LODE EOE EE

- My | | SS#/

Ke a

‘THE F3

» GARDENS’ BULLETIN

A SINGAPORE Y

oe eG

Vol. XXII, Part I 20th January, 1967

ae YD

CONTENTS

y PAGE

() Wuitmore, T. C.: Notes on the Systematy of Solomon Islands’

x Plants and some of their New Guinea Relatives, 1—VII - 1

re MarkearaF, F.: Ibid, VIII and IX - - - . 23,

& VAN RoyeEN, P.: Ibid, X_ - - - - . - ao

‘ A . Hoxtrum, R. E. & B. M. ALLEN: The Tree-ferns of Malaya - 41

\) GREEN, S.: Notes on the distribution of Nepenthes species in

PN _ Singapore - - - - - -' 53

re Hoittum, R. E.: Isaac Henry Burkill, 1870-1965 9 - Bros

Pr BurkKILL, H. M.: Ibid, a bibliography - - - - Mey | »

Q J. P. & H.M.B.: H. B. Gilliland, 1911-1965, an appreciation 107 §

N Hsuann KENG: Observations on Ancistrocladus tectorius - 113 \\

0) . Turner, G. J.: New records of plant diseases in Sarawak for

a5 the year 1965 - - - : - - - 123

To be purchased at the Botanic Gardens, Singapore

Price: $8

LEAD

LE LENO ODEO

K, Fy

—m® ‘ ¢ A

¢ . Ni

Mame fo. 5 Published by Authority | ‘)

D. q Printed by Lim Bian Han, Government Printer, Singapore \

\ or, ; ;

1e 9 << RED (

x Be 6 cet taal ‘

AY SIVED ‘Ap 2

Ad ; a el " \

a \¢ - s - - “ oS eo Fe f2_° pean

ReROPROPROPROPROFRO ROAR OSE RE BROPEAIBOA

pie

S200 7

MAR | 21968

THF

GARDENS’ BULLETIN

SINGAPORE

A scientific periodical for the publications of results of original

research in Botany in general and Plant Taxonomy in particular.

Edited by the Director of Botanic Gardens, Singapore, it is

published as material becomes available. Price ca. M$25 per

volume payable in advance. It is also available for exchange.

All manuscripts, enquiries, purchases and exchanges are to be

addressed to: —

The Director,

Botanic Gardens, —

Singapore 10,

Singapore.

COMPLIMENTARY

THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XXII, Part I 20th January, 1967

Notes on the Systematy of Solomon Islands’ Plants and

some of their New Guinea Relatives, I-VII .

T. C. WHITMORE

Forest Botanist, British Solomon Islands*

Summary

This paper adds to the bonfire of optimically proposed ‘paper-species

whose incineration is inevitable following a critical examination of the .

abundant collections now available from Malesia and Melanesia.

The amount of mophological variation differs from group to group

studied. Several classes may be distinguished and interpreted as stages in

the evolution of species, as follows:

1. Homogeneous groups, without morphological discontinuities; previously

optimistically divided:

(a) Campnosperma brassii reduced to C. brevipetiolata (p. 4).

(b) 3 Syzygiums transferred to Eugenia (p. 16).

(d) 2 spp. reduced to Tristiropsis acutangula (p. 17).

(e) Gmelina salomonensis reduced to G. moluccana (p. 18).

2. Evolution to a wide range of habitats but morphologically uniform,

various Calophyllums (p. 13).

3. ‘Ochlospecies’, ic. a group showing complex reticulate variation which

cannot be treated taxonomically:

4 spp. reduced to Schizomeria serrata (p. 5).

4. Widespread polymorphic species with variously distinct local popula-

tions:

(a) Buchanania arborescens (p. 2); of which B. novo-hibernica and

B. solomonensis are synonyms.

(b) Pometia pinnata (p. 17).

5. Closely related but distinct allopatric species pairs:

(a) Campnosperma brevipetiolata and C. auriculata (p. 4).

(b) Endospermum medullosum and E. malaccense (p. 8).

