 147
forma shimadana 148
constricta 151
major 154, 154, 155
major x pubescens /52, 153
monilifera sp. nov. 153, 154, 159
pubescens (Benth.)
C.E. Hubb 151, 1/52
pubescens auct. non C.E. Hubb /54
rariflora (F.M. Bailey) C.E. Hubb /58, 159, 160
rariflora auct. non C.E. Hubb 154
rubra sp. nov. 157, 157, 159
shimadana 148
striata sp. nov. 156, 157, 159
Thecostele alata 64
maculosa 64
zollingeri 64
Thelasis, carinata 64
decurva 64
elongata 64
micrantha 64
succosa 64
triptera 64
Thevetia, neriifolia 116
peruviana 116
Thrixspermum, album 64
amplexicaula 64
lilacinum 64
Thunia alba 66
Thysanolaena, agrostis 38
maxima 38
Toxocarpus griffithii 120
Trichaeta 128
ovata 128
Trichoglottis, fasciata 63
misera 64
retusa 65
winkleri var. minor 65
Tricholaena rosea 37
Trichotosia vestita 57
Triraphis madagascariensis
sensu Ridley 37
Trisetaria 125, 139
Trisetum 125-135, 139
subg. Heterolytrium 126
subg. Trisetum 126
sect. Sibirica 126
sect. Trisetaera 126
Index 5)
Trisetum, subg. Trisetum cont. Tropidia curculigoides 65
sect. Trisetum 126 Tupistra grandia 39
Trisetum species: Tylophora, asthmata 120
antarcticum 126, 130, 131, 134, 135 squarrosa 121
var. densum 134 tenuis 121
biaristatum 129
bifidum 126, 129
forma contractum 129
Tylostylis rigida 56
Typha angustifolia 72
var. biaristatum 129 Uncifera tenuicaulis 65
var. macranthum 129 Urceola, brachysepala 116
var. oshimense 129 lucida 116
var. papillosum 129 maingayi 117
var. viride 129 malaccensis 117
flavescens 125, 126, 128, 130 torulosa 117
var. bifidum 129
var. macranthum 129
var. papillosum 129
Vanda gigantea 65
Vandopsis gigantea 65
formosanum 132 Vallaris maingayi 114
latifolium Ridl. 126, 134 Vitaceae: emendata 124
melicoides 128 Vitis 213
ovatum 128 sect. Tetrastigma 213
parviflorum 125 species:
pratense 125, 128 curtisii 213, 223
sibiricum ssp. papillosum 129 dubia 219
spicatum (L.) Richt. 126, 128, 131 hookeri 223
var. formosanum 132 kunstleri 213, 222
subsp. alaskanum auct. non lanceolarium 213, 222, 223
Hulten 132, 133 lawsoni 213, 220
subsp. australiense 126, 135 macrostachya 124
subsp. formosanum comb. & stat. nov. peduncularis 213, 225
132, 133 quadrangularis 124
subsp. himalaicum 133, 135 scortechinii 213, 218
subsp. kinabaluense 133, 1/33 wrayi 213, 219
subsp. mongolicum 135
subsp. ovatipaniculatum 135
subsp. toluccense var. barbatipaleum 135
Walsura neuroides 19
Willughbeia, apiculata 117
spicatum auct. non Richt. 132, 133, 134 coriacea 117
Striatum 125 firma \17
subspicatum (L.) Beauv. 126, 131 flavescens 117
subspicatum auct. non Beauv. 132, 134 tenuiflora 117
taquettii 129 Wrightia religiosa 117
Tristellateia australasiae 123 Zanthoxyleae 94
Tristira Radlk. 19 Zingiber spectabile 75
Tristira sensu Ridley 19
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8.
PARKS AND RECREATION DEPARTMENT
PUBLICATIONS FOR SALE
The Gardens’ Bulletin, Singapore (Series IV).
Vols. 13-36, 1949-1983.
Price.
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. Selected Plants & Planting for a Garden City - Forty Shrubs, $1.20.
. Selected Plants & Planting for a Garden City - Forty Climbers, $3.00.
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. Malayan Orchid Hybrids by M.R. Henderson and G.H. Addison, $15 (1969).
. A Revised Flora of Malaya.
(a) Vol. 1, Orchids, by R.E. Holttum, $50 (3rd ed. 1980 Impr.).
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Boletus in Malaysia by E.J.H. Corner, $50 (1972).
Items 1-5 can be purchased from the Commissioner, Parks & Recreation Department,
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