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3 (CG) JUL 31 1995 ARBORETUM

The Gardens' Bulletin

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VOL. 46 (Part 1) June 1994 ISSN. 0374-7859

CONTENTS

CORNER, E.J.H.

On The Agaric Genera Hohenbuehelia and Oudemansiella

NE AED Oe ee ea PLT oe Se senicansotnbcdvmapendensys ys <nesvoas he 1 — 47

CORNER, E.J.H.

On the Agaric Genera Hohenbuehelia and Oudemansiella

ae: Ootemuansicila Spee. 28 Bassi en SU ths avian eget 49 — 75

KENG, H.

Flora Malesianae Precursores - LVIII, Part Four

Pee ens Scluma Ci ncarese) ty alewin. 2 acai bo en i RAE 77 — 88

LATIFF, A.

Pe ee ee ieee cha © ate ie 8 A 7 9 Re ieee 89 — 103

STONE, B.C.

Citrus Fruits of Assam: A New Key to Species, and Remarks on

Citrus assamensis Bhattacharya and Dutta, 1956 .00...........eccecceeseeeeeeeeeeeeeeeees 105 — 112

STONE, B.C.

Additional Notes on the Genus Glycosmis Correa (Rutaceae) ............::::c0e 113 — 119

STONE, B.C.

Supplement to the Rutaceae in Peninsular Malaysia ....................c0:c0000 121 — 140

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On The Agaric Genera Hohenbuehelia and Oudemansiella

Part I: Hohenbuehelia

E.J.H. CORNER

91 Hinton Way

Great Shelford

Cambridge CB2 5AH

England

Abstract

The construction of the fruit-body and the affinity of the genus are discussed. It is considered that it

comes between Pleurotus and Oudemansiella, having the basidia of the former and the pleurocystidia of the

latter. Eighteen species are described for Malesia, of which 15 are new. Notes are given on several extra-

Malesian species. New taxa: H. concentrica, H. griseipendens, H. incarnata, H. lanceifera, H. malesiana, H.

mellea, H. minutissima, H. pachyhyphata, H. pahangensis, H. perstriata, H. quadruplex, H. singaporensis, H.

suppapillosa, H. vermiculata. H. bullulifera Singer v. brasiliensis. New combinations: H. cystidioides

(C.G.Lloyd), H. subtorulosa (Cke).

Contents

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Be eRe ARCA BDEGIES oi oo ino is 55 iy- sal ca vtgule cas ode sbeapvecesesssapesscneses 38

ae es Neon aa yee a intn a kas oa dus Vag pier estab dient bao ase ntnnsed seasceascanierss 46

NS rds Oe NTA oe i th wan ois be Sa -ciee vos 4- “ch gaan chad det ab ds do wkagls Seda ecdanews donesensyss 75

General Account

A fruit-body with lateral pileus, an upper gelatinous layer to the flesh, and a

farinaceous smell of meal is a sign of Hohenbuehelia. It is checked microscopically

for the thick-walled pleurocystidia. However, there are specific variations in

most features and what seems to be a uniform genus has ragged fringes.

Fruit-body. Two European species have mesopodal fruit-bodies, namely H.

longipes (among moss in woodland) and H. culmicola (on remains of the coastal

dune-grass Elymus). At the other extreme, there are fruit-bodies which are

dorsifixed, more or less sessile, and at first cyphelliform, as H. bullulifera, H.

chevalieri, H. griseipendens, H. nigra, and H. singaporensis. Such species

resemble Resupinatus which lacks the thick-walled pleurocystidia.

Flesh. The upper gelatinous layer is usually thinner than the firm layer of the

flesh but it may be as thick or thicker. In a few species the flesh is almost

completely gelatinous. In contrast, the gelatinous layer is said to be poorly

developed or little differentiated in the South American H. roigii and H. spegazzinii.

Concerning the absolute thickness of the gelatinous layer, there must be some

caution in comparing descriptions. I measure its maximum thickness at the base

of the lateral pileus, as I do for the flesh, and this may be the customary procedure,

but Singer measures it towards the margin of the pileus. In this case, it is

necessary to state whether it is an old or young pileus because the old pilei

become marginally attenuate and may give a narrower measure than the young.

The gelatinous layer swells abnormally in material preserved in alchohol-formalin

and, also, in sections mounted in dilute potash. Sections of dried material

mounted in water give a more or less normal thickness.

Spores. Most species have smooth and thin-walled spores as seen under the

light microscope but there is one exception with verruculose to echinulate and

thick-walled spores, namely H. bursaeformis. For this, Singer (1969) made the

subgenus Reidia and Horak (1981) has treated it as the distinct genus Conchomyces

v. Overeem, while remarking oddly that Hohenbuehelia is characterised by

allantoid cylindric spores. Singer added a second species, H. dimorphocystis,

which Horak reduced to his C. bursaeformis. However, there are difficulties in

accepting this genus. Donoso (1981) showed that the spores of some species of

the genus have a thin mucilage sheath beneath which the endospore, under SEM,

is subverrucose, e.g. H. geogenia, H. heterosporica, H. petaloides and H. rickenii:

in the case of H. bursaeformis, the spines penetrate this sheath. Then, according

to Donoso, H. heterosporica has both smooth and subverrucose spores (as seen

under SEM), even on the same basidium. I find with Malesian material of H.

bursaeformis that the spores under the light microscope vary from echinulate to

merely asperulate and that these spines or warts contract in Melzer’s iodine and

disappear in dilute potash, thus unlike typical spines. In other words, there is a

gradation from echinulate spores (of a sort) to truly smooth spores, which nullifies

the main idea of a separate Conchomyces.

The spores of the genus are often said to be 1-guttate but, in my experience,

fresh spores are usually aguttate and, on drying, the contents contract into a sort

of gutta, though a few species have fresh spores with 1-2 minute guttulae.

Cheilocystidia. These make a sterile gill-edge and are nearly always thin-

walled. They vary in different species from clavate to cylindric-moniliform and

lecythiform, or ventricose. They would seem to supply useful specific features

but they may vary considerably even in the same fruit-body.

encrusted distally with granules or crystals. In most species they are not dextrinoid

but Stevenson (1964) has described several species from New Zealand with

Pleurocystidia. These are usually called metuloids because the thick wall is

dextrinoid walls. In H. bursaeformis the wall is scarcely thickened and not or

slightly encrusted. In two species, namely H. horrida and H. incarnata, the

pleurocystidia are extremely abundant. The wall thickens from apex to base,

young pleurocystidia being thin-walled. They are the first hymenial elements to

develop and have, in consequence, a deep tramal origin in the mature gill. Their

length may vary because the first to be formed are at the base of the gills and

between them and are the longest, and they shorten towards the gill-edge where

they may be almost hymenial and, even, mixed with the cheilocystidia.

Hyphae. These are short-celled and, usually, with little inflation though

strongly inflated in the flesh of some species. In others they develop thickened

walls and may become almost solid, as in H. pachyhyphata; this species has

ampulliform swellings on some hypae which resemble the inflated cells in

Chaetocalathus (Corner, in ed.). The gelatinous layer begins to form almost as

soon as the pileus. It separates a narrow superficial layer, one or two to ten

hyphae thick in different species, from the tissue of the flesh (Fig. 12). The

superficial layer develops a loose sterile hymenium with variously formed cells

and, in most species, narrow hyphae grow slowly out to project vertically to form

the villous surface; thus it is more evident in the proximal part of the pileus.

In H. quadruplex (Fig. 9) the gelatinous layer is double. In a few species, such as

H. suppapillosa (Fig. 11,12), there are many inflated cells in the gelatinous layer

as well as the usual uninflated hyphae. The gill-trama is constructed from

descending hyphae but in H. angustata there seems to be a vestige of the radial

construction seen in Panus s.str.

Surface of pileus. The loose sterile hymenium on the surface may develop

cells like cheilocystidia or pleurocystidia but in some species, especially those

with the fruit-bodies at first cyphelliform, there is a more or less close hymenioderm

of clavate cells, e.g. H. bullulifera, H. griseipendens, H. nigra, H. singaporensis

and H. subbarbatus (Dennis, 1953). Such a hymenioderm is used in modern

mycology to distinguish genera but has not aroused that attention in Hohenbuehelia.

Nematode-catching. This habit has been explained in detail by Thorn and

Barron (1986). They associate 9 species of the mycelial genus Nematoctonus

Drechsler, which has clamp-connections, with some 14 species of temperate

Hohenbuehelia. The nematodes are caught by a viscous excretion from short

hyphal processes, often lecythiform, and the mycelia may form solitary conidia

or chlamydospores. Tropical species remain to be tested.

Classification of the Species

Several ways have been proposed to arrange or to ally the species, but none

seems, as yet, to be satisfactory. Huijsman (1961) employed the relative thickness

of the gelatinous and firm layers of the flesh and the direction of the hyphae in

the gelatinous layer, whether it was ascending more or less vertically to the

surface, longitudinal (radiating) parallel with the hyphae of the firm layer, or

interwoven, His conclusions were drawn from rather few temperate species and,

as Singer has remarked, need extension to those of the world. Singer (1975)

distinguished subgen. Reidia with ornamented spores from subgen.

Hohenbuehelia with smooth spores but, as Donoso (1981a,b) has shown, the

distinction breaks down with SEM examination. For the Malesian species I have

used primarily the presence or absence of thick-walled cystidia like pleurocystidia

on the surface of the pileus because I find that it is a constant character. However,

though I have perused descriptions of other species, there remain numerous

doubts about its universal applicability.

Conchomyces was considered by Horak (1981) to be unrelated to any other

agaric genus on account of the echinulate, subglobose and rather thick-walled

spores, the large cheilocystidia, the rather thin-walled pleurocystidia and the

inflated hyphae of the firm tissue of the flesh. It is difficult to follow this

argument because all the points have their gradations in other species of

Hohenbuehelia with which it agrees in colour and gelatinous upper layer to the

pileus. The one species, H. bursaeformis, has been regarded as peculiar to the

southern hemisphere of Australasia and Chile, but its occurrence in Malaya and

north Borneo does not accord. Maybe it has traces of the stouter fruit-body of

Oudemansiella.

Basidiograph and Pleurocystidiograph

In Graphs 1 and 2, the ratio //w for basidia and pleurocystidia is plotted

against /, where / is the length and w the width of these structures as given in the

following descriptions for their mean values. The result is peculiar. The basidia

conform to a steep locus as drawn, roughly, by the line through the triangles, and

the pleurocystidia conform to a more gradual locus shown, roughly, by the line

drawn through the circles. However, there are two outstanding exceptions, for

the pleurocystidia of H. horrida (point h) and for one of the measures for H.

testudo.

In the equation to these loci //w = a + bl, bis 0.112 for the basidium and 0.015

for the pleurocystidium; the reciprocal of b gives the maximum width as 8.9 um

for the basidium and 66.7 um for the pleurocystidium. These values are based on

mean measures; the widest basidium that I have recorded was 10 um for H.

concentrica (mean 9 lum); the widest pleurocystidium was 28 Um in H. angustata.

Thus, the basidia of Hohenbuehelia are extremely narrow, even narrower that

the clavarioid basidium with b = 0.08 (1/b = 12.5 um). The pleurocystidia,

however, conform to the wider basidium seen in Amauroderma and Ganoderma

with b = 0.013 (Corner 1983). The hymenium of Hohenbuehelia seems to be

influenced in two ways; one is primitive and retains the wide or massive expression

and the other is far advanced towards the minimal narrowness of the uninflated

hypha.

R pleurocystidia

10 20 30 40 $0 6a 70 80 qo ido 10

Graph 1. Hohenbuehelia, basidiograph (triangles) and pleurocystidiograph (circles);

h, H. horrida; t, H. testudo. (Constructed from the data in this article).

basidia

10 é

20 40 69 80 100 120 {4Q {60

Graph 2. Hohenbuehelia, pleurocystidiograph for the longest pleurocystidia.

The situation in Pleurotus is instructive. In dealing with the Malesian species,

I distinguished species with narrowly ellipsoid to cylindric spores from those

with broadly ellipsoid to subglobose spores (Corner 1981). Graph 3 gives the

basidiographs of these two groups. That with narrow spores has the steeper slope

for which, roughly, b = 0.105 (1/b = 9.5 Um) which is comparable with

Hohenbuehelia; and that with broad spores has, roughly, b = 0.045 (1/b = 22.2

uum) which is intermediate to the large basidium of Oudemansiella with b = 0.02

(1/b = 50 um). That is to say, in comparision with Hohenbuehelia, Pleurotus is

progressing to the narrowness of the basidium with little tendency to retain the

pleurocystidium as the larger unit. The basidiograph of both genera corresponds

with that of the developing basidia of Oudemansiella with b = 0.11 (Corner,

1947).

These conclusions are tentative because they are based on mean measures and

not on true averages of many individual basidia. Moreover, in the case of the

pleurocystidium of Hohenbuehelia it is not always clear that the width refers to

that between the outer walls or to the overall width including the incrustation.

Nevertheless, it seems that the pleurocystidia of H. horrida conform with both

loci. The genus is less advanced in this respect than Pleurotus.

(0 20 30 40 $0 60 70 $0

Graph 3. Pleurotus, basidiograph. Species with narrow spores as triangles with locus

n. Species with broad spores as circles with locus b. L, Lampteromyces

japonicus. (Constructed from the data in Corner 1981).

Affinity

Modern taxonony places Hohenbuehelia along with Resupinatus (without

thick-walled pleurocystidia) in Tricholomataceae tr. Resupinateae. This

classification emphasizes the gelatinous texture of the flesh. As regards

Resupinatus, | have given reason to suppose that it is a mixture of marasmioid

fungi with subacerose basidioles and pleurotoid fungi without them (Corner, in

ed.). Concerning Hohenbuehelia, there are several species which fail in one way

or other the dual test of a gelatinous layer in the flesh and thick-walled

pleurocystidia. That its species are mostly pleurotoid seems not to matter because

H. culmicola and H. longipes are mesopodal.

Both diagnostic characters emphasize the difference from Pleurotus. An

analogy, however, seems to dispose of a family distinction. Polyporus s. str. and

Echinochaete have the same peculiar hyphal construction but Echinochaete has

an upper gelatinous layer to the pileus and thick-walled pleurocystidia, both

features lacking in Polyporus s. str. (Corner, 1984). Echinochaete has been

regarded as a synonym of Polyporus subgen. Asterochaete by Singer (1975).

Thus, the affinity of Hohenbuehelia may well concern Pleurotus, some species of

which have thin-walled pleurocystidia (Corner, 1981); moreover, Hohenbuehelia

has been regarded as a subgenus of Pleurotus by several authors. The faculty of

nematode-catching has been found, also, in several species of Pleurotus (Thorn

and Barron, 1986), but not reported for Resupinatus.

Another genus, in this connection, is Agaricochaete which has been re-

established with a third species by Pegler (1977). The fruit-bodies are mesopodal

and clitocyboid without gelatinous flesh but with the thick-walled pleurocystidia

of Hohenbuehelia. Emphasizing these pleurocystidia, Peglem placed the genus

in Resupinateae, and here enter two other species of South America, H. roigii and

H. spegazzinii. Their fruit-bodies have no distinctly gelatinous flesh but they

have the thick-walled pleurocystidia, and they are pleuropodal or sessile. If

emphasis is laid on pleurocystidia, then it is difficult to see how these two can be

separated from Agaricochaete, which would then become practically synonymous

with Hohenbuehelia.

The contrast is shown by H. bursaeformis with the gelatinous upper layer to

the flesh but with thin or but slightly thick-walled pleurocystidia. One is led to

think of Oudemansiella with mesopodal fruit-bodies and large spores, as if

antecedent, and it has a species with echinulate spores. Agaricochaete is evidently

rare and, maybe, there are other tropical species of this general alliance to be

discovered. The fourth species attributed to the genus, A. indica Natarajan et

Raman (1986), without clamp-connections, is excluded by Thorn and Barron who

think that it may be Lactocollybia.

Another alliance has been suggested with Geopetalum Pat. or Faerberia, as it

seems it must now be called (Pouzar, 1981), but I use the old name as that of past

literature. The fruit-body of G. carbonarium has the stature and colour of

mesopodal Hohenbuehelia and the thick-walled pleurocystidia, but it lacks the

gelatinous layer to the flesh. It is dimitic in the manner of several species of

Pleurotus and has cantharelloid gill-folds with thickening hymenium (Corner,

1966, 1968); moreover, it is not nematode-catching. A minor distinction may be

that, instead of the prevailing farinaceous smell of Hohenbuehelia, it disengages

HCN. Nevertheless, if the cantharelloid gill-fold preceded the agaric, Geopetalum

could have led to Pleurotus and, so via Agaricochaete to Hohenbuehelia. Whereas

Cantharellus is excluded from Agaricales by modern taxonomy, it consistently

admits the cantharelloid Geopetalum. It has been ranked with Hohenbuehelia in

Geopetalaceae, to the exclusion of Resupinatus, by Jiilich (1981). The common

character is the thick-walled pleurocystidium, not the gelatinous layer in the

flesh; the other characters, given by Jiilich, such as multiguttulate basidia and

spores, do not occur in my experience in Hohenbuehelia.

Three other pleurotoid genera have, at least in some of their species, thick-

walled pleurocystidia. Of these, Chaetocalathus and Panus lack the gelatinous

upper layer to the flesh but it is present in some species of Panellus. This genus

and Chaetocalathus, however, are marasmioid with subacerose basidioles. Panus

has, in the main, radiate gill-construction which may be a relic of the cantharelloid.

If one attempts to relate these genera, dispersed in various tribes or families,

in a phylogenetic tree, it becomes obvious how much must have disappeared in

the long course of evloution. I am forced to conclude that modern taxonomy still

needs modernising.

Hohenbuehelia Schulzer (1866) - Generic Character

Fruit-bodies pileate, mesopodal, pleuropodal or sessile to dorsifixed and at

first cyphelliform, gymnocarpic. Gills decurrent or radiating from the point of

attachment, thin, sometimes dichotomous. Flesh with an upper gelatinous layer

and a firm lower layer, in a few species almost wholly gelatinous. Smell often

farinaceous. Spores white, mostly smooth but varying verrucose to echinulate,

inamyloid. Hymenium not thickening, without subacerose basidioles; gill-edge

sterile with thin-walled cheilocystidia. Pleurocystidia typically thick-walled and

encrusted, usually not dextrinoid. Hyphae clamped, monomitic, short-celled,

inflating or not, inamyloid. Surface of pileus often villosulous with excrescent,

often faciculate, hyphae; pileocystidia thick-walled, present or not; in some species

hymenioderm with clavate cell. Colour of pileus usually fuscous brown, grey or

pallid subochraceous to whitish, either in the walls of the superficial hyphae or in

the sap of the upper hyphae of the firm layer of the flesh. Lignicolous or on

plant-remains, a few terricolous, associated (? always) with nematode-catching.

Worldwide.

Keys to the Species of Hohenbuehelia in Malesia

Surface of pileus with thick-walled cystidia ................ccssscceeeseeeeeeees Group 1

iio PIlCUS WIINOUL SUCK CYSLICGIA......................seccceelecnesecssossens Group 2

Group 1

Flesh almost wholly gelatinous. Pileus -14 mm wide, with subdiscoid base. Spores 4.5-6 x 3.3-4.3 um.

Selec Yella MOSY ClaVate. SOLOMON TSANG 4 oo05. c....00.sccccncinnecssersconssscconsceenaneesecssesetecta’ H. subdiscipes

Flesh with a distinct firm layer below the gelatinous layer. Cheilocystidia mostly ventricose with short,

often capitate appendage.

2. Gills distant, -3.5mm wide. Pileus -22mm in radius, subsessile, striate. Flesh -2mm thick. Spores 5-

a umanere RMN PMC UNLIURRINIR SN AMMCEIND RC ooh il da dct e cal sawci easvatacadivauncics sanpduetsroourveresbuves H. perstriata

2. Gills crowded. Pileus sessile or pleuropodal.

3. Pileus -7mm wide, greyish buff. Gills -0.7mm wide. Flesh with the gelatinous layer thicker than

the firm layer. Spores 4-6.5 x 3-4 um. Pileocystidia very heavily encrusted. Solomon Islands.

Ee Sout ae ERS Se Ay, RS I Fad RA alt “SEAR Seen et os iat, NER ke H. vermiculata

3. Pileus larger. Gills wider. Gelatinous layer thinner than the firm layer.

4. Cheilocystidia clavate to obtusely ventricose. Spores 3.5-4.3 x 3-3.5 um. Malaya..............

Lo kae Rl | ee GERAD Sit eee) 2) SUNS ear ee eS AONE SEP, hal. a H. pahangensis

4. Cheilocystidia often ventricose with capitate apex. Spores 5-7 x 2.5-3.5 um. S.E.Asia........

Dae ae aa as Se eR RN hy Sod A AR wend once chy cade vsenwndewe H. testudo

Group 2

Hyphae of the firm layer of the flesh and of the gill-trama with walls 1-3 thick or more. Spores smooth.

2. Firm layer of the pileus much thinner than the gelatinous.

3. Pileus -3mm wide, grey. Spores 5-7 x 2.7-3.5 um. New Guinea ...................6 H. minutissima

3. Pileus -45mm wide, greyish to brownish bistre, occasionally white. Gills waxy gelatinous. Spore

Sg Og eS aa eg) ee en ie a ee H. malesiana

2. Firm layer of flesh thicker than the gelatinous layer (at least in the proximal part of the pileus).

4. Pileus -30mm wide, white to dingy yellowish or fuscous vinaceous. Stem short, thick. Smell

farinaceous. Spores 6-7.5 x 3-3.5um. Hyphae of firm layer of flesh with ampulliform

CTO S.) PRONTNOG, CHRECTISIAIAG |.) s,s. éctery sotecsmn> cheat cima cbovobdetsinspsdesevsnsecassners H. pachyhyphata

4. Pileus - 13mm wide, white to greyish umber. Spores 4-5.5(-6) x 3.3-3.7 um. No ampulliform

fear Te EOIN 0 9S, do ud dne 9. <5 ms Re PSs CEPI MEL aol ol Sabccslssssesvusstbeosesss var. minor

Hyphae with thin or scarcely thickened walls.

5. Spores echinulate, subglobose. Pileus white to pale ochraceous. Malesia ................... H. bursaeformis

5. Spores smooth.

6. Pileus dorsifixed, at first cyphelliform, expanding to 25mm wide or less, fuliginous grey.

Flesh 1-2mm thick, gills -1.5mm, crowded. Cheilocystidia 15-30 x 6 -10(-15) um, clavate.

Surface of pileus more or less hymeniiform.

7. Spores 6-7 x 4-4.5 um. Pleurocystidia 12-25 um wide. Clavate cells on pileus 8-20 um wide,

ERM 0 Bb 550. 35s ethan Sica ahc tasty betas Nip) Wid i dsgavan vanilts casts pl gattonenesvo aes H. singaporensis

7. Spores 4-5 x 3.5-3.7 um. Pleurocystidia 7-14 um wide. Clavate cells -7 um wide, some

lobuiates scattered on pileus. Borneo ................:..0.<..0<.0.sessunee ne H. griseipendens

6. Pileus pleuropodal or laterally sessile from the first.

8. Pleurocystidia exceedingly numerous, lanceolate, 5-12(-14) lum wide. Gills more or less

distant, 4-6mm wide. Pileus - 7cm wide, shortly pleuropodal.

9. Wholly pinkish buff. Gelatinous layer of flesh - 150 um thick. Spores 4-7 x 4.5-6

uum, subglobose, smell slight, of fenugreek. Solomon Islands ............. H. incarnata

9. White to brownish ochrceous or greyish. Gelatinous layer 250-600 tum thick. Spores

5-6.5 x 3-4 um, ellipsoid. Smell farinaceous. Malesia .................04. H. horrida

8. Pleurocystidia numberous but not crowded, ventricose-fusiform. Gills 1.5-3mm

wide or -4mm.

10. Pleurocystidia with long attenuate apex, 40-160 x 12-20 um. Flesh almost

wholly gelatinous, firm layer 100-200 um thick. Gills pale ochraceous to

brownish. Pileus -20mm wide. Inodorous.

11. Gills crowded. Pileus white to pallid ochraceous. Spores 4-5.5 x

3. 7-423 Pins ROMEO ook os sco cee se <aek <acen steno H. lanceifera

11. Gills distant. Pileus honey yellow to brownish, not villous. Spores ? 5-6.5

XK 2.3-3 pee SMBAPOLE faceceicses:.-ccceccncnns codetee eee Renin meee ae H.mellea

10. Pleurocystidia not long attenuate, shorter. Gills crowded.

12. Gills -4mm wide. Pileus -8cm wide, ochraceous to orange. Flesh more

than Imm thick

13. Gills white. Pileus white to ochraceous and brownish, shortly

stipitate. Flesh 1.5-4mm thick, gelatinous layer - 1mm thick. Spores

6-7 x 4-5 um. Inodorous. Malaya, Sarawak ...... H. suppapillosa

13. Gills pale ochraceous buff. Pileus light orange to brownish

ochraceous, with concentric ridges. Flesh 3-5mm thick, gelatinous

layer 1-2mm thick. Spores 8-8.5 x 6-6.7 um. Smell farinaceous.

POAT BING i gs nh shai siceanadtes souedt stn sdb odaehemiaedanen inal H. concentrica

12. Gills -lmm wide, white to pale ochraceous buff. Spores 4-6.5 x 3-4 tum.

14. Pileus -8mm wide, greyish, sessile. Firm layer of flesh 50-100

uum thick, gelatinous layer much thicker. Solomon Islands ...

a2 schgi peat tg Xie saadllaiaiPan-diap Vsnsbieo ake aaa H. vermiculata

14. Pileus -30mm wide, pleuropodal, fuscous fawn, smooth. Flesh

with two gelatinous layers. Solomon Islands ..H. quadruples

Hohenbuehelia bursaeformis (Berk.) Reid Figure |

Kew Bull. 17 (1963) 304; Singer, Nova Hedwigia Beih. 29 (1969) 62.

Conchomyces bursaeformis (Berk.) Horak, Ann. Mycol. ser. II. 34 (1981) 109; New Zeal. J. Bot. 9

(1971) 458.

Pileus -6cm in radius, -8cm wide, sessile or with a short and more or less

lateral stem, at first convex or cucullate, generally descending, then applanate,

smooth except the shortly villous base, white to pale ochraceous bistre or drab

fawn buff, striate towards the incurved, minutely scurfy margin. Stem 0-5 x 1-3

mm, lateral or nearly so, villous. Gills decurrent, very crowded and thin, very

narrow, almost contiguous, 13-30 primaries 1-2mm wide, 5-7(-8) ranks, rarely

dichotomous, shining white, edge entire. Flesh 2-4mm thick at the base of the

pileus, fibrous firm then rather spongy, with a thin gelatinous layer 200-500 um

thick (100 um near the margin of the pileus). Smell sour or of fresh mushrooms

(RSS 1799), or none (RSS 844).

On fallen branches and trunks of various trees, and on dead palm-trunks, in

the forest. Malaya, Java, Borneo, New Guinea, Solomon Islands, New Caledonia,

Australia, New Zealand, Chile.

Figure 1. Hohenbuehelia bursaeformis. Spores, basidia, cheilocystidia and

pleurocystidia, x 1000. Surface of pileus, x 500.

Spores 5-6.7 x 4.5-6 um, 6-6.7 x 5-5.7 um (RSS 844, 1799), 6-8 x 5-5.6 um

(RSNB 5168, 8039), white, broadly ellipsoid to subglobose, closely and finely

echinulate with spines -1 um long, or with shorter spines -0.7 um (RSNB 5168,

8039), -0.5 um (Tembeling), or -03 um (RSS 1799), or merely asperulate (RSS

844), inamyloid, the spines often contracting in Melzer’s iodine and dissolving in

dilute potash. Basidia 17-22 x 7-8 um; sterigmata 4, 5 um long; no acerose

basidioles; subhymenium narrow, interwoven subgelatinous. Cheilocystidia 30-

60 x 6-20 um, subcylindric, clavate or ventricose, obtuse, not capitate, thin-

walled, smooth, as a sterille gill-edge but becoming disrupted or collapsing.

Pleurocystidia -65 x 9-20 um, clavate to ventricose with a short obtuse appendage

-16 x 3-6 um, thin-walled or the apex of the appendage with slightly thickened

wall -0.5 lum, smooth, drying with reddish brown vitreous-vacuolate contents

(dissolving and becoming colourless in KOH and Melzer’s iodine, or persistently

reddish brown in Melzer’s iodine). Hyphae monomitic, with minute clamps

(easily overlooked), inflating, with cylindric cells 16-105 (-140) x 3-23 um, not

or scarely constricted at the septa, branching rather loosely at a wide angle,

longitudinal and intewoven but more compact and 3-6 um wide and strictly

longitudinal in a thin layer 40-50 um thick over the gills and below the gelatinous

layer, the walls firm or thickened -0.5 um; in the gill-trama descending, short-

celled, inflated; in the gelatinous layer of the pileus longitudinal and interwoven.

Surface of pileus developing a villous layer -400 um thick and composed of

much entangled hyphae 3-8 Um wide, separate, sparingly branched, with smooth

wall -0.5 um thick, in the middle part of the pileus with clavate to subventricose

thin-walled cells 30-80 x 7-30 Um (? as a palisade in the young pileus, collapsing

in the villous part of old pile).

Collections.- Malaya, Pahang, Tembeling, Corner s.n. 4 Nov. 1930.- Borneo, Mt.

Kinabalu, 1100-1700m alt., on rotten wood, RSNB 607, 5167, 5167A 5167C,

§039.- Solomon Islands, Guadalcanal, Tsuva, on a rotten trunk of Areca, RSS

1799; San Cristobal, Warahito River, RSS 844, on a dead branch.

This is my description from Malesian material. It agrees well with that of

other authors. Horak gives the pleurocystidia as merely 20-35 x 7-16 Um with

the wall -1 um thick in the distal part, and covered with resinous incrustation or,

rather rarely, with crystals. Conchomyces is discussed on p. 2.

Hohenbuehelia concentrica sp. nov.

Pileus -6cm in radio, -8cm latus, sessilis flabelliformis convexo-planus, rugis concentris zonatus,

aurantiocervinus vel pallide cervino-ochraceus, sicco albivillosulus. Lamellae confertae subtenaces, 11-

19 primariae 2.5-3.5mm latae, ordinibus 6-8, pallide ochracei-alutaceae. Caro 3-5mm crassa, strato

gelatinoso 1-2mm crasso, strato villoso 0.3mm crasso, albida vel alutacea. Odor farinaceus fortis.

Sporae 8-8.5 x 6-6.7 Lim, laeves ovoidease, intus granuloso-opalescentes. Cheilocystidia vix evoluta,

lamellae acie hyphis -70 x 1.5-3 Lum haud vel laxe ramosis instructa. Pleurocystidia 30-80 x 7-15 Um,

ventricosa, tunicis -5 Um crassis, haud incrustata, solum apicibus acutis breviter projicientibus, sparsa

sed lamellae aciem versus numerosa. Subhymenium 15-20 Lim crassum, cellulis3-7 Lim latae. Superficies

pilei sine cystidiis. Ad truncum putridum Lophopetalum (Celastraceae) in silva. Singapore, Selitar,

Corner s.n. 4 Feb. 1944; typus, herb. Corner.

Pileus -6cm in radius, -8cm wide, sessile, convexo-plane, with narrow raised

concentric zones, wholly light orange fawn to pale fawn ochraceous, drying

white villous with the raised zones as narrow concentric ridges closely developed

towards the incurved margin. Stem practically none, as a finely villous pulvinus.

Gills crowded, rather tough, thin, not gelatinous, 11-19 primaries 2.5-3.5mm

wide, 6-8 ranks, entirely pale ochraceous buff. Flesh with the firm whitish to

pallid buff layer 2-3.5mm thick at the base of the pileus, the brownish gelatinous

upper layer 1-2mm thick, the villous layer 0.3mm thick. Smell strong, of meal.

On rotten wood of Lophopetalum (Celastraceae) in the forest. Singapore,

Seletar.

Spores 8-8.5 x 6-6.7 um, white, smooth, ovoid, wholly finely opalescent

granular. Basidia 35-43 x 8-10 um, clavate, finely opalescent granular, 3-3.5 um

wide at the base; sterigmata (2-3-)4, 6-6.5 x 2.5-3 um. Subhymenium 15-20 um

thick, composed of short-celled, rather regularly divergent hyphae 3-6 um wide,

at right angles to the descending hyphae of the gill-trama, distinctly corticate.

Cheilocystidia not formed, the gill-edge with many narrow emergent hyphae -70

x 1.5-3 um, not or sparsely branched, septate or not. Pleurocystidia 30-80 x 7-15

um, ventricose with a short acute process, wall -5 um thick, smooth, rather sparse

but more abundant towards the gill-edge, the tip shortly projecting. Hyphae

monomitic, clamped; in the firm layer of the flesh 3-7 um wide with rather firm

walls, longitudinal and interwoven, compact, 1.5-4.5 um wide and ascending in

the gelatinous layer; in the gill-trama 2-10 um wide, descending with thin or

scarcely thickened walls, not gelatinous. Surface of pileus with an often ill-

defined layer c. 20 um thick, composed of longitudinal hyphae 2-6 um wide,

overlain by a layer 20-40 um thick, composed of laxly interwoven hyphae 1.5-3

um wide, with excrescent and slightly thick-walled hyphae forming discrete or

anastomosing subagglutinated fascicles -350 um high at the base of the pileus; no

pileocystidia.

This species is very distinct in the orange-tinted and zoned pileus, the large

ovoid spores, the absence of cheilocystidia and the distinctly corticate

subhymenium.

Hohenbuehelia griseipendens sp. nov. Figure 2

Receptacula -20mm lata, sessilia cyphelliformia pendentia, pallide grisea. Lamellae confertae, 9-11

primariae -1.5mm latae, ordinibus 4-6. Caro 1-2mm crassa, ex integra gelatinosa firma. Odor nullus.

Sporae 4-5 x 3.5-3.7 Lum, laeves, late ellopsoideae. Cheilocystidia 15-30 x 6-10 Um, clavata sed nonnulla

ut pleurocystidia intermedia. Pleurocystidia 25-70 x 7-14 Lim, ventricosa breviter appendiculata, tunicis

-2.5 Um crassis, vel clavata tunicis haud vel vix incrassatis, etiam hymenialia ut cheilocystidia. Hyphae

in carne 1.5-3.5 Lm latae, gelatinosae. Superficies pilei sine cystidiis et cellulis clavatis, aut cum cellulis

clavatis paucis et saepe sublobatis. Ad lignum putridum in silva montana. Borneo, Mt Kinabalu, fl.

Liwagu, 1300m alt., RSNB 2562, 29 Aug. 1961; typus, herb. Corner.

var. nonsatisfacta var. nov.

Differt sporis angustioribus 4-5 x 2-2.3 Um, pleurocystidiis tunicis vix incrassatis -0.5 Lim, pilei

superficie acystidiata. Ad lignum putridum in silva. Insulae Solomonenses, Guadalcanal, RSS 511;

typus, herb. Corner

Fruit-bodies -20mm wide, sessile, cyphelliform pendent, pale grey, the

upperside drying finely white villous. Gills diverging from the central point,

crowded, thin, narrow, 9-11 primaries 1-1.5mm wide, 4-6 ranks, grey edge

entire. Flesh 1-2mm thick at the base, wholly gelatinous-firm. Smell none.

“4 bel

te rim es mis mee oe

MSR ape Pe

Figure 2. Hohenbuehelia griseipendens. Developing fruit-bodies in section, x 10.

Mature fruit-body in section, x 2. Cheilocystidia, pleurocystidia and

surface of pileus, x 500.

On rotten wood in montane forest. Borneo, Mt Kinabalu, Liwagu river

1300m alt.

Spores 4-5 x 3.5-3.7 mm, white, smooth, broadly ellipsoid, thin-walled,

inamyloid, (1-2 guttulate in material in alcohol-formalin). Basidia 5 um wide, 4-

spored. Cheilocystidia 15-30 x 6-10 um, mostly clavate, thin-walled, as a sterile

gill-edge, but with transitions to the pleurocstidia. Pleurocystidia 25-70 x 7-14

uum, ventricose and rather shortly appendaged to clavate, walls 0.5-2.5 tum thick

in the ventricose cells, not encrusted or slightly in the distal part, abundant, with

many smaller cystidia at the gill-surface like the cheilocystidia. Hyphae monomitic,

clamped, 1.5-3.5 um wide, gelatinous; in the gill-trama 3-5 um wide with firmly

gelatinous walls, descending. Surface of pileus with a layer 25-35 um thick,

composed of radiating and interwoven hyphae 2-5 um wide with firm (not

gelatinous) walls, some ending in a short subclavate cell -7 um wide and often

with shortly lobed apex, but not as a hymenioderm and without thick-walled

cystida; this layer giving rise with excrescent hyphae 2-4 um wide, often

es,

fasciculate, with slightly thickened walls, not or sparsely septate with clamps as

the villous layer -300 um thick.

var. nonsatisfacta

Pileus -9 mm wide, sessile, dorsifixed, unilaterally expanded and flabelliform

resupinate, striate , fuliginous grey, drying white villous over the centre. Gills

radiating from the excentric attachment, crowded, 8-10 primaries 1mm wide, 4-5

ranks, fuliginous grey. Flesh 1mm thick at the base, wholly gelatinous.

On rotten wood in the forest. Solomon Islands, Guadalcanal, Gallego, Monitor

Creek, 2 July 1965, RSS 5/1.

Spores 4-5 x 2.2-3 um, white, smooth, ellipsoid, inamyloid. Basidia 13-17 x

4-4.5 um, 4-spored; no subacerose basidioles; subhymenium 15-25 um thick,

composed of 1.5-3 um intewoven hyphae. Cheilocystidia 12-25 x 4-8 um,

clavate to ventricose, occasionally with a short appendage, not capitate, thin-

walled, smooth, as a sterile gill-edge. Pleurocystidia 30-55 x 7-9.5 um, base 3-4

um wide, fusiform, acute, wall slightly thickened -0.5 um, thinly encrusted distally,

not dextrinoid, abundant, varying more or less immersed to subhymenial and

projecting -18 um. Hyphae monomitic, clamped; in the flesh 1.5-3 um wide with

gelatinous walls in the gill-trama 2-4 um wide a discontinous layer, 1-2 hyphae

thick, of 2-4 um hyphae with firm walls, developing the thinly villous layer at the

base of the pileus, without cystidia or clavate cells.

This is close to H. singaporensis but it lacks the hymenioderm and the

pleurocystidia are not so thick-walled or so wide.

Hohenbuehelia horrida (Boedijn) comb. nov.

Acanthocystis horrida Boedijn, Bull. Jard. bot. Buitenz. ser. 3, 16 (1940) 400, f. 9.

Pileus -7 cm in radius, -6cm wide, shortly pleuropodal to laterally sessile,

spathulate to flabelliform and reniform, faintly striate, white then pale yellowish

drab or greyish bistre, drying white to greyish and radially sulcate and closely

white villous towards the base; margin becoming lobed and undulate in large

specimens. Stem -18 x 3-12mm, lateral varying rudimentary or subdiscoid,

occasionally absent, white villous. Gills decurrent, distant, rather thick, wide

interstices smooth, 11-25 primaries 4-6mm wide, 4-7 ranks, pale cream buff then

dingy pallid ochraceous, finally pinkish rufous (pale grey to greyish brown,

Boedijn). Flesh 1-2.5mm thick at the base of the pileus, 0.5-1mm halfway to the

margin, firm, subcoriaceous, with a gelatinous layer 250-600 um thick, watery

concolorous. Smell farinaceous, rather slight, or none (RSS 1044).

On fallen or standing dead trunks and rotten wood in lowland primary and

secondary forest. Malesia.

Spores 5-6 x 3-4 um, white in the mass, smooth, aguttate, inamyloid. Basidia

18-25 x 4-5 um; sterigmata 4, 4-5 um long; subhymenium 20-30 um thick, with

compactly interwoven hyphae 2.5-8 um wide, not gelatinous. Cheilocystidia

unformed, as clusters of small sterile basidia, some with a slender process ending

in an excreted globule -7 um wide, scattered among emergent hyphae 1.5-3 um

wide, often fasciculate in flattened clusters, the gill-edge sterile. Pleurocystidia -

140 x 12.5 um with pale yellowish walls -5.5 um thick, ventricose-lanceolate,

acute, lumen becoming linear, not or thinly encrusted, the larger subhymenial or

tramal and embedded or projecting -55 tum, the smaller 35-55 x 6-8 Um with thin

to slightly thickened wall hymenial, smooth or little encrusted, with all

intermediates, extremely abundant. Hyphae monomitic, clamped; in the firm

tissue of the flesh 3-11 tum wide, with thin or slightly thickened walls, longitudinal;

in the gelatinous layer 1.5-3 (-5) um, ascending; in the gill-trama 4-22 um wide,

often unevenly inflated, with firm to slightly thickened walls, loosely interwoven

and descending. Surface of pileus with a thin compact layer of longitudinal

hyphae 3-8(-15) um wide above the gelatinous layer and giving rise to a vague

outer layer c. 20 um thick of interwoven hyphae 1.5-3 um wide, variously

excrescent, septate, clamped, becoming aggregated or agglutinated into

fascicles -300 um long as the villous layer; pileocystidia absent.

Collections.- Malaya, Pahang, Tembeling, Corner s.n. Nov. 1930, common;

Johore, Sedili River, Corner s.n. 22 Oct. 1939.- Krakatau, leg. Boedijn.-

Solomon Islands, San Cristobal, Warahito river, 28 July 1965, RSS 855;

Kolombangara, 24 Aug. 1965, RSS 1044.

This is my description of this striking and widespread species. It agrees with

that of Boedijn except in the colour of the gills. The structure of the gill-edge

without distinct cheilocystidia suggests nematode-catching. H. incarnata is allied

and, perhaps, also the East African H. aurantiocystis Peglear (1977) though it has

crowded narrow gills and pleurocystidia often with reddish incrustation at the apex.

Hohenbuehelia incarnata sp. nov.

Receptacula pallide incarnata-alutacea sessilia vel breviter pleuropadalia. Pileus -6 cm latus,

spathulatus dein flabelliformis, substriatus, sicco tenuiter albidi-villosus vel subspiculosus. Lamellae

subdistantes, 7-13 primariae -5 mm latae, ordinibus 4(-5), raro connexae. Caro -2 mm crassa, strato

gelatinoso 100-150 Lum crasso. Odor subaromaticus. Sporae 5-7 x 4.5-6 Um, laeves, subglobosae.

Cheilocystidia 16-33 x 2-5 Lim, subcylindrica vel submoniliformia. Pleurocystidia 45-90 x 7-12 Lum,

lanceolata attenuata, apicibus acutis vel aciculiforminus, tunicis, -3 Lim crassis, apicem versus

subincrustata, copiosa conferta. Hyphae in carne firma 2-8 Lm latae. Superficies pilei sine cystidiis. Ad

lignum putridissimum, mycelio albo lignum infestanti. Insulae Solomonenses, Guadalcanal, 3 Jul. 1965,

RSS 529; typus, herb, Corner.

Fruit-bodies wholly pale pinkish buff, sessile to shortly pleuropodal. Pileus -6

cm wide, spathulate to flabelliform, ascending then horizontal, flexuous, substriate,

thinly white villous or spiculose on drying; margin incurved at first. Stem very —

short or none, white villous. Gills deeply decurrent, arising almost from the base

of the fruit-body, subdistant, with smooth interstices, 7-13 primaries -5 mm wide,

4(-5) ranks, occasionally joined. Flesh -2 mm thick at the base of the fruit-body,

rather firm, with a very thin gelatinous layer below the surface. Smell rather of

fenugreek.

On very rotten wood, the spongy white mycelium incorporating the wood.

Solomon Islands, Guadalcanal, Mt Gallego, frequent, RSS 529 and 529a.

Spores 5-7 x 4.5-6 um, white, smooth, subglobose. Cheilocystidia 16-33 x 2-5

um, subcylindric to submoniliform with slightly subcapitate apex. Pleurocystidia

45-90 x 7-12 um, lanceolate with long tapered acute to almost acicular apex, wall

-3 um thick, thinly encrusted near the apex or smooth, very abundant and closely

set. Hyphae monomitic, clamped; in the firm layer comprising most of the flesh

2-8 um, thin-walled, longitudinal, compact; in the gelatinous layer 100-150 Um

thick 1.5-3 um wide; in the gill-trama as in the firm tissue of the pileus. Surface

of pileus with more or less fasciculate cylindric hyphae 2-4 um wide, septate,

some with subclavate tips 3-5 tum wide, the fascicles -250 um long, arising from a

layer 20-70 um thick of similar, more or less radiating, appressed hyphae; no

pileocystidia.

This is close to H. horrida but with different colour and smell, thinner gelatinous

layer and distinct cheilocystidia.

Hohenbuehelia lanceifera sp. nov. Figure 3

Pileus -20 mm radio, sessilis lateralis spathulato-flabelliformis, rhizomorphis albis gracilibus affixus

vel ad basim discoideum subvillosum 3-7 latum, albus dein pallide ochraceus, opacus, basim versus

subvillosus. Lamellae confertae, 6-15 primariae -2.5 mm latae, ordinibus 4-5, albae dein pallide

subochraceae, aetate brunneolae vel subincarnatae. Caro -1.5 mm crassa, fere ex integra aquosi-gelatinosa,

subochracea. Inodora. Sporae 4-5.5 x 3.7-4.5 Lim, laeves subglobosae vel late ellipsoideae. Cheilocystidia

ut pleurocystidia parva vel ut basidia sterilia. Pleurocystidia 30-90 vel -140 x 7-20 Um ad lamellarum

basi, lamellae sciem versus 20-30 x 5-7 Lim, ventricoso-lanceolata, tunicis incrassatis, apicem versus

incrustata, copiosa. Caro strato firmo 150-200 Lim crasso, ex hyphis 3-7 Lim latis instructo. Superficies

pilei sine cystidiis. Ad ramos dejectos truncosque emortuos in silva montana. Borneo, Mt. Kinabalu,

1700m alt. RSNB 2891; typus, herb. Corner

Pileus -20 mm radius, lateral, sessile, spathulate to flabelliform, attached with

white mycelial fibrils or with a discoid subvillous base 3-7 mm wide, thinly

villous towards the base, minutely cottony over the limb, white then dingy watery

ochraceous; margin opaque, minutely fimbriate. Gills radiating from the base,

crowded, thin, 6-15 primaries 2-2.5 mm wide, 4-5 ranks, white then pallid dingy

ochraceous, finally brownish or pinkish. Flesh -1.5 mm thick at the base of the

pileus, more or less wholly watery gelatinous, subochraceous. Smell none.

On fallen rotten branches and dead standing trunks in montane forest. Borneo,

Mt Kinabalu 1700m alt.

Spores 4-5.5 x 3.7-4.5 um, white, smooth, subglobose to broadly ellipsoid.

Basidia 5-6 uum wide; aterigmata 4(-5), 3 um long. Cheilocystidia small to fairly

large, somewhat thick-walled as if immature pleurocystidia, with scattered sterile

basidia, thinly enveloped in mucilage, as a sterile gill-edge. Pleurocystidia -140 x

Figure 3. Hohebuehelia lanceifera. Surface of pileus and (below) firm layer of flesh,

and pleurocystidia, x 500.

20 um at the base of the gills, 30-90 x 7-14 um over the surface, 20-30 x 5-7 um

near the gill-edge, ventricose-lanceolate with long attenuate apex, thick-walled,

encrusted distally with crystals and towards the apex with granules, base more or

less tramal, abundant. Hyphae monomitic, clamped; in a firm layer 150-200 Um

thick over the gills 3-7 tum wide, compact, longitudinal; in the thick gelatinous

layer 1.5-3 um wide, longitudinal and ascending; in the gill-trama 3-10 tum wide,

descending, short-celled, not gelatinous; oleiferous hyphae 3-7(-10) um wide,

sparse in the tissue below the surface of the pileus. Surface of pileus with a layer

20-50 um thick composed of longitudinal, compact, rather short-celled hyphae 3-

7 um wide with firm walls, giving off fascicles 200 x 20-60 um of similar hyphae

forming the villous layer.

Collections.- Borneo, Mt Kinabalu, Tenompok, 8 Sept. 1961, RSNB 2891;

Mesilau, 22 April 1964, RSNB 8408.

This is near H. concentrica which has larger spores and fruit-bodies.

Hohenbuehelia malesiana sp. nov. Plate 1, Figure 4

Pileus -3 cm radio, -4.5 cm latus, lateralis, sessilis vel substipitatus, spathulatus dein flabelliformis,

ultimo lobatus, substriatus, sordide alutaceus vel albus, sicco albivillosus. Stipes vix evolutus. Lamellae

confertae vel subdistantes, nonnullae furcatae, ceracei-gelatinosae, 5-11 primariae 0.5-2 mm latae,

ordinibus 3-7, albidae vel concolores. Caro 0.3-1.5 mm crassa, fere ex integra gelatinosa. Inodora.

Sporae 7-9 x 3.5-4 Um, laeves subcylindricae, aguttatae vel 1-2 guttulatae (vivae). Cheilocystidia 15-25 x

5-10 Lum, clavata vel ventricosa, saepe capitulo 2-3.5 Lim lato substipitato praedita. Pleurocystidia 30-100

x 7-25 um, ventricoso-fusiformia, tunicis flavidulis crassis, apices versus tenuiter incrustata. Hyphae in

strato firmo tenuissimo 3-8(-10) Lim latae. Superficies pilei sine cystidiis. Ad lignum emortuum ramulosque

in silva. Peninsula Malayana, Borneo, Insulae Solomonenses. Typus Singapore Corner s.n. 17 March

1940: herb. Corner.

Pileus -3 cm in radius, -4.5 cm wide, sessile, lateral, spathulate then flabelliform,

finally lobate, smooth, translucent, faintly striate, watery greyish to pallid dingy

bistre, occasionally white, drying more or less wholly white villous and faintly

sulcate-striate. Stem very slight, generally none. Gills decurrent, subdistant to

Figure 4. Hohenbuehelia malesiana. Fruit-body in section, x 10. Spores, x 1000.

Cheilocystidia, pleurocystidia and surface of pileus, x 500.

crowded, sometimes dichotomous, 5-11 primaries 0.5-2 mm wide, 3-7 ranks,

waxy gelatinous, watery white to pale concolorous. Flesh 0.3-1.5 mm thick at the

base of the pileus, wholly and rather firmly gelatinous except a very thin layer

over the gills. Smell none.

On sticks and dead wood in the forest. Malay Peninsula to the Solomon

Islands.

Spores 7-9-x 3.5-4 um, white, smooth, subcylindric, often slightly curved,

aguttate or with 1-2 minute guttulae at the ends of the spore (living), inamyloid.

Basidia 20-30 x 6-8 um; sterigmata 4, 2.5 um long. Cheilocystidia 15-25 x 5-10

uum, clavate or ventricose with a short apical stalk -2 um long and a globose head

2-3.5 um wide, thin-walled, hyaline, as a sterile gill-edge. Pleurocystidia 30-100

x 7-25 tum, ventricose fusiform, walls thick and yellowish, roughened and minutely

encrusted over the distal third or smooth, acute to subacute. Hyphae monomitic,

clamped, not or scarcely inflated; firm layer of the flesh thinner than the gelatinous

layer, composed of hyphae 3-8(-10) um wide, 2-3 um in the gelatinous layer; gill-

trama as the firm layer of the flesh. Surface of pileus with a thin interrupted layer

of longitudinal hyphae 1.5-6(-10) um wide, developing excrescent, short-celled,

cylindric hyphae -300 um long at the base of the pileus, -50 um near the margin,

3-6 um wide, with firm walls, mostly unbranched, not or rarely fasciculate; no

thick-walled cystidia.

Collections.- Malaya, Perlis, Bukit Chupeng, Corner s.n. 29 Nov. 1929; Johore,

Mawai, Corner s.n. 21 Sept. 1934.- Singapore, Botanic Garden, Corner s.n.

March 1935; Bukit Timah, Corner s.n. 17 Sept. 1939, 17 March 1940.- Borneo,

Mt Kinabalu, 1700m alt., RSNB 5625 3 March 1964.- Solomon Islands,

Kolombangara, RSS 1162, 30 Aug. 1965.

This may be the commonest species in South East Asia. In 1934 and 1935 I

managed to raise some fruit-bodies from the log on which I had found them in

Johore. Many primordia started but merely three developed satisfactorily.

Measurements were begun when the primordial pileus was 2 mm in radius. In 6

days the largest reached its full size of 22 mm in radius, 25 mm wide, and then

lasted for 9 days before collapsing. During the first four days of observation, the

pileus increased in radius by 2, 3, 5 and 6 mm when it slowly declined in the next

2 days. The smallest pileus reached 8 mm in radius, 9 mm wide, in 3 days from

the same initial state and lasted for a further 7 days. The third fruit-body of

intermediate size, reaching 15 mm in radius, 20 mm wide, developed in much the

Same manner as the first and reached full size in 7 days; it collapsed 8 days later.

I reckoned the primordium of this third fruit-body was 48 hours old because it

developed near the other two but 2 days later. Thus the total life of the largest

fruit-body was about 16 days, of which the last 8-9 days were at full size. The

fruit-bodies eventually collapsed because they became sodden with water and

failed under their own weight.

Hohenbuehelia mellea sp. nov. Plate 1

Pileus -15 mm latus, sessilis spathulatus dein semiorbicularis, melleiflavus vel ochraceibrunneus,

squamulis flavidulis ornatus, striatus. Lamellae subdistantes, 6-8 primariae 1.5-2 mm latae, ordinibus 3-

4, ceraceimolles, pallide ochraceae dein pallide cinnamomei-ochraceae. Caro 1-2 mm crassa, fere ex

integra gelatinosa, concolor. Inodora. Sporae 5-6.5 x 2.5-3 Lim, laeves ellipsoideae, paucis visis.

Cheilocystidia 30-50 x 8-18 Lim, ventricosa, processu simplici vel 1-ramoso etiam subcapitato, tenuiter

tunicata. Pleurocystidia 50-160 x 11-20 Lim, ventricoso-fusiformia, longe attenuate acuta, tunicis -5 [um

crassis flavidis, incrustata. Stratum carnis firmum 100-150 Lim crassum, ex hyphis 4-15 Lim latis

instructum; in lamellae trama 5-18 Lim latae. Superficies pilei strato fere pseudoparenchymatico 15-20

Lum crasso obtecta, haud villosa, hinc inde cystidiis ut cheilocystidiis. Ad lignum emortuum in silva.

Singapore, Bukit Timah, Corner s.n. 24 Dec. 1940; typus, herb. Corner.

Pileus -15 mm wide, sessile, broadly spathulate to semicircular, slightly convex

becoming plane, honey yellow or ochraceous brownish, striate, with minute

yellowish fleck-like scales, drying radially rugulose. Gills radiating from the

base, subdistant, 6-8 primaries 1.5-2 mm wide, 3-4 ranks, waxy-soft, pale

ochraceous then pale cinnamon ochraceous. Flesh 1-2 mm thick at the base,

wholly gelatinous, waxy in the gills, concolorous. Smell none.

On dead wood in the forest. Singapore, Bukit Timah.

Spores 5-6.5 x 2.5-3 um, white, smooth ellipsoid, (few seen). Cheilocystidia

30-50 x 8-18 um, more or less ventricose with a simple or once branched,

cylindric or subcapitate appendage 2-3.5 um wide, some with an excretory blob,

rather irregular, thin-walled, colourless, as a narrow sterile gill-edge. Pleurocystidia

50-160 x 12-20 um, ventricose fusiform, long tapered, subacute to acute, the pale

yellow walls 2-5 um thick, heavily granular encrusted over the greater part,

abundant. Hyphae monomitic, clamped; in the very thin firm layer of the flesh,

—Va——<=

Plate 1. 1, Hohenbuehelia mellea. 2, H. malesiana.

3, H. suppapillosa.

v5 4

100-150 um thick, longitudinal 4-15 um wide; in the gelatinous tissue composing

most of the flesh, longitudinal and interwoven 3-5 Um wide; in the gill-trama 5-

18 um wide with firm yellowish walls. Surface of pileus with an almost

pseudoparenchymatous layer 15-20 um thick, composed of longitudinal hyphae

with cells 20-45 x 4-16 um, with scattered narrow hyphal ends shortly projecting

and scattered cystidia like the cheilocystidia; no villous layer, no thick-walled

cystidia, but at the base of the pileus numerous, shortly excrescent, slightly thick-

walled hyphae -200 x 2-4.5 um.

Superficially, this suggests Crepidotus. A sterile collection from Sarawak

seems intermediate between this and H. lanceifera. It had the pale colour of H.

lanceifera and the same structure to the pileus but the more distant gills and

cheilocystidia of H. mellea. Thus:- Pileus -30 mm wide, drab white tinged pale

ochraceous, thinly villous at the base. Gills distant, 7-9 primaries 2-3 mm wide,

4-5 ranks, pale ochraceous drab. Pleurocystidia -140 x 8-14 lum at the base of the

gills, fusiform, long attenuate, thick-walled, heavily encrusted with granules

between the swollen central part and the attenuate apex. Sarawak, Bako National

Park, Corner s.n. 31 Jan. 1959.

Hohenbuehelia minutissima sp. nov.

Pileus -3 mm latus subsessilis lateralis spathulatus griseus. Stipes -0.5 mm longus vel nullus. Lamellae

confertae, 7-9 primariae -0.5 mm latae, ordinibus 3-4, albae. Caro tenuissima, tenaciter gelantinosa.

Sporae 5-7 x 2.7-3.5 Lum, laeves subcylindricae. Cheilocystidia ut basidia sterilia clavata subventricosa,

saepe capitulata, vel ut pleurocystidia. Pleurocystidia 40-80 x 9-20 Lim, ventricosa subacuta, tunicis

flavidulis, apices versus tenuiter incrustata. Hyphae in strato firmo angusto et in lamellis 2-5 [im latae,

tunicis 1-2 Lim crassis, lumine lineari, subgelatinosae. Superficies pilei sine cystidia. Ad rami delapsi

corticem in silva. Nova Guinea, prope Lae, Corner s.n. 7 Sept. 1960; typus, herb. Corner.

Fruit-bodies very small, sessile or with a slight lateral stem. Pileus -3 mm in

radius and wide, spathulate, smooth, grey. Stem -0.5 mm long, white, pruinose,

or none. Gills crowded, narrow, acute, 7-9 primaries -0.5 mm wide, 3-4 ranks,

white. Flesh very thin, toughly gelatinous.

On the bark of a fallen branch in forest. New Guinea, near Lae.

Spores 5-7 x 2.7-3.5 um, white, smooth, subcylindric, thin-walled, aguttate.

Basidia 5.5-6 um wide, 4-spored. Cheilocystidia either as sterile basidia clavate

to subventricose and then often shortly apiculate and capitate, or as thick-walled,

thinly encrusted, obtuse to subacute cystidia, as a sterile gill-edge. Pleurocystidia

40-80 x 9-20 Lum, ventricose, subacute, with thick yellowish walls, thinly encrusted

distally with granular matter soluble in KOH. Hyphae monomitic, clamped, not

inflated; in the narrow firm layer of the flesh over the gills and in the gills 2-5 um

wide with walls 1-2 um thick and linear lumen, possibly subgelatinous; in the

gelatinous tissue, forming most of the flesh, 1.5-4 um wide, thin-walled. Surface

of pileus with a layer, 1-2 hyphae thick, of longitudinal hyphae 1.5-3 um wide,

thin-walled; no villous layer, no cystidia.

This is distinguished by the very small size of the fruit-body and the thick-

walled hyphae of the firm tissue.

Hohenbuehelia pachyhyphata sp. nov. Figure 5

Receptacula ex integro alba dein alutacea vel pallide fuscivinacea. Pileus 2-3 cm latus, breviter

pleuropodalis, flabelliformis, ultimo revolutus, opacus, sicco villosulus. Stipes -3 x 4 mm, sursum dilatatus.

Lamellae confertae, saepe furcatae vel prope stipitem reticulatae, 11-26 primariae 1-2 mm latae, ordinibus

4-7. Caro 2-4 mm crassa, strato gelatinoso 1 mm crasso. Odor farinaceus, fortis. Sporae 6-7.5 x 3-3.5

Lum, laeves, subcylindricae. Cheilocystidia -20 x 7 Lum, clavata vel ventricosa, saepe breviter capitulata,

etiam ut pleurocystidia. Pleurocystidia 30-80 x 8-20 Um, ventricosa, breviter acuta, tunicis crassis,

apicem versus tenuiter incrustata, copiosa. Hyphae in carnis strato firmo et in lamellis 3-10 Lim latae,

tunicis valde incrassatis, lumine quasi occluso, et ampullis -20 Um latis (-50 Lim in stipite). Superficies

pilei sine cystidiis crasse tunicatis. Ad lignum emortuum in silva. Borneo, Queensland. Typus, Borneo,

Corner s.n. 22 Jun. 1961; herb. Corner

PA TT ES

rit

' ’

Iny

yy .

Figure 5. Hohenbuehelia pachyhyphata. Cheilocystidia, pleurocystidia, surface of pileus

and hyphae of the firm layer of the flesh, x 500.

var. minor var. nov.

Differt pileo -13 mm lato, albo dein griseo-umbrino, strato gelatinoso 0.5 mm crasso, sporis 4.5-5.5(-

6) x 3.3-3.7 Lum ellipsoideis, hyphis haud ampulliformibus. Ad truncum delapsum in silva. Borneo, Mt.

Kinabalu, Mesilau 1700m alt., 26 Jan. 1964, RSNB 5141; herb. Corner.

Fruit-bodies wholly white, then pale yellowish buff to pale livid bistre or pale

fuscous vinaceous (RSNB 8691). Pileus 2-3 cm wide, lateral, convex then plane

to revolute, flabelliform, opaque, drying minutely villous or cottony flocculose;

margin incurved. Stem -3 x 4 mm, lateral, short, stout, widened upwards, pruinoso-

villosulous. Gills decurrent, crowded, thin, often forked or reticulate near the

stem, 11-26 primaries 1-2 mm wide, 4-7 ranks, edge entire. Flesh 2-4 mm thick at

the base of the pileus, the gelatinous layer 1 mm thick. Smell farinaceous, strong

on cutting.

On dead wood in the forest. Borneo, Queensland.

Spores 6-7.5 x 3-3.5 um, white, smooth, subcylindric, inamyloid. Basidia 18-

25 x 5 um, 4-spored. Cheilocystidia -20 x 7 um, clavate to subventricose, often

with a short capitate process, thin-walled, also mixed with some thick-walled

cystidia, as a sterile gill-edge. Pleurocystidia 30-80 x 8-20 um, ventricose with

rather short acute apex, thick-walled, arising deeply from the trama, immersed or

shortly projecting, the free part with thin incrustation (soluble in KOH), very

abundant. Hyphae monomitic, clamped; in the firm layer of the flesh 3-10 um

wide, becoming very thick-walled and nearly solid, but with ampulliform swellings

-20 um wide, -50 um in the stem, longitudinal, compact; in the gelatinous layer

1.5-3.5 um wide, walls thin or slightly thickened, ascending, also with a few

thick-walled hyphae; in the gill-trama as in the firm layer of the flesh, descending.

Surface of pileus with a layer 15-40 um thick of rather closely interwoven hyphae

2-4 uum wide, in places -10 um, with firm thin walls, producing fascicles -300 um

long of hyphae 2.5-5 tm wide with firm, often slightly thick-walled, often

subagglutinated, 0-2 septate and clamped, with scattered ill-formed and thin-

walled cystidia 4-8 um wide like the cheilocystidia, but no thick-walled cystidia.

Collections.- Borneo, Mt Kinabalu, 1100-1700 alt., east ridge, Corner s.n. 22

June 1961; Mesilau, 6 May 1964, RSNB 8691.- Queensland, Mary Cairncross

Reserve, Maleng, Corner s.n. 20 June 1964.

var. minor

Pileus -13 mm wide, pleuropodal, flabelliform, white then clouded greyish

umber towards the minutely villous base. Stem very short. Gills very crowded,

narrow, 13-17 primaries 1-1.5 mm wide, 5-6 ranks, white. Flesh 1-1.5 mm thick

at the base, the gelatinous layer 0.5 mm thick and generally thinner than the firm

layer. Smell ?

On a fallen trunk in the forest. Borneo, Mt Kinabalu, Mesilau 1700m alt., 26

Jan. 1964.

Spores 4.5-5.5(-6) x 3.3-3.7 um, white, smooth, ellipsoid, aguttate.

Cheilocystidia 12-23 x 5-9 um, clavate to ventricose with a short capitate process.

Pleurocystidia 30-90 x 7-20 um, ventricose-conical, acute, sometimes curved or

waisted, with thick yellowish walls thinly encrusted distally. Hyphae of the firm

layer of the flesh 2-8 lum wide, cylindric, clamped, with walls 1-3 um thick,

without ampulliform swellings. Surface of pileus as in var. pachyhyphata but the

excrescent hyphae often fasciculate.

The pale colour, farinaceous smell and very thick-walled hyphae of the firm

tissue distinguish this species.

Hohenbuehelia pahangensis sp. nov. Figure 6

Pileus -4 cm radio, -7 cm latus, sessilis vel breviter pleuropodalis, ascendens, spathulatus flabelliformis

opacus villosulus, fuscus dein pallide isabellinus. Stipes -5 x 2.5-4 mm, villosulus albidus. Lamellae alte

decurrentes, confertissimae, 12-16 primariae 2 mm latae, ordinibus 6-7, albae, acie serrulata. Caro 2-2.5

mm crassa, strato gelatinoso c. 300 im crasso. Odor farinaceus fortis. Sporae 3.5-4.3 x 3-3.5 Lim, laeves,

late ellipsoideae. Cheilocystidia 35-60 x 5-7 Lim, clavata vel obtuse subventricosa. Pleurocystidia 55-120 x

10.5-22 Um, ventricosa, breviter acuminata, tunicis flavidulis 2-7 Um crassis, laevia vel apicem versus

tenuiter incrustata. Hyphae in carnis strato firmo lamellisque 4-23 Lim latae. Superficies pilei cystidiis

40-90 x 6-8 Lim ut pleurocystidia praedita, etiam minoribus ut cheilocystidia. Ad terram muscosam in

silva montana. Malaya, Pahang, Cameron Highland, Corner s.n. 1 Aug. 1934; typus, herb. Corner.

Pileus -4 cm in radius, -7 cm wide, sessile or very shortly pleuropodal, steeply

ascending, spathulate-flabelliform, opaque, minutely villous from the base

outwards, fuscous, paler drab bistre with age. Stem -5 x 2.5-4 mm, thinly villous,

pallid white. Gills deeply decurrent, almost from the base of the fruit-body, very

crowded, 12-16 primaries 2 mm wide, 6-7 ranks, shining white, edge serrulate.

Flesh 2-2.5 mm thick at the base of the pileus, the gelatinous layer c. 300 um

thick. Smell farinaceous, strong.

On the ground among moss in montane forest, solitary, Malaya, Pahang,

Cameron Highlands, 1 Aug. 1934.

Spores 3.5-4.3 x 3-3.5 um, white, smooth, broadly ellipsoid, inamyloid. Basidia

14-19 x 4-4.5 um; sterigmata 4, 3 um long. Cheilocystidia 35-60 x 5-7 um,

clavate or obtusely subventricose, thin-walled, smooth, as a broad sterile gill-

edge. Pleurocystidia 55-120 x 10.5-22 um, ventricose, shortly acuminate, walls

2-7 uum thick yellowish, smooth or thinly encrusted distally, smaller towards the

gill-edge. Hyphae monomitic, clamped; in the firm layer of the flesh longitudinal

with cells 40-120 x 4-23 um, often rather constricted at the septa, thin-walled, in

the gelatinous layer 2-4 um wide and ascending; in the gill-trama as in the firm

layer of the flesh, descending. Surface of pileus with a discontinuous pellicle, 1-2

hyphae thick, of 1.5-3 um hyphae developing fascicles of the hyphae as the

villous layer; pileocystidia either as thick-walled pleurocystidia 40-90 x 6-8 um

or as the cheilocystidia.

This is close to the American H. angustata (p. 39) and differs in the larger

pileus, wider gills, thicker gelatinous layer in the pileus, the form of the

cheilocystidia, the strongly inflated hyphae in the firm layer of the flesh, the

descending hyphae in the gill-trama, and the smaller spores. The terricolous

habit, the fairly large fruit-body and the shining white gills are distinctive among

the Malesian species.

Figure 6. Hohenbuehelia pahangensis. Surface of pileus and (below) firm layer of

flesh, x 500.

{

Hohenbuehelia perstriata sp. nov.

Pileu -22 mm radio, spathulatus dein subreniformis, lateralis subsessilis, ex integro striatus, pallide

griseicervinus, sicco villosulus. Lamellae distantes, 4-9 primariae 2-3.5 mm latae, ordinibus 3-4, albae.

Caro tenuis, strato firmo 100-400 Um crasso, strato gelatinoso 60-200 Lim crasso. Odor farinaceus vel

nullus. Sporae 4.5-6 x 3.5-4.5 Lim, laeves, late ellipsoideae vel subglobosae. Basidia 2-3-4 sporigera.

Cheilocystidia 10-24 x 4-8(-10) Lum, clavata vel ventricosa, nonnulla capitulo breviter stipitato praedita.

Pleurocystidia (30-)50-140 x 9-17 Um, fusiformia acuta, tunicis -6 Um crassis, apices versus haud vel vix

incrustata. Hyphae in carnis strato firmo 3-8 Um latae Pileocystidia -85 x 4-7 Um ut pleurocystidia sed

angustiora, immersa vel superficialia erecta vel in tomenti fasciculis; superficies pilei basim versus

cellulae clavatae -26 x 10 Lum. Ad dejectamenta Aracearum Palmarumque. Insulae Solomonenses. Typus,

Kolombangara, RSS 1275; herb. Corner.

Pileus -22 mm radius, spathulate to subreniform, slightly ascending or

descending (RSS 1104), lateral, subsessile, wholly striate, pale fawn greyish,

drying finely villous-scurfy towards the base. Gills decurrent, distant, 4-9 primaries

2-3.5 mm wide, 3-4 ranks, interstices smooth, white. Flesh with a thin upper

gelatinous layer. Smell none or of meal (RSS 1/04).

On dead remains of aroids and palms (Caryota) in the forest. Solomon Islands.

Spores 4.5-6 x 3.5-4.5 um, white, smooth, ellipsoid to subglobose. Basidia

with 2-3-4 sterigmata (RSS /104). Cheilocystidia 10-24 x 4-8(-10) um, clavate or

subventricose, some with a shortly stipitate subcapitate swelling. Pleurocystidia

(30-)50-140 x 90-17 um, fusiform, acute, wall -6 um thick, smooth or slightly

encrusted distally. Hyphae monomitic, clamped; firm layer of the flesh 100-400

um thick, with thin-walled longitudinal hyphae, the cells -130 x 3-8 um; gelatinous

layer 60-200 um thick hyphae ascending; gill-trama with hyphae as in the firm

layer of the flesh. Surface of pileus with a thin interrupted layer of narrow, short-

celled hyphae 2-5 um wide with thin or slightly thickened walls, developing

cylindric, septate, projecting hyphae -160 um long and becoming more or less

fasciculate; pileocystidia -85 x 4-7 um, as the pleurocystidia, immersed or free

and appressed or erect, becoming incorporated in the excrescent fascicles, also

with a few thin-walled clavate cells -26 x 10 um near the base of the pileus.

Collections.- Solomon Islands, Kolombangara, RSS //04 on dead sheaths and

trunk of Caryota, 27 Aug. 1965; RSS 1275 on a dead aroid stem, 5 Sept. 1965.

The two collections differ slightly but are evidently conspecific.

Hohenbuehelia quadruplex sp. nov. Figure 7

Pileus -30 mm latus, pleuropodalis, spathulatus dein flabelliformis, laevis fuscicervinus, marginem

denticulatum versus substiatus pallidus. Stipes 3-10 x 2-4 mm, applanatus, concolor. Lamellae decurrentes

confertae hispidulosae, 15-26 primariae -1 mm latae, ordinibus 4-6, albae. Caro 1-2 mm crassa, strato

gelatinoso -250 Um crasso, concolor. Odor farinaceus, haud fortis. Sporae 4.5-6 x 3.3-4 Lim, laeves

ellipsoideae. Cheilocystidia 12-24 x 5-14 Um, clavata, nonnulla processu -7 Lim longo, saepe subcapitato,

praedita. Pleurocystidia 40-90 x 11-20 Lum, ventricosa acuta, tunicis flavidulis -6 Lim crassis, apicibus

saepe granuloso-incrustata, haud conferta. Hyphae in carnis strato firmo 3-14 Um latae. Stratum

gelatinosum duplex, superius 50-60 [im crassum, inferius -250 Lim, strato quasi firmo 20-30 Lim crasso

separatum. Superficies stipitis cystidiis crassitunicatis ut pleurocystidia in strato villoso immersis.

Superficies pilei sine cystidiis. Ad lignum putridum in silva. Insulae Solomonenses, San Cristobal, fl.

Warahito, 1 Aug. 1965, RSS 905; typus, herb. Corner.

Pileus -3 cm wide, pleuropodal, spathulate-flabelliform, smooth, fuscous fawn,

paler to the whitish substriate, incurved, minutely denticulate margin. Stem 3-10

x 2-4 mm, more or less flattened in the plane of the pileus, concolorous. Gills

decurrent, crowded, narrow, 15-26 primaries -1 mm wide, 4-5 ranks, hispidulous,

white. Flesh 1-2 mm thick at the base of the pileus, hygrophanous, concolorous,

with a thin gelatinous layer. Smell slight, farinaceous.