Gardens’ Bulletin, Singapore — XXII (1967)

Introduction

I have found it necessary in connection with the preparation

of a ‘Guide to the Forests of the Solomon Islands’ (Whitmore

1966, pp. 208, Oxford University Press) to investigate the taxo-

nomy of a number of the common, big trees of the archipelago.

These trees are important components of the rain-forests and

some of them have already or soon will enter commerce. For

both the student of ecology and the forester a sound taxonomic

base is essential. In these notes are set out the results of my

studies.

_In the Solomons, as in Malesia, it is being discovered that

many formerly recognised species cannot be maintained now

that abundant collections from many localities are to hand. Earlier

botanists, with few collections from widely separate localities,

saw distinct species where now various patterns of morphological

variation are revealed which show different stages in the evolution

of species (see summary). The early splitting was aided by a

firm preconception in the minds of the investigators of the

richness of tropical floras. Thus many of these notes follow the

current trend and involve the reduction of local “‘endemics” to

synonymy with the earliest described species of each group. In

each case I have pursued the problem as far as necessary to

provide what I hope will prove a stable name for the Solomons’

taxon, a name which future monographers will uphold. The

implications of this reduction of “species” for our conception of

the Malesian and Melanesian floras and on plant geography are

huge and important, and the time is coming when a reassessment

must be made.

_ The notes are based mainly on the set of specimens retained in

the Forest Hebarium Honiara, BSI[P. Replicates are at K, L, LAE.

SING and US; the last two sets are incomplete.

It is a pleasure to acknowledge the substantial assistance I

have received at Lae and Leiden with this work; in addition

Kew. British Museum, Singapore and Brisbane have kindly allowed

me to use their facilities and loaned me material for examination.

Dr. B. C. Stone kindly read a draft of this paper.

~ Professor H. Godwin F.R.S. very kindly allowed me to work in

the Cambridge University Botany School to complete these studies.

I. Anacardiaceae

A. Buchanania Roxburgh

There grows in the Solomons a species of Buchanania as a

scattered component of disturbed lowland rain-forest. It is a tree

of moderate size in the upper canopy and reaches about 6 ft.

girth and 120 ft. tall, with a tall, narrow, dense crown. It is

easily recognisable in the forest from various features (Whitmore

1966); and diagnostic is its red middle bark (phelloderm) revealed

when the papery superficial outer bark (phellem) is scraped off.

Whitmore — Solomon Islands’ Plants 3

In considering the name of this species | have examined the

following diagnostic features of all collections at Lae from the

Solomons and Bismarcks. The leaf apices are acute, sometimes

acuminate or, uncommonly, obtuse: both acute and obtuse leaves

are sometimes found on a single twig (BSIP 1472). The petiole

length varies from 15-SO mm.; on a single tree the variation

between leaves is about 10 mm. There are from 15-15 pairs of

secondary nerves whose spacing varies and whose parallelness

increases with closeness; they run more or less parallel towards

the margin but loop upwards well within it. The inflorescences

are commonly tomentose on the minor axes and have scattered

hairs on the main axes, sometimes they are almost glabrous but

only on BSIP 315 and BSIP 1087 could no hairs at all be found.

The carpels are hairy all over, or, more commonly, are glabrous

on the exposed side and hairy on the inner sides, the degree of

hairiness varies considerably but no completely glabrous carpels

were seen.

This Buchanania fits B. solomonensis Merr. et Perry (J. Arn.

Arb. 22, 530-1, 1941, based on Kajewski 1873 type, 2123 and

Brass 2745 which is the only species so far recorded from the Solo-

mons. I have seen an isotype at the British Museum; it matches my

own collections. Merrill and Perry based their species on three

fruiting collections. The remnants of a flower were found attached

to one of the fruits, and within this flower lies the only stated

difference from B. novo-hibernica Ltb. (Bot. Jahrb. 65, 349, 1921).

namely the gynoecium is pubescent not glabrous. In view of the

variation in hairiness of the carpels in my collections I do not

consider this character alone enough to establish a new species

and I cannot discover others in the descriptions. Accordingly I

consider B. solomonensis to be the same as B. novo-hibernica.

