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GARDENS’ BULLETIN"

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Leo W.H. Tan, B.Sc., Ph.D. (Singapore) | b

S.Y. Geh, B.Sc., M.Sc. (Singapore); Dip.Hort.Sc. (Massey)

Syed Yusoff Alsagoff

W.K. Tan, B.A. (Wms Coll., Mass.); M.Sc. (M. SU., Mich.);

Ph.D. (U.M., FI.)

Lawrence L.C. Leong, B.Sc., M.Sc., Ph.D. (Malaya) |

Jennifer Ng-Lim Cheo Tee, B.Sc., M.A. (Dublin)

The late R.E. Holttum

Hsuan Keng

A.N. Rao Sips Sal

B.L. Burtt kc ee

‘ " :

se St hs i

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oad

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THE GARDENS’ BULLETIN

SINGAPORE

VOL. 43 199]

CONTENTS

STONE, Benjamin C.:

New and Noteworthy Malesian Myrsinaceae, VI.

eh ine Mrenusel VINCHNGHGLG A DIC i, co iods bades jectenn oe son chs oka) sos ohio aig ests 1-17

WEST, John A.:

New Algal Records from the Singapore Mangroves .................::c::::eeeeseeeeeeees 19-21

KOCHUMMEN, K.M.:

Notes on the Systematy of Malayan Phanerogams

ES Se ee Bh a RR aC OA Aa tal gM ean Soh ad Gees + a eee a 23-26

LIM-HO, Chee Len, PHUA, Lek Kheng, LOW, Neok Chein

and GOH, Chong Jin:

Rename WU RUT 401 PCP ICG AUIS, occas vanes nccccsnsoscnccceaceees-ceecncdacsnereerceetsaqeceseees 27-38

WONG, K.M.:

Schizostachyum terminale Holtt., An Interesting

ee ee aESIT Uk OER T IDL SURE TACO 0 oe 2 Lo ois Sianeli dbwesnnyndes's?-ansiseeren- oe 39-42

Published by

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SINGAPORE 1025.

New and Noteworthy Malesian Myrsinaceae, VI.

Revision of the Genus Hymenandra A.DC.

BENJAMIN C. STONE

Department of Botany

B. P. Bishop Museum

P.O. Box 19000-A

Honolulu, Hawaii 96817

USA

EFFECTIVE PUBLICATION DATE: 23 MAR 1992

Abstract

Hymenandra A.DC., with eight species, is revised. Four new species are proposed and one species

originally placed in Ardisia is transferred. The genus is subdivided into two subgenera, Hymenandra and

Lacrimopila subg. nov. (the first with 6, the second with 2 species). A key to the species, new descriptions,

illustrations, and a list of exsiccatae examined are included.

Introduction

Hymenandra A.DC. (1841), first established as a section (Sectio 2) of Ardisia (De

Candolle 1834) was based entirely on Ardisia hymenandra Wall., first described in

1824. The genus was monotypic and was accepted as such by Mez in his monograph

(1902). It was not until 1958 that a second species was attributed to the genus when

Furtado (1958) transferred Ardisia iteophylla Ridl. 1924 to it. In “1975” (1976) Nayar

and Giri described another species from Burma. Subsequently there have been no

further reviews of the genus.

Furtado (1958) described Ardisia calcicola from Kalimantan. After re-examination

of the type material, this too is found to be a species of Hymenandra. The most

significant character of the genus, emphasized by De Candolle, is the staminal tube

in which the anthers are laterally connate by the thecal margins. Moreover, the in-

florescence is lateral either on a short and much reduced branch with one or two small

but otherwise normal leaves, or on a peduncle subtended by foliaceous bracts crowded

so as to form almost an involucre. The flowers, and the individual floral organs, are

often unusually slender and elongated.

The relationship of Hymenandra appears to be with Ardisia, and especially with

subgenera Pyrqus and Crispardisia, at least in regard to the position and form of the

inflorescence. It seems hypothetically possible that intermediate forms could form

a transition between Ardisia and Hymenandra. The other generic character, more

emphasized by Mez, is the ovular number, which in Hymenandra is “few” i.e. about

5-12. This too conforms with the situation in Ardisia subgenus Crispardisia, but not

with subg. Pyrqus. The tendency to lateral inflorescences is accompanied in several

species by the occurrence of papillae (suberect glandular trichomes) on the interior sur-

face of the calyx-lobes, which is also quite noticeable in Ardisia subgenus Crispardisia.

The genus Hymenanadra is divided here into two subgenera. Six species are retained

in the typical subgenus, and these agree with H. wallichii, the type species, in possess-

ing mostly obovate to oblanceolate leaves with rather short broad petioles; stature with

Z Gard. Bull. Sing. 43 (1991)

the main stem simple or very sparsely branched; and most also have the inner surface

of the calyx-lobes rather densely and minutely papillose. In addition, these species have

mostly oval to distinctly oblong glands in the perianth and stamens.

In subgenus Lacrimopila, the habit is more ramified, with the branch bases ampliate

(gussetted) as in Ardisia, and the plants may reach 3 m (10 ft.) height. The leaves are

elliptic to ovate, and distinctly petiolate. The inner surface of the calyx-lobes is not

papillose, but may be hirtellous. In one species, H. iteophylla, the inflorescence axes

and calyx are lepidote with stelliform to flabellate somewhat stalked scales, but the

anthers and ovary are glabrous. In the other species, H. diamphidia, the inflorescence

axes and vegetative innovations, the calyx on both surfaces, the backs of the corolla-

lobes, the anthers on both sides, and the ovary and style base, are all hirtellous with

shortly stalked ferrugineous hairs with twinned apical cells of a form resembling a tear-

drop (the subgeneric name being derived from this feature). In this species there are

usually 8 or 9 ovules in two series.

It is apparent that the two species assigned to subgenus Lacrimopila are rather more

different from each other than the species of subgenus Hymenandra are from one

another. Moreover there is a geographic difference, with H. iteophylla known so far

only from the Malay Peninsula, while H. diamphidia is only known from Sabah.

Possibly these two species should be distinguished further by establishing another

taxon for H. iteophylla alone.

As here construed Hymenandra should no longer be considered an exclusively

mainland Asian genus but may be termed an Indo-Malesian genus, with its centre

of diversity in Borneo. Five of the eight species are in fact confined to Borneo.

Hymenandra wallichii remains as a Himalayan species known from Sikkim, Bangla-

desh, Assam, and northwestern Burma, while H. iteophylla is the least known and is

restricted to the southernmost part of the Malay Peninsula. The most recently described

species, H. narayanaswamii Nayar & Giri, is known so far only from Burma.

All the species are more or less poorly represented in collections. Several species are

known only from single collections and the rest from a very few. However, Hymenandra

wallichii is represented in cultivation in some botanic gardens (e.g. Edinburgh). Other-

wise, further study will depend largely on the acquisition of newly collected materials.

Generic Distinctions

The tubular androecium is the chief feature on which the genus Hymenandra was

founded. In fact, the filaments are free from just below the anthers to the point where

each is adnate to the corolla-tube, so that the androecial tube is partly interrupted.

The connation of the anthers is minimal and not very tenacious but is quite real; yet

if the anthers were not connate, it would be difficult to distinguish these species from

Ardisia on technical grounds. They do, however, have certain habit characters in

common. The species in subgenus Hymenandra appear to form a coherent taxonomic

group. Those in subg. Lacrimopila deserve renewed study and comparison when

additional material is available. The strikingly different forms of indument of H.

diamphidia and H. iteophylla comprise a very noticeable character and the tendency

of the distal cells in the trichomes of the former species to be twinned (either adnate

or slightly separated) is noteworthy. Perhaps this is a character that may be regarded

as comparable to, but less elaborate than, the stalked scales of H. iteophylla.

Hymenandra narayanaswamii Nayar & Giri differs from H. wallichii in its distinctly

petiolate, elliptic leaves, apparent lack of conspicuous inflorescence bracts, and lack

of papillosity inside the calyx. The authors regrettably failed to report the ovule number.

Hymenandra wallichii, the type species of the genus, has a unique foliar character;

the crenate leaf margins. The crenation is here termed “double” because each lobe is

Revision of Hymenandra 3

notched, and a veinlet ending terminates in this notch; while the lobes themselves are

separated by a broad, shallowly scooped interval of the margin, without a comparable

veinlet termination. The remainder of the species all appear to have entire leaf margins.

Clearly, Hymenandra depends for its taxonomic status heavily upon the single

character of anther connation, and I find no other character which could supplant it.

Systematic Treatment

HYMENANDRA A.DC.

Ann. Sci. Nat. ser. 2, 2: 297. 1834; ibid. 16: 79. 1841; in DC. Prod. 8: 91. 1844. Endlicher, Gen. 736.

1836-40. Lindley, Veg. Kingd. 648. 1847. Hooker fil. in Bentham & Hooker fil. Gen. Pl. 2: 647. 1876.

Pax in Engler & Prantl, Die Pflanzenfamilien IV.1: 95. 1889. Baillon, Hist. Pl. 11: 332. 1892. Mez, in

Engler, Das Pflanzenreich Heft 9 (IV.236): 155. 1902. Furtado, Gardens’ Bull. Singapore 17: 306. 1958.

Nayar & Giri, Journ. Bombay Nat. Hist. Soc. 72: 818-821. “1975” (actually 1976). — Ardisia sectio 2.

Hymenandra A.DC., Trans. Linn. Soc. London 27: 126. 1834.

Type: Hymenandra wallichii A.DC. (= Ardisia hymenandra Wall. in Roxb. FI. Ind.

ed. Carey 2: 282. 1824). (Wallich’s epithet cannot be used under Hymenandra because

of tautology).