On rotten wood in the forest. Solomon Islands, San Cristobal.

ee SEER ele oe

“ = f Mt Sy 45-4

ayy ly

Ui Wl Wh ppl

ay ee

ATS I ih

ome eee 22000 OWT ——

—_—_—_—_——

——_— Oo

Figure 7. Hohenbuehelia quadruplex. Fruit-body in section, 2. Diagram of the

junction of stem and upper side of pileus, to show the origin of the

tissues in the pileus, x 50. Lower figure (from left to right), lower

surface of stem near its junction with the pileus, hymenium, and

surface of pileus, x 500.

Spores 4.5-6 x 3.3-4 um, white, smooth, ellipsoid. Basidia 17-20 x 5 um, 4-

spored. Cheilocystidia 12-24 x 5-14 um, clavate, some with a short, often

subcapitate, process -7 um long, thin-walled, as a sterile gill-edge. Pleurocystidia

c. 40-90 x 11-20 um, ventricose, acute, apex often granular encrusted, walls

yellowish -6 um thick, not crowded. Hyphae monomitic, clamped; in the firm

layer of the flesh with cells -200 x 3-12 um, in the stem with thin or slightly

thickened walls and the cells often unevenly inflated; in the gill-trama with thin-

walled hyphae 3-8 um wide, not mucilaginous; gelatinous layer of the pileus

double, consisting of an inner layer -250 um thick with ascending hyphae 3-5(-7)

tum wide, and a thinner upper layer 50-60 Um thick of ascending hyphae 1-2 um

wide, some of the hyphal ends projecting -60 um, the two layers separated by a

thin subgelatinous layer 20-30 um thick of longitudinal hyphae 2-5 um wide, all

layers devoid of thick-walled cystidia. Surface of stem covered by a thin villous

layer of 2-4 um hyphae -130 um long, 0-2 septate with clamps, often flexuous,

with many erect thick-walled cystidia 40-65 x 9-14 um embedded in the layer.

Surface of the pileus with the upper gelatinous layer pierced here and there by the

shortly excrescent hyphal tips -60 um long; no thick-walled cystidia.

This species is distinguished by the double gelatinous layer of the pileus,

which needs microscopic examination to ascertain, and by the thick-walled

caulocystidia which do not seem to encroach on the surface of the pileus.

Hohenbuehelia singaporensis sp. nov.

Receptacula ex integro fuligineigrisea dein fuligineifusca. Pileus -25 mm latus, sessilis cyphelliformis

dein orbicularis vel unilateraliter flabelliformis, striatus, sicco albidivillosulus. Lamellae subconfertae,

9-12 primarie -1.5 mm latae, ordinibus 5-7. Caro 0.5-2 mm crassa, ex integra gelatinosa. Sporae 6-7 x 4-

4.5 um, laeves, late ellipsoideae. Cheilocystidia 18-26 x 8-15 Um, clavata, tenuiter tunicata, laevia.

Pleurocystidia 45-85 x 12-25 Lim, ventricosa acuta, tunicis -5 Um crassis brunneis, plerumque immersa,

apicibus liberis incrustatis, etiam hymenialia breviora tunicisque tenuioribus. Hyphae 3-5 Um latae, hinc

inde 12 um. Superficies pilei plus minus hymeniidermis, cellulis clavatis -30 x 18 Um ut cheilocystidia,

sine cystidiis crasse tunicatis. Ad lignum in silva. Singapore, Bukit Timah, Corner s.n. 18 April. 1940;

typus, herb. Corner.

Fruit-bodies wholly dark fuliginous grey, becoming fuliginous fuscous. Pileus

-25 mm wide, sessile, at first cyphelliform, becoming subcircular and plano-

resupinate or laterally flabelliform, striate, drying finely whitish villous. Gills

radiating from the origin of the pileus, rather crowded, 9-12 primaries 1-1.5 mm

wide, 5-7 ranks. Flesh 0.5-2 mm thick at the base, wholly gelatinous.

On a dead log in the forest. Singapore, Bukit Timah.

Spores 6-7 x 4-4.5 um, white, smooth, ellipsoid, aguttate. Basidia 18-23 x 5-6

uum; sterigmata 4, 4 um long. Cheilocystidia 18-26 x 8-15 um, clavate, smooth,

thin-walled, with opalescent contents, as a narrow sterile gill-edge. Pleurocystidia

45-85 x 12-25 um, ventricose, acute, wall-5 um thick and brownish, mostly

immersed, some hymenia 38-50 x 11-18 Um with thinner walls, the emergent tip

thinly encrusted. Hyphae monomitic, clamped, 3-5 um wide, walls rather toughly

gelatinous, longitudinal but shortly ascending near the surface of the pileus, some

cells inflated -12 um wide, not forming a dry layer over the gills; in the gill-trama

similar. Surface of pileus more or less hymenioderm as a fairly compact palisade

of clavate cells -30 x 18 um, similar to the cheilocystidia, seated on a thin layer of

compact, short-celled, longitudinal hyphae 3-5 um wide; no thick-walled cystidia.

This fungus is very like H. bullulifera of South America (p. 40) in the clavate

cheilocystidia and the hymenioderm on the pileus, but it had no thick-walled

cystidia on the pileus. It is also like the American H. nigra (Schw.) Singer for

which an initial cyphelliform habit is figured by Waldo (1984), but H. nigra has

differently shaped cheilocystidia and lacks the hymenioderm (Singer and Digilio,

1952, Singer, 1969, Pegler, 1983). I describe the Singapore fungus as a different

species in the hope that further collections will establish its identity.

Hohenbuehelia subdiscipes sp. nov.

Pileus -14 mm latus, reniformis flabelliformis, ad basim subdiscoideum sessilis, pallide cervinibrunneus

vel cervinisubochraceus, tenuiter subtomentosus. Lamellae confertae, primariae 5-7, -1 mm latae, ordinibus

5-6, pallide cervinae. Caro 1-1.5 mm crassa, fere ex integra gelatinosa. Odor farinaceus fortis. Sporae

4.5-6 x 3.3-4.3 Lum, laeves late ellipsoideae. Cheilocystidia 10-18 x 5-8 Lim, clavata, nonnulla subconica.

Pleurocystidia 35-70 x 8-18 Lm, ventricosa lanceolata acuta, tunicis -4 Um, crassis, apices versus incrustata.

Hyphae in carnis strato firmo 60-100 Um crasso 3-7(-9) Lim latae, tenuiter tunicatae. Superficies pilei

cystidiis 25-55 x 4-12 Um ut pleurocystidiis, copiosis, haud immersis, praedita. Ad lignum delapsum in

silva. Insulae Solomonenses, Ysabel, Tetamba, 28 Sept, 1965, RSS 1466; typus, herb. Corner.

Pileus -14 mm wide, sessile to a short subdiscoid base, reniform flabelliform,

convex then plane, pale fawn brown or fawn subochraceous, drying thinly white

subtomentose. Gills crowded, narrow, 5-7 primaries -1 mm wide, 5-6 ranks, pale

fawn. Flesh 1-1.5 mm thick at the base of the pileus, almost wholly gelatinous.

Smell farinaceous, strong.

On a dead log in the forest. Solomon Islands, Ysabel, Tetamba.

Spores 4.5-6 x 3.3-4.3 um, white, smooth, ellipsoid. Cheilocystidia 10-18 x 5-

8 uum, mostly clavate, some subconic, not appendaged, thin-walled, smooth.

Pleurocystidia 35-70 x 8-18 um, ventricose-lanceolate acute, wall -4 Um thick,

granular encrusted at least distally, the incrustation insoluble in KOH, copious.

Hyphae monomitic, clamped; firm layer of flesh 60-100 tum thick, with thin-

walled longitudinal hyphae 3-7(-9) um wide; gelatinous layer much thicker; in

the gill-trama as in the firm layer but more or less descending, yet at the gill-edge

parallel with the edge. Surface of pileus with a thin layer 2-3 hyphae thick

composed of narrow, more or less longitudinal hyphae with cells 20-60 x 2-7 um,

walls thin or slightly thickened, some thinly encrusted, with divergent ends -250

um long near the base of the pileus; pileocystidia 25-55 x 4-12 tum as the

pleurocystidia, encrusted, copious in the superficial layer, not immersed.

This could be regarded as a small or young state of H. testudo but I prefer to

distinguish it at this stage of exploration on account of the fuller colour, the firm

layer of the flesh thicker than the gelatinous layer, and the shorter wider spores.

Hohenbuehelia suppapillosa sp. nov. Plate 1, Figure 8-10

Pileus -5 cm radio, -8 cm latus, pleuropodalis flabelliformis, albus dein pallide fuscibrunneus,

marginem substriatum versus subochraceus, sicco basim versus albivilloslus et papillis gelatinosis -0.5

mm altis sparsim ornatus. Stipes 1-7 mm longus latusque, villosulus concolor. Lamellae confertae, 9-17

primariae -4 mm latae, ordinibus 4-6, albae. Caro 1.5-4 mm crassa, strato gelatinoso c. 1 mm crasso.

Indodora. Sporae 6-7 x 4-5 Um, laeves ellipsoideae. Cheilocystidia (10-)15-30 x 6-10 Lim, ventricosa,

processu filiformi -20 x 1.5-2 Lum saepe subcapitato. Pleurocystidia (23-)55-100 x 12-22 Um, ad lamellarum

basim -130 Lim, ventricosa acuta, tunicis flavidis vel brunneolis 1.5-6 Lim crassis, copiosa. Hyphae in

carnis strato firmo 3-16 Lim latae. Superficies pilei sine cystidiis. Ad lignum emortuum in silva. Malaya,

Sarawak, Typus, Malaya, Pahang, Fraser’s Hill, 1300m alt., Corner s.n. 24 Sept. 1940.

Pileus -5 cm in radius, -8 cm wide, slightly ascending, pleuropodal, flabelliform,

hygrophanous, white then pale fuscous brownish, subochraceous towards the

whitish substriate margin, smooth or slightly papillose when moist, drying thinly

white villous especially near the base with scattered gelatinous papillae -0.5 mm

high. Stem 1-7 mm long and wide, short, thick, pale fuscous ochraceous, drying

wholly finely white villous. Gills decurrent, crowded, 9-17 primaries -4 mm

wide, 4-6 ranks, white. Flesh 1.5-4 mm thick at the base of the pileus, the

gelatinous layer c. 1 mm thick, concolorous drying whitish. Smell none.

On dead logs in the forest and on dead stilt-roots. Malaya, Pahang, Fraser’s

Hill; Sarawak, Bako National Park.

Spores 6-7 x 4-5 um, white, smooth, ellipsoid to pip-shaped, aguttate. Basidia

18-23 x 5-5.5 um; sterigmata 4, 2.5-3 um long. Cheilocystidia (10-)15-30 x 6-10

Figure 8. Hohenbuehelia suppapillosa. Spores, x 1000. Hymenium, pleurocystidia

and cheilocystidia, x 500.

Figure 9. Hohenbuehelia suppapillosa. Upper part of young pileus in section, x 500.

Spores, x 1000.

VE OAT

Figure 10. Hohenbuehelia suppapillosa. Basal part of mature pileus in section, x 250.

um, more or less ventricose with a short or long filiform, flexous or subcapitate

appendage -20 x 1.5-2 um, thin-walled, as a narrow sterile gill-edge. Pleurocrystidia

(23-)55-100 x 12-22 um, -130 um at the base of the gills, ventricose, acute, with

very thick yellowish to brownish walls -6 um thick, arising deeply in the trama

and projecting -50 um, sometimes slightly curved at the apex, smooth, abundant.

Hyphae monomitic, clamped; in the firm layer of the flesh 3-16 um wide, often

branched at a wide angle, longitudinal and interwoven with numerous H-

connections, compact near the gelatinous layer, thin-walled; in the gelatinous

layer mostly 2-5 um wide, some with a few cells 10-20 um wide, mostly ascending;

in the gill-trama as in the firm tissue of the pileus, descending and interwoven,

not gelatinous, the subhymenium rather thick with hyphae 2-4 um wide but not

corticate. Surface of pileus with a narrow, fairly compact, layer of longitudinal

and interwoven hyphae 3-5 um wide but some cells 8-20 um wide, giving rise to

narrow excrescent hyphae in fascicles -250 um high as the villous layer; no thick-

walled cystidia.

Collections.- Malaya, Pahang, Fraser’s Hill 1300m alt., Corner s.n. 24 Sept.

1940.- Sarawak, Bako National Park, on dead stilt-root, 27 Aug. 1972, Corner P-

178 (immature).

This has the tendency to inflate the hyphae in the pellicle of the pileus

as in H. mellea.

Hohenbuehelia testudo (Berk.) Pegler

Agaric Flora of Sri Lanka (1986) 173.

Acanthocystis testudo (Berk.) Boedijn, Bull. Jard. bot. Buitenz. ser. 3, 16 (1940) 402, f. 10.

Pileus 1-3(-6) cm wide, sessile or shortly pleuropodal, spathulate to flabelliform,

nearly white becoming dirty brown, paler to the whitish margin, finely villous

especially towards the base. Gills subdistant, 1-2 mm wide, 3-4 ranks, white.

Flesh c. 2 mm thick, the gelatinous layer 130-390 um thick. Smell farinaceous.

On dead wood. Ceylon, Malaya, Borneo, Krakatau.

Spores 5-6.5 x 3-3.5 um (6-7 x 2.5-3.5 um, Petch; 9-12.5 x 3.7-4.5 um,

Pegler). Basidia 20-24 x 5-6 um, 4-spored (20-25 x 5-6 um, Pegler). Cheilocystidia

15-25 x 4-7 um, narrowly ventricose, often shortly attenuate with or without a

subcapitate apex 1-2 um wide, thin-walled (Pegler). Pleurocystidia 44-78 x 12-18

lum, ventricose, acute, with thick yellowish walls, shorter towards the gill-edge,

the protruding part encrusted (Boedijn); 40-70 x 10-15 um, fusoid ventricose,

wall -7 um thick and deep yellowish brown (Pegler); 40-70 x 3-8 uum, fusiform,

more or less encrusted, (Corner). Hyphae of the firm layer of the flesh 4-8 um

wide, in the gelatinous layer 1-4 um wide, vertically arranged; in the gill-trama as

in the firm layer of the pileus. Villous hairs of the pileus as fascicles -200 x 100

um of encrusted hyphae 3-5 um wide, most of the hyphae ending in encrusted

ventricose-lanceolate pileocystidia 70-80 x 6-12 um; (pileipellis a disrupted repent

epicutis of non-gelatinised hyphae 2-4 um wide, sometimes forming short erect

fascicles, Pegler).

This description is taken mainly from that of Boedijn, with additions from that

of Pegler and observations of mine from a collection which I made at Kandy,

Ceylon. The most remarkable difference is the large size of the spores given by

Pegler who found them on the type-material. It is not clear that they were seen

attached and there is always the possibility that they were a deposit from another

fungus during collection. In recording the species from Malaya and Borneo,

Pegler does not mention the spores of these collections. He gives the basidia as

the same size as found by Boedijn, which seems unlikely with spores nearly

twice as big as Boedijn and I found.

A collection which I made in Brunei (Ulu Belalong 15 Feb. 1959), with pilei -

5 cm wide, differed macroscopically in the crowded white gills (1.5-2 mm wide

in c. 5 ranks). It had, also, the strong farinaceous smell which I noted in the

specimen from Kandy. There were also slight microscopic differences, thus:-

Spores 5-6 x 3.3-3.7 um. Basidia 20-24 x 6 um, 4-spored. Cheilocystidia

small, clavate to ventricose-capitate. Pleurocystidia 45-90 x 14-20 um, not or

slightly encrusted. Hyphae of the firm layer of the flesh -10 um wide, short-

celled, longitudinal in a narrow compact layer below the gelatinous layer, the

rest of the firm layer composed of laxly interwoven and longitudinal hyphae,

with walls -1 um thick in the layer just over the gills; in the gill-trama 3-7 um

wide, with submucilaginous walls -0.5 um thick. Surface of the pileus consisting

of a thin superficial layer 8-15 um thick, composed of longitudinal hyphae 2-

3.5 um wide with scattered processes 2.5-4 um wide as if sterile basidia;

fascicles of excrescent hyphae -200 x 70-100 um, with narrow lanceolate,

thick-walled, more or less encrusted pileocystidia -80 x 5-7 um towards the tips

of the fascicles, with rather thin-walled cystidia like the cheilocystidia sparsely

set on the sides of the fascicles.

var. glabra var. nov.

Differt pileo glabro, raro ad basim subvillosus; lamellis confertis, ordinibus 3-6, albis dein pallide

ochraceis vel pallide cervino-ochraceis. Typus, Borneo, RSNB 3011; herb. Corner.

Pileus 1-4 cm in radius, 1-6 cm wide, sessile or attenuate to a short lateral

stem, spathulate to flabelliform, at first descending then horizontal and finally

more or less ascending, undulate, smooth, subviscid, generally glabrous,

sometimes thinly subvillous or spiculose-villous at the base, white then pallid

ochraceous to pale fawn ochraceous, varying pale umber, greyish or fuscous

bistre, opaque or striate towards the margin. Stem -10 x 1.5-5 mm, lateral,

minutely villous. Gills decurrent, crowded, narrow, not dichotomous, 12-40

primaries (5-9 in small fruit-bodies) 1-3 mm wide, 3-6 ranks, white then pale

cream ochraceous to concolorous with the pileus, edge entire. Flesh 0.5-3 mm

thick at the base of the pileus, with a very thin gelatinous layer at the surface.

Smell farinaceous, slight or strong, or none (P-/65, RSS 1671), or slightly

fragrant (New Guinea 1 Oct. 1960).

On rotten wood, fallen trunks, and on the ground under rotten wood. Malaya,

Borneo, New Guinea, Solomon Islands.

Spores 5-6.5 x 3.5-4.5 um, white, smooth, ellipsoid to lacrymiform, aguttate

but drying 1-2 guttulate, inamyloid. Basidia 20-25 x 5-7 um or 18-22 x 5-7 um;

sterigmata 4; subhymenium 12-20 um thick, composed of 2-3 lm interwoven

hyphae. Cheilocystidia -25 x 4-8 uum, clavate or subventricose or subcylindric,

often with a short process with or without a subglobose head 1.5-2.5 um wide.

Pleurocystidia 30-90 x 7-20 um in some collections -120 x 25 lum, ventricose

with acute apex, wall -6 um thick, thinly encrusted or smooth, abundant. Hyphae

monomitic, clamped; in the firm layer of the flesh with cells 20-150 x 3-14 um

with wall thin or 0.5-1 um thick near the base of the pileus, longitudinal and

interwoven; gelatinous layer 100-250 um thick, with ascending hyphae 2-4(-6)

Lum wide; in the gill-trama 3-8 lum wide, thin-walled, descending; in the stem with

cells 30-150 x 3-20 um, often with ampulliform swellings at one end of the cell,

walls 0.5-1 um thick. Surface of stem with a sterile hymenium, becoming finely

villous with 2-3 um hyphae. Surface of pileus with a rather compact but thin

layer, 20-30 um thick, of appressed radiating hyphae 3-6(-8) Um wide, with firm

or slightly gelatinous walls, not developing excrescent hyphae except in some

cases at the very base of the pileus; pileocystidia 60-150 x 4-10 um, narrowly

fusiform, often with long stalk, acute, smooth or slightly encrusted, thick-walled,

generally immersed in the superficial layer and some arising from the outer

hyphae of the firm layer of the flesh and embedded in the gelatinous layer, not

projecting from the surface.

Collections.- Sarawak, Bako National Park, Corner P-165, 26 Aug. 1972.-

North Borneo, Mt Kinabalu, 1300-3300m alt., RSNB 30/1, 5183, 5226, 5226B,

5728 and Corner s.n. 30 June 1961, apparently throughout the year.- New Guinea,

Oomsis, Corner s.n. 1 Oct. 1960.- Solomon Islands, Guadalcanal RSS 1671];

Kolombangara RSS 1060; San Cristobal RSS 738, 971.

I treat this as a variety because I think that the whole complex about H.

testudo needs much more investigation, especially in regard to the size of the

spores, pleurocystidia and pileocystidia. Though the fruit-bodies usually smell

strongly of new meal, the smell may be weak or absent, and those of the New

Guinea collection were slightly fragrant. This collection also had the pilei becoming

slightly villous-spiculose at the base with the pileocystidia embedded in the

excrescent fascicles. Compare the Brazilian H. testudo (p. 45).

Hohenbuehelia vermiculata sp. nov. Figure 11

Pileus -8 mm radio, -7 mm latus, breviter pleuropodalis, vel sessilis, primo conchiformis fere

cyphelliformis, dein unilateraliter extensus, spathulatus griseo-alutaceus striatus, sicco basim versus

subvillosus. Stipes -1 x 2 mm. Lamellae subconfertae, primariae 5-7, -0.7 mm latae, ordinibus 3-4,

pallide subochraceae. Caro 1 mm crassa, fere ex integra gelatinosa, strato firmo 50-200 Lim crasso. Odor

farinaceus. Sporae 4-6.5 x 3-4 Um, laeves subcylindricae. Cheilocystidia 9-23 x 4-7 Um, clavata vel

ventricosa, nonnulla capitulo breviter stipitato praedita. Pleurocystidia 25-70 x 7-17 Lm, ventricosa

acuminata, dense incrustata. Hyphae in carnis strato firmo 2-6 Um latae. Pileocystidia 40-130 x 3-5(-6)

tum, dense incrustata, in toto -200 x 12-20 Um vermiformia. Ad ramulos truncosque delapsos in silva.

Insulae Solomonenses. Typus RSS 1160; herb. Corner.

Pileus -8 mm in radius, -7 mm wide, sessile or shortly pleuropodal, at first

convex-conchiform and almost cyphelliform, then unilaterally extending and

spathulate, pale greyish buff, translucent striate, drying thinly villous at the base

and pale bistre. Stem -1 x 2 mm, very short or none. Gills decurrent to the

attachment, rather crowded, 5-7 primaries -0.7 mm wide, 3-4 ranks, pale buff

ochraceous. Flesh 1 mm thick at the base of the pileus, mostly gelatinous, the

firm layer 150-200 um thick, or merely 50-100 um (RSS 681). Smell farinaceous.

On a fallen trunk in the forest, on sticks and on dead inflorescences of Ficus

subcongesta. Solomon Islands.

Figure 11. Hohenbuehelia vermiculata. Fruit-body in section, x 2. Spores, x 1000.

Cheilocystidia, pleurocystidia and surface of pileus, x 500.

Spores 4-6.5 x 3-4 um, white, smooth, subcylindric. Basidia 18-20 x 5-5.5

uum, 4-spored; no subacerose basidioles; subhymenium thin, with 2-3 um hyphae.

Cheilocystidia 9-23 x 4-7 um, clavate to ventricose with a short, often capitate,

appendage, smooth, thin-walled, as a sterile gill-edge. Pleurocystidia 25-70 x 7-

17 um, the smaller towards the gill-edge, ventricose, acuminate, heavily encrusted

(at least distally), wall -6 um thick. Hyphae monomitic, clamped; in the pileus

mostly gelatinous 1.5-2.5 um wide and ascending, in the firm layer 2-6 um wide

with slightly thickened walls, longitudinal; in the gill-trama as in the firm layer,

2-4 um wide. Surface of pileus with a thin layer of longitudinal hyphae 3-6 um

wide bearing scattered subclavate processes -25 x 4-7 um, and erect hyphal ends

-70 x 3-6 um, often 1-3 septate; pileocystidia 40-130 x 3-5(-6) um, appearing as

vermiform, often curved ropes of granular and crystalline material -200 x 12-20

um overall, thick-walled.

Collections.- Solomon Islands, Guadalcanal, 4 July 1965, RSS 681;

Kolombangara, 30 Aug. 1965, RSS 1160.

This is close to the alliance of H. testudo v. glabra and may be a depauperate

state.

Hohenbuehelia horakii Courtecuisse species incertae sedis

Doc. mycol. 14 (1984) 82.

Claudopus griseus Mass., Kew Bull. 1899, 169.- H. grisea (Mass.) Horak, Nova Hedwigia Beih. 65 (1980)

315, non H. grisea (Pk) Singer.

Massee gave: pileus 3-5 cm wide, conchiform with lateral stem, grey, often

corrugate; gills distant, narrow, connected by veins, grey; spores 8 x 5 um, rough;

cystidia 65-70 x 14-15 um, fusoid; on dead wood, Malay Peninsula, Perak,

Ridley 11.

Horak gave: spores 4.5-6 x 2.5-3 um, reniform, inamyloid; cystidia 30-50 x

12-18 um, fusiform, metuloid, wall -7 um thick, hyaline, partly encrusted with

crystals.

I do not recognise this fungus.

Notes on Extra-Malesian Species

The species here mentioned can be arranged according to my classification of

the Malesian species in the following way:

Species with thick-walled pileocystidia.

H. angustata, H. bullulifera (with hymenioderm), H. carbonaria, H.

cystidioides, H. petaloides, H. testudo.

Species without such pileocystidia.

a. With thick-walled hyphae in the firm layer of the flesh. H. atrocaerulea,

H. myxotricha, H. aff. reniformis.

b. Hyphae thin-walled - H. subtorulosa.

Hohenbuehelia angustata (Berk.) Singer

Lilloa 25 (1950) 109.

Pileus -3.5 cm in radius, 4.5 cm wide, pleuropodal, spathulate-flabelliform,

finely villous in the proximal half, striate near the margin, grey, pallid fuscous or

pale livid bistre. Stem 3-30 x 2-5 mm, lateral, fibrous, firm, finely villous, paler

concolorous. Gills deeply decurrent, very crowded, narrow, thin, 15-40 primaries

0.3-1 mm wide, 5-6 ranks, sometimes dichotomous especially near the stem,

white, edge entire. Flesh fibrous, firm, white, with a very thin gelatinous layer c.

50 um thick. Smell farinaceous, strong.

On the ground and on rotten wood in forest. North and South America.

Spores 6.5-8 x 4-5 um, ellipsoid, or 4-4.5 x 3-3.5 um, broadly ellipsoid to

subglobose (Manaus collection), white, smooth, thin-walled. Basidia 17-23 x 6

um, 4-spored.

Cheilocystidia 15-30 x 4-8 um, cylindric, clavate or subventricose with a short

obtuse to subacute, rarely capitate, apex, smooth, thin-walled, as a narrow sterile

gill-edge. Pleurocystidia 35-85 x 14-28 um, stoutly ventricose with short acute

apex, with thick yellowish walls, not or slightly encrusted, abundant. Hyphae

monomitic, clamped, 2-10 um wide in the firm tissue of the pileus, mostly

longitudinal; in the gill-trama 2-6 um, wide, thin-walled, not gelatinous, mostly

radiating and parallel with the gill-edge; gelatinous layer of pileus 20-50 um

thick. Surface of pileus with a compact layer c. 20 um thick composed of

longitudinal and interwoven appressed hyphae 2-4 um wide, with few excrescent

ends in fascicles -120 um wide of hyphae 2-8 tum wide; pileocystidia 28-70 x 6-

10 um, fusiform with prolonged acute apex and short, often subtruncate, base,

becoming overgrown by the villous layer, sometimes with thin-walled pileocystidia

in the fascicles of hyphae and hyphal ends like irregular, often lobate and contorted,

cheilocystidia and often with pale fuscous sap.

Collections.- Bolivia, Santa Cruz de la Sierra, on the ground, scattered or 2-3

together, 17 Jan. 1948, Corner 41/48. - Brazil, Amazonas, Manaus, on rotten

wood, Corner s.n. 24 Oct. 1948.

The stipitate pileus, very crowded narrow gills and the very thin gelatinous

layer on the pileus distinguish this species. Singer gave the spores as equally

variable, 3.5-6.7 x 3-4.5 um, ellipsoid to subglobose. Noteworthy is the radiate

construction of the gills, as in Panus, which probably explains why they are so

narrow. The limits between this species and H. testudo seem rather uncertain.

Thorn and Barron (1986), however, give the gelatinous layer as 180-250 um

thick.

Hohenbuehelia bullulifera Singer

Lilloa 25 (1952) 119.

var. brasiliensis var. nov. Figure 12

A typo differt pileo majori -20 mm lato, lamellis confertis, odore farinoso, sporis minoribus 4.5-5.5 x

3-3.5 Um. Typus, Brazil, Corner 296; herb. Corner.

Pileus -15 mm in radius, 20 mm wide, sessile, dorsally attached, at first

cyathiform then unilaterally spathulate to flabelliform, sometimes with a small

resupinate foot, wholly dark fuscous fuliginous, at first paler to the whitish and

persistently incurved margin, opaque or faintly striate near the margin, drying

white villous at least in the proximal part. Gills radiating from the base, crowded,

narrow, thin, tough, 7-11 primaries -1 mm wide, 4-6 ranks, concolorous; sometimes

with minute gills on the resupinate foot. Flesh 1-1.5 mm thick at the base, wholly

toughly gelatinous. Smell farinaceous, rather strong.

Figure 12. Hohenbuehelia bulluifera v. brasiliensis. Surface of pileus, x 500.

On dead wood in forest. Brazil.

Spores 4.5-5.5 x 3-3.5 um, white, smooth, aguttate. Basidia 18-23 x 5.5-6.5

um; sterigmata 4, 3 um long. Cheilocystidia 15-30 x 7-16 um, clavate, pyriform

to subglobose, with thin, colourless to pale fuscous walls, smooth, with a few

colourless plastid-like inclusions, as a broad sterile gill-edge. Pleurocystidia

(20-)40-90 x (6-)12-18 tum, fusiform to ventricose, with fuscous brown walls 2-5

um thick, often with a more or less distinct neck leading to the acute apex, arising

deeply in the subhymenium, thinly encrusted distally; the largest with deepest

origin near the bases of the gills, shorter towards the gill-edge, the shortest 20 x 6

um mixed sparsely with the cheilocystidia; also with thin-walled cystidia like the

cheilocystidia on the gill-surface especially towards the gill-edge. Hyphae

monomitic, clamped; in the flesh 2-5 um wide with rather toughly gelatinous

walls, longitudinal but shortly ascending near the surface, some with inflated

cells 8-16 um wide, compact but not forming a firm dry layer over the gills; in the

gill-trama similar. Surface of pileus more or less hymenioderm with cells -30 x

20 um like the cheilocystidia, some with pale fuscous walls, seated on a compact

layer of longitudinal hyphae 3-5 um wide with fuscous walls; pileocystidia like

the pleurocystidia but not so large, sometimes only near the base of the pileus;

excrescent hyphae 3-6 um wide in fascicles.

Collections.- Estado do Rio, Niteroi, Corner s.n. 7 Sept. 1947; Rio de Janeiro,

Corovado 500m alt., 21 Nov. 1948, Corner 296.

This agrees essentially with Singer’s description for H. bullulifera with spores

5.5-6.8 x 3-4 lum. Compare the very similar H. singaporensis (p. 29) without

thick-walled pileocystidia.

Hohenbuehelia atrocaerulea (Fr.) Singer

I give the following notes on an English collection.

Spores 7-9 x 3.5-4 um, subcylindric, aguttate. Basidia 25-30 x 5.5-6.3 um, 4-

spored. Cheilocystidia as sterile basidia, subclavata to ventricose and then often

with a short apical and sometimes capitulate process. Pleurocystidia -110 x 10-17

uum, encrusted, smaller towards the gill-edge. Hyphae in the firm layer of the

flesh 3-7 um wide, walls 0.5-1 um thick and pale brown near the gelatinous layer

0.5-1 mm thick with hyphae 1-3 um wide ascending in the upper part but

longitudinal in the lower part of the layer; in the gill-trama as in the firm layer of

the flesh but with thickened walls only in the central part of the trama, descending.

Surface of pileus without pileocystidia.

Hohenbuehelia carbonaria (Cke et Mass.) Pegler

Austral. J. Bot. 13 (1965) 327.

Panus carbonaria Cke et Mass., Grevillea 15 (1887) 94; Cooke, Handbook Austral Fungi (1892)

pl. 7, fr. 46.

Spores 7-9 x 4.5-5 um (Pegler), 8-10 x 5-5.5 um (Corner). Pleurocystidia 55-

80 x 9-14'um (Pegler), 60-100 x 9-16 um (Corner), fusiform acute, thick-walled,

distally encrusted. Pileocystidia -120 x 4-7 um, fusiform, acute. Australia.

When studying Panus some years ago, I examined the type at Kew and noted

that it had the construction of H. geogenia from which it differed in the larger

spores. However, I failed to note the direction of the hyphae in the gelatinous

layer of the pileus.

Hohenbuehelia cystidioides (C.G. Lloyd) comb. nov.

Cantharellus cystidioides C.G. Lloyd, Myc. Writ. 7 (1923) 1227, f. 2534, 2535.

Pileus sessile, dimidiate, smooth, brown (about the colour of Auricularia

auricula-judae). Gills dichotomous, thick, crowded, pruinose from the cystidia.

Flesh wholly gelatinous. Spores 4-5 x 2 um, hyaline, smooth, oblong ellipsoid,

thin-walled. Basidia c. 25 x 5 um. Cheilocystidia not seen, the gill-edge fertile.

Pleurocystidia 55-120 x 15-25 um, ventricose, the acute apex projecting 8-40 um

and rather thinly encrusted with crystals (often tetrahedral) insoluble in KOH,

walls 3-8 um thick, base attenuated and deeply inserted in the gill-trama, often

curved ascending, abundant. Hyphae monomitic, clamped, 2-4 um wide, the

gelatinous walls thin or slightly thickened, branched at a wide angle but with

many blind, secondarily septate, endings; in the gill-trama similar but not

gelatinous. Surface of pileus with densely interwoven, more or less contiguous,

hyphae with firm (not gelatinous) walls, in a layer 70-100 um thick; pileocystidia

as the pleurocystidia but shorter, scattered.

This was described from Japan, collected by J.E.A. Lewis. I examined the

collection at Kew, when I was studying cantharelloid fungi, and have added the

microscopical details. The species is remarkable for the dichotomous gills with

fertile edge, the wholly gelatinous flesh and the unusually thick pellicle.

Hohenbuehelia myxotricha (Lév.) Singer

The following are my notes on an English collection

Spores 9-14 x 4-5.5 um, cylindric, often slightly curved, aguttate or with a

few minute guttulae. Basidia 25-33 x 4-6 um; sterigmata 2, 5.5-7 um long.

Cheilocystidia -50 x 3.5-8 Um, mostly ventricose with an apical cylindric

appendage -30 x 1.5-2.5 um, some with a slightly swollen tip, others merely

clavate as sterile basidia even with 2 sterigmata, with all transitions to emergent

hyphae 1.5-2.5 um wide, as a sterile gill-edge. Pleurocystidia 30-70 x 7-10 um,

subclavate, fusiform or cylindric, thick-walled, encrusted with a distal conical

cap of compact crystals. Hyphae monomitic, clamped; in the stem, the firm layer

of the flesh and in the gill-trama 2.5-8 um wide, with thick walls; in the gelatinous

layer 1-2 um wide. Surface of pileus without cystidia.

Hohenbuehelia petaloides (Fr.) Singer Figures 13,14

Huijsman (1961); Murata (1979); Singer and Kuthan (1980); Doneso (1981); Watling (1985).

: seas = eo Ie ee

This is the north temperate complex of H. petaloides (type-species of the

genus), H. geogenia (Fr.) Singer, and H. repanda Huijsman. Its southerly allies

are H. angustata and H. testudo; all have the farinaceous smell. For H. geogenia,

I note the following points.

Hyphae in the firm layer of the flesh with cells 40-130 x 3-12 um, thin-walled

but in the stem with walls 0.5-1 um thick; in the gelatinous layer (up to 700 um

thick) 3-6 um wide, ascending; in the rhizomorphs monomitic, clamped, 1.5-

3.5(-5) um wide, thin-walled but the outer hyphae with slightly thickened and

slightly encrusted walls; in the gill-trama 3-7 um wide, descending. Surface of

pileus with pileocystidia 50-110 x 4-7 um, narrowly fusiform, with thick brownish

walls, thinly encrusted.

a= s

Se SRN \

== 57]

: i /

VLG wm

(i yy

jets y My z

Wat WAG SEZ

ay (| S77, J

SSW /

Figure 13. Hohenbuehelia geogenia. Margin of pileus in section, to show the upper

gelatinous layer of the flesh with ascending hyphae and thick-walled

pileocystidia (thick lines), and the pleurocystidia (thin lines); x 45.

Hohenbuehelia aff. reniformis (Fr.) Singer

Pileus -12 mm in radius, sessile or with a very short vague stem attached by a

thin white narrow byssoid mycelium, reniform semicircular, fuscous cinereous,

paler and substriate near the margin, drying pallid fuscous and shortly white

;

Figure 14. Hohenbuehelia geogenia. Surface of pileus with thick-walled pileocystidia

arising in the gelatinous layer; x 1000.

villous, pruinose towards the margin, strigose near the base. Gills decurrent,

crowded, narrow, tough, 8-13 primaries -1 mm wide, 4-6 ranks, white, in age

yellowish. Flesh 0.3-0.5 mm thick, gelatinous except a layer c. 100 um thick over

the gills. Inodorous.

On a dead branch in forest. Brazil, Estado do Rio, Niteroi, Corner s.n. 31

Aug. 1947.

Spores 8-10 x 4-5 um, white, smooth, subcylindric, aguttate, inamyloid. Basidia

22-30 x 7-8 um; sterigmata 4, 3-4 um long. Cheilocystidia 16-32 x 5-10 um,

ventricose with a subcapitate, mostly subhastiform, appendage 2-3 um wide on a

stalk 1.5-3 um long, thin-walled, as a sterile gill-edge, mixed with pleurocystidia.

Pleurocystidia 25-75 x 7-16 um, lanceolate fusiform, some waisted, walls 1-3.5

um thick and colourless, rather thinly encrusted at the acute apex, abundant and

also on the gill-edge. Hyphae monomitic, clamped; in the firm layer of the flesh

and in the gill-trama 3-7 um wide with walls 0.5-1.5(-2.5) um thick, descending

in the gills; in the gelatinous layer 1-3 um wide. Surface of pileus with subclavate

processes -35 x 4-10 um in a loose layer, with pale fuscous sap, arising from a

loose pseudoparenchyma of cells 12-25 x 7-12 um with fuscous walls and fuscous

granular incrustation; villous layer composed of loose fascicles of excrescent

hyphae; without thick-walled cystidia.

This fits the description of the temperate H. reniformis, yet it fits also diminutive

H. malesiana. I describe it because too little is yet known of these fungi.

Hohenbuehelia subtorulosa (Cke) comb. nov.

Panus subtorulosus Cke; Pegler, Kew Bull. Add. Ser. X (1983) 258.

I give the following notes from my examination of the type-specimen at Kew,

namely Glaziou 9153, Dec. 1878, Rio de Janeiro.

Fruit-bodies closely caespitose-merismatoid from a short thick common trunk,

dried wholly fuliginous fuscous and horny. Pileus -25 mm in radius, often with a

prolonged lateral stem, finely villous over the central part. Gills crowded, very

narrow, 0.5 mm broad. Spores 4-5 x 2.5-3 um, colourless, smooth. Pleurocystidia

35-48 x 9-16 um, ventricose, subacute to acute, short, very thick-walled, apparently

smooth, very abundant. Hyphae of the flesh wholly agglutinated, no structure

discernible. Surface of pileus with the villous layer -80 um thick, composed of 3-

5 um hyphae, clamped, more or less erect; no pileocystidia seen.

Hohenbuehelia testudo

The following description refers to a fungus that I found in Brazil and seems

almost identical with the Asian H. testudo (p. 34). Both H. testudo and H.

angustata relate to the complex of H. petaloides.

Pileus -5 cm in radius, -7 cm wide, sessile, lateral, ascending, spathulate to

flabelliform, pale watery ochraceous to pale fuscous brownish, whitish towards

the substriate margin, drying white villous. Gills decurrent, crowded, narrow, 14-

20 primaries 2 mm wide, 5-6 ranks, subreticulate at the base, white to dingy

cream. Flesh 5-6 mm thick at the base, with a thin gelatinous layer above the

softly floccose firmer flesh, white. Smell farinaceous.

On fallen trunks in forest. Brazil.

Spores 5-6 x 3-3.5 um, white, smooth, broadly ellipsoid, thin-walled. Basidia

16-21 x 5-5.7 um, (adherent in mucilage), 4-spored. Cheilocystidia -23 x 5-9 um,

subventricose with a more or less prolonged apex, as a sterile gill-edge but

collapsing. Pleurocystidia 35-90 x 9-23 tum, with thick yellowish walls, smooth

or slightly encrusted, ventricose acute. Hyphae monomitic, clamped; in the firm

layer of the flesh with cells 20-140 x 3-24 Um, thin-walled; in the gelatinous layer

2-7 um wide but some -18 um wide, ascending or vertical, some of the wider

hyphae with a secondary septum; in the gill-trama as in the firm layer of the flesh

but with slightly thickened walls, descending. Surface of pileus with a more or

less disrupted pellicle of longitudinal and interwoven hyphae 2-6 (-12) um wide,

1-3 hyphae thick, developing excrescent clamped hyphae 4-7 Lum wide in fascicles

-100 x 10-50 um; pileocystidia -90 x 7-11 um as smooth thick-walled

pleurocystidia, becoming decumbent, scattered.

Collections.- Rio de Janeiro, Corcovado, 2 Dec. 1948, Corner 316; Amazonas,

Manaus, near Flores, 3 Oct. 1948, Corner 1235.

References

Corner, E.J.H. (1947). Variation in size and shape of spores, basidia and cystidia

in Basidiomycetes. New Phytologist 46: 195-228.

— (1966). A monograph of cantharelloid fungi. Ann. Bot. Mem. n. 2. Oxford

University Press.

— (1968). Mycology in the tropics - Apologia pro monographia sua secunda.

New Phytologist 67: 219-228.

— (1981). The agaric genera Lentinua, Panus and Pleurotus. Nova Hedwigia

Beiheft 69.

— (1983). Ad Polyporaceas I. Amauroderma and Ganoderma. Nova Hedwigia

Beiheft 75.

— (1984). Ad Polyporaceas Il. Nova Hedwigia Beiheft 78.

Dennis, R.W.G. (1953). Some pleurotoid fungi from the West Indies. Kew

Bull. 1953: 31-45.

Donoso, J. (1981a). Contribution a la connaissance des Agaricales lignicoles du

Chili. Bull. Soc. Mycol. Fr. 97: 23-31.

— (1981b). Hohenbuehelia S. Schulz. Definition et affinités taxonomiques.

Cryptog. Mycol. 2: 153-162.

Horak, E. (1981). Conchomyces v. Overeem - an independent genus within the

Agaricales Ann. Mycol. Ser. Il, 34: 109-114.

Hujsman, H.S.C. (1961). Observations sur le genre Hohenbuehelia.

Persoonia 2: 101-107.

Jiilich, W. (1981). Higher taxa of Basidiomycets. Bibliotheca Mycologica 85.

Lazo, W. (1984). Introduccion al estudio de los hongos superiores, III. Boletin

Mycologica 2: 27-66.

Murata, Y. (1979). New records of gill-fungi from Hokkaido (5). Trans. Mycol.

Soc. Japan 79: 317-324.

Natarajan, K. and N. Raman (1980). South Indian Agaricales VIII. Agaricochaete

indica. Mycologia 72: 630-632.

Pegler, D.N. (1966). Tropical African Agaricales. Persoonia 4: 73-124.

— (1977). A preliminary Agaric Flora of East Africa. Kew Bull. Add Ser. VI.

— (1983). Agaric Flora of the Lesser Antilles. Kew Bull. Add. Ser. IX.

— (1986). Agaric Flora of Sri Lanka. Kew Bull. Add Ser. XII.

Pouzar, Z. (1981). Faerberia, a genus of cantharelloid fungi. Ceska

Mykologie 35: 185-188.

Singer, R. (1969). Mycoflora Australis. Nova Hedwigia Beiheft 29.

— (1975). Agaricales in Modern Taxonomy. J. Cramer, Vaduz.

— and A.P.L. Digilio (1952). Prodromo de la Flora Agaricina Argentina. Lilloa

25: 5-462.

— and J. Kuthan (1980). Comparison of some lignicolous white-spored American

agaries with European species. Ceska Mykologie 34: 57-73.

Stevenson, G. (1964). The Agaricales of New Zealand, V. Kew Bull. 19: 1-59.

Thorn, R.G. and G.L. Barron (1986). Nematoctonus and the tribe Resupinateae in

Ontario, Canada. Mycotaxon 25: 321-453.

Watling, R. (1985). Hymenial surfaces in developing agaric primordia. Bot. J.

Linn. Soc. 91: 273-283.

On the Agaric Genera Hohenbuehelia and Oudemansiella

Part II: Oudemansiella Speg.

E.J.H. CORNER

91 Hinton Way

Great Shelford

Cambridge CB2 5AH

England

Abstract

This is mainly an account of Malesian species, of which O. crassifolia, O. lianicola and O. submucida are

new, but notes are added on several temperate and South American species. The structure of the pileus

introduces a new criterion into the specific classification of the genus. The connection with marasmioid Xerula

is discussed. A species from Japan, referred tentatively to O. radicata var. hygrophoroides (Sing. et Clemengon)

Pegler et Young is intermediate in that it has the marasmioid subacerose basidioles in the hymenium. It is re-

affirmed that the pleurocystidia conform with the basidiograph locus of Oudemansiella and it is suggested that

the narrow basidia of Hohenbuehelia and Pleurotus and the subacerose basidioles of Marasmius correspond

with the developing and uncharged basidia of Oudemansiella.

Contents

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Recent publications on this genus have enabled me to prepare this account of

five Malesian species. In 1929, soon after I arrived in Singapore, I was intrigued

by two common agarics. One I identified as Collybia apalosarca, now known to

be one of the many synonyms of O. canarii. The other I called uncertainly C.

radicata and it has passed unidentified, but I know now that it is C. altissima

Mass., described from Singapore; the trouble in those days was the inadequacy of

conventional description, as well as mistrust of the erratic mycologist George

Massee. I have added three new species, mainly from later collections which I

made in Borneo.

Oudemansiella, though very unlike the majority of marasmioid fungi in general

appearance, now appears as the remains of the ancestry of Marasmieae and as a

connection with Pleurotus. The genus is distinguished by the fleshy fruit-body,

commonly with viscid brown to white pileus, the central stem, the large white

inamyloid spores with a single large oil-drop or many small ones, the large

cheilo- and pleurocystidia, the long-celled thin-walled hyphae with clamps, the

palisade of clavate cells on the pileus, and the lignicolous habit, though there may

be exception in one point or another. Seventeen species, some with several

varieties, are recognised by Pegler and Young (1986), to which Redhead, Ginns

and Shoemaker (1987) add four others, but some of these are reduced by Boekhout

and Bas (1986). The ultra-structure of the spore-wall has been studied in detail by

Pegler and Young.

Classification of Oudemansiella

For various reasons but chiefly because it omits the structure of the surface of

the pileus, I find that the revised classification of the species by Pegler and

Young (1986) is unsatisfactory. My criticism arises mainly from consideration of

the tropical species.

Subgen. Oudemansiella is separated from subgen. Xerula by reason of habitat,

gelatinisation of the pellicle on the pileus, and manner of development. The

habitat is given as lignicolous versus radicicolous but, as roots are also lignified,

it is a distinction between growing on stem-wood rather than root-wood. This

distinction holds in the main, as with Amauroderma contrasted with Ganoderma,

but there may be anomalies such as that concerning O. brunneimarginata

mentioned under O. raphanipes. Imai (1938) reported O. radicata on decayed

wood above ground in Hokkaido. With this distinction there is coupled the

presence of the pseudorhiza, said to be peculiar to subgen. Xerula, but there is O.

radicata f. arrhiza Lange (1936) as well as the state of the species on stem-wood

without the pseudorhiza, and in subgen. Oudemansiella there is O. canarii f.

radicans that grows on wood above ground. The pseudorhiza is a facultative

rhizomorph indicated, probably, by the basal plug by which all the fruit-bodies

are attached to stem-wood. Concerning the gelatinisation of the pellicle, it occurs

in both subgenera and is absent from some species in both, as in O. lianicola,

which could be placed in subgen. Oudemansiella because it grows on stem-wood

without a pseudorhiza.

Development of the fruit-body is described as bivelangiocarpic in subgen.

Oudemansiella, according to Reijnders (1948) and as gymnocarpic in subgen.

Xerula. The term bivelangiocarpic in this context is misleading because it implies

a universal veil and a partial veil and, at most, so far as I know, there is only a

partial or marginal veil in the genus. Indeed, Reijnders (1952) called O. canarii,

which is supposed to be conspecific with the type of the genus O. platensis,

metavelangiocarpic because what slight veil it has is formed by the union of

outgrowths from the margin of the pileus and the base of the stem (Corner, 1934).

Supposedly, the palisade on the pileus and the loose hyphal outgrowth on the

stem are taken to be the universal veil, but I regard them as just the surface of

pileus and stem in a gymnocarpic primordium; to regard them as adnate veils, as

velangiocarpic implies, is like regarding the hair on one’s head as a veil. This

marginal veil certainly supplies the ring on the stem of O. mucida and that, not

always present, of O. submucida. In O. canarii it is vestigial, and there may be a

trace of it in O. radicata of subgen. Xerula according to Reijnders (1952), as in

O. lianicola, but with no vestige in the expanded fruit-body. In fact, the presence

or absence of this marginal veil can be proved only by microscopic study of very

young primordia and it is not clear that this has been accomplished. It is

unreasonable to ascribe two veils to O. mucida and none to O. radicata when

both have the same kind of structure for the surface of the pileus. Thus, I regard

Oudemansiella as gymnocarpic with or without a marginal veil.

Surface-structure of the Pileus

This matter seems to me to supply the most important means for specific

arrangement in the genus. I distinguish five states, as follows.

1. The double palisade, with an outer palisade separated from an inner palisade

by a thick gelatinous layer.

A. The outer palisade developing moniliform rows of cells (mainly by

secondary septation) to form a trichoderm, e.g. O. platensis (subgen.

Oudemansiella).

B. The outer palisade remaining a single layer, e.g. O. canarii (subgen.

Oudemansiella).

2. A single palisade, with or without a thin mucilaginous hypodermis.

A. The palisade developing moniliform rows of cells (mainly by secondary

septation), e.g. O. lianicola (subgen. Oudemansiella in habitat, subgen.

Xerula in lacking a gelatinous layer).

B. The palisade remaining a single layer of clavate cells, with or without

ventricose-filiform pileocystidia. e.g. O. mucida and O submucida (subgen.

Oudemansiella) and probably most species of subgen. Xerula.

3. No palisade but a fairly thick gelatinous layer, e.g. O. crassifolia (subgen.

Oudemansiella in habitat).

Nevertheless, it is not clear that all these differences are absolute. Thus, in O.

altissima, the structure of the central part of the pileus places it in Group 2A and

that of the outer part in Group 2B (Corner, 1934, figures 8, 9). Then, the lack of

palisades in Group 3 may not hold for the unexpanded primordial pileus which

may have a slight palisade of clavate cells soon to become disrupted. Indeed,

disruption of the outer palisade on expansion in pilei of Groups 1A and 1B causes

the flecks on the gelatinous surface which are easily washed off by rain so as to

give the appearance of Group 3 but with the inner palisade intact. Thus, in O.

platensis and O. canarii, the true construction is not usually evident in mature

fruit-bodies.

Figure 1. Oudemansiella altissima (left) and O. radicata (right). Surface of central

part of pileus, x 500.

It emerges that O. mucida and O. submucida have the structure prevalent in

subgen. Xerula though, in respect of habitat and development, they agree with

subgen. Oudemansiella. The surface-structure cuts up the distinction between the

two subgenera.

Another issue concerns the presence of clamp-connections. They occur in all

the hyphae of the fruit-body but in most species that I have studied they are

absent from the hyphae of the one or two palisades. They are described as present

in the palisade for X. furfuracea, X. megalospora and X. rubrobrunnescens by

Redhead, Ginns and Shoemaker (1987), and some of the hyphae of the palisade

in O. lianicola are clamped. Whether the absence of clamps reflects an antecedent

state with moniliform secondary septation to form a trichoderm is not clear, but I

certainly consider that the trichoderm of O. platensis is antecedent to the single

outer palisade of O. canarii, just as the trichoderm resolves into the hymenioderm

in other basidiomycetes.

Actually, what appears as a single palisade in O. altissima, perhaps also in X.

caussei and X. kuehneri according to Boekhout and Bas, is more complex and, in

its way, triple (Corner, 1934, figure 8). There is the main layer of large clavate

cells beyond which project the ventricose-filiform pileocystidia, and between the

stalks of these cells there are shorter, narrower, clavate or cylindric cells with

brown sap. From this construction, there emerge the other main kinds. In the

simplest case, the palisade reduces to a truly single layer of cells. That of O.

canarii adds a gelatinous layer between the small, inner and brown cells and the

others, and in O. platensis this is complicated by secondary septation of the

hyphal ends in the outer palisade.

The Stem

In O. mucida and O. submucida the ring lies at or above the middle of the

stem but the vestige of the ring or marginal veil is on the upper side of the

bulbous base of the stem in O. canarii and O. platensis. The explanation is

provided by O. canarii var. perstipitata. The stem of O. canarii corresponds with

the supra-annular part of the stem of O. mucida (Corner, 1934, figure 11). That

illustration showed how the aerial stem of species with a pseudorhiza may

similarly correspond with the supra-annular part of O. mucida. Thus, it may be

that the primordia of the fruit-bodies of subgen. Xerula are formed at the surface

of the ground in a button-stage similar to that of O. canarii with or without a

trace of marginal veil. The idea is supported by the fact that the pileus is not

acutely conical in Oudemansiella, for such is usually the case with pilei initiated

below ground and forced through the soil to the surface. However, Buller (1931)

found the primordia of O. radicata were formed on the root and were pushed by

the pseudorhiza to the surface of the ground. I think that the problem still needs

investigation.

Basidia and Pleurocystidia

In 1947 I showed that both the mature basidia and the pleurocystidia of

Oudemansiella conformed to the same locus on the basidiograph that had for its

definition //w = 1.7 + 0.02/, giving a maximum width of 50 um. More data have

come to hand and the revised basidiograph is shown in Graph 1, along with the

original locus. I see no reason to alter its slope though an upward shift in the

value of the ordinates to 1.8 or 1.9 might assimilate more points. They, however,

are species-points, based on mean values, whereas my original data were based

on averages; thus, a basidium not fully expanded will give a higher value for //w

than one that is, and it is very easy to be deceived by this detail in a specific

description. Moreover, as the basidia in some species are dimorphic, more detailed

enquiry is needed to discover if there is any difference in this matter between the

long and short basidia. However, Buller gave the basidia of O. radicata as

monomorphic. For the initial stage of enlargement of the basidium up to 27 um

long and before it began to be charged with dense cytoplasm, the locus was //w =

1 + 0.11/, and this corresponds with that of the narrow basidia of Hohenbuehelia

and Pleurotus (Corner).

20 4o &Q %0 100 120 1h0 160 Iga 200 220 240

Graph 1. Oudemansiella, basidograph (triangles) and pleurocystidiograph (circles),

based on the data in this article (solid triangles and circles) and on those from

the authors cited. 1947, the locus based on average data (Corner, 1947). y,

the basidiograph for developing basidia.

The presence of subacerose basidioles in the hymenium of the fungus here

described as O. radicata var. hygrophoroides necessitates closer study because,

in this respect, it connects with Xerula based on X. longipes (Corner, in ed.). If its

species-point on the basidiograph is reliable, then its basidia are tending, like

those of X. longipes, to the narrow basidia of Marasmius, but its pleurocystidia

are typically those of Oudemansiella. It seems that the subacerose basidiole,

comparable in size with the developing and uncharged basidia of Oudemansiella,

have supplied the steeper locus for the basidia and pleurocystidia of Marasmius,

where //w = 1 = 0.21/ to | = 0.0711 with maxima as 5 um and 14.1 um respectively.

I note that the flattened or subtruncate apex of the pleurocystidium in some

species may be the effect of pressure upon the adjacent gill in the crowded state

of the primordial pileus, as seen in Coprinus and Pluteus.

Monomitic or Sarcodimitic

The hyphal construction of the fruit-body in Oudemansiella has always seemed

to me to be monomitic with long-celled inflating hyphae together with relatively

few uninflated hyphae. Thus, the texture is fleshy and the stem solid. Recently,

however, it has been proposed by Boekhout and Bas (1986) and by Redhead

(1987) that the stem and gills are sarcodimitic, as I described for Trogia. I have

discussed this matter in a second account of Trogia (Corner, 1991) and prefer my

original conclusion. The waxy-cartilaginous consistency and hollow stem of

Trogia are not features of Oudemansiella and, if the long cells of this and some

other genera of agarics may be the precursors of the inflated fusiform skeletal

cells of Trogia, they are not specially differentiated.

Affinity of Oudemansiella

It is generally agreed that Oudemansiella should be referred to Marasmiaceae

(Romagnesi, 1977) or Tricholomataceae tr. Marasmieae (Singer, 1975). It is put

in an allied family Xerulaceae by Jiilich (1981), which covers Oudemansiella,

Xerula and Lampteromyces. This last genus is put in Tricolomataceae tr.

Clitocybeae by Singer but, as I have shown (Corner, 1981), it is impossible to

distinguish clearly Lampteromyces from Pleurotus, and the custom now is to put

Pleurotus far away in or near Polyporaceae, though it lacks entirely the peculiar

hyphal construction of its key-genera Polyporus s. str. and Lentinus s. str. (Corner,

1984).

The idea of Xerulaceae has been greatly enlarged by Redhead (1987) who

considers it to be essentially a sarcodimitic family that ranges into the monomitic.

Thus, he places in it Trogia, Oudemansiella, Xerula and Mycena, but not

Lampteromyces or Marasmius. If the sarcodimitic origin is so essential, then the

polyporoid Meripilus must be added or given an allied family, though polyporoid

allies of Mycena are, apparently, not to be excluded. However, as already

mentioned, I am far from convinced of this aggregation, and the affinity of

Xerula with Marasmius is confirmation.

Xerula is based on X. longipes or, as it is now called X. pudens. It has the

marasmioid dry pileus, subagglutinated surface to the stem, acerose basidioles,

and thick-walled caulo- and pileo-cystidia, but it has, also, the fleshier texture,

large guttate spores, long-celled stem-hyphae, and the lack of clamps from the

palisades of stem and pileus as in Oudemansiella. In size of basidia and

pleurocystidia it comes between Marasmius and Oudemansiella. While this

confirms the alliance of Oudemansiella with Marasmiaceae, it still leaves the

fleshy consistency and lack of acerose basidioles as distinctive. This last point is

bridged by the fungus here described as O. radicata v hygrophoroides. It joins

Oudemansiella with Xerula which joins with Marasmius, without the intrusion of

Trogia. Lampteromyces links Pleurotus and Oudemansiella. Here is a range from

lignicolous on root and stem to humicolous and foliicolous fungi, and from

typically agaricoid to stereoid, in the exploitation of the saprophytic habitats of

the forest, parallel with the alliance of Trogia.

Here is confusion in the modern classification of Agaricales, which I do not

attempt to amend while there are still so many gaps in knowledge and so much

more to be discovered in the tropical flora.

Since this paper was written, a European fungus, first described as Hydropus

mediterraneus Pacioni et Lalli (1985), has been transferred to Flammulina by

Bas and Robich who give a detailed description (Persoonia 13, 1988, 489), and

most recently to Oudemansiella by Ortega, Vizoso and Contu (Documents

mycologiques f. 82, 1991, 25). This genus was rejected by Bas and Robich on the

ground that Redhead had decided that Oudemansiella was sarcodimitic, which I

regard as a mistake. The species seems to come between Xerula and

Oudemansiella. If it can be fitted into Flammulina, here is another instance of

superfluous genera.

Key to the Malesian species of Oudemansiella

1. Terricolous with rooting base from buried wood. Pileus dry. Stem fuscous brown scurfy fibrillose ..........

nplnsds aha ewaauwaaiies «sivctipaybep-oucans aa des des dunes pice ah Reatekapmatba leas do in Seieg yond sineae tn lSaa i ie mae O. altissima

1. Lignicolous above ground.

2. Pileus at first with a gelatinous pellicle 0.3-1 mm thick, with white to greyish flecks washing off with

rain, dark umber, paler to whitish on expansion. COMMON .............::cssccesseesseeeeeeesneeeeneeeneees O. canarii

2. Pileus dry or with a much thinner gelatinous pellicle, without superficial flecks.

3. Stem with a membranous ring. Pileus smeary viscid, white to pale ochraceous. Borneo...............

gis carenadhansvesteandpansiatn cuban nanan ss Odigt st tea eaten «aad ch paleo aici deta ie ease a O. submucida

(pilots GEy™ ..6...0.i\yccsebeks as detnn bgt Me etree kbd acis ss etanaceds tse e cau ses eee var. persicca)

3. Without such a ring.

4. Gills very thick, obtuse. Pileus -3 cm wide, with a thin viscid pellicle, white. Spores 21-28 x

19-24 um, Bormeo like see eeecnaccreeccaess cvs hota shs tacaks cl eer te eae eee O. crassifolia

(pileus -6 cm wide, dry, pale pink as the stem); spores 19-24 tum. Malaya ....... v. incarnata)

4. Gills rather thin. Pileus -3 cm wide, dry, pallid ochraceous with subferruginous and strongly

reticulate centre. Spores 10.5-13 x 8.5-9.5 tum. On dead lianes. Borneo ........... O. lianicola

Oudemansiella altissima (Mass.) comb. nov. Figures 1, 2

Collybia altissima Mass., Kew Bull. 1914, 358.- C. radicata, Corner in Trans. Brit. mycol. Soc. 19 (1934)

64, f. 8-10.

Pileus 1-12 cm wide, convex then plane or concave, often subumbonate, dry

(slightly viscid in decay), smooth or more or less extensively regulose-reticulate

or rugulose rivulose, pale fuscous brownish or pale umber with darker centre;

margin substriate, often slightly sulcate-crenulate. Stem 6-14 cm x 3-8 mm at the

apex, 5-16 mm at ground level, with tapering root -16 cm long arising from a

slender white rhizomorph 0.5 mm wide, fibrous, dry, concolorous, wholly fuscous

brownish scurfy pruinose or subfibrillose but the white apex pruinose; veil none.

Gills sinuate to adnexo-adnate, subdistant, rather thick and waxy, 13-33 primaries

4-12 mm wide, 3-5 ranks, not veined, white, sometimes with umber brown edge.

Flesh 3-7 mm thick in the centre of the pileus, sappy, without a gelatinous layer,

white. Smell sour.

On the ground, solitary or occasionally 2-3 together, from buried roots in the

forest. Malesia to the Solomon Islands, uncommon, lowland to 1700 mm alt.

my |

Figure 2. Oudemansiella altissima. Surface of lower part of stem, x 500.

Spores 15-19 x 12-15 um, in some collections 11.5-15.5 x 11-14 um, white,

smooth, subglobose, the wall 0.3 um thick, multiguttulate. Basidia 58-75 x 13-18

uum; sterigmata (2-)4, 6-10 x 3-4 um, 9-13 x 4-6 um on 2-spored basidia and

sometimes with an abortive third sterigma; no acerose basidioles; subhymenium

not corticate. Cheilocystidia as in O. canarii, clavate to ventricose and shortly

appendaged, in some collections with brown sap. Pleurocystidia 80-190 x 18-50

um, clavate to ventricose with a long obtuse appendage as in O. canarii, thin-

walled. Hyphae clamped in both 2- and 4-spored fruit-bodies, but without clamps

in the palisades on pileus and stem; in the stem strictly longitudinal with firm

walls, the cells 100-1200 x 5-38 um, cylindric with broad septa or with tapered

ends, with few uninflated interweaving hyphae; oleiferous hyphae 3-9 um wide,

scattered. Surface of stem with loosely appressed and entangled longitudinal

hyphae 3-9 um wide, without clamps, with brown sap, ending with clavate to

ventricose, often secondarily septate, cells 25-75 x 8-25 um with brown sap,

compacted into clusters towards the stem-apex, developing filiform processes

1.5-3 um wide in the lower part of the stem (as on the pileus), these processes

often bifid in 2-spored fruit-bodies. Surface of the pileus covered over the centre

with a compact palisade with 3 kinds of cell (with intermediates); 1, narrowly

ventricose colourless cells with the cell-body 30-60 x 4-12 um prolonged into a

filiform unbranched, aseptate, tapering process 70-350 x 1.5-3.5 um, rarely 1-

septate, eventually mucifying in old fruit-bodies; 2, large clavate or ventricose

hyaline cells 40-70 x 8-20 um, without a process; 3, small clavate or subcylindric

cells 15-40 x 5-12 Um with brown sap, packed between the stalks of the other

cells and forming the dense inner part of the palisade, without a gelatinous

substratum. Surface of the pileus over the limb with a palisade consisting mostly

of large clavate cells with brown sap, the subterminal cells often inflated with

brown sap, with a thin gelatinous substratum.

This fungus, which I called C. radicata for convenience in 1934, is C. altissima

Mass, according to the unmistakeable illustration of the type-collection E.M.

Burkill 112 and the specimen in the Singapore herbarium. It comes in the key of

Pegler and Young (1986) to sect. Albotomentosae where it should come next to

the otherwise very dissimilar O. xeruloides. A much closer species seems to be

O. japonica (sect. Radicatae) of which v. colensoi may be identical. Once, when

I was delayed in the town of Goiania in Brazil in January 1968, I found two fruit

bodies in different places which seemed identical with O. altissima, but I had no

means of making a herbarium specimen.

Oudemansiella canarii (Jungh.) v. Hoehn. Figures 3, 7

Pegler and Young (1986) 589.

Collybia apalosarca Berk. et Br.; Corner, Trans. Brit. mycol. Soc. 19 (1934) 39-88.

Pileus 4 mm-15 cm wide, convex then plane, often gibbous, never revolute,

sometimes regulose, with a smeary viscid gelatinous pellicle at first spotted with

greyish flecks, striate in small specimens, dark umber becoming paler fuscous

brown to dirty white when old, rarely pallid white from the first; margin slightly

incurved at first, thin, acute, entire. Stem 3 mm-9 cm x 0.5-9 mm at the apex, I-

14 mm at the base, gradually tapered upwards or subcylindric, central to slightly

excentric, straight or incurved, thinly pruinose or subvillous, fibrous, solid, apex

often subcostate, white or, when young, pale clear yellow, sometimes with pale

rufous streaks; base dilated, generally with a narrow ridge or zone round the

upper margin, attached by a microscopic root; veil marginal, slight, evanescent.

Gills rounded adnate, adnexed or nearly free, often separating free, sometimes

with a subdecurrent tooth, more or less ventricose, rather crowded, often broad,

thick, waxy to submucilaginous, not veined, 11-39 primaries 1-15 mm wide. 1-5

ranks, white, often with the edge pale clear yellow at first, becoming powdered

with the spores and brownish in decay. Flesh 0.5-5 mm thick in the centre of the

pileus, 0.3-3 mm thick halfway to the margin, at first firm and fleshy then rather

cottony spongy, white, unchanging, with a gelatinous layer 0.3-1 mm thick next

the surface of the unexpanded pileus. Smell of fresh fish, sometimes strong.

On dead trunks and branches in the forest and in the open, common.

Palaeotropics.

Figure 3. Oudemansiella canarii. Hymenium with pleurocystidium and multigittulate

basidia, x 500.

Spores 16-23 x 16-22 um, white in the mass, smooth, subglobose to broadly

ellipsoid, wall -0.5 um thick, densely granular guttulate. Basidia 54-75 x 18-25

um, monomorphic, densely multiguttulate; sterigmata 4, 5-9 x 3.5-5 um at the

base, stout, arcuate; acerose basidioles none. Cheilocystidia 45-95 x 8-28 um,

mostly 10-15 um, 2.5-4 um at the base, clavate, rarely subventricose, often with a

long stalk, thin-walled, vacuolate, colourless or in some specimens with yellowish

oily masses as exudates. Pleurocystidia 110-280 x 23-40 um, 10-16 um at the

apex, 3-4.5 um at the base, very large, broadly fusiform with obtuse apex to

ventricose with prolonged subcylindric apex, thin-walled, vacuolate, colourless.

Caulocystidia as thin-walled terminal cells 6-17 um wide with finely vacuolate-

reticulate colourless cytoplasm, on lax superficial hyphae, in some specimens

with yellowish oily masses as on the gill-edge; basal disc with a lax palisade of

uninflated hyphae but the terminal cells 5-8 lum wide and often thinly encrusted

with yellowish granules. Surface of the pileus pelliculose with two palisade

layers separated by the gelatinous layer; outer palisade consisting of rather laxly

arranged, more or less ventricose-appendaged, thin-walled cells 70-200 x 6-11

um, mostly with pale umber sap, the hypodermic of laxly interwoven hyphae

with cells 35-200 x 2-5(-12) um, this palisade splitting into the greyish flecks on

the pileus; inner palisade c. 100 um high, composed of narrow subcylindric

hyphae with cells 25-50 x 2.3-5 um, in places -6 Um wide, with pale fuscous

umber sap. Hyphae clamped except those of the palisades on pileus and stem and

of the gelatinous layer of the pileus; in the pileus with cells 60-500 x 8-30 um and

some 3-5 um uninflated; in the stem with cells 100-1200 x 8-30 um, a few

uninflated, at the base of the stem 5-12 uum wide and densely interwoven, in the

basal plug 3-5 um wide with slightly thickened walls and very densely interwoven.

forma radicans Corner (1934)

Stem with a rooting base as a pseudorhiza one to several cm long, from the

rotten wood. Frequent with typical specimens in Malaya.

var. perstipitate Corner (1934)

Stem without a basal disc, the basal part elongating and bearing an indistinct

zone or slight ring (as the remains of the marginal veil) about the middle.

Fruit-bodies small, with pileus -5 cm wide, stem -6 cm long. Much less

common than the typical state in Malaya. .

This description is condensed from the extended account that I gave in 1934.

It is the commonest species of the genus in Malesia, and I do not regard it as the

same as O. platensis of South America for the reasons given under that species.

Both are placed in subgen. Oudemansiella because of the lack of pseudorhiza but

f. radicans belies this distinction from subgen. Xerula. The veil in both species is

so slight that, to be sure of its presence, it is necessary to section the unexpanded

primordium.

There occurs on fallen branches in Tjibodas Botanic Garden, Java, a fungus

which I class temporarily with f. radicans but it requires fuller investigation. The

fruit-bodies are rather small with a pinkish to rufous tan pileus and a rather strong

sour smell. It agrees with O. canarii microscopically except that the outer palisade

of the pileus consists of rows of 2-3 or more inflated cells with the terminal

cell clavate, in the manner of O. lianicola.

Oudemansiella crassifolia sp. nov. Figure 4

Receptacula alba. Pileus 1.2-2.5 cm latus, convexus dein planus, viscidus laevis. Stipes 12-50 x 2.5-

3.5 mm, basim abruptum vel subattenuatum versus 2.5-5 mm latus, subfloccosus, apicem versus pruinosus;

velo annuloque non viso. Lamellae adnatae rotundatae, subconfertae dein subdistantes, crassae obtusae,

acie saepe undulata, 14-28 primariae 2-3 mm latae, ordinibus 2-3. Caro tenuis, sub pilei superficie

tenuiter gelatinosa. Sporae 21-28 x 19-24 Lum, subglobosae laeves multiguttulatae. Basidia 70-130 x 25-28

um. Cheilocystidia 55-130 x 13-38 Um, ut in O. canarii. Pleurocystidia 105-165 x 30-50 Lim, clavata vel

ventricosa, copiosa. Superficies pilei strato gelatinoso 300-500 Lum crasso, hyphis efibulatis 2-5 Lum latis

pervaso, haud vallato, hypharum apicibus decumbentibus 3-8 [im latis vel in cellulis sparsis -35 x 16 [im

expansis. Ad lignum emortuum ramulisque solitaria in silva montana. Borneo, Mt. Kinabalu 1700m alt.

Typus RSNB 5483B; herb. Corner.

var. incarnata var. nov.

Differt pileo stipiteque primo pallide incarnato; sporis minoribus 19-24 Lim; cheilocystidiis brevioribus

40-65 x 10-38 Um; pilei superficie strato gelatinoso tenuiori 100-150 Lim crasso. Ad truncos emortuos

Mangiferae et Pini in silva montana. Malaya, Pahang, Cameron Highland 1500m alt. Typus, Corner s.n.

1 Oct. 1966; herb. Corner.

Fruit-bodies entirely white. Pileus 1.2-2.5 cm wide, convex then plane, smooth,

with a viscid pellicle. Stem 1.2-5 cm x 2.5-3.5 mm above, 2.5-5 mm below,

pruinose upwards, fibrillose and subfloccose downwards to the abrupt or

subattenuate base, not discoid; veil and ring none. Gills rounded adnate, rather

crowded then subdistant, with a very thick obtuse and often undulate edge, 14-28

primaries 2-3 mm wide, 2-3 ranks. Flesh thin, with a thin gelatinous layer on the

upper side.

On dead wood and sticks in the forest, solitary. Borneo, Mt Kinabalu 1700m

alt.

Figure 4. Oudemansiella crassifolia. Spore, x 1000. Pleurocystidia and surface of

pileus, x 500.