Unfortunately this decision.must be made without seeing the

type and only cited collection of B. novo-hibernica (Peekel 812)

which appears to have been lost; but I have examined both the

figure in Peekel’s manuscript flora (Nachtr. 84—5) and the descrip-

tion, which he bases on many observations of living material, of

what he describes as a very common species, and can see no

difference between B. solomonensis and B. novo-hibernica.

Lauterbach based B. novo-hibernica on a single collection. In

the same paper he mentions B. arborescens Bl. (Mus. Bot. Lugd.

— Batav., 183, 1826), a widespread polymorphic species, which

he calls B. florida Schauer. Unfortunately he does not give a dia-

gnosis for B. arborescens and the differences from B. novo-

hibernica must be taken from his key. There is in fact only one,

namely side nerves ‘unter sich. siemlich parallel’ versus ‘nicht

parallel’. This is a very slight distinction (see above) which I

consider inadequate to maintain two species, so B. novo-hibernica

becomes a synonym of B. arborescens.

There is a number of species described since B. arborescens

in this group. In W. Malesia I note solely B. lucida Bl. which

appears superficially very similar from the many sheets at Kew.

4 Gardens’ Bulletin, Singapore — XXII (1967)

In Queensland, E. Malesia and Melanesia are B. florida Schauer,

B. mangoides F. Muell.*, B. longifolia Span.*, B. montana Ltb.*,

B. nabirensis Kan. et Hat.*, B. monticola Kan. et Hat.*, B. papuana

C. T. White* and B. vitiensis Engl. (from Fiji). I have examined

at Kew and/or Leiden (on loan from Bogor) type or authentic

collections of the species starred (*) and descriptions of all those

listed. It appears to me that we are dealing with a single, wide-

ranging, polymorphic species with local morphological variants

which may or may not be distinct enough to be considered separate

species. The problem must await a monographer. As far as the

Solomons and Bismarcks are concerned however it looks as

though the regional variants, B. novo-hibernica and B. solomonen-

sis, are not sufficiently distinct from each other or from the main

plexus to be given specific rank and | think they can both be

considered as part of B. arborescens.

BSIP 1656 from Vanikoro, Santa Cruz Islands, matches the

type, viewed at Kew, of B. attenuata A. C. Smith, from Fiji. This

appears distinct from B. arborescens which has not been collected

from the Santa Cruz islands yet.

Buchanania specimens examined from Solomons

and Bismarcks

SOLOMONS (BSIP series unless indicated otherwise).

Bougainville Kajewski 1873, 2123.

Choiseul 4038, 5215.

Rob Roy 5311.

New Georgia Islands Baga 1327, 1375 Kolombangara 1472

New Georgia 275, 3277, 4701, Waterhouse 227 Gizo 3046,

5636 Gatukai 315 Vangunu 915, 948, 983.

Guadalcanal 49, 745, 786, 1072, 1087, 1137, 2558.

Malaita 71, 3442, 3496, 4457.

Santa Ysabel 2479, 2627, 2737, 3614, 4075.

San Cristobal 4276.

BISMARKS.

New Britain NGF 1838, NGF 10067.

B. Campnosperma Thwaites

C. brevipetiolata Volkens Bot. Jahrb. 31, 446, 1902.

C. brassii Merr. et Perry J. Arn. Arb. 22, 535, 1941.

The type and only collection of C. brassii, Brass 3355, comes

from Santa Ysabel, Garona. I revisited this locality and with some

difficulty penetrated the coastal swamp on to the foothills where,

as Brass states, it is a conspicuous tree. Brass 3355 (isotype from

Brisbane viewed on loan) has leaves very hairy below and has

dried a rich chestnut brown. The very many collections we have

now made of C. brevipetiolata have variable tomentum on the

Whitmore — Solomon Islands’ Plants 5

lower leaf surface but some sheets including the collection from

Garona are as hairy as C. brassii. As my material was all collected

into spirit, using Mr. Womersley’s technique, and Brass’s was.

dried in the field the colour of the specimen is not significant. In

all other respects C. brassii is identical with C. brevipetiolata.