Revised description: Flowers hermaphrodite, 5-merous. Sepals quincuncial, spreading

at anthesis, connate shortly at base, mostly papillose on interior surface (or hirtellous),

often rather elongated, valvate or scarcely imbricate. Corolla with short tubular base

adnate to tubular (filamentous) base of androecium, lobes slightly imbricate, ovate

to narrowly ovate, glabrous or sparsely papillose within toward base. Internal glands

in sepals, petals, and anthers roundish or oval or frequently elongated (oblong to

sublinear). Stamens connate, in the lower part as a filament tube adnate to the base

of the corolla tube, but the filament tips free between the base of the anther and

the point of adnation to the corolla; anthers connate laterally by the thecal margins,

narrowly ovate-acuminate or oblong-ovate and acuminate, dorsally glandular. Gynoe-

cium with ovoid ovary and slender, elongated style; stigma punctiform. Placenta ovoid

to turbinate, with 5-12 ovules in | or 2 series. Fruit 1-seeded, globular, sometimes

oblate, exocarp glandular, (sometimes?) subtended by the persistent clayx.

Key to the Species

la. Inflorescence condensed umbelliform or compactly cymose, at first conspicuously bracteate. Calyx,

corolla and anther connective with oval or oblong-linear immersed glands. Calyx lobes papillose on

inner surface. Filaments short, less than half as long as anther. Leaves obovate, petioles short and

rather complanate, margins entire to slightly crenulate. Main stem short suberect or ?decumbent, not

pe Ee, PC ee en oe Pen Pe Bere oe subgenus Hymenandra

2a. Ovary tomentellous; leaves larger, to over 30 cm long, obovate, gradually narrowed to base, the

broad petiole 5 mm long; lateral veins numerous; margins of blade distally crenulate, each lobe

notched with a veinlet termination in it; ovules biseriate; sepals c. 3-4 mm long. Distrib. Sikkim,

rintRe Maer be SON > BOSS. EEE el ceil ils ce bins tie ee. 2a ave tres (1) Hi. wallichii

2b. Ovary glabrous; leaf margin entire.

3a. Leaves as in H. wallichii, large obovate, to 40 cm long and 10 cm wide, subsessile with broad,

short petioles; sepals over 6 mm long, ovate-oblong and acute; ovules uniseriate. Distrib.

Ne ee ee ue wens ccm d,s: wie ss (2) H. lilacina

3b. Leaves smaller, sometimes to 35 cm long, but often narrower, obovate or oblanceolate; petioles

rather distinct; sepals narrowly ovate or ovate, less than 6 mm long; ovules uni- or biseriate.

4a. Lateral fertile branch very short, c. 1 cm long; leaves elliptic-lanceolate; anthers 3 mm

long; placenta with 5 uniseriate ovules. Distrib. Borneo ............... (3) H. rosea

4 Gard. Bull. Sing. 43 (1991)

4b. Lateral fertile branch longer, usually 9-12 cm long; leaves oblanceolate to obovate;

anthers 3-4.5 mm long; placenta with 8-10 uniseriate ovules.

Sa. Leaves obovate, at base obtuse to almost subcordate, at apex obtuse, 12-18 cm long,

7-9 cm wide; petioles 5-6 mm long; flowers 8 mm long; sepals ovate-subrhombic,

c. 5.5 mm long... Distrib. ,.BorneG...... + 2. 1.0:. .>a4. oe (4) H. beamanii

5b. Leaves oblanceolate, 30-35 cm long, 4-5 cm wide; petioles 1-2 cm long; flowers 5-6 mm

long; sepals elliptic ovate, 4.5 mm long. Distrib. Borneo ........ (5) H. calcicola

lb. Inflorescence compact or open, up to 3 times pinnate, paniculate, axes somewhat to considerably

elongated; bracts caducous; calyx-lobes marginally hirtellous and their interior surfaces not papillose,

but either hirtellous or glabrous; taller, more ramified plants.

6a. Inflorescence condensed, axillary and terminal, the peduncles 10-15 mm long.

7a. Calyx lobes triangular-ovate, 1.5 mm long; calyx and pedicels with scattered substelliform or

flabellate scales; filaments elongate, about 1.6 mm (c. one-half as long as the anther); anther

glabrous; ovary glabrous. Distrib..Malaya «~ ... 2 ....:..«<.2@ laws samt (8) H. iteophylla

7b. Calyx lobes and pedicels densely puberulous; filaments short, c. 1 mm long, much shorter

than the anther; anther glabrous; ovary glabrous; style 8-10 mm long. Distrib. Burma

be winiein die 6p vu « mis so snip a 5 5"em eee Sa ld dene ee (6) H. narayanaswamil

6b. Inflorescence open, racemose-paniculate, tripinnate, the peduncles to 20 mm long; calyx lobes

narrowly ovate, c. 3.5 mm long; calyx and pedicels, dorsal midline of corolla lobes, anthers (both

faces) and ovary all hirtellous with ferrugineous-tipped glandular hairs; filaments very short, c.

0.5 mm long; ovary and lower part of style hirtellous. Distrib. Borneo ... (7) H. diamphidia

Subgenus Hymenandra

1. H. wallichti A.DC. Fig. 1.

Ann. Sci. Nat. ser. 2, 16: 79. 1841; in DC. Prod. 8: 91. 1844. Clarke, in Hook. fil., Fl. Brit. Ind. 3:

532. 1882. Mez, Myrsinaceae, in Engler, Pflanzenr. Heft 9 (1V.236): 155, fig. 24. 1902. Kanjilal & Das,

Flora of Assam 3: 186. 1939. Nayar & Giri, Journ. Bombay Nat. Hist. Soc. 72: 819. “1975” (1976).

Ardisia hymenandra Wall. in Roxb. FI. Ind. ed. Carey 2: 282. 1824; Pl. Asiat. Rar. 2: 57, t. 175. 1831.

A. DC., Trans. Linn. Soc. Lond. 27: 126. 1834; Ann. Sci. Nat. ser. 2, 2: 12. 1834.

Leaves c. 30 cm long, to 11 cm wide; petiole 5-12 mm long, broad, complanate.

Leaf margins along distal half double-crenulate, each crenation-lobe notched, with a

veinlet termination in the notch. Blade with rather numerous (about 25) pairs of lateral

veins. Inflorescence terminal on short lateral fertile branch, compact, overall c. 10 cm

long, at first with pseudowhorled conspicuous bracts, decurved. Flowers c. 10 mm long,

calyx-lobes about 2 mm long, narrowly ovate, gland-dotted, the margins finely ciliate;

corolla-lobes narrowly long-ovate, the margins subacuminate, distinctly papillose within;

staminal tube elongate, the filaments very short, anthers much elongated, 7.5 mm

long, slenderly acuminate, laterally connate but the tips free, the connectives dorsally

set with oblong glands. Ovary tomentellous, | mm high, the style slender, 6.75 mm

long; stigma small, punctiform-truncate. Placenta about 0:5 mm high, with about

10 to 12 ovules in two rows.

Type: Wallich Cat. 2226 (from Sylhet, mountains of Juntiyapoor, now in Bangladesh),

K! holotype; CAL isotype.

Specimens examined: ASSAM: Khasia, Griffith 3596 (K!). (Two sheets, here designated

3596a and 3596b; the former bears a label with the annotation “Frutex humilis, foltis

subcarnosis; inflorescentia carnea nutans sub hemisphaerica”’; towards Beesale Collieae;

this sheet possesses old flowers, the second has fruits.) Cachar, “shrub of shade, with

nearly simple stem and reddish flowers borne in shortly pedunculate cluster close to

the somewhat succulent stem”; Gopal teetah B. masa, 5 June 1873, coll. Memang

Sirdar for Keenan (K!). Meghalaya, 1,000 m alt., June 1876, coll. ignot. 3/2 (CAL, not

seen, cited by Nayar & Giri). Naga hills, May 1899, Prain’s collector 100021 (CAL, cited

Fig. 1.

Hymenandra wallichii A.DC. a. Flower in profile. b. Calyx-lobe. ¢. Corolla-lobe. d. Detached

stamen, dorsal view. e. Ovary and style. f. Placenta, with 12 ovules. g. Capitate glandular

trichomes from ovary. h. Fruit. i. Fruit in transverse section (dried, the seed omitted). j. Leaf

undersurface. k. Portion of leaf margin, showing double crenulation. I. Peltate scale from leaf

undersurface. (a-i, from De Silva, Wallich list no. 2266, type collection; j-l, from Keenan.)

6 Gard. Bull. Sing. 43 (1991)

by Nayar & Giri). BANGLADESH: Sillet (Sylhet), in mountains of Juntiyapoor, flower-

ing in March, coll. De Silva for Wallich, Wall. List 2266 (K!), holotype, isotype CAL.

Cultivated in some botanic gardens/greenhouses (ref. Edinburgh).