Spores 21-28 x 19-24 um, subglobose, smooth, multiguttulate. Basidia 70-130

x 25-28 um, 4-spored; subhymenium -30 um thick, composed of 2-3 um hyphae,

interwoven, not corticate. Cheilocystidia 55-130 x 13-38 um, as in O. canarii,

forming the very thick gill-edge and widest in the transitional zone to the

hymenium. Pleurocystidia 105-165 x 30-50 um, clavate or ventricose with obtuse

or scarcely prolonged apex, stalk narrow, abundant, strongly projecting. Hyphae

clamped except in the mucilage on the pileus; stem-cells 80-600 x 5-20 um,

cylindric or with tapered ends. Surface of stem with hyphae 3-6 um wide, here

and there entangled into fibrillose flocci, with scattered subclavate ends 5-9 um

wide. Surface of pileus with a gelatinous layer 300-500 um thick, pervaded by 2-

5 um hyphae without clamps, branching but without forming any palisade, the

hyphal ends at the surface more or less shortly decumbent on the mucilage with

cylindric to subventricose ends 3-8 um wide, also with a few scattered clavate

cells -35 x 16 um.

Collections.- Borneo, Mt Kinabalu, Bembangan and Mesilau valleys, 1700m

alt.; RSNB 5060, 22 Jan. 1964; RSNB 5483A, 26 Feb. 1964; RSNB 5483B, 12

March 1964; RSNB 5483C, 16 March 1964.

var. incarnata

Pileus -6 cm wide, convex to plane, smooth, dry, opaque, pale pink, fading

white from the centre outwards. Stem 1-1.5 cm x 2-3.5 mm, 3-4.5 mm at the

abrupt base, subcylindric, thinly white villous downwards, pale pink then white.

Gills adnate, separating free, thick, waxy, subdistant, 14-17 primaries -5 mm

wide, 3 ranks, white. Flesh 2-3 mm thick in the centre of the pileus, floccose-

tough, with a thin gelatinous layer at the surface, white. Smell rather sour and of

cucumber.

On fallen trunks of Mangifera odorata and Pinus sp. Malaya, Pahang, Cameron

Highlands, 1500m alt.

Spores 19-24 um, subglobose. Basidia 60-88 x 20-24 um; sterigmata 2-4.

Cheilocystidia 40-65 x 10-38 um, clavate to ventricose, obtuse, not appendaged,

thin-walled, smooth, as a thick sterile edge of the gill, often with the subterminal

cell or two inflated and ellipsoid to subglobose. Pleurocystidia 120-220 x 22-40

um as in O. canarii, often slightly waisted, wall 0.5 um thick. Structure as 1n var.

crassifolia but the gelatinous layer on the pileus 100-150 um thick with 3-6(-7)

uum hyphase often disarticulating.

The fruit-bodies of this species seem characteristically small and solitary. The

pileus has the gelatinous layer on the surface as in O. canarii but without distinct

palisades unless the primordial pileus may have a single layer of clavate cells.

The very thick gills with undulate edge are characteristic. I saw no sign of veil or

annulus.

Oudemansiella lianicola sp. nov. Figures 5, 6

Pileus -28 mm latus, convexus siccus, primo furfuraceus subvillosus, in centre valde ruguloso-

reticulatus, pallide ochraceus centro subferrugineo. Stipes 30-40 x 3-4 mm, ad basim abruptum

subbulbosum 5-7 mm, fistulosus, sursum albidus pruinosus, deorsum appresse fibrillosi-subfloccosus

brunneus. Velum marginale vix evolutum, annulo nullo. Lamellae adnatae rotundatae subdistantes,

haud vel vix venosae, 19-23 primariae 3-5 mm latae, ordinibus 3, albidae dein pallide ochraceae. Caro 3-

6 mm crassa in pilei centro, firma, haud gelatinosa, flavidi-albidula. Sporae 10.5-13 x 8.5-9.5 Lim, laeves,

ellipsoideae multiguttulatae. Basidia 46-58 x 11-12.5 Lim. Cheilocystidia 40-100 x 12-28 Lum, clavate vel

subfusiformia, ut acie lata sterili. Pleurocystidia 50-85 x 18-38 Lim, clavate, nonnulla subtruncata, raro

submucronata, copiosa. Superficies pilei e cellulis clavatis 70-120 x 15-40 Lum, tunicis brunneis, et cellulis

25-70 x 8-38 Lum subglobosis, tunicis brunneolis, in seriebus moniliformibus -300 Lim longis instructa,

sine fibulis. Ad lianos emortuos gregaria vel subcaespitosa, in silva montana. Borneo, Mt. Kinabalu

1400m alt. Typus, RSNB 5006; herb. Corner.

Pileus -28 mm wide (not fully expanded), convex, dry, at first furfuraceus

subvillous, strongly rugulose reticulate in the centre, minutely furfuraceous towards

the substriate incurved margin, pallid bistre ochraceous or tan ochraceous, brownish

towards the subferruginous centre. Stem 3-4 cm x 3-4 mm 5-7 mm at the abrupt

subbulbous base, slightly attenuate upwards, hollow, fibrillose, whitish and

subpruinose upwards, fawn brown and appressedly fibrillose-floccose or

subfurfuraceous downwards, the base madder brown subvillous. Veil marginal,

Pret nS

Fete toe tng

Figure 5. Oudemansiella lianicola. Fruit-bodies in section, x 2. Spores, x 1000.

Pleurocystidia (above) and cheilocystidia (below), x 500.

Figure 6. Oudemansiella lianicola. Surface of pileus, with several large clavate

cells of the orighinal palisade and moniliform cell-rows of incursive

hyphae, x 500.

very slight, fibrillar, leaving no trace on the stem. Gills rounded adnate, separating

free, subdistant, not or scarcely veined, 19-23 primaries 3-5 mm wide, 3 ranks,

cream white to pallid subochraceous. Flesh 3-6 mm thick in the centre of the

pileus, rather firm, homogeneous, without a gelatinous layer, pallid yellowish

white. Smell slight, subacrid.

On a dead hanging liane, gregarious to subcaespitose. Borneo, Mt Kinabalu,

Mesilau 1400m alt, 18 Jan. 1964.

Spores 10.5-13 x 8.5-9.6 um, white, smooth, ellipsoid, multiguttulate,

inamyloid. Basidia 46-58 x 11-12.5 um, 4-spored; no acerose basidioles;

subhymenium narrow, composed of 2-4 um hyphae, not corticate, becoming

submucilaginous. Cheilocystidia 40-100 x 12-28 um, clavate to subfusiform with

obtuse apex, rarely shortly appendaged, thin-walled, smooth, as a broad sterile

gill-edge. Pleurocystidia 50-85 x 18-38 um, clavate, mostly subtruncate, not

appendaged or occasionally submucronate, thin-walled smooth, very abundant,

(the contents contracting in alcohol-formalin into a granular mass with fine

membrane). Hyphae clamped; in the stem, with cells 70-500 x 5-36 Um or -50 Um

wide and then often in moniliform rows at the base of the stem, septa broad and

transverse to oblique. Surface of stem with loosely interwoven, longitudinal

hyphae 3-12 tum wide, short-celled, with pale brown walls, the end-cells subclavate

or subfusiform as on the pileus, these slight caulocystidia here and there loosely

clustered into the furfuraceous particles especially towards the base of the stem

with a loose tomentum of hyphae c. 200 lum thick. Surface of pileus covered by a

fairly compact pile of hyphal ends 200-300 um high in the centre or the pileus,

consisting of large clavate cells 70-120 x 15-40 lm with dark brown walls and

more or less moniliform rows of ellipsoid, pyriform or subglobose cells 25-70 x

8-38 um with pale brown walls, with (and without) clamps; the young pileus

covered with the large clavate cells, then on expansion developing the moniliform

rows and causing the reticulation in the centre of the pileus; without a gelatinous

layer.

The habitat of this fungus seems particular because, if it grew on ordinary

wood fallen from trees, I would surely have found it in such places. The pileus

develops the trichoderm of Physocystidium, though it begins with the hymenioderm

as in O. platensis.

Oudemansiella mucida (Fr.) v. Hoehn. Figure 10

I have the following notes on English collections:—

Gills slightly veined at the base, 15-26 primaries 4-16 mm wide, 3-6

ranks. Flesh 3-10 mm thick in the centre of the pileus, 1-2 mm halfway to the

margin, gelatinous in the gill-trama and at the surface of the pileus. Spores

14-18.5 um, subglobose, with distinctly thickened wall. Cheilocystidia -95 x

7-12 um, subfusiform subacute to clavate, rarely subventricose, thin-walled.

Pleurocystidia none seen (collapsing), none (Ricken), -200 x 40 um (Boursier

1926), 90-140(-180) x 15-30(-40) um (Rea 1922). Surface of pileus with a

single palisade of clavate cells 18-35 x 5-10 um, occasionally obtusely

ventricose, rather small, compact, derived from gelatinous hyphae 1.5-2.5 um

wide without clamps (other hyphae with clamps).

I note the account of physiological factors in the development of the fruit-

body of this species by Semerdzieva and Musilek (1970).

Oudemansiella platensis Speg. Figure 7

It is now customary to regard this species, described and widely reported from

south and tropical America, as identical with O. canarii of the Old World. I have

seen these fungi commonly in Brazil and Malesia and I am not convinced that

they are conspecific even though the fruit-bodies are extremely similar. Thus, I

never gathered the Malesian fungus in South America or that in Malesia. They

PAN

{

’

Figure 7. Oudemansiella platensis. (left) and O. canarii (right). Outer palisade on

surface of pileus, x 500.

differ in the construction of the surface of the pileus which develops an almost

pseudoparenchymatous tissue in what I regard as O. platensis. Therefore, the

greyish flecks, which form from the disruption of this layer on the expansion of

the pileus, are thicker and more conspicuous in the American species. I have the

following details concerning O. platensis:-

Spores 15-25 um wide, subglobose, wall slightly thickened. Basidia 70-80 x

18-21 um. Cheilocystidia as in O. canarii. Pleurocystidia 170-280 x 30-46 um,

clavate to ventricose with a long obtuse appendage (as in O. canarii). Hyphae

clamped except in the surface tissue of pileus and stem, but some collections of

fruit-bodies apparently devoid of clamps. Surface of pileus with the double

palisade of O. canarii but the outer palisade developing into an almost

pseudoparenchymatous layer 200-300 tum thick, composed of moniliform rows

of ellipsoid to subglobes cells 30-100 x 5-70 um but smaller towards the surface

and with end-cells 10-30 x 5-8 tm, mainly secondarily septate, thin-walled,

colourless, this pellicle breaking into the flecks on the gelatinous layer; inner

palisade with many clavate cells -40 x 7-15(-20) um.

Collections from Niteroi, Chavantina and Manaus in Brazil; Rick 286, det. O.

platensis, herb. Mus. Nac. Rio de Janeiro.

The thick superficial layer develops hypodermally from hyphal branches arising

below the initial and simple outer palisade the cells of which persist among the

stalks of the moniliform rows. Likewise, hyphal branches grow from just below

the inner palisade into the gelatinous layer where they may also produce rows of

more or less inflated cells before continuing to the outer palisade.

This complicated surface relates with that of O. lianicola. There may be

another species in Amazonia resembling O. platensis but without gelatinous

layer to the pileus, which I collected near Manaus 17 October 1948.

O. radicata (Fr.) Singer Figure |

For this common species I gave the following notes from collections made

about Cambridge, England:-

Spores 14-16 x 10-11.5 um. Cheilocystidia 60-130 x 10-25 um, clavate to

ventricose, obtuse, as a broad sterile gill-edge. Pleurocystidia similar, ventricose,

obtuse, mostly capped by an oily brownish globule 10-28 um wide. Surface of

pileus with a single compact palisade of elegantly clavate cells 30-90 x 8-17 um,

with slender stalks, walls -1 tm thick (at least in the stalks), seated on interwoven

submucilaginous hyphae 3-7 um wide, no thick gelatinous layer, no ventricose-

filiform pileocystidia. Stem hyphae with cells 250-700 x 5-45 um (Boekhout and

Bas), with broad transverse septa, clamped; some oleiferous hyphae 3-10 um

wide. Caulocystidia clavate, scattered.

Oudemansiella ? radicata var. hygrophoroides Figure 8.

(Singer et Clemencon) Pegler et Young (1986).

The following description refers to a fungus which I collected in northern

Japan and which certainly belongs to the complex of O. radicata in Hokkaido

(Imai, 1938). However, it presents two problems. First, the presence of long

pileocystidia as well as clavate cells in the palisade on the pileus, just as in the

Malesian O. altissima, raises the question whether it belongs in sect.

Figure 8. Oudemansiella? radicata var. hygrophoroides (from Japan). Spores, x 1000.

Cheilocystidia and pleurocystidium, x 500. Caulocystidia in a cluster, x 500.

Albotomentosae or sect. Radicatae. If referred to the former, it does not fit any

known species; if to the latter, it comes out as the European O. radicata var.

hygrophoroides. It has the spores of Xerula megalospora Redhead, Ginns et

Shoemaker but not its capitate cystidia. Second, there are scattered among the

immature clavate basidia fusiform to subacerose basidioles which are one of the

marasmioid features of Xerula longipes. | have never seen such basidioles in

other species of Oudemansiella, to which genus all other features of its fruit-body

conform.

Fruit-bodies as those of O. radicata with viscid regulose pileus 2-10 cm wide;

stem finely brown scurfy from minute clusters of hairs. Rooting in the forest.

Hokkaido, pr. Yamabe, 18 Sept. 1966.

Spores 16-19.5 x 9.5-11.5 tum, mango-shaped, even subapiculate, smooth,

rather thin-walled, oily opalescent (alc. formalin material). Basidia 46-58 x 10-13

tum; sterigmata (3-)4, 7.5-9.5 um long; acerose basidioles scattered among the

young basidia. Cheilocystida 60-120 x 14-26 um, ventricose, acute, not

appendaged, thin-walled, but in some cases with the apex thinly oily-granular

encrusted, as a sterile gill-edge. Pleurocystidia 90-160 x 18-28 um, ventricose

with prolonged subcylindric obtuse appendage 7-16 um wide, more or less thickly

oily-encrusted over the appendage, often with a brown oily resinous apical globule,

thin-walled. Hyphae clamped, in the stem -14 um wide and long-celled. Surface

of stem with clusters of subcylindric or gradually tapering caulocystidia 40-220 x

6-13 um, simple, aseptate, with smooth and firm or slightly thickened walls, the

obtuse apex 3-6 Lim wide, arising in clusters from the ends of 1-3 hyphae. Surface

of the pileus with a palisade of clavate cells 40-70 x 8-20 um, thin-walled or the

stalks slightly thick-walled and apparently buried in mucilage, also many

pileocystidia similar to the caulocystidia but narrower, 5-8 tum with the apex 3-5

um, rarely -16 Um.

Oudemansiella raphanipes (Berk.) Pegler et Young (1986)

O. brunneomarginata L. Vassiljeva; Endo and Hongo, Trans. mycol. Soc. Japan 17 (1976) 345, f.1;

Yokoyama, Fungi of Japan (1989) 121; Imazeki and Hongo, Coloured Illustrations of mushrooms of

Japan (1987) f. 154.

Pegler and Young give O. brunneomarginata as a synonym of O. raphanipes

in sect. Radicatae of subgen. Xerula. This subgenus was defined in Pegler and

Young’s key as comprising fruit-bodies arising from a pseudorhiza growing from

buried roots. O. raphanipes was distinguished in their key by the minute reflexed

squamules on the stem. Yet, the Japanese fungus described and illustrated by the

Japanese authors, grows on dead trunks of Acer mono with a pseudorhiza evidently

penetrating the rotten wood, much as in O. canarii f. radicans, and it does not

appear to have the recurved squamules on the stem. Whatever the true identity of

the Japanese fungus may be, it transgresses the distinction in habit between the

two subgenera.

Oudemansiella steffenii (Rick) Singer Figure 9

Lilloa 26(1953) 66; Pegler and Young (1986) 599.

O. echinospora Singer, Mycologia 37 (1945) 439.

Pileus 3-13 cm wide, convex to plane, opaque, dry, very smooth or radially

rugulose towards the striate margin, fuscous fuliginous then fuscous ochraceous

to fuscous bistre over the limb. Stem 6-16 cm x 3-10 mm upwards, 4-20 mm at

ground level, white and fuliginous scurfy pruinose or fibrillose to more or less

entirely fuliginous; rooting base rather short or up to 15 cm long. Veil and ring

not formed. Gills adnexed, scarcely crowded, c. 25 primaries 4-11 mm wide, 3

ranks, white with pale fuscous edge. Flesh 3-10 mm thick in the centre of the

pileus, firmly fleshy, without a gelatinous layer.

On the ground in forest, mostly solitary. South America.

Figure 9. Oudemansiella steffenii. Spores, x 1000. Pleurocystidia (large) and

pileocystidia, x 500.

Spores 11-15 lum wide (spore-body), white, subglobose, varying rather sparsely

to closely verrucose to echinulate with conical, obtuse to acute, warts 1.5-3 x 1-

1.5 um, thin-walled, guttate, inamyloid. Basidia 65-80 x 18-24 um, evidently

dimorphic; sterigmata (2-3-)4. Cheilocystidia 40-75 x 15-30 um, clavate, thin-

walled, smooth, with fuscous sap, as a sterile gill-edge. Pleurocystidia 100-180 x

25-35 um, ventricose-fusiform, often with subcapitate to subtruncate apex 13-17

um wide, thin-walled, smooth. Caulocystidia as in O. canarii but with fuscous

sap as in many superficial hyphae of the stem. Surface of pileus with a single

palisade of clavate to ventricose cells 45-75 x 15-30 um, thin-walled, with

fuscous sap, arising from a firm layer of narrow, interwoven hyphae with fuscous

sap, without a gelatinous layer, without long pileocystidia. All hyphae in the

fruit-body with clamps.

These are my notes on the fungus as I found it, rather commonly, in Brazil.

Oudemansiella submucida sp. nov. Figure 10

Pileus 1-10 cm latus, convexus dein planus, glutinoso-pelliculosus laevis, albus dein subochraceus,

marginem versus substriatus. Stipes 1.5-7 cm x 1.5-10 mm, ad basim vix incrassatus, fibrosus

firmus siccus, primo subfloccoso-fibrillosus, albus dein subochraceus; annulo 2-6 mm lato, prope

stipitis apicem, membranaceo pendenti albo, etiam aliquando deficienti. Lamellae adnexae vel

adnatae, ventricosae subdistantes crassae ceraceae puberulae, haud venulosae, primariae 16-28 2-14

mm latae, ordinibus 3, albae. Caro 2-10 mm crassa in pilei centro, hygrophana, ad auperficiem

tenuiter glutinoso-pelliculosa, alba. Sporae 18-25 x 17.5-23 Lim, subglobosae laeves. Cheilocystidia et

pleurocystidia ut in O. canarii. Pellicula pilei e cellulis clavatis 25-35 x 15-25(-30) Lim, e strato

subgelatinoso angusto orientibus, instructa. Ad truncos delapsos in silva montana. Borneo,

Mt. Kinabalu c. 1700m alt. Typus, RSNB 5201; herb. Corner.

Figure 10. Oudemansiella mucida (left) and O. submucida (right).

Spores of pileus, x 500.

var. persicca var. nov.

Differt pileo sicco sine strato mucilaginoso, etiam pilei hymeniodermate laxe evoluto. Borneo,

Mt. Kinabalu, Mesilau 1700m alt., Aprile 1964. Typus, RSNB 5201A; herb. Corner.

Pileus 1-10 cm wide, convex then plane, smeary viscid, pelliculose, smooth,

without flecks of veil, white to pale butter ochraceous, substriate towards the

margin. Stem 1.5-7 cm x 1.5-10 mm, base slightly thickened, fibrous, firm, dry,

at first slightly floccose-fibrillose, white then pale butter ochraceous from below

upwards. Ring 2-6 mm wide, near the stem-apex, membranous, pendent, collapsing,

white, but some specimens without the ring. Gills adnexed to adnate, subdistant,

not veined, thick, waxy, puberulous, 16-28 primaries 2-14 mm wide, 3 ranks,

white. Flesh 2-10 mm thick in the centre of the pileus, hygrophanous, thinly

viscid-gelatinous at the surface but bounded by a pellicle, no conspicuous

gelatinous layer, white.

On fallen trunks in montane forest. Borneo, Mt. Kinabalu.

Spores 18-25 x 17.5-23 um, white, subglobose, smooth, wall slightly thickened.

Cheilocystidia and pleurocystidia as in O. canarii. Hyphae clamped except those

in the pellicle of the pileus; in the stem with cells 90-850 x 5-34 um, the longer

often tapered; in the pileus -45 um wide and rather spindle-shaped; in the ring 3-

8 um wide, unspecialised. Pellicle on the pileus composed of a fairly compact

palisade of smooth, more or less clavate, cells 25-35 x 15-25(-30) um, seated on a

narrow subgelatinous layer of hyphae 2-4 um wide and without clamps.

Collections.- Borneo, Mt Kinabalu, Tenompok, 8 Sept. 1961, RSNB 2890;

Mesilau 2 Feb. 1964, RSNB 5201.

This species comes between O. canarii and O. mucida, having the fruit-body

and surface structure of the pileus as in O. mucida and the large spores and

cystidia of O. canarii. It differs from O. mucida, also, in the dry stem and larger

clavate cells on the pileus. The dryness is emphasized by var. persicca.

References

Boekhout, T. and C. Bas (1986). Notulae ad floram agaricinam neerlandicam -

XII. Some notes on the genera Oudemansiella and Xerula. Persoonia 13: 45-

56.

Bon, M. (1975). Agaricales de la cote atlantique frangaise. Doc. Mycol. IV, fasc.

17: 1-40.

Boursier, M. (1926). Note sur le genre Mucidula Pat. Bull. Soc. Mycol. Fr. 40:

332-333.

Buller, A.H.R. (1924). Researches on Fungi III. Longmans, Green & Co., London,

New York, Toronto.

— (1931). Researches on Fungi IV. Longmans, Greens & Co., London, New

York, Toronto.

Corner, E.J.H. (1947). Variation in size and shape of spores, basidia and cystidia

in Basidiomycetes. New Phytologist 46: 195-228.

— (1981). The Agaric genera Lentinus, Panus and Pleurotus. Nova Hedwigia

Beih. 69.

— (1984). Ad Polyporaceas Il. Nova Hedwigia Beih. 78.

— (1991). Trogia (Basidiomycetes). Gard. Bull. Sing., Suppl. 2. Singapore

Corner, E.J.H. (1994). On Hohenbuehelia (Agaricales). Gard. Bull. Sing. 46(1):

1-31.

Imai, S. (1938). Studies on the Agaricaceae of Hokkaido, /. J. Fac. Agric.,

Hokkaido Imp. Univ. 43, pt. 1: p. 115.

Jiilich, W. (1981). Higher taxa of Basidiomycetes. Bibliotheca Mycologica 85.

Kobayasi, Y., Y. Otani and T. Hongo (1973). Some higher fungi found in New

Guinea. Rept. Tottori Mycol. Inst. (Japan) b. 10.

Lange, J.E. (1936), Flora Agaricina Danica, II: 9.

Pegler, D.N. (1986). Agaric flora of Sri Lanka. Kew Bull. Add. Ser. XII.

— and T.W.K. Young (1986). Classification of Oudemansiella (Basidiomycota:

Tricholomataceae). Trans. Brit. Mycol. Soc. 87: 583-602.

Rea, C. (1922). British Basidiomycetae. Cambridge University Press.

Redhead, S.A. (1987). The Xerulaceae (Basidiomycetes), a family with

sarcodimitic tissues. Can. J. Bot. 65: 1551-1562.

Redhead, S.A., J. Ginns and R.A. Shoemaker (1987). The Xerula (Collybia,

Oudemansiella) radicata complex in Canada. Mycotaxon 30: 357-405.

Reijnders, A.F.M. (1948). Etudes sur le developpement et |’ organisation

histologique des carpophores dans les Agaricales. Rec. Trav. Bot. Neerlandais

41: 213-396.

— (1952). Recherches sur le developpement des carpophores dans les Agaricales.

Verh. Kon. Nederl. Akad. Wetensch. Natuurkund., ser. 2, 48 n. 4; 1-116.

Romagnesi, H. (1940). Mycenella et Xerula. Bull. Soc. Mycol. Fr. 56: 61-65.

Romagnesi, H. (1977). Sur la multiplication excessive des genres en mycologie.

Bull. Soc. Mycol. Fr. 93: 248, 254.

Sathe, A.V. and J. Daniel (1980). Agaricales (Mushrooms) of South West India.

Maharashtra Association for the Cultivation of Sciences, Research Institute,

Monograph 1. Pune, India.

Semerdzieva, M. and V. Musilek (1970). Wachstum und Entwickelung des

Basidiomyzeten Oudemansiella mucida. Ceska Mykologia 24: 44-53.

Singer, R. (1975), The Agaricales in modern taxonomy. J. Cramer, Vaduz.

Plate 1.

Oudemansiella altissima. Specimens from Singapore.

On the Agaric Genera Hohenbuehelia and Oudemansiella

Acanthocystis

horridus

testudo

Agaricochaete

Amauroderma

Cantharellus

cystidioides

Chaetocalathus

Claudopus

griseus

Collybia

altissima

apalosarca

radicata

Conchomyces

bursaeformis

Crepidotus

Echinochaete

Faerberia

Ganoderma

Geopetalum

Hohenbuehelia

subg. Reidia

angustata

atrocaerulea

aurantiocystis

bullulifera

v. brasiliensis

bursaeformis

carbonaria

chevalieri

concentrica

culmicola

cystidioides

dimorphocystis

geogenia

griseipendens

v. nonsatisfacta

heterosporica

horakii

horrida

incarnata

lanceifera

longipes

malesiana

mellea

minutissima

myxotricha ©

nigra

pachyhyphata

v. minor

pahangensis

perstriata

petaloides

quadruplex

reniformis

repanda

rickenii

Index to Parts I and II

15, 34 roigil 2

15 singaporensis 1, 3, 9, 29, 41

34 spegazzini pe

7,8 subbarabtus 5

4,50 subdiscipes 9, 30

subtorulosa 1, 38

8 suppapillosa 13 136 32: 33

42 testudo 4, 5, 9, 34, 36, 38, 39, 43, 44

+8 v. glabra 9, 35, 38

s vermiculata 1, 9, 10, 36, 37

49 Lactocollybia fj

49 56, 58 Lampteromyces 6, 55

49, 58 :

49, 58 Marasmiaceae 53

2,4 Marasmius 49, 55

23250

22 Nematoctonus 3

7 Oudemansiella

altissima 31.52, 56, 57,/58, Gi, 74

é brunneimarginata 50, 69

4, 50 canafrli ANCA D2. Sof50,1,;

7,8 58, 59, 61, 66, 67, 72

f. radicans 50, 60

V. perstipitata 53, 60

4 crassifolia 49, 56, 60, 61, 62

3, 4, 25, 38, 39, 43, 44 v. incarnata 56, 61, 62

38. 41 echinospora 69

16 japonica . 58

1,3, 30, 38,40; 41 lianicola 49, 51, 52, 56, 63, 64, 67

40 mucida BA 2 33, GV 1s 72

2,4, 7-910: 141 platensis 50) Sk S2...53. 65,66; 67

38, 41 radicata SO 52, 33, 67; 68

1 f. arrhiza 50

1, 4, 10, 12, 18 v. hygrophoroides 49, 54, 55, 67, 68

1,6 raphanipes 50, 69

1, 38, 42 steffenii 69, 70

2 submucida 49°51, 52, 53; 70, 71

2, 43, 44 Vv. persicca 56 Tic h2

£310, 43.414 xeruloides 58

ny - Panellus 8

38 Panus 3,8

3, 4, 5 6, 10. 15 carbonarius 4]

L. 3. 10, 16 subtorulosus 44

SF 6a Veg fb 6 Pleurotus 6, 8, 49, 53, 55

1, 9, 19, 44 Polyporus hs 55

1, 10, 21 Resupinatus IR

t,o. 22

38, 42 Tricholomataceae 55

lL, 3,20 Trogia 34:55

he ; 3 Xerula AS, 502.51, 52753; 55; ss

caussei 5

L, 3 “7 furfuracea 52

2-38 49 43, 44 kuehneri Sz 54

1; 3. 10, ah 28 longipes 55, 68

38, 43, 44 megalospora 52

43 pudens 55

| rubrobrunnescens 52

MS ae It 5 a

3 yohay,

~~ 7

Flora Malesianae Precursores - LVIII, Part Four*

The Genus Schima (Theaceae) in Malesia

HsuAN KENG

Singapore

Abstract

This is a taxonomic treatment of 3 species of Schima (Theaceae) found in the Malesian region. Two,

S. brevifolia and S. monticola, were at one time reduced to the status of subspecies.

Introduction

The binomial Schima noronhae Reinwardt first appeared in Blumes’ Catalogus

(1823); descriptions for the genus and species were added and thus validated

their status in his Bijdragen two years later. The generic name is derived from the

Greek word skiasma (shadow), generally interpreted as referring to the dense

crown of the plant.

Several Schima species were previously described under the genus Gordonia.

Both genera are characterized by the showy, typical theaceous flowers and the

usually 5-valved capsules. Blume, in defining the new genus, pointed out that

Schima differs from Gordonia in their calyx and fruit characters. Subsequent

authors also noted their difference in seed characters. These can be summarized

as follows.

Gordonia:—

Sepals 5 or 6, usually large, unequal, overlapped and imbricate, free, often

deciduous; capsules ellipsoid to cylindric-oblong, often angulate and sometimes

sulcate; seeds ovoid or ellipsoid, flattened, with a large obliquely attached knife-

shaped apical wing.

Schima:-

Sepals 5, rarely 6, smaller, almost equal, weakly imbricate and seemingly

valvate in a fully expanded flower, united below, persistent; capsules globose or

nearly so; seeds broadly reniform, surrounded by a narrow membranous wing

except near the funiculus.

Species of Schima are distributed from E. Himalayas eastwards through

Myanmar, S. China to Taiwan, Ryukyu and Bonin (Oganawara) Islands, and

southwards to Thailand, Indo-China and W. Malesia. No general consensus on

the number of species of the genus has been reached. For example, Melchior

(1925) cited 18 species; Airy-Shaw (1973), 15 species; How (1982), 30 species;

and Mabberley (1987), 1 species.

* Part One to Three: The genera Pyrenaria, Gordonia, and Camellia (Theaceae) in Malesia, in Gardens’

Bulletin, Singapore 33 (1980), 254-289; 37 (1984), 1-47; and 42 (1989), 65-69, respectively.

The last-named author obviously based his opinion on the conclusion drawn

by the late Dr. S. Bloembergen (1952). In that treatment, the genus Schima

contains only 1 species, namely, Schima wallichii (DC.) Korth., which, in turn, is

divided into 9 subspecies. The main arguments as to why Schima is considered

monotypic are: (1) the variation of the ‘vegetative parts’ (leaf-blades and

petioles) varies between clear-cut and rather narrow limits, and (2) the

generatives (flowers and fruits) could never be used in the delimitation of

species, as they vary merely in dimensions and not in number or forms of the

composing parts (Bloembergen, 1952; p. 141). He also frankly admitted that

he only examined ‘few examples from the area outside Indonesia’, but his

‘study of the scanty amount of specimens, literature and the drawings seen

appeared more than sufficient covering’ (p. 133).

It is a fact that size of the flower of Schima, like many other plants, can be

affected to a certain extent by the environment (e.g., soil condition, altitude, etc.)

and the age and condition of the tree, and also even small, immature fruits can be

dehiscent after the processes of pressing and drying. Nevertheless, the extreme

view of totally disregarding all reproductive characters appears to deviate from

the traditional practise of generations of taxonomists. Furthermore it is perhaps

rather imprudent to extrapolate the conclusion based on the study in a limited

area to the whole range of the genus.

Incidently, should Bloembergen’s broad species concept be accepted it would

be probably necessary to revise almost all the existing monographs of theaceous

genera, such as Kobuski’s Eurya (1938), Adinandra (1947), Sealy’s Camellia

(1958), etc. and drastically reduce the number of species of each genus.

Dr. Bloembergen divided the monotypic ‘species’ into nine ‘subspecies’ and

presented these ‘subspecies’ in a map (his Fig. A, on p. 150) which shows that

most of them are geographically isolated. The fact is that these subspecies are of

rather different qualifications: some of them are almost indistinguishable, while

others appear to be perfectly good species in the traditional sense.

For instance, as shown in his map, subsp. noronhae is found in N.W. Borneo,

and subsp. crenata, in E. Borneo. Their difference is largely based on the lamina

margins: margins mostly completely entire vs. margins mostly crenate-dentate.

Exceptions to this are explained in his key and again in the descriptions under

each subspecies. His definition of subspp. noronhae and wallichii is even more

deficient, practically all the characters mentioned in the diagnoses of both

subspecies overlap. Dr. Bloembergen conceded that subsp. wallichii ‘is very

close to subsp. noronhae, but is evidently much less polymorphous, a typical

character being the prominent nervation and the generally forked lateral nerves’.

Unfortunately even the final point does not always stand.

On the other hand, in his discussion under subsp. brevifolia, Dr. Bloembergen

(p. 176) reported that Prof. C.G.G.J. Van Steenis once erroneously mentioned the

occurrence of this plant from Sumatra. He predicted that the specimens misquoted

(Steenis 8636 and 9653) belong either to Laplacea or Gordonia. This plant in

question, in fact was first named Laplacea vulcanica Korthals, and later, renamed

Gordonia vulcanica (Korth.) H. Keng (Keng, 1984; p. 42). It appears to be

inconceivable that these two plants so strikingly similar in many aspects should

be accorded two different taxonomic status, one a species and the other a

subspecies, in two closely related genera. Its small, subsessile, rounded-ovate

leaves and prominent flowers are outstanding. Another of his subspecies, subsp.

monticola, with thick coriaceous leaves and strongly thickened pedicels of flowers

is deemed to be a good species in the traditional sense.

In this treatment, most of Bloembergen’s subspecies as they occur in

Malesia are merged into the species, with the exception of two, namely

subspp. brevifolia and monticola, which are resurrected to their original specific

status. Because of the limitation of knowledge and availability of materials,

this treatment is confined only to the Malesian region.

General Account of the Taxonomic Characters

The Malesian Schima species are mostly medium to tall evergreen trees,

sometimes shrubby at higher altitudes. The branches and branchlets are generally

glabrous or glabrescent, rarely pubescent.

The leaves are simple, alternate and spirally arranged on the branchlets. The

leaf margins are entire, or partly or totally, weakly or strongly undulate or serrate.

Petioles are long or short, or subsessile, mostly slender, sometimes thickened.

The flowers are borne in axillary, solitary or more often in an apical cluster

resembling a cymose or corymbose inflorescence. Following the activation and

elongation of the central dormant bud in the cluster, it becomes clear that each

individual flower is actually borne in the axil of a caducous scale. Sometimes

these flower clusters can be further aggregated into a terminal paniculate

conglomerate.

The flowers are hermaphrodite. Pedicles long or short, slender or stout, generally

2-bracteolate at the apex; the bracteoles are caducous. Calyx and coralla are

clearly differentiated. Calyx is 5-, rarely 6-lobed, the lobes deltoid-rounded, +

equal, weakly imbricate in bud, persistent. Petals are 5, rarely 6 or 4, of unequal

sizes, shortly connate below; the most exterior one is the smallest, strongly

concave and tightly enveloping the other petals in bud. They are white in colour,

often with pink tinge on the outer surface near the base or at the tip.

The androecium consists of numerous stamens which are in 3 to 4 whorls. The

stamens are briefly united below and also adnate to the base of the corolla and

they shed together after anthesis.

The gynoecium consists of a globose ovary, a stout style and a club- or disc-

shaped stigma. The ovary is silky tomentose, usually 5-, sometimes 6-loculate,

with 3(2-5) ovules in each locule.

The fruit is a woody capsule, mostly depressed globose, rarely slightly

elongated, loculicidally dehiscent into 5 valves. The valves eventually break off,

leaving a persistent thick, grooved, central columella. Seeds generally 1 or 2,

sometimes 3 or more in each locule, broadly reniform, flattened, narrowly winged

all round except near the point of attachment. The embryo is large, fleshy and

slightly curved; the endosperm is a thin layer surrounding the embryo.

Taxonomic Treatment

Schima Reinwardt ex Blume

Cat. (1823) 80, nom. nud., Bijdr. (1825) 129; Benth. in B. & H. Gen. PI. 1 (1862) 185; Melchior in E.

& P. Pflanzenfam. ed. 2, 21 (1925) 138; Bloembergen in Reinwardtia 2 (1952) 134.

Small to tall trees, rarely shrubs. Leaves simple, alternate and spirally arranged,

entire or shallowly crenate or serrate. Flowers bisexual, axillary or subterminal,

solitary or many congested into a racemose or cymose cluster, sometimes

paniculate; pedicels long or short, with 2 bracteoles at or near the apex, caducous;

calyx-lobes 5, sometimes 6, deltoid-rounded, subequal, persistent; petals 5-6,

shortly connate below, unequal, the outermost one, oblong and concave; stamens

numerous, briefly united and adnate to corolla at base; ovary spherical, mostly 5-

loculate, with 2 to 5 (mostly 3) ovules per locule, on axile placentation; style

solitary, stout, enlarged and shallowly lobed above into a stigma. Capsule globose

or slightly depressed above, woody, dehiscing loculicidally into 5 (rarely 6 or 4)

valves, with a thick and grooved, persistent central columella. Seeds reniform,

strongly flattened, narrowly winged around except near the funicule; embryo

large and fleshy, slightly curved; endosperm in a thin layer enveloping the

embryo.

A genus with probably around 10-15 species occurs in East and Southeast

Asia. Three species are found in western Malesia.

There is a nomenclatural complication of the generic name Schima. In 1823,

Blume (Catalogus. p. 80) listed Schima noronhae Reinw. and a new species,

Schima excelsa B1. Only the new species was provided with a very brief

description. Later he realized that S. excelsa belongs to the genus Gordonia, and

to which it was duely transferred (Bijdr. 3, p. 130). Meantime, he also prepared

generic and specific descriptions for Reinwardt’s naked names of Schima and

Schima noronhae, thus validating both.

The crucial point is the validity of the earlier Schima in 1823 as it mentioned

two species of which only one was described. Lacking a generic description, the

description of one species (namely S. excelsa) could therefore be maintained as a

combined generic and specific description. Thus if the Schima Reinw. ex B1.

1823 was validly published, then the 1825 name was a later homonym and could

only be maintained by conservation. The late Professor C.G.G.J. van Steenis (in

Taxon 2 (1953)115) therefore proposed to conserve Schima Reinw. ex B1. 1825.

A majority of the members of the Committee for Spermatophyte Conservation of

generic names of IAPT at a 1959 meeting, however, decided that the 1823

publication was invalid and the 1825 name could stand without conservation

(Taxon 9 (1960) 15).

Key to the species

A. Petioles of leaves subsessile, generally 2-3mm long; leaf-blades mostly rounded ovate to obovate. ..........

I ee 9 La aa Ma rag I is 5 aac sien asnsas and uensseonphsnnnspuseoscuspeonsacbe S. brevifolia

A. Petioles of leaves usually more than | cm long; leaf-blades generally lanceolate to oblong.

B. Leaf-blades generally thick coriaceous; pedicels, especially the upper part strongly thickened (usually

GEE IE SARIS Seg te PONE ol oA Pa” OA EE 2 SS Rs S. monticola

B. Leaf-blades generally chartaceous to coriaceous; pedicels mostly slender (generally less than 2 mm

ia eg ese se 5 eg dy son dackdaip saps asunvahenecploacbsuespe cogs rcbuitees S. wallichii

Schima brevifolia (Hook. f.) Stapf

Hook, Icon, Pl. 23(4)(1893) pl. 2264; in Trans. Linn. Soc. Bot. 2(4) (1894) 135; Gibbs in J. Linn. Soc.

Bot. 42 (1914) 60; Airy-Shaw in Kew Bull. 1914 (1914) 498: Melch. in E. & P. Pflansenfam. ed.

2, 21 (1925) 139.

Gordonia brevifolia Hook. f. in Trans. Linn. Soc. 23 (1860) 162; Burk. in J. Str. Br. As. Soc. Beng. 76

(1917) 158.

Schima noronhae Reinw. ex B1. subsp. brevifolia (hook. f.) Steenis in Bull. Jard. Bot. Btzg. III. 13

(1936) 50.

Schima wallichii (DC.) Korth. subsp. brevifolia (Hook. f.) Bloemb. in Reinwardtia 2 (1952) 177, f. C 8

& 10. J. 5-9.

A spreading shrub, 1.5-2m high, rarely a small tree, to 5m tall. Leaves generally

crowded on the tips of branches; leaf-blades coriaceous, broadly ovate or ovate-

elliptic, sometimes suborbicular, 2.5-5(-8) cm long, 2-4 (-6.5) cm wide, apex

obtuse or rounded, sometimes briefly subcaudate, base rounded or subcordate;

margin subentire or finely crenulate; lateral veins 6-9 pairs, merged and looped

into submarginal reticulation; young leaves soft white silky pubescent beneath,

mature ones glabrous or glabrescent on both surfaces; petiole short and stout,

usually 2-3 mm long. Flowers 4-5 cm across, solitary in upper leaf-axils, often

congested at the top of branchlets, in cyme-like or raceme-like clusters; pedicels

1.2-1.6 (-3) cm long, slightly thickened toward the apex, 2-bracteolate, glabrous

or pubescent; calyx mostly 5-lobed, the lobes deltoid or subcordate, 3-5 mm

long; petals 5, rarely 6, creamy white, often with a tinge of pink near the base,

obovate or rounded, subequal, 2-2.5 cm long with a cuneate base, soft pubescent

or pilose on the outer surface especially near the base ovary globose, hirsute.

Capsule depressed globose, 1.6-2.5 cm across, glabrous or velvet.

Schima brevifolia (Hook. f.) Stapf

Distribution. Borneo (Sabah and Sarawak).

Sabah. Ranau, Mt. Kinabalu (numerous collections, only representative

specimens cited), J. & M.S. Clemens 32444, 32637; G. Mikil SAN 31772, 41769;

J. Sinclair et al. 80383.

Sarawak, Kalabit Highlands, Mt. Murud, P. Chai 02045; H.P. Nooteboom &

P. Chai 02272.

Ecology. In montane oak forest, mossy forest, or on sandstone at the summit

or on slopes; from 1800 to 3500m.

Schima monticola Kurz

J. As. Soc. Beng. 43(2)(1874) 93, 181; For. F1. Burm. 1 (1877) 107; Szyszyl. in E. & P. Pflanzenfam.

3(6) (1895) 186.

Schima noronhae Reinw. ex B1. var. rigida Ridl. Fl. Mal. Pen. 1 (1922) 202.

Schima forrestii Airy-Shaw in Kew Bull. 1936 (1936) 496.

Schima wallichii (DC.) Korth. subsp. monticola (Kurz) Bloemb. in Reinwardtia 2 (1952) 176, f. C9 &

11, J4 a-c.

Small, medium-sized to tall tree, 5-10 up to 35m high, much branched; bark

dark brown, scaly, or coarsely irregularly dippled; the branches and branchlets

glabrous or silky pubescent. Leaf-blades thick-coriaceous, elliptic or oblong

lanceolate, 6-10 (-15) cm long, 4-7 (-8) cm wide, apex acute, acuminate or

subrounded, base obtuse, margin crenulate to crenate-serrate, lateral veins 9-12

pairs, less distinct, glabrous above, glabrescent or silky pubescent beneath; petiole

1-2 cm long. Flowers 4.5-7 cm across, axillary, solitary, often congested above in

raceme- or cyme-like clusters; pedicels 1.5-3 (-5) cm long, prominently swollen

over their entire length especially the upper part, shallowly two-keeled on the

dried specimens; bracteoles deltoid, 3-6 mm wide; calyx reddish, mostly 5-lobed,

the lobes subrounded, 3-6 mm long, silky pubescent externally; corolla 4.5-6 cm

across, white with pink tinge on the outer surface. Fruit depressed globose, 1.5-

2.5 cm across, silky tomentose.

Distribution. Myanmar, S.W. China and Malesia (The Malay Peninsula and

Borneo).

Malay Peninsula: G. Tapis, Pahang, Y.C. Chan FRI 19858, P.F. Cockburn

FRI 11004; Cameron Highlands, J.F. Maxwell 78193; G. Ulu Kah, Pahang and

Selangor Border, T7.C. Whitmore FRI 12210; G. Padang, Trengganu, Whitmore

FRI 12660, 12664, 12724.

Borneo (Sarawak, Brunei, Sabah and Kalimantan)

Sarawak. Ulu Sg. Dapur, Mt. Murud, /lias Paie S$ 26446; Kalabit Highlands,

H.P. Nooteboom & P. Chai 01721, P. Chai §35927.

Brunei. Ulu Tanburong, Medaint Watershed Ridge, P.S. Ashton 2553.

Schima monticola Kurz

Kalimantan. G Paris, South of Long Bawan, Krayan, Koto Okamoto & Ueda

Walujo B. 7488.

Sabah. (Representative specimens) Mt. Kinabalu, W.L. Chow & E.J.H. Corner

RSNB 4082, J. & M.S. Clemens 31987, 32435, 33197; G.H.S. Wood SAN 16717.

Ecology. It is usually found in dipterocarp or oak forests at 1400 to 1800m,

sometimes can reach as high as 2200m on top of the ridges. It also occurs in

Agathis forest or heath forest at 500-700m in certain coasts of Borneo.

Schima wallichii (DC.) Korth.

Kruidk (1842) 143; Choisy in Zoll. Syst. Verz. 2 (1854) 144; Mig. FI. Ind. Bat. 1 (1859) 492; Foxw. in

Philip. J. Sc. 4 (Bot.) (1909) 503; Bloemb. in Reinwardtia 2 (1952) 136, p.p.; Back. & Bakh. f. Fl.

Java 1 (1963) 321.

Gordonia Wallichii DC. Prodr. 1 (1824) 528.

Schima noronhae Reinw. ex B1. Cat. (1823) 80, nom. nud., Bijdr. (1825) 130; Mig. FI. Ind. Bat. 1

(1859) 492; Ridl. Fl. Mal. Pen. 1 (1922) 201.

Schima crenata Korth. Kruidk (1842) 143, pl. 29.

Schima antherisosa Korth. Kruidk (1842) 145; Mig. FI. Ind. Bat. 1 (1859) 492.

Schima hypoglauca Migq. Sum. (1862) 190, 484.

Schima bancana Miq. Ann. Mus. Bot. Lugd.-Bat. 4 (1868) 113.

Schima rigida Miq. |.c. 4 (1868) 113.

Schima sulcinervia Migq. l.c. (1868) 113.

Schima lobbii (Hook. f.) Pierre, Fl. For. Cochinch. 2 (1887) pl. 121.

Schima pulgarensis Elm. Leafl. Philip. Bot. 5 (1915) 1843.

Small to large tree, 5-30 (-45)m tall; young branchlets appressed pubescent,

old ones glabrous or glabrescent; bark pale reddish to dark brown, fissured. Leaf-

blades usually chartaceous to thin coriaceous, lanceolate to oblong-elliptic, 7-15

(-24) cm long, 1.5-5 (-7.5) cm wide, apex acute or acuminate, base acute, margin

entire, shallowly crenate to serrate, glabrous or glabrescent on both surfaces,

glaucous or sometimes pubescent below, especially along the midrib; petiole 1-

2.5 (-3) cm long. Flowers 4.5-7 cm across; pedicel slender, often slightly thickened

near the tip, 1.5-4 (-6) cm long; calyx-lobes subrounded, 2.5-3 mm across; petals

ovate or obovate, unequal, white, sometimes with a tinge of pink near the base

externally, 2-3.5 cm long, labrous or glabrescent on the inner surface, pubescent

or partly so on the outer surface; ovary densely pubescent. Capsule spherical or

depressed globose, silvery pubescent, 1.5-2 cm across, generally bearing a style-

base.

Distribution. From Nepal, Sikkim, Assam, Myanmar, S. China to Malesia

(The Malay Peninsula, Sumatra, Java, Borneo, Celebes and the Philippines.)

(only representative specimens are cited)

The Malay Peninsula. Fraser’s Hill, Pahang, van Balgooy 2071; Penang Hill,

Penang, S. Chelliah FRI 98134; Maxwell’s Hill, Taiping, B. Everrett FRI 13570;

G. Ledang, Johore, H.S. Loh FRI 19200; Taman Negara, Pahang, E. Soepadmo

910.

Sumatra. Banka, Biinanmeijer 1409, 1903; G. Malintang, Biinnanmeijer 3624;

Palembang, F.H. Endert 314; G. Leuser, Atjeh, de Willde & de Willde-Duyfjes

15614.

Java. G. Salak, Preanger, Bakh. v./d. Brink 1728; G. Papandajan, S.H. Caerb

635; Tijbodas, Danser 5932; G. Malabar, H.O. Fobres 959d, 1072a.

Borneo. Sarawak. Bt. Tibong, Kapit, J.A.R. Anderson et al. S28679, Bt.

Goram, P. Chai $36/03; Bt. Pantau, Melinau, /lias Paie $25723; Telok Bandung,

Santubong, /lias & Jugah S38685. Brunei. Bt. Teraya, P.S. Ashton $7894. Sabah.

Bt Tangunan, Telupid, Abd. Rahim et al. SAN 93288, Mt. Tambayukan, Renau,

Aban Gibot SAN 55427; Sosopedon, Renau, G. Mikil SAN 34515. Kalimantan.

Grayau, Selim Bau, W. Borneo, J.J. Afriastini 1122, Central E. Borneo, F.H.

Endert 3635; Loa Byanan, W. of Samarinda, Dostermans 6400; G. Niut, Ponitanak,

G. Shea 28145.

The Philippines. Mt. Pugar, Palawan, ADE. Elmer 13191 (isotype of Schima

pulgarensis Elm.); Baguio, Mt. Province, M.L. Steiner 2110.

Celebes. N.E. of Makassar, W. Meijer 10747.

Ecology. In oak-laurel forest or mossy forest, or on slopes of dry hill side,

from coastal heath forest near the sea level to 1500 m or sometimes reaching

2500 m in montane forest in Malesia.

Acknowledgements

I am grateful to the Executive Director, National Parks Board, Singapore for

the herbarium and library facilities provided, and to the Director and staff of the

Rijksherbarium, Leiden for the loan of the entire collection of Schima specimens.

I would also like to extend my thanks to Dr. Ding Hou for his effort to

search some literature which are not available in Singapore, to Mrs. Ng

Siew Yin and Dr. Chin See Chung, Inche Md. Shah and other staff of the

Herbarium of the Singapore Botanic Gardens for their willing assistance in

many ways, and to my wife, Mrs. Ro-siu Ling Keng for preparing the

illustrations reproduced in this paper and for her constant encouragement.

References

Airy-Shaw, H.K. (1973), A dictionary of the flowering plants and ferns. (J.C.

Willis). Rev. 8th ed. Cambridge Univ. Press.

Bloembergen S. (1952). A critical study in the complex-polymorphous genus

Schima (Theaceae). in Reinwardtia 2: 133-183.

Blume, C.L. (1823), Catalogus van eenige der merkwaardigste zoo in- als uit-

heemsche gewassen, te vinden in ‘s Lands Plantentuin te Buitenzorg. Batavia.

— (1825). Bijdragen tot de Flora van Nederlandsch Indié. 17 parts (1825-27).

Batavia.

How, F.C. (1982). A dictionary of the families and genera of Chinese seed plants

(rev. by T.L. Wu et al.) 2nd ed. Beijing: Science Press.

Keng, H. (1984), Florae Malesianae Precursores LVIU, pt. 2. The genus Gordonia

(Theaceae) in Malesia. Gard. Bull. Sing. 37: 1-47.

Kobuski, C.E. (1938). Studies in Theaceae. III. Eurya subgenera Euryodes and

Penteurya. Ann. Mo. Bot. Gard. 25: 299-359.

— (1947). Studies in Theaceae XV. A revision of the genus Adinandra. Jour.

Arnold Arb. 28: 1-98.

Mabberley, D.J. (1987). The Plantbook. Cambridge Univ. Press.

Melchior, H. (1925). Theaceae. In: A. Engler and K. Prantl, Die Natiirlichen

Pflanzenfamilien. 2nd ed. 21: 109-154. Leipzig.

Sealy, J.R. (1958). A revision of the genus Camellia. London.

BENJAMIN CLEMENS STONE

1933-1994

Photograph taken at Philippine National Herbarium, Oct. 1992 by Ali Ibrahim.

Dr Stone arrived in Peninsular Malaysia in 1965 to take up a

teaching position in Botany, at the University of Malaya, Kuala

Lumpur. It was from there that he began his long association

with the Singapore Herbarium and the Botanic Gardens. In the

pursuit of his many botanical interests, including what he termed

“the big game of the plant world,” members of the Pandanaceae,

he made repeated visits to the Herbarium and the Gardens.

Periodically over the years he would send seeds to the Gardens

from his field trips around the region. More frequently than

seeds came manuscripts. Twenty of his papers, including those

posthumously published in this issue, have been accepted by

this Bulletin. These reflect his wide interests in botany and

covered topics from the families, Pandanaceae, Rutaceae,

Araliaceae, Joinvilleaceae, Nymphaeaceae and Myrsinaceae.

The staff of the National Parks Board who knew Dr Stone,

some since his first visit in 1967, were shocked and grieved

when news of his fatal heart attack arrived. His enthusiasm and

intellect have been inspirational to many who knew him; to

some of us he was a teacher and friend as well. We will all miss

the sharing of his ideas and this Bulletin will miss his

contributions.

NATIONAL PARKS BOARD

SINGAPORE

In Memoriam B. C. Stone (1933-1994)

A. LATIFF

Department of Botany

Universiti Kebangsaan Malaysia

43600 UKM Bangi, Malaysia

As was almost simultaneously announced by Drs. Domingo Madulid in Manila

and Sy Sohmer in Fort Worth, Texas, Dr. Benjamin Clemens Stone died about

three months before his 61st birthday, on 19 March 1994. His sudden death took

everybody who knew him by suprise. His first publication about Malesian botany

and plants was in 1960. He has thus devoted some 29 years of his life to the study

of the Malesian flora. His first interest, however, was in the Hawaiian Flora with a

first publication in 1957. He has also worked on other flora, notably that of the

Pacific Basin.

Ben was born as the only child to an English father and American mother, in

Shanghai, China on 26 May 1933. Before the war, his mother took Ben to live in

the San Diego area in California, sensing the coming of troubles in the Far East.

The Stone family was united after the war, after his father was interned in a camp

in Japan. Ben married Michiko and they had a son, David and a daughter, Sylvia.

Ben received his primary and secondary education in the San Diego area, and

his B.A. in Botany from Pomona College, in Claremont, California in 1954. He

was so sentimentally attached to that college that he supported David studying

there with great pride. He wrote a thesis on the pandans and earned his Ph.D

from the University of Hawaii in 1960. His major professor in Hawaii was Dr.

Harold St. John, a fixture in Hawaiian botany from 1926 to 1991, when he passed

away at the age of 99. Ben’s career path then took him to Washington, where he

was a research assistant in the Department of Botany at the U.S. National Museum

of Natural History at the Smithsonian Institution, from 1960-61. From there he

went to Guam where he was made Professor of Biology at the College (now

University) of Guam from 1961 to 1965. It was during this period that he wrote

the Flora of Guam single-handedly and established Micronesica, the journal of

the University of Guam.

While in Guam he showed his initial interest in Flora Malesiana (1960) when

he confessed that he wished to master the flora of the adjacent Malesian region

after mastering the flora of Polynesia and Micronesia. He grabbed the opportunity

to work on the Malesian flora by taking a lecturer position in the Department of

Botany, University of Malaya at Kuala Lumpur in 1965.

From 1965 to 1983 he taught botany and in particular plant taxonomy,

supervised numerous under- and post-graduate students, researched on plants and

vegetation, looked after the herbarium (KLU) and contributed intellectual and

scientific integrity, in the Department of Botany at the University of Malaya. His

inspired scholarship and untiring dedication contributed towards putting Malaysian

taxonomy in particular, and botany in general, at its height to-day. Scores of

students were attracted to botany as a result of his mentoring and he developed an

understanding about the culture, custom and tradition of the region, especially of

Polynesia, Micronesia, Malaysia and later the Philippines.

I entered the University of Malaya in 1969 and only in my final year, came to

know him closely. I remembered him as a lecturer, plant taxonomist, botanist

extraordinaire, story teller, jazz flautist, devoted model airplane builder and plant

collector. As a botanist he saw the necessity for herbaria to facilitate taxonomic

works (1965), strategies for conservation of ecosystems and plants (1968), national

parks as reservoirs of biodiversity (1969, 1968), understanding plant genetic

resources (1974, 1979), taxonomic man-power to conduct research (1974),

scholarly publications as an extension of mental exercise (1975), botanical gardens

to complement the herbarium (1978), natural history museums for general

education (1981), and excellent plant collecting techniques to ensure collections

useful to future botanists (1983, 1987). As a plant taxonomist he was not as

conservative as the late Dr. M. Jacobs and Dr. P.W. Leenhouts of Rijksherbarium,

Leiden, for instance, in his recognition of Glycosmis calcicola (1972), and some

myrsinaceous taxa in Malesia (1982). He was a Malesian taxonomist extraordinaire

who mastered four difficult plant families, viz. initially the Pandanaceae, then the

Rutaceae, the Araliaceae and lately the Myrsinaceae. He also worked on the

Theophrastaceae, Alangiaceae, Orchidaceae, and many others including some

ferns.

Apart from taxonomy, his research and publications covered evolutionary

biology, ecology, plant geography, cytology, phytochemistry and vegetation study.

Although he gave far greater importance to gross morphological characters as the

basis for classification, he had a great interest in the contributions given by

comparative anatomical, palynological and cytological studies. I noticed he had

not used the work ‘cladistic’ at all in his publications which prompts me to

believe that he had no interest in that subject.

As a man I found him very honest and liberal in his thoughts. He always

smiled and never once had I observed him to lose his cool though there were

many occasions when others would have shouted or split hairs. He had wanted to

call me “Lat” a shorter form of my second name which I disliked because that

was the name of a cartoonist! He was very helpful in the field teaching me many

Latin names and diagnostic features of various plant families that we encountered.

I remembered very well in 1982 that he was overjoyed when he could at last buy

himself a fancy electric typewriter. The man loved to type his correspondence

though he had excellent hand-writing.

In 1983 after 18 years in humid Kuala Lumpur, he pondered seriously about

the college education of his children, David and Sylvia. He talked about his aged

mother and a flat he had bought in California. In 1984 he decided to move back

to the U.S. by taking an optional retirement as a Reader in Botany from the

University of Malaya and took the position of Chairman, Department of Botany

of the Philadelphia Academy of Natural Science, a position he held until 1990

when he joined the Flora of the Philippines Project as the Principal Investigator

for the Philippine Plant Inventory at the urging of Dr. Sy Sohmer at Hawaii.

When the latter moved to BRIT at Fort Worth, Texas, he gladly joined the staff as

Senior Research Botanist.

In Philadelphia he felt wasted, the department was too spacious but under-

staffed and under-equipped. The place had no IBM PC system to store type

collections or word processing. He was very ambitious indeed to start working

on Malesian plants, but the department had no interest in Malesia. Thus he tried

to build the Asiatic collection in order to initiate research development. He had

wanted to arrange exchange programmes for plant specimens and also develop a

joint proposal between his department and Malesian or Asian institutions and

herbaria, especially the Tree Flora of Sabah and Sarawak Project. The programme

should have an adequate opportunity for post-graduate training and post-doctoral

exchange. He had wanted his friends in the universities (Prof. Ruth Kiew, Prof.

E. Soepadmo and myself) to look after the former aspects.

From 1990 to the day when he passed away he lived at intervals in the U.S.

and Manila. On that fateful Saturday, 19th March 1994 he routinely went to the

herbarium to take some plant presses off the driers. He collapsed and the security

guard on duty failed to revive him. So ended a life that began in Shanghai, China

and ended in Manila, where he had began his second intimate affair with the

Malesian plants through the Philippine Flora Project.

Ben was the author or editor of more than 300 articles and books. He was a

specialist in four difficult tropical families, the Pandanaceae, Rutaceae, Araliaceae

and Myrsinaceae. He wrote or co-authored numerous monumental papers in these

families and his reputation as a Pandanus worker was unsurpassed. Unfortunately,

he died before he could finish his monograph of Freycinetia and contribute the

family Myrsinaceae for the Tree Flora of Sabah and Sarawak as well.

Ben has had some influence on the development of Malaysian botany, of a

magnitude large enough for his name to be forever linked to it. He has also been

quite diverse in all fields in which he has been active. A great botanist has passed

away after 61 long, well-spent and useful years, of which almost 34 years was

devoted to botany alone. He was a friend to many all over the world, and he was

always prepared to help people with his vast knowledge.

He will be missed very much.

Bibliography of Malesian Botanical Publications of B.C. Stone

1960

Flora Malesiana : A review. Pac. Sci. 14 (4) : 423-424.

1965

On the purpose of a University Herbarium. The Malayan Scientist 2 : 23-26.

Pandanus Stickm. in the Malayan Peninsula, Singapore and lower Thailand. Mal.

Nat. J. 19 (4) : 203-213.

1966

Pandanus Stickm. in the Malayan Peninsula, Singapore and Lower Thailand.

Mal. Nat. J. 19 (5) : 291-301.

1967

A new species of Pandan from Sarawak, with notes on Sect. Mutidens St. John.

Fedn. Mus. J. n.s. 11 : 113-119.

A new species of Pandanus from East Malaysia. The Malayan Scientist 3 : 24-

25h

Materials for a monograph of Freycinetia (Pandanaceae) I. Gard. Bull. Sing. 22

(2) : 129-152.

Studies of Malesian Pandanaceae, I. Gard. Bull. Sing. 22 (2) : 231-257.

1968

The genus Pandanus in Malaya, Singapore and Lower Thailand. Part 3. Mal.

Nat. J. 21 (1) : 3-16.

The genus Pandanus in Malaya, Singapore and Lower Thailand. Part 4. Mal.

Nat. J. 21 (2) : 125-141.

Studies in Malesian Pandanaceae, II]. Two new species of Pandanus Sect.

Fusiforma St. John. Reinwardtia 7 (4) : 411-420.

Materials for a monograph of Freycinetia (Pandanaceae) III. Freycinetia corneri,

a new species from Malaya. Brittonia 20 (3) :198-202.

Conservation of vegetation. The Malayan Scientist 4 : 42-48.

Typification of Schizostegeopsis Copeland. Philipp. J. Sci. 95 (4) 425.

(with J. J. Gaudet). Plant life of Malaysia. Book Review. Quart. Rev. Biol. 43

(3) : 308-310.

Materials for a monograph of Freycinetia (Pandanaceae) IV. Subdivision of the

genus, with fifteen new sections. Blumea 16 (2) : 361-372.

Morphological studies in Pandanceae I. Staminodia and pistillodia of Pandanus

and their hypothetical significance. Phytomorphology 18 (4) : 498-509.

1969

National Parks a National Resource. In Stone, B.C. (Ed.). Symposium on natural

resources and conservation in Malaysia and Singapore. University of Malaya,

Kuala Lumpur. pp 1-8.

Studies in Malesian Pandanaceae III. Notes on Pandanus Section Solmsia, sect.

nov. Fedn. Mus. J. n.s. 12 : 105-110, 4 plates.

Studies in Malesian Pandanaceae IV. A revision of Asterostigma and Asterodontia,

two sections of the genus Pandanus. Fedn. Mus. J. n.s. 12 : 111-116.

Studies in Malesian Pandanaceae V. Two new subsections of Pandanus sect.

Acrostigma Kurz, Scabridi and Dimissistyli. Fedn. Mus. J. n.s. 12 : 117-121.

Additions to the Malayan Flora I. Zanthoxylum acanthopodium. Mal. Nat. J. 23 :

31-32, fig. 1.

Materials for a monograph of Freycinetia Gaud. IX. A revised list of Philippine

species with critical notes and some new taxa. Webbia 23 (2) ; 597-607.

Materials for a monograph of Freycinetia Gaud. X. Chronological list of all

binomials. Taxon 18 (6) : 672 680.

Studies in Malesian Pandaceae VI. The identity of Pandanus pilaris Ridley.

Webbia 23 (2) : 607-610.

1970

Malayan climbing pandans: the genus Freycinetia in Malaya. Mal. Nat. J. 23 ;

84-91. 3 plates.

Morphological studies in Pandanaceae II. The ‘coniferoid’ habit in Pandanus

section Acanthostyla. Bull. Torrey Bot. Club 97 (3) : 144-149.

Fruits and seeds of the frangipani (Plumeria). Mal. Nat. J. 23 : 143-144, Plates

18.

Materials for a monograph of Freycinetia Gaud. (Pandanaceae) XIII. A new

species from Ternate island, Moluccas. Pac. Sci. 24 (3) : 417 419.

Materials for a monography of Freycinetia Gaud. (Pandanaceae) V. Singapore,

Malaya and Thailand. Gard. Bull. Sing. 25 (2) : 189-207.

Materials for a monograph of Freycinetia Gaud. (Pandanaceae) VI. Species of

Borneo. Gard. Bull. Sing. 25 (2) : 208-233.

The botanical identity of mengkuang, a common Pandanus in cultivation in

Malaya. Malayan Agriculturist 9 : 34-44, 4 plates.

1971

Additions to the Malayan Flora II. Scirpus confervoides. Mal. Nat. J. 24 ; 91-94.

(with T.C. Whitmore). Notes on the systematy of Solomon Islands and some of

their New Guinean relatives. Reinwardtia 8 (1) : 3-11.

Studies in Malesian Pandanaceae VIII. Some new and little-known sections of

Pandanus. Fedn. Mus. J. n.s. 13 : 138-149.

Studies in Malesian Pandanceae IX. Taxonomic notes on Pandanus species and

synonyms. Fedn. Mus. J. n.s. 13 : 150-154.

Materials for a monograph of Freycinetia Gaud. XII. Fedn. Mus. J. n.s. 13 : 155-

165.

(with A.L. Lim). Notes on systematic foliar anatomy of the genus Freycinetia

(Pandanaceae). J. Japan Bot. 46 (7) : 14-24.

(with C.H. Cheah) . Reports on Pandanaceae. In IOPB Chromosome number

reports XXXII. Presented by Askell Love. Taxon 20 : 356.

Lov 2

A reconsideration of the evolutionary status of the family Pandanaceae and its

significance in monocotyledon phylogeny. Quart. Rev. Biol. 47 (1) : 34-35.

Rutaceae. In T. C. Whitmore (Ed.). Tree Flora of Malaya 1| : 367-387. Longmans.

Studies in Malesian Pandanaceae VII. A review of Javanese Pandanaceae, with

notes on plants cultivated in the Hortus Bogoriensis. Reinwardtia 8 (2) : 309-

318.

A new wild Citrus species from Malaya. The Planter (Kuala Lumpur) 48 :

90-92.

The genus Pandanus in the Solomon Islands, with notes on adjacent regions.

Malays. J. Sci. 1A : 93-132.

Studies in Malesian Pandanaceae X. Four new Malesian species of Pandanus and

notes on P. sarawakensis. Fedn. Mus. J. n.s. 14 : 127-136.

Arthrodactylis and Pandanus : A brief comment on the “Characteres Generum

Plantarum” of J.R. Forster. Gard. Bull. Sing. 26 (1) ; 113-114.

Notes on the systematy of Malayan phanerogams XVI. Glycosmis calcicola,

n.sp. (Rutaceae). Gard. Bull. Sing 26 (1) : 55-57.

Studies in Malesian Pandanaceae XI. A new species of Pandanus. Fedn. Mus.

te. Says 99-202.

Materials for a monograph of Freycinetia Gaud. (Pandanaceae) . XV. The

Sumatran species. Fedn. Mus. J. n.s. 15 : 203-207.

Additions to the Malayan Flora III. Gastonia papuana. Mal. Nat. J. 25 : 164-165.

Pandans. Malaysian Panorama 2 (2) : 15-18. Ministry of Foreign Affairs, Kuala

Lumpur.

Bellucia : A South American fruit tree found in Malaya. The Planter (Kuala

Lumpur) 48, (588) : 276-278.

The interface between teaching and research. In: D.S. Nijhar (Ed.). The role of

Asian universities in a changing world. University of Malaya, Kuala Lumpur. pp

29-32.

1973

(with S.P.Chu). Morphological studies in Pandanaceae V. A further survey of

folliar anatomy in the genus Pandanus. J. Japan Bot. 48 (2) : 55-64.

The genus Pandanus in the Solomon Islands with notes on adjacent regions.

Malays. J. Sc. 24 : 59-80.

(Book review). P. F. Allen. Natural rubber and the synthetics. The Planter

(Kuala Lumpur) 49 (566) : 208-209.

(with J.B.Lowry, R.W. Scora & K. Jong). Citrus halimii : A new species from

Malaya and Peninsular Thailand. Biotropica 5 (2) : 102-110.

(with C.H. Cheah). Studies in Malesian Pandanaceae XIV. Pandanus Bot.

Jahrb. 93 (4) : 498-529.

(with D.W. Lee & J.B. Lowry). Effect of drought on Kinabalu. Mal. Nat. J. 26

(3&4) : 178-179.

Copelandiopteris, a new genus of Philippine ferns. Webbia 28: 491-494.

Citrus. Malaysian Panorama 3 (4) : 2-5. Ministry of Foreign Affairs, Kuala

Lumpur.

1974

(with K. Jong & E. Soepadmo). Malaysian tropical under-exploited genetic

reservoir of edible fruit tree species. In E. Soepadmo & K.G. Singh (Eds.) .

Proceedings of the symposium on biological resources and national development.

pp. 113-121.

Studies in Malesian Pandanaceae XII. Pandanus sects. Solmsia and Rykiopsis

and Rykia subsect. Gressittia. Fedn. Mus. J. n.s. 17 ; 99-163.

Studies in Malesian Pandanaceae XIII. New and noteworthy Pandanaceae from

Papuasia. Contrib. Herb. Austral. 4 : 7-40.

A white-flowered variant of the beach morning glory, jpomoea pescaprae. Mal.

Nat. J. 27 (1&2) : 17-19.

(Book Review). Proceedings of the symposium on biological resources and

national development. The Planter (Kuala Lumpur) 50 (576) : 114-115.

Towards an improved infrageneric classification in Pandanus (Pandanaceae).

Bot. Jahrb. Syst. 94 : 459-540.

Taxonomists for Malaysia : A vital need. The Planter 50 (580) : 199-200.

National ecosystem reserves : key to success in nature conservation. In W.C.

Lim & R. Singh (Eds.). Battle for the environment : the Malaysian experience.

Malayan Nature Society & Consumers Association of Penang. pp. 44-46.

Langkawi landscape. Malaysian Panorama 4 (2) : 2-7.

1975

Quinine : Time for a revival? The Planter 51 (586) : 14-15.

(with K.M. Kochummen). A new Alangium (Alangiaceae) from Sarawak. Blumea

22 (2( : 35-38.

Studies in Malesian Vitaceae XV. Two new species of Pandanus from

Mt. Kinabalu, Sabah (Borneo), with notes on the Pandanaceae of Mt.

Kinabalu. Malays. J. Sci. 3A : 69-74.

Appendix to Henderson’s Malayan Wild Flowers. Mal. Nat. J. 28 : 57-83.

Scholarly serial publications of academic institutions and societies in Malaysia

to-day : A review and commentary. In B. Lim (Ed.) . Scholarly publishing in

S.E. Asia. Universiti Malaya Press, Kuala Lumpur. pp. 36-42.

(with C.H. Cheah). Embryo sac and microsporangium development in Pandanus

(Pandanaceae). Phytomorphology 25 92) : 228-238.

1976

Sandbur (Cenchrus echinatus), an addition to the Malaysian grass flora. The

Planter (Kuala Lumpur) 52 (607) ; 397-399.

Germination and early seedling growth in Melanorrhoea woodsiana, a Malaysian

species of rangas. Malayan Naturalist 3 (1&2) : 49.

Copelandiopteris endoneura (Price) Stone, comb. nov. Kalikasan 5 (3) : 329-

ASF,

1977

(Review). T.C. Whitmore. Tropical Rain Forests of the Far East. Mal. Nat. J. 30

(1) : 115-118.

(Review). T.C. Whitmore (Ed.). Tree Flora of Malaya 1. Quart. Rev. Biol. 52 (1) ;

91-93.

(Review). C.F. Symington. Forester’s manual of dipterocarps. Quarts. Rev.

Biol. 52 (1) : 91-93.

(with D.W. Lee, M. Ratnasabapathy & T.T. Khoo). The natural history of Pulau

Tioman. Merlin Samudera Tioman Sdn. Bhd., Kuala Lumpur. 69 pp.

(Review). P.F. Cockburn. Trees of Sabah 1. Quart. Rev. Biol. 52 (1) : 91-93.

On the identity of the Nicobar bread-fruit and a new record of Pandanus leram in

beach drift on Pulau Langkawi. Mal. Nat. J. 30 (1) : 93-102.

(with H.J.M. Bowen & J.F. Veldkamp). European weeds introduced to Gunung

Ulu Kali, Pahang, Malaya. Mal. Nat. J. 30 (1) : 103-108.

(Review). The pterocarpus. Philippine Science Journal of Forestry. Mal. Nat. J. 30

(1) : 109.

The morphology and systematics of Pandanus (Pandanaceae) to-day. Gard.

Bull. Sing. 29 : 137-142.

Notes on the systematy of Malayan Phanerogams XXV. Araliaceae. Gard. Bull.

Sing. 30 : 275-291.

1978

A proposal for a Malaysian national Park service. Mal. Nat. J. 29 (4) : 257-263.

(Book review). W. Jack, Description of Malayan Plants I-III. J. Mal. Br. Roy.

Asiat. Soc. 51 (1) : 122-124.

(Book review). K. Paijmans, New Guinea Vegetation. Quart. Rev. Biol. 53 (2) :

171-172.