Like so many of the species founded by Merrill and Perry on

one or two collections this ‘Solomons’ endemic’ is revealed for

what it is and must be sunk into synonymy with a widespread

species.

C. brevipetiolata is very similar in the herbarium to C. auriculata

Hk. f. of western Malesia, also a species of disturbed places.

The two species look rather different in the forest, inter alia

young trees of C. auriculata have very much bigger leaves. Pending

a monographic study I prefer to leave the two distinct.

II. Cunoniaceae

Schizomeria D. Don

Schizomeria serrata Hochreutiner Ann. Cons. Jarb. Bot Genéve

10, 188, 1907.

S. floribunda Schlechter Bot. Jahrb. 52, 156, 1915.

S$. pulleana O. C. Schmidt Nova Guinea 14, 150, 1924.

S. brassiti Mattfeld J. Arn. Arb. 20,435, 1939.

S. whitei Mattfeld loc. cit. 435-6.

In the lowland rain-forests of the Solomons is a Schizomeria.

It is a variable tree and the problems are whether it is one or

more species and then what to call it. The bark varies from

smooth to scaly to fissured, a wide range for a single species,

and there is considerable variation in the characters of herbarium

specimens.

From twenty-six collections we have made in B.S.I.P. great

variation can be seen in leaf margin, leaf base, inflorescence size

and shape. Concerning leaf characters, at one extreme are collec-

tions with lanceolate, cuneate-based, closely serrate leaves (BSIP

313, 1225, 1397, 1455, 2891, 3565) and at the other extreme

collections with broadly ovate, round-based, crenate leaves (BSIP

308, 1144, 2764, 4050). There is a series between these extremes,

and some collections even have both cuneate and round-based

leaves (BSIP 1144, 2764, 4050, 5613); nor is there a sharp dis-

junction between serrate and crenate margin. Concerning inflore-

scence characters the panicle varies very much in size. At one

extreme are BSIP 976, 1225, 1397, 2748 with compact panicles.

less than 3 cm. cross and these are connected via BSIP 2764

5269, 5270 with panicles 10 cm. long x 7 cm. across to the very

large open panicles of BSIP 1144, 18x15 cm. There are inter-

mediates between these stages. There is a rough but not invariable

correlation between the series or spectrum of leaf characters and

those of the inflorescence. However it also appears to matter how

6 Gardens’ Bulletin, Singapore — XXII (1967)

mature the inflorescence was at collection, the flowers on the

compact inflorescences are mostly in bud suggesting that these

are young and would later have expanded. Concerning fruit shape,

one collection, BSIP 3140, has spherical fruits, some. BSIP

893, 976, 2891, 3383, 3565, 5613, NGF 8445, have ellipsoidal

fruits, and a few collections have both spherical and ellipsoidal

fruits, BSIP 821, 1328, 1397, 4050. Fruit shape is not correlated

with the other characters described and probably depends on

degree of maturity. The fruits sometimes reach 15 mm. long but

are usually 8-10 mm. In none of these three series of characters

are there sharp disjunctions and the series are not correlated with

geography.

The petals have fallen off the open flowers of all collections

examined. The flowers vary in size, from 1-3 mm. across but

only reach 2-3 mm. when the fruit has begun to develop (BSIP

1144).

Because the various characters vary fairly independently of

each other and are each and every one connected by intermediate

stages I think all these collections must be placed in a single

species.

White (in ‘Taxonomy and Geography’, 1962 p. 79 et seq.) has

coined the useful and appropriate term ochlospecies (Gr. oklos-an

irregular crowd, a mob; also ‘trouble’, ‘annoyance’) for species

showing a complex reticulate pattern of variation of this kind and

in which the morphological disjunctions are insufficient for us to

recognise taxonomic categories. In the Buchanania arborescens

group analysed above morphological differentiation has proceeded

further to produce distinct local forms which can be given taxono-

mic status. In other words evolution has gone further in the

Buchanania arborescens group than in Schizomeria serrata.

Several species of Schizomeria have been described which all

fit our material. At Kew I have examined authentic material of

the following species of this group.