2. Hymenandra lilacina B.C. Stone, sp. nov. Fig. 2.

Suffrutex glaber, stipite brevi, crasso, suberecto ad 20-30 cm alto, ad 10 mm diametro, cortice 2 mm

crasso; foliis spiraliter dispositis, magnis, obovato-oblanceolatis, usque ad 40 cm longis et 10 cm latis,

coriaceis, integris, acutis vel obtuse acuminatis, basi longe angustatis decurrentibus demum abrupte sub-

truncatis vel subrotundatis, petiolo breve, lato, 5-10 mm longo et 4-6 mm lato, dorso rotundato, infra

subcanaliculato; costa supra leviter et infra valde elevato; venis lateralibus numerosis (c. 10-12—-paribus),

venis secundariis et tertiariis distinctis prominulentibus; pagina infra in sicco brunneo. Ramus fertilis lateralis

angustis c. 15 cm longis, 1.5-2 mm diametro, nudo termini excepto dein 1-2-foliati, foliis valde reductis,

ad 10 cm longis, subsessilibus. Inflorescentia bisumbellata, compacta bracteis lanceolatis pseudo-involucello

formantibus; pedunculis 5 (postea —9) mm longis; pedicellis 3-5 mm longis. Flos c. 7 mm longus, calycis

lobis anguste ovato-oblongis, copiose glandulosis (glandulis oblongo-sublineatis), 5-venosis, glabris sed

margine minute ciliatis; pagina intus copiose et ubique papilloso (papillis subcapitatis); corolla in parte

basali quaterni tubulosi, in toto 7 mm longo, lobis ovatis acuminatis, in basi subauriculatis, integris,

copiose glandulosis, utrinque glabris vel intus basaliter persparse papillosis; tubo staminorum 5 mm longo

antheris 4.5 mm longis apiculatis, dorso bilineatim glandulosis (glandulis c. 12-15 nigris); ovario conico

glabro, pustulato-glanduloso, 1 mm alto, stylo erecto 4.5 mm longo gracillimo (glandulis internalibus

evidentibus), stigma albo subtruncato; placenta 0.75 mm alto, ovulis 8 vel 9 uniseriatis. Fructus globosus

apiculatus 6 mm longus, 5.5 mm latus, glandulosis, monospermus, lobis calyci persistentibus.

Type: BORNEO: East Kalimantan, Berouw, Mapulu, foot of Mount Ilas Mapulu,

sandstone, 300 m alt., locally common, subshrub c. 30 cm high, flowers pale purple,

fruit red, 19 September 1957, A.J.G.H. Kostermans 13951 (K! holotype; 2 sheets; iso-

types CANB, L).

A striking plant with considerable horticultural potential, similar in gross appearance

to H. wallichii, but with entire leaf margins, somewhat shorter flowers, more briefly

apiculate anthers, glabrous ovary, and uniseriate ovules.

3. Hymenandra rosea B.C. Stone, sp. nov. Fig. 3.

Fruticulus 30 cm altus, stipite suberecto-decumbente, ad 4 mm diametro, cortice atro, ubique glabro

lobis calycis ciliatis excepto. Folia pauca, subconferta. Folia petiolis 12-16 mm longis, laminis lanceolato-

ellipticis, ad 13 cm longis, 3.4 cm latis, acutis non-acuminatis, basi angustatis, marginibus integris vel

perobscure crenulatis apicem versus, glabris; costa media infra prominente; venis lateralibus gracilibus,

subobscuris, c. 10-12-paribus, venis tertiariis vix manifestis; pagina infra pallidiora, lineis brevibus oblique

et perdissite dispersis atris evidentis; glandulis perminutis. Inflorescentia terminalis in ramulo laterale

brevissimo c. 1 cm longo, 1-2-foliato (foliis valde reductis) situata; bracteis lanceolatis 3-5 mm longis

atroglandulosis; umbello composito umbellis secundariis 1 vel 2 involucellatis, pedicellis 5-6 mm longis.

Flos c. 7 mm longus, calyce profunde lobato, lobis ovato-ellipticis subacutis, 4.5 mm longis, 2 mm latis,

margine ciliatis (trichomiis anguste clavatis 0.05 mm longis), 5-venatis, glandulis ovalibus vel oblongis

atris; corolla profunde lobato, lobis late ovatis acutis, 6 mm longis, 3.5 mm latis, integris, 7-venatis, glan-

dulosis; filamentis distinctis 1 mm longis glabris; antheris angustis subacuminatis 3 mm longis, dorso ad

connectivo glanduloso; ovario glabro subconico glanduloso, c. 1 mm alto, stylo gracile 4.5 mm longo

stigma truncato; placenta conico acuminoideo substipitato 0.75 mm alto, ovulis 5 uniseriatis.

Type: BORNEO: SABAH: Sandakan district, Gomantong Hill, primary forest, black soil,

flowers pink, 27 June 1963, J Ah Wing (Awing) SAN 38111 (K! holotype, isotype SAN).

It is not clear if the anthers are all united in this species. The flowers studied had

apparently free anthers, but they may have separated in the drying or boiling process.

The overall habit and general floral features of this plant are highly concordant with

Hymenandra.

Fig. 2.

or thn ie

t er.

[Dede

—

=~ ve Te

aries

Hymenandra lilacina n. sp. a. Flower in profile. b. Calyx-lobe, interior view; entire surface with

suberect papilliform glandular trichomes. ¢. Corolla-lobe, interior view; antistaminal subbasal

zone sparsely and infrequently with a few papillae. d. Staminal tube, with glandular connectives.

e. Ovary and style with dermal glands. f. Ovary in longitudinal section displaying placenta.

g. Placenta with 9 ovules. From type collection, Kostermans 1395].

7Tmm j

HISE “NYS

Fig. 3.

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Hymenandra rosea n. sp. a. Flower in profile. b. Calyx-lobe, interior view, showing surface

almost uniformly set with suberect papilliform glandular trichomes. ¢. Corolla-lobe, interior

view, the glands are subdermal. d. Stamen, dorsal view, the dotted lines showing attachment to

base of corolla-tube. e. Ovary and style; subdermal glands only in ovary. f. Placenta in profile

(lower) and top views; ovules 5. All from type collection, SAN 38/11.

Revision of Hymenandra 9

4. Hymenandra calcicola (Furt.) B.C. Stone, comb. nov.

Ardisia calcicola Furtado, Gardens’ Bulletin, Singapore 17: 296. 1958.

Glabrous subshrub to 50-75 cm tall; upper part of stem 5 mm diam. Main leaves

oblanceolate or obovate, subacute to acute, toward the base gradually narrowed, to

20 cm long and 5 cm wide. Petioles rather distinct and somewhat long, up to 20 mm

long and less than 2 mm thick. Lamina subcoriaceous, entire, obscurely undulate or

entire, midrib beneath prominent; undersurface darker than midrib; lateral veins

obvious, about 6-10 pairs with subequal intercalated veins, prominulent beneath;

tertiary veins mostly obscure; glands very minute, reddish, scattered. Lateral fertile

branches slender, 9-12 cm long, 1 mm thick, at apex with one much reduced leaf (this

subsessile, narrowly elliptic-lanceolate, acute at both ends). Inflorescence an umbel of

umbels, about 2 cm long and equally wide, peduncles 5-8 mm long, bracts lanceolate-

acuminate, to 5-6 mm long; pedicels 5-7 mm long, slender. Flower c. 6 mm long,

calyx-lobes 4 mm long, 1.8 mm wide, subentire, but sparsely ciliate near base, copiously

black-glandular with oval-oblong glands; interior surface entirely papillose with sub-

erect glands. Corolla tubular in basal fifth, lobes ovate-acuminate, entire, internally

and externally quite glabrous, rather sparsely glandular with oblong reddish glands.

Stamen tube 5 mm long, anthers 4.5 mm long, connective rather sparsely glandular

(about 8 glands in each of the two rows); ovary glabrous, distinctly glandular, conic,

almost 1 mm high, style slender, glandular, 3 mm long; stigma truncate. Placenta with

10 ovules in one series, c. 0.7 mm high. Fruit not seen.

Type: BORNEO: KALIMANTAN: West Koetai subdistrict, 30 m alt., edge of limestone

rockface in flat country, in forest, shrub to 75 cm high, flowers dark red, 22 November

1925, FH. Endert 5134 (SING! holotype, isotypes K, L).

Additional specimen examined: BORNEO: KALIMANTAN: Berau, Mt. Njapa on Kelai

River, 1,000 m alt., shrub 1 m, flowers and fruits red, 25 Oct. 1963, Kostermans

21518 (L!)

Although placed in Ardisia by Furtado, this species appears to have a staminal tube;

as noted in the caption to the illustration provided by Furtado in his protolog, which

states “E, Flos juvenilis sepalis desumptis ut stamina tubiformiter collata appareant.”

Furtado’s diagnosis also specifies that the calyx-lobes are lepidote on both sides; |

equate this term in this case with papillose. This species clearly is very near to H.

lilacina and H. rosea.

5. Hymenandra beamanii B.C. Stone, sp. nov. Figs. 4, 5.

Suffrutex suberectus humilis ad 50 cm altus, stipite simplici, ramuli unifoliati fertili excepta; foliis

caulinibus magnis obovatis obtusis basi truncatis vel subcordatis, petiolis crassis 5-6 mm longis, laminis

12-17 cm longis, 7-9 cm latis, glabris, sed in pagina infra minute et dissite lepidotis. Folia in ramuli laterali

solitaria multo minores subsessilia. Inflorescentia compacta congeste bisumbellatis, apicalia in ramulis

lateralibus, axibus valde contractis, pedicellis 5-11 mm longis. Flores numerosi conspersi, 7-8 mm longi,

bracteis lanceolatis c. 5 mm longis. Calyx cupulari-urceolatis profunde lobatis, scarioso-marginatis, extus

glabris, margine minute et sparsiter capitato-ciliatis, glandulis immersis ovalibus vel oblongis; intus minute

et perdense papillosis. Corolla 7.5 mm longa in basi tubuloso per 1.5 mm, lobis ovatis acutis leviter

asymmetricis, glandulis immersis ovalibus vel oblongis distaliter auctis, integris, utrinque glabris. Stamina

fere 6 mm longa, per antheras coalitis basi per filamentas tubulosa, filamentis liberis ad basi corollae

adnatis 1.5 mm, parte libera 0.5 mm longo, antheris lanceolatis 4 mm longis, marginaliter connatis

introrsis, dorso glandulosis glandulis paucis (c. 12) atris rotundatis, apiculis antherarum breviter liberatis.

Ovarium conico-ovoideum, obscure glandulosum, c. 1 mm altum, in stylo elongato 6 mm longo produc-

tum, stigma minute punctiforme. Placenta turbinata minute apiculata, 0.75 mm alta, ovulis 6-9 uniseriatis.