Revisio Pandanacearum Part I. Pandanus subgenera Coronata and Acrostigma;

Flora Malesiana Precursores. Fedn. Mus. J. n.s. 23 : 1-74.

Studies in Malesian Rutaceae I. Notes towards a revision of the genus Glycosmis

Correa. Fedn. Mus. J. n.s. 23 : 75-110.

Studies in Malesian Rutaceae II. New records and new taxa of Aurantoideae

from Borneo. Fedn. Mus. J. n.s. 23 : 111-116.

A new orchid from Gunung Ulu Kali, Pahang. Fedn. Mus. J. n.s. 23 : 116-123.

Additions to the Malayan Flora, no. 6. The genus Ornithochilus (Orchidaceae)

new to Malaysia. J. Malay. Br. Roy. Asiat. Soc. 51 (2) : 139-142.

(Book review). E.J.H. Corner, The freshwater swamp forest of South Johore and

Singapore. Mal. Nat. J. 30 (3&4) : 613-616.

Araliaceae. In F.S.P. Ng (Ed.) Tree Flora of Malaya 3 : 10-35. Longmans.

(with R. Kiew). Ilex micrococca Maxm. and J. polyneura (Hd. Mat. ) Hu

(Aquifoliaceae) : new records for Malaya and Thailand. Mal. Nat. J. 32(2) : 149-

156.

1979

Barclaya, the Malaysian waterlily. Malay. Naturalist : 20-22.

(with D.W. Lee & J.B. Lowry). Abaxial anthocyanin layer in leaves of tropical

rainforest plants; enhancers of light capture in deep shade. Biotropica 11 (1) :

70-77.

Studies in Malesian Pandanaceae XVII. On the taxonomy of ‘Pandan wangi’ a

pandanus cultivar with scented leaves. Economic Botany 32 : 285-293.

(with S. Bien). Genetic resources of essential oil plants in Malaysia. Malays.

Appl. Biol. 8 : 53-58.

Protective coloration of young leaves in certain Malaysia palms. Biotropica 11

(2): 126.

| 1980

Balanophora elongata (Balanophoraceae) new to the Malay Peninsula. Mal.

Wat. J: 3342) 129-832.

A new species and new section of the genus Freycinetia (Pandanaceae) from

New Caledonia. Pac. Sci. 33(2) : 149-151.

Additions to the Malayan Flora VIII. The Malaysian Forester 43 (2) : 244-262

The vegetation and plant communities of Pulau Balambangan, Sabah, East

Malaysia. J. Roy. Asiat. Soc. Mal. Br. 53 (1) : 68-89.

(with W.R. Philipson et al.). The systematic position of Aralidium Miq. - A

multi-disciplinary study. Taxon 29 (4) : 391-416.

Rediscovery of Thismia clavigera (Becc.) F.V.Muell. (Burmanniaceae). Blumea

26 : 419-425.

(with M. Ratnasabapathy). Thismia clavigera (Burmanniaceae), a saprophyte

new to the Malayan flora, and notes on other Thismia species in the Peninsula.

Sains Malaysiana 9 (2) : 209-214.

A nomenclatural change in Malaysian Araliaceae : Eleutherococcus malayanus

(Henderson) comb. nov., instead of Acanthopanax malayanus Henderson.

Malaysian Forester 43 (3) : 395.

The significance and future prospect of wild orchids in Malaysia. In Proceedings

of 3rd ASEAN Orchid Congress, Kuala Lumpur. pp. 132-144.

1981

The need for Natural History Museum reference collection research facilities in

Malaysia. In H.M. Collier (Ed.). Proceedings of Silver Jubillee Congress on

Science and Technology in Resource Development, Malaysian Scientific Association,

Kuala Lumpur. pp. 199-206.

(with K.L. Huynh). On a new subsection of Pandanus section Cauliflora

(Pandanaceae) with paniculate inflorescence structure, distinctive leaf anatomy

and pollen morphology. Bot. J. Linn. Soc. 83 : 213-220.

1982

New Guinea Pandanaceae : first approach to ecology and biogeography. In J. L.

Gressitt (Ed.). Biogeography and Ecology of New Guinea I : 401-436.

Additions to the Malayan Flora no.5. A new Geostachys (Zingiberaceae) from

Gunung Ulu Kali, Pahang, Malaysia. Malays. J. Sci. 6A : 75-81.

The summit flora of Gunung Ulu Kali, Pahang, Malaysia. Malays. J. Sci. 6A:

75-81.

Nomenclature of Joinvillea (Joinvilleaceae). Gard. Bull. Sing. 34(2) : 223-225.

In Memoriam : Dr. Monte Gregg Manuel, Bryologist, 1947-1981. Gard. Bull.

Sing. 34 (2) : 227.

New and noteworthy Malaysian Myrsinaceae I. The Malaysian Forester

45(1) : 101-121.

A new combination for Barclaya kunstleri of the Nymphaeaceae. Gard.

Bull. Sing. 35 (1) : 69-71.

Two new species of Pandanus from Gunung Mulu National Park. In A.C. Jermy

& K.P Kavanagh (Eds.) Notulae et Novitae Muluenses. Bot. J. Linn. Soc. 85

: 31-35.

1983

(with K.L. Huynh). The identity, affinities and staminate floral structure of

Pandanus pendulinus Martelli (Pandanaceae). Gard. Bull. Sing. 35 (2) : 199-

207.

(Book Review). Handbooks of the Flora of Papua New Guinea. Vol. II (E.E.

Henty, Ed.). Gard. Bull. Sing. 35(2) : 231-232.

Studies in Malesian Pandanaceae 19. New species of Freycinetia and Pandanus

from Malesia and Southeast Asia. J. Arnold Arb. 64 92) : 309-324.

Revisio Pandanacearum Part II. Pandanus subgenus Rykia (De Vr.) Kurz (section

Gressitia, Rykiopsis and Solmsia excepted). Fedn. Mus. J. n.s. 28: 1-100.

(Review). K. M. Kochummen, Effects of elevation on vegetation on Gunung

Jerai, Kedah. Mal. Nat. J. 36 (4) : 289-291.

Studies in Malesian Rutaceae III. Melicope suberosa, a new species and new

generic record for the Malayan flora. Gard. Bull. Sing. 36 (1) : 93-100.

(with D.T. Jones). Two new Rutaceae from Genting Highlands. Nature

Malaysiana 6 (3) : 4-11.

100

(Review). P.S. Ashton, Dipterocarpaceae. Fl. Males. 9 (2). Taxon 32 (4) :

694-696.

A guide to collecting Pandanaceae (Pandanus, Freycinetia and Sararanga). Ann.

Missouri Bot. Gard. 70 : 137-145.

Some new and critical Pandanus species of Subgenus Acrostigma I. Supplement

to Revisio Pandanacearum. Gard. Bull. Sing. 36 (2) : 205-212.

1984

(with R. Kiew). A new species of J/lex (Aquifoliaceae) from Gunung Ulu Kali,

Pahang, Peninsular Malaysia. Mal. Nat. J. 37 : 193-198.

Additions to the Malayan Flora IX. New and noteworthy species, wild and

introduced in Peninsular Malaysia and Sarawak. Mal. Nat. J. 37 : 185-191.

Pandanus from Ok Tedi region, Papua New Guinea, collected by Debra Donoghue.

Economic Botany 38 (3) : 304-313.

A field key and enumeration of the species of Pandanaceae in Gunung Mulu

National Park. In A.C. Jermy (Ed.). Studies on the flora of Gunung Mulu

National Park, Sarawak, Forest Department, Kuching. pp. 77-83.

A preliminary survey of Rutaceae of Gunung Mulu National Park. In Jermy,

A.C. (Ed.). Studies on the flora of Gunung Mulu National Park, Sarawak. pp.

137-143. Kuching Forest Department Headquarters.

A new species of Pandanus (Pandanaceae) from New Caledonia, with a synopsis

of Pandanus sect. Veillonia. Bull. Mus. Natn. Hist. J. (Paris), ser. 4,6 sect. B,

Adansonia | : 57-62.

1985

Studies in Malesian Rutaceae III. A conspectus of the genus Glycosmis Correa.

Proc. Acad. Nat. Sci. Phila. 137 (2); 1-27.

New and noteworthy paleotropical species of Rutaceae. Proc. Acad. Nat. Sci.

Phila. 137(2) : 213-228.

On the second collection of Pandanus halleorum. Kew Bull. 40 (2) : 287-289.

Forest by the sea. Creatures of the mangroves. In National Geographic Society:

A resource guide to the 1986 National Geographic Specials (TV). National

Geographic Society Educational media Division, Washington, DC. p. 8-9.

1986

The human equation in species extinction. Letter to Editor. Bioscience 36(8) :

524-525. |

The genus Pandanus (Pandanaceae) on Christmas island, Indian Ocean. Gard.

Bull. Sing. 39(2) : 193-202.

101

Yellow stars : a brief introduction to the Hypoxidaceae. Herbertia 42 : 51-57.

1987

New taxa of Pandanus (Pandanaceae) from Malesia and Papuasia. Blumea 32 (2) :

427-441.

Collecting botanical specimens : do’s and don’t’s. Orang Asli Studies Newsletter

(Dartmouth College) 6 : 5-11.

(with A. Weber). A new species of Phyllagathis (Melastomataceae) from the

Endau-Rompin proposed national park, Malaysia. Prod. Acad. Nat. Sci. Phila.

139°: 307-313:

1988

Notes on the genus Labisia Lindl. (Myrsinaceae). Mal. Nat. J. 42 (1) : 43-51.

New and noteworthy Malesian Myrsinaceae II. Emblemantha, a new genus from

Sumatra. Proc. Acad. Nat. Sci. Phila. 140 (2) : 275-280.

1989

(with T.G. Hartley). Reduction of Pelea with new combinations in Melicope

(Rutaceae). Taxon 38 (1) : 119-123.

Myrsinaceae. In Ng, F.S.P. (Ed.). Tree Flora of Malaya. Vol. 4 : 264-284.

New and noteworthy Malesian Myrsinaceae, III. On the genus Ardisia in Borneo.

Proc. Acad. Nat. Sci. Phila. 141 : 263-306.

New and noteworthy Malesian Myrsinaceae, IV. Two new species in Embelia

Burm. f. and Maesa Forssk. from Borneo. Proc. Acad. Nat. Sci. Phila. 141 :

307-311.

1990

Studies in Malesian Myrsinaceae 5. Additional new species of Ardisia Sw.

Proc. Acad. Nat. Sc. Phila. 142 : 21-58.

1991

The genus Loheria Merrill (Myrsinaceae). Micronesica 24 (1) : 65-80.

(with E.W.M. Verheij). Citrus. In PROSEA 2 : 119-126.

Pandanus parkinson. In PROSEA 2 : 240-243.

1992

The New Guinea species of Pandanus section Maysops St. John (Pandanaceae).

Blumea 37 (1) : 31-61.

New evidence for the reconciliation of floral organisation in Pandanaceae

with normal angiosperm patterns. In P. Baas et al. (Eds.). The plant

102

diversity in Malesia. Leiden pp. 33-55.

New and noteworthy Malesian Myrsinaceae 6. Revision of the genus Hymenandra

A. DC. Gard. Bull. Sing. 43 : 1-17.

Myrsinaceae as an example of plant diversity in Malesia with special reference to

the species in Borneo. Mal. Nat. J. 45 (1-4) : 230-237.

A note on the repaged reprint by William Griffith of Jack’s Description of

Malayan plants (Calcutta, 1843). Bartonia 57 : 32-35.

(Book review). Orchids of Java, by J.B. Comber and Orchids of the Solomon

islands and Bougainville by B.A. Lewis & P.J. Cribb. Micronesica 25 (1) : 133-

136.

(Book review). Orchids of Borneo. P.J. Cribb (Ed.). Micronesica 25 : 219.

A revision of the genus Tetrardisia Mez (Myrsinaceae). Malay. Nat. J. 46: 13-

24.

Systellantha, a new genus of Myrsinaceae from Borneo. Malay. Nat. J. 46: 13-

24.

Pee,

New and noteworthy Malesian Myrsinaceae 6. Scherantha, a new subgenus of

Ardisia. Pac. Sci. 47 (3) : 276-294.

Studies in Malesian Pandanaceae 21. The genus Pandanus in Borneo.

Sandakania 2 : 35-84.

Reduction of the genus Parardisia (Myrsinaceae). Nordic J. Bot. 13 (1) : 55-57.

1994

(with P.A. Cox & K.L.Huynh). Evolution and systematics of Pandanaceae (in

press).

Revisio Pandanacearum III. Pandanus section Cristata Martelli (Synonym,

Jeanneretia) of subgenus Kurzia (in press).

Supplement to the Rutaceae in Peninsular Malaysia (in press).

103

A list of taxa named in honour of Dr. B. C. Stone

Pelea stonei Degener & I. Degener, Phytologia 19 (1970) 315

(Rutaceae)

. Cryptocoryne stonei Rataj, Studie CSA V, 3 (1975) 95

(Araceae)

. Pandanus sect. Stonedendron Huynh, Bot. Jahrb. 97,1 (1976) 91

(Pandanaceae)

. Marcgravia stonei J.F. Utley, Brenesia 9 (1976) 52

(Marcgraviaceae)

. Arthrophyllum stonei A.L. Lim, Mal. Forester 43,2 (1980) 263

(Araliaceae)

. Pterisanthes stonei Latiff, Fed. Mus. J. n.s. 27 (1982) 51

(Vitaceae)

. Phyllagathis stonei A. Weber, Pl. Syst. Evol. 157, 3-4 (1987) 192

(Melastomataceae)

. Cyrtandra stonei B.L. Burtt, Edinburgh J. Bot. 47,3 (1990) 229

(Gesneriaceae)

is TANT batt oe RRA ee

‘

Citrus Fruits of Assam: A New Key to Species, and Remarks on

Citrus assamensis Bhattacharya and Dutta, 1956

BENJAMIN C. STONE

Botanical Research Institute of Texas

509 Pecan St., Fort Worth, TX 76102

USA

Abstract

The revision of Citrus in Assam by Bhattacharya and Dutta, 1956, is a work on citriculture and citrus

taxonomy the importance of which is not limited to the Asean region; it contains a new species, C. assamensis,

and a key to all the Assam citrus, as well as full descriptions of all taxa including the floral features. The utility

of this work is enhanced by illustrations but constructional errors in the key have prevented its correct and

effective use. A new key expressing the authors’ intentions, as well as remarks on Citrus taxa and relationships

and notes on the typification of C. assamensis are presented in this paper.

Introduction

S.C. Bhattacharya and S. Dutta published their important study of Assam

Citrus “Classification of the Citrus Fruits of Assam” (1956) in Monograph 20 of

the Indian Council of Agricultural Research. It is a major work in several important

respects, as it includes not only full descriptions, including floral features, of all

taxa, but it provides numerous vernacular names, a key to the species, and the

diagnosis of a newly proposed species, C. assamensis. This contribution is thus

valuable both to agriculture and to botany. It is also noteworthy for its adoption

of several Citrus species which were suppressed (regarded as synonyms or hybrids)

by Swingle in his classic revision of the Orange subfamily Aurantioideae (1943,

1967). While the authors generally follow Swingle’s classification, they accept

Six species not recognized by Swingle among the 17 species recorded for Assam.

These six species are C. jambhiri, C. karna, C. limetta, C. nobilis, C.

megaloxycarpa, and their new C. assamensis.

Unusual for agricultural literature devoted to Citrus, Bhattacharya and Dutta

include full technical data on the flowers of all taxa treated, omitting information

only when their data was incomplete. Thus they provide fuller descriptions than

is often the case, and the rather liberal use of illustrations also assists the botanist

or agriculturist seeking information or identification. As experienced agriculturists

their opinions are valuable, and their field knowledge, much gleaned in tribal

villages with citriculture little influenced by modern commerce, presented in a

generally judicious balance with botanical and “varietal” (cultivar) data, enhances

the work.

Taxonomically Bhattacharya and Dutta are much closer in their taxonomic

outlook to the conservative Swingle system than to the much more elaborate and

finely divided system of Tanaka (1954). Reece (1967) in his editing of Swingle’s

106

work hardly modified it, though Hodgson (1965) accepted several more species

than did Swingle. In this respect, Bhattacharya and Dutta agree rather closely

with Hodgson. The controversial taxa of Citrus are virtually all cultivated, and

differences in classification therefore particularly involve citriculturists.

Key to Species

One of the worthy features of the Bhattacharya & Dutta study is, rightly, the

inclusion of a key to the species. For practical purposes, such a key is always

desirable, whether for agricultural or purely botanical purposes.

However the reader attempting to use this key soon encounters almost

insuperable difficulties due partly to some peculiarities of its construction. In

effect it is not so much a key as a collection of short, partial diagnoses. It is not

possible at this date to understand why the key was prepared in this manner. ©

Though in part dichotomous, the key later on becomes polychotomous. In effect

the key is difficult to use at best and often enough is unworkable.

Because of this, it seemed beneficial to prepare a new key, based on the same

data that Bhattacharya and Dutta used, which would be strictly dichotomous and

effective. Without necessarily accepting all the taxa of Citrus that they recognize,

any botanist or citriculturist will benefit from having a functional key permitting

identification of the taxa as construed by these authors. Furthermore, a usable key

may stimulate collecting and further investigations of Citrus both in Assam and

perhaps elsewhere.

Correction of the Key.- An analysis of the key shows some functional couplets.

The initial couplet divides the genus into the two groups recognized by Swingle

as “Eucitrus” and “Papeda” (Bhattacharya & Dutta, 1956, p. 11, postscript).

These two groups are subgenera in Swingle’s classification. Within subg. Papeda

the couplet N/NN separates Citrus ichangensis from C. latipes; subsequently

however there is a solitary lead O, with a subordinate P, rather than couplets (P

describes but does not fully discriminate C. macroptera). Within subg. Citrus

(“Eucitrus”), C. medica is discriminated, but lead B is one of a trichotomy B/BB/

BBB. Under BB, C. limon is discriminated, but leads D, E, G, and H are solitary,

though there is a couplet F/FF. Lead E seems to indicate that the peel of both C.

limetta and C. megaloxycarpa is loose, which is not true. In the last group under

BBB the first “couplet is another trichotomy K/KK/KKK, each ending with a

species; yet KKK is followed by 3 more leads, L, LL, and M (L and LL do not

form a true couplet, as the half-couplet which should have been marked MM is

annexed as the latter part of LL). Moreover C. grandis (from lead M) can be

compared to C. aurantifolia (from LL), but lead L is inaccessible from the

preceding KKK. Faults such as these prevent a user from successfully operating

this key in a manner which is both baffling and misleading.

107

TABLE I

Species sequence in KEY Species sequence in TEXT

C. medica

C. limon

C. jambhiri

C. karna

C. reticulata

C. indica

C. limetta

C. megaloxycarpa

C. aurantium

C. sinensis

C. assamensis

C. nobilis

C. aurantifolia

C. grandis

C. latipes

C. macroptera

C. medica

C. limon

C. jambhiri

C. karna

C. aurantifolia

C. limetta

C. reticulata

C. nobilis

C. indica

C. sinensis

C. aurantium

C. grandis

C. megaloxycarpa

C. ichangensis

C. macroptera

C. assamensis

Sequence of species in the classification.- The sequence of species in the key

is not the same as that in the following systematic treatment. This is not necessarily

very important, but it is of interest to compare the two sequences (Table 1).

The TEXT sequence is basically like the sequence in Swingle (1943, 1967),

though it differs in two ways - first by the insertion of species not accepted by

Swingle (these are in positions which at least implicitly indicate their taxonomic

relationship); and second by the transposition of some species (e.g. in Swingle’s

arrangement, C. aurantium is no. 4, C. sinensis is no. 5, and C. reticulata is no. 6;

in the Bhattacharya and Dutta sequence, C. reticulata with the immediately

following C. nobilis - not in Swingle - precedes C. aurantium and C. sinensis,

which are in reverse order. The pomelo, C. maxima, is in position no. 7 in both

treatments, but in Bhattacharya & Dutta it is followed by C. megaloxycarpa (not

in Swingle). Finally, the members of subg. Papeda come last. The last species in

Bhattacharya & Dutta’s sequence is C. assamensis, which is clearly asserted

NOT to be a member of subg. Papeda; it ought to be inserted just after C.

megaloxycarpa, its nearest (putative) relative. In fact in the key, C. assamensis is

found between C. sinensis and C. nobilis, yet the authors take pains to show it is

not closely related to C. hystrix, which is certainly a member of subg. Papeda. By

108

inference one may conclude that they thought that C. assamensis was related

closely to either C. sinensis or C. nobilis, or both. In a short chapter on hybrid

forms, the authors describe a cultivar called “hash-khuli’ which, they suggest,

may be a hybrid of C. assamensis and C. maxima (C. grandis). In my opinion, C.

assamensis may be related to C. megaloxycarpa and to C. maxima, evidence

being the similarity of leaf and petiole and in the former also the purplish corolla.

In fact, Singh & Nath (1969) relegated C. assamensis to synonymy under C.

maxima, or more precisely to one of the synonyms, C. megaloxycarpa var.

pennivesiculata.

REVISED KEY TO CITRUS OF ASSAM

1. Pulp-vesicles lacking acrid oil droplets. Petiole unwinged, or marginate, or with a small to moderate wing

never much more than 1/4 as long as the blade and less than 1/3 as wide. Stamens more or less connate or

polyadelphous.

Subg. CITRUS

2. Petiole unwinged, not articulated. Petals tinged purplish. Flowers dimorphic, staminate and perfect.

Fruit large, usually elongated, yellow, the rind thick, hard, usually somewhat sweet or palatable.

CTT: ..<.-10s050secxcnesannahsneBidiade naga iiade seat ak esiteas tate het haiga tna nal Citrus medica L.

2’ Petiole unwinged, moderately winged, or noticeably winged, and articulated.

3. Petals noticeably purplish-tinged. Fruit greenish to yellow, ellipsoid to ovoid or elongated, often

nippled at the end.

4. Rind rather bumpy or warty, moderately to weakly adherent. Carpels rather easily separable.

Petals 12-21 mm long. Stamens 21-28 per flower. Fruit up to 9.5 cm diameter. ROUGH

LEMON 6.....<:occssondepcnnssibcnsvguaptliatessadigd us ledinid ances ttptn taint A ssth eedt ines mien Citrus jambhiri Lush.

4' Rind relatively smooth, rather strongly adherent. Carpels strongly adherent. Petals 18-29 mm

long. Stamens 19-49 per flower.

5. Tree flowering all year round. Fruit rind rather thin, 3-10 mm thick, firm, not sweet, pulp

and juice sour. Fruit subglobose to oblong, nippled, up to 9 cm diameter. Stamens 26-49

per flower. Petals 18-29 mm long. LEMON. ..............::000+ Citrus limon (L.) Burm. f.

5' Tree flowering but once a year, or only one major crop per year. Rind moderately thin,

5-7 mm thick, or thick to very thick, up to 35 mm. Stamens 19-42 per flower.

6. Rind 10-35 mm thick. Fruit rather to quite large, often 12-14 cm diameter, sometimes

smaller (7-12 cm diam.), not scented with ginger-like or eucalyptus-like odor.

7. Fruit usually 7-12 cm diameter, rind soft and spongy, sweet, smooth or slightly

pitted. Petiole wing 8-16 mm long, 2-3 mm wide. Stamens 24-29 per flower.

KARNA. .........:-cosconsnvnachonsceginashecthcibedstdalies sik iaaan aan Citrus karna Raf.

7' Fruit usually 10-14 cm diameter, rind firm, leathery, or brittle, not sweet. Petiole

wing usually 10-30 mm wide (rarely only 5-10 mm wide), about 1/7 to 1/4 as

long as blade. Stamens 19-38 per flower. AMILBED.| ............ccccesceecsesseecseeeeeeees

resaenieorsdptignbassititnesinldeddessiiiidedsanan name Citrus megaloxycarpa Lushington

6' Rind 6-7 mm thick, leathery. Fruit 7-10 cm diameter, subglobose to almost turbinate.

Rind oil with gingery or eucalyptus scent. Rind light yellow or pale greenish, almost

smooth. Petiole wings somewhat broad, obovate, about 1/4 as long as wide as the

blade. Stamens 26-42 per flower.

ADA-JAMIR. ....:isujuucds>osctaasamstooiaiene Citrus assamensis Bhattach. & Dutta

3' Petals pure white. Fruit green, yellow, orange, or reddish, not lemon-like except in some forms of

C. aurantifolia.

109

8. Fruit with a loose, easily detached or separable rind. Cotyledons green. Leaves often rather

narrowly elliptic or oblong-elliptic or somewhat rhombic.

9. Seeds large orbicular flattened, 14-15 mm long, smooth. Fruit small, 2.5-4 cm diameter,

oblate, the rind red, with scanty, slimy, very sour juice.

Filaments pubescent. Leaf apex somewhat caudate. Petiole scarcely winged.

ee Bk 2 2 EERE ahs Ge aie Ee Citrus indica Tanaka

9' Seeds small, medium, or large (to 16 mm long), somewhat cuneate, slightly rough, or

somewhat clavate with fin-like projection of testa. Fruit orange to red, often oblate, sour

to quite sweet, juice not slimy, rather copious and palatable. Filaments glabrous. Leaf apex

not caudate. Petiole wing small and narrow (or nil) or broad and obovate-oblanceolate.

10. Petiole wing conspicuous, broadly spathulate, up to nearly half as long as the blade,

but only one-fourth as wide. Stamens mostly 22-32 per flower. Rind 5-9 mm thick.

Pe a eS oe pu ettstd scar esv den cedendaanasebaseetnseneshecebiccensetennd Citrus nobilis Lour.

10' Petiole wing small, very narrow, or nil. Stamens 14-24 per flower. Rind about 5 mm

Bg | Et 2, pan Ae ea ES ea ee ee Ca ee ee Citrus reticulata Blanco

8' Fruit with tightly adherent rind. Seeds with white cotyledons. Leaves various in shape, but not

rhombic.

11. Petiole short and virtually wingless. Rind light yellow, glossy. Fruit subglobose, lemon-

like. Pulp vesicles whitish. Juice sweet. Chalazal cap ochre-yellow. SWEET LIME......

os hae © pis See ECE Sea ES 9S ee Rae ME tpl Pek Citrus limetta (Risso) Lush.

11' Petiole winged, wings narrow to broad.

12. Petiole wing small, narrow, up to 12-15 mm long and 5-10 mm wide.

13. Fruit 4-5 cm diameter, subglobose to oblong. Rind very thin. Pulp vesicles

greenish to whitish. Juice sour. Chalazal cap brown. LIME...............0.....:.:.200

TE cad oie creel a egw set gc comin de oti Citrus aurantifolia (Christm.) Swingle

13' Fruit 5-9 cm diameter, subglobose. Rind 4-7 mm thick, yellow to orange. Pulp

vesicles yellowish to orange. Juice usually sweet. Chalazal cap “Indian red.”

Petiole wing 10-20 mm long, 1-3 mm wide. SWEET ORANGE..........0..0.........

ene <2 OS hE 5. SOR Ass SANE se, Pee ey BER a Pere Citrus sinensis L.

12' Petiole wing larger, broader, up to 7 cm long and 5 cm wide.

14. Inflorescence glabrous, few-flowered. Stamens 22-24 per flower. Fruit 6.5-

8.5 cm diameter, deep orange to scarlet. Rind 7-10 mm thick. Pulp vesicles

yellow to orange. Juice very sour and somewhat bitter. Seeds 12-16 mm

long. Chalazal cap “Indian red.” SOUR, BITTER, or SEVILLE ORANGE

SA a se acne Silane bei ig lal EE ene ot Citrus aurantium L.

14' Inflorescence racemose-glomerate, pubescent, with up to about 10 flowers.

Stamens mostly 30-40 (rarely as few as 22) per flower. Fruit often 10-15 cm

diameter, green to yellow, sometimes tinged pinkish, the rind often 10-20

mm thick or more. Juice rather bland, or mildly tart or sweet. Seeds 15-23

mm long. Chalazal cap brown or reddish-brown. POMELO.......................

NEE Sa RO SESE TRAE PED Citrus maxima (L.) Merr.

l' Pulp vesicles containing acrid oil droplets. Petiole long, broadly winged, the wing almost as wide or as wide

as the blade, and from half as long to longer than the blade. Stamens usually free, rarely coherent.

Subgenus PAPEDA

15. Leaf apex acuminate-caudate. Stamens coherent. Pulp vesicles globose to obovoid, white. Juice sour

and scanty. Seeds numerous, 12-20 mm long, 10-28 mm wide. Chalazal cap broad, brown. ..............

I Od, ons on cPnvn tanltlinnpnmenacdy pul divedinnnalbcadieng Citrus ichangensis Swingle

15'. Leaf apex not caudate. Stamens free. Seeds somewhat smaller. Chalazal cap light red.

110

16. Leaf apex acute to subcaudate. Fruit about 5-10 cm diameter. Style 4-8 mm long. ..................0.

sdasectaenatniesyecsosenteasshunavaeteduasdsiassay abiipns Giallasnerae- sale Akane a ese aie i Citrus latipes Swingle

16' Leaf apex rounded to obtuse. Fruit usually over 10 cm diameter. Style 1-4 mm long. MELANESIAN

PAPEDIA. .:0is0-sns-nsi ag sugenh outatioeyanetiveaps.gqaseess aaah dada nae ee Citrus macroptera Montrouzier

Vernacular Names

Bhattacharya and Dutta provide a wealth of vernacular names applied to

various forms and cultivars of the Citrus species in Assam. For completeness

these are repeated here; those capitalized are the “preferred’ names.

Citrus medica. (CITRON). Birajora; mithajora; soh-manong; bakol-

khowatenga; soh-madeh; jaara-jamir; tumehan-thor; haijange; naya-changney;

bhimra; mokari; mohalung; sutrung; madh-kunkur; madh-kakri; maulung; natterun.

Citrus jambhiri (ROUGH LEMON). Soh-myndong; soh-jalia; kata-jamir;

sinduri-nemutenga; mithu-tulia; nemu-tenga.

Citrus limon (LEMON). Naya-changney; pati-lebu; katajamuri; elachi-lebu;

soh-long; soh-synteng; pani-jamir.

Citrus karna (KARNA). Karna; soh-sarkar.

Citrus megaloxycarpa (AMILBED). Amilbed; bor-tenga; hukma-tenga; holong-

tenga; jama-tenga.

Citrus assamensis (ADA-JAMIR). Ada-jamir.

Citrus indica (INDIAN WILD ORANGEB). (No vernacular names recorded).

Citrus nobilis (KING ORANGE). Jeneru-tenga.

Citrus reticulata (MANDARIN). Sweet forms: soh-niamtra; soh-umkdai; naga-

santra. Sour forms: soh-siem; kapura-tenga.

Citrus limetta (SWEET LIME). Mitha-kagzhi; mou-muri; soh-bakhlein.

Citrus aurantifolia (LIME). Kagzhi.

Citrus sinensis (SWEET ORANGE). Soh-niangriang.

Citrus aurantium (SOUR ORANGE). Karun-jamir; gondh-kuntra.

Citrus maxima (POMELO). Rebab-tenga; soh-myngor; mat.

Citrus ichangensis. (No vernacular names recorded).

Citrus latipes. Soh-kympho-shrieh.

Citrus macroptera (MELANESIAN PAPEDA). Sat-kara; tith-kara.

111

Note On Typification of Citrus assamensis

In the protologue, Bhattacharya and Dutta do not specifically mention a

“type” but they remark (1956, p. 787) “The original specimen had . . . been sent

to Kew Herbarium...” and it may be assumed that by this they indicated the

type. I have examined the relevant material at Kew and choose the fuller sheet as

the lectotype. The label of this specimen reads as follows: “Herbarium, Citrus

Fruit Research Station, Burnihat, Assam. - Fam. Rutaceae. Citrus assamensis

Dutta & Bhattacharya, sp. nov. Vernac. name, ada-jamir. - Plant medium-sized,

very thorny, stout; leaves coriaceous, glossy; petiole spathulate, margin revolute;

flower purple; fruit spherical smooth; aroma eucalyptus smell; pulp free from oil

droplets.” The locality data is: ASSAM, Karimganj, alt. about 300 ft., 6 November

1938, S. Dutta & S.C. Bhattacharya no. 2365. K! (lectotype).

Because of the significance of this species, and to provide an example of the

thoroughness of the descriptions employed by Bhattacharya and Dutta, the original

description is suggested as a model of its kind. From the original publication, the

authors note is here quoted. “The specimen of ada-jamir as it is known locally

has been collected from an interior village in Karimganj subdivision of the

district of Cachar, Assam . . . identical specimens have also been found to occur

particularly in Sylhet, North Cachar hills, and Khasi hills. It is sporadically

grown in home gardens.” (Note that Sylhet now is within Bangladesh).

Bhattacharya & Dutta showed that C. assamensis could not be assigned to C.

hystrix (a table of differences is presented) but nevertheless referred to subgenus

Papeda which includes C. hystrix. However, the absence of acrid oil droplets in

the pulp-vesicles must exclude C. assamensis from subg. Papeda.

More probably C. assamensis is closely related to the Pomelo, C. maxima (C.

grandis, C. decumana), and to the Amilbed or Sour Pomelo, C. megaloxycarpa -

with which the Ada-jamir shares its purplish petals and sour juice. The placement

of C. assamensis in the original key (of Bhattacharya & Dutta) between C.

sinensis and C. nobilis apparently does not indicate relationship.

The “ginger or eucalyptus odor” specified for C. assamensis is noted as very

characteristic; “the fruits are valued locally for their peculiar aromatic flavour

and intense sour juice. The aroma of the rind approaches to eucalyptus smell but

people characterize it to be similar to that of the ginger, Zingiber officinale, and

hence the name “ada-jamir” (ada = ginger; jamir = citrus). It is also called Soh-

sying (soh = soft; sying = ginger) in the Khasi hills of Assam.”

Postscript

This paper clarifies the key structure in the work by Bhattacharya & Dutta;

clarifies the typification and posited relationship of their new species Citrus

112

assamensis; and commends their descriptive work to botanists and agriculturists

as a suitable model. The status of C. assamensis remains controversial; renewed

study of it is recommended. Good specimens of C. indica, C. assamensis, C.

ichangensis, C. latipes, and C. jambhiri are worth obtaining in the Assam area.

References

Bhattacharya, S.C., and S. Dutta (1956). Classification of the Citrus Fruits of

Assam. Indian Council of Agricultural Research. Scientific Monograph No.

20. 110 pp.

Hodgson, R.W. (1965). Taxonomy and nomenclature in Citrus fruits. International

Organization of Citrus Virologists, Proceedings, 2: 1-7.

Reece, P.C. See Swingle, (1967).

Swingle, W.T. (1943). The Botany of Citrus and its wild relatives of the Orange

Subfamily. pp. 129-474, in Webber, H.J. and L.D. Batchelor, eds., The Citrus

Industry, I: History, World Distribution, Botany, and Varieties. University of

California Press, Berkeley.

— (1967). Ibid. Pp. 190-430, in Reuther, W., H.J. Webber, and L.D. Batchelor,

eds., The Citrus Industry, I: History, World Distribution, Botany, and Varieties.

Revised edition; Botany revised by P.C. Reece. University of California Press,

Berkeley.

Tanaka, T. (1954). Species Problem in Citrus. Revisio Aurantiacearum IX.

Japanese Society for the Promotion of Science, Ueno, Tokyo, pp. 1-152.

Additional Notes on the Genus Glycosmis Correa (Rutaceae)

BENJAMIN C. STONE

Botanical Research Institute of Texas,

509 Pecan Street,

Fort Worth, TX 76102

USA

and

Philippine Plant Inventory,

Herbarium, National Museum,

Box 2659, Manila,

Philippines

Abstract

Updating of the Conspectus of the genus Glycosmis of Stone, 1985 is required as certain additions and

corrections have to be made, and comments on recent publications that have dealt with this genus are required

in the context of a monographic perspective. Three new proposed species are discussed, as well as various

nomenclatural and taxonomic questions.

Introduction

Since the appearance of the author’s “Conspectus of the Genus Glycosmis”

(Stone, 1985) two new taxonomic contributions have been made to the genus

(Tao, 1984; Huang, 1987) dealing chiefly with Chinese species. Also some

anomalies have been detected in the Conspectus that require some taxonomic or

nomenclatural adjustments. This paper is therefore intended as an extension of

the Conspectus, to bring recent information and conclusions into the monographic

perspective that is required.

Comments on three proposed new species of Glycosmis

1. Glycosmis motuoensis D.D. Tao

Acta Botanica Yunnanica 6(3): 285-287, 1984; ex Huang, Flora Xizangica 3: 30, 1986.

This tentatively accepted species seems to be closely allied to G. cyanocarpa

var. cymosa Kurz, differing in (i) petals glabrous but very slightly ciliolate, and

(ii) lateral veins somewhat more numerous and slender. These differences are

minor at best, but may prove to be correlated with others when better material is

available. If so, the taxon may stand. However, the status of G. cyanocarpa itself

requires review. Currently, several varieties are distinguished within G. cyanocarpa

(B1.) Sprengel, including var. cymosa Kurz. This variety has been regarded as a

distinct species by Narayanaswamy (1941), who made a new combination (G.

cymosa (Kurz) Narayan.). Unfortunately, as previously discussed in the

Conspectus, Narayanaswamy conceptualized this species as jointly comprising

the taxon G. longifolia Tanaka, which had been published earlier; this name

114

should therefore have been adopted by Narayanaswamy for his species. Despite

this, the combination G. cymosa is validly published when G. longifolia is

conceptually excluded. In other words, Narayanaswamy made a viable binomial

with his combination of Kurz’s varietal epithet, but incorrectly applied it to the

taxon he recognized. At any rate, it is to Kurz’s variety that Tao’s G. motuoensis

seems most similar, and not to G. longifolia (thus not to G. cymosa sensu

Narayanaswamy!).

The type collection of G. motuoensis (Qinghai-Xizang Expedition no. 74-

4540), was kindly made available on loan from Dr Tao and the South China

Institute of Botany, Kunming, P.R.C. The type collection is from Medog, Tibet,

at an altitude of about 800 m. It is thus quite possible that the plants formed part

of a scattered population or series of populations that extend into Sikkim Himalaya.

Plants of this series in other herbaria (especially in India) may have been previously

identified as G. cyanocarpa var. cymosa.

The material seen is in an early state of flowering before anthesis; the floral

parts are therefore smaller than their potential dimensions, which are estimated as

being about twice those evident in the specimen. The stamens, although described

as subequal, are definitely alternately longer and shorter, as is normally the case

in most species of Glycosmis. The anthers do have connective glands, although

they are smaller than the apical gland. Little else can be added to Tao’s diagnosis,

though it may be noted that the leaflet undersurfaces are pale grayish-green with

a faint pinkish-brown tinge. More significantly, the dissection of the flowers

shows that the ovary may be either 4- or 5-merous. Ovary locule number is an

important character in the genus and should always be determined. In my synoptic

key, G. motuoensis would key out to Group D, next to species 7 (G. cyanocarpa).

The protologue states that G. motuoensis is related to G. erythrocarpa Hay.

(of Taiwan), probably on the basis of the shared trifoliolate character; but that

relationship is not very close. For a fuller understanding of the proposed new

species it appears necessary to obtain good material with ripe flowers and fruits,

and to search for it also in adjacent areas (e.g. Bhutan and Nepal).

2. Glycosmis lucida Wall.

ex Huang, Guihaia 7(2): 119-120, 1987.

C.C. Huang has here validated Wallich’s nomen nudum and applied it to a

taxon conceptually identical to G. cyanocarpa var. cymosa Kurz. The synonymy

has already been stated by Kurz (1876) and Narayanaswamy (1941), who used

the binomial G. cymosa (Kurz) Narayan. However as mentioned above, because

Narayanaswamy included G. longifolia Tanaka (1928) in his concept of G.

cymosa, he should have adopted Tanaka’s existing name for it. G. longifolia

Tanaka is in fact conceptually identical to G. cyanocarpa var. simplicifolia Kurz.

Huang apparently typifies G. lucida Wall. ex Huang by a Grifith specimen

115

(no. 523 in K). This collection is mentioned by Kurz in his description of G.

cyanocarpa var. cymosa Kutz.

Huang excludes G. longifolia from his concept of G. lucida; therefore the

name G. cymosa (Kurz) Narayan. is the correct name to be adopted for this taxon.

The name G. cymosa Zipp. ex Span. (1841) is mentioned by Huang, but that

name, being a nomen nudium, has no validity and cannot pre-empt the usage of

the epithet ‘cymosa.’ If this taxon deserves species rank, it must be called

Glycosmis cymosa (Kurz) Narayan. (1941), and G. longifolia Tanaka (G.

cyanocarpa var. simplicifolia Kurz) must be excluded.

If however the taxon is regarded as having varietal rank under a different

species name, then Kurz’s varietal epithet must be retained. If the variety is

considered to be conspecific with (even if not convarietal with) G. cyanocarpa

var. simplicifolia then the binomial ‘G. cymosa cannot be used, and the earlier

valid name G. longifolia Tanaka must be used. This has the same lectotype as G.

cyanocarpa Var. simplicifolia.

Huang also mentions other synonyms, which with one exception are also

previously cited in the Conspectus (1985), viz. G. oxyphylla Wall., nom. nud.; G.

tetraphylla Wall., nom. nud.; G. pentaphylla var. yunnanensis Huang, Icon.

Corm. Sin. Suppl. 2: 159, 1983, nom. nud.

3. Glycosmis oligantha Huang

Guihaia 7(2): 122-123, 1987.

This validly published name (Latin diagnosis is provided) denotes a plant

from Guangxi; the designated type specimen is S.C. Chen 3153. It is described as

being similar to G. gracilis (Huang writes ‘G. gracilis Tanaka’ which is an

illegitimate name; the correct name is G. gracilis Stone, 1985). It seems however

even closer to G. craibii Tanaka. It apparently belongs to Group D (in the key in

the Conspectus), and according to this classification, G. craibii is better regarded

as a variety of G. puberula Lindl. which is found in Group E, as it generally has

unifoliolate, though sometimes 2-3-foliolate, leaves. Glycosmis oligantha is

described as having mainly 4-7 leaflets; leaflet number being variable, there is

probably no serious impediment to the implied relationships. More problematical,

however, is the question of ovary-locule number, which is omitted in Huang’s

diagnosis. If for example the ovary is 3-locular, a relationship to G. gracilis

would be more definitely supported; but if not, then a relationship with G.

puberula var. craibii (Tan.) Stone could be supported. The diagnosis of G.

oligantha also lacks a specification of the number and position of the anther

glands, features which are often useful in the taxonomy of Glycosmis.

Huang cites seven specimens, all from Guangxi, mostly from forest habitats

between 250 and 560 m altitude. For the present, judgment on the status of this

species is reserved.

116

Other taxa discussed by Huang (1987)

In the same paper in which he describes G. oligantha, Huang discusses

several other taxa for which he attempts some elucidation (he remarks that he. . .

“attempted to elucidate those of the doubted and confused species so far. . .

recorded from China.”) To facilitate reference to these, I take these up here in the

same order and add commentary as deemed appropriate.

Glycosmis cochinchinesis (Lour.) Pierre ex Engl.

Huang (1987) and Stone (1985) clearly agree on the delimitation, typification,

and synonymy of this species.

Glycosmis montana Pierre

F1. For. Cochinch. Pl. 285b. 1893.

This species is included under G. lanceolata (B1.) Sprengel in the Conspectus.

On further consideration this now seems incorrect, and I believe that Huang’s

interpretation of this taxon is correct or at least preferable. The short synonymy

he gives is in agreement, so far as it goes, with my concept, except for G.

tonkinensis Tanaka. The latter name, as cited (Tanaka ex Guillaumin, in Humbert,

Fl. Gen. Indoch. Suppl. 1: 629. 1946) is an invalidly published name, without

nomenclatural standing; it lacked a Latin diagnosis. It properly belongs as a

synonym of G. tetracronia Stone (1985), and is definitely not the same as G.

montana Pierre. The latter has a predominantly 3-locular ovary, the former a

predominantly 5-(or 4-) locular ovary, an important difference in this genus. If G.

lanceolata and G. montana are to be kept apart, a more exacting discrimination

should be undertaken to establish reliable differentiating features. The latter

seems to be the same taxon as G. greenei var. simplex Stone. The application of

the name G. lanceolata as used in the Conspectus is incorrect; research on this

matter is still in progress.

Glycosmis pseudoracemosa (Guill.) Swingle

Not. Syst. 2: 162, 1911.

Both Huang (1987) and Stone (1985) fully agree on the status and synonymy

of this species.

Glycosmis longifolia Tanaka

Bull. Soc. Bot. France 75: 709. 1928.

This taxon has figured in the earlier discussions (see above). By recognizing

this species, Huang implicitly accepts a division of the broad species concept of

117

G. cyanocarpa (B1.) Sprengel; i.e. he agrees with Tanaka that G. cyanocarpa var.

simplicifolia Kurz deserves separate status as a species. Moreover, he accepts

species rank for G. cyanocarpa var. cymosa, although the name G. lucida which

he applied to it is unnecessary; the correct name is G. cymosa (Kurz) Narayan.

Taxonomically, a subdivision of G. cyanocarpa sens. lat. is not at all

objectionable, although in the Conspectus it was retained in the broad sense (but

with ten recognized varieties!). If G. cyanocarpa is reinterpreted as a narrower

concept, it becomes a strictly Malesian taxon; the Indian, Sri Lankan, Thai,

Burmese, Chinese, and Tibetan populations would be excluded. This approach

would reinstate both G. longifolia and G. cymosa as distinct species. However,

such reinstatement does not fully satisfy the problem.

Huang cites the authority of G. longifolia as “(Oliver) Tanaka” but this is

incorrect. Tanaka (1928) clearly distinguishes when he is publishing a new

species and making a new combination. For example, see his paragraph on G.

esquirolii which he clearly designates: ‘G. esquirolii (Levl.) Tanaka, n. comb.’ In

contrast, for G. longifolia, the form is: ‘G. longifolia Tanaka, n. sp.’ This is a

perfectly clear indication of Tanaka’s nomenclatural meaning. Also, he distinctly

includes a Latin diagnosis- something not ordinarily provided for a new

combination.

It may also be noted that Oliver published the name ‘/ongifolia’ at the rank of

subvariety. This rarely used rank explains why Kurz’s taxon, var. simplicifolia,

bears a legitimate name. Oliver does not cite a holotype, but he does cite four

specimens (i.e. syntypes) to typify the subvariety. Jenkins’ Assam specimen has

already been designated as lectotype (Stone, 1985) for both subvar. longifolia

and G. longifolia. Tanaka states only “Type: Herb. Kew” without specifying a

particular specimen.

Glycosmis pentaphylla (Retz.) DC.

Huang (1987) and Stone (1985) essentially agree on the interpretation of this

historically confused and much abused name. However, Huang attributes the

contribution to Correa (Ann. Mus. Paris 6: 384, 1805), but Correa never actually

made this combination; it was first made, in fact, by De Candolle (Prodr. 1: 538.

1824), although De Candolle misapplied the name. Both Huang and Stone include

in G. pentaphylla the serrulate-margined plants originally named Limonia arborea

Roxb. (i.e. G. arborea (Roxb.) DC.).

Glycosmis esquirolii (Levl.) Tanaka

The synonymy for this species as given by Huang (1987) is correct, essentially

the same as that given in the Conspectus (1985), but I have also included G.

winitii Craib, of Thailand, as the same species.

118

Glycosmis parviflora (Sims) Little

Phytologia 2: 463. 1948.

Huang attributes this name to Kurz (he cites Journ. Bot. n.s. 5: 40. 1876).

However, Kurz did not mention Sims as author, nor did he specifically cite

Limonia parviflora as a basionym, though he gave a reference to Bot. Mag. t.

2416; in any case, Kurz did not accept the name, as he only mentions it as a

synonym (under G. citrifolia, just the opposite of our modern conclusion as to the

relative nomenclatural status of the two binomials). I do not believe that Kurz

effectively made this combination, and prefer to attribute it to Little, who most

explicitly did make it.

Glycosmis craibii Tanaka

Bull. Mus. Hist. Nat. Paris ser, 2, 2: 159, 1930.

Huang (1987) accepts this in the original conception of Tanaka; in the

Conspectus, I have placed it as a variety of G. puberula LindL., a relatively minor

difference of interpretation. We agree on the delimitation of the taxon.

Glycosmis craibii var. glabra (Craib) Tanaka

Lc. 1930.

Huang (1987) accepts this variety in Tanaka’s original sense. In the Conspectus,

I consider it rather as a synonym of G. ovoidea Pierre, a species not discussed by

Huang.

Corrections to the Conspectus

The correct name for G. lanceolata sensu Stone

In the Conspectus (p. 10) the name G. lanceolata is used for a rather broad

concept covering 11 synonyms. It is now increasingly apparent that, while the

taxon intended is comparatively homogeneous, the binominal G. lanceolata should

not have been applied to it. Current research suggests that the correct name for

this taxon is G. trifoliata (Bl.) Sprengel.

Correspondingly, the taxon denoted by G. lanceolata in the original sense is

probably best suggested by Narayanaswamy (1941) in his interpretation. The

relationship is probably near G. pentaphylla. A full resolution of this problem is

still required.

The correct name for G. sapindoides is G. macrophylla

By some inexplicable oversight, the name G. sapindoides Lindl. in Wall. ex

119

Oliv. was retained despite the clear citation of an earlier binomical in the synonymy.

The long usage of G. sapindoides would in another situation argue for its

conservation, but it must be relegated to synonymy. The correct name and

synonymy for this species are presented here, to serve as a replacement for

species no. 34, in Proc. Acad. Nat. Sci. Philad. 137: 18. 1985. In addition, two

new combinations are required, as shown below.

[34.] Glycosmis macrophylla (Blume) Miquel

Fl. Ned. Ind. 1, 2: 522, 1859. Type: Java, Tjanjor; Blume, L!

Syn. G. sapindoides Lindl. in Wall. ex Oliver, J. Linn. Soc. Bot. 5, Suppl. 2:38, 1861. Type: Penang,

Wallich cat. 6376, K!

Syn. G. cyanocarpa var. sapindoides (Lindl.) Kurz, J. Bot. 14: 34. 1876.

Syn. G. elata Ridley, J. Fed. Mal. St. Mus. 10: 130. 1920. Type: Malaya, Kelantan, Chaning woods;

Ridley, SING!

Syn. Sclerostylis macrophylla Blume, Bijdr. Fl. Ned. Ind. 3: 135. 1825. Type: Java, Tjanjor; Blume,

[34b.] Glycosmis macrophylla var. microphylla (Stone) Stone, comb. nov.

Syn. G. sapindoides var. microphylla Stone, Proc. Acad. Nat. Sci. Philad. 137: 18. 1985. Type: Flores

Island, Kostermans 22059, AAU!

[34c.] Glycosmis macrophylla var. australiensis (Stone) Stone, comb. nov.

Syn. G. sapindoides var. australiensis Stone, Proc. Acad. Nat. Sci. Philad. 137: 18. 1985. Type:

Western Australia, Augustin Island, Wilson 19775, PERTH!

References

Huang, Cheng-Chiu (1987). New Taxa and Notes on Glycosmis Corr. (Materials

for Chinese Rutaceae IV). Guihaia 7 (2): 115-124.

Kurz, S. (1876). On the species of Glycosmis. J. of Botany (ed. Trimen) n.s. 5:

33-40, tabs, 174-175.

Narayanaswamy, G. (1941). A Revision of the Indo-Malayan species of Glycosmis

Correa. Records of the Botanical Survey of India 14 (2): 1-iv, 1-72, 1-11, figs. 1-30.

Stone, Benjamin C. (1985). A Conspectus of the Genus Glycosmis Correa. Studies

in Malesian Rutaceae III. Proceedings of the Academy of Natural Sciences of

Philadelphia 137: 1-37.

Tanaka, T. (1928). Revisio Aurantiacearum I. Bulletin de la Societe de Botanique

de France 75: 708-715.

Tao, De-Ding. (1984). Taxa Nova Rutacearum. Acta Botanica Yunnanica 6(3):

285-287.

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Supplement to the Rutaceae in Peninsular Malaysia

B.C. STONE

Botanical Research Institute of Texas

509 Pecan Street, Fort Worth, TX 76102

USA

with account of Melicope

T.G. HARTLEY

Australian National Herbarium, CSIRO Division of Plant Industry,

P.O.Box 1600, Canberra, Australia 2601.

Abstract

Since the publication of the Rutaceae in the “Tree Flora of Malaya” (vol.1,1972), there have been a

number of changes both bibliographic and taxonomic which ought to be accommodated in this account. The

genus Terminthodia has been subsumed within Tetractomia; a new genus Maclurodendron has been established

to include some species formerly placed in Acronychia; and Tetradium has been revived to hold certain species

of Euodia. The remaining species of Euodia are now believed to be most correctly placed in Melicope. They

are herein revised by T.G. Hartley. His account shows that Melicope is represented in Peninsular Malaysia by

10 species. Among these, M. corneri T. Hartley is proposed as a new species, M. pahangensis T.Hartley as a

new name, and M. pachyphylla (King) T. Hartley and M. macrocarpa (King) T. Hartley as new combinations.

In the Aurantioideae, there are now improved treatments of Glycosmis and Citrus, and the plant referred to as

“Citrus sp. A” in the 1972 treatment was subsequently described as a new species, C.halimii. Some controversial

matters such as the discrimination of certain genera and species are discussed herein. Finally the key to

genera, which was imperfect, has been reworked and corrected. This account thus summarizes these details

and presents, where useful, a new treatment to substitute for the old; it should be used in conjunction with the

1972 treatment. To facilitate use, the keys, generic, and specific accounts are set out here in the same form as

in that volume.

ANNOTATED KEY TO GENERA OF MALAYAN RUTACEAE

(Based chiefly on vegetative characters)

1. Woody climbers, never trees or erect shrubs; stems with recurved or hooklike spines .................::0cee 2

a PO EEE PURER Be et Ee eg cca en Se cct ggg sel eocnncncshsuducccescecccancoaces 4

2. Leaves simple, shortly stalked; flowers solitary or clustered in axils, with 4 or 5 petals; fruits globose to

obovoid, sometimes lobed, smooth to rough, lacking pulp-vesicles; seeds few ........... (some) Paramignya

2. Leaves compound, pinnate or trifoliolate (sometimes unifoliolate in juveniles) ................:ceeeeeeeeeeeeeee 3

3. Leaves with 3-9 leaflets; twigs and leaf-stalks prickly; fruits dry, splitting open; medium sized climbers.

nal ONE EET se eT a POP rr WD oy: ee Zanthoxlyum nitidum

3. Leaves with 3 leaflets; leaf stalk not winged; twigs and leaf-stalks not prickly, but axils with usually

recurved spines. Flowers in axillary recemes or panicles; fruits globose to ellipsoid, mucilaginous, without

OER RRM hn 22 LYS ssc csien sagnesearininyasnenncubdins sbenna sang Luvunga Buch.-Ham. RIDLEY (1922)

354. SWINGLE (1943)244. Indomalaya,

c. 12 spp., 2 spp. in Malaya. Fruits look

and smell like small limes.

4. Erect shrub of mangrove or Nipa fringe; fruit 3-4-celled and angled, resembling an angled lemon; seeds very

large, to 35mm long; ovules usually 4 per cell, but one maturing. Leaves simple; spines single or paired,

a 8 ci can nsa Ansa ssciech sk Sodas eds psa atensokabuatnhvbhe Ab okendaae ep nvaseivscusiche'uadaseessoses Merope

5. Buds densely covered with minute rusty-reddish-brown hairs; leaves usually alternate, but rarely opposite,

simple or compound pinnate with up to about 9 leaflets. Ovary 2-5-celled, the cells 1-ovulate. Fruit slightly

ee Se Ta aca Sivas oh Quualebudedn teen uss vdtha DoiaG ghiddasekobayucuyhos eabhvusshbacsvadhs chddansddcae Glycosmis

172

5. Buds glabrous, or if hairy, the hairs whitish, tawny, or grey. Leaves simple or pinnate, opposite or alternate.

Ovary cells usually with 2 or more ovules (except Paramignya). Fruit dry or fleshy ..............cccccsseeeseeees 6

6. Twigs and often also trunk prickly, with scattered prickles ..............:.esscssecesceseeereseteceneceteeenes Zanthoxylum

6. Twigs unarmed, not prickly, sometimes with solitary or paired spines in the axils .............ccesseeeseeeeeeeeees y |

7. Leaves mostly with 3 to matry L6atlets 25.050. <é..csccanececateoneesnovessverensnanse sures oneeacsiatalis a alesis tae nnn 8

7. Leaves simple or at most with 3 leaflets ooo. .....:5:c0c.o0seseencsscseoasnsososonss puewerssepemsaMlnis eee pane tna 15

8. Leaves with 9-13 leaflets, those at base smallest, in a graded series with those at leaf tip the largest; leaf stalk

flattened, narrowly winged; fruit large, ellipsoid, firm, the pulp fleshy and resinous; seeds numerous, large

sessasdenvesnessnssnaysustines Sena cds'debai su wniadsehs nniomtietees ghade aues puMeREM cfs ious beach lieae rete tee aie tie rr Merrillia

8. Leaves with subequal leaflets, leaf-stalk winged or not; fruit VaTIOUS ..............:ssessecsseeeseesseeeeceseesaeesneeees 9

9. Leaves with 5-31 leaflets, rarely a few with only 3; leaves sometimes pubescent ...............c::cceeeeeeeeeeeees 10

9. Leaves usually with 3-7, often 5, leaflets rocco siin 60.202 caccccscnessnnacckcbeonen eau oh eels eet ale ean i)

10. Inflorescence terminal, large, open, to 30cm long; flowers racemosely disposed, well spaced; crushed

tissues foul-smellinig ...........5.:.3202..cecosetnncs aahaslebenbeetete RE tees ws «fi as desta ant noone eee re Clausena excavata

10. Inflorescence up to 15cm long; flowers cymosely disposed, crowded; crushed parts fragrant or not ..... 11

11 Leave opposite; trees; fruit a dehiscent. capsule of follicles. .......,..,...:-tvcee-netes eee Tetradium

11. Leaves alternate; trees or shrubs; firmitaerrye a3: 10scic.2:.iedsces-sec-dceensas0i ogusssoutashnevadgueadasshel aaa tz

12 Leaves faintly scented if crushed; unripe fruits oblong; wild plants ............. ccc eeeeeeseeeseeeeeee Micromelum

12. Leaves spicy-scented if crushed; fruits globose; cultivated plants ..............ecseeseeeeeeteeee Murraya koenigii

13 Leaflets 5, large, c. 12-18cm long; flowers very small (2.5mm long), in panicles c. 30 cm long; ovary 4-

celled, glabrous, with bulging glands in.:520 chi teeth ee Clausena macrophylla

13. Leaflets 3-7, smaller, usually somewhat rhomboid to obovate; ovary 2-6-celled ...........eeceeeeeseeeeneeeeeeees 14

14. Leaf stalk broadly winged; fruit large, woody, globular, 5-7.5 cm wide; deciduous, cultivated tree ..........

ns tvbeavennnctecusessrsntnrcunnoyseducrns~starecidus ds:ta ecg iaaReha Sete name a aaa cheaters dal eae an Limonia acidissima

14. Leaf stalk not winged; fruit small, berrylike, to 12mm wide; seeds 1 or 2, hairy; wild or cultivated,

evergreen, sinall tree or Shrub. ..,....;.s.cccuselcdesnncoe Mosaneae opusei ne aigl ly sate tanatd is Peas ee ee Murraya

15 Leaves with usually 3, sometimes only 1 or. 2, leaflets, .......0...:.<d0:0135002snpacheonse<ourae koageseysuny nthe aeeneneae an 16

15. Leaves simple or umifoliolate <2... .,..i:..ssseasectesstnesbais anaes betendadnruirsadslnap ancaldonteasiie: posts oaat ad aie 18

16. Twigs unarmed; leaves opposite; fruit indemiacent \ 2.0. .ac2.5.-0.6.5.ndonnndeas tevin boehasdeteles ss bocteeb uote eae Melicope

16. Twigs armed with axillary spines; leases alternate; fruit indehiscent ..................c:ccccesseeeenteeeseneeeeneeeeseneees 17

17. Flowers 4- or 5-merous; fruit many-seeded, with hard woody shell; deciduous cultivated tree ..................

sisesileyesnensncesondedsinensencasedsukaseysseiThons derdnedagedey cov senealgaa ah talalt Mies hina na Aegle marmelos

17. Flowers 3-merous; fruit 1-3-seeded, fleshy; evergreen cultivated shrub .................:00000 Triphasia trifolia

18. Twigs unarmed; leaves Opposite .......+.<0u-cesndsndecens tosses is agstise sdenanenvepevensaenstiaaeectiaiaanel tense i ianrnnn nnn 19

18. Twigs armed with axillary spines; leaves altermate 1.1. <ccscnsssincn<nsssoonrsesanch opsakegisntetasens dace euneeeen mannan 22

19. Fruit dry, deliiscent .............0::oocressessesoaneassuseeduesabanybinnsadnaiieaachdus sen salen nek agit namie canes 20

19. Prost fleshy, indehiscent ..............:sseocss-saseesscspouadeneasncnctetse dnupinanuns elie sunime siete aaiia iain ant sana 21

20. Terminal bud glabrous; flowers with 4 stamens alternating with 4 staminodes; seeds winged Tetractomia

20. Terminal bud nearly glabrous to velutinous; flowers with 4 or 8 stamens (or staminodes); seeds not winged.

spsueutenbaeae cpamsdvescasevensdesvovseucencs ssvessscarsed@bedholiaspaduraraecws ed tunh'ehqauty ols deni iniemesiane tie tian ttn Melicope

21. Flowers bisexual; petals valvate; staminal filaments hairy ................... Acronychia pedunculata (L.) Migq.,

Fl. Ind. Bat. Suppl. (1861) 532,

including A. laurifolia Bl. fide

Hartley, J. Arnold Arbor. 55 (1974)

549.

21. Flowers unisexual; petals imbricate; staminal filaments glabrous .............:::s:cesseeeseeseneens Maclurodendron

22. Axillary spines short, curved, paired; fruit small, round, 2-seeded; shrubby plant.. Paramignya cuspidata

22. Axillary spines longer, single or paired, straight; fruit usually 4cm wide or more, seeds usually more than

2s BUGWD OF TOS 0....2..cccveccccsocrecsesconsensescoes sons seuslaceooosedess caddie vacdeseuele eslesdetel sy naiesneiieasnnininan iain 23

123

23. Fruit 3-5-celled, each cell with 1 or 2 seeds; stamens twice as many as petals; small, wild trees Atalantia

23. Fruit 3-18-celled, each cell with 2-12 seeds; stamens 4 (or more) times as many as petals .................... 24

24. Fruit 3-7-celled, each cell with 1 or 2 seeds; stamens connate basally; fruit rind thin, soft, palatable;

i ce cs arn a a vaio aeclna chan sceoevdiipnesoimeavewenngsanendenegivancowesionsdcomdonssoeb Fortunella

24. Fruit usually 8-18-celled, each cell with 4-12 seeds: stamens free or basally connate,the filaments irregularly,

often loosely, connate; fruit rind firm but flexible, somewhat fleshy, not or marginally palatable; wild or

i goatee ta si anit aha cancnubich bps angabaubceetSeaiomeeneas Citrus

Supplementary notes:

Herbs of the genus Ruta (R.graveolens L.), the Rue plant, are occasionally grown as pot plants; they are readily

recognizable by their herbaceous habit, glaucous foliage of compound leaves, pungent aroma, yellow flowers,

capsular fruits, and angular seeds.

There is a recent revision of Clausena in : Molino, J.-F. (1994). Revision du genre Clausena Burm. f.

(Rutaceae). Bull. Mus. Hist. Nat. (Paris), ser. 4, Sect. B, Adansonia 16(1), 105-153

Citrus L.

Citrus halimii Stone, in Stone, Lowry, Scora & Jong, Biotropica 5 (1973) 102.

(after Sultan Abdul Halim Mu’azzam Shah ibni Almarhum Sultan Badlishah of Kedah.

Citrus sp. A: Stone, in Tree Flora of Malaya 1 (1972) 375.

A tree up to 23m tall, trunk cylindric, straight, with ascending branches; bark

smooth gray, thin; wood white. Leaves elliptic or narrowly elliptic, thin coriaceous,

margins entire to obscurely and minutely subcrenulate, undersurface slightly

paler than upper, somewhat olivaceous when dry, the blades 8-15cm long, 3.5-

7.5 cm wide, copiously glandular but the adult leaves not highly fragrant; petioles

articulated at both ends, usually 1-2cm long, the margins distinctly but narrowly

winged. Main lateral nerves 7-11 pairs. Flowers solitary, in axils, pedicels to

3.5mm long; calyx of 5 deltate sepals with minutely ciliate margins; petals white,

3-5 (or more) mm long. Stamens 18-20, free or the filaments loosely connate in

small groups of 2 or 3, glabrous, the anthers yellow. Ovary 6-10-celled, on a flat

disc; style columnar, with 5-6-angled flat stigma; ovary cells with 1-3, rarely 5,

ovules, Fruit about 5 x 5 cm but variable in size, subglobose to pyriform, at apex

slightly concave, the rind glossy deep yellow, bumpy, copiously glandular, firmly

adherent, about 6mm thick, with thin white endocarp; pulp vesicles numerous,

subglobose to pyriform with slender stalk, pale greenish to yellowish-white

translucent, containing acid juice. Seeds numerous, monoembryonic, to 18+ per

fruit, large, flat, about 20 x 9mm, 3-3.5mm thick, narrowed at base, veiny

rugulose, the chalazal cap pale magenta-pink; cotyledons white, the hypocotyl

pale greenish-white.

S. Peninsular Thailand and Peninsular Malaysia, and Borneo (Sabah). In

Malaya mostly in the Main Range at moderate altitudes typically between 2000

and 4500 feet (640-1450m). on ridges in submontane forest.

N.v. “limau kedut kera” or “Limau Kedangsa.”

124

Glycosmis Correa

(Gr. glucus = sweet; osmion=smell)

Ann. Mus. Paris 6 (1805) 384. Ridley, Flora 1 (1922) 348-51. Burkill, Dictionary (1935) 1086. Corner,

Wayside Trees (1940) 571. Swingle, Citrus Ind. (1943) 153. Phoenicimon Ridl. (under Sapindaceae),

Flora 5 Suppl. (1925) 301. Stone, in Whitmore, Tree Fl. Malaya 1 (1972) 380.

A genus of about 45 species from India and Sn Lanka through Burma,

Thailand, southern China, Indochina, Taiwan, throughout Malesia E to NW &

NE Australia, and introduced/naturalized in Florida, W.Indies,and elsewhere.

Continuing studies of this genus (Stone, Proc. Acad. Nat. Sci. Philadelphia

137, 1985) have greatly changed the taxonomy and nomenclature of this group

and thus of the earlier account in the Tree Flora of Malaya, so the present account

must completely replace the earlier one.

Key to Peninsular Malaysia species of Glycosmis

1. Leaves all simple .............-.22:2--+--nsesesrsqcnegrendnsncipesces deinen psinvap7=sse 3 3niieh= eae bas ae mp y

1. Leaves with 3 or more leaflets, or rarely with 2 or 1 leaflet, these mingled with multifoliolate leaflets on the

SANE PAU . 2.2 -.-eesearvensnnesegaceninnndcbiasdlas sie madieds Meditel it onguhle pt debi g eweisioc sundae neh Seale ees ey a 3

2. Leaves leathery, oblong-obovate, to 6cm wide, with obscure veins, alternate; inflorescences much reduced,

PEM IOY oo nesdiciserncstnnosponsith:Susvvanracecddemneee aie ae ea ae G. crassifolia Ridl. J. Roy. Asiat. Soc. Str.

Br. 75 (1917) 14. Endemic, rare; known

from Malacca and Pahang (Taman Negara).

2. Leaves thin coriaceous, elliptic to ovate, wider than 6cm, with obvious veins, sometimes opposite ..........

ssnapaanusnandyesengpvesucesnpetnst apestabentp raed qeipicae mene oe G.chlorosperma (B1.) Spr. var lindleyana

(Swingle) Stone, Proc. Acad. Nat. Sci. Phila.

137 (1985) 3. (G.lindley-ana Swingle,

Citrus Ind. 1 (1943) 155; G. macrophylla

Lindl.: Ridl. Flora 1 (1922) 349. Penang,

Perak.

3. Leaves of 3 leaflets, or sometimes only 2 Or} Or PAOD Y 4 ice cies: cockcnseveseaunsss saseesianes<hnrserasses37400) ee 4

3. Leaves with 5 or more leafletis rapely 4 Ot 3.3505. 5.1ys cncnandetslahagaaints ong Thcote ld phe teguracbeas tyieagtas agen eae 7

4. Ovary reddish pubescent;leaflets usually 1-3, elliptic, with about 6 pairs of secondary veins. Sometimes on

Fimmeatomie Nett s |... 205: -.-néninadesegs odo onsepeigndedicbina oni ane G. puberula Lindl. ex Oliver J. Linn Soc.

Bot. 5 Suppl. 2 (1861) 39. Penang and

Perak; S. Thailand, and a variety in Vietnam

4. Ovary glabrous or with a very few, ephemeral, scattered Wairs .................:cccssesccceeseeeseseessseseesencecseneseseseees 5

5. Fruits to 2cm diam.; leaflets large, to 25 x 6cm, ovate, acuminate with 5-8 pairs of secondary veins;

inflonescence pscudoterminal «......0;.cicssveiecssttanstascabansdaaseerine G. collina Stone Proc. Acad. Nat. Sci. Phila.

137 (1985) 4. G. macrocarpa sensu Ridl.

Flora 1 (1922) 349, non Wight 1840. Perak

(G.Bubu), Pahang (Fraser’s Hill). Montane.

Endemic.

5. Fruits smaller; leaflets smaller and/or of different form, with 4-15 pairs of secondary veins; inflorescences

AMMUNALY oon cc cccssseccsssccssnensconccssesenssesnsecesssorveceseedboasoce ssa Heuehesiedbobpaies ds /aibasiinnts ie antedeananaTt Tannin 6

6 Leaflets up to c.7.5 cm long, elliptic, with rather few (4-6) secondary veins; fruits c. 4-Smm diam. Ovary

glabrate, at first usually with a very few ephemeral scattered hairs. Coastal areas, especially limestone, and

EL a nna, eet fe BL 8” G. mauritiana (Lam.) Tan., Bull. Soc Bot.

France 75 (1928) 708. G. rupestris Ridl. J.

Roy. Asiat. Soc. Str. Br 59 (1911) 81; Flora

1 (1922) 350. Perlis, Kedah, Perak, Penang,

Kelantan. Widespread and variable; also in

10.

11.

125

India, Sri Lanka, Andaman Islands, Nicobar

Islands, Mauritius, Laos, Hainan, and SE

Borneo.

Leaflets often 12-15+ cm long, elliptic-lanceolate, with up to 15-16 pairs of secondary veins; fruit 10-12 mm

diam. Ovary quite glabrous. Coastal areas, and secondary vegetation, sometimes in cultivation .............

a G. pentaphylla (Retz.) DC. Prod. (1824)

538. G. citrifolia sensu Ridley, Flora 1

(1922) 349, non Lindley, 1826. In Malaya

chiefly in the north. Also in India, Sri

Lanka, Burma, Thailand, SW China, and

Indochina.

Leaflets 5-7, leathery, each with about 18 pairs of secondary veins, glabrous above, reddish-scurfy beneath,

large (to 23cm long and 10cm wide, the petiole 6mm long. Panicles elongated, to 20 cm long, densely

reddish-pubescent, almost spikelike, with densely crowded small flowers; ovary 3-celled ........................-

CEE BES RES PR REE - G. decipiens Stone, Tree Fl. Mal. 1 (1972)

381. Phoenicimon rubiginosa Ridl. Flora 5

Suppl. (1925) 301; not G. rubiginosa Ridl.

Kew Bull. (1925) 78. Negri Sembilan,

Pahang, Johore, Trengganu; Pulau Tioman

and Anambas Islands. Rare but distinctive.

Leaflets large or considerably smaller and with few pairs of secondary veins; inflorescences much shorter,

I a ec tans a rs Bn cnet nwt cir discal o Sunnsinaawcecienicain antznntabaice 8

Ovary reddish-puberulent, 3-celled; leaflets 5 to 7, up to 34 cm long, but with only 6-9 pairs of secondary

mane WChaiss CNMIET NENT OUNSIIS 5. <.05...<5.<05 6p cnng stasis wnnstinsntbes e002 G. macrophylla (B1.) Mig. Fl. Ned. Ind. 1

(1859) 522. Sclerostylis macrophylla BI.

Bijdr. (1825) 135. G. sapindoides Lindl.

ex Oliver J. Linn. Soc. Bot. 5 Suppl. 2

(1861) 38. G. elata Ridl. J. Fed. Mal. St.

Mus. 10 (1920) 130. Kedah, Kelantan,

Penang. Andaman and Nicobar Islands,

Thailand, Sumatra, Java, Sunda Isl., Kei

Is]., Papua and W. Australia.

PE 4 ek a Re SR SE rae Oe eae". Seen A ee ce Se a ST aE 9

Leaflets 7-12, linear-lanceolate; rheophytic shrub ..................... G. perakensis Narayanaswamy, Rec. Bot.

Surv. India 14 (1941) 59. Perak, Pahang.

Endemic.

Leaflets mostly 5-7, rarely 9 ot only 3 or 4, not linear-lanceolate; not rheophytic «................:cceeeseeeeees 10

. Leaflets small, usually 3-5, rarely 1 or 7, mostly 2.5-5 cm long, rounded and slightly notched at apex:

RS SS it SRR ae TRS pe Ae G. trichanthera Guillaumin Bull. Bot. Soc.