S. serrata: Hochreutiner 103 (isotype); also two recent collections

from same tree (VII a 12 at Bogor): Soepadmo 10, 23.xi.59

(sheet also at Lae); Rastini 164, 14.11.60.

S. floribunda: Ledermann 9763 (isotype), Ledermann 9664.

iY pulleana: Lam 1474 (isotype).

S. brassii: Brass 713 (isotype).

S. whitei: Kajewski 1336 (isotype) 1135.

The characters on which these species have been differentiated

are mainly those demonstrated above to vary more or less

independently and continuously. The collections from the Solomons

cover the whole range of morphological variation of all these

species. I have also examined twenty-one collections from New

Guinea and the Moluccas; they all appear similar. Now that

abundant material is available I do not think any of these species

can be maintained, the formerly perceived disjunctions break

down.

Whitmore — Solomon Islands’ Plants 7

I conclude that the wide-ranging lowland Schizomeria of Sahul-

land is probably best regarded as one polymorphic species for

which the earliest name is S. serrata Hochr. and that S$. floribunda,

S. pulleana, S. brassii, and S. whitei must be reduced to synonmy

now. In all probability when material can be traced S. katastega

Mattf. (J. Arn. Arb. 20, 434, 1939), S. tegens Mattf. (loc. cit.) and

S. homaliiformis Kan. et Hat. (Bot. Mag. Tokyo, 56, 109, 1942)

they will also prove synonymous.

Two further observations must be made: —

(a) S. pulleana is said to differ from S. floribunda in various of

the features demonstrated inconstant above but also in possessing

longer and irregularly 3-lobed petals. However as the petals are

early caducous in Schizomeria this character cannot be considered

satisfactory or adequate to uphold a species.

(b) S. serrata: the type and the figure in Ic. Bog. 3, t. 228, 68-70,

1907 have big lanceolate leaves about three times as long as broad

whereas |S. floribunda usually has ovate leaves about twice ‘as

long as broad. The difference is so striking that I at first was

inclined to keep the species separate, but BSIP 1397, 1455, 2891,

3565 and 4731 have lanceolate leaves, although rather smaller

than the type of S. serrata. Finally however I viewed Rastini 164

from the type tree of S. serrata and this has both shapes of leaves

on a single specimen. The flowers of this tree are only 2-3 mm.

across on the specimens although Hochreutiner states them to be

7-8 mm. diam. The biggest dried fruit is 15 x 12 mm., about the

same size as that of S. floribunda. Thus the material, but not his

description, falls within the range of all the other species in the

group. I take it he must have viewed and described living

specimens.

Schizomeria serrata material examined at Kew and/or Lae, loci

from west to east: in addition to authentic material listed above.

MOLUCCAS

Solea bb 28867.

Morotai Kostermans 996, 1081, 1084, 1205, 1278.

Ambon Robinson 603, Teysmann s.n. (cult. in Hort. Bog.)

NEW GUINEA

West W. New Guinea Div. BW 4472, 5781, 7604

| Hollandia Div. BW 7995

Biak Dijk 914 ,

East Morobe Distr. NGF 3145, 8445, 19045

Sepik Ledermann 9664

Papua Sorong Pleyte 996

Sogeri NGF 4200

Garitar Brass 713

Misima Isl. Brass 27534, 27653

Oriomo R. Brass 5804

8 Gardens’ Bulletin, Singapore — XXII (1967)

BISMARCKS

New Britain NGF 1893

SOLOMONS (BSIP series unless indicated otherwise)

Bougainville Voyce B3, D8; Waterhouse K 733

Choiseul 4050, 5269, 5270

Rob Roy 5343, 5356

New Georgia Islands Baga 1328, 2891 Gizo 5613 Kolombangara

821, 1397, 1455 New Georgia 3140, 3789, 4731, 4765 Vangunu

155,893; 976;1225

Santa Ysabel 2748, 2764, 3565, Brass 3428

Guadalcanal Tina 1105, 1144, Rere 308, 313, 3383

II. Euphorbiaceae

Endospermum Bentham

Endospermum medullosum L. S. Smith Proc. R. Soc. Queensl. 58,

53, 1947.