Fructus ruber globosus, c. 6 mm diam., minute apiculatus.

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Fig. 4. Hymenandra beamanii n. sp. a. Flower in profile. b. Calyx-lobe, interior view; surface with

small papillae; immersed glands reddish or blackish. ec. Corolla-lobe, interior view; note basal

area (dotted) to which filament is adnate. d. Stamen tube; filaments adnate to corolla-tube on

dotted areas. e. Stamen, ventral view. f. Gynoecium. g. Placenta (ovules 8). h. Placenta in

transverse section. From SAN 77243, type.

Type: BORNEO: SABAH: Ranau district, about 3 miles north of Kampong Takutan,

primary forest on ridge at 2,800 ft. alt., flowers whitish, 28 May 1973, G. Shea & Aban

Gibot SAN 77243 (K! holotype, isotypes L! SAN).

Additional specimen examined: BORNEO: SABAH: Ranau district, southwest side of

Lohan River, 700-900 m alt., on cliffs and slopes over ultramafic substrate, low stature

forest, 5 April 1984, JH. Beaman 9229 (PH! MSC, UKMS).

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Fig. 5.

B.cademy of Natural Sciences of Phi

Ardisia beamanii.B.C.S5toue

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Det Benjamin C. Stone

PLANTS OF BORNEO

Ardisia

Malaysia. Sabah, Ranaut District: bank,

slopes, and chiffs on SW side of Lohan

: River. 6°00°N, L1641'E. Elev. 700 -

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: Tiny treelet, fruits red.

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Beaman, L.

Universi Kebangre™ Malaysia, 284"

Hymenandra beamanii n. sp. From Beaman 9229 in PH

Uhramafic geology

12 | Gard. Bull. Sing. 43 (1991)

The very compact umbelliform inflorescences borne on slender subterminal fertile

branches, each with a strongly reduced leaf, the elongate sepals with densely papillose

inner surfaces, and the laterally joined anthers readily distinguish this species as a

Hymenandra. The floral details in the description above are taken from the SAN

collection, those of the fruits from the Beaman collection, and the habit and leaf

information from both.

The epithet commemorates the collector, Professor John Beaman of the Michigan

State University, who collected this and numerous other Sabah plants during the course

of a Fulbright-sponsored stint at the National University of Malaysia, Sabah campus.

6. Hymenandra narayanaswamii Nayar & Giri

Journ. Bombay Nat. Hist. Soc. 72: 818-821. “1975” (actual date of publication 13 Aug 1976).

Shrub with terete, glabrous stems. Leaves oblong to oblong-elliptic, 8-18 cm long,

3-11 cm wide, acute at both apex and base, entire, pellucid-glandular-punctate,

membranaceous, the midrib prominent beneath; lateral veins about 30 with about as

many nearly as prominulent intersecondaries; reticulations evident; petiole 10-12 mm

long, canaliculate above. Inflorescence axillary, 4-7 cm long, subcorymbose paniculate,

shorter than the leaf, densely puberulous, glandular-dotted; flowers 5-merous, corym-

bosely to subumbellately arranged; pedicels 4-8 mm long; calyx lobes triangular-

lanceolate, densely puberulous and glandular-punctate; corolla lobes shortly united at

base, lanceolate, 7-8 mm long, 1.5-2 mm wide, long acuminate, glandular-punctate,

dextrorsely imbricate; stamens with very short (0.5 mm) free filaments, anthers linear-

lanceolate 6-7 mm long, laterally connate, glandular-punctate dorsally along the con-

nective; ovary subglobose, scarcely 1 mm high, glandular, abruptly narrowed at apex

into the slender 8-10 mm long style; stigma inconspicuous; placenta ... and ovules

... (not reported).

Type: BURMA: Tavoy, PT? Russell 2105 (CAL, not seen).

Nayar & Giri state of this species that it is allied to Hymenandra wallichii but

differs in having oblong or oblong-elliptic leaves with cuneate base, acute apex, entire

margin, and membraneous texture, triangular lanceolate calyx lobes and abruptly

attenuated apex of anther; whereas in H. wallichii, the leaves are obovate-lanceolate or

oblanceolate with attenuate base, rotund apex, dentate margin, and fleshy texture;

calyx lobes are ovate; and apex of anther is gradually attenuate. Most unfortunately,

these authors do not report on certain critical characters such as whether or not the

interior faces of the calyx lobes are papillose, and what the number and disposition

of the ovules may be on the placenta. The species is illustrated but these features are

not shown in the drawings either, nor are there descriptions or drawings of the trichomes.

No mention of the bracts is made. The authors do not redescribe AH. wallichii or

H. iteophylla.

Subgenus Lacrimopila B.C. Stone, subg. nov.

Inflorescentia compacta vel laxe tripinnatim paniculata, axibus pedicellisque satis vel bene elongatis,

bracteis cito caducis; calycis lobis marginaliter hirtellis vel lepidotis, intus epapillosis (sed in species typicum

hirtellis); indumento ferrugineo, pilibus cellulis apicalibus capitulatis lacrimiformibus, vel lepidis flabelli-

formis vel stellatiformis. Arbusculae vel frutices modice erectae ramis pluralibus basi ampliati alternatim

foliatis. Typus: Hymenandra diamphidia Stone.

7. Hymenandra diamphidia B.C. Stone, sp. nov. Fig. 6.

Arbuscula usque ad 3 m alta, sparse ramosa, ramis ramulisque gracilibus, subteretibus, ad 3-4 mm

diametro, cortice atro, subdensiter ferrugineo-tomentellis, demum glabrescentibus; indumento trichomiis

Revision of Hymenandra 13

stipite cellulis 1 vel 2 pellucidis cellulis terminalibus ovoideis plerumque gemellatis translucide brunneis

acutis vel bifidis c. 0.05-0.075 mm longis dissite dispersis in superficiem pedicelli, calycis, corollae,

antherae, et gynoecii, et axium omnium inflorescentii; partibus aliis glabris. Folia exacte anguste ovatae

vix acuminatae, 3-20 cm longae, 1.5-6.5 cm latae, basi rotundata vel subcordata, valde petiolatis, petiolis

gracilibus 10-30 mm longis, subdensiter minuteque tomentellis; lamina integra submembranacea, per-

conspicue multiglanduloso glandulis atris orbicularibus, pagina supra glabra (marginibus et costa tomentellis

excepta), infra pallidiora sueto purpurata ad venis tomentellis; costa supra vadose impresso, infra valde

carinato, venis lateralibus manifestis, 6-9-paribus, curvato-patentibus, praeter margos sublonge continuatis;

reticulationibus tenuibus sed manifestis. Inflorescentia in ramulis lateralibus terminalis, pendula, laxe

tripinnatim racemoso-paniculatis, 10-15 cm longis, multifloris, axibus omnibus gracilibus minuteque

tomentellis, floribus in pedunculis laxe dispositis, pedicellis 5-8 mm longis in statu floriferi (vel usque ad

10 mm longis in statu fructiferi). Flores angusti c. 6 mm longi purpurei 5-meri tomentelli. Calyx profunde

lobatus, lobis peranguste triangularibus 3-5 mm longis, basi 0.6-0.7 mm latis utrinque et marginaliter

tomentellis, basem versus persparse glandulosis. Corolla profunde lobata 6.5 mm longa, lobis anguste

ovatis subacuminatis apicem versus leviter convolutis, medialiter persparse glandulosis, integris, extus

dorso tomentellis, intus glabris, obscure paucivenosis, basi minime tubulosis. Stamina per antheris

connata filamentis liberis brevissimis complanatis 0.5 mm longis perminime ad tubo corollae adnatis,

antheris lanceolatis acuminatis 4.5 mm longis, utrinque sparse tomentellis, haud glandulosis, rimose

aperientibus, apiculis brevissime liberatis, polline luteo. Ovarium breviter conicum vix 1 mm altum in

stylo gracile albo 4 mm longo basem versus sparse tomentello productum, stigma punctiforme perminute

sparseque subpapilloso; placenta turbinato ovulis 8-9 irregulariter biseriatim dispositis. Fructus globosus

tomentellus, miniatus vel scarlatinus, apiculatus, c. 6 mm diametro, calycis lobis patentibus persistentibus

et pedicellis gracilibus auctus.

Type: BORNEO: SABAH: Ranau District, Mount Kinabalu, Ulu Liwagu and Ulu Mesilau

(6 N, c. 116 35’ BE), c. 5,000 ft. alt., shrub 3-5 ft. tall, sparingly branched, flowers

purplish, berries pink, young leaves purple beneath, inflorescence and infructescence

hanging, 6 November 1961, W.L. Chew, E.J.H. Corner and J.D. Stainton RSNB 2809

(K! holotype).

Additional specimens examined: BORNEO: SABAH: Mount Kinabalu, Ulu Liwagu and

Ulu Mesilau, Tenompok, 5,000 ft. alt., tree 10 ft. tall, in oak-podocarp forest, fruit red,

8 September 1961, Chew, Corner and Stainton RSNB 1454 (K!). Same locality, Ulu

Langanani, Sungei Mamut (6 04’ N, 116 40-44’ E), 4,500 ft. alt., shrub 8 ft. tall, fruit

scarlet, in montane oak forest, 10 August 1961, Chew, Corner and Stainton RSNB

1265 (K!).

A most extraordinary species, with numerous peculiar features, differing from all

other species of Hymenandra other than H. iteophylla in habit, indument, shape and

size of glands, and type of inflorescence. The comparatively slender and elongated

petioles, narrow leaves, with rather large scattered black pustular glands, the open,

pendulous, slender-stalked inflorescence, the characteristic form of the glandular tri-

chomes, the tomentellous ovary, and the presence of hirtellous indument on the anthers,

are all manifest and remarkable features of this species. The epithet “diamphidia”

meaning (Greek) “utterly different” seems appropriate for this plant.