France 91 (1945) 216. Limestone hills in

various states; W of Main Range as var.

trichanthera (syn. G.calcicola Stone, Gard.

Bull. Sing. 26 (1972) 55); in Kelantan, var.

kelantanica (Stone) Stone, Proc. Acad. Nat.

Sci. Phila. 137 (1985) 22, with much larger

leaflets. Restricted to karst limestone. The

typical variety occurs in Vietnam; another

is found in Burma, and one more in

Sumatra. The notched leaflet tips together

with small 4-merous flowers are reliable

characters. This is a sharply distinct species.

Leaflets somewhat larger, not rounded and notched at the tip. Flowers 5-merous ................:.:0000008 11

Leaflets usually elliptic to ovate, with wide-spaced rather few veins prominent beneath; inflorescences

5s Diab cbiistuling eannn concen cn esaseniigsstsualpinhsKiwaasuaieinswiesyweannvceégrh cwsiecwuneencehnaxhyss 12

Leaflets mostly elliptic-lanceolate, with rather numerous fine secondary veins; inflorescences axillary ...

eo Sa dns ian ak vas ncbenyn sescsvtabnckseaces ¢hasnpeqanseumnnnss G. pentaphylla (Retz.) DC. (see above)

126

12. Inflorescences glabrous or nearly SO 2,.........-csncssossscsnasseesseseseoes G. chlorosperma (B1.) Sprengel, Syst. Veg.

ed. 16, 4 (1827) 162. Cookia chloro-sperma

Bl. Bijdr. 3 (1925) 135. G.malayana Ridl.

J. Roy. Asiat. Soc. Str.Br. 75 (1917) 12. G.

monticola Ridl. 1.c. Widespread through

the Malay Peninsula, lowlands and hills, in

forest, common in W. Malesia. (1) var.

chlorosperma: Widespread in the Peninsula.

(2) var. angustifolia Narayana-swamy, Rec.

Bot. Surv. India 14 (1941) 43. Perak, very

local; S.Borneo. With narrow leaflets. (3)

var. paraphyllophora Stone, Proc. Acad.

Nat. Sci. Phila. 137 (1985) 3. Penang,

Kedah, S. Thailand. Inflorescence bases

with conspicuous paraphylls. (4) var.

lindleyana (Swingle) Stone 1.c. Penang.

With large ‘simple’ opposite leaves.

12. Inflorescences densely reddish-pubescent ...............eeseeeeeeeeeeeees G.tomentella Ridl. J. Roy. Asiat. Soc. Str.

- Br. 75 (1917) 14; Flora 1 (1922) 350.

Selangor (G.Nuang), rare in montane forest.

Also in Sumatra.

Limonia L.

Limonia acidissima L. is the correct name for Feronia limonia (L.) Swingle,

as used in Tree Flora Mal. 1 (1972) 370 (F. elephantum Corr. is another synonym).

The genus Feronia is a synonym of Limonia which is monotypic. This species is

exotic in the Malayan flora, and occurs only in cultivation. The woody-shelled

fruits, the leaves with strongly winged rachis and usually 5 leaflets, and the

deciduous habit distinguish it readily. For a fuller explanation of the nomenclature,

see Stone & Nicholson, Taxon 27 (1978) 551.

Maclurodendron Hartley

Gard. Bull. Sing. 35 (1982) 1-19.

(After F.A MaClure, American botanist specialized in bamboos)

Small to medium trees, all apparently dioecious. Indumentum brownish to

rusty, of simple or fascicled hairs. Leaves opposite, unifoliolate, blades pinnately

veined, pellucid glandular-punctate. Inflorescences axillary in upper axils, cymose,

paniculate, or racemose. Flowers unisexual, 4-merous, the sepals valvate, petals

narrowly imbricate, deciduous; stamens 8, free, unequal, those opposite petals

shorter than the others, the longer ones about as long as the petals; filaments

glabrous; anthers dorsifixed, sterile in pistillate flowers. Gynoecium 4-carpellate,

rudimentary in staminate flowers, with an irregularly, shallowly 8-lobed disc;

carpels 2-ovulate, the ovules collateral to subcollateral; style straight, with 4-

lobed capitate stigma. Fruit a syncarpous drupe of 4 cells, with glandular

leathery exocarp, pergamentaceous endocarp, and | or 2 ovoid to reniform glossy

black seeds per cell; endosperm copious; embryo straight; cotyledons flattened.

A genus of 6 species from Burma, E to Indochina and S China, in W Malesia

from the Malayan Peninsula to the Philipines; two species in Peninsular Malaysia.

127

Key to Malaysian species of Maclurodendron

1. Flower buds 2.5-3 mm wide, sepals and petals densely pubescent outside; style scars distinct on fruit as 4

crowded dots. Leaflet blades 23.5-18.5cm long ............0.0..... M. magnificum Hartley, Gard. Bull. Sing. 35

(1982) 15. Pahang (Genting Highlands, Ulu

Kali, 1500 m). Endemic. Montane forest.

1. Flower buds 1-2mm wide, sepals and petals pubescent to glabrous outside; style scar unitary (scars

confluent) at drupe apex. leaflet blades 5.5 - 24cm long........ M.porteri (Hook.f.) Hartley, Gard. Bull. Sing.

35 (1982) 8. Acronychia porteri Hook. f. FI.

Brit. India 1 (1875) 498. All states and

Singapore; Burma; W.Malesia.

Melicope J. R. & G. Forster

by T. G. HarRTLEY

Char. Gen. PI. (1775) 28, ed. 2 (1776) 55. Engler in Engler & Prantl, Nat. Pflanzenfam. ed. 2.19a (1931)

231. Hartley, Sandakania 4 (1994) 47.

(Gr. meli, honey, and kope, a cutting, referring to the emarginate lobes of the nectar-secreting,

intrastaminal disc)

Shrubs (rarely scandent) or trees. Leaves opposite or whorled, digitately

trifoliolate or unifoliolate. Inflorescences cymose or thyrsiform or sometimes

reduced to solitary flowers, axillary or ramuligerous (1.e., on branchlets below

leaves) or rarely terminal, ramigerous, or cauligerous. Flowers small, bisexual or

functionally unisexual; sepals 4; petals 4, distinct, valvate or narrowly imbricate;

stamens (rudimentary in ? flowers) 8 or 4 or rarely 8-4, distinct; disc intrastaminal,

pulvinate to annular or cupular; gynoecium (rudimentary or sometimes obsolete

in ¢ flowers) 4-carpellate, the carpels connate basally or up to their full length,

with a common apical or subapical style or the stylar elements rarely becoming

divergent, the stigma punctiform to capitate, peltate, or 4-branched, the ovules 2

or | per carpel. Fruit of 1-4 basally connate follicles (the abortive carpels, if any,

persistent) or grading to syncarpous (carpels united into a 4-locular, loculicidally

dehiscent capsule); endocarp pergamentaceous to cartilaginous, adnate to or

separate from epicarp in mature fruit but not expelled at dehiscence. Seeds

solitary or in pairs, remaining attached in dehisced fruit; testa with thick inner

layer of dense, black sclerenchyma and spongy outer layer bounded externally by

a shiny, black pellicle; endosperm copious; embryo straight or slightly curved,

the cotyledons + flattened, elliptic, planate, the hypocotyl superior.

Malagasy and Indo-Himalayan regions eastward to the Hawaiian and

Marquesan Islands and south to New Zealand. About 230 species, 10 of which

occur in the Malayan Peninsula (i.e., in Peninsular Malaysia and/or Singapore).

Melicope is characterized mainly by its combination of opposite or whorled,

digitately trifoliolate or unifoliolate leaves, dehiscent fruit, and shiny, black,

pelliculose seeds which remain attached in the dehisced fruit. Euodia J.R.& G.

Forster, with which Melicope was long confused, consists of seven species and

128

ranges from New Guinea and northeastern Australia east to Samoa, Tonga, and

Niue. Like Melicope, it has opposite, digitately trifoliolate or unifoliolate leaves

and dehiscent fruit, but its seeds are neither shiny nor pelliculose and they are

forcibly expelled, along with the endocarp, when the fruit dehisces. This

classification of the two genera was first proposed in a revision of the southeast

Asian genus Tetradium Lour. (Hartley, Gard. Bull. Sing. 34 (1981) 91-131), which

itself was long confused with Euodia, and was followed in a recent account of the

Bornean species of Melicope (Hartley, Sandakania 4 (1994) 47-74).

The manner of attachment of the seed in the dehisced fruit or Melicope is

variable and provides a useful taxonomic character. In some species, the

attachment is either by a partially detached axile strip of pericarp tissue or by a

partially detached raphe, or by both. This kind of attachment, which is seen in

species 1, as enumerated below, is designated as Type A. In other species,

neither the pericarp nor the raphe detaches and the two are connected by a

funiculus which ranges up to about 3 mm in length. This manner of attachment,

which is seen in species 2-10 below, is designated as Type B.

Seeds of Melicope are often irregularly angled when two develop in a single

fruiting carpel. These shapes, which are caused by crowding, are not given in the

descriptions.

In the above-mentioned account of the Bornean species of Melicope, full

descriptions were given for five species that also occur in the Malayan Peninsula,

namely, M. accedens (Blume) T. Hartley, M. denhamii (Seem.) T. Hartley, M.

glabra (Blume) T. Hartley, M. hookeri T. Hartley, and M. lunu-ankenda (Gaertn.)

T. Hartley. Those descriptions are not repeated here. The synonymies given

herein are intended to be relevant only to the Malayan Peninsula. Melicope

accedens, M. denhamii, M. glabra, and M. lunu-ankenda have additional synonyms

outside this region. With one exception (see Evodia sensu Gaertner, mentioned

in the synonymy of M. lunu-ankenda), the original spelling Euodia is used

throughout the synonymies, correcting the orthographic variant Evodia, which

was used by most of the authors.

Key to species of Melicope in the Malayan Peninsula

1. Leaves, or most of them, unifoliolate’ ............és0sssosssecsacdces dens eunsuus suvuey doeroce uaeeoleuy Valu: ih fen ue ean yf.

1. Leaves, or most of them, trifoliolate ....;........-s0sssscssacoononensodeapaceemseacess conhbesuaeubinnsceeseeueensseiiniannaatann===I 3

2. Flowers with 8 stamens; endocarp separate in mature fruit; young branchlets 4-5 mm wide in third

internode, becoming manifestly Corky |........:cbseosea:ssanesis)voygcedsseluach apne Uae te esa 1. M. suberosa

2. Flowers with 4 stamens; endocarp adnate in mature fruit; young brachlets 2.5-3.5 mm wide in third

internode, not becoming manifestly Corky «....:-0.sessssnssivsnerdesnsynssenentdeanniohedibue manne 10. M. pahangensis

3. Leaflet blades (as seen in the Malayan Peninsula) lobed, sinuate, or repand; plants cultivated or possibly

CRCEDOG sichienicnsind soihronscnd scons anos ensis'ebsnouss eapupides cabot deapuasatecs thes ¥idag dah bodies iaeennananrnnnn 2. M. denhamii

9s) DCAMRCEPIAGES CMUNPE osc piispsinsencisacnsso0: S...esscssasoeresapedpvelepsnsdsaesaeeoeseiiien asin elinne > ihieet Liisi titan 4

4... Trichomes mostly fasciculate or stellate ..........:..0ss0s0s+és00ssdsessbenasos pulbsebalansenicaeyenstatananananannn 7. M. hookeri

4. Trichomes, or most of them, simple ............:.01::s0cescesssssssecssorensdiesis0binvatehacsssllumayiinl ios iin anne 5

129

5. Petals (2.4-) 3.3-4 mm long, the abaxial surface appressed-pubescent; fruiting carpels 8-10 mm long,

the endocarp glabrous; indumentum of leaflet blades mostly restricted to midrib and margin.............

sa ccna Es oie vepewe ceca ewenneacnccocnsnseoensunvene-sheseotensenstnannosaannecs 9. M. pachyphylla

5. Petals (as seen in the Malayan Peninsula) 1-3 mm long, the abaxial surface glabrous or with sparse

indumentum; fruiting carpels (as seen in the Malayan Peninsula) 2.5-5 mm long, or, if larger, then the

endocarp pubescent; indumentum of leaflet blades, if present, not as aboVE ..............:csceeseeseeeseeeeeeeeneeees 6

6. Fruiting carpels (7.5-) 10-11 mm long; endocarp and locules of ovary with indumentum8. M. macrocarpa

6. Fruiting carpels 2.5-5 mm long; endocarp and locules of ovary glabrous ............0....cccceesseeeseeeseeeseeeeeeeeees 7

7. Petals 1-1.5 mm long, glabrous, persistent in fruit; fruiting carpels 2.5-3 mm long .............. 5. M. corneri

7. Petals (1.5-) 2-2.5 (-3) mm long, with indumentum, especially adaxially, or glabrous, deciduous in fruit;

aN IE > NNN NERD RES See Hn Ea sh os ~L da oavn su navsenceasedtn ope sh nsdoseaty ssasecsndacasthiesaaupetecatustenes 8

8. Terminal leaflet blades obovate or broadly so, (7.5-) 9-16.5 cm long, the main veins plane or impressed above,

9-15 on each side of midrib, divergent at angle of 50-60°, the apex usually abruptly and obtusely short-

acuminate; main branches of inflorescences ascending; fruiting carpels 3-4 mm long .............. 3. M. glabra

8. Terminal leaflet blades (as seen in the Malayan Peninsula) elliptic to obovate, 6-30 cm long, the main

veins prominulous to impressed above, 11-22 on each side of midrib, divergent at angle of 60-70°, the

apex usually acuminate; main branches of inflorescences spreading or ascending; fruiting carpels 3.5-

9. Leaflet blades glabrous or nearly so, up to 22 cm long, the main veins prominulous above; main branches

of inflorescences usually ascending; fruiting carpels about 5 mm long........................ 4. M. lunu-ankenda

9. Leaflet blades nearly glabrous to pubescent below, up to 30 cm long, the main veins usually

impressed above; main branches of inflorescences usually spreading; fruiting carpels 3.5-4.5 mm

SLA ee SI SES Ries 16 sane 9 Re oe Oe Ee EOE ey I ERE ee RIT 6. M. accedens

1. Melicope suberosa B. Stone

Gard. Bull. Sing. 36 (1983) 94, fig. 1,2; tab. 1. Type: Peninsular Malaysia: Pahang: Genting Highlands,

Gunong Ulu Kali, Stone & Lowry 15338 (CANB, Isotype).

Tree to 10 m high, trunk to 15 cm diam., like the branches with rugose pale

corky bark; trichomes simple or fasciculate. Young branchlets like the petioles

glabrous or nearly so, becoming manifestly corky, 4-5 mm wide in third internode;

terminal bud densely puberulent. Leaves opposite, unifoliolate, 7.5-21.5 cm

long; petiole 1-4 cm long, 1-2 mm wide at middle; petiolule obsolete; leaflet

blade chartaceous, glabrous, elliptic or elliptic-obovate, 6.5-17.5 x 3-9.5 cm, the

base obtuse to acute, the margin entire or in occasional leaves few-crenulate

toward apex, the apex obtuse to subacuminate, the midrib plane above, the main

veins prominulous above, 10-15 per side, divergent at angle of 65-70°, the veinlet

reticulation + obscure. Inflorescences axillary, few-flowered, 1.5-2 x 1.3-1.5 cm,

the peduncle nearly glabrous or sparsely puberulent, 0.8-1.3 cm long, the pedicels

puberulent, 1.5-2 mm long (about 3 mm long in fruit). Flowers unisexual (only ?

seen), plants probably dioecious; sepals puberulent or sparsely so abaxially,

glabrous adaxially, connate at base, ovate-triangular, about 1.5 mm long, persistent

in fruit; petals greenish white, narrowly imbricate, sparsely puberulent abaxially,

glabrous adaxially, ovate-elliptic, about 4 mm long, deciduous in fruit; stamens

8, infertile, the antesepalous ones about 2 mm long, the filament glabrous, narrowly

obtuse at apex, the anther 0.6-0.8 mm long; disc glabrous; gynoecium 2.2-2.7

mm long, the ovary puberulent or sparsely so, the carpels 2-ovulate, the style

puberulent in proximal 1/2, including stigma 1.5-2 mm long, the stigma capitate,

130

weakly 4-lobed, about 0.6 mm wide. Fruiting carpels connate at base, divaricate,

subglobose to broadly ellipsoid, about 8 mm long, the exocarp dry, glabrate, the

endocarp glabrous, separate; seed attachment Type A; seeds ellipsoid, about 7

mm long.

Known only from the type locality in central Peninsular Malaysia; forest at

1550 m.

Melicope suberosa is most nearly related to M. jugosa T. Hartley and M.

sororia T. Hartley, which are endemic to Borneo. It differs from those species

mainly in having manifestly corky branchlets, persistent sepals, puberulent petals

and ovary, and smaller fruiting carpels. Among its congeners in the Malayan

Peninsula, it is the only species with 8-staminate flowers, separate endocarp, and

Type A seed attachment.

The specific epithet (from the Latin suwberosus, corky) refers to the thick bark

of the branches and trunk.

2. Melicope denhamii (Seem.) T. Hartley

Sandakania 4 (1994) 57. Picrasma denhamii Seem., FI. Vit. (1865) 33.

Type: New Hebrides : Aneitum [Aneityum], McGillivray 46 (BM, Holotype).

Aralia quercifolia Anon., Gard. Chron. (1881) 785, fig. 140, nom, prov.; hort. ex Truff., Rev. Hort.

(1891) 224. Euodia quercifolia (hort. ex Truff.) Ridl., Gard. Chron., ser. 3, 76 (1924) 303. Type

not designated. The illustration of Aralia quercifolia is reasonably adequate for the identification

of this plant.

Euodia ridleyi Hochr., Icon. Bogor 2 (1905) tab. 151. Euodia schullei var. ridleyi (Hochr.) Lauterb.,

Bot. Jahrb. Syst. 55 (1918) 230. Euodia suaveolens var. ridleyi (Hochr.) Bakh. f., Blumea 6

(1950) 365. Probable type: Java: Bot. Gard. Bogor (ex Bot. Gard. Singapore), Backer, Dec.

1904 (U, Isotype).

Borneo east to the southern Philippines and Caroline Islands and southeast

throughout eastern Malesia to the Solomon Islands, New Hebrides, and Fiji. In

the Malayan Peninsula, only putative cultigens of the species are known. These

are represented by collections made from Bot. Gard. Univ. Malaya (Kuala Lumpur)

and Bot. Gard. Singapore.

From their congeners in the Malayan Peninsula, the cultigens of Melicope

denhamii (which apparently orginated in Papuasia) are immediately recognizable

by their lobed, sinuate, or repand leaflet blades. The specific epithet commemorates

H. M. Denham, a British sea captain.

3. Melicope glabra (Blume) T. Hartley

Sandakania 4 (1994) 60. Fagara glabra Blume, Catalogus (1823) 40. Euodia glabra (Blume) Blume,

Bijdr. (1825) 245. Ampacus glabra (Blume) Kuntze, Revis. Gen. Pl. 1 (1891) 98. Type: Java: Blume

(US, Lectotype, designated by Hartley, opp. cit.).

Euodia kingii Engl. in Engl. & Prantl. Nat. Pflanzenfam. III. 4 (1896) 121. Type: Malakka [Malaya

sensu Engler]: “E. glabra King in-herb.” The B material of this was presumably lost. It was

probably the same plant that King (J.Asiat. Soc. Bengal, pt. 2, Nat. Hist. 62 (1893) 208) correctly

identified as Euodia glabra (Blume) Blume.

131

Malayan Peninsula to Sumatra and western Java. In the Malayan Peninsula,

known from Penang, Perak, Trengganu, Selangor, Pahang, Johore, and Singapore;

primary and secondary forest from near sea level to 450 m.

Melicope glabra is most nearly related to M. lunu-ankenda, differing

mainly in the following combination of features: terminal leaflet blades obovate

or broadly so, (7.5-) 9-16.5 x 4-12 cm, the apex abruptly and usually obtusely

acuminate or sometimes rounded, obtuse, or emarginate, the midrib impressed

above, the main veins plane or impressed above, 9-15 per side, divergent at

angle of 50-60°; fruiting carpels 3-4 mm long.

4. Melicope lunu-ankenda (Gaertn.) T. Hartley

Sandakania 4 (1994) 61. Fagara lunu-ankenda Gaertn., Fruct. Sem. Pl. 1 (1788) 334, tab. 68, fig. 9.

Fagara zeylanica J.F. Gmelin, Syst. Nat. 2 (1791) 258 (not seen); Syst. Veg. 1 (1796) 258, nom. illeg.

Zanthoxylum zeylanicum (J. F. Gmelin) DC., Prodr. 1 (1824) 728, nom. illeg. Euodia lunu-ankenda

(Gaertn.) Merr., Philipp. J. Sci. (Bot.) 7 (1912, publ. 1913) 378, as lunur-ankenda. Type: Ceylon,

Konig (L, Holotype).

[Fagara triphylla sensu Roxb., Fl. Ind. 1 (1820) 436, excl. syn., non Lam. 1798.] Zanthoxylum

roxburghianum Cham., Linnaea 5 (1830) 58. Euodia roxburghiana (Cham.) Benth., Fl. Hongk.

(1861) 59. Ampacus roxburghiana (Cham.) Kuntze, Revis. Gen. Pl. 1 (1891) 98. Euodia malayana

Ridl., Fl. Malay Penins. 1 (1922) 342, nom, illeg. Type: cult. Bot. Gard. Calcutta; introduced by

Roxburgh from Penang (not seen).

Roxburgh’s description, apparently drawn up from living material at Calcutta,

is reasonably adequate for the identification of this plant. It is doubtful if a type

was preserved. Roxburgh incorrectly gave Linnaeus as the author of Fagara

triphylla. In the reference he cited (Sp. Pl. ed. Willd. 1 (1798) 666). Willdenow

correctly referred the species to Lamarck.

Himalaya southward to Ceylon, Java, Celebes, and southwestern Philippines.

In the Malayan Peninsula, known from Province Wellesley, Perak, Selangor,

Pahang, Negri Sembilan, Malacca, Johore, and Singapore; coastal and inland

primary and secondary well-drained forest and peat swamp; to 60 m.

Melicope lunu-ankenda is most nearly related to M. glabra (q.v.). The specific

epithet is a Ceylonese name for the plant.

5. Melicope corneri T. Hartley, sp. nov.

Type: Peninsular Malaysia: Selangor: Ulu Gombak, Carrick 1468 (L, Holotype).

Arbor 4.5-7.5 m alta, trichomatibus pro parte maxima simplicibus; foliis trifoliolatis, 34-51 cm

longis; foliolorum laminis chartaceis, saltem subtus in costa et venis primariis et supra in costa +

sparse pubescentibus, in foliolo terminali ellipticis usque obovatis, 20-31 x 7-12 cm; inflorescentiis

axillaribus, multifloris, 4-10 x 3-8 cm, ramis principalibus patentibus; floribus unisexualibus (plantae

dioeciae); sepalis ca. 0.5 mm longis, in fructu persistentibus; petalis glabris, 1-1.5 mm longis, in

fructu persistentibus; staminibus 4, in floribus {7 ca. 2.5mm longis (in floribus ? 1-1.5 mm longis),

filamento glabro, apice subulato usque filiformi, anthera ca. 0.6 mm longa (in floribus ¢ ca. 0.3 mm

longa); gynoecio in floribus ? ca. 1 mm longo (in floribus (J ca. 0.3 mm longo), stigmate capitato,

inconspicue 4-lobato; carpellis fructificantibus basi connatis, 2.5-3 mm longis, endocarpio glabro,

saltem apicem versus adnato; seminibus per Type B affixis, ca. 2 mm longis.

132

Tree 4.5-7.5 m high, trichomes mostly simple. Young branchlets like the

petioles pubescent, 5.4-7 mm wide in third internode; terminal bud velutinous.

Leaves opposite, trifoliolate, 34-51 cm long; petiole 12-19 cm long, 2-4 mm wide

at middle; petiolule in lateral leaflets obsolete or up to 3 mm long, in terminal

leaflet 2-5 mm long; leaflet blades chartaceous, + sparsely pubescent, at least on

midrib and main veins below and on midrib above, in lateral leaflets elliptic or

elliptic-obovate, in terminal leaflet elliptic to obovate, 20-31 x 7-12 cm, the base

in lateral leaflets obtuse to acute, in terminal leaflet acute to subattenuate, the

margin entire, the apex acuminate, the midrib and main veins slightly impressed

above, the main veins in terminal leaflet 15-20 per side, divergent at angle of 60-

70°, the veinlet reticulation prominulous to obscure. IJnflorescences axillary,

many-flowered, 4-10 x 3-8 cm, the axis and branches pubescent, the peduncle 1-

5 cm long, the main branches spreading, the pedicels puberulent or sparsely

pubescent, 0.7-2 mm long (1.5-2 mm long in fruit). Flowers unisexual, plants

dioecious; sepals sparsely puberulent abaxially, glabrous adaxially, connate at

base or up to 1/4 their length, ovate or ovate-triangular, about 0.5 mm long,

persistent in fruit; petals cream or greenish, narrowly imbricate, glabrous, ovate-

elliptic, 1-1.5 mm long, persistent in fruit; stamens 4, in ¥ flowers about 2.5 mm

long (1-1.5 mm long in ? flowers),the filament glabrous, subulate to filiform at

apex, the anther about 0.6 mm long (about 0.3 mm long in ? flowers); disc

pubescent or sparsely so; gynoecium in ¢ flowers about 1 mm long (about 0.3

mm long in ¢ flowers),. the ovary pubescent or sparsely so, the carpels 2-ovulate,

the style pilosulose, at least in proximal 1/2, including stigma about 0.7 mm long,

the stigma capitate, weakly 4-lobed, 0.25-0.3 mm wide. Fruiting carpels connate

at base, divaricate, subglobose or broadly ovoid to obovoid, 2.5-3 mm long, the

exocarp subfleshy, glabrate, the endocarp glabrous, adnate, at least toward apex;

seed attachment Type B, the funiculus about 0.6 mm long, 0.3-0.5 mm wide at

middle; seeds ellipsoid, about 2 mm long.

Endemic to Peninsular Malaysia; primary forest and borders; 18-750 m.

Paratypes: Peninsular Malaysia: Selangor: Gunong Bunga Buah, Whitmore

FRI 337 (A, L); Ulu Gombak, Stone 6892 (CANB); genting Simpah, Poore 236

(CANB). Pahang: Genting Highlands, Chung ] (KLU). Negri Sembilan: Jelebu,

Everett KEP 104947 (A, L), Johore: Gunong Belumut, Whitmore FRI 8771 (L);

Kota Tinggi, Sungai Pelepah, Corner SF 31433 (A, BO). Peninsular Malaysia

without precise locality, Herb. Maingay Kew Distrib. No. 277 pro parte (L).

Melicope corneri is most nearly related to M. accedens, differing mainly in its

smaller anthers and seeds and its combination of usually smaller, glabrous,

persistent petals and usually smaller fruiting carpels. The specific epithet

commemorates Edred J. H. Corner.

6. Melicope accedens (Blume) T. Hartley

Sandakania 4 (1994) 67. Euodia accedens Blume, Bijdr. (1825) 246. Zanthoxylon accedens (Blume)

133

Mig., Fl. Ned. Ind. 1(2) (1859) 671. Ampacus accedens (Blume) Kuntze, Revis. Gen. Pl. 1 (1891) 98.

Type: Java, Blume (L, Lectotype, designated by Hartley, opp. cit.).

Euodia pilulifera King, J. Asiat. Soc. Bengal, pt. 2, Nat. Hist. 62 (1893) 210. Syntypes: Peninsular

Malaysia: Perak: Larut, King’s collector (Kunstler) 6275 (CAL, Holosyntype; L, US, Isosyntypes);

without precise locality, Scortechini 360 (not seen), Wray 2995 (not seen).

Andaman Islands east to Vietnam and south to Java. In the Malayan Peninsula,

known from Kedah, Perak, Trengganu, Selangor, Pahang, Malacca, and Johore;

primary and secondary forest from near sea level to 1980 m.

Melicope accedens is most nearly related to M. corneri (q.v.). The specific

epithet (from the Latin accedo, approaching or resembling) was most likely

intended by Blume to refer to the relationship of the species to Euodia macrophylla

Blume. The latter is now considered to be conspecific with M. accedens (Hartley,

Sandakania 4 (1994) 67).

As Hartley noted in Sandakania 4 (1994) 69, two variants of Melicope accedens

occur in Peninsular Malaysia. Plants described in that account as Variant A

correspond with specimens centering around the type of M. accedens, whereas

those described as Variant B corresponds with specimens centering around the

type of M. pilulifera.

7. Melicope hookeri T. Hartley

Sandakania 4 (1994) 70. Euodia robusta Hook. f., Fl. Brit. Ind. 1 (1875) 488. Ampacus robusta

(Hook.f.) Kuntze, Revis, Gen. Pl. 1 (1891) 98. Syntypes: Malayan Peninsula: Penang, Phillips (not

seen); Singapore, Herb. Maingay Kew Distrib. No. 278 pro parte (GH, Lectotype, designated by

Hartley, opp. cit.). A Leiden sheet with this Maingay number is Melicope glabra.

Peninsular Malaysia (Negri Sembilan: Berembun Forest Reserve), Singapore

(Bukit Timah Forest Reserve), Sumatra, and Borneo; primary and secondary

forest and borders; at lower altitudes throughout the range and ascending to 1600

m in Borneo.

Melicope hookeri is very closely related to M. incana T. Hartley, which

occurs in east-central Sumatra, Borneo, and northern Celebes. The latter species

differs mainly in its densely whitish-tomentose leaflet blades. From its trifoliolate

congeners in the Malayan Peninsula, M. hookeri is readily distinguishable (with

adequate magnification) by its mostly fasciculate or stellate trichomes.

8. Melicope macrocarpa (King) T. Hartley, comb. nov.

Euodia macrocarpa King, J. Asiat. Soc. Bengal, pt.2, Nat. Hist. 62 (1893) 209. Syntypes: Peninsular

Malaysia: Perak: Larut, King’s collector 7489 (L, Lectotype, here designated; US, Isolectotype);

without precise locality, Wray 2618 (not seen), 3266 (not seen).

Tree 5-21 m high, trichomes mostly simple. Young branchlets like the petioles

and inflorescences velutinous or minutely so or rarely sparsely puberulent, 5-8

mm wide in third internode; terminal bud velutinous or rarely appressed-pubescent.

Leaves opposite, trifoliolate (occasional leaves unifoliolate), 18-43 cm long;

petiole 4.5-16.5 (-20) cm long, 2-4 mm wide at middle; petiolule in lateral

134

leaflets 1-9 mm long, in terminal leaflet 2-12 mm long; leaflet blades subcoriaceous

or coriaceous, sparsely puberulent to pubescent below, especially on midrib and

main veins, puberulent on midrib or glabrous above, elliptic or narrowly so to

obovate or in lateral leaflets sometimes narrowly ovate-elliptic, in terminal leaflet

11-32 x 5-15 cm, the base acute to subattenuate, the margin entire, the apex

acuminate or abruptly so or sometimes rounded, the midrib plane or slightly

impressed above, the main veins prominulous to slightly impressed above, in

terminal leaflet 14-22 per side, divergent at angle of 60-75°, the veinlet reticulation

prominulous or plane below or obscure. /nflorescences axillary, many-flowered,

7-14 x 3.5-13cm, the peduncle 1-6 cm long, the main branches ascending, the

pedicels 1.5-2 mm long (2.5-3 mm long in fruit). Flowers unisexual, plants

dioecious; sepals nearly glabrous to puberulent abaxially, glabrous adaxially,

connate at base or up to 1/4 their length, ovate, 0.6-1 mm long, persistent in fruit;

petals white, narrowly imbricate, glabrous or rarely sparsely strigillose abaxially,

glabrous or rarely in proximal 1/3-1/2 sericeous adaxially, ovate-elliptic, 2.7-3

mm long, deciduous in fruit; stamens 4, in ¥ flowers 4-4.5 mm long (1.5-2 mm

long in 9 flowers), the filament glabrous abaxially, sparsely pilosulose in proximal

1/4 adaxially, subulate to filiform at apex, the anther about 1.5 mm long (0.8-1 mm

long in 9 flowers); disc pubescent or villosulous; gynoecium in ? flowers about 2.5

mm long (about 1 mm long in ¥ flowers), the ovary pubescent, the carpels 2-

ovulate, the style pubescent to pilosulose, including stigma about 1.5 mm long,

the stigma capitate, weakly 4-lobed, about 0.6 mm wide. Fruiting carpels connate

at base, divaricate, ellipsoid to obovoid, (7.5-) 10-11 mm long, the exocarp

subfleshy, puberulent to velutinous, the endocarp pubescent, adnate, at least

toward apex; seed attachment Type B, the funiculus 1.5-2.5 mm long, 0.5-0.6

mm wide at middle; seeds ovoid to ellipsoid or rarely subglobose, (4-) 5.5-7 mm

long.

Peninsular Malaysia (Penang, Perak, Selangor, and Pahang) and northern

Sumatra; primary and secondary forest; mostly at lower altitudes but ascending

to 1500 m in Pahang.

From its congeners in the Malayan Peninsula, Melicope macrocarpa differs

mainly in having indumentum on its endocarp and in the locules of its ovary.

9. Melicope pachyphylla (King) T. Hartley, comb. nov.

Euodia pachyphylla King, J. Asiat. Soc. Bengal, pt. 2, Nat. Hist. 62 (1893) 210; Ann. Roy. Bot. Gard.

(Calcutta) 9 (1901) 12, tab. 15. Syntypes: Peninsular Malaysia: Perak: Gunong Batu Puteh, Wray

229 (L, Lectotype, here designated); Gunong Babu, King’s collector (Kunstler) 7432 (L, Isosyntype),

Scortechini 732 (not seen), Wray 3835 (not seen). Pahang: Gunong Berumbun, Wray 1571 (UC,

Isosyntype).

Shrub or tree 1.2-4.5 m high, trichomes mostly simple. Young branchlets like

the terminal bud, petioles, and inflorescences pubescent to velutinous, 3-5.5 mm

wide in third internode. Leaves opposite, trifoliolate, 6-25 cm long; petiole 2-11

cm long, 1.5-3 mm wide at middle; petiolules 1.5-15 mm long; leaflet blades

135

subcoriaceous or coriaceous, puberulent or pubescent on midrib below and,

toward base, on midrib above and margin, otherwise glabrous or nearly so,

elliptic to obovate, in terminal leaflet 4-12.5 x 2-5.5 cm, the base acute to

attenuate or in lateral leaflets obtuse, the margin entire, the apex rounded or

emarginate to short-acuminate, the midrib impressed above, the main veins

prominulous above, in terminal leaflet 9-13 (-17) per side, divergent at angle of

70-75°, the veinlet reticulation obscure. /nflorescences axillary, several- or many-

flowered, 3-9 x 2-3.5 cm, the peduncle (0.35-) 1.5-6 cm long, the main branches

ascending, the pedicels 1-3 mm long (3-4 mm long in fruit). Flowers unisexual,

plants dioecious; sepals pubescent to velutinous abaxially, glabrous adaxially,

connate at base, ovate or ovate-triangular, (1-) 1.5-2 mm long, persistent in fruit;

petals white or cream, narrowly imbricate, rather fleshy, appressed-pubescent

abaxially, sericeous-pubescent adaxially, ovate-elliptic, (2.5-) 3.3-4 mm long,

deciduous in fruit; stamens 4, about 3 mm long, the filament glabrous abaxially,

sparsely pilosulose in proximal 1/6-1/3 adaxially, narrowly obtuse or sometimes

acute at apex, the anther in ¥ flowers 1.5-1.7 mm long (0.7-0.8 mm long in 9

flowers); disc glabrous; gynoecium in ? flowers 1.2-2.5 mm long (about 0.6 mm

long in ¢ flowers), the ovary and style pubescent, the carpels 2-ovulate, the style

including stigma 0.6-1.5 mm long, the stigma clavate or rarely capitate and

weakly 4-lobed, 0.3-0.4 mm wide. Fruiting carpels connate at base, divaricate,

ellipsoid, 8-10 mm long, the exocarp dry, puberulent or glabrate, the endocarp

glabrous, adnate, at least toward apex; seed attachment Type B, the funiculus

about 1 mm long, about 1 mm wide at middle; seeds subglobose or ellipsoid, 4.5-

5 mm long.

Endemic to Peninsular Malaysia, where known from Perak and Pahang; forest

and open ridges, 1350-2000 m.

From its trifoliolate congeners in the Malayan Peninsula, Melicope

pachyphylla is distinguishable mainly by its petals, which are rather fleshy,

appressed-pubescent abaxially, sericeous-pubescent adaxially, and comparatively

large; by its stigma, which is usually clavate; by its fruiting carpels, which are

comparatively large and have glabrous endocarp; and by its leaflet blades, in

which the indumentum is mostly restricted to the midrib and margin.

10. Melicope pahangensis T. Hartley, nom. nov.

Euodia simplicifolia Ridl., J. Linn. Soc., Bot. 38 (1908) 306. Type: Peninsular Malaysia: Pahang:

Gunong Tahan, Wray & Robinson 5492 (BM, Holotype; SING, Isotype). The specific epithet

simplicifolia is pre-empted in Melicope.

Shrub about 2 m high, trichomes simple. Young branchlets like the petioles

nearly glabrous to puberulent and + glaucous, 2.5-3.5 mm wide in third internode;

terminal bud sparsely puberulent to appressed-pubescent. Leaves opposite,

unifoliolate (rarely occasional leaves bifoliolate), 6-11.5 cm long; petiole 1-2 cm

long, 1-1.5 mm wide at middle; petiolule obsolete; leaflet blade subcoriaceous or

136

coriaceous, glabrous, ovate to elliptic, 5-10 x 2.5-4.5 cm, the base rounded to

acute, the margin entire, the apex acute to acuminate, the midrib prominulous or

plane above, the main veins prominulous above, 8-11 per side, divergent at angle

of 60-70°, the veinlet reticulation prominulous or + obscure. Inflorescences

axillary, puberulent, several-flowered, 2-2.5 x 0.8-1 cm, the peduncle 1-1.8 cm

long, the main branches ascending, the pedicels 1.5-2.5 mm long (1.5-3.5 mm

long in fruit). Flowers unisexual (only ? seen), plants probably dioecious; sepals

sparsely puberulent abaxially, glabrous adaxially, connate at base, ovate-triangular,

0.7-0.8 mm long, persistent in fruit; petals (colour unknown) narrowly imbricate,

glabrous abaxially, sparsely and minutely puberulent in proximal 1/3 adaxially,

ovate-elliptic, about 2 mm long, deciduous in fruit; stamens 4, infertile, 0.6-0.8

mm long, the filament glabrous, acute at apex, the anther 0.3-0.5 mm long; disc

pubescent; gynoecium 2-2.5 mm long, the ovary pubescent, the carpels 2-ovulate,

the style pubescent in proximal 1/3, including stigma 1.5-2 mm long, the stigma

capitate, weakly 4-lobed, 0.5-0.6 mm wide. Fruiting carpels connate at base,

ascending, ellipsoid, 6.5-8.5 mm long, the exocarp dry, puberulent or glabrate,

the endocarp glabrous, adnate, at least toward apex; seed attachment Type B, the

funiculus about 0.4 mm long, about 0.6 mm wide at middle; seeds subglobose to

ellipsoid, 4.5-5 mm long.

Endemic to Peninsular Malaysia, where known only from Pahang (Gunong

Tahan and Gunong Brinchang); forest from 1500-2100 m.

From its congeners in the Malayan Peninsula, Melicope pahangensis differs

mainly in its combination of unifoliolate leaves, 4-staminate flowers, adnate

endocarp, and Type B seed attachment.

Excluded species

In his treatment of Euodia in Tree Fl. Malaya 1 (1972) 376-379, Stone

included two species that are now believed not to occur in the Malayan Peninsula,

namely, E. latifolia DC. and E. euneura (Miq.) Mig. The former, which has been

transferred to Melicope (Hartley, Sandakania 4 (1994) 72), ranges from Java

northward to Borneo and the Philippines and eastward to Samoa. It is superficially

similar to M. accedens and M. corneri, with which Stone apparently confused it.

The latter, which is to be transferred to Melicope, is believed to be endemic to

southern Sumatra. Stone apparently confused it with pubescent-leaved plants of

M. macrocarpa, which is one of its close relative, although his description of the

fruit fits neither species.

Index to numbered collections of Melicope

from the Malayan Peninsula

137

The numbers in parentheses refer to the corresponding species in the text.

Anderson 25 (4). Burkill 798 (9); SF 1441, SF 6607 (4). Burkill & Haniff

13248 (6). Carrick 1468 (5). Chew 894 (6). Chung / (5); 2, 3 (6). J. & M.S.

Clemens 5028 (4); 22429 (2). Cockburn FRI 82/1] (9). Corner SF 21200 (4); SF

31433 (5); SF 31927 (3). Cuming 2270 (4). Curtis 2428 (3). Derry 110, 182 (4);

1016 (6). Everett FRI 14005 (4); FRI 14315, FRI 14355 (3); KEP 104947 (5).

Franck J058 (4). Furtado SF 34842 (2). Herb. Griffith 1175 (4); 1176 (6).

Haniff SF 6947, SF 10318 (6). Henderson //289 (8); 11477 (6); SF 17892, SF

23315 (9). Hooker 837 (8). Hou 694 (4). Ismail KEP 104882 (7). Ismail &

Sauji KLU C-42 (2). Jones & Baya 2050, 2053, 2054 (8). Jones et al. 202/d (6).

Kadim & Mamud 97 (8). Kadim & Nur 326 (4). King’s collector (mostly leg.

Kunstler) 698 (8); 775, 1034 (4); 4390 (3); 5269, 5491, 5649 (4): 6130, 6190 (3):

6241 (8); 6275 (6); 7432 (9); 7489 (8); 7556, 7573 (4); 10367, 10630 (3).

Kochummen FRI 2064 (4); KEP 76685 (3); KEP 79122 (6); KEP 93139 (4).

Kunstler (see King’s collector). Herb. Kuntze 6088 (4). Liew SF 37255 (3).

Littke 589 (10). Loh FRI 13374, FRI 13422, FRI 13545 (6). Mahmud Univ.

Malaya 13358 (4). Herb. Maingay Kew Distrib. No. 276 (4); Kew Distrib. No.

277 (GH=6, L=5); Kew Distrib. No. 278 (GH=7, L=3). Maxwell 78-245 (4).

Moysey & Kiah SF 33799 (3). Ng FRI 5699 (4); FRI 5955 (9). Nur SF 2155 (4)

SF 32951 (6); SF 3369] (4). Ogata KEP 105160 (4). Poore 236 (5). Ridley

1633 (6); 5104 (4); 6327, 6329 (3); 6767 (4); 15949 (10). Scortechini 84 (6).

Shah & Nur 776 (4); 899 (8). Sinclair SF 40632 (3). Singh 10/2 (4). Soepadmo

967 (10). Soepadmo & Mahmud 1/033 (4). Sow KEP 71485 (4). Spare SF

36250 (4). Stone 5551, 6436 (6); 6892 (5); 7207, 7235 (9); 14526 (8). Stone &

Lowry 15338 (1). Suppiah FRI 11388, FRI 11827, FRI 11934 (3). Symington

KEP 32207 (9); FMS 36298 (6). Teo & Pachiappan 92, 3/6 (4). Univ. Malaya

8007 (9). Whitmore FRI 337 (5); FRI 3214 (6); FRI 3416 (8); FRI 8771 (5); FRI

12591 (6); FRI 12615, FRI 15163 (3). Wray 229, 1571 (9); 1885 (6); 2295, 2510,

2716 (4); 3161 (3); 3978 (6). Wray & Robinson 5492 (10).

Acknowledgements

Thanks are extended to the directors and curators of A, AAU , BISH, BM,

BO, BRI, CAL, CANB, GH, K, KLU, L, MEL, NSW, NY, P, SING, U, UC,

UKMB, US, and W for making specimens in their care available for this study.

Tetractomia Hook.f.

This genus has been revised by Hartley (J. Arnold Arb. 60 (1979) 127) with

Terminthodia Ridley being reduced to synonymy under it. The species

Terminthodia viridiflora Ridley, previously accepted in the Tree Flora account

(p. 385) has also been reduced to a synonym of Tetractomia tetrandrum (Roxb.)

Craib. Hartley recognizes three forms in the Malayan Peninsula, the “generalized

138

form,” the “Malayan mountain race,” and the “peat swamp race.” He regards

Terminthodia as a representative of the Malayan mountain race, in which he also

includes Tetractomia holtumii (already synonymized in the Tree Flora account.)

Hartley states that “ a complete range of intermediates between the generalized

form and the Malayan mountain race exists in Malayan forests between 900 and

1400 m altitude, and it is evident that the latter is merely a high mountain

extreme of the former. The peat swamp race is reasonably distinct from the

generalized form in the Malay Peninsula in having inflorescences shorter than the

subtending leaves and a tendency toward smaller petals and follicles.” Tetractomia

majus is upheld as a distinct species. The account of Tetractomia therefore

remains as originally published except for the inclusion of Terminthodia, and the

reduction of Terminthodia viridiflora to the synonymy of Tetractomia tetrandrum.

It may be emphasized that the gender of the name Tetractomia is neuter, not

feminine. The name 7. majus is correct but deceptive, as the epithet is the

comparative form of the adjective for large, i.e. equivalent to ‘larger’.

Tetradium Lour.

Loureiro, Fl. Cochinch. (1790) 91; not of Dulac, 1867. HARTLEY Gard. Bull, Sing. 34 (1981) 91.

Gr. tetradion, quaternion, referring to the 4-merous flowers and fruit of Tetradium trichotomum Lour.

Trees or shrubs, usually dioecious, deciduous or evergreen. Indument of

simple hairs. Leaves opposite, pinnate, mostly with a terminal leaflet, the

lateral leaflets usually stalked, their blades usually oil-dotted. Inflorescences

corymbose to occasionally paniculate, terminal or both terminal and in the

upper axils. Flowers unisexual (but rarely bisexual), 4-5-merous, with valvate,

persistent sepals; petals narrowly imbricate in bud, deciduous; stamens opposite

the sepals, longer than the petals in staminate flowers; filaments usually villous

up to the middle, anthers ovoid, dorsifixed; stamens rudimentary or obsolete in

pistillate flowers. Intrastaminal disc present. Gynoecium in pistillate flowers

about as long as the petals, carpels free or connate at base, forming a lobed to

subglobose or obovoid ovary; carpels 2- (or 1-) ovulate; style apical with

peltate 4-5-lobed stigma, Carpels in staminate flowers rudimentary, fingerlike.

Fruit follicular, the follicles 1-2-seeded, dehiscent adaxially, apically and partly

down the abaxial surface; endocarp cartilaginous, persistent. Seeds dark brown

to glossy black, smooth, retained within the follicle; endosperm fleshy; embryo

straight; cotyledons plano-convex; hypocotyl terminal.

A genus of 9 species of India, Indochina, China, Korea, Japan and the Ryukyu

Islands, W.Malesia including the Philippines and Sumatra, but not in Borneo.

Most of the species had originally or subsequently been placed in Euodia and

there is a general resemblance. Two species in Peninsular Malaysia; both very

rare and only in the northern region.

139

Key to the Peninsular Malaysian species of Tetradium

1. Follicles 2-seeded though ofter seemingly 1-seeded; flowers 4-merous; leaflets greenish, hairy ................

Coenen nn ee copie tb vn une deadeaavngenceunnbvansnsvccendeg Tetradium sambucinum

1. Follicles 1-seeded; flowers 5-merous; leaflets glabrous or slightly pubescent, usually glaucous beneath...

i ee che Se Rh a oc onthe By osinnsegonaeane dun’ op ea scgcvansnevaeneupwanal Tetradium glabrifolium

Tetradium sambucinum (B1.) Hartley

(Like Sambucus, Caprifoliaceae)

Gard. Bull.Sing. 34 (1981) 100. Philagonia sambucina BI. Cat. Pl. Buitenz. (1823) 21; Bijdr. (1825) 250.

Euodia sambucina (Bl.) Hook.f.ex Koord. & Val. Med. Lands Plantent. 17 (1896) 216.

Medium to large tree to 34 m tall, branchlets puberulent but glabrate. Leaves

to 36 cm long; of 3-6 pairs of leaflets; stalks of lateral leaflets to 10 mm long;

blades elliptic-oblong to ovate, lanceolate, or oblanceolate, 6-18 cm long, 2-5

cm wide, at base usually somewhat oblique (except in terminal leaflet), with

13-16 pairs of main secondary veins, the margins crenulate distally, the apex

acuminate; conspicuously gland-dotted, drying pale green or brownish, usually

with appressed pubescence on midrib and main veins. Inflorescences to 25 cm

long, the axes slightly pubescent, pedicels less than 1.5 mm long. Flowers 4-

merous, petals yellow-green drying brownish, to 3 mm long, somewhat villous

inside. Fruit of 4 follicles, each 3-4 mm long and high, free to base or nearly

so; endocarp sparsely pubescent; seeds 2 per follicle, coherent (like a single

one), 1.5-2 mm long.

W. Malaysia, Sumatra, Java, and Sumbawa. In Peninsular Malaysia, known

only in Trengganu, G. Andi Mangin (Whitmore FRI 12/39).

Tetradium glabrifolium (Champ. ex Benth.) Hartley

(glabrous leaved)

Gard. Bull. Sing. 34 (1981) 109. Boymia glabrifolia Champ. ex Benth. in Hook. J. Bot. Kew Gard. Misc.

3 (1851) 330. Megabotrya meliaefolia Hance ex Walp. Ann. Bot. Syst. 2 (1852) 259. Evodia meliaefolia

(Hance ex Walp.) Benth. Fl. Hongkong (1861) 58. Phellodendron burkillii Steenis, Gard. Bull. Sing. 17

(1960) 357.

Small or medium tree to 20 m tall, branchlets finely pubescent but fully

glabrate. Leaves to 38 cm long, with 2-9 (rarely but 1) pairs of leaflets, the

blades elliptic to oblong-elliptic, 4-15 cm long, 1.7-6 cm wide, the lateral ones

acute to subtruncate at base, usually oblique, the apex acuminate, the margins

entire to crenulate, with 8-18 pairs of secondary veins, gland-dots inconspicuous,

undersurface whitish or pale green and usually distinctly glaucous; lateral

leaflet stalks 3-15 mm long. Inflorescences to 19 cm long, the axes finely

pubescent to glabrous, pedicels to 4 mm long. Flowers mostly 5-merous;

petals greenish to yellowish or white, to 4 mm long. Fruit usually 5-carpellate,

all or only 1-4 carpels maturing as follicles, these puberulent on the sides, 1-

seeded. Seed black, subglobose or ovoid, 2.5-4 mm long, usually paired with a

smaller aborted seed.

140

NE India and Sikkim Himalaya to S China and Indochina, N to J apan, Taiwan,

S to Peninsular Malaysia, Sumatra, and NE to the Philippines.

In Peninsular Malaysia, known from Kedah (Enggang Forest Reserve,. KEP

78904, type of “Phellodendron burkillii’”) and Gunung Jerai (Kedah Peak, FRI

021715); recorded as a tree to 16 m (50 ft.) tall with smooth slightly lenticellate

bark. Rare.

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The Gardens' Bulletin

Singapore

VOL. 46 (Part 2) June 1994 ISSN 0374-7859

NATIONAL PARKS BOARD

Singapore Botanic Gardens Cluny Road Singapore 259569 Tel: 4741165 Telefax: 4754295

THE GARDENS’ BULLETIN

EDITORIAL COMMITTEE

Chairman: Bernard T.G. Tan, B.Sc.Hons. (Singapore); D. Phil. (Oxon.)

Members: Syed Yusoff Alsagoff

C.J.Goh, B.Sc.Hons. (Singapore); Ph.D (Newcastle upon Tyne)

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

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

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

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

T.W. Foong, B.Sc.Hons., Ph.D. (Cant.)

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

Editors : Chin See Chung, B.Sc.Hons (Singapore); M.Sc: (Malaya); M.S.,

M. Phil., Ph.D.( Yale)

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

Reviewers: F.B. Sampson, Victoria University, Wellington, New Zealand.

J.M. Herr, University of South Carolina, Columbia, U S A.

I.M. Turner, National University of Singapore.

K.M. Kochummen, Forest Research Institute of Malaysia,

Kuala Lumpur, Malaysia.

S.C. Chin, National Parks Board, Singapore.

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

Singapore. Neither the National Parks Board nor the Editorial Committee is respon-

sible for the opinions or conclusions expressed by the contributing authors.

The Gardens’ Bulletin is priced at Singapore $40.00 excluding postage. Overseas

subscribers are required to make paymentin the form of bank drafts or internationalmoney

orders in Singapore currency payable to the Executive Director, National Parks Board,

Singapore. }

Instructions for contributing authors are found behind the Contents Table.

= :

APR 0.9 1995 ARNOLD <4

wi TUBS

The Bardens’ Bulletin

Singapore

VOL. 46(Part 2) Dec 1994 ISSN. 0374-7859

CONTENTS

TURNER, I.M., TAN, H.T.W., CHUA, K.S., METCALFE, D.J.

Recent Botanical Collections from the

SN ee NGA Pe ee MR NE, SERMON EN oc fa no cana nn coe nook «ir ce ans ann sacvanbeen- 1 — 36

WONG YEW KWAN, CHEW PING TING, ALI BIN IBRAHIM

The Tree Communities of the Central Catchment Nature Reserve.

a as dane hse ako wap hem cudnt 37 — 78

LIM, A.L., PRAKASH, H.

Embryology and Development in the Winged Bean

Reem RE ENT INES SPP DIREPLIDERIIS, a a ps Losin Pac ncaa a ahd cage na db wg n cnet cs 79 — 92

WONG YEW KWAN, WHITMORE, T.C.

Carallia brachiata cv. Honiara,

a eco Pastisiate Ornamental Tree ©......2.05¢ ccc he tse ates 93 — 98

SAUNDERS, R.M.K.

The Angiosperm Flora of Singapore Part 2

NUNN Oi Ma Re sy 0S ER Pe en tae, ders ch seen sad aceon canbennttcatios 99 — 102

CHUA, K.S., TAN, H.T.W., TURNER, I.M.

The Angiosperm Flora of Singapore Part 3

ere OS ocach bel bn was aban ive cnnaicnnpwasegwavanwascunuen 103 — 107

TAN, H.T.W., TURNER, I.M., CHUA, K.S.

A Botanical Survey of Pulau Jong, Singapore ................::ccceeeeeeees 109 — 123

TURNER, I.M.

Notes on the Flora of Malaya: New Records,

Overlooked Records and some Nomenclatural Clarification ........ 125 — 130

TURNER, I.M., TAN, H.T.W., CHUA, K.S.

Agaitions to the Flora of Singapore, IT. ....................s.0.escsssesseceeeseees 131 —135

TAY, E.P.

Book Review :

Flora of Australia. Vol. 49: Oceanic Islands 1 .......2.........0....00cce0.00000. 137— 138

CHIN SEE CHUNG

Book Review:

PERE ALE Ow as Sd hc ds as bas pnbd dsuanabpasntapabannbve 138 — 140

Published by

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Recent Botanical Collections

from the Nature Reserves of Singapore

I.M. Turner, H.T.W. TAN, K.S. CHUA AND D.J. METCALFE*

Department of Botany,

The National University of Singapore

Singapore 119260

*Department of Plant Sciences

University of Cambridge

Cambridge CB2 3EA

United Kingdom

Abstract

A botanical survey of the Nature Reserves in Singapore conducted in 1992-93 resulted in the

gathering of more than 2,600 vascular plant specimens. More than 600 species were represented,

including two species not previously recorded from Singapore. The results of the survey are discussed

in the paper.

Introduction

A botanical survey of the Singapore Nature Reserves was conducted

in order to obtain a better understanding of their current flora. Emphasis

was placed on the Central Catchment Area rather than Bukit Timah Nature

Reserve as the former has been much less well investigated than the latter.

Methods

The survey used the technique of collecting fertile plant material to

make voucher herbarium specimens which could be accurately identified

and then used as a permanent record of the contemporary flora of the

Nature Reserves. Teams of collectors visited all parts of the Central

Catchment Area in the three month period of April-June 1992. More

collections were made in the same months of the following year. Further

sporadic collecting trips were made by the authors. When in the field, the

teams endeavoured to collect specimens from all species of plants they

found flowering, fruiting or sporulating. The teams used extension cutters

which aid in the collection of material from heights of up to 5m. Clearly

this meant that it was rarely possible to obtain fertile material from tall

trees.

The specimens gathered were usually pressed in the late afternoon

of the day of collection, or on the morning of the next day. Unpressed

specimens were stored at 4 °C if they were kept overnight. The specimens

were dried at 60 °C in a drying oven and then poisoned with 2.5 % (w/v)

mercuric chloride in 95 % ethanol, redried and then mounted on acid-free

paper. A few specimens were preserved in 70 % ethanol. The specimens

were numbered in one Nature Reserves Survey (NRS) sequence (with the

exception of a minority of collections recorded under personal series) and

identified largely by matching to named specimens in the herbarium of the

Singapore Botanic Gardens. Sets of specimens have been deposited in the

herbarium of the Department of Botany, National University of Singapore

(SINU) and the herbarium of the Singapore Botanic Gardens (SING).

- A numbered grid square system was devised (Fig. 1) to localise the

collections. Information on the NRS specimens has been entered into a

computerized database.

Results

The NRS series reached 2613 over the collecting period, with a large

number of duplicates gathered. Over 600 species (see Appendix 1) have been

identified from the collections. Most of these species are small trees and shrubs,

climbers, herbs and some aquatics reflecting our inability to make good collections

from tall trees. Plants typical of secondary forest, which covers much of the

Central Catchment Area, were found e.g. Macaranga spp., Trema spp., Adinandra

dumosa and Rhodamnia cinerea, and a fair number of introduced and escaped

plants e.g. Hydrocera triflora, Ochna kirkii, Canna indica, Dioscorea

sansibarensis, Wikstroemia ridleyi, Heliconia psittacorum and Cardiospermum

halicacabum. However, many native species more clearly associated with primary

forest were also collected. These included trees and shrubs such as Dracaena

spp., Campnosperma squamatum, Canarium patentinervium, Lindera lucida,

Litsea spp., Sterculia spp., Horsfieldia polyspherula, Gymnacranthera forbesii,

Matthaea sancta, and a number of species in the Rubiaceae and Euphorbiaceae.

Climbers e.g. Uvaria cordata, Pyramidanthe prismatica, Salacia grandiflora,

Gnetum spp., Dissochaeta spp., Stephania capitata, a number of members of the

Vitaceae and many rattan species; and herbs such as Amischotolype gracilis,

Aglaonema spp., Mapania spp., Globba leucantha, Hanguana malayana,

Peliosanthes teta, Stachyphrynium griffithii, Labisia pumila and Ophiorrhiza

singaporensis are all good indicators of primary forest status wherever they ar

found. |

ole eel

Withee | ery

CIE Rae

during the botanical survey of the Nature Reserves. Based on 1:50 000

topographical map produced by the Singapore Mapping Unit of the Ministry

Fig. 1. Map showing the grid system used to localise specimens collected

of Defence (1983).

Discussion

Without doubt the area that provided the most interesting specimens

was the primary swamp forest around the SAF Firing Range at Nee Soon.

It was the only site where several of the orchid species, such as

Bulbophyllum macranthum, Plocoglottis javanica, and Nephelaphyllum

pulchrum were collected. Other interesting species probably restricted to

this, the last patch of swamp forest in Singapore, include Kopsia

singapurensis and Luvunga crassifolia. The forest around the range still

maintains a primary nature and contains some very large trees, though an

alarming number are dead possibly reflecting a change in the drainage

regime. Only Bukit Timah Nature Reserve can boast a vegetation as

botanically exciting. It certainly merits the highest conservation status.

Elsewhere in the Central Catchment Area there are other patches of

forest that approach the primary lowland forest type, most notably around

MacRitchie Reservoir. Though most of the rest of the Catchment Area is

covered with secondary forest, or even so degraded as to be reduced to

lallang or resam stands, interesting plants can still be found. These areas

are also of value in that they may represent future areas of colonisation for

species currently restricted to tiny, fragmented primary areas. An expansion

of range that is much to be encouraged as it will help ensure the future

survival of those species in Singapore.

A number of the species collected were of particular interest.

Bulbophyllum gusdorfii and Iodes cirrhosa are new records for Singapore (Turner,

Tan & Chua, 1994). Daphniphyllum laurinum was found to occur when it

had been reported as probably extinct in Singapore (Keng, 1990).

Acknowledgements

This study was supported financially by the National Parks Board

and in part by the National University of Singapore through research grants

RP3880301 and RP3890339. We would like to thank Robert Teo, Ali

Ibrahim, Chew Ping Ting, Sharon Chan and Wendy Kan from NParks for

their help with the project. We are most grateful to the following for their

assistance in the field and laboratory: Chee Wee San, Goh Mei Fong,

Sixtus Goh, James Gan, Jennifer Hardie, Hin Sock Ee, Kevin Khng, Koh

Kah Kiong, Koh Wei Kee, Liaw Su Eng, Lim Lee Yee, Daniel Lim, Vincent

Lim, Walter Lim, Marinah Marzuki, Mok Wei-Ching, Angeline Ng, Ng

Weng Fun, Jeanette Ong, Brendon Phuah, Soong Beng Ching, Tan Hoe

Teck, Gilbert Tan, Paula Thomas, Woh Siew Leong, Yap Hwee Kheng,

Michelle Yap and Jean Yong. Haji Samsuri Haji Ahmad, Dr D. J. Middleton

and Dr T.C. Whitmore are acknowledged for their help with determination

of specimens.

References

Keng, H. (1990). The concise flora of Singapore: Gymnosperms and

Dicotyledons, Singapore University Press; Singapore.

Turner, I.M., Tan, H.T.W., & Chua, K.S. (in press). Additions to the flora

of Singapore, II. Gardens’ Bulletin, Singapore.

Appendix

List of collections made during the survey with their grid locality

(CCA number). Specimens collected in Bukit Timah Nature Reserve are

indicated as such by the appearance of the code BTNR.

LYCOPODOPHYTA

Lycopodiaceae

Huperzia phlegmaria (L.) Rothm. - NRS0544 CCA 47; NRS1457 CCA 47

Selaginellaceae

Selaginella argentea (Wall.) Spr. - NRS1972 CCA 18

Selaginella atroviridis (Wall.) Spr. - NRS1782 CCA 48; NRS2299 CCA 7; NRS2477 CCA 7; NRS2540

CCA 44; NRS2607 CCA 6

Selaginella intermedia (Blume) Spring - NRS1285 CCA 43

Selaginella willdenowii (Desv.) Bak. - NRS0606 CCA 17

FILICINOPHYTA

Adiantaceae

Syngramma alismifolia (C. Presl) J. Sm. - NRS0632 CCA 17; NRS1610 CCA 18; NRS1951 CCA 18;

NRS1968 CCA 18

Taenitis blechnoides (Willd.) Sw. - NRS0348 CCA 30 and many others

Taenitis interrupta Hook. & Grev. - NRS0247 CCA 17

Aspleniaceae

Asplenium batuense Alderw. - NRS1185 CCA 18; NRS1518 CCA 13; NRS1621 CCA 18; NRS1924

CCA 25; NRS1944 CCA 18

Asplenium longissimum Blume - NRS0394 CCA 30; NRS0630 CCA 17

Asplenium nidus L. - NRS0349 CCA 30

Asplenium tenerum Forst. - NRS1611 CCA 18

Blechnaceae

Blechnum finlaysonianum Hook. & Grev. - NRS0343 CCA 30

Blechnum orientale L. - NRS0267 CCA 17; NRS2111 CCA 12

Stenochlaena palustris (Burm.) Bedd. - NRS0343 CCA 30; NRS1218 CCA 7; NRS1589 CCA 18;

NRS1649 CCA 46; NRS1838 CCA 16; NRS2086 CCA 17; NRS2295 CCA 7

Cyatheaceae

ee errr

Cyathea glabra (Blume) Copel. - NRS1526 CCA 13; NRS1893 CCA 16; NRS1913 CCA 25; NRS1950

CCA 18

Cyathea latebrosa (Wall. ex Hook.) Copel. - NRS1508 CCA 13; NRS1600 CCA 18

Dennstaedtiaceae ‘

Histiopteris incisa (Thunb.) J.Sm. - NRS0634 CCA 17 4

Lindsaea doryophora Kramer - NRS1519 CCA 13 |

Lindsaea ensifolia Sw. ssp. coriacea (Alderw.) Kramer - NRS2281 CCA 28

Lindsaea parasitica (Roxb. ex Griff.) Hieron. - NRS1075 CCA 17; NRS0158 CCA 18; NRS0631 CCA 17 | ;

Microlepia speluncae (L.) Moore var. hancei (Prantl) C.Chr. - NRS1664 CCA 46 |

Pteridium caudatum (L.) Maxon ssp. yarrabense (Domin) Parris - NRS0156 CCA 18; NRS0633 CCA 17

Dryopteridaceae

Pleocnemia olivacea (Copel.) Holttum - NRS1792 CCA 48

Tectaria singaporeana (Wall. ex Hook. & Grev.) Copel. - NRS0274 CCA 17; NRS1483 CCA 49;

NRS1814 CCA 48; NRS2156 CCA 17; NRS2520 CCA 15

Gleicheniaceae

Gleichenia truncata (Willd.) Spr. - NRS1868 CCA 16; NRS1981 CCA 24

Hymenophyllaceae

Trichomanes christii Copel. - NRS0867 CCA 18

Trichomanes javanicum Blume - NRS1295 CCA 43

Trichomanes obscurum Blume - NRS1094 CCA 33; NRS0258 CCA 17

Lomariopsidaceae

Teratophyllum aculeatum (Blume) Mett. ex Kuhn - NRS0051 CCA 13

Teratophyllum ludens (Fée) Holttum - NRS0132 CCA 18

Teratophyllum rotundifoliatum (Bonap.) Holttum - NRS0675 CCA 17

Oleandraceae

Nephrolepis acutifolia (Desv.) Chr. - NRS1403 CCA 7; NRS1491 CCA 49; NRS1830 CCA 15; NRS1922

CCA 25; NRS2602 CCA 6

Nephrolepis biserrata (Sw.) Schott - NRS2263 CCA 28

Parkeriaceae

Ceratopteris thalictroides (L.) Brongn. - NRS0720 CCA 10

Polypodiaceae

Drynaria quercifolia (L.) Sm. - NRS1327 BTNR

Lecanopteris sinuosa (Wall. ex Hook.) Copel. - NRS0157 CCA 18; NRS0221; NRS1193 CCA 7; NRS0529

CCA 36; NRS0455 CCA 46

Platycerium coronarium (Koenig) Desv. - NRS1603 CCA 18; NRS1652 CCA 46; NRS2439 CCA 7

Pyrrosia longifolia (Burm.) Morton - NRSO131 CCA 18

Pyrrosia piloselloides (L.) M.G. Price - NRS2441 CCA 7

Pteridaceae

Pteris ensiformis Burm. - NRS2383 CCA 45

Salviniaceae

Salvinia molesta D.S. Mitchell - NRS0834 CCA 11

Schizaeaceae

Lygodium longifolium (Willd.) Sw. - NRS1439 CCA 44; NRS1647 CCA 15; NRS1699 CCA 45; NRS1745

CCA 47; NRS1770 CCA 47; NRS1906 CCA 25; NRS2021 CCA 24; NRS2116 CCA 12; NRS2227 CCA

36; NRS2368 CCA 45; NRS2216 CCA 43

Lygodium microphyllum (Cav.) R.Br. - NRS0243 CCA 17; NRS2248 CCA 28

Schizaea dichotoma (L.) Sm. - NRS0479 CCA 17; NRS1850 CCA 16; NRS2088 CCA 17; NRS2561

CCA 43

Schizaea digitata (L.) Sw. - NRS0384 CCA 17; NRS0171 CCA 18; NRS2055 CCA 24; NRS2359 CCA

Thelypteridaceae

Christella dentata (Forssk.) Brownsey & Jermy - NRS0263 CCA 17

Christella parasitica (L.) Lév. - NRS0629 CCA 17; NRSO0145 CCA 18

Cyclosorus interruptus (Willd.) H. It6 - NRSO242 CCA 17

Mesophlebion motleyanum (Hook.) Holttum - NRS1750 CCA 12 t;

Pronephrium triphyllum (Sw.) Holttum - NRS0286 CCA 17 %

Vittariaceae i

Vittaria ensiformis Sw. - NRS1832 CCA 15; NRS2406 CCA 45 {

GNETOPHYTA

Gnetaceae

Gnetum gnemonoides Brongn. - NRS0781 CCA 18

Gnetum macrostachyum Hook.f. - NRS1262 CCA 29; NRS0325 CCA 30; NRS0244 CCA 17; NRS0196

CCA 18

Gnetum microcarpum Blume - NRS0915 CCA 6

CONIFEROPHYTA

Podocarpaceae

Podocarpus polystachus R.Br. ex Endl. - NRS0428 CCA 47; NRS0344 CCA 30

ANGIOSPERMOPHYTA

Acanthaceae

Asystasia nemorum Nees - NRS0400 CCA 30; NRS0820 CCA 11

Hygrophila ringens (L.) R.Br. ex Steud. - NRS2438 CCA ?

Ruellia repens L. - NRS1049 CCA 41; NRS0029 CCA 13

Ruellia tuberosa L. - NRS0965 CCA 26

Staurogyne setigera Kuntze - NRS0166 CCA 18

Thunbergia alata Boj. ex Sims - NRS0741 CCA 10; NRS0916 CCA 6

Thunbergia grandiflora Roxb. - NRS0454 CCA 46

Amaranthaceae

Alternanthera sessilis (L.) R.Br. ex DC. - NRS0496 CCA 17

Amaranthus tricolor L. - NRS0913 CCA 6

Anacardiaceae

Buchanania arborescens (Blume) Blume - NRS1932 CCA 25

Buchanania sessifolia Blume - NRS0241 CCA 17; NRS1372 CCA 10

Campnosperma squamatum Ridl. - NRS1010 CCA 18; NRS1413 CCA 7; NRS2427 CCA 13

Ancistrocladaceae

Ancistrocladus tectorius (Lour.) Merr. - NRS1523 CCA 13

Anisophylleaceae

Anisophyllea disticha (Jack) Baill. - NRSO591 CCA 53; NRS0488 CCA 17; NRS0217 CCA 25; NRS0307

BTNR

Annonaceae

Artabotrys costatus King - NRS1358 CCA 18

Artabotrys suaveolens (Blume) Blume - NRS0694 CCA 45; NRS0827 CCA 11; NRS0249 CCA 17;

NRS0341 CCA 30; NRS1412 CCA 7

Cyathocalyx ramuliflorus (Maingay ex Hook.f. & Thomson) Scheff. - NRS0585 CCA 53; DJM177

BTNR

Cyathostemma viridiflorum Griff. - NRS0651 CCA 17; NRS0483 CCA 17; NRS0703 CCA 45; NRS0885

CCA 54

Desmos dasymaschala (Blume) Saff. - NRS1235 CCA 54; NRS1293 CCA 43

Ellipeia cuneifolia Hook.f. & Thomson - NRS1983 CCA 24

Fissistigma fulgens (Hook.f. & Thomson) Merr. - NRS0666 CCA 17

Friesodielsia latifolia (Hook.f. & Thomson) Steenis - NRS0216 CCA 25

Goniothalamus macrophyllus (Blume) Hook.f. & Thomson - NRS1171 CCA 18; NRS0313 BTNR

Goniothalamus ridleyi King - NRS0160 CCA 18

Mitrella kentii (Blume) Mig. - NRS0119 CCA 18; NRS0546 CCA 47; NRS1042 CCA 41

Phaeanthus ophthalmicus (Roxb. ex G.Don) J. Sinclair - NRS1208 CCA 7; NRS1276 CCA 5; NRS1402

CCA 7; DJM179 BTNR

Polyalthia angustissima Ridl. - NRS2133 CCA 17

Polyathia cauliflora Hook.f. & Thomson - NRS1234 CCA 54; NRS1233 CCA 54; NRS1787 CCA 48

Polyalthia lateriflora (Blume) King - NRS1419 CCA 43

Polyathia macropoda King - NRS1069 CCA 17; NRS1311 CCA 35; NRS1219 CCA 54; NRS2137 17

Popowia fusca King - NRS1307 CCA 43

Popowia pisocarpa (Blume) Endl. - NRS0695 CCA 45; NRS1389 CCA 49; NRS1415 CCA 43

Pyramidanthe prismatica (Hook.f. & Thomson) Merr. - NRS0277 CCA 17; NRS0470 CCA 46; CCA 17

Uvaria cordata (Dunal) Alston - NRS0120 CCA 18; NRS0355 CCA 30; NRS0543 CCA 47

Uvaria hirsuta Jack - NRS1334 CCA 49

Xylopia ferruginea (Hook.f. & Thomson) Hook.f. & Thomson - NRS1222 CCA 54

Xylopia malayana Hook.f. & Thomson - NRS1335 CCA 49; NRS2054 CCA 24; DJM100 CCA 18

Apocynaceae

Allamanda cathartica L. - NRSO796 CCA 11

Alstonia angustifolia Wall. ex A.DC. - NRS0905 CCA 6; NRS1207 CCA 7; NRS1142 CCA 36

Kibatalia maingayi (Hook.f.) Woodson - NRS0224 CCA ?