Endospermum malaccense Muell. Arg.* Flora 47, 469, 1864.

The common lowland Solomons tree A’asa (Kwara’ae) has been

called E. medullosum from Walker’s early collections (BSIP 97,

304), determined at Brisbane, onwards. It is a perfect match for

the abundant collections at Lae, including the type and other

sheets cited by Smith. The species is distinctive in forest and

herbarium.

I have examined at Kew the type of E. malaccense (Griffith

4721) and Maingay 1392 which also is cited in the “Pflanzenreich’

account, as well as many old and recent western Malesian gather-

ings. E. medullosum is extremely similar to this species and sterile

trees are indistinguishable. Smith makes it clear that he saw no

western Malesian material when he discribed his species. There

is a slight yet consistent difference in the inflorescence and flower

stalk, which can be summarised as a key:

Male and female inflorescences paniculate; branches often them-

selves branching;; branch systems usually at least 2.5 cm. long:

flowers and fruits sessile, or on very short stalks 1 mm. long.

—E. medullosum

Male and female inflorescences spikes or narrow panicles; branches

simple, to 2.5 cm. long. Flowers and fruits on distinct stalks c. 10

mm. long.

—E. malaccense

me Airy Shaw (Kew Bull. 14, 395, 1960) considers ‘it highly probable that

the correct name for this species is E. diadenum (Miq). may Shaw, and I

agree with him.

Whitmore — Solomon Islands’ Plants . 9

On the basis of these differences I propose that provisionally

the two species are kept distinct. When more fertile gatherings

come to hand from the region of ‘“‘Wallacea’’” between western

and eastern Malesia it may be found that, as in so many other

cases, we have just one widespread variable species.

Collections of E. medullosum, and state of material examined,

from Solomons (all BSIP series): —

Shortland 5744 (fr.)

Wagina 5470 (very young inflorescence)

New Georgia Islands Baga 4202 (seedling) Kolomban-

gara 1431 (fr.) New Georgia

1978 (fr.) 2543 (fr.)

Santa Ysabel 2124, 2202 (it.)

Guadalcanal 1169 (very young inflorescence}

Malaita 3451 (very young inflorescence)

IV. Guttiferae

Calophyllum Linnaeus

A. Big Coriaceous Leaves Species

C. kajewskii A. C. Smith J. Arn. Arb. 22, 353, 1941.

C. solomonense A. C. Smith loc. cit., 346.

We have in the Solomons a big-leafed, big-fruited Calophyllum

which is a common tree of lowland rain-forest. This note is to

establish its identity and also that of a much less common species.

C. kajewskii is based on Kajewski 2024, a sterile branch with

a single detached fruit. Waterhouse 201 is also cited by Smith;

the sheet seen was sterile but the description stated ‘orang-like

fruit’. Smith states ‘the foliage closely resembles that of C. inophy-

llum’, but the species is unique in the region with its extremely

large fruit reaching 5-6 cm. diameter.

C. solomonense is based on Kajewski 2469. Brass 3447 was

also seen. Like C. kajewskii the foliage ‘bears a close resemblance

to C. inophyllum, it differs obviously in the greatly contracted

inflorescence’ (Smith).

The two species are not at all clearly distinguished in Smith’s

latin descriptions. I have examined all the sheets Smith saw

(except Waterhouse 201), kindly lent me at Lae by the Brisbane

herbarium and at Cambridge by the Singapore herbarium, and

our own extensive collections from the Solomons. I am able to

distinguish the species on the following diagnostic characters.

C. kajewskii

Twigs and petioles commonly blue-black, or sometimes golden,

glaucous. Leaf blade obovate or ovate, rarely broadly ovate,

4x11 to 9x 19 cm. (ratio 2.7—2.1), secondary veins coarser than in

10 Gardens’ Bulletin, Singapore — XXII (1967)

C. solomonense, raised on both surfaces (felt by finger tips),

60-80° to midrib. Flowers in racemes to 7 cm. long up to 10 per

raceme. Fruit globose 5-6 cm. across at maturity. Very big timber

tree with coarsely fissured bark. Kwara’ae name Ba’ula. Common

throughout Solomons, also in Bismarcks and with two collections

from northwest New Guinea.