The glandular trichomes have a short stalk of one or two cells, and terminate in one

or usually two cells which are laterally adnate or more or less separated at the tip, thus

appearing “twinned”. These apical cells are in the shape of a teardrop (lacrimiform)

and are translucently ferrugineous.

While placing this species in a subgenus separate from that including the type

species of Hymenandra, the tubular androecium and the ovular number conform

well to the generic characters. Yet there are several peculiar or unique features of this

species. Including H. iteophylla with its very different kind of indument, but similar

habit, may be incorrect, but I would prefer newer collections to become available for

study before reassessing this relationship.

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Fig.6. |Hymenandra diamphidia n. sp. a. Flower in profile. b. Calyx with one lobe removed. ¢. Corolla-

lobe, interior view. d. Stamen tube. e. Single detached stamen, ventral face oblique. f. Ovary

and style. g. Placenta with 9 ovules. h. Tip of style showing sparsely papillate stigma. i. Trichomes

showing the twinned apical cells of lacrymiform shape, and 1-2-celled stalks; representative

of the indument on all parts. j. Leaf undersurface, the tertiary veins omitted for clarity. Scat-

tered dots are immersed glands. k. Branch base (gusset). All from type collection, Chew, Corner

& Stainton RSNB 2809. (All illustrations original.)

Revision of Hymenandra 15

8. Hymenandra iteophylla (Ridl.) Furtado Fig. 7.

Gardens’ Bulletin, Singapore 17: 306. 1958. Nayar & Giri, Journ. Bombay Nat. Hist. Soc. 72: 821.

“1975” (actually published 13 Aug 1976).

Ardisia iteophylla Ridley, Journ. Bot. 62: 298. 1924; Fl. Malay. Penins. 5: 318. 1924.

Entirely resembling an Ardisia (Ridley); branchlets with broad triangular gussets at

base; shrubby with ascending branches; leaves lanceolate-elliptic, thinly coriaceous,

slightly acuminate, cuneate, tapered to base, 5-10 cm long, 1-2 cm wide, the petiole

5-10 mm long; young leaves brown-lepidote (or golden) beneath; scales irregular, sub-

stellate; indument of young branches ferrugineous, with the scales erect, flabellate to

substellate, later evanescent; midrib elevated beneath, lateral and tertiary veins obscure,

numerous, close; undersurface densely lepidote. Inflorescence terminal and axillary

subterminal, with foliaceous bracts; umbels with lepidote-puberulent axes and bracts;

pedicels 6-7 mm long, slender; calyx tuberculate-glandular, the lobes narrowly ovate,

1.5 mm long, copiously glandular; ciliate; corolla-lobes pink, long and narrow, im-

bricate, acute, entire, glabrous, not or very obscurely glandular, 4-5 mm long, the tube

0.75 mm long; stamens lanceolate, the filaments much elongated (more than half as

long as the anthers), distinct; anthers lanceolate, 4.25 mm long, connate marginally,

but tips free; outer thecae with fertile portion about *% as long as anther, inner thecae

with fertile portion shorter, about half as long as anther; connective set dorsally with

two rows of large convex glands; ovary subglobose, 1.5 mm high, glabrous, with the

slightly glandular style 6.5 mm long and projecting beyond the corolla. Placenta

0.5 mm high, with about 6 to 8 ovules in one row.

Type: MALAYSIA: JOHORE: Gunung Besidong, 1,500 ft. alt., hill forest, R.A. Holttum

SF 10973 (SING! holotype).

Additional specimens examined: MALAYSIA: JOHORE: Gunung Belumut, northwest,

1,200 ft. alt., rocky stream bed, monopodial treelet 6 ft. tall, flowers purplish-pink,

leaves golden beneath, 15 May 1968, 7:C. Whitmore FRI 8782 (KEP!) TRENGGANU:

Bukit Bauk Forest Reserve, 1,000 ft. alt., shrub 6 ft. tall, on ridge, fruits green, 13

November 1968, K.M. Kochummen FRI 260] (KEP!).

This species is very distinctive in its small leaves with a golden-brown indument

beneath, the subterminal umbels, the scattered multiform flabellate to substellate erect

or suberect scales on the axes (while the scales on the leaves are peltate and appressed),

the dark roundish glands of the calyx and connectives, and the long, slender filaments.

Its relationship to H. diamphidia is perhaps rather dubious, though in habit it cor-

responds well. The highly different type of indument, glabrous anthers and ovary,

elongated filaments, and smaller, lepidote leaves are obvious distinguishing features.

A character worthy of note is the unequal length of the thecae in each anther. The

inner adjacent thecae are about half the anther length; the outermost thecae are about

two-thirds the length of the anther.

References

De Candolle, A (1834). A review of the natural order Myrsineae. 7rans. Linn. Soc.

Lon. 17: 95-138.

. (1841). Deuxieme memoire sur les Myrsinacees. Annales des Sciences Naturelles,

ser. 2, 16:65-97.

Furtado, C.X. (1958). Some new or noteworthy species of Malaysia. Gard. Bull. Sing.

17: 279-311.

Fig. 7.

Hymenandra iteophylla (Ridl.) Furt. a. Flower in profile. b. Corolla interior, showing two lobes

with filaments intact but anthers removed. ec. Androecium, detached from corolla. d. Ovary

and style. e. Placenta. f. Placenta, enlarged, with 8 ovules. g. Placenta in transverse section.

h. Portion of inflorescence showing one branch, with axillant bract and bracts of the branch.

i. Various flabellate and stelliform scales of the indument. (All from Holttum SF. 10973, type

collection.)

Revision of Hymenandra 17

Mez, C. (1902). Myrsinaceae. In Engler, A., Das Pflanzenreich, Heft 9 (I1V.236): 1-437.

Nayar, M.P. and Giri, G.S. “1975” (1976). A synopsis of the genus Hymenandra A.DC.

(Mysinaceae) and a new species from Burma. J. Bom. Nat. Hist. Soc. 72: 818-821.

(Actual date of publication: 13 August 1976.)

Roxburgh, W. (1824). Flora Indica, Ed. W. Carey 2: 282. Cf. Ardisia hymenandra

Wallich.

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New Algal Records from the Singapore Mangroves

JOHN A. WEST

Department of Plant Biology

University of California

Berkeley CA 93720 USA

EFFECTIVE PUBLICATION DATE: 23 MAR 1992

Abstract

The following marine algae are newly recorded for Singapore mangroves: Bostrychia pinnata Tanaka

et Chihara, Bostrychia simpliciuscula Harvey ex J. Agardh (Rhodophyta, Ceramiales, Rhodomelaceae),

Caloglossa angustalata nom. prov. (Rhodophyta, Ceramiales, Delesseriaceae) and Boodleopsis carolinensis

Trono (Chlorophyta, Caulerpales).

Collections and Observations

The rapidly dwindling mangroves of Singapore are an interesting and diverse habitat

of various estuarine algae. Johnson (1979), Teo and Wee (1983) and Chapman (1984)

recorded many of these. In June 1989 I had the opportunity to collect specimens in

several sites for laboratory culture investigations on their reproductive biology. Below

are listed four taxa previously unrecorded in Singapore. The classification, reproductive

condition, date of collection, specific location, habitat and culture observations are

given for each record.

Chlorophyta, Caulerpales

Boodleopsis carolinensis Trono

Johnson (1979) recorded Derbesia fastigiata Taylor as the only genus of the Cauler-

pales in the Singapore mangroves, but the vegetative and reproductive features of the

present specimens are similar to B. carolinensis. A dense green turf is formed by erect

filaments 15-21 um diam., di- to tri-chotomously branched and with prominent con-

strictions present at the nodes. This system arises from a basal system comprised of

lighter pigmented, more irregularly shaped and branched filaments of a greater dia-

meter. Subspherical sporangia up to 110 wm diam. are present occasionally on the

erect filaments. These features generally fit the description provided by Trono (1971)

for B. carolinensis. Boodleopsis pusilla (Collins) Taylor, Joly ef Bernatowicz (Taylor,

et al. 1953, Calderon-Saenz and Schnetter 1989) is a more widely distributed species

but the filament diameter is greater (23-45 ym). Boodleopsis siphonaea A. et E.S.

Gepp also is larger in filament diameter (30-60 um) according to the comments of

Tanaka and Chihara (1988). Boodleopsis hawaiiensis Gilbert also conforms to the

general appearance of these specimens except for the greater range in filament diameter

(60-110 »m) and the apparent lack of sporangia (Gilbert, 1965).

Collections were made on 12 June 1989 at Kranji and on 13 June 1989 at Mandai.

In both sites Boodleopsis formed a dense turf at the +3.0 m level above chart datum

on decaying logs and on mud.

20 Gard. Bull. Sing. 43 (1991)

Rhodophyta, Ceramiales, Rhodomelaceae

Bostrychia pinnata Tanaka et Chihara

This species, described in 1984 by Tanaka and Chihara from Okinawa, appears to

be distributed widely in Asia (Indonesia: Tanaka and Chihara, 1988, Australia: King

and Puttock, 1986, West et a/, 1992). It was found in two localities in Singapore: Lim

Chu Kang, mixed with Catenella and Caloglossa, 13 June 1989 and Lim Chu Kang,

on Avicennia bark at +3.0 m above chart datum, 16 June 1989. All field plants were

nonreproductive, but both isolates have developed tetrasporangia in culture. Tetraspores

developed into unisexual and bisexual gametophytes. Both female and _ bisexual

gametophytes formed well developed carposporophytes that released viable carpospores

that again developed into tetrasporophytes.