Kopsia singapurensis Ridl. - NRSO126 CCA 18 ae

Leuconotis griffithii Hook.f. - NRSO900 CCA 6

Leuconotis maingayi Dyer ex Hook.f. - NRS0169 CCA 18

Tabernaemontana corymbosa Roxb. - NRS0975 CCA 26; NRS1085 CCA 20

Tabernaemontana pauciflora Blume - NRS0903 CCA 6; NRS1270 CCA 5; NRS1110 CCA 33; NRS1316

CCA 35

Willughbeia coriacea Wall. - NRSO048 CCA 13; NRS0077 CCA 24; NRS2335 CCA 47

Willughbeia tenuiflora Dyer ex Hook. f. - NRS0245 CCA 17

Araceae

Aglaonema nebulosum N.E.Br. - NRS0673 CCA 17; NRS1225 CCA 54; NRS0161 CCA 18; NRS1475

CCA 47

Aglaonema nitidium (Jack) Kunth - NRS0766 CCA 18; NRS0009 CCA 13; NRS0994 CCA 13; NRS0142

CCA 18; NRS0604 CCA17; NRS0367 CCA 30

Aglaonema simplex Blume - NRS0868 CCA 18; NRS1103 CCA 33; NRS1060 CCA 41; NRS1038 CCA

18; NRS1019 CCA 18; DJM315 BTNR

Alocasia denudata Engl. -NRS0712 CCA 45; NRS0297 BTNR; NRS0453 CCA 46; NRS1465 CCA 47

Alocasia macrorrhizos (L.) G.Don - NRS0970 CCA 32; NRS0989 CCA 32

Anadendrum montanum (Blume) Schott - NRSO081 CCA 24; NRS1470 CCA 47; NRS2418 CCA 13;

DJM158 BTNR

Cryptocoryne griffithii Schott - NRS1525 CCA 13; NRS1978 CCA 18

Cyrtosperma merkusii (Hassk.) Schott - NRS1091 CCA 33; NRS1090 CCA 33; NRS0889 CCA 18;

NRS1500 CCA 13

Homalomena griffithii (Schott) Hook.f. - NRS2519 CCA 58

Homalomena sagittifolia Jungh. ex Schott - NRS0538 CCA 47; NRS1263 CCA 41; NRS2419 CCA 18

Pothos latifolius Hook.f. - NRS1415 CCA 43; NRS1450 CCA 47; NRS1819 CCA 48; NRS2013 CCA

24; NRS2148 CCA 17; NRS2503 CCA 15

Rhaphidophora korthalsii Schott - NRS1591 CCA 18; NRS1969 CCA 28

Rhaphidophora lobbii Schott - NRS1485 CCA 49

Rhaphidophora sylvestris (Blume) Engl. var. montana (Blume) Nicols. - NRS0992 CCA 13

Schismatoglottis wallichii (Roxb.) Hook.f. - NRS1604 CCA 18; NRS1721 CCA 43

Scindapsus hederaceus (Zoll. & Moritzi) Mig. - NRS1509 CCA 13

Scindapsus pictus Hassk. - NRS1756 CCA 47; NRS1815 CCA 48; NRS2169 CCA 17; NRS2323 CCA 6;

NRS2514 CCA 15

Syngonium podophyllum Schott - NRS1371 CCA 10; NRS2430 CCA 12

Aristolochiaceae

Thottea grandiflora Rottb. - NRS1377 CCA 49; NRS1458 CCA 47; NRS1458 CCA 47

Asclepiadaceae

Dischidia major (Vahl) Merr. - NRS0810 CCA 11

Dischidia nummularia R.Br. - NRS0576 CCA 46; NRS0547 CCA 47

Hoya lacunosa Blume - NRS0640 CCA 17; NRS1356 CCA 24

Hoya sp. - NRS0948 CCA 28

Hoya parasitica (Roxb.) Wall. ex Wight - NRS0519 CCA 36; NRS0584 CCA 53; NRS0414 CCA 47;

NRS0446 CCA 47; NRS0561 CCA 46; NRS1147 CCA 36; NRS2468 CCA 7

Hoya ridleyi King & Gamble - NRS2063 CCA 24

Balsaminaceae

Hydrocera triflora (L.) Wight & Arnott - NRS0480 CCA 17; NRS0742 CCA 10; NRS1216 CCA 7

Bignoniaceae

Jacaranda filicifolia D.Don. - NRS0327 CCA 30

Spathodea campanulata Beauv. - NRS0760 CCA 18

Bombacaceae

Neesia synandra Mast. - NRS0320 BTNR; NRS0356 CCA 30; NRS0996 CCA 13

Boraginaceae

Cordia cylindristachya Roem. & Schultz - NRS0934 CCA 6; NRS1129 CCA 33

Heliotropium indicum L. - NRS1159 CCA 16; NRS0032 CCA 13

Bromeliaceae

Ananas comosus (L.) Merr. - NRS0628 CCA 17

Burmannia coelestis D.Don - NRS1323 48

Burseraceae

Canarium patentinervium Mig. - NRS0581 CCA 53

Butomaceae

Limnocharis flava (L.) Buchenau - NRS0019 CCA 13; NRS1089 CCA 20

Cabombaceae

Cabomba aquatica Aubl. - NRS0017 CCA 13

Cannaceae

Canna indica L. - NRS0963 CCA 32

Capparaceae

Cleome rutidosperma DC. - NRS0174 CCA 18; NRS0912 CCA 6

Celastraceae

Bhesa paniculata Arn. - NRS0170 CCA 18; NRS0521 CCA; NRS0579 CCA 53; NRS0057 CCA 25;

NRS1202 CCA 7; NRS1030 CCA 18

Kokoona reflexa (Laws.) Ding Hou - DJM346 BTNR

Salacia grandiflora Kurz - NRS0128 CCA 18

Commelinaceae

Amischotolype gracilis (Ridl.) ILM. Turner - NRS0272 CCA 17; NRS0179 CCA 18; NRS0377 CCA 17;

NRS0875 CCA 18; NRS1306 CCA 43; NRS2521 CCA 15

Commelina diffusa Burm.f. - NRS0238 CCA 17

Compositae

Ageratum conyzoides L. - NRS0084 CCA 24

Blumea lacera (Burm.f.) DC. - NRSO151 CCA 18

Crassocephalum crepidiodes (Benth.) S.Moore - NRS1215 CCA 7

Eclipta prostrata (L.) L. - NRS0926 CCA 6

Elephantopus scaber L. - NRSO773 CCA 18; NRS0873 CCA 18

Emilia sonchifolia (L.) DC. ex Wight - NRS0928 CCA 6

Erechtites hieracifolia (L.) Rafin. ex DC. - NRS0939 CCA 28

Mikania micrantha Kunth - NRS0226 CCA 25; NRS0396 CCA 30; NRS0736 CCA 10

Sparganophorus sparganophora (L.) C. Jeffrey - NRS1206 CCA 7

Synedrella nodiflora (L.) Gaertn. - NRS0918 CCA 6

Connaraceae

Agelaea borneensis (Hook.f.) Merr. - NRS0641 CCA 17; NRS0638 CCA 17; NRS2422 CCA 12

Agelaea macrophylla (Zoll.) Leenh. - NRS1538 CCA 28

Cnestis palala (Lour.) Merr. - NRS2129 CCA 13

Rourea fulgens Planch. - NRSO860 CCA 18

Rourea mimosoides (Vahl) Planch. - NRS1553 CCA 28

Convallariaceae

Peliosanthes teta Andr. ssp. humilis (Andr.) Jessp. - NRS1169 CCA 18; NRS1031 CCA 18

Convolvulaceae

Aniseia martinicensis (Jacq.) Choisy - NRS0951 CCA 28; NRS0977 CCA 26

Argyreia ridleyi Prain ex Ooststr. - NRSO001 CCA 13

Cuscuta australis R.Br. - NRS1364 CCA 5

Erycibe malaccensis C.B. Clarke - NRS1354 CCA 24; NRS1577 CCA 15

Erycibe tomentosa Blume var. tomentosa - NRS0423 CCA 47; NRS0536 CCA 47; NRS0405 CCA 30;

NRS0398 CCA 30; NRSO181 CCA 18; NRS0365 CCA 30; NRS0205 CCA ?; NRS0800 CCA 11;

NRS0968 CCA 26; NRS0552 CCA 46

Ipomoea congesta R.Br. - NRS2425 CCA 13

Ipomoea pes-caprae (L.) Sweet ssp. brasiliensis (L.) Ooststr. - NRS0625 CCA 17

Merremia hederacea (Burm.f.) Hallier f. - NRSOO03 CCA 13

Merremia tridentata (L.) Hallier f. - NRS0891 CCA 6; NRS0971 CCA 26; NRS0219 CCA 25

Costaceae

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Costus lucanusianus J. Braun & K. Schum. - NRS0826 CCA 11 *

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Costus speciosus (J. K6nig) Sm. - NRS0967 CCA 32 3

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‘ | Cucurbitaceae

| i Momordica charantia L. - NRS0895 CCA 6

: Trichosanthes celebica Cogn. - NRS0392 CCA 17

E Trichosanthes wawraei Cogn. - NRS0078 CCA 24; NRS0842 CCA 11

: Cyperaceae

B Cyperus digitatus Roxb. - NRS0717 CCA 10

1 Cyperus halpan L. - NRS0594 CCA 53

3 Fimbristylis globulosa (Retz.) Kunth - NRS0919 CCA 6

| f Gahnia tristis Nees - NRS1138 CCA 33; NRS1820 CCA 16; DJM320 BTINR

| . Hypolytrum nemorum (Vahl) Spreng. - NRS0961 C29/37; NRS1176 CCA 18; NRS1158 CCA 16;

i NRS1236 CCA 54; NRS0040 CCA 13; NRS0607 CCA 17; NRSO0082 CCA 24; NRS0093 CCA 24;

| (= NRS1477 CCA 47

2 Mapania cuspidata (Miq.) Uittien - NRS1290 CCA 43; NRS0304 BTNR:; NRS1620 CCA 18; NRS1743

ea CCA 47

| i Mapania palustris (Hassk. ex Steud.) Fern.-Vill. - NRS1278 CCA 43; NRS0369 CCA 17

: 4 Mapania squamata (Kurz) C.B. Clarke - NRS1505 CCA 13

F Rhynchospora corymbosa (L.) E. Britton - NRS0095 CCA 24

‘ Scleria biflora Roxb. - NRS2483 CCA 7

; Scleria ciliaris Nees - NRS0620 CCA 17

Scleria levis Retz. - NRS0831 CCA 11; NRS1927 CCA 25; NRS2301 CCA 7; NRS2372 CCA 45

Thoracostachyum bancanum (Mig.) Kurz - NRS0362 CCA 30; NRS2396 CCA 45

Daphniphyllaceae

Daphniphyllum laurinum (Benth.) Baill. - NRS0416 CCA 47

Dilleniaceae

Dillenia suffruticosa (Griff.) Martelli - NRS0197 CCA ?

Tetracera indica (Christm. & Panz.) Merr. - NRSO184 CCA 18; NRS0287 CCA 17; NRS0485 CCA 17;

NRS0946 CCA 28; NRS0711 CCA 45; NRS0415 CCA 47; DJM178 BTNR

Dioscoreaceae

Dioscorea laurifolia Wall. ex Hook.f. - NRS0443 CCA 47; NRS0042 CCA 13

Dioscorea prainiana R.Knuth NRS2270 CCA 28

Dioscorea pyrifolia Kunth - NRS0481 CCA 17

Dioscorea sansibarensis Pax - NRS1224 CCA 54

Dioscorea stenomeriflora Prain & Burkill - NRS1632 CCA 33

Dipterocarpaceae

Dipterocarpus sp. - NRS0691 CCA 17

Dracaenaceae

Dracaena aurantiaca Wall. - NRS0292 CCA 17; NRS0113 CCA 24; NRS0139 CCA 18

Dracaena fragrans (L.) Ker-Gawl. - NRS0909 CCA 6; NRS0813 CCA 11

Dracaena granulata Hook.f. - NRS1104 CCA 33

Dracaena porteri Baker - NRS0334 CCA 30; NRS0059 CCA 25; NRS1260 CCA 29; NRS1221 CCA 54;

NRS0956 C29/37; NRS0067 CCA 25

Dracaena umbratica Ridl. - NRS1167 CCA 18; NRS1146 CCA 33; NRS1066 CCA 17

Ebenaceae

Diospyros lanceifolia Roxb. - NRS2014 CCA 24

Elaeocarpaceae

Elaeocarpus ferrugineus (Jack) Steud. - NRSOO88 CCA 24

Elaeocarpus mastersii King - NRSOOS55 CCA 25; NRS0328 CCA 30; NRS0435 CCA 47; NRS0475 CCA

46; NRS0713 CCA 10; NRS0693 CCA 45

Elaeocarpus petiolatus (Jack) Wall. - NRSO525 CCA 36; NRS0582 CCA 53; NRS0643 CCA 17

Elaeocarpus polystachyus Wall. - NRSO978 CCA 26; NRS0906 CCA 6; NRS1107 CCA 33

Elaeocarpus stipularis Blume - NRSO060 CCA 25

Eriocaulaceae

Eriocaulon longifolium Nees - NRS0436 CCA 47

wey ees i if tf

Actephila excelsa (Dalz.) Miill.Arg. var. javanica (Miq.) Pax & K. Hoffm. - NRS0299 BTNR

Euphorbiaceae

Agrostistachys longifolia (Wight) Benth. ex Hook.f. - NRS0378 CCA 17; NRS0189 CCA 18; NRS0605

CCA 17

Alchornea villosa Mill.Arg. - NRS0239 CCA 17

Antidesma cuspidatum Miill.Arg. - NRS0856 CCA 18; NRS0270 CCA 17; NRS0639 CCA 17; NRS0648

CCA 17; NRS0880 CCA 18; NRS0473 CCA 46

Antidesma neurocarpum Mig. - NRS0116 CCA 18; NRS0882 CCA 18

Aporusa benthamiana Hook.f. - NRS1231 CCA 54

Aporusa confusa Gage - NRS1359 CCA 18

Aporusa frutescens Blume - NRS0193 CCA 18

Aporusa microstachya (Tul.) Miill.Arg. - NRS2018 CCA 24

Aporusa prainiana King ex Gage - NRS0094 CCA 24; NRS0570 CCA 46

Aporusa symplocoides (Hook-.f.) Gage - NRS0899 CCA 6; NRS402 CCA30; NRS0838 CCA 11; NRS1096

CCA 33; NRS0876 CCA18; NRS0333 CCA 30; NRS0340 CCA 30; NRS0459 CCA 46; NRS0425 CCA

47; NRS0904 CCA 6; NRS0567 CCA 46

Baccaurea kunstleri King ex Gage - DJM232 BTINR

Baccaurea parviflora (Mill.Arg.) Miill.Arg. - NRS0315 BTINR

Baccaurea racemosa (Reinw.) Miill.Arg. - DJM209 BTNR

Breynia coronata Hook.f. - NRS1115 CCA 28

Breynia reclinata (Roxb.) Hook.f. - NRS0461 CCA 46

Claoxylon indicum (Reinw. ex Blume) Endl. ex Hassk. - NRS0718 CCA 10

Croton laevifolius Blume - NRS0872 CCA 18; NRS0859 CCA 18

Endospermum diadenum (Migq.) Airy Shaw - NRS1351 CCA 24

Glochidion littorale Blume - NRS0941 CCA 28; NRS0441 CCA 47

Glochidion superbum Baill. - NRSO007 CCA 13; NRS0358 CCA 30; NRS0943 CCA 28

Hevea brasilensis (Willd. ex A. Juss.) Miill.Arg. - NRS1255 CCA 29; NRS0958 CCA 29/37

Koilodepas longifolium Hook.f. - NRS1305 CCA 43

Macaranga conifera (Zoll.) Miill.Arg. - NRS1250 CCA 29

Macaranga gigantea (Rchb. f. & Zoll.) Miill.Arg. - NRS0O957 C29/37; NRS0954 CCA 28

Macaranga heynei I.M. Johnston - NRS0492 CCA 17

Macaranga punticulata Gage - NRS0716 CCA 10; NRS0984 CCA 32

Macaranga triloba (Reinw.) Miill.Arg. - NRS0037 CCA 13; NRS0599 CCA 17; NRS0194 CCA 18;

NRS0213 CCA 25

Mallotus paniculatus (Lam.) Miill.Arg. - NRS0211 CCA 25; NRS0715 CCA 10; NRS0984 CCA 32;

NRS0716 CCA 10; DJM129 BTNR

Manihot glaziovii Mill.Arg. - NRSO707 CCA 45; NRS0223 CCA 25; NRS0565 CCA 46; NRS0728

CCA 10

Neoscortechinia kingii (Hook-.f.) Pax & K. Hoffm. - NRS0771 CCA 18; NRS2611 CCA 6

Sauropus androgynus (L.) Merr. - NRS0232 CCA 25

Trigonostemon longifolius Baill. - NRS1279 CCA 43

Fagaceae

Castanopsis lucida (Nees) Soepadmo - DJM141 BTNR

Lithocarpus cantleyanus (King ex Hook.f.) Rehder - NRS1163 CCA 16; NRS0808 CCA 11

Lithocarpus conocarpus (Oudem.) Rehder - NRS0539 CCA 47; NRS0564 CCA 46; NRS0498 CCA 17

Lithocarpus encleisacarpus (Korth.) A. Camus - NRS0668 CCA 17

Lithocarpus lucidus (Roxb.) Rehder - NRS0080 CCA 24

Flacourtiaceae

Flacourtia rukam Zoll. & Moritzi - NRS0448 CCA 46; NRS0419 CCA 47; NRS0851 CCA 18; NRS1015

CCA 18; NRS0855 CCA 18; NRS0559 CCA 18; NRS0557 CCA 46

Flagellariaceae

Flagellaria indica L. - NRSO015 CCA 13; NRS2121 CCA12; NRS2458 CCA 7; DJM165 BTNR

Gesneriaceae

Aeschynanthus albidus (Blume) Steud. (= A. purpurascens Hassk.) - NRS1012 CCA 18; NRS1037

CCA 18

Gramineae

Bambusa tulda Roxb. - NRS0346 CCA 30

Centotheca latifolia (Osbeck) Trin. - NRS0615 CCA 17

Coix lacryma-jobi L. - NRS0921 CCA 6

Dendrocalamus asper (Schult.f.) Baker ex Heyne - NRS0523 CCA 36

Imperata cylindrica (L.) P. Beauv. - NRS2277 CCA 28

Isachne globosa (Thunb.) Kuntze - NRS0438 CCA 47

Isachne pulchella Roth ex Roem. & Schultz - NRS0168 CCA 18

Leptaspis urceolata (Roxb.) R.Br. - NRS0039 CCA 13

Thysanolaena latifolia (Roxb. ex Hornem.) Honda - NRS0020 CCA 13; NRS0587 CCA 53

Guttiferae

Calophyllum lanigerum Miq. var. austrocoriaceum (T.C. Whitmore) P.F. Stevens - NRS0278 CCA 17;

NRS2336 CCA 48

Calophyllum tetrapterum Mig. - NRSO578 CCA 53; NRS0440 CCA 47

Calophyllum teysmannii Mig. - NRS1118 CCA 28

Cratoxylum cochinchinense (Lour.) Blume - NRS0172 CCA 18

Garcinia forbesii King - NRS0359 CCA 30; NRS0937 CCA 28; NRS0887 CCA 18; NRS0858 CCA 18;

NRS0352 CCA 30; NRS1404 CCA 7; NRS2420 CCA 17

Garcinia griffithii T. Anderson - NRSO0108 CCA 24; NRS1938 CCA 24; DJM101 CCA 18

Garcinia mangostana L. - NRS1410 CCA 7

Garcinia parvifolia (Miq.) Mig. - NRS2002 24; NRS2059 24

Garcinia urophylla Scort. ex King - NRS1326 BTNR

Ploiarium alternifolium (Vahl) Melch. - NRS0098 CCA 24; NRS1143 CCA 36

Hanguanaceae

Hanguana malayana (Jack) Merr. - NRS0298 BTNR; NRS0035 CCA 13; NRS1077 CCA 17; NRS0598

CCA 53; NRS1862 CCA 16; NRS2145 CCA 17

Hydrocharitaceae

Hyadrilla verticillata (Roxb.) Royle - NRS0395 CCA 30; NRS0016 CCA 13

Hypoxidaceae

Curculigo latifolia Dryand. - NRS0371 CCA 17; NRS0025 CCA 13; NRS1144 CCA 33; NRS1095 CCA

33; NRS1132 CCA 33

Icacinaceae

Gomphandra quadrifida (Blume) Sleumer var. ovalifolia (Ridl.) Sleumer - NRS1309 CCA 35; NRS1226

CCA 54; NRS0777 CCA 18; NRS0751 CCA 18

Iodes cirrhosa Turcez. - NRS0026 CCA 13

Iodes ovalis Blume - NRS0004 CCA 13

Phytocrene bracteata Wall. - NRSO714 CCA 10; NRS0980 CCA 32; NRS2039 CCA 24

20 oa

Iridaceae

Trimezia martinicensis (Jacq.) Herbert - NRS0134 CCA 18; NRS0993 CCA 13; NRS0595 CCA 53

Ixonanthaceae

Ixonanthes icosandra Jack - NRS0099 CCA 24; NRS0901 CCA 6

Ixonanthes reticulata Jack - NRS0783 CCA 18

Labiatae

Hyptis brevipes Poit. - NRSO551 CCA 46

Hyptis capitata Jacq. - NRS0974 CCA 26

Lauraceae

Cassytha filiformis L. - NRS1080 CCA 20; NRS0723 CCA 10

Cinnamomum iners Reinw. ex Blume - NRS0236 CCA 25; NRS1259 CCA 29

Cinnamomum javanicum Blume - DJM230 BTNR

Cryptocarya ferrea Blume - NRS1005 CCA 13; DJM175 CCA 18

Lindera lucida (Blume) Boerl. - NRS0337 CCA 30

Litsea costata (Blume) Boerl. - DJM310 BTNR

Litsea elliptica Blume - NRS0936 CCA 28

Litsea ferruginea Blume - NRS0572 CCA 46

Litsea ridleyi Gamble - NRS1183 CCA 18

Leeaceae

Leea indica (Burm.f.) Merr. - NRS0214 CCA 25; NRS0969 CCA 26; NRS1044 CCA 41; NRS1119

CCA 28; NRS0046 CCA 13; NRS0439 CCA 47

Leguminosae

Archidendron clypearia (Jack) 1.C. Nielsen - NRS0261 CCA 17; NRS0212 CCA 25; NRS0421 CCA 13; —

NRS0254 CCA 17; NRS0574 CCA 46; NRS1058 CCA 414

Bauhinia semibifida Roxb. - NRS0255 CCA 17; NRS0990 CCA 13; NRS0290 CCA 17; NRS1050 CCA

41; NRS0772 CCA 18; NRS0511 CCA 36; DJM154 CCA 18 Sulgl paki fs

Cassia siamea Lam. - NRS1078 CCA 20 oye Lee

Centrosema pubescens Benth. - NRSO0164 CCA 18; NRS0925 CCA 6; NRS0733 CCA 10

Chamaecrista mimosoides (L.) Greene - NRS0735 CCA 10

Clitoria laurifolia Poir. - NRS0133 CCA 18; NRS0182 CCA 18; NRS1079 CCA 20

Crotalaria retusa L. - NRS1082 CCA 20

Derris amoena Benth. var. maingayana (Baker) Prain - NRS0655 CCA 17; NRSO118 CCA 18; NRS0545

CCA 47

Desmodium heterocarpon (L.) DC. - NRS0518 CCA 36

Entada spiralis Ridl. - NRS1130 CCA 33; NRS0510 CCA 17

Indigofera hirsuta L. - NRS1084 CCA 20

Milletia atropurpurea (Wall.) Benth. - NRS1261 CCA 29

Mimosa pigra L. - NRS0542 CCA 47

Parkia speciosa Hassk. - NRS1232 CCA 54

Saraca thaipingensis Cantley - NRS0779 CCA 18

Senna alata (L.) Roxb. - NRS0730 CCA 10; NRS0522 CCA 36

Spatholobus ferrugineus Benth. - NRS2414 CCA 13

Lentibulariaceae

Utricularia bifida L. - NRS1342 CCA 48

Utricularia caerulea L. - NRS1324 CCA 48: NRS1343 CCA 48

Utricularia gibba L. - NRS1344 CCA 48

Linaceae

Indorouchera griffithiana (Planch.) Hallier f. - NRSO814 CCA 11; NRS1257 CCA 29: NRS0462 CCA

46; NRSO0100 CCA 24; NRS0155 CCA 18

Loganiaceae

Fagraea fragrans Roxb. - NRS0269 CCA 17; NRS0745 CCA 10; NRSO0908 CCA 6

Fagraea racemosa Jack ex Wall. - NRS1068 CCA 17; NRS1065 CCA 17; NRS1027 CCA 18

Loranthaceae

Dendrophthoe pentandra (L.) Miq. - NRSO388 CCA 17; NRS0987 CCA 26; NRS0614 CCA 17

Macrosolen cochinchinensis (Lour.) Tiegh. - NRS0387 CCA 17

Scurrula ferruginea (Jack) Danser - NRS0293 CCA 17; NRS0426 CCA 47; NRS0323 CCA 30; NRS0609

CCA 17

Malvaceae

Sida cordifolia L. - NRS0509 CCA 36

Sida rhombifolia L. - NRS0931 CCA 6

Urena lobata L. - NRS0932 CCA 6; NRS0898 CCA 6

Marantaceae

Phrynium parvum (Ridl.) Holttum - NRS1228 CCA 54

Stachyphrynium griffithii (Baker) K. Schum. - NRS1283 CCA 43

Melastomataceae

Clidemia hirta (L.) D. Don. - NRS0043 CCA 13

Dissochaeta celebica Blume - NRS1056 CCA 41; NRS1043 CCA 41; NRS0065 CCA 25

Dissochaeta pallida (Jack) Blume - NRS0659 CCA 17

Melastoma malabathricum L. - NRS0427 CCA 47; NRS2122 CCA 12

Pachycentria maingayi (C.B. Clarke) J.F. Maxwell - NRS1723 43

Pternandra coerulescens Jack - NRSO754 CCA 18; NRS0264 CCA 17; NRS0861 CCA 18; NRS0645

CCA 17

Pternandra echinata Jack - NRS0192 CCA 18; NRS0474 CCA 46; NRS0420 CCA 47

Pternandra tuberculata (Korth.) Nayar - DJM251 BTNR

Sonerila heterostemon Naud. - NRS0778 CCA 18; NRS2412 CCA 18

Meliaceae

Dysoxylum cauliflorum Hiern. - NRSO381 CCA 17; NRS1348 CCA 49

Menispermaceae

Fibraurea tinctoria Lour. - NRS0S93 CCA 53; NRS0143 CCA 18; NRS0644 CCA 17; NRS0004 CCA

30; NRS0087 CCA 24; NRS0408 CCA 47; NRS0401 CCA 30; NRS1035 CCA 18; NRS0332 CCA 30;

NRS1047 CCA 41; NRS0864 CCA 18; NRS0817 CCA 11; NRS0761 CCA 18; NRS1341 CCA 49

Limacia scandens Lour. - NRS0229 CCA 25; NRS0147 CCA 18; NRS0713 CCA 10; NRS0627 CCA 17;

NRS1822 CCA 15

Stephania capitata (Blume) Spreng. - NRSO159 CCA 18; NRS0794 CCA 11 . ad

- D

Tinomiscium petiolare Miers. ex Hook.f, & Thomson - NRS0091 CCA 24; NRS1062 CCA 17; NRS0671

CCA 17 | ee

Tinospora macrocarpa Diels - NRS0849 CCA 18 q

Menyanthaceae

Nymphoides indica (L.) Kuntze - NRS0897 CCA 6; NRS0732 CCA 10; NRS0832 CCA 11; NRS1272

CCA 5

Monimiaceae

Matthaea sancta Blume - NRS1003 CCA 13; NRS0012 CCA 13; NRS1024 CCA 18; NRS1023 CCA 18;

NRS1034 CCA 18; NRS1503 CCA 13

Moraceae

Artocarpus gomezianus Wall. ex Tréc. - NRSO513 CCA 36; NRS2437 CCA?

Ficus apiocarpa Miq. - NRS0811 CCA 11; NRS1559 CCA 28; NRS2128 CCA 13; NRS2160 17

Ficus aurantiacea Griff. - NRSO318 BINR

Ficus aurata Miq. var. aurata - NRS0324 CCA 30; NRS0933 CCA 6; NRSO825 CCA 11; NRS0612 CCA

17; NRS0284 BTNR

Ficus aurata Miq. var. longipilosa Corner - NRS0209 CCA ?; NRSO797 CCA 11; NRSO803 CCA 11:

NRS1140 CCA 36

Ficus binnendykii Miq. var. coriacea Corner - NRS0250 CCA 17; NRS0279 CCA 17

Ficus bracteata Wall. ex Mig. - NRS1532 CCA 13

Ficus chartacea Wall. ex King - NRS0295 CCA 17; NRS0623 CCA 17; NRS1469 CCA 47; DJM130

BTNR

Ficus consociata Blume var. murtoni King - NRS0514 CCA 17

Ficus deltoidea Jack - DJIM12 BTNR

Ficus fistulosa Reinw. ex Blume var. fistulosa - NRSOO10 CCA 13; NRSO195 CCA 18; NRS0234 CCA

25; NRS0137 CCA 18; NRS0757 CCA 18; NRS1123 CCA 28; NRS1128 CCA 28

Ficus globosa Blume - NRS0183 CCA 18; NRSO0130 CCA 18; NRS0430 CCA 47; NRS0424 CCA 47;

NRS0434 CCA 47; NRS0959 CCA 29/37; NRSO888 CCA 18; NRS1253 CCA 29; NRS0680 CCA 17

Ficus grossularioides Burm.f. - NRSO750 CCA 10; NRSO0802 CCA 11; NRS0747 CCA 10

Ficus heteropleura Blume - NRS0202 CCA ?; NRS0203 CCA ?; NRS1254 CCA 29; NRS0960 C29/37;

NRS1180 CCA 18

Ficus laevis Blume - DJM362 BTNR

Ficus pisocarpa Blume - DJM318 CCA 18

Ficus sagittata Vahl - NRS0618 CCA 17

Ficus scortechinii King - NRS1174 CCA 18

Ficus sinuata Thunb. - NRS0186 CCA 18; NRS1000 CCA 13

Ficus sundaica Blume - NRS0014 CCA 13

Ficus trichocarpa Blume - DJM171 CCA 18

Ficus villosa Blume - NRS1618 CCA 18; NRS1896 CCA 16; NRS1954 CCA 18; NRS2120 CCA 12

Streblus elongatus (Miq.) Corner - NRS0512 CCA 36; NRS0308 BTNR; NRS1370 CCA 10

Musaceae

Heliconia bihai L. - NRS0835 CCA 11

Heliconia psittacorum L.f. - NRS0045 CCA 13; NRS0O701 CCA 45

Myristicaceae

Gymnacranthera farquhariana (Hook.f. & Thomson) Warb. - NRS1052 CCA 41

Gymnacranthera forbesii (King) Warb. - NRS1190 CCA 18; NRS0464 CCA 46

Horsfieldia polyspherula (Hook.f. emend King) J. Sinclair var. sumatrana (Miq.) W.J. de Wilde -

NRS0520 CCA 36

Horsfieldia punctatifolia J. Sinclair - DJM132 BTNR

Knema latericia Elm. - NRS0869 CCA 18; NRS0769 CCA 18; NRS0866 CCA 18; NRS1026 CCA 18;

NRS0086 CCA 24; NRS0637 CCA 17

Knema laurina (Blume) Warb. - NRS0074 CCA 24

Knema malayana Warb. - NRS0030 CCA 13

Myristica elliptica Wall. ex Hook.f. & Thomson - NRS0391 CCA 17; NRS0049 CCA 13; NRS1243

CCA 54

Myristica maingayi Hook.f. - DJM309 BTNR

Myrsinaceae

Ardisia colorata Roxb. - NRS0018 CCA 13; NRS0685 CCA 45; NRS1161 CCA 16; NRS1124 CCA 28;

NRS0596 CCA 53; NRS0312 BTNR; NRS0610 CCA 17; NRS0819 CCA 11; NRS0795 CCA 11; NRSO0792

CCA 11; NRS0886 CCA18

Ardisia miqueliana Scheff. - NRSO767 CCA 18

Ardisia singaporensis Ridl. - NRS2079 CCA 17

Ardisia villosa Roxb. - NRS0883 CCA 18

Embelia canescens Jack - NRS0947 CCA 28

Embelia lampani Scheff. - NRS0468 CCA 46; NRS0613 CCA 17

Embelia ribes Burm. - NRS0463 CCA 46; NRS0589 CCA 53; NRS0452 CCA46; NRS0208 CCA 25

Labisia pumila (Blume) Fern.-Vill. - NRS0686 CCA 45; NRS1186 CCA 18; NRS1039 CCA 18; NRS0140

CCA 18; NRS0376 CCA 17; NRS1014 CCA 18; NRS0180 CCA 18; NRS0753 CCA 18; NRS1384 CCA

49: NRS1466 CCA 47; NRS1800 CCA 48

Maesa ramentacea Wall. ex Roxb. - NRS0724 CCA 10; NRS1249 CCA 29; NRS0467 CCA 46 aye )

Myrtaceae

Aphanomyrtus rostrata Mig. - NRS0878 CCA 18

Decaspermum fruticosum J.R. Forst. & G. Forst - NRS0727 CCA 10; NRS0602 CCA 17: NRS0791

CCA 11; DJM174 CCA 18

Eugenia cerina M.R. Hend. - NRS0235 CCA 25

Eugenia cumingiana Vidal - NRS1332 CCA 49

Eugenia filiformis Duthie var. filiformis - NRS1267 CCA 5; NRS0537 CCA 47

Eugenia filiformis Duthie var. clavimyrtus (Koord. & Valeton) M.R. Hend. - NRS1994 CCA 24

Eugenia grandis Wight - NRS0231 CCA 25; NRS0350 CCA 30; NRS0924 CCA 6

Eugenia longiflora (Presl) Fern.-Vill. - NRSO013 CCA 13; NRS0062 CCA 25; NRS0237 CCA 17:

NRS0894 CCA 6; NRS0938 CCA 28

Eugenia oblongifolia Duthie - NRS0497 CCA 17

Eugenia pachyphylla Kurz - NRS0138 CCA 18

Eugenia polyantha Wight - NRS0562 CCA 18; NRS1274 CCA 5; NRS1361 CCA 18

Eugenia pustulata Duthie - NRS1347 CCA 49

Eugenia ridleyi King - NRS0555 CCA 5

Eugenia spicata Lam. - NRS0743 CCA 10; DJM322 CCA 48

Psidium guajava L. - NRS0890 CCA 6

Rhodamnia cinerea Jack - NRS0257 CCA 17; NRS0491 CCA 17; NRS0910 CCA 6; NRS0929 CCA 6;

NRS0114 CCA 28; NRS0892 CCA 6; NRS1059 CCA 41; NRS0418 CCA 47

Rhodomytrus tomentosa (Ait.) Hassk. - NRS0981 CCA 26

Najadaceae

Najas indica (Willd.) Cham. - NRS0403 CCA 30; NRS0838 CCA 18

Najas malesiana Delile - NRS0090 CCA 24

Nepenthaceae

Nepenthes ampullaria Jack - NRS0345 CCA 30

Nepenthes gracilis Korth. - NRS0422 CCA 47; NRS0603 CCA 17; NRS1102 CCA 33

Nepenthes rafflesiana Jack - NRS0268 CCA 17; NRS0601 CCA 17; NRS1093 CCA 33

Nymphaeaceae

Nymphaea pubescens Willd. - NRSOOO8 CCA 13

26 “"

Ochnaceae

Gomphia serrata (Gaertn.) Kanis - NRS0301 BTNR

Ochna kirkii Oliv. - NRS0793 CCA 11

Olacaceae

Erythropalum scandens Blume - NRS0390 CCA 17; NRS0657 CCA 17; NRS0845 CCA 18; NRS1017

CCA 18; NRS0566 CCA 46; NRS0844 CCA 18; NRS1876 CCA 18; NRS2423 CCA 12

Strombosia ceylanica Gardn. - NRS1392 CCA 49; NRS1714 CCA 43; NRS1946 CCA 18

Strombosia javanica Blume - DJM333 BTNR

Onagraceae

Ludwigia adscendens (L.) Hara - NRSO729 CCA 10

Ludwigia hyssopifolia (G. Don) Exell - NRS0033 CCA 13; NRS0489 CCA 17 -

Ludwigia octovalvis (Jacq.) Raven - NRS0410 CCA 47

Opiliaceae

Champereia manillana (Blume) Merr. - NRS0265 CCA 17; NRS0465 CCA 46; NRS0600 CCA 17;

NRS0588 CCA 53; NRS1125 CCA 28; NRS0442 CCA 47; NRS0698 CCA 45; NRS0335 CCA 30;

NRS0699 CCA 45

Lepionurus sylvestris Blume - NRS2337 CCA 47

Orchidaceae

Arundina graminifolia (Don) Hochr. - NRS0115 CCA 18; NRS0966 CCA 26; NRS0737 CCA 10

Bromheadia finlaysoniana (Lindl.) Rchb.f. - NSRO985 CCA 26; NRS0399 CCA 30

Bulbophyllum gusdorfii J.J.Sm. - NRS1683 CCA 45

Bulbophyllum macranthum Lindl. - NRS1320 CCA 18

Bublophyllum vaginatum (Lindl.) Rchb.f. - NRS1682 CCA 45

Calanthe pulchra (Blume) Lindl. - NRS2089 CCA 16

Claderia viridiflora Hook.f. - NRSO002 CCA 13; NRS1113 CCA 28

Cymbidium finlaysonianum Lindl. - NRS2175 CCA17

Dendrobium crumenatum Sw. - NRS0O110 CCA 24; NRS0200 CCA ?; NRS1081 CCA 20

Eulophia graminea Lindl. - NRS0124 CCA 18

Lecanorchis malaccensis Ridl. - NRS1533 CCA 13 : te ;

Nephelaphyllum pulchrum Blume - NRSO775 CCA 18; NRS0847 CCA 18; NRS1936 CCA 18

Plocoglottis gigantea (Hook-.f.) J.J.Sm. - NRS0122 CCA 18; NRS0125 CCA 18; NRS1002 CCA 13

Plocoglottis javanica Blume - NRS0999 CCA 13; NRS1516 CCA 13

Pteroceras pallidum (Ridl.) Holttum - NRS1935 CCA 18

Spathoglottis plicata Blume - NRS0021 CCA 13; NRSO738 CCA 10; NRS0548 CCA 47; NRS1923 CCA

Taeniophyllum obtusum Blume - NRS1318 CCA 18

Thrixspermum trichoglottis (Hook-f.) Kuntze - NRS1319 CCA 18

Vanilla griffithii Rchb.f. - NRSO955 CCA 28; NRS1112 CCA 28; NRS1353 CCA 24; NRS2140 CCA 17

Oxalidaceae

Averrhoa carambola L. - NRS0922 CCA 6

Oxalis barrelieri L. - NRS0437 CCA 47: NRS0776 CCA 18: NRS0914 CCA 6

Palmae

Areca alicae F.v.M. - NRS0962 C29/37

Areca catechu L. - NRS1088 CCA 20

Calamus diepenhorstii Mig. - NRS1245 CCA 54; NRS1239°CCA 54

Calamus insignis Griff. - NRS1302 CCA 43

Calamus lobbianus Becc. - NRS1300 CCA 43; NRS0294 BTNR

Calamus oxleyanus Teijsm. & Binn. - NRS1299 CCA 43; NRS0316 BTINR

Caryota mitis Lour. - NRS0457 CCA 46; NRS0407 CCA 47

Daemonorops angustifolia (Griff.) Martelli - NRS0677 CCA 17; NRS0833 CCA 11; NRS0683 CCA 17:

NRS0031 CCA 13; NRS2431 CCA 13; NRS0505 CCA 17

Daemonorops didymophylla Becc. - NRS0360 CCA 30; NRSO818 CCA 11; NRS1298 CCA 43; NRS0682

CCA 17; NRS0824 CCA 11; NRS1240 CCA 54; NRS0676 CCA 17; NRS0311 BTNR: NRS0366 CCA

Daemonorops geniculata (Griff.) Martelli - NRS0541 CCA 47; NRS0450 CCA 46

Daemonorops grandis (Griff.) Martelli - NRSO0283 CCA 17; NRS0503 CCA 17; NRS0506 CCA 17

Daemonorops hystrix (Griff.) Martelli - NRS0502 CCA 17; NRS0056 CCA 25; NRS0806 CCA 11;

NRS0815 CCA 11

Daemonorops leptopus (Griff.) Martelli - NRS1109 CCA 33

Daemonorops longipes (Griff.) Martelli - NRSO386 CCA 17; NRS0505 CCA 17; NRS0361 CCA 30:

NRS0385 CCA 17; NRS0681 CCA 17; NRS0280 CCA 17; NRS2424 CCA 13

Daemonorops periacantha Miq. - NRS0684 CCA 17

Iguanura wallichiana (Wall. ex Martelli) Hook.f. - NRS0177 CCA 18; NRS1313 CCA 35

Korthalsia echinometra Becc. - NRS1175 CCA 18

Licuala ferruginea Becc. - NRS0550 CCA 47; NRS1177 CCA 18; NRS0597 CCA 53

Nenga pumila (Martelli) Wendl. - NRS0364 CCA 30; NRS0821 CCA 11; NRS0744 CCA 10; NRS1248

CCA 54

Oncosperma horridum (Griff.) Scheff. - NRS0342 CCA 30

Pinanga malaiana (Martelli) Scheff. - NRSO109 CCA 24; NRS0135 CCA 18; NRS0382 CCA 17; NRS0879

CCA 18

Plectocomia elongata Martelli ex Blume - NRS0107 CCA 24; NRS1111 CCA 33; NRS0549 CCA 47;

NRS1266 CCA 41; NRS0678 CCA 17

Rhopaloblaste singaporensis (Becc.) Hook.f. - NRS1308 CCA 35; NRS1444 CCA 43; NRS1455 CCA

Pandaceae

Galearia fulva (Tul.) Mig. - NRS0654 CCA 17; NRS1230 CCA 54; NRS0881 CCA 18

Pandanaceae

Freycinetia angustifolia Blume - NRS2057 24; NRS2131 CCA 17

Freycinetia imbricata Blume - NRS1420 CCA 43

Frecinetia javanica Blume - NRS1527 CCA13; NRS1597 CCA 18; NRS1959 CCA 18

Pandanus atrocarpus Griff. - NRS1993 CCA 24; NRS2097 CCA 17; NRS2198 CCA 25

Pandanus houlletii Carriere - NRS1418 CCA 43

Pandanus monotheca Martelli - NRSO710 CCA 45; NRS1303 CCA 43; NRS1241 CCA 54

Pandanus motleyanus Solms - NRS2183 CCA 25

Pandanus parvus Ridl. - NRS1776 CCA 48

Pandanus scortechinii Martelli - NRS2072 CCA 16

Passifloraceae

Adenia macrophylla (Blume) Koord. var. singaporeana (Wall. ex G. Don) W.J. de Wilde - NRS0058

CCA 25; NRS0447 CCA 46; NRS1028 CCA 18; NRS1357 CCA 25

Passiflora foetida L. - NRSO726 CCA 10

Passiflora laurifolia L. - NRS1237 CCA 54

Passiflora suberosa L. - NRSO260 CCA 17; NRS0527 CCA 36

Phormiaceae

Dianella ensifolia (L.) DC. - NRS1136 CCA 33; NRS0281 CCA 18

Piperaceae

Piper caninum Blume - NRS1472 CCA 47; NRS1432 CCA 43; NRS2042 CCA 24

Piper muricatum Blume - NRS1187 CCA 18

Piper pedicellosum Wall. ex C. DC. - NRS1265 CCA 41

Polygalaceae

Polygala paniculata L. - NRS0952 CCA 28

Xanthophyllum affine Korth. - NRSO816 CCA 11; NRS1213 CCA 7; NRS0995 CCA 13; NRS1762

CCA 47; NRS1783 CCA 48; NRS2016 CCA 24

Xanthophyllum ellipticum Korth. - NRS0575 CCA 46; NRS2338 CCA 47

Salomonia cantoniensis Lour. - NRSO105 CCA 24

Polygonaceae

Polygonum barbatum L. - NRS0022 CCA 13; NRS1198 CCA 7

Polygonum chinense L. - NRS0976 CCA 26; NRS0964 CCA 26

Polygonum orientale L. - NRS1105 CCA 33

Pontederiaceae

Monochoria hastata (L.) Solms - NRS1083 CCA 20

Monochoria vaginalis (Burm.f.) Pres] - NRS0036 CCA 13; NRS0658 CCA 17

Rhizophoraceae

Gynotroches axillaris Blume - NRS0432 CCA 47; NRS1053 CCA 41; NRS0207 BTNR; NRS0608 CCA

Pellacalyx axillaris Korth. - NRS0218 CCA 25; NRS0146 CCA 18; NRSO0411 CCA 47; NRS1258 CCA

29; NRS1362 CCA 18

Rosaceae

Prunus polystachya (Hook f.) Kalkman - NRS0201 BTNR; NRS0191 CCA 18; NRS0150 CCA 18;

~ NRS0347 CCA 30; NRS0106 CCA 24

Rubus moluccanus L. - NRS1117 CCA 28; NRS1170 CCA 18; NRS1273 CCA 5; NRS1310 CCA 35;

NRS0530 CCA 47

Rubiaceae

Aidia wallichiana Tirveng. - NRS1339 CCA 49

Borreria laevicaulis (Miq.) Ridl. - NRS10 CCA41

Canthium confertum Korth. - NRS0251 CCA 17

Canthium glabrum Blume - NRS2342 CCA 48

Canthium horridum Blume - NRS0044 CCA 13

Cephaelis singaporensis Ridl. - NRS2223 CCA 43

Chasalia chartacea Craib - NRS0788 CCA 11; NRS0768 CCA 18; NRS1048 CCA 41; NRS1099 CCA

33; NRS1229 CCA 54; NRS1314 CCA 35; NRS1025 CCA 18

Chasalia curviflora (Wall.) Thwaites - NRS0271 CCA 17; NRS0649 CCA 17

Diodia ocymifolia (Willd. ex Roem. & Schultz) Bremek. - NRS0429 CCA 47

Diplospora malaccensis Hook.f. - NRSO763 CCA 18; NRS1264 CCA 41

Gaertnera grisea Hook.f. ex C.B. Clarke - NRS0302 BTINR

Gaertnera obesa Hook.f. ex C.B. Clarke - NRS0586 CCA 53; NRS0590 CCA 53; NRS0073 CCA25;

NRS0061 CCA 25; NRS1707 45; NRS1989 CCA 24

Gaertnera viminea Hook.f. ex C.B. Clarke - NRS0303 BTNR; NRS1312 CCA 35

Gynochthodes sublanceolata Mig. - NRS0445 CCA 47; NRS1471 CCA 47

Hedyotis auricularia L. - NRSO141 CCA 18

Hedyotis capitellata Wall. ex G. Don - NRS0076 CCA 24; DJM166 CCA 18

Hedyotis congesta R.Br. ex G. Don - NRS0577 CCA 53; NRS0809 CCA 11; NRS1145 CCA 36;

NRS1108 CCA 33; NRS0540 CCA 47; NRS0101 CCA 24; DJM139 CCA 18

Hedyotis herbacea L. - NRSOOSO CCA 13; NRS0911 CCA 6; NRS1214 CCA 7

Hydnophytum formicarium Jack - NRS0444 CCA 47; NRS0650 CCA 17; NRS1723 CCA 43

Ixora congesta Roxb. - NRS0330 CCA 17; NRS0517 CCA 36; NRS0372 CCA 17; NRS0451 CCA 46;

NRS0516 CCA 36; NRS0841 CCA 11; NRS0725 CCA 10; NRS0721 CCA 10; NRS0979 CCA 26;

NRS0972 CCA 26; NRS1133 CCA 33; NRS1135 CCA 33; NRS1134 CCA 33; NRS1098 CCA 33;

NRS1101 CCA 33; NRS1092 CCA 33; NRS0246 CCA 17; NRS0053 CCA 13; NRS1481 CCA 47;

DJM239 BTNR

Ixora javanica (Blume) DC. - NRS0230 CCA 17; NRS0380 CCA 17; NRS1065 CCA 17

Ixora lobbii King & Gamble - NRS2410 CCA 13; NRS2091 CCA 17; NRS2064 CCA 16; NRS1667 r

CCA 5 sae

Ixora pendula Jack - NRS0240 CCA 17; NRS0379 CCA 17; NRS1238 CCA 54; NRS1281 CCA 43;

NRS1294 CCA 43; NRS310 BTNR; NRS0104 CCA 24

Jackiopsis ornata (Wall.) Ridsdale - NRS0O749 CCA 10

Lasianthus appressus Hook.f. or L. attenuatus Jack - NRS1291 CCA 43; NRS0998 CCA 13; NRS1178

CCA 18; DJM203 BTNR

Lasianthus attenuatus Jack - NRSO023 CCA 13

Lasianthus constrictus Wight - NRS0034 CCA 13; NRS0482 CCA 17; NRS1008 CCA 18; NRS1006

CCA 13; NRS1011 CCA 18; NRS1182 CCA 18; NRS1172 CCA 18

Lasianthus cyanocarpus Jack - NRS1464 CCA 47; NRS1897 CCA 16

Lasianthus densifolius Mig. - NRS0829 CCA 11; NRS0756 CCA 18; NRS1157 CCA 16; NRS1073 CCA

17; NRS1196 CCA 7; NRS1200 CCA 7; DJM312 BTNR

Lasianthus griffithii Wight - NRS1949 CCA 18

Lasianthus perakensis King & Gamble - NRS0103 CCA 24; NRS1155 CCA 16

Lasianthus ridleyi King & Gamble - NRS0306 BTNR

Lasianthus scabridus King & Gamble - NRS2048 CCA 24

Lasianthus tomentosus Blume - DJM6 BTNR

Lucinaea membranacea King - NRS0862 CCA 18; NRS1022 CCA 18; NRS0188 CCA 18

Morinda citrifolia L. - NRS0167 CCA 18; NRS0669 CCA 17; NRS0986 CCA 32

Morinda umbellata L. - NRS0940 CCA 28; NRS1121 CCA 28; NRS2411 CCA 18

Mussaenda erythrophylla Schum. & Thonn. - NRS1401 CCA 7

Mussaenda glabra Vahl - NRS0266 CCA 17; NRS0041 CCA 13; NRS0047 CCA 13; NRS0755 CCA 18;

NRS1086 CCA 20; NRS1188 CCA 18; NRS0326 CCA 30

Mussaendopsis beccariana Baill. - NRS0991 CCA 13

Ophiorrhiza singaporensis Ridl. - NRS0165 CCA 18; NRS1975 CCA 18; DJM160 BTNR

Paederia scandens (Lour.) Merr. - NRS0982 CCA 26; NRS0983 CCA 32

Pavetta wallichiana Steud. - NRS0647 CCA 17; NRS0642 CCA 17; NRS0123; NRS0828 CCA 11;

NRS0759 CCA 18; NRS0854 CCA 18; NRS1141 CCA 33; NRS1072 CCA 17; NRS0692 CCA 45;

NRS1013 CCA 18

Porterandia anisophylla (Jack ex Roxb.) Ridl. - NRS0852 CCA 18; NRS0507 CCA 36; NRS0764 CCA

18; NRS0839 CCA 11; NRS0850 CCA 18; NRS0616 CCA 17; NRS0532 CCA 47

Psychotria sp. - NRS1067 CCA 17

Psychotria helferiana Kurz - NRSO789 CCA 11; NRS1405 CCA 7; NRS1565 CCA 15; NRS1803 CCA

48; NRS1861 CCA 16

Psychotria maingayi Hook.f. - NRS1863 CCA 16

Psychotria obovata Wall. - NRS1148 CCA 33

Psychotria ovoidea Wall. - NRS0469 CCA 46; NRS0329 CCA 30; NRS0460 CCA 46; NRS1440 CCA

43; NRS1543 CCA 28; NRS1623 CCA 33; NRS1659 CCA 46; NRS2215 CCA 43; NRS2284 CCA 28;

NRS2529 CCA 44

Psychotria penangensis Hook.f. - NRSO770 CCA 18; NRS0874 CCA 18; NRS1400 CCA 7; NRS1497

CCA 49; NRS1629 CCA 33; NRS2596 CCA 6

Psychotria rostrata Blume - NRS2153 CCA 17; NRS2331 CCA 6

Psychotria sarmentosa Blume - NRS0173 CCA 18; NRS1071 CCA 17; NRS0626 CCA 17; NRS1663

CCA 46; NRS1825 CCA 15; NRS2045 CCA 24; NRS2125 CCA 12; NRS2192 CCA 25; NRS2266 CCA

28; NRS2303 CCA 7; NRS2352 CCA 45; NRS2566 CCA 43

Psydrax sp. 10 - DJM250 BTNR

Rothmannia macrophylla (R.Br. ex Hook.f.) Bremek. - NRS1268 CCA 5; NRS2003 CCA 24; NRS2046

CCA 24

Tarenna odorata (Roxb.) B.L. Rob. - NRS0083 CCA 24; NRS0871 CCA 18; NRS2138 CCA 17

Tarenna stellulata (Hook.f.) Ridl. - NRS2065 CCA 16

Timonius wallichianus (Korth.) Valeton - NRS0857 CCA 18; NRS0092 CCA 24; NRS0210 CCA 25;

NRS0762 CCA 18; NRS0920 CCA 6; NRS0840 CCA 11; NRS1269 CCA 5; NRS1628 CCA 33; NRS1644

CCA 15; NRS2060 CCA 24; NRS2208 CCA 43; NRS2391 CCA 45

Uncaria cordata (Lour.) Merr. - NRS0556 CCA 46; NRS0233 CCA 25; DJM172 CCA 18

Uncaria gambir (Hunter) Roxb. - NRS1164 CCA 16; NRS1355 CCA 24

Uncaria lanosa Wall. var. glabrata (Blume) Ridsdale - NRS0508 CCA 36

Uncaria longiflora (Poir.) Merr. - NRS1898 CCA 16, NRS2224 CCA 43

Urophyllum blumeanum (Wight) Hook.f. - NRSO752 CCA 18; NRS0799 CCA 11; NRS0617 CCA 17;

NRS1531 CCA 13; DJM204 BTNR

Urophyllum glabrum Wall. - NRS0O162 CCA 18; NRS0148 CCA 18; NRS0163 CCA 18; NRS0253 CCA

17; NRS0296 BTNR; NRS0592 CCA 53; NRS0413 CCA 47; NRS0531 CCA 47; NRS0528 CCA 36;

NRS0330 CCA 30; NRS0621 CCA 17; NRS1380 CCA 49; NRS1813 CCA 48; NRS1885 CCA 16

Urophyllum griffithianum (Wight) Hook.f. - NRS0688 CCA 45; NRS1247 CCA 54; DJM384 BTNR

Urophyllum hirsutum (Wight) Hook.f. - NRS0687 CCA 45; NRS0300 BTNR; NRS1315 CCA 335;

NRS1284 CCA 43; DJM383 BTNR

Urophyllum sp. 2 of Wong (Tree Flora Vol. 4) - NRS0096 CCA 24; NRS0176 CCA 18; NRS0790 CCA

11; NRS1192 CCA 7; NRS1162 CCA 16; NRS1191 CCA 7; NRS1057 CCA 41; NRS0812 CCA 11;

NRS1595 CCA 18; NRS2141 CCA 17; DJM125 BTNR

Urophyllum streptopodium Wall. ex Hook.f. - NRS2609 CCA 6; DJM152 BTNR

Rutaceae

Citrus medica L. - NRS0071 CCA 25

Clausena excavata Burm.f. - NRS0524 CCA 36; NRS0973 CCA 26

Glycosmis chlorosperma (Blume) Spreng. - NRS0262 CCA 17; NRS0798 CCA 11

Luvunga crassifolia Tanaka - NRS0635 CCA 17

Murraya koenigii Spreng. - NRSO988 CCA 26

Santalaceae

Dendrotrophe varians (Blume) Mig. - NRSO0085 CCA 24; NRS1210 CCA 7

Sapindaceae

Cardiospermum halicacabum L. - NRS0902 CCA 6

Guioa pubescens (Zoll. & Moritzi) Radlk. - NRS0935 CCA 28; NRS1122 CCA 28

Nephelium cuspidatum Blume var. eriopetalum (Mig.) Leenh. - NRS2432 CCA ?

Nephelium lappaceum L. - NRS0006 CCA 13; NRS0927 CCA 6; NRS0075 CCA 24; NRS1403 CCA 18

Sapotaceae

Palaquium obovatum (Griff.) Engl. var. obovatum - NRS0351 CCA 30

Planchonella maingayi (C.B. Clarke) van Royen - NRS0321 BTNR

Planchonella obovata (R.Br.) Pierre - NRS0412 CCA 47

Scrophulariaceae

Adenosma javanicum (Blume) Koord. - NRS0225 CCA 25

Limnophila sessiliflora (Vahl) Blume - NRS0836 CCA 11; NRS1195 CCA 7; NRS1194 CCA 7; NRS1211

CCA 7

Limnophila villosa Blume - NRS0063 CCA 25; NRS0068 CCA 25

Lindernia crustacea (L.) F.Muell. - NRS0024 CCA 13

Lindernia elata (Benth.) Wettst. - NRS1212 CCA 7; NRS0484 CCA 17

Striga asiatica (L.) Kuntze - NRS0494 CCA 17

Smilacaceae

Smilax bracteata Presl var. barbata (Wall. ex DC.) Koyama - NRS0129 CCA 18; NRS0944 CCA 28;

NRS0417 CCA 47; NRS1545 CCA 28; NRS2269 CCA 28

Smilax calophylla Wall. - NRS0314 BTNR; NRS0997 CCA 13

Smilax leucophylla Blume - NRS1576 CCA 15

-———

. Solanaceae

Solanum torvum Sw. - NRS0907 CCA 6; NRS0739 CCA 10

Sterculiaceae

Commersonia bartramia (L.) Merr. - NRS0652 CCA 17; NRS1018 CCA 18; NRS1156 CCA 16; NRS0558

CCA 46

Melochia corchorifolia L. - NRS0945 CCA 28; NRS0487 CCA 17

Sterculia sp. - NRS0O804 CCA 11; NRS1021 CCA 18

Sterculia coccinea Jack - NRS0370 CCA 17; NRS0696 CCA 45; NRS0700 CCA 45; NRS1275 CCA 5;

NRS0580 CCA 53; NRS0054 CCA 13; NRS0339 CCA 30; NRS1205 CCA 7; NRS1199 CCA 7; NRS0515

CCA 36; NRS0121 CCA 18; NRS0569 CCA 46; NRS1152 CCA 16

Sterculia rubiginosa Vent. - NRS0719 CCA 10

Symplocaceae

Symplocos adenophylla Wall. ex D. Don - NRS0748 CCA 10

Taccaceae

Tacca integrifolia Ker-Gawl. - NRS0456 CCA 46; NRS0336 CCA 30; NRS1599 CCA 45; NRS1929

CCA 25

Theaceae

Adinandra dumosa Jack - NRS0198 CCA 18; NRS0353 CCA 30; NRS0734 CCA 10; NRS0930 CCA 6;

NRS0949 CCA 28

Eurya acuminata DC. - NRS0052 CCA 13; NRS0486 CCA 17; NRS0222 CCA 25; NRS0563 CCA 46;

NRS0865 CCA 18

Gordonia singaporiana Wall. ex Ridl. - NRS0319 BTNR

Pyrenaria acuminata Planch. ex Choisy - NRS1016 CCA 18; NRS0619 CCA 17; NRS0705 CCA 45

Thymelaeaceae

Wikstroemia ridleyi Gamble - NRS0248 CCA 17; NRS2340 CCA 47

Tiliaceae ret:

Grewia acuminata Juss. - NRS0185 CCA 18; NRS0646 CCA 17; NRS0117 CCA 18; NRS0801 CCA 11; a) ;

NRS0870 CCA 18; NRS0848 CCA 18 41

; eee:

Microcos blattaefolia (Corner) Rao - NRS0653 CCA 17; NRS0822 CCA 11 a re ‘i “ie

Ulmaceae

Gironniera nervosa Planch. - NRS0066 CCA 25: NRS0622 CCA 17

Trema cannabina Lour. - NRS0338 CCA 30; NRS0153 CCA 18; NRS0397 CCA 30; NRS0746 CCA 10:

NRS0611 CCA 17

Trema tomentosa (Roxb.) Hara - NRS0740 CCA 10; NRS0896 CCA 6; NRS0152 CCA 18

Umbelliferae

Centella asiatica (L.) Urb. - NRSO288 CCA 17

Urticaceae

Poikilospermum suaveolens (Blume) Merr. - NRS0305; NRS0472 CCA 46; NRS1020 CCA 18; NRS0526

CCA 36; NRS0837 CCA 11

Verbenaceae

Callicarpa longifolia Lam. - NRS0893 CCA 6

Clerodendrum deflexum Wall. - NRS0533 CCA 47; NRS1009 CCA 18; NRS0830 CCA 11; NRS1289

CCA 43; NRS1160 CCA 16; NRS1131 CCA 33; NRS1097 CCA 33; NRS1150 CCA 33

Clerodendrum laevifolium Blume - NRS0154; NRS0363 CCA 30; NRS0823 CCA 11; NRS0690 CCA

45; NRS0758 CCA 18; NRS0072 CCA 25; NRS0154 CCA ?; NRS0466 CCA 46

Clerodendrum paniculatum L. - NRS0846 CCA 18; NRS1001 CCA 13

Clerodendrum villosum Blume - NRS0097 CCA 24: NRS0005 CCA 13; NRS0038 CCA 13; NRS0950

CCA 28

Lantana camara L. - NRS0765 CCA 18

Stachytarpheta indica (L.) Vahl - NRS0175 CCA 18

Vitex pinnata L. - NRS0070 CCA 25: NRS0731 CCA 10; NRS0136 CCA 18

Vitex vestita Wall. ex Schau. - NRS0220 CCA 25; NRS1139 CCA 33; NRS1137 CCA 33

Viscaceae

Viscum ovalifolium Wall. ex DC. - NRS0389 CCA 17

Vitaceae

Ampelocissus elegans (Kurz) Gagnep. - NRS0317 BTNR; NRS0431 CCA 47; NRS0215 CCA 25;

NRS0722 CCA 10; NRS1149 CCA 36; NRS0807 CCA 11; NRS0011 CCA 13; NRS0554 CCA 46

Ampelocissus gracilis (Wall.) Planch. - NRS1296 CCA 43; NRS1117 CCA 28; NRS1170 CCA 18;

NRS1273 CCA 5; NRS1310 CCA 35; NRS0530 CCA 47

Cayratia novemfolia (Wall.) Burkill - NRS0393 CCA 17

Cissus hastata (Miq.) Planch. - NRS0O187 CCA 18; NRS0853 CCA 18

Cissus repens Lam. - NRS0884 CCA 18

Pterisanthes polita (Miq.) Laws. - NRS0127 CCA 18; NRS1288 CCA 43; NRS1179 CCA 18; NRS1040

CCA 18; NRS1184 CCA 18; NRS1120 CCA 28; NRS1292 CCA 43; NRS0112 CCA 24

Zingiberaceae

Alpinia galanga (L.) Sw. - NRS0697 CCA 45

Alpinia purpurata (Vieill.) K. Schum. - NRS0709 CCA 45

Amomum xanthophlebium Baker - NRS0636 CCA 17

Globba leucantha Migq. - NRS0368 CCA 17

Hornstedtia leonurus (Koenig) Retz. - NRS1100 CCA 33

Hornstedtia scyphifera (Koenig) Steud. - NRS0409 CCA 47; NRS1304 CCA 43; NRS1151 CCA 36:

NRS1061 CCA 41; NRS0259 CCA 17; NRS1734 47

Zingiber griffithii Baker - NRSO787 CCA 18; NRS1173 CCA 18; NRS1127 CCA 28; NRS1004 CCA 13

Zingiber puberulum Ridl. - NRSO309

The Tree Communities of the Central Catchment Nature

Reserve, Singapore

Wonc YEw Kwan!, CHEw PING TING? and ALI BIN IBRAHINZ

- 89 Soo Chow Garden Road,

Singapore 575526

eB; National Parks Board,

Singapore Botanic Gardens,

Singapore 259569

Abstract

A sample survey was conducted to study the tree communities of the Central Catchment

Nature Reserve. The forests were stratified into types using vertical aerial photographs. Some 62

sampling units, each about 0.2 ha in size, were laid down in 3 forest types, consisting mainly of

secondary forests, of different degrees of maturity, and relatively undisturbed patches of primary

forests. The sampling percent was 0.8

The trees were measured for girths down to 30 cm and identified down to species. In all 7,462

trees were sampled and these were found to belong to 499 species, 46 of which could not be identified.

The sample netted in some 20 species of dipterocarps with 154 individuals. A surprising discovery is

the presence of 3 Shorea curtisii in a patch of primary Lowland Dipterocarp Forest, sensu Symington

(1941) north of MacRitchie Reservoir. The species is not known to be associated with this forest type

in Peninsular Malaysia and Singapore. Another distributional record is the discovery of 2 trees of

Shorea ochrophioia in another patch of primary forest, though not within the sample. This belongs to

the Heavy Hardwood (Balau) Group of the genus Shorea and so far none of its members has been

recorded in Singapore.

Based on the trends of the species-area curves, the sample appears to have netted in most of

the secondary forest species but the primary stands are likely to yield many more species if an inventory

of a higher intensity of sampling is carried out.

Stand tables are given to show the distribution of the species in each forest type. Fifty-two

species were found to be common to all the three forest types, there being no dipterocarps amongst

them, as expected.

The stands from the relatively undisturbed patches of primary forests were compared with

those at Bukit Timah. In terms of species complexity some stands of forests of the two places compare

well with one another, but in terms of stand densities, and absolute number of species per unit area,

the stands of the Catchment Reserve appear to be better than those of the Bukit Timah forests.

The secondary forests of the Reserve are supposed to have been developed on degraded soil.

The present edaphic conditions are good.

Introduction

The Central Catchment Nature Reserve (hereafter referred to as the

Catchment Reserve or simply the Reserve), estimated in this study to be about

1,660 ha in extent, occupies a central position on Singapore Island (see Fig.1). The

vegetation is mostly of a secondary nature but patches of primary forest are scattered

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Nature Reserve. Insert shows the position of the Nature Reserves within —

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within the mosaic of secondary forests, undergoing different stages of

succession.

Many qualitative observations concerning the plant communities

within the Reserve have been made in the past. Gilliland (1958) was

probably the first ecologist to have made some quantitative study over a

small area of the forests. Recently Corlett (1991) while studying the

succession in the secondary forests sampled selected areas of the more

matured forests. So far as is known no quantitative sampling has ever been

done to study the tree populations of the patches of primary forests, which

are known to be dipterocarp bearing.

The National Parks Board (NParks), now administering all nature

reserves in Singapore, has in the past few years commissioned various

scientists and specialists to make studies on both the physical features &

biological components of the Reserve. We were asked to study in particular

the tree communities. The study covers an area of about 1,530 ha, with an

overall sampling intensity of 0.8 %, but with much higher proportions of

the sampling units located in older forest types and none in the open areas

with early stages of ecological succession (see Table 1). Three of our

sampling units are located in the Nee Soon Swamp Forest, estimated to be

about 96 ha in extent. Sampling began in early 1992 and a detailed technical

report was submitted to NParks in early 1993 (Wong, 1993).

The sampling points are permanently marked at site for future follow-

up studies. Fig.1 shows the approximate positions of the sampling points.

Method of Study

1. Stratification of the Study Area into Forest Types

The forests were stratified using black and white vertical aerial

photographs of scale 1:20,000 and taken in 1990. Four strata, based on the

structures of the forests, could be recognised and were delineated. The

phototypes, designated as Forest Types 1 - 4 (FT 1, FT 2, FT 3 & FT 4)

were then traced out and a vegetation map produced. The general

characteristics of these forest types, based on both photo appearance &

ground checks, are briefly described below and the estimated area of each

type is given in Table 1.

FT 1. Vegetation of early succession with few scattered trees or groups

of trees, the ground being covered with thick Resam ferns

(Dicranopteris spp.), or tall shrubs, tall grasses and/or sedges. Tangles

of woody climbers are common. The climber Smilax bracteata var.

barbata could be rampant locally, smothering tree crowns.

FT 2. Vegetation with many small trees, 8 to 15 m tall. There is

general closure of the canopy, but gaps with Resam and climber tangles

are still quite common, although in the areas with a canopy cover, the

climbers and ferns may be on their way out. The tree crowns are

small; distinct tree crowns may not be discernible in the aerial photos.

Smilax may still be rampant in places, with tangles of their stems

carpeting the forest floor. The tree population has a high proportion

of Adinandra dumosa and Rhodamnia cinerea. Myrica esculenta may

_also be locally abundant. Advanced growth of sizable trees may be

scattered amongst the smaller trees.

FT 3. Vegetation with larger trees and higher density. Canopy is

generally continuous. In the aerial photos, distinct tree crowns are

discernible, as the larger trees have larger crowns. The canopy may

range from 10 to 20m high. Advanced growth or relics of larger trees

may be present. Rhodamnia cinerea still assumes a high proportion,

although that for Adinandra dumosa may be reduced (see Appendix

2). The ground may still have some Resam ferns, but if present they

exist in a sparse condition. Climbers may still be present. However,

no tangles may be on the ground, except in open gaps.

Composition of the trees assumes a more complex nature with many

more species to the unit area. High forest trees like species of

Calophyllum & many species of Eugenia may be in the admixtures.

Garcinia species are also common.

FT 4. Vegetation with a continuous canopy and much taller trees. The

profile is typically multi-storey. Some areas may have structures and girth

class distribution resembling those of primary forests. Relics left behind by

previous fellings would have attained large to enormous sizes. Isolated

crowns of emergents are clearly visible in the aerial photos. To this

type also belong some patches of near-primary jungles, with the family

Dipterocarpaceae showing some degree of structural and family

dominance.

Included in this type is also the Nee Soon Swamp Forest.

2. Estimation of Areas of the Reserve and Forest Types

A dot-grid was used to make the estimate and the results are

presented in Table 1.

Table 1. Area of Catchment Reserve, with Breakdown into Forest Types

(FT) and Sampling Percentage Therein

Area (Ha) ; 1531.6*

No. of 62

sampling

units

Equivalent

Hectares

Percentage

Sampling

*Note: Including an area of 130 ha not studied in this survey, the total area of the Reserve appears to be in the

region of 1,660 ha. No sampling was done in FT 1, hence the 0’s.

3. Sampling Method

The sampling units were located subjectively in the different FT’s.

As the present study is both an ecological and botanical study, our bias

was to locate more of the sampling units in the more matured forests and

the near-primary forests (FT’s 3 & 4). This is the reason why we did not

sample FT 1, and located only 5 clusters in FT 2, which was found in

essence to be an earlier stage of FT 3. We also tended to put the sampling

units where the trees were.

The sampling unit each consists of a cluster of 4 circles. Each circle is

given a radius of 12.6 m; its area is therefore 449 square metres. The 4

circles together therefore have an area of 1,996 square metres or very near

to 0.2 ha. For some clusters, the 4 circles were laid systematically in the

directions of the cardinal points about, and equidistant from, a centre. This

is done when the forest was uniform. When such a systematic layout

would hit gaps, the 4 circles were sited subjectively where the trees were.

Half the number of the clusters were sited in this way.

In view of the subjective siting of the clusters and also of some of the

circles of the cluster, it has therefore to be noted that the results as presented

here may be on the optimistic side for each forest type and all interpretations of

the results or extra-polation of the data would have to be viewed in this light.

4. Enumeration And Plant Identification

Within each circle of a cluster, all trees with girths equal to or larger than

30 cm were measured for girths at 1.3 m from the ground (referred to as girth

breast height or gbh). A tree with buttresses higher than 1.3 m from the

ground was measured above the buttresses or if these were too high, then

the girth of the tree had to be estimated. For a tree with multiple stems, or

with coppice shoots, if the bifurcation or splitting occurred at below 1.3 m,

then each stem was measured and booked as though it was a separate tree,

if its girth met the minimum requirement. Such cases are very common for

species like Rhodamnia cinerea, Adinandra dumosa and Gynotroches

axillaris. Timonius wallichianus and even Eugenia grandis also occasionally

exhibit such a phenomenon, which is likely to have been induced by fire

during early stages of succession. Vestiges of fire damage of very recent

occurrences could be seen in areas of FT 1.

After measuring the girth, the tree was notched or lightly blazed with

a knife, so as to avoid double accounting. The tree was then identified as far

as possible down to species. As most of the trees are sterile at any one

period of the year, identification in most cases are based on leaf and bark

characteristics, including exudates from within the bark. If the tree could not

be identified fully in the field then collection of leaf specimens had to be

made for further identification in the herbarium.

The book was closed for every circle.

For the collection of leaf specimens, in most cases it was fairly easy

to pick the right leaves on the forest floor. However, in some cases when

the tree crown was smothered with heavy climbers, this could prove a

difficult task. Indeed in some cases we just failed to know which could be

the right leaves. For these, one would just have to give up and record such

a tree as “unknown”. In the list of trees presented in Appendix 2, the class

at the end of the list labelled “ZU” shows such trees. There were in all 19

such individuals, each assumed to be a species. In other cases, although we

had good specimens from the field, all matching work in the herbarium

nevertheless failed. There were 37 such individuals and we have placed

them under 27 species. The numerals prefixed with a “Z” at the end of the

plant list in Appendix 2 show these unidentifiable plants.

(42

There are thus in all some 46 species of trees, with 56 individuals,

which could not be identified. This is a small percentage (0.73) considering

that there are 7,462 trees in the sample of 62 clusters.

Results and Discussions

1. Floristic Composition

(a) General

The sample of 62 clusters netted in a total of 7,462 individual trees. A

breakdown showing the number of clusters in each forest type and the number of

species and individuals sampled therein is summarised in Table 2. FT 4, with 22

clusters, included three clusters located in the Nee Soon Swamp Forest. As

stated earlier, 46 species with 56 individuals could not be identified. Of the

7,406 individuals which were identified, they have been found to belong to 453

species and these in turn fall under 63 families.

The list has included a couple of new records for dipterocarps. A few

more of the non-dipterocarp species may also turn out to be new records for

Singapore and checking is continuing at the Herbarium to confirm this.

The species are listed in Appendix 2 with indications of the number of

individuals occurring in each forest type. Table 2 gives a summary of some of

the stand attributes.

FT 2 and FT 3 consist of stands with an abundance of secondary forest

species. The ubiquitous species is Rhodamnia cinerea. It occurs in all the forest

types and in 50 of the total sample of 62 clusters. Even in the 15 relatively

undisturbed primary forest stands of FT 4, some 60 individuals are found in 6 of

them. It is, however, not present in the 3 clusters located within the Nee Soon

Swamp Forest.

One reason why a secondary forest species like Rhodamnia cinerea appears

to be so overwhelmingly present is our treatment of coppiced stems as

“individuals” in our enumeration under certain conditions stated earlier. If we

had considered only rooted frequency, then its overall numbers would be

substantially less. We have, however, not compiled the data in that manner.

Table 2. Species And Individuals In Different Forest Types (FT)

(Species include the 46 unidentifiable species with 56 individuals)

No. of a gfe 22 62

Clusters

No. of

Individuals

Total No. of

Species

No.of identified

species

Families of the

identified species

*Note : The totals for species & families do not agree with the sum of the individual values because of overlap

in species distribution in the different forest types.

The other dominant secondary species are Adinandra dumosa,

Timonius wallichianus and Macaranga conifera. All these secondary species,

appear to diminish in proportion in the population as the secondary forest

gets older. Reference to Appendix 2 will show that Adinandra forms 11.5%

of the population in FT 2 and it drops to 7.1% in FT 3. In FT 4 the

percentage is only 1.3. Rhodamnia drops from 31.3%, through 27.9% to

7.3%. The drop for Macaranga conifera is dramatic from 17.1%, through

1.0% to 0.22%. Timonius wallichianus drops from 4.3% to 2.7% and persists

with the latter proportion in FT 4. A species like Gynotroches axillaris,

which is of frequent occurrence in old secondary forests and primary forests,

on the other hand shows about 1.0% in FT 2, 2.4% in FT 3 and 2.0% in FT

One species which is not actually a secondary forest species but

nevertheless registers strongly in FT 3 is Garcinia parvifolia. Its presence

in FT 2 is only 4 individuals out of a population of 823, or less than half a

percent, but its proportion goes up to nearly 5%, or 10 times more, in FT

3, showing that originally fruits of this species from perhaps the primary

forests had come in to seed up the young secondary forests and as the

resultant stands and individuals mature they in turn produce seeds to enable

the species to proliferate further. Seedlings of Garcinia parvifolia are found

in large numbers on the forest floor. The fruits of Garcinia are eaten by

bats and rodents and these must have been responsible for spreading the

species. Reference to Appendix 2 will show similar pattern of succession

from FT 2 to FT 3 for some of the Calophyllum species, Gynotroches axillaris,

Elaeocarpus mastersii, and Litsea elliptica. Whether the same agents are

responsible for their spread & proliferation is uncertain.

We present in Table 3 a list of the species common to all the three forest

types. There are 52 of them. The list also shows their relative abundance in each

of the forest types. Noticeably, but not surprisingly, there are no dipterocarps in

the list. However, quite a number of these are high forest species and they are

making their presence felt in FT 2 and FT 3, which are essentially secondary

forests. Looking at the totals of the list, we note that the total number of

individuals of these 52 species amounts to 4,814, or 64.5% of the total individuals

in the sample population. The total of 52 is only about 11.6% of the species total

of 499.

(b) The Dipterocarps

The dipterocarps are perhaps the most important tree family in the primary

lowland forests in Malaysia and Singapore. Twenty-five clusters, 18 in FT 4 and 7

in FT 3, have species of dipterocarps. Taking the whole girth range of => 30cm,

there are 154 individuals in the 25 clusters. These belong to 20 species of

this family, and the most widespread, though not the most abundant, species

is Vatica ridleyana, with 15 individuals occurring in 11 of the 25 clusters.

The distribution of the other species is shown in the list in Table 4. Vatica

ridleyana has individuals which are relatively small trees compared with

other dipterocarps. If we take the greater girths of the sample, say => 61

cm, we have a population of 114 individuals, then the list is topped by

Shorea pauciflora with 13 individuals distributed over 10 clusters. Specimens

of S. pauciflora are huge, the largest encountered has a girth of 386 cm. In

contrast the largest tree of V. ridleyana has a girth of only 140 cm.

Concerning the dipterocarps, one very interesting and indeed

surprising find is the presence of Seraya (Shorea curtisii) in Cluster 13 of

the Catchment Reserve. The forest type in which this cluster occurs is

essentially Lowland Dipterocarp Forest (LDF), sensu Symington (1941)

and in Peninsular Malaysia this species is not known to grow in LDF.

Table 3. Species Common to All Forest Types

Adinandra dumosa 95

Alstonia angustifolia

Alstonia angustiloba

Antidesma cuspidatum

Aquilaria malaccensis

Archidendron clypearia

| Arthrophyllum diversifolium

Artocarpus rigidus

Co wOon nan nan ff WO NO

Artocarpus scortechinii

Beilschmiedia madang

bia

Buchanania sessilifolia

Calophyllum pulcherrimum

Calophyllum tetrapterum

—

Campnosperma auriculatum

book

Campnosperma squamatum

—

Castanopsis wallichii

eosk

Cratoxylum arborescens

Decaspermum fruticosum

eak

Dysoxylum cauliflorum

Elaeocarpus ferrugineus

Elaeocarpus mastersii

Elaeocarpus petiolatus

Endospermum diadenum

iw)

Eugenia glauca

Eugenia grandis

Eugenia longiflora

Eugenia microcalyx

Euodia glabra

Fagraea fragrans

Ficus lamponga

Garcinia parvifolia

Gironniera nervosa

Gynotroches axillaris

Horsfieldia polyspherula

Ixonanthes reticulata

Knema intermedia

Lithocarpus ewyckii

Litsea elliptica

Litsea firma

Litsea grandis

Macaranga conifera

Macaranga triloba

Porterandia anisophylla

Prunus polystachya

Pternandra echinata

Rhodamnia cinerea

Scorodocarpus borneensis

Streblus elongatus

Styrax benzoin

Timonius wallichianus

Vitex pinnata

Xanthophyllum ellipticum

Table 4. Dipterocarps in 25 Clusters, 18 in FT 4 and 7 in FT 3.

(All species => 30 cm girth)

(15 clusters in FT 4 are relatively undisturbed Primary Forests)

(INDI = No.of individuals, CSP = No. of clusters in which the species occur.)

i ee

Vatica ridleyana

Shorea pauciflora

Shorea macroptera

Dipterocarpus sublamellatus

Shorea parvifolia

Anisoptera megistocarpa

Hopea griffithti

Hopea mengarawan

oO oo SV NHN AM Bw WN =

Shorea gibbosa

—

Shorea leprosula

—

Vatica maingayi

—_

Dipterocarpus grandiflorus

—

Shorea bracteolata

pond

—

Shorea ovalis

Dipterocarpus cornutus

Shorea gratissima

Shorea platycarpa

Dipterocarpus apterus

Shorea curtisii

= WD KSF& NYO Lf NN W

Vatica ?ridleyana

(Same Stands but with species => 61 cm girth)

Shorea pauciflora

Shorea macroptera

Shorea parvifolia

Vatica ridleyana

Anisoptera megistocarpa

Dipterocarpus sublamellatus

Shorea gibbosa

Shorea leprosula

Oo wont nd nr Ff WY NY

Hopea griffithii

am)

Dipterocarpus grandiflorus

jem

inn

Hopea mengarawan

—_

Shorea bracteolata

jamal

oS)

Shorea gratissima

cls

Shorea ovalis

faery

Vatica maingayi

Dipterocarpus apterus

feak

—

Dipterocarpus cornutus

—_

Shorea curtisii

= WO Lf —-|& WH LF LH NY LL

= = me me NN NHN NO NW NHN HNO WO Lf LH Hh Hh HO OW CV

Shorea platycarpa

There, the species is found in the hill forests in the Main Range and other

localities, generally beginning to occur at an elevation of about 300 m asl

and rising up to 800 m asl, although in some Coastal Hill Forests, sensu

Symington (1941) it begins to occur at much lower elevations.

In Singapore hitherto Seraya was known only from Bukit Timah.

The forest at Bukit Timah is a Coastal Hill Forest according to Symington’s

classification.

Four Seraya trees were found, one of which was dead. The elevation

of the site as shown in a topographic map is about 30 - 40 m asl. All four

trees have attained a fair size. The three living ones have girths of 210, 216,

and 267 cm. The trees are growing on a slight slope and regeneration

appears to be quite numerous on the ground, with the taller ones having

reached a height of about 3 to 4 m.

We also stumbled upon two Shorea ochrophloia, one near Cluster 21

and another near Cluster 55. This belongs to the Balau (Heavy Hardwood)

Group of the genus Shorea. This is a new record for Singapore. Subsequent

to this discovery, two more trees were found in a sample plot in Bukit

Timah Nature Reserve. The plot has been used in a study undertaken by

the Smithsonian Tropical Research Institute in collaboration with the

National Institute of Education in Singapore. Hitherto, the heavy hardwood

Shoreas have not been found in Singapore, although they are of common

or sporadic occurrences in the Malaysian jungles, both in the lowland and

in the hill forests. It is a matter for conjecture as to why we do not see

more of the balaus in the Catchment Reserve. One reason could be that

their timbers are naturally durable and were therefore continuously sought

for constructional purposes during the early days of timber utilisation,

when the technology for preservation had not been developed, taking not

only big trees for conversion into sawn timbers but also pole sized timbers

for rustic uses, perhaps also for use as firewood in the cooking of gambier,

because dense timbers, such as they are, normally have higher calorific

values. If the balaus were in the Catchment forests before, it is perhaps the

continuous exploitation of small poles in addition to big sized timbers that

had spelt their doom. They are very slow growing and conceivably poles

were taken out even before they had reached reproductive age.

Another interesting find is Dipterocarpus apterus. Although not

exactly a new record, only one specimen had been collected near MacRitchie

Reservoir in 1957. We found the only specimen in our sample in Cluster

58, at the extreme west of Seletar Reservoir.

As pointed out earlier FT 3, essentially forests of a secondary nature,

also has 7 stands with dipterocarps. This may give the impression that they

are making a comeback in the secondary forests. This, however is not the

case; it is more the outcome of our having used structure to delineate the

forests into photo-types. In this process, highly disturbed forests with

remnants of dipterocarps have been classified as FT 3.

2. Degrees of Complexity of the Tree Flora

We use the conventional Mischungsquotient (Richard, 1964), which is

simply the ratio of the number of individuals per species of a population,

to show the complexity. Under normal stand densities, the smaller the

ratio, meaning few individuals to the species, the more complex is the

specific composition of a forest. The ratio has been worked out for each

cluster by forest type and the results are presented in Appendix 1. Looking

at the mean values of the ratios for the forest types, as expected, one notes

a gradation from a high to a low value as the forests mature from FT 2,

through FT 3 to FT 4. The mean value for FT 2 is 7.3, that for FT 3, 3.9,

and that for FT 4, 2.1. This shows higher complexity as the forests mature.

In simple terms, for the stands of FT 2, for every species we encounter,

there may be over 7 individuals in the forests, and the respective figures

for FT 3 & FT 4 are about 4 and 2 individuals.

3. The Species-Area Curve

In our sample we have netted in 499 species of the trees with the

minimum girth of 30 cm. To what extent have we exhausted the species list

or are we likely to find more species of the same girth range? To give us

some indications of this, we have plotted two species-area curves, one for

FT 4 alone and the other for FT 2 & FT 3 combined. The reason for

combining FT 2 & FT 3 is that these stands are located in secondary or

highly disturbed, but not clear felled, regenerated forests, while FT 4 are in

patches of primary forests, although 5 of these stands also have species

lists suggesting they are matured secondary forests.

There are several ways of plotting a species-area curve (Greig-Smith,

1964). The method we have used here is perhaps the least efficient according

to him. The area of a particular point in the graph is simply the cumulative

total of areas of clusters added up to that point. Likewise the corresponding

cumulative total for the species of the clusters is used. For the toting up we

followed the numerical order in which the clusters were sampled in each

forest type. FT 2 & FT 3 has a combined population of 5,209, and FT 4

2,253, individuals. The respective number of species are 293 and 417.

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No.of species

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400

300

200

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Sample Area (na)

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Fig. 2. Species / area curves

FT 2/3: S’dary foreests, 40 plots, 293 spp

FT 4: Mainly primary,, 22 plots, 417 spp

The two species-area curves are presented in Fig. 2. They show very

characteristic trends. The curve for FT 2/3 shows not only a more gradual

gradient, indicating more gradual species recruitment as the area increases, but

also a definite gradual flattening out, indicating that our sample has perhaps

netted in most of the species. Contrasting, the curve for FT 4 rises much more

steeply and at the end still shows no flattening out, indicating that our sample of

22 clusters has in no way yet got most of the species. As a corollary, if we

sample FT 4 more thoroughly, we are likely to net in many more species.

4. Stand Density Attributes

Table 5 shows the stem densities and basal areas by girth classes. We are

showing separately these attributes for FT 2 and FT 3 combined (40 clusters)

and FT 4 by itself (22 clusters) for comparison. In both cases, the whole range of

girth classes of =>30 cm is used.

It can be seen that the structure, in terms of girth class distribution,

is typical of uneven-aged forests, with a high proportion of small stems,

and as girth size increases, the number of trees drops rapidly. For the

secondary forests one would expect the girth distribution to show a

truncation after a certain point. The fact that FT 2/FT 3 have individuals

dragging into the higher girth classes could be due to FT 3 having a few

stands with relics or advanced growth.

In the FT 2/FT 3 forests, the strong presence of young poles in the

30 - 60 cm class shows that recruitment for the forest as a whole is good

and this augers well for the Catchment Reserve.

As expected, FT 4 does show that the population has many individuals with

very large girths, far outnumbering those of the younger stands in FT 2 & FT 3.

5. Vegetation dynamics

Although we did not sample FT 1, there is no doubt that within the

Catchment Reserve this open type of vegetation would in time develop into the

FT 2 type of forests, thence to FT 3, in the natural succession. The speed with

which this process will take place would no doubt depend on edaphic, aerial and

biotic factors. From FT 1 to FT 2, incidence of bush fire could play an important

role. While it may destroy a stand of small trees, it may on the other hand burn up

an existing climber or Resam thicket to enable the area to be seeded up by tree

species. From FT 2 to FT 3, the presence of nearby seed sources would certainly

speed up succession. The existing stands of FT 4 and the more matured stands of

FT 3 will form such sources. Dispersal of such high-forest genera like Garcinia,

Calophyllum, Eugenia and Gynotoches from these sources is definitely helped by

birds, rodents and bats.

In general, at the moment, compositions of the FT 2 and FT 3 stands

bear close similarities to Stage 3 & Stage 4 respectively of the succession

described by Corlett (1991).

The influence of proximity of a seed source in succession is very

much borne out by the observations of Sim et al (1992). They laid down 7

sample plots in secondary forests now dominated by Adinandra dumosa,

perhaps much like FT 2 in our present study. All but one were isolated

areas, the exception being in the Bukit Timah Nature Reserve. They have

stated that succession in these plots has been arrested due inter alia to low

pH, poor soil nutrients and the lack of a seed source to add new species to

such communities. However, scrutiny of their plant lists reveals the presence

of a substantial proportion of high forest tree species in their Bukit Timah

plot. These species occur exclusively in that particular plot. The species are

Calophyllum pulcherrimum, Calophyllum sp.1, Calophyllum sp.2, Eugenia

sp.l1, Eugenia sp.2, Eurycoma_ longifolia, Gynotroches axillaris,

Ochanostachys amentacea, Palaquium gutta, Psychotria sp., and Santiria

apiculata. It is clear evidence that succession has progressed beyond the

Adinandra/Rhodamnia (equivalent to our FT 2) stage and the factor that is

responsible is undoubtedly a nearby source of seeds of such high-forest

species from within the Bukit Timah Nature Reserve.

6. Forests Of Bukit Timah and Catchment Reserve Compared

(a) Floristic compositions

As the Coastal Hill Forest, sensu Symington (1941), on Bukit Timah

is the only other dipterocarp forest of a primary nature found in Singapore

we would like to compare it with the stands of the primary forest of the

Catchment Reserve which essentially is typical Lowland Dipterocarp Forest.

For this comparison we have taken the sampling units sited in the

relatively undisturbed primary forest stands of the Catchment Reserve,

but excluding the Nee Soon Swamp Forest, and those on Bukit Timah

reported by Wong (1987). There are respectively 15 and 16 clusters. As the

Bukit Timah sampling was for trees =>24 inches (61 cm) corresponding

data of the Catchment Reserve were used.

Table 6. Comparison of Dipterocarps In Bukit Timah (BT)

and the Central Catchment Reserve (CR)

(All trees with girth =>61 cm. The sample at BT has 16 clusters, at CR 15 clusters.)

No.of individuals

SPECIES

Anisoptera costata

Anisoptera megistocarpa

Dipterocarpus apterus

Dipterocarpus caudatus

ssp penangianus

Dipterocarpus cornutus

Dipterocarpus grandiflorus

Dipterocarpus sublamellatus

Hopea griffithii

Hopea mengarawan

Shorea bracteolata

eae

Shorea curtisit

Shorea gibbosa

Shorea gratissima

Shorea leprosula

Shorea macroptera

Shorea ovalis

Shorea parvifolia

Shorea pauciflora

Vatica maingayi

Vatica ridleyana

Oo ww wo kr On ff fF We fF WO ~

Vatica sp.A

Table 6 shows that the Catchment Reserve has 18 species of dipterocarps

and Bukit Timah 11 species, this despite the fact that the trees of Bukit Timah

were sampled with 16 clusters while those of the Catchment Reserve with 15.

However, this is to be expected as those clusters sited in the Catchment Reserve

are spread over a much wider area, whereas those in Bukit Timah are located

within a solid block of forests of 75 ha. In terms of individuals, the Bukit Timah

stands appear to have more dipterocarps, there being 102 individuals compared to

88 in the Catchment Reserve sample. This superiority in numbers is due to the

presence of large numbers of Shorea curtisii and Dipterocarpus caudatus ssp

penangianus in the Bukit Timah stands.

(b) Relative floristic complexity & stand densities of the two Areas

We now compare the Mischungsquotients of the two areas. The comparison

is presented in Table 7. It can be seen that the average stand of the Bukit Timah

forest has a smaller quotient of 1.5, compared to 1.9 of the average stand in the

Catchment Reserve. As stated earlier under normal stand densities, the smaller

the quotient, the more complex is the stand. However, it has to be said that a low

stocking, concomitant with the number of species being constant, could also give

rise to small quotients. Looking at Table 7 and 8, the species per cluster of the two

areas did not vary much (BT = 22.9 & CR = 24:5 species/cluster) but the stand

densities of the Bukit Timah forests appear to be consistently much below those

of the Catchment forests, showing that the reason for the lower quotients in Bukit

Timah is exactly what has been just stated. So despite their smaller mean, the

complexity of the forests in absolute terms appears to be not as good as that of the

Catchment forests. To put it in another way, the Catchment forests have overall

denser stocking and a higher number of species per unit area. The Catchment

Reserve being more species rich is also borne out by the fact that for the 16 stands

in the Bukit Timah forests there are 178 species of trees with girths => 61 cm, but

the number in the Catchment Reserve is 215 species.

Our first reaction to the higher stocking density in the Catchment Reserve,

when compared to stands at Bukit Timah, is that the stands in the Catchment

Reserve may have smaller trees, because it is quite common to have young stands

with a high density but with the numbers made up of small trees. To check on this

point we present a comparison of the girth class distribution of the forests of the

two places in Table 8. Looking at this comparison one is amazed by the fact that

for the girth distributions of the two areas, class for class the number of trees per

ha for the Catchment Reserve outnumbers that obtained for the Bukit Timah

forests. And looking at the basal area per tree figures, class for class the size of the

average tree of CR is remarkably similar to that of BT, showing that the higher

stocking of the forests in the Catchment Reserve is achieved not through having a

population of small trees.

Table 7. Mischungsquotients or Number of Individuals per Species

(Comparing the 15 less disturbed clusters of FT 4 with 16 clusters of primary forests of Bukit Timah. All trees

are with gbh => 61 cm. The relevant stands of the Catchment Reserve contain 215 spp., the Bukit Timah

stands only 178 spp.)

Indiv. | Species | quotient Indiv | Species | quotient

28 19 2k

42 28 1.4

38 fas 1.9

42 27 1.6

37 ZS 1.8

39 27 2.1

30 25 2.0

32 20 dik

24 23 1.4

24 20 2.0

32 19 2.0

39 24 at

50 25 ie

27 21 Six

33 23 1.6

23 19 1.2 - -

*Note: the two totals are exactly the same; this is entirely fortuitous.

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actually 48 trees in the sample of the Catchment Reserve; 16 of these trees

have girths exceeding 300 cm. The corresponding numbers for the Bukit

Timah sample are 26 trees and 5 trees. However, the largest tree in the

Bukit Timah sample (a Shorea curtisii) has a girth of 194 inches (or about

490 cm), whereas that in the Catchment Reserve (a Dyera costulata) is

only 424 cm. The largest dipterocarp in the FT 4 in the sample is a Shorea

pauciflora with a girth of 386 cm. (Note: the largest tree we came across in

the Catchment Reserve, not in the sample, but not far from Cluster 13

northwest of MacRithchie Reservoir, is a Dyera costulata with a girth at

breast height of 615 cm.)

From the above comparison of the less disturbed dipterocarp bearing

stands of the two areas, one can conclude that such patches of forests in the

Catchment Reserve are in some ways superior to the stands at Bukit Timah.

Conclusions

The relatively undisturbed stands of the primary forests of the Central

Catchment Nature Reserve are indeed valuable natural assets because of their

very diverse specific compositions and they therefore still contain a very large

gene pool. The stands of secondary forests with very varied specific compositions

also constitute a valuable scientific asset. They have redeveloped by themselves

after the original forests were cleared and the land parcels used for long periods

of cultivation until they were declared as protected catchment areas when

cultivation was stopped. For the areas now with a tree cover, the regrowth period

may vary from SO to well over 130 years. The stands offer a good insight on

plant succession under such conditions and could be used for scientific

comparisons with vegetation developed in other parts of the Tropics.

The present study shows beyond doubt that some of the forest stands have

floristic compositions and structural characteristics similar to those of primary

forests in the Malaysian region. We do not know the exact history of these

stands. Some of these could have been undisturbed; others could have been

reserved for the supply of fuel wood for the gambier plantations and were

therefore exploited to different degrees during the last Century. However,

if they were so disturbed before, the vestiges of disturbance are now

completely absent. Some of these areas surrounding the MacRitchie

Reservoir must have been protected since the construction of this reservoir

in 1867 (Anon., PUB publication, 1985). Over this long period of time,

even if the stands had been disturbed, natural regeneration aided by the

relics, including the dipterocarps, would have made such stands recover

completely, ensuring also their biological diversity.

Stadt, mae i ms, bata tiag ed bie nea

These primary stands are classified under Forest Type 4 (FT 4).

Although according to the aerial phototyping and estimate of areas, this

type amounts to some 280 ha, some of these have been found to be matured

stands of secondary forests with big advanced growth or relics, including

species of dipterocarps. The exact extent of the really primary patches is

likely to be somewhat less than this.

Analysing the stand attributes of these primary stands, one is of the

view that some stands are superior to those on Bukit Timah, traditionally

regarded as the only place in Singapore with primary forests.

The species/area curve of the stands of FT 4 and that for the Bukit

Timah forests show similar form with a sharp gradient without any flattening

out, indicating that in both places more species are expected to be found, if

a more intensive inventory is done.

The stands of secondary forests show the dynamics of succession in

the Catchment Reserve and are developing well. The older of the truly

secondary stands which have developed on land abandoned after prolonged

cultivation (Corlett, 1991) though with superior stocking and with many

high forest species now, yet do not have any species of dipterocarps. As

dipterocarps have very inefficient seed dispersal, seeding under natural

conditions may not happen and such stands might eventually mature into

non-dipterocarp forests.

Corlett (1991) reckons that the secondary forests within the

Catchment Reserve have developed from land severely degraded or

exhausted by cultivation (gambier, pepper & pineapple being important

crops). Degradation and exhaustion, however, were not defined.

Agronomically we would regard severe sheet erosion, so much so that

substantial layers of the solum are gone, and with severe gully formations,

as severe degradation. We have during the 10 months of field work not

found any evidence of severe gully formation and now that the forest

cover is so good with a good litter on the forest floor and a good organic

layer beneath, even if sheet erosion had occurred before, it would be

difficult to discern now. We did, however, see some excavated spots,

trenches here and there, and vestiges of roads and rides.

Looking at some of the trenches present (dug presumably during the

War), the soil is deep and the profile is just as good as any one could see in

a Rengam Series, an Ultisol of granitic origin, which is what the soil in the

Reserve is, excepting of course the swamps and riparian fringes.

Acknowledgements

We are indebted to the National Parks Board and its Executive

Director, Dr Tan Wee Kiat, for having given us the golden opportunity of

conducting the survey for the Central Catchment Nature Reserve and for

having given their permission for the data to be published in this article.

Dr Leong Chee Chiew, Mr Tay Eng Pin and Mr Robert Teo, all staff

members of the Botanic Gardens did much of the spade work to have the survey

initiated and also provided assistance during the field work for which we should

like to express our grateful thanks.

We should also like to express our appreciation for the assistance provided

by Dr Ian Turner and Dr Hugh Tan of the Botany Department, NUS, in checking

the authenticities of many of the plants and plant names.

Last but not least, all of us involved in the survey wish to express our

thanks to the PUB for having made available a boat and boatman for ferrying us

to and from some of our sampling sites.

References

Anon. (1985). Yesterday and today - the story of public electricity, water and gas

supplies in Singapore. Published by the Times Book International for the

Public Utilities Board, Singapore.

Corlett,R.T. (1991). Plant succession on degraded land in Singapore. J. of

Trop. Forest Sc. 4:151-161.

— (1992). The ecological transformation of Singapore, 1819-1990. J. of

Biogeography. 19:411-420.

Gilliland,H.B.(1958). Plant communities on Singapore Island. Gdns’ Bull,

Sing. 17:82-90.

Greig Smith,P.(1964). Quantitative plant ecology. Butterworths, London.

Pg 151 et seq.

Richards, P.W. (1964). The tropical rain forest. Cambridge University Press.

Pg 250.

Sim, J.W.S., Tan, H.T.W., Turner, I.M. (1992). Adinandra belukar: an

anthropogenic heath forest in Singapore. Vegetatio 102:125-137.

}

Symington, C.F. (1941). Malayan Forest Records No.16. Foresters’ Manual

of Dipterocarps. Forest Research Institute of Malaysia.

Wong, Y.K. (1993). The forests and tree populations of the Central

Catchment Nature Reserve. Unpublished technical report of a survey

commissioned by the National Parks Board, Singapore.

— (1987). Ecology of the trees of Bukit Timah Nature Reserve. Gdns’

Bull., Sing. 40:45 76.

Appendix 1. Mischungsquotients or No. of Individuals/Sp.

for All Clusters of the Catchment Reserve

(All trees => 30 cm gbh)

No. of Indiv | No. of Species Michungsquotient

FT No. of Indiv

Appendix 2. Distribution of Species In Forest Types (FT)

(All 62 Clusters, with 5 in Ft2, 35 in Ft3 and 22 in FT4)

(At end of list, Z=trees with collected leaf specimens but could not be identified in the herbarium, ZU=trees

for which we failed to collect leaf specimens & could not be identified)

SPECIES No. of Individuals

rv a

1. | Acronychia porteri Hook. f. 0 9 11 20

2. | Actinodaphne glomerata (Bl.) Nees 0 i 0 I

3. | Actinodaphne malaccensis Hook. f. 0 a 2

4. | Actinodaphne pruinosa Nees 0 6 2

5. | Adenanthera bicolor Moon 0 10 14

6. | Adinandra dumosa Jack O98 Pale 29

7. | Aglaia exstipulata (Griff.) Theob. 0 0 1

8. | Aglaia leucophylla King 0 1 0

9. | Aglaia maingayi (Hiern) King 0 0 6

Aglaia malaccensis (Ridl.) Pannell 0 0 6

Aglaia odoratissima Bl. 0 0 2

Aglaia rubiginosa (Hiern) Pannell 0 0 2

Aglaia sp. 0 0 2

Aidia wallichiana Tirv. 0 13 4

Alangium nobile (Clarke) Harms 0 0 2

Albizia splendens Mig. 0 1 is

Alphonsea maingayi Hook. f. & Thoms. 0 0 5

Alseodaphne bancana Migq. 0 ) 6

Alseodaphne intermedia kosterman 0 ) 1

Alstonia angustifolia Wall. ex A. DC. 19 39 6

Alstonia angustiloba Miq. 1 s) 2

Anisophyllea griffithii Oliv. ) 0 1

Anisoptera megistocarpa Sloot. 0 1 6

Antidesma coriaceum Tul. 0 Ps 2

Antidesma cuspidatum M.A. 1 2 10

Antidesma neurocarpum Miq. 0 0 1

Antidesma salicinum Rid. 0 0 1

Aphanomyrtus skiophila (Duthie) Valeton ) i 0

Aporusa ?nervosa 0 4 1

Aporusa ?penangensis 0 0 1

Aporusa benthamiana Hook. f. 0 3 6

Aporusa bracteosa P. & H. 0 0 3

SPECIES No. of Individuals

Aporusa frutescens Bl.

Aporusa miqueliana M.A.

Aporusa nervosa Hook. f.

Aporusa penangensis (Ridl.) Airy Shaw

Aporusa symplocoides (Hook. f.) Gage

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Aquilaria malaccensis Lamk.

Archidendron clypearia (Jack) I. Niels

Archidendron ellipticum (Bl.) Niels.

Archidendron globosum (B1.) Niels.

Ardisia colorata Roxb.

Arthrophyllum diversifolium Bl.

Artocarpus ?kemando

Artocarpus anisophyllus Miq.

Artocarpus dadah Mia.

Artocarpus fulvicortex Jarrett

Artocarpus heterophyllus Lamk.

Artocarpus integer (Thunb.) Merr.

Artocarpus kemando Miq.

Artocarpus lanceifolius Roxb.

Artocarpus lowii King

Artocarpus nitidus Trec.

Artocarpus rigidus B1.

Artocarpus scortechinii King

Baccaurea ?sumatrana

Baccaurea hookeri Gage

Baccaurea kunstleri King ex Gage

Baccaurea maingayi Hook. f.

Baccaurea minor Hook. f.

Baccaurea parviflora (M.A.) M.A.

Baccaurea racemosa (Reinw.) M.A.

Baccaurea reticulata Hook. f.

Baccaurea sumatrana M.A.

Beilschmiedia kunstleri Gamble

Beilschmiedia madang B1.

Bhesa paniculata Arn.

Bhesa robusta (Roxb.) Ding Hou

Blumeodendron ?tokbrai

Blumeodendron tokbrai (B1.) J.J. Smith

Bouea oppositifolia (Roxb.) Meisn.

Brackenridgea hookeri (Planch.) A. Gray

Buchanania arborescens (B1.) B1.

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Appendix 2 (Continued)

SPECIES No. of Individuals

Buchanania sessifolia B1.

Calophyllum ?ferrugineum

Calophyllum ?rufigemmatum

Calophyllum dispar P.F. Stevens

Calophyllum ferrugineum Rid.

Calophyllum lanigerum Miq. v.austrocoriaceum *:

(T.C. Whitmore) P.F. Steven 1

Calophyllum macrocarpum Hook. f. :

Calophyllum pulcherrimum Wall. ex Choisy |

Calophyllum rigidum Mia.

Calophyllum rubiginosum

Hend. & Wyatt-Smith

Calophyllum rufigemmatum :

Hend. & Wyatt-Smith

Calophyllum sundaicum P.F. Stevens

Calophyllum tetrapterum Mia.

Calophyllum teysmannii Miq.

Calophyllum wallichianum Planch. & Tr.

v. incrassatum

(Hend. & Wyatt-Smith) P.F. Stevens

Campnosperma auriculatum (B1.) Hook. f.

Campnosperma squamatum Ridl.

Canarium ?grandifolium

Canarium grandifolium (Ridl.) Lam

Canarium littorale B1.

Canarium patentinervium Miq.

Canarium pilosum Benn.

Canthium glabrum B1.

Carallia brachiata (Lour.) Merr.

Castanopsis megacarpa Gamble

Castanopsis nephelioides King ex Hook. f.

Castanopsis schefferiana Hance

Castanopsis wallichii King ex Hook. f.

Cheilosa malayana (Hook.f.)

Corner ex Airy Shaw

Chisocheton patens B1.

Chisocheton pentandrus (Blanco) Merr.

Chisocheton sarawakanus (C. DC.) Harms

Cinnamomum iners Reinw. ex B1.

Cleistanthus sumatranus (Miq.) M.A.

Clerodendron laevifolium B1.

Cocos nucifera L.

Cratoxylum ?maingayi

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Appendix 2 (Continued)

SPECIES No. of Individuals

Cratoxylum arborescens (Vahl) B1.

Cratoxylum cochinchinense (Lour.) B1.

Cratoxylum formosum (Jack) Dyer

Cratoxylum maingayi Dyer

Croton laevifolius B1.

Crypteronia griffithii Clarke

Cryptocarya ferrea B1.

Cryptocarya impressa Miq.

Cryptocarya rugulosa Hook. f.

Ctenolophon ?parvifolius

Ctenolophon parvifolius Oliv.

Cyathocalyx ramuliflorus

(Maingay ex Hook. f. & Thoms.) Scheff.

Cyathocalyx ridleyi (King) Sinclair

Dacryodes costata (Benn.) Lam

Dacryodes laxa (Benn.) Lam

Dacryodes rostrata (B1.) Lam

Dacryodes rugosa (B1.) Lam

Decaspermum fruticosum J.R. & G. Forst.

Dehaasia incrassata (Jack) Kostermans

Dialium ?maingayi Baker

Dialium ?platysepalum

Dialium indum L. v. bursa (de Wit) Rojo

Dialium platysepalum Baker

Dillenia grandifolia Wall. ex Hook. f. & Thoms.

Diospyros ?ridleyi Bakh.

Diospyros buxifolia (B1.) Hiern

Diospyros lanceifolia Roxb.

Diospyros maingayi (Hiern) Bakh.

Diospyros pilosanthera Blanco v. oblonga

(Wall. ex G. Don) Ng

Diospyros sp. 1

Diospyros styraciformis King & Gamble

Diplospora malaccensis Hook. f.

Dipterocarpus apterus Foxw.

Dipterocarpus cornutus Dyer

Dipterocarpus grandiflorus Blanco

Dipterocarpus sublamellatus Foxw.

Drypetes pendula Rid.

Durio griffithii (Mast.) Bakh.

Durio singaporensis Rid.

Dyera costulata (Miq.) Hook. f.

Dysoxylum cauliflorum Hiern

pooch

eS oeew o co co eceaocc =

— Go Oo Gor Sa So Si ST NS

WRN PR WABRNR AN FR

o>)

pum

DCDOowWddOWrRrROONRFRNOCN AD

NO Bh ~] WR UO ~) RP RR WOO ~ATR ON WN

Se Cae ee Soe. Ss. Sra = ss) SS & Cre

NO h BR WON OR RR OR UNM FR WwW ~l

aa & O =

a’

—

WOhRPANMO fF NOOO FN -

WwW

NNN N RK OO

Nm Uo

Appendix 2 (Continued)

SPECIES

Dysoxylum densiflorum (B1.) Mig.

Dysoxylum excelsum BI.

Dysoxylum flavescens Hiern

Elaeocarpus ferrugineus (Jack) Steud.

Elaeocarpus floribundus Bl.

Elaeocarpus masterii king

Elaeocarpus nitidus

Jack v.salicifolius (King) Ng

Elaeocarpus palembanicus (Miq.) Corner

Elaeocarpus petiolatus (Jack) Wall.

Elaeocarpus rugosus Roxb.

Elaeocarpus stipularis Bl.

Endospermum diadenum (Miq.) Airy Shaw

Enicosanthum sp. 1

Eugenia ?microcalyx

Eugenia ?nigricans

Eugenia ?pseudosubtilis King

Eugenia cerina Hend.

Eugenia chlorantha Duthie

Eugenia cumingiana Vidal

Eugenia duthieana King

Eugenia filiformis Duthie v.clabimyrtus

(Koord. & Valet.) Hend.

Eugenia glauca King

Eugenia grandis Wight

Eugenia longiflora (Presl) F.-Vill.

Eugenia microcalyx Duthie

Eugenia muelleri Miq.

Eugenia nemestrina Hend.

Eugenia ngadimaniana Hend.

Eugenia nigricans King

Eugenia oblongifolia Duthie

Eugenia pachyphylla kurz

Eugenia papillosa Duthie

Eugenia pauper Ridl.

Eugenia pendens Duthie

Eugenia polyantha Wight

Eugenia ridleyi King |

Eugenia spicata Lamk.

Eugenia subdecussata Duthie

Eugenia tumida Duthie .

Euodia glabra (B1.) BI.

Eurya acuminata DC.

No. of Individuals

S50". OO OO aS te =

NWWrRrPOoOOCOhRN NF ON O

1 3

2 3

37 4

22 on

0 0

) 3

0 6

0 10

0 a) 1

0 2 1

0 0 1

0 6 i

0 0 2

) 3 2

) 4 2

) 1 0

) 5 2

0 0 3

13 1 4

0 1 5

Appendix 2 (Continued)

SPECIES No. of Individuals

Eurycoma longifolia Jack 0

Fagraea fragrans Roxb. 1 : ; :

Fahrenheitia pendula (Hassk.) Airy Shaw 0 0 1 1

Ficus glandulifera (Wall. ex Miq.) King 1 0 1 2

Ficus kerkhovenii Val. 0 ) 1 1

Ficus lamponga Mia. 1 1 1 3

Galearia fulva (Tul.) Mig. 0 1 0 1

Galearia maingayi Hook. f. 0 1 1 2

Ganua kingiana (Brace) Van Den Assem 0 0 5 5

Ganua motleyana (De Vr.) Pierre ex Dubard 0 0 4 4

Garcinia atroviridis Griff. & T. Anders. 0 0 2 2

Garcinia eugeniaefolia Wall. ex T. Anders. 0 88 8 96

Garcinia forbesii King 0 0 = 5

Garcinia griffithii T. Anders. 0 6 1 7

Garcinia maingayi Hook. f. v. stylosa King 0 1 0 1

Garcinia nervosa Miq. 0 0 1 1

Garcinia parvifolia (Mig.) Miq. 4 | 213 40 Dor

Garcinia scortechinii King 0 0 3 3

Gardenia griffithii Hook. f. 0 0 1 1

Gardenia tubifera Wall. 0 0 3 3

Gironniera ?nervosa 0 0 1 1

Gironniera nervosa Planch. 11 60 47 118

Gironniera parvifolia Planch. 0 1 20 21

Gironniera subaequalis Planch. 0 2 8 10

Glochidion superbum Baill. 1 1 0 2

Gluta wallichii (Hook. f.) Ding Hou ) 5 19 24

Gnetum gnemon L. 0 0 1 1

Gonystylus confusus Airy Shaw 0 10 5 15

Gonystylus maingayi Hook. f. 0 0 2 2

Gordonia ?singaporiana Wall. ex Rid. 0 1 0 1

Gordonia multinervis King 1 11 0 12

Grewia blattaefolia Corner 0 1 8 9

Guioa pleuropteris (B1.) Radlk. 0 1 0 1

Guioa pubescens (Z. & M.) Radlk. 0 18 2 20

Gymnacranthera bancana (Miq.) Sinclair 0 0 1 1

Gymnacranthera farquhariana 0 1 12 3

(Hook. f. & Thoms.) Warb.

Gymnacranthera forbesii (King) Warb.

Gynotroches axillaris B1.

Helicia petiolaris Benn.

Heritiera ?javanica B1.

Heritiera elata Ridl.

Heritiera borneensis (Merr.) Kostermans

—

om)

=H OoOonna AN

NNN kK WN ©

WM NM WR DW

Appendix 2 (Continued)

e SPECIES

Heritiera simplicifolia (Mast.) Kostermans

Hevea brasiliensis (Willd. ex A. Juss.) M.A.

Hopea griffithii Kurz

Hopea mangarawan Maiq.

Horsfieldia crassifolia

(Hook. f. & Thoms.) Warb.

Horsfieldia polyspherula

(Hook. f. emend. King) J. Sinclair

No. of Individuals

OO On &

Horsfieldia sucosa (King) Warb. 0 2 6 8

Horsfieldia superba (Hook. f. & Thoms.) Warb. 0 0 1 1

Horsfieldia wallichii (Hook. f. & Thoms.) Warb. | 0 0 2 2

Hymenaea courbaril L. 0 1 0 1

Ilex cymosa B1 1 0 0 11

Ilex macrophylla Hook.f. 0 1 0 it

Irvingia malayana Oliv. & Benn. 0 0 1 1

Ixonanthes icosandra Jack 0 0 5 65

Ixonanthes reticulata Jack 9 8 1 58

Jackiopsis ornata (Wall.) Ridsdale 0 0 1 1

Kibara coriace (B1.) Tul. 0 0 1 e

Kibatalia maingayi (Hook. f.) Woodson 0 1 3 4

Knema communis Sinclair 0 2 6 8

Knema conferta (King) Warb. 0 3 1 4

Knema curtisii 0 3 2 5

(King) Warb. v. paludosa J. Sinclair

Knema furfuracea (Hook. f. & Thoms) Warb.

Knema hookeriana

(Wall. ex Hook. f. & Thoms.) Warb.

Knema intermedia (B1.) Warb.

Knema latericia Elm.

Knema laurina (B1.) Warb.

Knema malayana Warb.

Koompassia malaccensis Maingay ex Benth.

Kopsia singapurensis Rid.

Lansium domesticum Correa

Licania splendens (Korth.) Pranc e

Lindera lucida (B1.) Boerl.

Lithocarpus ?ewyckii

Lithocarpus bennettii (Miq.) Rehd.

Lithocarpus conocarpus (Oudem.) Rehd.

Lithocarpus encleisacarpus (Korth.) A. Camus

Lithocarpus ewyckii (Korth.) Rehd.

Lithocarpus lucidus (Roxb.) Rehd.

Lithocarpus sundaicus (B1.) Boerl.

—

a =

—_

eS)

—_

POF BRR NRK OR RK KN OA WN

ww CO OooOoonajcoeoCcooom

NWrAIWrooetwoooftrNreR BD

—

Appendix 2 (Continued)

Litsea accedens (B1.) Boerl.

Litsea castanea Hook. f.

Litsea costalis (B1.) Kostermans

Litsea elliptica B1.

Litsea erectinervia Kostermans

Litsea ferruginea B1.

Litsea firma Hook. f.

Litsea grandis Hook. f.

Litsea maingayi Hook. f.

Litsea ridleyi Gamble

Litsea robusta B1.

Lophopetalm multinervium Rid.

Lophopetalum wightianum Arn.

Macaranga conifera (Zoll.) M.A.

Macaranga gigantea (Rchb. f. & Zoll.) M.A.

Macaranga lowii King ex Hook. f.

Macaranga triloba (Bl.) M.A.

Madhuca korthalsii (Pierre) Lam

Madhuca malaccensis (Clarke) Lam

Madhuca sericea (Miq.) Lam

Magnolia candolii (B1.) H. Keng

Magnolia elegans (B1.) H. Keng

Mallotus penangensis M.A.

Mangifera foetida Lour.

Mangifera griffithii Hook. f.

Mangifera indica L.

Mangifera subsessilifolia Kostermans

Maranthes corymbosa B1.

Mastixia trichotoma B1.

Melanochyla auriculata Hook. f.

Melanochyla caesia (B1.) Ding Hou

Meliosma lanceolata B1. v. lanceolata

Meliosma simplicifolia (Roxb.) Walp.

Memecylon edule Roxb.

Memecylon floridum Rid.

Memecylon lilacinum Z. & M.

Memcylon megacarpum Furtado

Memecylon paniculatum Jack

Mezzettia parviflora Becc.

Microdesmis caseariifolia Planch.

Monocarpia marginalis (Scheff.) Sinclair

Mussaendopsis beccariana Baill.

Macaranga hypoleuca (Rchb. f. & Zoll.) M.A.

No. of Individuals

OnOrOrONGCOCCCC OF OF COrFR FR Or FP Or OFrP UMNMNrFRCoOooeoe 8 OR Re NOW W

o>)

—_"

mRreRNN RK NRK OR RK RK NOI WNOrR OWDWrROoOdDdanwnorh wOoonwWwWMnNnNnonwndre £& WN ~] NY

jommd

—

~~)

—

tn? Jae a

‘Nn

—"

Ww Ame Ore ff

WNMNR RR DOR OR OO Re

WDQ— Ne NO eR — — bo

i bh

Appendix 2 (Continued)

SPECIES No. of Individuals

Myrica esculenta Buch.-Ham.

Myristica ?guatteriifolia A. DC.

Myristica ?lowiana

Myristica ?maingayi

Myristica cinnamomea King

Myristica elliptica Hook. f. & Thoms.

Mpyristica iners B1.

Myristica lowiana King

Myristica maingayi Hook. f.

Myristicaceae 1

Myristicaceae 2

Myristicaceae 3

Nauclea officinalis —

(Pierre ex Pitard) Merr. & Chun

Neesia synandra Mast.

Neolitsea ?zeylanica

Neolitsea zeylanica Merr.

Neoscortechinia kingii (Hook. f.) P. & H.

Nephelium cuspidatum

B1. v. eriopetalum (Miq.) Leenh.

Norrisia maior Soler.

Norrisia malaccensis Gardn.

Nothaphoebe umbelliflora (B1.) B1.

Ochanostachys amentacea Mast.

Osmelia philippina (Turcz.) Benth.

Palaquium ?rostratum

Palaquium hexandrum (Griff.) Baill.

Palaquium microphyllum King & Gamble

Palaquium obovatum (Griff.) Engl.

Palaquium rostratum (Miq.) Burck

Palaquium sp.1

Palaquium xanthochymum (De Vr.) Pierre

Parartocarpus bracteatus (King) Becc.

Parinari oblongifolia Hook. f.

Parishia maingayi Hook. f.

Parkia speciosa Hassk.

Payena lucida (G. Don) DC.

Payena obscura Burck

Pellacalyx axillaris Korth.

Pellacalyx saccardianus Scott.

Pentace triptera Mast. .

Pertusadina eurhyncha (Miq.) Ridsdale

Phoebe grandis Merr.

Se oeqocqqooqoceqaqq =

ODOOoOrRNNFR NWN R KF LN

RB RRR RNR MN ONrF FR CO

ota oa & ©

mo GO

NNrFR CO CO

NRF WN

—

/ * .- = o-

we ee er ee eee

b ee

KFKNNWOANK ON KN

RS Ke nO MRK rR Or NY OW WO NR DNR NNW WR eR

Orowct ost Ho Orownrndorwrc o&

Appendix 2 (Continued)

SPECIES No. of Individuals

358. | Pimeleodendron griffithianum (M.A.) Benth.

359. | Pithecellobium jiringa (Jack) Prain

360. | Planchonella maingayi (Clarke) van Royen

361. | Ploiarium alternifolium (Vahl) Melchior

362. | Polyalthia ?hookeriana King

363. | Polyalthia glauca (Hassk.) Muell.

364. | Polyalthia jenkensii

(Hook. f. & Thoms.) Hook.f. & Thoms.

365. | Polyalthia macropoda King

366. | Polyalthia rumphii Merr.

367. | Polyalthia sumatrana (Miq.) Kurz

368. | Pometia pinnata Forst. f. alnifolia

369. | Popowia fusca King

370. | Popowia pisocarpa (B1.) Endl.

371. | Porterandia anisophylla (Jack ex Roxb.) Ridl.

372. | Pouteria malaccensis (Clarke) Baehni

373. | Prunus arborea (B1.) Kalkm.

374. | Purnus polystachya (Hook. f.) Kalkm.

375. | Pseudoeugenia singaporensis King

376. | Psydrax sp. 10 of Wong (1989)

377. | Psydrax sp. 11 of Wong (1989)

378. | Pternandra coerulescens Jack

379. | Pternandra echinata Jack

380. | Pyrenaria acuminata Planch. ex Choisy

381. | Rhodamnia cinerea Jack

382. | Sandoricum beccarianum Baill.

383. | Sandoricum koetjape (Burm. f.) Merr.

384. | Santiria ?griffithii

385. | Santiria apiculata Benn.

386. | Santiria griffithii (Hook. f.) Engl.

387. | Santiria laevigata B1.

388. | Santiria rubiginosa B1.

389. | Santiria tomentosa B1.

390. | Sapotaceae? 1

391. | Sarcotheca griffithii

(Planch. ex Hook. f.) Hall. f.

392. | Sarcotheca laxa Knuth v. sericea

(Ridl.) Veldk.

393. | Scaphium macropodum

(Miq.) Beumee ex Heyne

394. | Scleropyrum wallichianum (Wight & Arn.) Arn.

395. | Scorodocarpus borneensis Becc.

396. | Shorea bracteolata Dyer

= = © ©. ©

SS See tS o.S

Sowocr & Soorhre WN WN

b — _

hRreMNBRN WN NH

WBE RUNOWDRORPREPANUNBROUNPRPNE YH

SOonN WOR Orr OnNonN Wr aDAO worl

Appendix 2 (Continued)

SPECIES No. of Individuals

Shorea curtisii Dyer ex King

Shorea gibbosa Brandis

Shorea gratissima Dyer

Shorea leprosula Miq.

Shorea macroptera Dyer

Shorea ovalis B1.

Shorea parvifolia Dyer

Shorea pauciflora King

Shorea platycarpa Heim.

Sindora coriacea Maingay ex Prain

Spathodea campanulata P. Beauv.

Stemonurus scorpioides Becc.

Sterculia ?shillinglawii F.V. Muell.

Sterculia cordata B1.

Sterculia edelfetii F.v. Muell.

Sterculia macrophylla Vent.

Sterculia parviflora Roxb.

Sterculia rubiginosa Vent.

Streblus elongatus (Miq.) Corner

Strombosia ceylanica Gardn.

Strombosia javanica B1.

Styrax benzoin Dryand.

Swintonia schwenkii (T. & B.) T. & B.

Symplocos adenophylla Wall. ex G. Don

Symplocos fasciculata Zoll.

Symplocos rubiginosa Wall. ex DC.

Tarenna costata (Miq.) Merr.

Tarenna mollis (Wall. ex Hook. f.) B.L.

Robinson

Tarenna odorata (Roxb.) B.L. Robinson

Teijsmanniodendron ?holophyllum (Baker)

Kostermans

Teijsmanniodendron coriaceum (Clarke)

Kostermans

Terminalia subspathulata King

Ternstroemia penangiana Choisy

Timonius wallichianus (Korth.) Valeton

Triomma malaccensis Hook. f.

Tristaniopsis merguensis (Griff.) Wilson &

Waterhouse

Turpinia sphaerocarpa Hassk.

Vatica ?ridleyana

Vatica maingayi Dyer

—

Www non RN Ww

= . WW —

FE WHA OUNN WN RRR kK WH BN

ee ee ae

Appendix 2 (Continued)

SPECIES No. of Individuals

Vatica ridleyana Brandis 0 3 2 15

Vitex pinnata L. 1 6 1 8

Xanthophyllum ?affine 0 l 0 1

Xanthophyllum affine Korth. 0 2 4 16

Xanthophyllum amoenum Chodat 0 l 1 2

Xanthophyllum ellipticum Korth. l 2 3 6

Xanthophyllum eruhynchum Mia. 0 3 2 5

Xanthophyllum griffithii Hook. f. ex Benn 0 0 1 l

Xanthophyllum obscurum Benn. 0 0 2 2

Xanthophyllum stipitatum Benn. 0 0 3 3

Xanthophyllum vitellinum (B1.) Dietr. 0 8 6 14

Xerospermum noronhianum B1. 0 l 3 4

Xylopia caudata Hook. f. & Thoms. 0 8 2 10

Xylopia ferruginea (Hook. f. & Thoms.) 0 44 4 48

Hook. f. & Thoms.

Xylopia ferruginea v.oxyantha 0 14 0 14

(Hook. f. & Thoms.) Sinclair

Xylopia fusca Maingay ex Hook. f. & Thoms.

Xylopia magna Maingay ex Hook. f. & Thoms.

Xylopia malayana Hook. f. & Thoms.

—

eee Le ee ee ae a are

0 ee ee ee ee ee ee ne oe

)

ee ee ee

Appendix 2 (Continued)

SPECIES No. of Individuals

bok (peek

DOrRrPrRPROOR RFP RP RP OC OR RP RP rR rR OO

en a SO NO oe

rota Pec

Embryology and Seed Development in the Winged Bean,

Psophocarpus tetragonolobus

A. L. Lim! AND N. PRAKASH?

Department of Botany,

University of Malaya,

59100 Kuala Lumpur,

Malaysia.

2Department of Botany,

University of New England,

Armidale, N.S.W., 2351,

Australia.

Abstract

In Psophocarpus tetragonolobus (L.) DC the anther is tetrasporangiate. Its wall development

usually conforms to the Dicotyledonous type but occasionally to the Basic type. Simultaneous cytokinesis

in microsporocytes results in tetrahedral microspore tetrads. The mature pollen grains are triporate

and 2-celled with a reticulate exine. The mature ovule is campylotropous, bitegmic and crassinucellate.

The micropyle is zig-zag and formed by both the integuments. The embryo sac development follows

the monosporic Polygonum type. Fertilization is porogamous and triple fusion precedes syngamy. The

endosperm development is of the Nuclear type and free-nuclear endosperm haustoria develop in both

the micropylar, and chalazal parts. The first two divisions of the zygote are transverse producing a

linear 4-celled proembryo, but the subsequent divisions are in various planes. At the early globular

stage of the embryo, the suspensor cells become hypertrophied and haustorial. In mature seed, the

inner integument remains 2 or 3 layered, but the outer becomes 15-30 layers and is particularly

massive around the micropyle. The thick-walled palisade cells of the seed coat are derived from the

outer epidermis of the outer integument, while all the other layers remain parenchymatous.

Introduction

Psophocarpus tetragonolobus (L.) DC., the winged bean, is a

perennial climber of the tribe Phaseoleae of the legume subfamily

Papilionoideae. The geographical origin of the species is uncertain with

Africa (where all the other five species of the genus are indigenous), India,

South East Asia (particularly Indonesia, where there are numerous local

names for the plant), and Papua New Guinea having been suggested

variously as_ possible places of origin (Hymowitz and Boyd, 1977).

Hymowitz and Boyd (1977) themselves favoured a Papua New Guinea

origin and stated that “it would be difficult to find another high rainfall-

adapted tropical legume crop with as many desirable characteristics as

Psophocarpus tetragonolobus.” Virtually all parts of the plant are edible.

Besides the unripe fruits, which are popular as'a green vegetable, the

leaves, flowers, seeds and tuberous roots are consumed as well. The protein

and oil content (37% and 18% respectively) of the seeds compares

favourably with that of soybeans, and even the leaves, flowers, pods and

tubers have protein levels of 10-15% (Abbiw, 1990). The plants are also

considered as a valuable source of green fodder and manure. Detailed

embryological information, which might be useful in any attempts to

improve this crop of considerable potential, has so far been lacking.

Materials and Methods

Flowering and fruiting materials of suitable ages were collected

between 1990 and 1992 from glasshouse-grown plants at the Department

of Botany, University of New England. These were fixed in FPA (formalin-

propionic acid- 70% ethyl alcohol - 5:5:90 v/v) and stored after 48 hr in

70% ethyl alcohol. Following standard procedures of microtechnique (Sass,

1958), the specimens were embedded in Paraplast, sectioned, and stained

in safranin and fast green. Permanent slides were made after dehydration

in an ethyl alcohol series, treatment with Histoclear (a xylene substitute),

and mounting in Eukitt.

Observations

Anther:

The anther is tetrasporangiate (Fig. 1A). Hypodermal archesporial

cells in each sporangium divide periclinally to form the primary parietal

and primary sporogenous cells (Fig 1B). The latter divide repeatedly in

various directions and differentiate into microsporocytes. The primary

parietal cells divide periclinally to form the outer and inner secondary

parietal cells. Usually, only the outer secondary parietal cells continue

further divisions whereas the inner do not (Fig. 1D). The derivatives of the

outer secondary parietal cells constitute the endothecium and the two

middle layers while the inner secondary parietal cells directly develop into

the tapetum. Occasionally the inner secondary parietal cells also show

mitosis (Fig. 1C) suggesting that although the anther wall formation usually

conforms to the Dicotyledonous type (Davis, 1966), the Basic type may

also occur. Rarely, the cells of a middle layer may further divide resulting

in a third middle layer (Fig. 1E). The tapetum is secretory; its cells remain

uninucleate and develop one or two large vacuoles before meiosis occurs

in the microsporocytes (Fig.1F,G). The mature anther wall, lying under

the single-layered epidermis, consists of one layer each of endothecium

and tapetum and 2 or 3 middle layers lying in between.

At the time of meiosis, the microsporocytes become ensheathed in

callose. Simultaneous cytokinesis accompanies meiosis (Fig. 1H-J) and the

resultant microspore tetrads are tetrahedral (Fig. 1G, K). Different stages

of development, ranging from late prophase to the microspore tetrad phase,

can be observed within the same flower bud. In addition, different

microsporangia of the same anther may exhibit different stages of

development though usually all microsporocytes within a single sporangium

show synchrony. However, in one instance while the microsporocytes in the

lower part of the sporangium were at the late prophase stage, those in the

middle were at diakinesis, and in the upper part they were at metaphase I.

The microspore walls are formed by furrowing and the microspores

are initially isolated from each other by callose (Fig. 1J,K). Dissolution of

callose releases individual microspores into the sporangium by which time

a thin exine and incipient pores become evident (Fig. 1L). At this stage,

the tapetum is still intact though large vacuoles appear in the cells and the

cytoplasm appears sparse. By then all but one of the middle layers

disintegrate. The uninucleate microspore enlarges, the exine thickens, and

its nucleus migrates to the periphery of a large central vacuole, giving the

microspore the characteristic signet-ring appearance. Next the tapetum

also degenerates while the cells of the endothecium enlarge and become

radially elongated. The microspore undergoes mitosis to form the 2-celled

pollen grain. In the anther wall fibrous thickenings appear in the

endothecium and the single persistent middle layer still remains distinct

(Fig. 1M). By the time the microspores become mature pollen grains, the

tapetum and the remaining middle layer completely break down leaving

only the fibrous endothecium and the epidermis, the cells of which

accumulate brownish-staining crystals (Fig. 1N). The mature pollen grains

are triporate, 2-celled, and show a reticulate exine.

Ovule:

Mature ovules are campylotropous, bitegmic and crassinucellate. Ovular

primordia, 10-18 in number, are initiated as homogeneous hemispherical

protuberances from the marginal placenta (Fig. 2A). All the ovules in an ovary

develop synchronously. The primordia begin to undergo curvature even before

the integuments arise and finally assume the campylotropous form (Fig. 2A-C).

Fig. 1. Psophocarpus, anther wall and pollen development. All cross

sections of anther. A, tetrasporangiate anther with pollen tetrads; B-E,

stages in anther wall development; F, microspore mother cells and wall

layers; G, anther wall at pollen tetrad; H-K, microsporogenesis; L,M,

differentiation of the mature anther wall and pollen; N, anther wall and

mature 2-celled pollen. (ar, archesporial cell; en, endothecium; ep, epidermis;

g, generative nucleus; is, inner secondary parietal cell; mi, microsporocyte;

ml, middle layer; os, outer secondary parietal cell; sc, sporogenous cell; ¢

tapetum; v, vegetative nucleus).

Fig. 2. Psophocarpus, ovule and embryo sac development. All longitudinal

sections of ovule. A-D, stages in ovule development; £, pollen tube entering

ovule; F, initiation of the inner and outer integuments; G, asingle sporogenous

cell; H, twin sporogenous cells; /, megasporocyte; J, megasporogenesis; K-

M, megagametogenesis; N, mature female gametophyte; O, degeneration

of antipodals in the female gametophyte. (an, antipodal; eg, egg; fm,

functional megaspore; ii, inner integument; ob, obturator; oi, outer

integument; pn, polar nuclei; pt, pollen tube; sg, starch grains; sn, secondary

nucleus; sy, synergid).

Both integuments differentiate simultaneously from the nucellus at

the sporogenous cell stage (Fig. 2F). However, the development of the

inner integument is slower than that of the outer so that at the

megasporocyte stage the outer integument of 3 layers encloses the inner

integument of 2 or 3 layers and the nucellus (Fig. 2B). The outer integument

continues to grow rapidly and completely envelops the nucellus at the end

of megasporogenesis while the inner reaches only half its length. By the 2-

nucleate stage of the female gametophyte, the inner integument, which is

still 2 or 3 cells thick, reaches the micropyle while the cells of the outer

integument undergo periclinal and anticlinal divisions in the micropylar

part so that it becomes 8-10 cells thick at the tip (Fig. 2C). As the ovule

matures, the tip of the outer integument continues to show repeated

divisions to form a massive structure of 13-20 layers of cells, growing towards

the funiculus and covering the inner integument and the micropylar end of

the nucellus. As a consequence, the micropyle becomes zig-zag with the

exostome surrounded by the outer integument and the endostome bounded

on all sides by the inner integument of 2 or 3 layers of cells (Fig. 2D).

Starch grains accumulate in the cells of the outer and inner integuments in

the region surrounding the micropyle.

Usually one of the hypodermal cells differentiates into the archesporium

which divides periclinally to form a primary parietal cell and a sporogenous

cell (Fig. 2G). A single instance of twin sporogenous cells was observed

(Fig. 2H). As the cytoplasm becomes denser and the nucleus enlarges, the

sporogenous cell becomes the megasporocyte while the primary parietal

cell divides giving rise to 2 or 3 nucellar layers above it (Fig. 21). Cytokinesis

following the first meiotic division produces a dyad, of which only the

lower member undergoes meiosis II. Thus a linear triad is formed at the

end of meiosis (Fig. 2J). The smaller upper dyad cell degenerates

simultaneously with the middle megaspore, leaving only the chalazal

megaspore to continue development (Fig. 2K). After three successive

nuclear divisions a mature eight-nucleate female gametophyte is formed

(Fig. 2L-N); its development thus conforming to the monosporic Polygonum

type. As the female gametophyte matures, an egg flanked by two synergids

- each with a filiform apparatus - differentiates at the micropylar end, the

polar nuclei migrate toward the funicular side of the female gametophyte

and the antipodals are organised in the chalazal region (Fig. 2N). Soon

after formation, the two polar nuclei fuse into a large secondary sooner 1:

which becomes located just below the egg (Fig. 2E,O).

During its development the female gametophyte rapidly increases in size

at the expense of the nucellar tissue, especially at the sides and in the

Fig. 3. Psophocarpus, embryo development. All longitudinal sections of

seed. A, division of endosperm nucleus before syngamy; B,C, division of

zygote; D, linear 4-celled proembryo and nuclear endosperm showing

beginning of endosperm haustoria; E-M, stages in the development of

globular embryo, note the extensive haustorial suspensor; N, section of whole

seed showing globular embryo and endosperm haustoria; O, globular embryo;

P, early heart-shaped embryo. (c, cellular endosperm; ds, degenerating

synergid; eh, endosperm haustorium; em, embryo; /s, haustorial suspensor;

ma, male gamete; n, endosperm nuclei; sg, starch grains).

micropylar region. Hence the mature female gametophyte is bordered by

the inner integument on the sides, the single-layered nucellar epidermis at

the micropylar end, and the hypostase in the chalazal region. An

endothelium is not differentiated. Starch grains are first deposited in the

nucellar epidermis at the 4-nucleate female gametophyte stage. They

become abundant in not only the two integuments but also the mature

female gametophyte (Fig. 2N,O) and persist even after fertilisation.

Fertilisation:

Pollen tubes grow along the placenta towards the nucellus through the

micropyle (Fig. 2E, 4A). The epidermal cells of the placenta become

papillate, assume a glandular appearance, and possibly act as a a pollen

tube guide. The entry of the pollen tube is porogamous and through one of

the synergids; the other synergid degenerates soon after fertilization. One

male gamete fuses with the secondary nucleus to form the primary

endosperm nucleus whereas the second enters the egg but does not

immediately fuse with its nucleus. The two nuclei within the egg remain

separate while the primary endosperm nucleus divides to form free

endosperm nuclei (Fig. 3A). Syngamy occurs after the formation of eight

endosperm nuclei.

Endosperm:

The development of the endosperm is of the Nuclear type. Just before the

primary endosperm nucleus divides, the embryo sac enlarges significantly

and large starch grains accumulate, particularly in the micropylar region.

At least the first three mitotic divisions are synchronous. The free nuclei,

resulting from repeated divisions, mainly occupy the micropylar region

and along the periphery of the enlarging endosperm in the vicinity of the

funiculus (Fig. 3D). At the 16 nucleate stage of the endosperm, the zygote

divides (Fig. 3B). At the time of a linear proembryo there are about 60

endosperm nuclei. As the proembryo develops, the number of free nuclei

continues to increase rapidly.

A tubular coenocytic haustorium develops from the upper end of the

endosperm on the side of the funiculus. It grows towards the chalazal

nucellar tissue causing many of the cells to degenerate (Fig. 3D, 4B). The

narrow haustorium, with a sac-like tip, is discernible at the linear proembryo

stage and becomes very distinct by the early globular stage. As the embryo

develops, the haustorium grows in between the inner integument and the

nucellus (Fig. 3N). In addition, the chalazalmost free-nuclear part of the

endosperm also shows haustorial activity. Both haustoria remain free-

nuclear throughout the life of the endosperm.

The main body of the endosperm remains completely free-nuclear until

the early globular embryo stage. The number of nuclei counted at this

stage was 658 with 274 situated mainly in the micropylar quarter of the

endosperm and showing either mitotic metaphase or anaphase. The

remaining occur as free nuclei mostly along the periphery of the endosperm.

Cell-formation in the endosperm begins from the micropylar region and

proceeds towards the chalazal part at the late globular embryo stage (Fig.

4C). By now, because of the breakdown of the nucellus, the endosperm is

completely surrounded by the inner integument except at the chalazal

region where a butt of the nucellus persists until the cotyledons differentiate

in the embryo. The endosperm is almost totally consumed by the embryo

in the mature seed and the reserve foods are transferred to the cotyledons.

Embryo:

The zygote enlarges and divides after the formation of 16 nuclei in the

endosperm (Fig. 3B). The first division is transverse forming an apical cell

and a basal cell after which both further divide transversely to form a

linear 4-celled proembryo (Fig. 3C, E, F). The next division may be

transverse in one of the cells forming a linear 5-celled proembryo (Fig.

3G); or vertical and oblique in one or more cells (Fig. 3H-K). Subsequent

oblique divisions result in the early globular embryo while the derivatives

of the basal cell continue to divide in an irregular and variable manner to

produce the cells of the suspensor (Fig.3K, L). The suspensor cells become

hypertrophied and elongate significantly, extending from the embryo sac

into the erstwhile endostome and finally touching the outer integument

(Fig. 3M-O). By the heart-shaped stage of the embryo (Fig. 3P), the

endosperm is mostly used up. The massive suspensor eats into the

integuments and apparently assumes a major, if not the sole, responsibility

for the continued nutrition of the embryo.

Seed:

The fertilised female gametophyte undergoes gross enlargement at the

expense of cells in the micropylar part of the nucellus so that only 7 or 8

nucellar epidermal cells eventually persist. The cells of the inner integument

divide anticlinally and undergo radial elongation. Meanwhile, the endosperm

haustoria rapidly disorganise and destroy the chalazal nucellus which is gradually

resorbed, thus increasing the size of the endosperm. At the linear 4-celled stage

of the proembryo, the endosperm is bordered by the inner integument along half

its length (Fig. 3D); while by the late globular embryo stage the entire length

Fig. 4. Psophocarpus, longitudinal sections of seeds. A, division of zygote

with degenerating synergid and pollen tube at the micropyle; B, linear 4-

celled proembryo with nuclear endosperm forming haustorium; C, globular

embryo with haustorial suspensor and endosperm haustoria. (ds, degenerating

synergid; eh, endosperm haustorium; em, embryo; hs, haustorial suspensor;

pt, pollen tube; z, zygote. Scale: A,B = 40mm; C = 0.5mm).

except the chalaza is lined by the inner integument (Fig. 3N). Throughout

this development the inner integument continues to divide anticlinally to

expand its length but remains 2 or 3 cells thick. The outer integument, on

the other hand, becomes 15-30 layers thick through both anticlinal and

periclinal divisions and is particularly massive around the micropylar region

(Fig. 4C). Further, the outer epidermal cells of the outer integument divide

anticlinally, the resultant cells elongate, lose their nuclei and differentiate

into a palisade tissue of thick-walled columnar cells. The cells below the

epidermis remain parenchymatous (Fig. 4C) without showing the

differentiation of a layer of hour-glass cells as in certain other legumes.

Discussion

Dicotyledonous type of anther wall development was the only type recorded

in the family (Davis, 1966; Prakash, 1987) until the present report of the

occasional presence of the Basic type as well. Besides the secretory tapetum

of uninucleate cells, the anther wall occasionally shows three middle layers

which is one layer more than that reported in the other members of the

family (Prakash, 1987). The morphology of the ovule and its development

reaffirm the earlier observations on the family (Prakash, 1987; Prakash

and Chan, 1976; Prakash and Herr, 1979). The female gametophyte

development conforms to the monosporic Polygonum type, which seems

to be a common feature throughout the Leguminosae - except for members

of the Australian endemic papilionoid tribe Mirbelieae which exhibit a

variety of novel patterns (Cameron and Prakash, 1993). Also, the antipodals

are normal in P. tetragonolobus unlike certain Australian genera of the

tribes Indigofereae, Bossiaeeae and Mirbelieae which possess giant

antipodals (Cameron and Prakash, 1990). An instance of twin sporogenous

cells has been observed; and this too is known in the family (Rembert,

1969b; Prakash, 1987). Presumably only one sporogenous cell continues

development because twin female gametophytes, which have been recorded

in certain other members of the family, have never been found in the

present material. Often, during megasporogenesis, the second division of

the micropylar dyad is arrested and a triad is formed as in Cassia (Rembert,

1969a), and Vicia villosa (Rembert, 1969b). An integumentary endothelium,

which occurs in many other members of the family (Prakash, 1987), does

not seem to differentiate in any member of the tribe Phaseoleae. However,

an endothelium of nucellar origin has been reported in Phaseolus

aconitifolius (Deshpande and Bhasin, 1974).

The egg nucleus fuses with the male gamete only after the division of the

primary endosperm nucleus. This delayed syngamy has been described in

one other legume i.e., Acacia leucophloea (Ugemuge,1982), in which it is

achieved after the formation of four endosperm nuclei. As in all other

legumes studied so far, the endosperm is of the Nuclear type and it becomes

cellular only after several hundred free nuclei are formed. The presence of

a chalazal haustorium which remains free-nuclear even after cell-formation

had occurred in the micropylar half of the endosperm is known in several

legumes (Johri and Garg, 1959) including Atylosia, Glycine, Teramnus (Rau,

1953) Canavalia, Hardenbergia violacea, Kennedia rubicunda and Vandasia

retusa (Cameron, 1988) of the tribe Phaseoleae. However, in Psophocarpus

there is an additional tubular lateral haustorium which also remains

coenocytic. The zygote divides when there are 16 free endosperm nuclei as

has also been reported in Acacia leucophloea (Ugemuge,1982).

As in many other groups of the Leguminosae (Lersten, 1983), in Phaseoleae

also the morphology of the suspensor is variable; from being filamentous

in species of Erythrina (McNaughton, 1976) to massive in Phaseolus.

Cameron (1988) found a short suspensor in Glycine clandestina,

Hardenbergia violacea and Kennedia rubicunda but a long, filamentous,

uniseriate suspensor of inflated cells in Canavalia. However, unlike P.

tetragonolobus, in most species the suspensor is contained wholly within

the endosperm.

The pattern of embryogeny in Psophocarpus tetragonolobus is essentially

similar to that in Glycine (Ho, 1963; Souéges, 1949). The structure of the seed

coat is similar to that described for the family by Corner (1951, 1976) who

believed that the functioning of the exposed surface as a protective layer suggests

a primitive construction. However, unlike certain other members of the family

(Corner, 1976; Prakash and Chan, 1976), a layer of hour glass cells is not

distinguished in the mesophyll of the winged bean seed.

Acknowledgements

We are grateful to Professor Abdul Latiff Mohamed of the Department of

Botany, National University, Malaysia for the supply of seed and some

fixed materials on which this investigation is based. We appreciate the

help of Professor Goh Chong Jin, Head of Botany, National University of

Singapore, for providing facilities to Prakash so that the manuscript could

be finalised. We also wish to record our thanks to the University of

Malaya, Kuala Lumpur, for granting Lim study leave at the University of

New England where the experimental work for this project was completed.