Bostrychia simplicituscula Harvey ex J. Agardh

- This species also is widely distributed in Asia (King and Puttock, 1989). It was

obtained in two localities in Singapore: Tetrasporangiate plants occurred at Mandai in

a Boodleopsis turf at +3.0 m above chart datum on 13 June 1989. Vegetative and male

plants were present in mud at +3.1 m at Lim Chu Kang on 16 June 1989. The Mandai

isolate in culture has completed a Polysiphonia-type life history with self-compatible

unisexual and bisexual gametophytes. The Lim Chu Kang isolate in culture is a male

that is cross-fertile with the Mandai isolate.

The B. pinnata and B. simpliciuscula collected in Singapore and elsewhere contain

D-dulcitol and D-sorbitol as compounds important in osmotic acclimation (Karsten,

et al. 1992).

Rhodophyta, Ceramiales, Delesseriaceae

Caloglossa angustalata nom. prov.

Vegetative plants were collected in the upper turf areas at +2.0-3.0 m above chart

datum at Mandai on 13 June 1989 and Lim Chu Kang on 16 June 1989 mixed with

Caloglossa stipitata Post. Both isolates in culture have completed a Polysiphonia-type

life history and remain uniform in morphology.

A careful survey of the literature and herbarium specimens indicate this species is

undescribed. The narrow axes with wing cells often absent makes this most distinctive

in comparison with other Caloglossa species. The final description of Caloglossa

angustalata will be published in Botanica Marina (West, in preparation).

A similar species, C. ogasawaraensis Okamura, was isolated into culture from col-

lections made from mud-covered Avicennia pneumatophores in Darwin, Northern

Territory, Australia and from Rhizophora prop roots in Ilha do Cardoso, Sao Paulo,

Brazil and Tumbes, Peru (West, 1991). Caloglossa ogasawaraensis produces 3-5 wing

cells along each side of the midrib in the upper region of each internode whereas

C. angustalata rarely forms more than 1-2 wing cells even at its widest. All Caloglossa

species including C. angustalata produce mannitol for osmotic acclimation (Karsten,

et al. 1992).

For all of the above-mentioned species, herbarium voucher specimens have been

deposited at the University of California, Berkeley, Herbarium (UC). The culture

isolates are deposited in The Culture Collection of Algae, Department of Botany,

University of Texas, Austin, TX 78713-7640.

New Algal Records from Singapore Mangroves 21

Acknowledgements

A travel grant from the University of California Committee on Research made this

work possible. Tim Lowrey, Paddy Murphy, Shawn Lum, Johnny Wee and many

other colleagues at the Departments of Botany and Zoology, National University of

Singapore, generously provided facilities and assistance.

References

Calderon-Saenz, E. and Schnetter, R. (1989). The life histories of Boodleopsis vauch-

erloidea sp. nov. and B. pusilla (Caulerpales) and their phylogenetic implications.

Phycologia 28: 476-490.

Chapman, V.J. (1984). Botanical surveys in mangrove communities, pp. 53-80. In

S. Snedaker and J. Snedaker (eds.) The Mangrove System: Research Methods.

UNESCO. Paris.

Gilbert, W.J. (1965). Contributions to the marine Chlorophyta of Hawaii, II. Addi-

tional records. Pac. Sci. 19: 482-492.

Johnson, A. (1979). Algae of Singapore Mangroves. Proc. BIOTROP Symp. on

Mangrove and Estuarine Vegetation in SE Asia. No. 10: 45-49.

Karsten, U., West, J. and Zuccarello, G. (1992). Polyol content of Bostrychia and

Stictosiphonia (Rhodomelaceae, Rhodophyta) from field and culture. Bot. Mar.

35: in press.

Karsten, U., West, J., Mostaert, A., King R., Barrow, K. and Kirst, G. (1992). Mannitol

in the red alga Caloglossa. J. Plant Physiol. (submitted).

King, R. and Puttock, C. (1986). Bostrychia pinnata J. Tanaka et Chihara in Australia.

Bull. Nat. Sci. Mus., Tokyo, Series B. 12: 17-24.

. (1989). The morphology and taxonomy of Bostrychia Montagne and Sticto-

siphonia J.D. Hooker et Harvey (Rhodomelaceae/Rhodophyta). Aus. Sys. Bot.

2: 1-73.

Tanaka, J. and Chihara, M (1984). Taxonomic studies of Japanese mangrove macroalgae.

I. Genus Bostrychia. Bull. Nat. Sci. Mus., Tokyo, Series B. 10: 115-126.

. (1988). Macroalgae in Indonesian mangrove forests. Bull. Nat. Sci. Mus., Tokyo,

Series B. 14: 93-106.

Taylor, W.R., Joly A. and Bernatowicz, M. (1953). The relation of Dichotomosiphon

pusillus to the algal genus Boodleopsis. Michigan Acad. Sci., Arts and Letters. 38:

97-107 + 3 plates.

Teo, Lee Wei and Wee, Yeow Chin (1983). Seaweeds of Singapore. Singapore University

Press. iv + 123 pp.

Trono, G. (1971). Some new species of marine benthic algae from the Caroline Islands,

Western-Central Pacific. Micronesica 7: 45-77.

West, J. (1991). New records of marine algae from Peru. Bot. Mar. 34: 410-416.

West, J., Hommersand, M. and Calumpong, H. (1992). Reproductive biology of

Bostrychia pinnata J. Tanaka et Chihara in laboratory culture. Bot. Mar. (submitted).

yt z

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Notes on the Systematy cf Malayan Phanerogams

XX XI Lauraceae

K.M. KOCHUMMEN

c/o FRIM, Kepong, Malaysia

EFFECTIVE PUBLICATION DATE: 23 MAR 1992

Abstract

Two new species, Cinnamomum pubescens and Endiandra scrobiculata, and two new varieties,

Actinodaphne sesquipedalis var. glabra and Lindera concinna var. reticulata are described.

Introduction

Malayan Lauraceae was studied for incorporation in the 4th volume of Tree Flora

of Malaya. On completion of this study, two new species, Cinnamomum pubescens

and Endiandra scrobiculata and two new varieties, Actinodaphne sesquipedalis var.

glabra and Lindera concinna var. reticulata were recognised which are described here.

Actinodaphne

Actinodaphne sesquipedalis Hook. f. var. glabra Kochummen var. nov.

A varietate typica in foliis glaberrimis differt.

This new variety has glabrous leaves with 4-6 cm long petioles and large 2-4 cm

long glabrous bud scales. They are found in lowland and hill forests along the east

coast of Malaya.

Cinnamomum

Cinnamomum pubescens Kochummen sp. nov. Fig. 1.

Ramuli juveniles paulo angulosi flavo-brunnee pubescentes. Folia alterna vel opposita 3-nervata, petiolo

1-2 cm longo longe velutino. Lamina coriacea lanceolata 7.5-16 cm longa 2.5-6 cm lata viride-lutea, apice

acuto, infra subtiliter pubescenti, supra costa nervis secundariisque applanatis ad immersis, infra elevatis,

nervis secundariis nervulis lateralibus paucis ad marginem radiantibus nervum infra marginalem incon-

spicuum arcuatum formantibus, nervis tertiariis scalariformibus confertissimis, infra distinatis, supra

inconspicuis. Inflorescentia axillaris c. 10 longa, pubescens. Alabastrum globosum c. 4 mm longum,

pedicello c. 2 mm longo dense pubescenti, periantho subequali fere globoso utrique dense pubescenti.

Stamina 9 in seriebus 3, seriebus primis secundariisque extrosis tertiis glandulis 2 magnis cordatis in medico

filamenti affixis, filamentis dense pubescentibus, staminodiis stipitatis cordatis. Ovarium paulo rugosum,

stigmate decurrenti prominenti. Fructus immaturus in periantho infundibuliforme tube indurato rugoso

lobis persistentibus pubescentibus sessilis, pedunculo c. 8 mm longo. (TYPUS FRI 35248 [KEP]).

This is a small tree known only from 3 collections from the mountain forests at

Cameron Highlands in Pahang at about 1500 m.

Fig. 1.

Cinnamomum pubescens Koch. FRI. 35248.

Systematy of Malayan Phanerogams XX XI Lauraceae 25

Endiandra

Endiandra scrobiculata Kostermans ex Kochummen sp. nov. Fig. 2.

Arbor excelsa ad 33 m alta, caule 70 cm diam. cortex cinereo-brunnea laevis, intus rubra granulata

suaveolens, ligno juvenali luteo. Ramuli atro-brunnei ad nigri, juvenales leviter angulosi. Laminae coriacea

ellipticae ad lanceolatae 6.5-13 cm longa 2.5-5.5 cm latae, apice acuto, basi cuneata, costa supra applanata

nigrescenti, secundariis 7-9 paribus utrinque inconspicussime elevatis, reticulationibus inconspicue mani-

festis, petiolo 1-1.5 cm longo. Paniculae axillares. Flores ignoti. Fructus oblongi ca. 7 cm longi 4.5 cm

lati laeves atro-brunnei in sicco. (TYPUS: KEP 45477, 93366 [KEP]).

This is a rare tree known only from mountain forests at Frasers Hill, Pahang at

about 1200 m.

Kostermans gave this name to the collections at Kepong years ago, however he did

not publish it.

Lindera

Lindera concinna var. reticulata Kochummen var. nov.

A varietate typica in laminarium nervatura valde reticulata supra distincta quoque in ramulis dense

velutinis differt.

Differs from the type variety in the strongly reticulate venation which is distinct on

the upper surface of the leaf and in the densely velvety hairy twigs.

Locally common in mountain forests in Selangor and Pahang.

KEP. 45477

KEP. 93366

Fig. 2. Endiandra scrobiculata.