References

Abbiw, D. (1990). Useful plants of Ghana. Intermediate Technol. Publ.,

London, & Royal Bot. Gard., Kew, Surrey.

Cameron, B.G. (1988). Embryology and Floral Morphology of Australian

Papilionoideae. Unpubl. Ph.D. thesis, Univ. New England, Armidale.

— and Prakash, N. (1990). Occurrence of giant antipodals in the female

gametophytes of Australian Bossiaeeae, Indigofereae and Mirbelieae

(Leguminosae). Australian J. Bot. 38, 395-401.

— and —— (1993). Ovule characters of systematic value in papilionoid

legumes. Proc. Symp. 3.8.1., 15th Int. Bot. Cong., Yokohama. Abstracts,

p.96.

Corner, E.J.H. (1951). The leguminous seed. Phytomorphology 1, 117-150.

—— (1976). The Seeds of Dicotyledons. Vols 1 & 2. Cambridge Univ. Press,

Cambridge.

Davis, G.L. (1966). Systematic Embryology of the Angiosperms. John Wiley,

New York.

Deshpande, P.K. and Bhasin, R.K. (1974). Embryological studies in Phaseolus

aconitifolius Jacq. Bot. Gaz. 135, 104-113.

Ho, M.Y. (1963). (Embryological studies of soybeans 1. the development of the

embryo and albumen.) Acta Bot. Sinica 11, 318-328. (In Chinese).

Hymowitz, T. and Boyd, J. (1977). Origin, ethnobotany and agricultural potential

of the winged bean - Psophocarpus tetragonolobus. Econ. Bot. 31, 180-

188.

Johri, B.M. and Garg, S. (1959). Development of endosperm haustoria in some

Leguminosae. Phytomorphology 9, 34-46.

Lersten, N.R. (1983). Suspensors in Leguminosae. Bot. Rev. 49, 233-257.

McNaughton, J.E. (1976). Embryology of Erythrina caffra Thunb. Sporogenesis

and gametogenesis. J. South Afr. Bot. 42, 395-400.

Prakash, N. (1987). Embryology of the Leguminosae. /n C.H. Stirton (ed).

Advances in Legume Systematics. Part 3., Royal Botanic Gardens, Kew. pp

242-278.

— and Chan, Y.Y. (1976). Embryology of Glycine max. Phytomorphology

26, 302-309.

— and Herr, J.M. Jr. (1979). An embryological study of Glottidium

vesicarium through the use of clearing technique. Phytomorphology

29,71-77.

Rau, M.A. (1953). Some observations on the endosperm in Papilionaceae.

Phytomorphology 3, 209-222.

Rembert, D. (1969a). Comparative megasporogenesis in Caesalpiniaceae.

Bot. Gaz. 130, 47-52.

—— (1969b). Comparative megasporogenesis in Papilionaceae. Amer. J.

Bot. 56, 584-591.

Sass, J.E. (1958). Botanical Microtechnique. lowa State Univ. Press, Ames.

Souéges, R. (1949). Embryogénie des Papilionacées. Développment de

l’embryon chez le Glycine soja Sieb. et Zucc. (Soya hispida Moench.).

C. R. Acad. Sci. Paris 229, 1183-1185.

Ugemuge, (1982). Embryological studies in Acacia leucophloea Willd. J.

Indian Bot. Soc. 61, 278-285.

- Carallia brachiata cv. Honiara,

a Beautiful Fastigiate Ornamental Tree

WoncG YEW Kwan! and T.C. WHITMORE?

189 Soo Chow Garden Rd., Singapore 575526

2Geography Department, Cambridge University,

Cambridge CB2 3EN, England

Carallia is an inland genus of the mainly mangrove family Rhizophoraceae.

The genus extends from Madagascar to the Solomon Islands and has c. 10 species

(Hou, 1958), eight of them in Malesia. Carallia brachiata occurs throughout the

Asian range of the genus. It is a common, but never abundant, rather nondescript

tree of lowland tropical rain forest, reaching 50 m tall.

In a few villages of the Solomon Islands and in the capital Honiara

there occurs an exceptionally beautiful fastigiate form of C. brachiata, with

weeping limbs and the crown shape of the Lombardy Poplar (Populus

nigra cv. Italica) of Europe, see Fig. 1. This form reaches 18 m tall and 25

cm bole diameter but is usually smaller. Numerous fruits are produced at

frequent intervals and germinate easily.

The trees planted in Coronation Gardens, Honiara, came from Auki District,

Malaita, where the form is reputed to occur wild. A tree seen at Lodomae village,

north Kolombangara, is reputed to be grown from Choiseul seeds.

Several attempts to export this fine tree by G.F.C. Dennis, doyen of

horticulture in the Solomons, have all failed. But of 25 seeds from a tree in

his garden in Honiara, collected on 2 February 1986 by one us (TCW) and

planted in Singapore on 12 February 1986 by WYK all but one germinated

between 7 March and 28 March 1986. From this batch of seedlings six trees

have now been planted out along Cheang Hong Lim St. and twelve trees

along Cross St. in the Central Business District of Singapore, on very poor

compacted soil. These were 8 m tall in July 1994, (Fig. 2) and they bore

copious young fruit. Three previous attempts to germinate seeds from

separate fruitings failed, but recently the Singapore Botamic Gardens

managed to collect seeds and germinate them successfully. The seedlings

are now being nurtured for planting out at a later date.

f "gue Mme |:

4 IW an hag a ota ,

Co el

Fig. 1. Carallia brachiata cy. Honiara in Coronation Gardens, Honiara,

Solomon Islands, 1964, with John Sore.

——

Herbarium specimens of this form are identical to typical C. brachiata and

in his Flora Malesiana revision Hou (1958) gave it no taxonomic recognition.

We believe this elegant fastigiate tree has high potential as a wayside tree,

and that now a small population exists in Singapore, the form should be

grown as a superior replacement or supplement to the other introduced

fastigiate tree Polyalthia longifolia cv. Pendula.

To give identity to this form we propose the cultivar name Honiara.

Carallia brachiata (Lour.) Merr. cultivar Honiara

Differs from the wild tree in its tall, narrow columnar habit with pendulous

branches. Representative herbarium collections (all seen at SIN): Solomon

Islands: Honiara, Coronation Gardens: BSIP 503, 3 Aug. 1962, Forester

collector; BSIP 7870, 23 Feb 1967, G.F.C. Dennis. Kolombangara, Lodomae

village: BSIP 4413, 26 July 1964, T.C. Whitmore, ‘dautoli’.

The seedlings of typical C. brachiata are described by Burger (1972). Our

observations concur except that the cultivar has entire, not serrate

cotyledons (Fig.3). Ng (1991-2) illustrated seed (Fig. 466B) and seedling

(Fig. 768) of the typical form. These are also similar to the cultivar but Ng

noted germination occurred between 52 and 106 days, much slower than

we observed. Our cultivar very soon develops weeping branches (Fig.4),

and grows rapidly in height (4 mo 15 cm, 5 mo 25 cm, 7 mo 40 cm, 10 mo

60 cm).

A second batch of the 1986 seed lot, sent to the Forest Research Institute

of Malaysia, failed to germinate. It seems likely from this and the earlier

failures that the seeds soon lose viability, perhaps due to desiccation, i.e. they

are recalcitrant sensu Roberts (1973).

Acknowledgements

We thank Mr G.F.C. Dennis for the seeds and Mdm. Ohn Set for caring for

the seedlings and then arranging their planting out. The journey to re-

measure permanent sample forest plots in the Solomons which made this

enterprise possible was funded by the National Geographic Society of

Washington DC, USA.

96 :

eed joes

Fig. 2. Carallia brachiata cy. Honiara as astreet tree in Singapore, July 1994.

References

Hou, D. (1958). Rhizophoraceae. Flora Malesiana series I, 4 429-93.

Burger, D. Hzn. (1972). Seedlings of some tropical trees and shrubs mainly

of southeast Asia. PUDOC, Wageningen.

Ng, F S P (1991-2). Manual of forest fruits, seeds and seedlings. 2 vols.

Malayan Forest Record 34.

Roberts, E.H. (1973). Predicting the storage life of seeds. Seed Science &

Technology 1, 499-574.

Fig. 3. Young seedlings showing cotyledons with entire margins.

Seedling at 10 months age, 65 cm tall, scale marked every 30 cm.

ig. 4.

The Angiosperm Flora of Singapore Part 2

PHILYDRACEAE

R.M.K. SAUNDERS

Department of Ecology & Biodiversity,

The University of Hong Kong

Pokfulam Road ,

Hong Kong

Philydrum Banks & Sol. ex Gaertn

Fruct. sem. pl. 1 (1788) 62, t. 16; Ridl., Fl. Malay Penins. 4 (1924) 347; Skottsb., Bull. Jard. bot.

Etat Brux. ser. 3, 13: (1933) 111; Skottsb., Fl. Males. ser. 1, 4:1(1948) 5.

Erect, perennial, caespitose herbs with a short rhizome. Leaves

densely rosulate, equitant, 2-ranked; linear, fleshy, parallel-veined, sheathing

at base. Inflorescence a simple or paniculate terminal spike; scape 1 m or

longer, with few cauline leaves gradually replaced by alternate bracts.

Flowers bisexual; zygomorphic; sessile, solitary in axil of spathaceous bracts;

bracts enclosing flower buds, reflexed at anthesis, later embracing the fruit:

perianth corolline, 4-segmented, 2-seriate, persistent as fruit cover, yellow,

2 outer tepals larger, adaxial and abaxial respectively, 2 inner tepals smaller,

lateral; stamen single, filament flattened, adnate with base of inner and

adaxial tepals, anther dorsifixed, 2-loculate, spirally twisted, extrorse,

opening lengthwise by slits, pollen grains in tetrads, staminodes cuneate,

acute, shorter than fertile stamen; ovary single, superior, 3-loculate, with

parietal placentation, ovules many per locule, anatropous; style simple.

Fruit a persistent triangular-oblong loculicidal capsule with 3 valves. Seeds

with corona and spirally-striate testa, many per locule; embryo straight.

Distribution - Monotypic genus, occurring in South Japan, Taiwan,

South-East China, Indo-China, Malay Peninsula, Guam, South New Guinea

and North, East and South-East Australia (Hamann, 1966a). P. lJanuginosum

is reported to be extinct in Singapore (Keng, 1987) but was previously

collected in Bedok.

Ecology - P. lanuginosum occurs in freshwater ponds, marshes and

ricefields at low altitudes in its natural range (Skottsberg, 1948).

Uses - P. lanuginosum has no known economic importance.

100

0.25 mm

2cm 2mm 1m

Fig. 1. Philydrum lanuginosum Banks & Sol. ex Gaertn. a. Habit. b. Flower

with one lateral tepal removed. c. Flower with carpel and adaxial tepal

removed. d. Stamen, with spirally twisted anthers. e. Carpel. f. Ovary

(transverse section), with parietal placentation. g. Seed, with spirally striate

testa. [a. H.N. Ridley 5907 (SING); b.-g. drawn from fresh material. Scale

bars: a =2cm,b-e =2 mm; f= 1mm; g=0.25 mm.] Del. R.J. Nicholls.

101

Notes - The most comprehensive account of the Philydraceae is by

Hamann (1966a), who studied various aspects of taxonomic relevance,

including morphology, anatomy and embryology. The embryology has

been the subject of considerable research (Hamann, 1963, 1966a, 1966b:

Kapil and Walia, 1965), and has assisted with the clarification of the

phylogenetic relationships of the family. Similarities are apparent with the

Pontederiaceae and Haemodoraceae, although a relationship with the

Burmanniaceae has also recently been suggested.

1. P.lanuginosum Banks & Sol. ex Gaertn

Fruct. sem. pl. 1 (1788) 62,t. 16; Ridl., J. Straits Br. R. Asiat. Soc. 33 (1900) 169; Ridl., Mat. FI.

Malay Penins. 2 (1907) 110; Ridl., Fl. Malay Penins. 4 (1924) 347; Skottsb., Bull Jard. bot. Etat Brux.

ser. 3,13 (1933) 111; Skottsb., Fl. Males. ser. 1, 4:1(1948) 5; M.R. Hend., Malayan Wild Flowers,

Monocotyledons (1954) 192; H. Keng, Gdns’ Bull., Singapore 39 (1987) 123; I.M. Turner, K.S. Chua &

H.T.W. Tan, J. Singapore Nat. Acad. Sci. 18 & 19 (1990) 79; ILM. Turner, Gdns’ Bull., Singapore 45

(1993) 188.

Leaves 40-80 cm long (including basal sheath), glabrous, thick,

aerenchymatous; sheath 14-30 by 1-1.5 cm. Jnflorescence 20-60 cm long;

scape slender, terete, glabrate below, villous towards the flowers; bracts

ovate to subulate, 2-10 by 0.75-1 cm, woolly on abaxial side, short-acuminate

to attenuate. Flowers with thin perianth, outer tepals 10-15 by c. 10 mm,

many-veined, long villous outside, the upper with 2 stronger veins and

bidentate, acute, margins inflexed, inner tepals spathulate, c. 8 by 2 mm,

membranous, 3-veined, with base hairy outside; stamen c. 9 mm long,

glabrous, anther + spherical; ovary densely long-woolly; style glabrous;

stigma broad-triangular, long-papillose. Fruit 9-12 by 4-5 mm. Seeds dark

reddish-brown, bulb-shaped, c. 1 mm long; n = 8 (Hamann, 1966a) and 2n

= 16 (Briggs, 1966).

Acknowledgements

I would like to thank the Director, Botanic Gardens, Singapore for

the herbarium specimen borrowed.

References

Briggs, B.G. (1966). Chromosome numbers of some Australian

monocotyledons. Contr. N.S.W. natn. Herb. 4, 24-34.

Hamann, U. (1963). Die Embryologie von Philydrum lanuginosum

(Monocotyledonae - Philydraceae). Ber. dt. bot. Ges. 76, 203-208.

102

Hamann, U. (1966a). Embryologische, morphologisch-anatomische und

systematische Untersuchungen an Philydraceen. Willdenowia 4, 1-

178.

— (1966b). Nochmals zur Embryologie von Philydrum lanuginosum .

Beitr. Biol. Pfl. 42, 151-159

Kapil, R.N.& Walia, K. (1965). The embryology of Philydrum lanuginosum

Banks ex Gaertn. and the systematic position of the Philydraceae.

Beitr. Biol. Pfl. 41, 381-404.

Skottsberg, C. (1948). Philydraceae. Fl. Males. ser. 1,4:1, 5-7.

Keng, H. (1987). Annotated list of seed plants in Singapore (XI). Gdns’

_ Bull., Singapore 40, 113-132.

The Angiosperm Flora of Singapore Part 3

PLANTAGINACEAE

K.S. Cuua, H.T.W. TAN and I.M. TuRNER

Department of Botany,

The National University of Singapore

Singapore 119260,

Plantago L.

Sp. pl. (1753) 112; Gen. pl. ed. 5 (1754) 52; Ridl., J. Straits Br. R. Asiat. Soc. 33 (1900) 125:

Ridl., Fl. Malay Penins. 2 (1923) 225; M.R. Hend., Malayan Wild Flowers, Dicotyledons (1959) 268; H.

Keng, Gdns’ Bull., Singapore 36 (1983) 111; H. Keng, Concise Flora of Singapore (1990) 140; IM.

Turner, K.S. Chua & H.T.W. Tan., Journal of the Singapore National Academy of Science 18 & 19

(1990) 80; I.M. Turner, Gdns’ Bull., Singapore 45 (1993) 189.

Annual or perennial herbs, rarely small shrubs. Leaves simple; usually

spirally arranged; lamina venation parallel, margin entire or toothed; petiole

forming a sheath at the base; exstipulate. /nflorescence spicate or capitate;

axillary; pedunculate; bracts persistent. Flowers 4-merous; unisexual or

bisexual; regular; calyx lobed or deeply cleft; corolla scarious, lobes

imbricate in bud; stamens as many as and alternating with corolla lobes,

filaments long, anthers 2-loculate, exserted, versatile, opening by

longitudinal slits; ovary single, superior, usually 2-loculate, placentation

axile, ovules, with an integument several cells thick, 1 to many per locule;

style 1, bifid. Fruit a circumscissile capsule, dehiscing transversally with the

top segment falling off as a lid. Seeds, with a translucent testa, | or more

per locule; endospermous.

Distribution - The genus Plantago, with about 250 species (Mabberley,

1990), is naturally distributed in Europe and temperate Asia. According to

Holm et al. (1977), the widespread introduction of P. major and P.

lanceolata, to various parts of the world has resulted in the almost

cosmopolitan distribution of the genus; both species are found in all

continents, except the Antarctic and Arctic. Man has played a prominent

role in the widespread distribution of this genus. Only Plantago major

occurs in Singapore.

Ecology - Mostly weeds of disturbed areas such as agricultural land,

wasteland, cultivated ground, roadsides, and open fields. Allard (1965)

regarded P. lanceolata as one of the 12 most successful naturally growing

104

Fig. 1: Plantago major L. a. Habit. b. A segment of the inflorescence showing

a flower and a flowering bud. c. Longitudinal section of flower. d.

‘Exploded’view of the cie=rcumscissile capsule. [K. S. Chua 413]

105

colonizing plants. P. major and P. lanceolata are among the world ’s most

noxious weeds (Holm et al., 1977).

Uses - When soaked in water, the seeds of some species produce

copious mucilage which are medicinal (Burkill, 1966). The testa (‘husk’) of

several species, notably P. ovata, P. psyllium and P. indica, are prepared

by the pharmaceutical industry and widely used as a bulk laxative (Leung,

1980) and remedy for chronic diarrhoea. The husk mucilage is also used

as a thickener in some food products.

1. Plantago major L.,

Sp. pl. (1753) 112; Ridl., J. Straits Br. R. Asiat. Soc. 33 (1900) 125; Ridl., Fl. Malay Penins. 2 (1923)

225-226; M.R. Hend., Malayan Wild Flowers, Dicotyledons (1959) 268; H. Keng, Gdns’ Bull., Singapore 36

(1983) 111; H. Keng, Concise Flora of Singapore (1990) 140; I.M. Turner, K.S. Chua & H.T.W. Tan, J.

Singapore Nat. Acad. Sci. 18 & 19 (1990) 80; I.M. Turner, Gdns’ Bull., Singapore 45 (1993) 189.

P. asiatica L.

Perennial herb. Leaves simple; in a radical rosette; lamina broadly ovate to

narrowly lanceolate, 3-22 by 1-22 cm, with 5-9 longitudinal veins, glabrous or

pubescent, apex rounded, obtuse or acute, margin entire, shallowly or deeply

toothed, base abruptly narrowed; petiole 1-25 cm long. /nflorescence an upright

spike, 10-30 cm long; peduncle 4-50 cm long, terete or shallowly ribbed, glabrous

or pubescent; bracts 1-3.25mm long. Flowers greenish-white, bisexual; calyx lobes

oval, oblong, obtuse or acute, 1.25-2 mm long, margin scarious; corolla lobes

spreading, 1-1.5 mm long; style 4-6 mm long. Fruit ovoid, 3-4 mm long. Seeds

brown or black, oblong, 1 by 0.75 mm, rugose, 4-21 per locule.

Distribution - Singapore: widely distributed and locally common; Nee

Soon Swamp Forest, Kent Ridge, Dover Crescent, Punggol, Tanglin, etc.

Ecology - P. major can be found as a naturalized weed in disturbed

areas such as wasteland, public car parks, gardens and lawns. The plant is

remarkable in its ability to adapt to a wide range of growing conditions. In

Singapore, P. major can be found growing in periodically-mown lawns, in

waterlogged areas beside drains, and in cracks in concrete pavements. In

Europe and Morocco, P. major has been found growing in regions of 2,000

metres a.s.l. Its most northerly location is Spitsbergen, a Norwegian island

at latitude 77 °N (Sagar and Harper, 1964). The wide distribution of the

plant has led Holm et al. (1977) to suggest that its “habitat is probably not

restricted by climate.” Being of temperate origin, the plant’s adaptability

and tolerance to adverse growing conditions are severely tested in the low

altitude and equatorial climate of Singapore. Holm et al. (1977) mentioned

106

that P. major is rarely a troublesome weed in the equator as the tropical

heat and competition from the taller and more vigorous-growing plants

help to keep P. major under control. The mature plants of P. major flower

and fruit frequently throughout the year. Ridley (1930) noted that the

seeds are wind-dispersed.

Uses - According to Burkill (1966), P. major was originally introduced

from China for use as a medicine for diarrhoea and dysentery and for the

seeds in making jelly, while the leaves are occasionally eaten as a vegetable.

He also added that the Malays use the plant in the treatment of cough,

dysentry and gonorrhea.

Notes - P. major is extremely variable in form. Backer (1965) stated

that “they might be taken for different species, were it not that they are

connected by series of intergrades.” For a key to the numerous varieties

and taxa of Plantago major, see Pilger (1937).

Acknowledgements

This project was supported by The National University of Singapore

Research Grants RP 880301 and RP 930325.

References

Allard, R.W. (1965). Genetic systems associated with colonizing ability

in predominantly self-pollinated species. In: (Baker, H.G. and Stebbins, G.

L. eds.) The genetics of colonizing species. Academic Press; New York;

(49-75.)

Backer, C.A. and Bakhuizen van den Brink, R.C. Jr. (1965). Flora of Java

Vol. 2, Noordhoff; Groningen; 445-446.

Burkill, I.H. (1966). A dictionary of the economic products of the Malay

Peninsula Vol. 2, Ministry of Agriculture and Co-operatives; Kuala

Lumpur; 1797-1798.

Holm, L.G., Plucknett, D.L., Pancho, J.V. and Herberger, J.P. (1977). The

world’s worst weeds: distribution and biology, The University Press of

Hawai; Honolulu; 385-393.

107

Leung, A.Y. (1980). Encyclopedia of common natural ingredients used in

food, drugs, and cosmetics. John Wiley & Sons, New York; 272-273.

Mabberley, D.J. (1990). The plant book: a portable dictionary of the higher

plants. Cambridge University Press; Cambridge; 460.

Pilger, R. (1937). Plantaginaceae. In Engler, A., Das Pflanzenreich, IV, 269,

102, 1-466.

Ridley, H.N. (1930). The dispersal of plants throughout the world. L. Reeve

& Co.; Ashford, Kent; 28-31.

Sagar, G.R. and Harper, J.L. (1964). Biological flora of the British Isles:

Plantago major L., Plantago media L., and Plantago lanceolata L. J.

Ecol. 52: 189-221.

suid nee

as im

ics Ma

A Botanical Survey of Pulau Jong, Singapore

H.T.W. Tan, I.M. TuRNER and K.S. CHUA

Department of Botany

The National University of Singapore

Singapore 119260

Abstract

A botanical survey of Pulau Jong of the Republic of Singapore, a 0.6 ha island off the south

coast of Singapore Island, recently found at least 38 native vascular plant species which are listed here.

Previous botanical records for the island are also collated to bring the recorded number of species now

to 52. The contemporary flora is dominated by beach and secondary forest species. The slight change

in the species composition and decline in number compared to observations made by Holttum (1925)

are typical of the random fluctuations seen in small island floras.

Introduction

Pulau Jong is an island within the Republic of Singapore. Its vascular

plant flora was described by Holttum (1925) in some detail. More recent

records include collections made by J. Sinclair in 1950 and Johnson (1977)

who probably quoted Holttum (1925). It was chosen for a botanical survey

because it is relatively untouched largely owing to its reputation of being

haunted (Brooke 1925), its very steep terrain, small area and lack of water,

all of which would make it extremely difficult for settlement. It would thus

be interesting to examine the flora in detail after a lapse of about 70 years.

This survey is also part of the on-going Angiosperm Flora of Singapore

Project

Site

Pulau Jong (N1° 12' 56.2", E 103° 47' 18.2") is a small, dome-shaped

island (Fig. 1) of about 0.6 ha (Resource Centre, Ministry of Information

and the Arts 1994) lying off the south coast of Singapore Island, surrounded

by Pulau Bukum to the north-west, Pulau Sakeng to the south-west and

Pulau Sebarok to the south-east (Fig. 5). Jong, is Malay for junk and is

probably an allusion to the silhouette of the island being similar to that

type of vessel. The island is generally undisturbed and activities of man are

hardly evident. Brooke (1925) mentioned that half a dozen tombstones

were found near the water’s edge when he visited the island and a rusty,

still-standing flagpole was found by us at the summit of the island which is

23.4 m in altitude.

Geologically, the island is the type locality of the Jong Facies and

110

“contains alternating beds of roundstone conglomerate and sandstone, and

less frequently, beds of mudstone” (Anonymous, 1976).

Although there are no records of the climate for the Island, it would

be very similar to that for mainland Singapore which has an equatorial

climate with uniformly high temperatures, humidity and rainfall year-round

(Chia and Foong, 1991).

Methods

All the specimens cited in Table 1 as “Collected Recently” were

obtained by a team of five researchers who made a thorough survey of the

island on 13 Nov 1992. These collections were made into herbarium sheet

specimens which were identified largely by comparison with named

specimens in the Herbarium, Singapore Botanic Gardens (SING).

Nomenclature follows those of Turner, Chua and Tan (1990) and Turner

(1993). Voucher specimens of all the species reported were deposited in

the Herbarium, Department of Botany, The National University of

Singapore (SINU) and replicates when available, distributed to other

herbaria, mostly to SING. A survey of specimens from previous collections

found in SING and SINU was also made to compile an historical list of

species found on Pulau Jong.

Results and Discussion

The species that were recorded in the past and more recently are

given in Table 1. Found recently were a total of 38 vascular plants

comprising nine ferns, two gymnosperms and 27 angiosperms. All can be

considered native species.

From herbarium sheet labels, Holttum had made some collections of

the species he mentioned in his 1925 paper. Collection dates were 9 Jun, 11

Jun and 13 Jul 1924. Sinclair also made collections on 22 Sep 1950. From

the survey of past collections and literature, the total number of vascular

plants including those recently collected is 52 with 13 ferns, two

gymnosperms and 37 angiosperms.

The Pulau Jong flora is approximately 1.5% of the total flora for the

Republic of Singapore in about 0.0009% of the area. The species list is

made up largely of beach and secondary forest elements with some

mangrove species.

Most of the species are common with some exceptions considered

endangered, vulnerable or rare in the Singapore context according to Turner

et al. (1994). A few trees of Podocarpus polystachyus at the highwater

mark were

111

observed. This is an endangered species with only a few other

individuals known in Singapore from Kampong Mandai Kechil and Sentosa

(Tan, Turner and Chua, 1994; Turner et al., 1994). One treelet of Pongamia

pinnata which has a conservation status of vulnerable (Turner et al., 1994)

was seen. Another taxon currently found on the island and categorized as

vulnerable is Preridium caudatum ssp. yarrabense (Wee, 1994). Schefflera

elliptica, a rare species, which Keng (1990) noted to be probably extinct,

has now been rediscovered here as well as in Pulau Ubin (Turner et al.,

1992). Gnetum microcarpum, a rare species in Singapore (Turner et al.,

1994), said to be of the rain forest (Markgraf, 1951), was growing well in a

few locations on the island climbing on trees almost to the sea edge and

bearing many strobili in seed. There were many fully blooming individuals

of the rare Tarenna fragrans (Turner et al., 1994). This species is probably

absent on the mainland but also seen in Pulau Ubin (Turner et al., 1992).

Other recently collected species considered to be rare according to Turner

et al. (1994) include Ficus globosa, Memecylon edule and Xylocarpus

granatum. In view of the significant number of species classified endangered,

vulnerable or rare, the island is worth conserving as it is one of the few

localities for these species.

The flora has changed from that which Holttum (1925) described. Of

the sea shore plants he mentioned, Desmodium umbellatum, Heritiera

littoralis, Hibiscus tiliaceus, Intsia bijuga (called Afzelia retusa by Holttum),

Premna corymbosa (Premna_ integrifolia), Pterocarpus indicus and

Sonneratia griffithii are now extinct. Found too was a seedling of the

mangrove-dwelling Xylocarpus granatum which was not previously recorded

by Holttum (1925).

The slopes of the island have changed little in species composition

(Figs. 2 and 3). The extant species are very similar to those mentioned by

Holttum (1925) except for the extinction of Commersonia bartramia

(Commersonia platyphylla). Eugenia spicata and Eugenia grandis are much

more plentiful than described by Holttum (1925). At the more rocky areas,

patches of Davallia species (Fig. 3) and Dicranopteris linearis (Fig. 2) occur

between the beach forest areas. Davallia solida was collected by Holttum

(1925) but we collected only Davallia denticulata. As both species occur in

this type of habitat and are very similar, it is quite possible that in both our

cases, one could easily overlook the presence of the other.

The vegetation of the summit of the island (Fig. 4) has changed

considerably. Holttum (1925) described it as being quite open, mostly covered

by bracken fern (probably Preridium caudatum ssp. yarrabense) and the grass,

Eriachne pallescens with stunted bushes of Melastoma malabathricum (Melastoma

112

Fig. 1. View of Pulau Jong from the east-south-east. Most of the island is

covered by beach forest.

Fig. 2. Patch of Dicranopteris linearis (c) growing to the edge of the sea.

Fig. 3. Pandanus odoratissimus plants in the foreground with a patch of

Davallia species (v) behind.

Fig. 4. Beach forest at the summit of Pulau Jong. The tallest trees are

about 8 m in height. I.M. Turner is seen about centre.

ae aod

113

polyanthum). Currently, the summit has mostly a dense growth of trees

and is similar in species composition to the slopes with small patches of

Dicranopteris linearis in the few open areas. Most of the trees are Eugenia

spicata with lesser numbers of Eugenia grandis and Myrica esculenta. This

cover of trees has occurred contrary to the predictions of Holttum (1925)

who felt that the high exposure and drainage of the summit area would not

be conducive to growth of taller vegetation. The tallest trees at the summit

and slopes are about 8 m in height.

The extant species of epiphytes and lithophytes are similar to those

found by Holttum (1925). The pigeon orchid plants (Dendrobium

crumenatum) however, appeared to be on the verge of extinction with a

few tiny, half-dried individuals on the rocks almost at the sea edge, growing

with Pyrrosia lanceolata.

The present vegetation of the island consists of patches of beach

forest with patches of Davallia species (Fig. 3) or Dicranopteris linearis

(Fig. 2) in the open areas with poorer or no soil. Presumably, in time, with

the build up of soil, the whole island will become covered with beach

forest.

The ten newly recorded species may have either been overlooked by

Holttum (1925) who had stated that the 41 vascular plant species he noted “‘is

probably by no means all that are present”, or, they may have since arrived by

various means.

The two newly recorded ferns, young plants of Acrostichum speciosum

and Vittaria ensiformis have wind-dispersed spores. Ridley (1930) has noted that

Xylocarpus granatum (called Carapa moluccensis by Ridley) has corky, buoyant

seeds which are sea-dispersed. Ridley (1930) quoting H.B. Guppy has noted that

the legumes of Pongamia pinnata (Pongamia glabra) are buoyant and can float

for months in the sea. He also mentioned that Derris trifoliata (Derris uliginosa)

has similar fruits and dispersal mode.

The other species are probably bird-dispersed as their fruits are of the

colours Ridley (1930) listed, that attract birds. Ridley (1930) noted that seeds of

Gnetum species are mostly dispersed by birds but may be dispersed by water

and have been seen in sea-drift. Gnetum microcarpum has pink, ripe seeds

which are probably bird-dispersed. Arthrophyllum diversifolium (Arthrophyllum

ovalifolium) is probably bird-dispersed (Ridley, 1930). Breynia reclinata has

slightly fleshy, red capsules and red is the most attractive colour to birds (Ridley,

1930). Memecylon edule has fleshy, black berries when ripe and black is the

third most attractive colour to birds (Ridley, 1930). Lastly, Ficus globosa which

114

Singapore

Island

Fig. 5. Maps of Singapore and Pulau Jong. In the bottom map, the contour

lines are at 5 m intervals. This map is based on the 1:2,500 topographic

map published by the Chief Surveyor, Singapore, 1970 and by courtesy of

the Chief Surveyor, Singapore.

115

has a 1.5 cm diameter fruit may be bird or bat-dispersed like many other

members of its genus.

The 16 species which have become extinct, may have become so for

any of the factors including a lack of adaptation to the island habitat, the

‘founder principle’, greater susceptibility to random non-adaptive changes

in genomes because of the small population size, biological interactions

between species, currently present such as competition or co-evolutionary

effects (Cox and Moore, 1985). The slight decline in number of species,

extinction of some and the immigration of others are typical of the random

fluctuations of small island floras.

Holttum (1925) mentioned that he found Prerocarpus indicus and

Morinda citrifolia on the island. Morinda citrifolia was also found in the

recent survey. In view of the relatively untouched nature of the island, in

all probability Pterocarpus indicus was, and Morinda citrifolia is, growing

naturally. Turner, Chua and Tan (1990) and Turner (1993) omitted

Pterocarpus indicus from their lists of Singapore vascular plants because it

was not considered a native or naturalised species but one that is only

cultivated and Morinda citrifolia was considered an alien species. Whitmore

(1972) commented that Pterocarpus indicus occurs naturally “in coastal

areas and tidal creeks along the east coast of Johor(e) and the Rompin

district of Pahang” in Peninsular Malaysia. It thus seems very possible that

this species also occurs naturally in Singapore which is immediately south

of Johore. Sinclair had also collected a specimen of Pterocarpus indicus on

22 Sep 1950 at Pulau Sakeng (Specimen - J. Sinclair SFN 39009). Although

Pulau Sakeng is close to Pulau Jong, it has been inhabited even before the

founding of Singapore by the British, so whether that tree from which the

specimen was taken, was cultivated, cannot be ascertained. To confuse

issues further, this species was introduced as a street tree so the vast majority

of trees in Singapore have been planted. Wong (1989) described Morinda

citrifolia to be “cultivated in villages and wild on lowland and rocky coasts’

so the specimens on Pulau Jong are likely to be wild plants. This species is

difficult to designate as native or alien because it is cultivated as well as

occurs naturally.

Acknowlegements

We are grateful for the assistance of Soong Beng Ching, Jean Yong Wan

Hong and Siti Dahlia binte Mohd. Dali as well as Avijit Gupta for information on

the geology of the area. We would like to express our appreciation to the Director

of the Botanic Gardens, Singapore for allowing us the use of the Herbarium and

116

the Chief Surveyor, Singapore for allowing us to reproduce in part, the 1:2500

topography map of Pulau Jong. This survey was supported by The National

University of Singapore Research Grant RP 880301.

References

Anonymous (1976). Geology of the Republic of Singapore, Public Works

Department, Singapore; Singapore.

Brooke, G.E. (1925). Notes on the Flora of Pulau Jong: Introduction. The

Singapore Naturalist 5, 47.

Chia, L.S. and Foong. S.F. (1991). Climate and weather. In: The biophysical

environment of Singapore edited by Chia, L.S., Rahman, A. and Tay,

D.B.H. Singapore University Press, Singapore. pp. 13-49.

Cox, C.B. and Moore, P.D. (1985). Biogeography: an ecological and

evolutionary approach. 4th ed., Blackwell Scientific Publications;

Oxford.

Holttum, R.E. (1925). Notes on the Flora of Pulau Jong: The vegetation.

The Singapore Naturalist 5, 48-50.

Johnson, A. (1977). A student’s guide to the ferns of Singapore Island.

Singapore University Press; Singapore.

Keng, H. (1990). The concise Flora of Singapore: Gymnosperms and

dicotyledons. Singapore University Press; Singapore.

Markgraf, F. (1951). Gnetaceae. Flora Malesiana I, 4, 337-347.

Ridley, H.N. (1930). The dispersal of plants throughout the world. L. Reeve;

Ashford, Kent.

Resource Centre, Ministry of Information and the Arts (1993). Singapore

facts and pictures 1993. Ministry of Information and the Arts;

Singapore.

Tan, H.T.W., Turner, I.M. and Chua, K.S. (1994). Flora-seed plants. In: The

Singapore Red Data Book of endangered plants and animals. Ed: Ng,

P.K.L. and Wee, Y.C. Nature Society (Singapore); Singapore; 18-49

Turner, I.M. (1993). The names used for Singapore plants since 1900. Gdns’

{ere >

rer ead FR

117

Bull., Singapore 45, 1-287.

Turner, I.M., Chua, K.S. and Tan, H.T.W. (1990). A checklist of native and

naturalized vascular plants of the Republic of Singapore. J. Singapore

Nat. Acad. Sci. 18 & 19, 58-88.

Turner, I.M., Tan, H.T.W., Ali bin Ibrahim and Corlett, R.T. (1994).

Checklists of threatened species - seed plants. In: The Singapore Red

Data Book of endangered plants and animals. Ed: Ng, P.K.L. and

Wee, Y.C. Nature Society (Singapore); Singapore; 273-313

Turner, I.M., Tan, H.T.W., Chua, K.S., Haji Samsuri bin Haji Ahmad and

Wee, Y.C. (1992). A botanical survey of Pulau Ubin. Gdns’ Bull.,

Singapore 44, 51-71.

Wee, Y.C. (1994). Checklists of threatened species - ferns and fern allies. In:

The Singapore Red Data Book of endangered plants and animals. Ed:

Ng, P.K.L. and Wee, Y.C. Nature Society (Singapore); Singapore;

269-272

Whitmore, T.C. (1972). Leguminosae. Tree Flora of Malaya 1, 237-304.

Wong, K.M. (1989). Rubiaceae. Tree Flora of Malaya 4, 324-425.

118

Table 1. The vascular plants of Pulau Jong.

_ This list includes those mentioned by Holttum (1925), Johnson (1977) and collections made by Holttum,

J. Sinclair and us. The specimens and nomenclature of Holttum and Sinclair were redeterminded and updated, respectively.

S/No.

Species

PTERIDOPHYTA

Blechnaceae

Blechnum orientatle L.

Davalliaceae

Davallia denticulata (J. Burm.) Mett.

Davallia solida (G. Forst.) Sw.

Humata heterophylla (Sm.) Desv.

Gleicheniaceae

Dicranopteris linearis (Burm.f.) Underw.

Hypolepidaceae

Pteridium caudatum (L.) Maxon ssp.

yarrabense (Dommin) Parris

Nephrolepidaceae

Nephrolepis biserrata (Sw.) Schott

Polypodiaceae

Phymatosorus scolopendria (Burm.f.)

Pic. Serm.

Pyrrosia lanceolata (L.) Farw.

Pyrrosia piloselloides (L.) M.G. Price

Johnson Collected

Specimen

R.E.Holttum s.n. 11 Jun 1924

KS. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 16

R.E. Holttum s.n. 9 Jun 1924

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 27

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 31

R.E. Holttum s.n. 13 Jul 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 19

KS. Chua, B.C. Soong, H.T.W. Tan,

L.M. Turner & J.W.H. Yong JONG 17

R.E. Holttum s.n. 9 Jun 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turmer & J.W.H. Yong JONG 11

R.E. Holttum s.n. 13 Jul 1924

119

S/No. Species Holttum Johnson Collected Specimen

(1925) (1977) Recently

Pteridaceae

ll. Acrostichum speciosum Willd. - - + K.S.Chua, B.C. Soong, H.T.W. Tan,

IM. Turner & J.W.H. Yong JONG 20

Schizaeaceae

12. Lygodium microphyllum (Cav.) R.Br. + - - R.E. Holttum s.n. 13 Jul 1924:

KS. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 13

Vittariaceae

13. Vittaria ensiformis Sw. var. ensiformis - - ~ KS. Chua. B.C. Soong, H.T.W. Tan,

IM. Turner & J.W.H. Yong JONG 45

PINOPHYTA

Gnetaceae

1. — Gnetan microcarpn Blume - - + KS. Chua, B.C. Soong, HT-W. Tan.

£ microcarpum I.M. Tumer & J.W.H. Yong JONG 28

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 38

Podocarpaceae

2. Podocarpus polystachyus R.Br. ex Endl. - - - KS. Chua, B.C. Soong, H.T.W. Tan,

1.M. Turmer & J.W.H. Yong JONG 26

MAGNOLIOPHYTA

Araliaceae

1. Arthrophyllum diversifolium Blume - - - K.S.Chua, B.C. Soong, H.T.W. Tan,

I.M. Turmer & J.W.H. Yong JONG 42

2. Schefflera elliptica (Blume) Harms - - > KS. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 4

Asclepiadaceae

3. Dischidia major (Vahl) Mert. - - - Nil

S/No. Species Holttum Johnson Collected Specimen

(1925) (1977) Recently

Euphorbiaceae

4. — Breynia reclinata (Roxb.) Hook. - - * KS. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 22

5. Macaranga heynei 1.M. Johnst. + - + R.E. Holttum s.n. 11 Jun 1924

Goodeniaceae

6. Scaevola taccada (Gaertn.) Roxb. + - - KS. Chua, B.C. Soong, H.T.W. Tan,

ILM. Turner & J.W.H. Yong JONG 6

Gramineae

7. Eriachne pallescens R.Br. + - + R.E. Holttum s.n. 13 Jul 1924;

KS. Chua, B.C. Soong, H.T.W. Tan,

LLM. Turner & J.W.H. Yong JONG 9

8. Imperata cylindrica (L.) P.Beauv. + - + KS. Chua, B.C. Soong, H.T.W. Tan,

var. major (Nees) C.E. Hubb. ex I.M. Turner & J.W.H. Yong JONG 12

C.E.Hubb. & R.E. Vaughan

Leguminosae

9. Dalbergia candenatensis (Dennst.) Prain + - + R-E. Holttum s.n. 13 Jul 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.WH. Yong JONG 35

10. Derris trifoliata Lour. - - + K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 7

11. Desmodium umbellatum (L.) DC. + - - R.E. Holttum s.n. 11 June 1924

12. Intsia bijuga (Colebr.) Kuntze + - - R.E. Holttum s.n. 13 Jul 1924;

J. Sinclair SFN 39001

13. Pongamia pinnata (L.) Pierre - - + K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 39

14. Pterocarpus indicus Wild. + - - Nil

Liliaceae

15. Dianella ensifolia (L.) DC. + - + KS. Chua, B.C. Soong, H.T.W. Tan,

j LM. Turner & J.W.H. Yong JONG 18

| ‘Pea

ie Ar Pee FS

>) Pe edi i

a >

121

S/No. Species Holttum Johnson Collected Specimen

(1925) (1977) Recently

Malvaceae

16. Hibiscus tiliaceus L. + - - Nil

Melastomataceae

17. Melastoma malabathnicum L. - - - KS. Chua, B.C. Soong, H-T.W. Tan,

[M-Tumer & J.W.H. Yong JONG 40

18. Memecylon edule Roxb. var. edule - - - J. Sinclair SFN 39004:

K.S. Chua, B.C Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 1;

K.S. Chua, B.C. Soong, H.T.W. Tan.

LM. Turner & J.W.H. Yong JONG 43

Meliaceae

19. Xylocarpus granatum J.Konig - - + K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 3

Moraceae

20. Ficus globosa Blume - - + K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 41

21. Ficus grossularioides Burm.f. + - + K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turer & J.W.H. Yong JONG 25

Myricaceae

22. Myrica esculenta Buch.-Ham. - - ~ K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 23

Myrtaceae

23. Eugenia grandis Wight + : + K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 29;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turuer & J.W. Yong 33.

24. Eugenia spicata Lam. + - + R.E. Holttum s.n. 11 Jun 1924

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 24

122

S/No.

aS.

26.

$40

28.

29,

30.

31.

(1925)

Species Holttum

Orchidaceae

Dendrobium crumenatum Sw. +

Pandanaceae

Pandanus odoratissimus Lf. +

Rubiaceae

Gynochthodes sublanceolata Miq. +

Morinda citrifolia L. -

Morinda umbellata L. +

Tarenna fragrans (Nees) +

Koord. & Valeton

Sapindaceae

Guioa pleuropteris (Blume) Radlk. +

Johnson Collected

Specimen

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 44

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 21

R.E. Holttum s.n. 11 Jun 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 2

R.E. Holttum s.n. 11 Jun 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 10;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 36

R.E. Holttum s.n. 11 Jun 1924;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 32

R.E. Holttum s.n. 13 Jul 1924;

J.Sinclair SFN 39005;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 4;

K.S. Chua, B.C Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 5;

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 37

R.E. Holttum s.n. 11 Jun 1924;

J. Sinclair SEN 39003; a

K.S. Chua, B.C. Soong, H.T.W. Tan,

. ee

IM. Tumer & WH. Yong JONG 15.

} Ry’

_— aa

Py i

S/No.

i 78

33.

34.

35.

36.

37.

Species

Sonneratiaceae

Sonneratia alba Sm.

Sterculiaceae

Commersonia bartramia (L.) Merr.

Heritiera littoralis Dryand. ex Aiton

Theaceae

Adinandra dumosa Jack

Verbenaceae

Premna corymbosa (Burm.f.)

Rottler & Wild.

Vitex pinnata L.

Holttum

(1925)

Johnson Collected

(1977)

Recently

123

Specimen

Nil

R.E. Holttum s.n. 13 Jul 1924

Nil

K.S. Chua, B.C. Soong, H.T.W. Tan,

LM. Turner & J.W.H. Yong JONG 30

Nil

K.S. Chua, B.C. Soong, H.T.W. Tan,

I.M. Turner & J.W.H. Yong JONG 8

Notes on the Flora of Malaya: New Records,

Overlooked Records and some

Nomenclatural Clarification

I.M. TURNER

Department of Botany

The National University of Singapore

Singapore 119260

Abstract

Browallia americana L., Persicaria nepalensis (Meisn.) H. Gross, Sonchus oleraceus L. and

Verbena bonariensis L. must be added to the flora of Malaya, being established as weeds in the

highlands. Ranunculus cantoniensis DC. is recorded for the first time for Malaya. Mitracarpus hirtus

(L.) DC. is probably also an established member of the weed flora of the lowlands. Desmodium

obcordatum (Migq.) Kurz is native to the far north of Peninsular Malaysia. Alsomitra macrocarpa

(Blume) M. Roem. was not included in earlier Malayan floras, but has been collected from the lowland

forests of several states in the Peninsula. Two species of Maclura occur in Malaya. Begonia perakensis

var. rotundata Irmsch. is reduced to the type variety of the species. Phrynium pubinerve Blume is the

correct name for the widespread lowland forest maranta referred to previously as Phrynium malaccense

Ridl. or Phrynium capitatum Willd.

Introduction

Whilst working on the compilation of a checklist of the vascular plants of

Malaya (Peninsular Malaysia and Singapore) I have come across a number of

records of species native or naturalized that have either not been published before,

or have been published rather obscurely making it unlikely that any but the

extremely persistent would come across the record. The purpose of this paper

therefore, is to list these new or overlooked records and also to attempt to clarify a

few nomenclatural problems in the flora.

New Weed Records from the Highlands

A high proportion of the new or overlooked records are weedy species

that have become naturalized around the hill resorts in the Peninsula. I

have not been able to find earlier records of their presence in the Malay

Peninsula in the literature for some. Others appear only in Stone’s Swmmit

Flora of Gunung Ulu Kali (Stone, 1981) which is not easy to obtain.

126

Browallia americana L. [Solanaceae]

Stone (1981 p. 144) reports this species as being naturalized on the

summit zone of Gunung Ulu Kali in the Genting Highlands. I have also

seen it growing as a weed of tea in the Cameron Highlands. A description

of this blue-flowered herb is given by Backer and Bakhuizen van der

Brink (1965 p. 482). This species is originally from South America but is

now naturalized in the palaeotropics.

Persicaria nepalensis (Meisn.) H. Gross [Polygonaceae]

This species has probably been accidently introduced into the

Cameron Highlands from the Himalayan region. It was first collected in

the Boh Plantations by Md. Nur (SFN 32847) on 3 April 1937. J. Sinclair

collected it in two localities the Cameron Highlands in August of 1956.

The species is described in Backer and Bakhuizen van den Brink (1963 p.

222) and figured in van Steenis (1972 plate 41-8) under the synonym

Polygonum nepalense Meisn.

Ranunculus cantoniensis DC. [Ranunculaceae |

I was surprised to see a buttercup growing on a grassy roadside

verge in Tanah Rata, Cameron Highlands. A visit to Kew allowed me to

identify my collection [I.M. Turner 94-3/] as Ranunculus cantoniensis DC.,

a species widespread in temperate and subtropical Asia. A single collection

from one small patch of plants is not sufficient to confirm naturalization of

the species, but it seems likely that this species will eventually be added to

the list; the first Ranunculus for Malaya.

Sonchus oleraceus L. |Compositae |

This softly-spiny yellow-flowered composite is now quite a common

weed of cultivation around the towns of the Cameron Highlands. Native to

temperate Eurasia, the earliest Malayan collection (in SING) was that of J.

Sinclair (9959) made on 4 November 1958 at the junction of Batu Brinchang

Road and Sungei Palas Tea Estate Road. For a detailed description see

Backer and Bakhuizen van den Brink (1965 p. 435).

Verbena bonariensis L. |[Verbenaceae |

Stone (1981 p. 150) reported this plant from Gunung Ulu Kali. It has

been collected a number of times from the Cameron Highlands also (e.g.

127

H.M. Burkill 2869, J. Sinclair 9931). It appears to persist readily as

an escape from cultivation as an ornamental herb. A detailed description

is given by Yeo (1990 p. 105).

New Records from the Lowlands

A few species found in the lowlands also appear to have been

overlooked:

Desmodium obcordatum (Miq.) Kurz [Leguminosae]

A twining subshrub with characteristic obcordate leaves, placed in

the monotypic genus Hegnera by Schindler but now generally treated within

Desmodium sensu stricto. | have seen three collections from Peninsular

Malaysia. The earliest (M.R. Henderson, $FN23079) was made on 22 Nov

1929 from Gua Nangka in Perlis. Two others come from rubber estates in

Kedah. Ohashi (1973) cites the distribution of this species as Indochina,

South Sumatra and Java, so its presence in the driest parts of northern

Peninsular Malaysia is not unexpected.

Mitracarpus hirtus (L.) DC. [Rubiaceae]

This small herb appears to have become naturalized from tropical

America. There are three collections in SING by J. Sinclair of this species

from near Kepong in Selangor (SFN40076), Telok Paku Road, Singapore

(10768) and the Scudai River in Johore (10825). All apparently from open

dry sandy sites. Inspection of such sites in other places may reveal this

species elsewhere in Malaya. Detailed descriptions are given by Verdcourt

(1976 under the synonym M. villosus (Sw.) DC., and 1989). It is now a

pantropical weed which originated from tropical South America.

Alsomitra macrocarpa (Blume) M. Roem. [Cucurbitaceae]

Reid (1953), in a short article, noted the presence of this

cucurbitaceous vine in the lowland forests of Peninsular Malaysia. There

are collections in SING from Pahang, Negri Sembilan and Johore. The

plant is remarkable for its football-sized fruits which contain winged seeds.

The fruits, or at least the empty fruit shells, are featured in Davison (1988

p. 127) being used as playthings by kampung kids in Johore. Winged seeds,

probably of this species, are shown gliding down through a rain-forest

128

canopy in Borneo in a spectacular film sequence in the BBC The Private

Life of Plants series.

Maclura in the Malay Peninsula

In an annotated key to genera of the Moraceae in Malaya,

Kochummen (1978 p. 120) states that Maclura amboinensis Blume is only

one native species belonging to this genus. Ridley (1923) also included

only one species referable to Maclura. This was Cudrania javensis Tréc.,

which is a synonym of Maclura cochinchinensis (Lour.) Corner. Kochummen

cites Cudrania javensis sensu Ridley as a synonym of Maclura amboinensis

but I wondered whether Ridley might have actually correctly identified at

least some of the Malayan material. Revisions of the genus (Corner, 1962;

Berg, 1986) do not contain any specific reference to Maclura cochinchinensis

in Malaya, but inspection of material in SING showed that both species

are present. Maclura amboinensis Blume is a spiny climber of hill and

montane forest collected from Perak and Pahang. Maclura cochinchinensis

(Lour.) Corner is a small tree found in Perlis, Kedah and Penang with a

majority of collections from limestone sites.

The varieties of Begonia perakensis King

In his monograph of Begonia in the Malay Peninsula, Irmscher (1929)

described two varieties of Begonia perakensis King. However, he made no

allusion to any type variety for the species. In describing the species, King

referred to three specimens of Kunstler’s, numbers 10338, 10506 and 10951.

Irmscher included /0338 in the list of specimens of his variety rotundata

and he rejected Kunstler 1/0566 from Begonia perakensis. I imagine that

King’s 10506 is the same as Irmscher’s 1/0566 but typographical errors have

occured somewhere. The third syntype given by King is not mentioned by

Irmscher. Thus there is a strong case for regarding var. rotundata as the

type variety of Begonia perakensis, a conclusion that would become

incontravertible if Kunstler 1/0338 were declared the lectotype of Begonia

perakensis. As | have not seen this specimen I refrain from lectotypification,

but provisionally I reduce var. rotundata to the type variety.

129

Begonia perakensis King, J. Asiat. Soc. Bengal, Pt. 2 Nat Hist. 71 (1902) 64

var. perakensis

Begonia perakensis var. rotundata Irmsch., Mitt. Inst. Allg. Bot.

Hamburg 8 (1929) 129.

var. conjungens Irmsch., Mitt. Inst. Allg. Bot. Hamburg 8 (1929) 129

The correct name for Phrynium capitatum

In his monograph of Malayan Marantaceae Holttum (1951) used the

name Phrynium capitatum Willd. for the common lowland forest maranta.

Recently, Suresh and Nicolson (1986) have shown that Phrynium capitatum

Willd. is an illegitimate name. They go on to provide a new name in

Phrynium for the oldest combination available for this species Pontederia

ovata L.; the new name being necessary as Phrynium ovatum was already

occupied by a species decribed by Nees and Martelli in 1823. Suresh and

Nicolson’s new name, Phrynium rheedei, would be the correct one for the

species if no other valid combinations in Phrynium existed for it. However,

it appears that the species is widespread and at least two earlier

combinations are available. The oldest being Phrynium pubinerve Blume

as given below:

Phrynium pubinerve Blume, Enum. PI. Javae (1827) 38; Back. & Bakh.f.,

Fl. Java 3 (1968) 79; Noltie, Fl. Bhutan 3(1) (1994) 214.

Pontederia ovata L., Sp. Pl. (1753) 288. Phrynium rheedei Suresh &

Nicolson, Taxon 35 (1986) 355.

Phrynium capitatum Willd., Sp. Pl. 1 (1797) 17, nom. illeg. Holttum, Gdns’

Bull. Sing. 13 (1951) 287.

Phrynium malaccense Ridl., J. Str. Br. R. Asiat. Soc. 32 (1899) 180.

Acknowledgements

I am grateful to the Director of the Singapore Botanic Gardens for

permission to use the facilities of the Gardens’ Herbarium and Library.

130

References

Backer, C.A., and R.C. Bakhuizen van den Brink (1963). Flora of Java, Vol.

I,N.V. P. Noordhoff, Groningen.

— (1965). Flora of Java, Vol. IT. N.V. P. Noordhoff, Groningen.

— (1968). Flora of Java, Vol. III. N.V. P. Noordhoff, Groningen.

Berg, C.C. (1986). The delimitation and subdivision of the genus Maclura

(Moraceae). Proc. Ned. Akad. Wet. ser. C Biol. Med. Sci. 89: 241-247.

Corner, E.J.H. (1962). The classification of Moraceae. Gdns’ Bull., Sing. 19:

187- 252.

Davison, G.W.H. (1988). Endau-Rompin: a Malaysian Heritage. Malayan

Nature Society, Kuala Lumpur.

Holttum, R.E. (1951). The Marantaceae of Malaya. Gdns’ Bull., Sing. 13:

254-296.

Irmscher, E. (1929). Die Begoniaceen der Malatischen Halbinsel. Mitt. Inst.

Allg. Bot. Hamburg 8: 86-160.

Kochummen, K.M. (1978). Moraceae. Tree Fl. Malaya 3: 119-168.

Ohashi, H. (1973). The Asiatic species of Desmodium and its allied genera

(Leguminosae). Ginkgoana 1: 1-318.

Reid, J.A. (1953). A remarkable winged seed. Malay. Nat. J. 8: 59-61.

Ridley, H.N. (1924). The Flora of the Malay Peninsula, Vol. IIT, Reeve &

Co., London.

Stone, B.C. (1981). The summit flora of Gunung Ulu Kali (Pahang, Malaysia).

Fed. Mus. J. 26: 1-157.

Suresh, C.R., and D.H. Nicolson (1986). Two nomenclatural novelties based

on Rheede’s “Hortus Malabaricus”. Taxon 35: 354-355.

van Steenis, C.G.G.J. (1972). The Mountain Flora of Java. E.J. Brill, Leiden.

Verdcourt, B. (1976). Rubiaceae (Part 1). Fl. Trop. East Africa 1-414.

— (1989). Rubiaceae. Fl. Zambesiaca 5(1): 1-210.

Yeo, P.F. (1990). A re-definition of Verbena brasiliensis. Kew Bull. 45: 101-120.

—

es 4

q f

‘

Additions to the Flora of Singapore, II.

I.M. TuRNER, H.T.W. TAn and K.S. CHua

Department of Botany,

The National University of Singapore,

Singapore 119260

Abstract

Seven species of vascular plants not previously reported for the flora of the Republic of

Singapore are listed in this paper. Three of these, the epiphytic orchid Bulbophyllum gusdorfii J.J.Sm..,

the icacinaceous liana Jodes cirrhosa Turcz. and the leguminous sea-shore shrub Sophora tomentosa L.,

are apparently overlooked native species. The other four, the sedge Cyperus difformis L., the fern

Pteris semipinnata L. and the leguminous shrubs Macroptilium lathyroides (L.) Urban and Sesbania

cannabina (Retz.) Poir., are naturalized exotic species.

The Republic of Singapore has a native and naturalized vascular

plant flora containing some 2,500 species (Turner, 1993). Despite fairly

intensive botanical inventory for more than a century it is still possible to

find additions to the flora. These are either overlooked native species or

exotics that have become naturalized after accidental or deliberate

introduction. In this paper we list seven new records.

1. Bulbophyllum gusdorfii J.J.Sm. [Orchidaceae]

An epiphyte of forest trees consisting of a horizontally creeping

rhizome bearing ovoid, slightly four-angled pseudobulbs up to 2.5 cm long,

each with a single coriaceous, oblanceolate-obovate leaf which can reach

16.5 by 3.5 cm in size. The pinkish scape arises from the base of the

pseudobulb, and is from 7 to 9 cm long. At the tip of the scape, the 6-8

flowers are found in an umbel-like arrangement. Each flower has a dorsal

sepal and about equal-sized lateral petals. The lateral sepals are joined for

almost all their upper edges, are up to 26 mm long and 8 mm at the widest

point, and are pale yellow flushed purple at the base. The identification

was confirmed from a plant (K.S. Chua et al., NRS/683, 29 Apr. 1993)

collected from a fallen branch in the forest to the north of MacRitchie

Reservoir near Thompson Ridge and grown until it flowered at the National

University of Singapore.

Bulbophyllum gusdorfii has been reported from Selangor, Pahang

and Johore in the Malay Peninsula, as well as from Sumatra and possibly

the Philippines (Seidenfaden & Wood, 1992).

2. Cyperus difformis L. [Cyperaceae]

A small annual, about 50 cm tall, with tufted stems, well-developed

but weak, leaves and reddish fibrous roots. The inflorescence is simple or

compound. It is made up of 5-9 spreading primary branches of up to 4 cm

long, often bearing shorter secondary branches radiating from their tips.

Each branch or branchlet ends in a spike consisting of a globose head of

up to 40 spikelets stellately arranged.

A specimen (K.S. Chua 1059, 11 Oct. 1994) of Cyperus difformis was

recently collected from a clump of sedges in a patch of waterlogged open

ground beside the road in Lim Chu Kang. Ridley (1925) reported that

Cyperus difformis was rare in the Malay Peninsula, occurring as a weed in

the ricefields of Kelantan. Elsewhere it is widespread across the Tropics

and Subtropics of the Old World, being found from southern Europe and

Africa to Australia, but in Malesia is only common on Java (Kern, 1974).

Expanses of waste land in Singapore appear to have favoured the

establishment of weedy species, such as this one, endemic to more seasonal

tropical climates.

3. odes cirrhosa Turcz. [Icacinaceae]

A liana with woody stems to 8 cm in diameter climbing with the aid

of tendrils. The simple leaves are roughly ovate and hairy beneath. The

tiny yellowy-white flowers are borne in much-branched lax cymes to 15 cm

long. The fruits are red drupes about 1.5 cm long. In Singapore the first

record of this species is a specimen (I.M. Turner et al. NRSO026, 1 Apr.

1992) collected from the margin of the Nee Soon Swamp Forest near

Seletar Reservoir Park. The species is found over a wide area of both the

Malay Peninsula and Malesia (Sleumer, 1971), so it does not seem unlikely

that it is native to Singapore.

4. Macroptilium lathyroides (L.) Urban [Leguminosae]

The phasey bean or kacang batang is a small shrub reaching about

1.5 m tall. In the open it remains erect but among other plants the young

branches may twine round adjacent stems allowing the plant to climb. The

leaves are trifoliate with ovate-lanceolate to elliptic leaflets. The “dried

blood” red flowers are borne in racemes up to 15 cm long with a 40 cm

peduncle. The narrow pods are more or less cylindrical, about 10 cm long

and given to abrupt dehiscence.

133

This species, native to Tropical America, has been used quite widely

in the Tropics as a forage legume (Jones & ‘t Mannetje, 1992) and probably

arrived in Singapore as an escape from an agricultural trial somewhere in

the region. It has been observed in a number of localities in Singapore,

seemingly preferring open sandy sites, particularly newly reclaimed land.

Collections include: Ali Ibrahim 95 (25 Jun 1987) from reclaimed land at

Marina South, I.M. Turner 93-/2 (17 Jan 1993) from reclaimed land at

Marina East and K.S. Chua et al. SB 3063 (13 Oct 1993) from Sungei

Buloh Nature Park, along roadsides at the Lim Chu Kang boundary.

5. Pteris semipinnata L. [Pteridaceae]

A terrestrial fern with fronds 30-40 cm long. The fronds are pinnate;

the terminal pinna being deltoid with lateral lobes incised close to the

rachis. The 4-7 pairs of subopposite lower pinnae are sessile with lobes

along their lower margins. The edges of the pinnae are minutely and

irregularly toothed. The pale brown spores are produced from sori running

round the margins of the fertile fronds.

Pteris semipinnata was first collected in Singapore by J. Sinclair in

1950 (SFN 39124) on Pulau Brani. The field note on the herbarium sheet

describes the locality as a clayey slope near the jetty and declares the fern

to be common in that site. More recently this species has been collected

from Labrador Park at the base of a Eugenia grandis tree (I1.M. Turner 92-

50, 23 Aug. 1992) and from a similar habitat in Mount Faber Park (I.M.

Turner 93-2, 1 Jan. 1993). Holttum (1968) noted that it had not been

recorded south of Malacca in the Malay Peninsula, and was commonest in

the more seasonal North East, where it was often found in light shade

around towns and villages. It occurs across Indochina and southern China

(Tagawa & Iwatsuki, 1989). It would seem likely therefore, given the

thorough plant collecting that has taken place in Singapore, that Preris

semipinnata has undergone a recent range extension. It appears to be a

species that is suited to the park habitat and development has favoured its

establishment in new areas.

6. Sesbania cannabina (Retz.) Poir. [Leguminosae |

An erect shrub to 4 m in height, found growing in wasteland and

newly cleared areas. The branches are characteristically held more or less

horizontally and bear paripinnate leaves. The flowers, borne in axillary

racemes, have yellow petals streaked with brown. The slender, straight to

curved, legumes may be up to 20 cm long. This species appears to have

134

spread across Singapore in the last 15 years or so and has now been

recorded from areas as various as Boon Lay Way, Bukit Batok Road,

Hindhede Drive, Marina East and Turut Track (collections include K:S.

Chua & H.T.W. Tan 353 and K.S. Chua and I.M. Turner 655).

Sesbania cannabina is probably native to Australia and parts of

Malesia, but has become established across the Old World Tropics including

Africa (Gillet, 1963).

7. Sophora tomentosa L. [Leguminosae]

A sea-shore shrub to 5 m tall characteristically with most parts more

or less covered with a dense grey silky tomentum. The pinnate leaves

possess 5-9 opposite pairs plus a terminal leaflet and are up to 30 cm long.

The bright yellow flowers are borne in racemes up to 30 cm long. These

give rise to pods which look like short strings of black beads because of

constrictions between each seed in the pod.

Sophora tomentosa is a pantropical coastal plant (Rudd, 1980). In

the Malay Peninsula it is common on the East Coast but on the west found

only north of Lumut in Perak (Corner, 1988). There are no earlier records

of this species from Singapore, yet it was recently collected (Turner et al.

LAZ 36, 29 June 1993) on Pulau Sakijang Pelepah (Lazarus Island) where

a single bush was found growing on the rocky southern shore of the island.

It seems that it is a resident species that has eluded detection by earlier

Singapore-based botanists. Seed was collected from the plant and

successfully germinated in the Department of Botany, The National

University of Singapore. It is hoped to establish the young plants at various

suitable localities in Singapore to expand the population of this attractive

native plant.

Acknowledgements

Dr Frits Adema and Dr Wee Yeow Chin are thanked for their

assistance with the identification of specimens.

References

Corner, E.J.H. (1988) Wayside trees of Malaya. Malayan Nature Society, Kuala

Lumpur.

Gillett, J.B. (1963) Sesbania in Africa (excluding Madagascar) and southern Arabia. —

Kew Bulletin 17: 91-159.

135

Holttum, R.E. (1968) A revised flora of Malaya. Vol. II. Ferns of Malaya, Second

Edition, Government Printing Office, Singapore.

Jones, R.M., & ‘t Mannetje, L. (1992) Plant resources of South-East Asia No 4

Forages. Pudoc Scientific Publishers, Rotterdam.

Kern, J.H. (1974) Cyperaceae. Flora Malesiana, Series 1 7(3): 435-753.

Ridley, H.N. (1925) Flora of the Malay Peninsula. Vol. V.L. Reeves & Co., Ashford,

Kent.

Rudd, V.E. (1980) Fabaceae. Revised Handbook of the Flora of Ceylon 1: 428458.

Seidenfaden, G., & Wood, J.J. (1992) The orchids of Peninsular Malaysia and

Singapore. Olsen & Olsen, Fredensborg, Denmark.

Sleumer, H. (1971) Icacinaceae. Flora Malesiana, Series 1 7(1): 1-87.

Tagawa, M., & Iwatsuki, K. (1989) Pteridaceae. Flora of Thailand 3(2): 231-260.

Tumer, I.M. (1993) Names used for Singapore plants since 1900. Gdns’ Bulletin,

Singapore 45: 1-287.

Book Reviews

Flora of Australia Volume 49 : Oceanic Islands 1.

Pp xxili + 681. 1 black-and-white photograph, 16 colour plates (63 colour photographs), 3

maps, 1 coloured and 41 line drawings. Australian Government Publishing Service, Canberra, 1994.

ISBN 0 644 29384 5 (pbk.). Price $A 54.95 (paperback), $A 64.95 (hardcover). Available by mail from

AGPS Mail Order Sales, GPO Box 84, Canberra ACT 2601, Australia.

Though listed as Volume 49, this first volume on the flora of Australia’s

Oceanic Islands came one year after the second volume (Volume 50), covering

Part 2 of Oceanic Islands, was published. It covers Norfolk Island and Lord Howe

Island in the Tasman Sea. Written almost entirely by Mr Peter Green, formerly a

botanist at the Royal Botanic Gardens, Kew, this volume includes 136 families,

455 genera and 706 species and subspecific taxa.

The layout of the book is excellent and the descriptions are brief but

diagnostic. The clever use of different font and type sizes is easy on the

eyes and enables the reader to use the book with ease. Keys to the families,

genera and species are clear and well set-out.

The separate checklists of plants in Norfolk Island and Lord Howe

Island in the introductory chapter are very useful. Endemic and naturalised

species listed are marked with different symbols to allow readers a quick

survey of the flora at the specific level.

The inclusion of a glossary towards the end of the book is most

welcome for readers who are not specialists. This is one of the useful

features in Flora of Australia and should set the trend for modern flora

writing.

There are a few minor errors and omissions in the book. The colour

photograph of Fig. 21 (p. xxi) appears to be inverted while those of Fig. 11

(p. xviii) and Fig. 58 (p. 283) would look more natural sideways. It would

have been useful to include a list of illustrations and photographs. About

63% of the families covered are illustrated with line drawings though I feel

that more could have been incorporated.

In the Locality Map (fig. 32), the name Vanuatu should be adopted

rather than New Hebrides. This is more so when Vanuatu is mentioned in

the text (p. 2), The name New Hebrides could have been included under

parenthesis to inform readers who are familiar with it.

138

On page 2, it is stated that the degree of endemism of vascular

plants in Norfolk and Lord Howe Islands is 44.9 %. This percentage of

endemism should actually be 43.2 considering there are 149 endemics in

the 345 indigenous species. This is a very high percentage of endemism for

these islands with a total area of about 51.2 sq. km. The explanation of the

specific epithet for Ricinus communis has been omitted. The genus

Sanseviera on p. 523, 525 and 676 should be spelt as Sansevieria.

Despite these minor errors and omissions, I have no reservation in

recommending this book to anyone interested in the flora or even the

geography, climate, physical features and history of human habitation of

these two islands. Considering the amount of time and effort put into the

preparation of this book and the excellent production, not forgetting the

impeccable editing, the cost is very reasonable. ;

;

‘

Tay Eng Pin

National Parks Board, 5

Singapore. f

Rattans

Dransfield, J. & Manokaran, N. (eds.), 1993. Plant Resources of South-East Asia (PROSEA)

No. 6. Pudoc Scientific Publishers, Wageningen. 1993. 137pp. ISBN 90-220-1057-0. Hardbound. Dfl.

120.00. Available from, PROSEA Foundation Publication Office, Wageningen Agricultural University,

P.O. Box 341, 6700 AH Wageningen, The Netherlands. For developing countries a cheap paper

edition (about US$ 10.00) is available from, PROSEA Network Office, P.O. Box 234, Bogor 16122,

Indonesia.

This is the sixth volume from PROSEA, an international programme

on the documentation of information on plant resources of Southeast Asia.

This volume by 17 contributors provides details on 23 species and one

genus of rattan that are commercially important or have potential to be so.

Another 105 less important species are briefly discussed. It is not an

identification manual; keys are not provided.

Over virtually all of Southeast Asia the use of rattan in village life is

ubiquitous and significant.. Traditional cultures would not be the same

without rattan. This product has also contributed to the building of cities

in this part of the world as the required binding material for wooden

139

scaffoldings. Several loops of rattan strips at each junction and joint, hold

together massive scaffoldings. A demonstration of its toughness and high

coefficient of friction?

The first part of the book is a concise introduction to the subject. It

covers a wide range of topics including, origin and geographical distribution,

uses, history of the rattan trade, morphology, growth and development.

ecology, exploitation of wild resources, cultivation and research priorities

and development. It is made clear that serious rattan research in response

to a rapidly diminishing natural resource dates back only 15 years. This

perhaps is the cause for unclear statements in the text. For example in the

bottom paragraph of page 34, “There may be little control over the

collection of rattans from the wild in many countries.” Is there any doubt

that there is little control? In the bottom paragraph on page 35, after

saying that large-diameter canes have to be cured with a hot oil mixture,

the sentence continues, .....” this treatment is said to make the canes durable

by removing gums, resins and water.” Yet on page 52 it is quite clearly

stated (by a contributor) that this curing is to protect the canes from attack

by staining fungi and the powder-post beetle. In the bottom paragraph on

page 36 it is stated that, “Even where licences are issued and royalties paid

to forest departments, there is evidence to suggest that harvesting is carried

out with little thought for sustainability.” The author of this is overly

optimistic to expect that because licences are issued and royalties collected,

the licensee will do something about sustainability (one is reminded about

the logging industry). In any case it is expressed that the basic data required

for understanding possible rates of harvest of wild populations are still

being compiled (top paragraph on page 37).

Less explicable are slips in the citations. On page 16 there is reference

to Brown (1941-1943), in the literature listing this is Brown (1951-1957).

The years for some of the citations in the text (pages 34, 37 & 38) are

absent; neither are these references listed in the literature. However,

overall, the introduction is useful and very interesting.

Part Two, the main part of the book, is an alphabetical treatment of

major species. All available information (some species have little) seems

to have been neatly summarized under the following headings; vernacular

names, origin and geographical distribution, production and international

trade, properties, description (or sometimes, botany), growth and

development, other botanical information, ecology, propagation and

planting, husbandry, diseases and pests, harvesting, yield, handling after

harvest, genetic resources, breeding, prospects, and literature. Although

the contents emphasize cultural practices and handling of the product, the

botanical description of each species is detailed and accompanied by a

140

good quality quarter-page line drawing showing all important parts.

Part Three of the book treats the minor species alphabetically.

Information for each species appears under these headings; vernacular

names, distribution, uses, and observations (which normally include notes

on the botany, distribution and ecology).

The literature is listed in three different places: after each species in

Part Two, where, as if emphasizing that these are important species, the

reference is spelled out in full; at the end of Part Three where the references

are numbered and only the numbers are cited after each species entry; and

at then end of the book under the heading, “Literature.” In this final list

not all items found in the first two parts appear.

This is a useful addition to the growing literature on this important

resource. The availability of a cheaper, “developing country” edition is

most welcome, especially as this edition, though with a soft cover, appears

identical in the paper, binding and printing to the hardbound one. The

editors and editorial staff have done a marvellous job creating a uniform

product from the contributions of so many.

Chin See Chung

National Parks Board,

Singapore.