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Embryo Culture of Howea Palms

LIM-HO CHEE LEN, PHUA LEK KHENG, LOW NEOK CHEIN

Tissue Culture Laboratory, Singapore Botanic Gardens

AND

GOH CHONG JIN

Department of Botany, National University of Singapore

EFFECTIVE PUBLICATION DATE: 23 MAR 1992

Abstract

This paper reports the findings of an experiment on embryo culture of Howea belmoreana and Howea

forsteriana. It was demonstrated that growth regulators significantly affect the development of embryo

culture. The medium containing 1 mg/~! of 2,4-D and 0.5 mg/~! of 2iP was found to give the best

results for both of the Howea palms. Conventional germination of Howea palm seeds generally takes a

year or more (Reynolds 1982). In our experiments, the embryo culture required only 12-20 weeks to

develop fully rooted plantlets. This saving in time could be significant for commercial production.

Introduction

The Howea palms are ornamental plants which are arborescent monocotyledons

with solitary vegetative shoots. They are relatively slow growing and consequently

require much time and effort in production. Reynolds (1982) found that it normally

took a year or more to germinate a Howea seed in a special germination bed.

Because of its lack of a tendency to form branches, Howea palms cannot be multi-

plied by conventional methods of vegetative propagation. In 1986, the Department of

Botany, National University of Singapore, and the Parks and Recreation Department,

Ministry of National Development, undertook a joint research project on the tissue

culture propagation of tropical trees, palms and shrubs. The Tissue Culture Laboratory

of the Botanic Gardens was assigned the task of studying the tissue culture propaga-

tion of Howea palms.

The study proceeded in two stages. In the first stage, embryo culture was produced

from imported seeds of Howea palms. In the second stage, the plantlets and callus

derived from the embryo culture were used for tissue culture experiments. This paper

summarises our findings and observations of the first stage of the project: the develop-

ment of Howea palm embryo culture under the influence of different combinations

of growth regulators.

Materials and Methods

(a) Plant Materials

One thousand seeds each of Howea belmoreana and Howea forsteriana were im-

ported from Australia. These seeds were excised for the embryos. Before excision, the

seeds were soaked in tap water for 24 hours and then surface-sterilised by soaking for

15 minutes in 1% (w/v) sodium hypochlorite solution. The seeds were then sliced open

with an hand cutter and the exposed embryos were carefully removed with sharp-tipped

vA |

28 Gard. Bull. Sing. 43 (1991)

forceps. The excised embryos were then sterilised in a 5% chlorox solution for a few

minutes, rinsed with distilled water, and cultured on various culture media.

(b) Nutrient Media

For germination of embryos, the basal medium used was the Murashige and Skoog

salts (1962) supplemented with 170 mg/~' NaH,PO:2H,O, 100 mg/~' meso-inositol,

0.4 mg/~! thiamine-HCl, 3% (w/v) sucrose, 40 mg/~' adenine sulphate-2H,0, 50 mg/~!

activated charcoal, and 0.8% agar. Three types of auxin; 2,4-dichlorophenoxyacetic acid

(2,4-D), wnapthaleneacetic acid (NAA), and Indole-3-acetic acid (IAA), and two types

of cytokinin; N®-[A? isopentyl] adenine (2iP) and 6-benzylaminopurine (ae were

used in the following combinations in our experiments:

(i) 1 mg 2,4-D and 0.5 mg 21P;

(ii) 1 mg 2,4-D and 0.5 mg BAP;

(iii) 1 mg NAA and 0.5 mg 2iP;

(iv) 1 mg NAA and 0.5 mg BAP;

(v) 1 mg IAA and 0.5 mg 2iP;

(vi) 1 mg IAA and 0.5 mg BAP.

Three months after germination, the embryos were transferred onto an auxin/

cytokinin-free medium which contained only the basal salts as described above, but

at half the concentration.

The pH value of all the media was adjusted to 5.5 to 5.7. The media were dispensed

into 25 mm xX 150 mm culture tubes (10-20 ml of medium per tube) or 100 ml conical

flasks (20-30 ml of medium per flask) and sterilised for 15 minutes at 120°C under

1.5 ke cm? pressure.

The cultures were kept under 2-3 kIx lighting provided by true-lite tube on a 16-hour

photoperiod throughout the experiment. The environmental temperature was kept at

264120,

(d) Experiments Conducted

One thousand embryos each of Howea belmoreana and Howea forsteriana were

excised. For each of the two palms, 150 embryos were cultured in each of the six media

described in Section II (b). The remaining 100 embryos were cultured in the basal

medium without growth regulators. These last 100 embryos were used as controls for

the other experiments.

After three months, the germinated embryos were transferred to the auxin/cytokinin-

free medium. Thereafter, transfers at 4-6 week intervals were necessary to maintain

embryo/plantlet growth until they developed their first true leaf.

Results and Discussions

During the first three months of the experiments, the embryos exhibited 10 growth

patterns: no growth/contaminated; developed into swollen heads only (Fig. 1); deve-

loped into cotylendonary sheaths without roots (Fig. 2); developed into callus (Fig. 3);

developed into cotyledonary sheaths with roots (Fig. 4); developed into callus with

roots only (Fig. 5); developed into callus with shoots only (Fig. 6); developed into

shoots only (Fig. 7); developed into callus with shoots and roots (Fig. 8); and developed

directly into shoots and roots (Fig. 9). Some of these growth patterns had also been

Fig. 1 Embryos develop into

swollen heads only.

Fig. 2. Embryo develops into

cotyledonary sheaths without

roots.

Fig. 3. Embryo develops into

callus.

Fig. 4 Embryo develops into

cotyledonary sheaths with

root.

Fig. 5 Embryo develops into callus

with roots only.

Fig. 6 Embryo develops into callus with

shoot only.

Fig. 7 | Embryo develops into

shoot only.

Fig. 8 | Embryo develops into callus with shoot and

roots.

Fig. 9. Embryo develops into shoot and roots directly.

observed in date palm culture (Tisserat 1979, Tisserat 1982, Gabr and Tisserat 1985)

and oil palm culture (Hodel 1977, Nwankwo and Krikorian 1986).

Table la lists the percentages of Howea belmoreana embryos that took on each of

the growth patterns when cultured in the various media. Table 1b lists the correspond-

ing figures for Howea forsteriana. Both of the tables show that growth regulators are

important as the percentages of no-growth embryos for the control experiments are

very much larger than those for the other experiments. For the various media except

the control, the percentages of Howea belmoreana (forsteriana) embryos that showed

no growth are close to each other, ranging from 46.00-49.33% (13.33-30.00%). There

is a Slight hint that the media with 2,4-D/2iP, 2,4-D/BAP, and NAA/BAP favour direct

development into shoots and roots. However, at this stage, it is still not clear whether

the differences in the growth regulators significantly affect the success of the embryo

culture.

After the first three months, all the germinated embryos were transferred on the

auxin/cytokinin-free medium. After repeated transfers, some of the embryos developed

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34 | Gard. Bull. Sing. 43 (1991)

into normal plantlets, while others remained as roots, shoots, or callus only, or even

browned off and died. There was however a strong dependence of this later develop-

ment on the initial growth pattern of the embryo as shown in Table 2a, b. In general,

only those embryos which initially developed both shoots and roots stood a good

chance of developing into normal plantlets, and nearly all that initially developed into

swollen heads, cotyledonary sheaths, or callus with shoots or roots only, had browned

off and died.

Table 2a

Further developments of Howea belmoreana embryos in percentages

Browned off Callus Roots Shoots Normal

Embryos develop into and died only only only plantlets

swollen heads only 100

cotyledonary sheaths

without roots 85 5 10

callus 60 20 10 10

cotyledonary sheaths

with roots 98 2

callus with roots only 100

callus with shoots only 98 Ps

shoots only 70 30

callus with shoots and

roots 100

shoots and roots directly 100

Table 2b

Further developments of Howea forsteriana embryos in percentages

Browned off Callus Roots Shoots Normal

Embryos develop into and died only only only plantlets

swollen heads only 100

cotyledonary sheaths

without roots 92 3 5

callus (b 10 10 5

cotyledonary sheaths

with roots 95 5

callus with roots only 100

callus with shoots only 99 I

shoots only 80 20

callus with shoots and

roots 5 95

shoots and roots directly 5 95

ee)

Embryo Culture of Howea Plant 35

Considering only development into normal plantlets as success, we may work out

the success rate figures for each culture medium by summing the products of the

figures of the last column of Table 2a(b) and the corresponding figures of the columns

of Table la(b). For instance, the success rate figure for Howea belmoreana in the

medium IAA/BAP is given by

eet ee 6.07 4 01x 5.33: + 0: x 3.33 + 0 x 4.00

eee & 200 4 05. x 8.00.4 1.x 6.67 + 1x 6.67)% = 16.98%.

The success rate figures for the various media are tabulated in Table 3 and the cor-

responding bar-chart is shown in Fig. 10. It is evident that the media 2,4-D/2iP and

2,4-D/BAP yielded the best results, especially for Howea forsteriana. Howea belmoreana

appears to be not as sensitive to the media as Howea forsteriana. Our results agree

with Tisserat (1981) and Gabr and Tisserat (1985) who observed increased germination

rate and accelerated growth of date palms in 2,4-D/2iP medium, and also Nwankwo

and Krikorian (1986) who made similar observations for oil palms. However, both of

these previous studies used very much higher concentrations of 2,4-D (100 mg/~') and

2iP (3 mg/~'). Fig. 11 shows the various stages of normal growth of a healthy embryo

of Howea forsteriana over a period of eight weeks.

Table 3

Success rates of the media for the two Howea palms in percentages

Control 2,4-D/2iP 2,4-D/BAP NAA/2iP NAA/BAP_ IAA/2iP IAA/BAP

Howea

belmoreana 2.82 23.90 28.30 16.31 21.43 20.83 16.98

Howea

forsteriana 3.36 44.23 37.03 14.20 25.28 12.89 22.24

Our observations suggest that the size of the embryo could be a factor affecting

its growth. Most of the embryos that died or showed no growth at all were very small

in size. Those smaller embryos that survived generally took a long time to show any

sign of growth, and they often developed into swollen heads, callus, or shoots or roots

only. On the other hand, the large embryos in suitable media tended to grow within

three months into normal plantlets. It is possible that the size of the embryos is an

indication of the age of the embryos. The age of the embryo and the condition of the

explant have been shown to have an effect on the growth pattern of date palm and

oil palm culture (Jones 1974, Hodel 1977, Zaid and Tisserat 1983). Unfortunately,

since the seeds were imported, we did not have information on the age and quality

of the embryos used in our experiments. We therefore cannot draw a definite conclu-

sion on this aspect about Howea palms.

Conclusions

This study demonstrates that growth regulators are important for embryo culture of

Howea belmoreana and Howea forsteriana. Of the combinations tested, the medium

containing 1 mg/~! of 2,4-D and 0.5 mg/~! of 2iP gave the best results for both Howea

palms.

dVd/VVI

‘sued OM} S42 JO JINI[ND OAIQUID DY} JO} BIPSUI dy] JO areI ssadonG = QT. “BI

DUDIAAaJSAO{ DAMOY NK]

d!¢/V VI

dVd/VVN

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. \ ,

Fig. 11. | The development of an excised embryo of Howea forsteriana in the presence of growth regulator

over a 40-day period.

Germination of Howea palm seeds normally takes a year or more (Reynolds 1982).

In our experiments, the embryo culture required only 12-20 weeks to develop fully

rooted plantlets. This saving in time could be significant for commercial production.

Similar results were obtained by Hodel (1977) for oil palm, which took 20-24 weeks

to grow from seed to seedling, but only 10-12 weeks to achieve the same stage of

development in embryo culture.

The age of the embryos may also affect the growth of the Howea embryos. This

possibility is worth a further study. A better knowledge of the physiology and the

biochemistry of the embryo will also be helpful in understanding the requirements of

embryo growth in vitro.

References

Gabr, M.F. and Tisserat, B. (1985). Propagating Palms in Vitro with Special Emphasis

on the Date Palm (Phoenix dactylifera L.) Sci. Hort., 25, 255-262.

Hodel, D. (1977). Notes on Embryo Culture of Palms. Principes, 2, 103-108.

Jones, L.H. (1974). Propagation of Clonal Oil Palm by Tissue Culture. Oi/ Palm News,

17, 1-9.

Murashige, T. and Skoog, T. (1962). A Revised Medium for Rapid Growth and Bio-

assays with Tobacco Cultures. Physiologia Planta, 15, 473.

Nwankwo, B.A. and Krikorian, A.D. (1986). Morphogenetic Potential of Embryo- and

Seedling-Derived Callus of Elaeis guineensis Jacq. var. pisifera Becc. Ann. Bot.,

51, 65-76.

Reynolds, J.F. (1982). Tissue Culture in Forestry. Martinus Nijhoff/Dr. W. Junk

Publishers.

Tisserat, B. (1979). Propagation of Date Palm (Phoenix dactylifera L.) in Vitro. J. Exp.

Bot., 30, 1275-1283.

38 Gard. Bull. Sing. 43 (1991)

. (1981). Date Palm Tissue Cultures in Advances in Agricultural Technology,

Western Region, Ser. No. 17, USDA, ARS, 1-50.

_____. (1982). Factors Involved in the Production of Plantlets from Date Palm Callus

Cultures. Euphytica, 31, 201-214.

Zaid, A. and Tisserat, B. (1983). In Vitro Shoot Tip Differentiation in Phoenix

dactylifera L. Date Palm J., 2, 163-182.

Schizostachyum terminale Holtt., An Interesting

New Bamboo Record for Borneo

K.M. WONG

Forestry Research Centre

Forestry Department

P.O. Box 1407

90008 Sandakan, Sabah

EFFECTIVE PUBLICATION DATE: 23 MAR 1992

Abstract

The bamboo Schizostachyum terminale Holtt., first recorded and for more than 30 years known only

from Peninsular Malaysia, has been documented for north Borneo. It has a clambering, thicket-forming

growth habit and an unusual preference for seasonally inundated swampy riverbanks and alluvial flats.

Introduction

In 1956, Holttum described Schizostachyum terminale, known only from the type

specimen Nauen SFN 35831 (K, SING), collected on the banks of the Krian River in

Kedah state, Peninsular Malaysia. A second Malayan collection was made only in 1982

(Wong FRI 32399 — K, KEP, L, SING) from a damp logged-over patch of vegetation

beside a stream at Rantau Panjang in Selangor state. Both collections were from the

west coast of Peninsular Malaysia and the evidence then indicated a rather restricted

distribution. Since 1982, searches along the Krian River and various other rivers near

the Malayan west coast have ended in vain, suggesting the species was rare and that

its survival was possibly threatened. In 1988, a third collection (Saw FRI 36283) was

made from riverine swampy ground at 70 m above sea level, quite inland at the Krau

Game Sanctuary near Kuala Lompat in Pahang state.

Recently this species has been discovered in north Borneo, and investigations of wild

populations there in the East Malaysian state of Sabah (voucher: Wong FRI 35151 —

K, KEP, SING) (Fig. 1) and in Brunei Darussalam (voucher: Wong WKM 2096 —

BRUN, K, L, SING) have yielded more information on the growth habit, habitat

requirements and general distribution which were hitherto poorly understood. These

are discussed in the present paper.

Growth Habit

S. terminale is a small clambering bamboo occurring as clumps, each with 3-10

culms of 8-15 mm diameter. As in other species of the genus, the culms are white

appressed-hairy on the internodes, slightly white-waxy, and arise from a sympodially

constructed rhizome system. This species differs from other Malayan and Bornean

Schizostachyum species in having culms which clamber over the surrounding vegeta-

tion. This clambering habit is possible through elongation of the primary branch at

several nodes, which reiterates the original culm in morphology and habit, and which

Fig. 1. | Flowering leafy branch of Schizostachyum terminale.

in turn produces further such branches of a higher order. This system of reiterating

branches allows the bamboo to clamber and entangle with nearby tree branches, a

situation akin to that in Dinochloa bamboos (Wong, 1986), except that the culms do

not twine.

Occasionally the base of the reiterating branch is thickened and produces roots as

does a rhizome, and it is conceivable that when such parts come into contact with the

ground, vegetative reproduction is facilitated. In other Schizostachyum species, the

branch complement also arises from a single primary branch axis at the node, but

when fully developed all branches at a node are of subequal size and a dominant is

not easily distinguishable; however, development of “aerial rhizomes” (arising from the

rooting of thickened branch bases) is also known in other species such as S. /atifolium

Gamble, especially when the culm tip is damaged (Wong, 1990). In S. terminale, the

branch complement consists of a primary axis which frequently remains dormant

initially but later develops to reiterate the culm, and several small leafy branches arising

from the basal nodes of this primary axis. In addition, reiterative branch elongation

et ee

Schizostachyum terminale 4]

occurs even without any damage to the original culm. In this way, thickets or curtains

of this bamboo arise.

Habitat Conditions

In Sabah, S. terminale grows on the seasonally flooded banks of the Kinabatangan

River near Tanjung Bulat, at about 15 m above sea level (Fig. 2). Thickets of it drape

the vegetation in riverbank forest dominated by the trees Octomeles sumatrana and

Terminalia copelandii. From the mud markings on the vegetation, this forest can be

flooded some 2-3 m high. Inundation can last up to several weeks at a time (C.F. Tan,

pers. comm.)

In Brunei, S. ferminale is abundant along low wide stretches of the Belait River

(Fig. 3) but above the mangrove or nipah (Nypa fruticans) zones, at 15-20 m above

sea level. There it grows in the seasonally inundated swampy banks and alluvial flats

together with Syzygium spp., and the common riverbank rattan Daemonorops fissa.

The waters can rise some 2 m higher during heavy rains, especially around May-June

and September-November. The species is absent in forest away from such seasonally

inundated riverine zones.

Bamboos are generally known to occur only away from swamp habitats, but S.

terminale is a clear exception. Its distribution only in the habitats recorded above

suggests its restriction to swampy ground and a tolerance of prolonged inundation.

Elsewhere, Chusquea paludicola Clark is known to inhabit highland Sphagnum peat

bogs above 2,000 m altitude in Costa Rica (X. Londono, pers. comm.; Clark, 1986),

although in those situations such prolonged severe inundation does not occur.

“i

.-%

= -_-

- — w

Fig. 2. Undergrowth thickets of Schizostachyum terminale along the flood-prone bank of the

Kinabatangan River, Sabah.

Fig. 3. Tangles of Schizostachyum terminale on the bank of the Belait River, Brunei.

Acknowledgements

This account is based on part of the research carried out for a Ph.D. programme

under the supervision of A.L. Lim at the University of Malaya. I am grateful to the

curators of the herbaria at Kepong and the Singapore Botanic Gardens for permission

to consult the collections there, and to P.S. Shim and C.F. Tan for the opportunity to

visit the Kinabatangan River in Sabah.

References

Clark, L.G. (1986). Eight new species of Chusquea (Poaceae: Bambusoideae). Jowa State

J. Res. 61(1): 99-122.

Wong, K.M. (1981). Flowering, fruiting and germination of the bamboo Schizostachyum

zollingeri in Perlis. Mal. For. 44: 453-463.

. (1986). The growth habits of Malayan bamboos. Kew Bull. 41(3): 703-720.

. (1987). The bamboos of the Ulu Endau area, Johore, Malaysia. Mal. Nat. J.

41: 249-256.

. (1990). The growth architecture and ecology of some tropical bamboos. J. Amer.

Bamboo Soc. 8(1 & 2): 31-46.

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