C. solomonense

Twigs and petioles always golden. Leaf blade oblong ,7 x 16

to 8x28 cm. (ratio 2.3-3.5). Secondary veins finer than in C.

kajewskii, not or scarcely raised on upper surface (not felt by

finger tips), slightly raised but not prominent below, 80-90° to

midrib. Flowers unknown. Fruits single or paired, on 5 mm.

pedicels and a 10 mm. peduncle; only 2 cm. across at maturity.

Smaller (our collections and pace Kajewski 2469), and much less

common tree than C. kajewskii, with smooth, slightly fissured

bark. When bark is cut superficial phellem comes away to reveal

red phelloderm and its own inner surface which is yellow (BSIP

2580, 2791); no other Solomons Calophyllum does this. Branches

more or less pendent with leaf bases overlapping, like /mocarpus

fagiferus, (BSIP 2580). Kwara’ae confused by this tree and call

it Gwaragwaro (C. vitiense) or Oleole (C. paludosum).

The leaf differences between these two species are subtle and

I am unable to place one sterile collection (BSIP 1940) in one

species or the other, and the 20 ft. tall young tree BSIP 1925

from under C. kajewskii BSIP 1921 has very solomonense-like

leaves.

In passing I observe that Smith apparently made a mistake

when he described C. solomonense. He says that on Brass 3447

‘the fruits appear to be solitary in the leaf axils, in the type

(Kejewski 2469) . . . (they) are . . . on short racemes’ whereas

in the Brisbane sheets it is the other way round, the fruits are

solitary in the type and racemose in the Brass collection.

Collections examined (BSIP series unless otherwise stated)

Calophyllum kajewskii

SOLOMONS

Bougainville Tonelai Hbr. NGF 13029 Buin Kajewksi 2024 type.

Mono 4178

Choiseul 5268a

Rob Roy 5363

New Georgia Islands Baga 1290, 1321, 3065, 4207 Gizo 5610

Kolombangara 818, 1407, 4092 Rendova 1899, 1921, 1925

New Georgia 1941, 3111 Gatukai 318, 1248b.

Santa Ysabel 2722, 2738, Brass 3447

Guadalcanal 2790

Malaita 3415, 3846

San Cristobal 4213, 4345

Whitmore — Solomon Islands’ Plants Ao 11

BISMARCKS

New Britain NGF 24, 1827, 7007, 10914, 10922, 17048

New Ireland NGF 12352

Umbo NGF 9623

NEW GUINEA

NW. Hollandia Res. Tam. BW 764, 1641

Calophyllum solomonense

SOLOMONS

Santa Ysabel 2580, Brass 3447

Guadalcanal 2794, 3317, Kajewski 2469 type

C. kajewskii or solomonense

New Georgia 1940

It remains to demonstrate that these recently described

Solomons’ species are distinct from the earlier described big-

leafed Calophyllums of the Bismarks and New Guinea. None of

the latter is known at all well. Each was described from only a

few collections and in some cases flowers or fruits or both were

not known. Recent collections at Lae are sterile. Nevertheless

there are distinctive features for each specis, which I enumerate

below and which enable me to place the collections at Lae with

some confidence and to consider them different from C. kajewskii

and C. solomonense, as well as from each other.

C. peekelii Lauterbach, Bot. Jahrb. 58, 11, 1923.

Cited specimens: BISMARCKS: New Ireland: Namatanai,

Peekel 132, fruits (but lost); Namarodu, Peekel 781, flowers. NEW

GUINEA: Kei Islands: Warburg 20041, sterile.

To these I add: BISMARCKS: Admiralty Islands: Los Negros

Isl. NGF 536, sterile; St. Matthias Group, Mussau Isl. W 293

(A. C. Richardson), sterile.

The fruit shape and size are only known from Peekel’s manu-

script flora of the Bismarcks p. 1233.

Diagnostic characters. Leaf blade 32 x 11 cm.; flowers big, 2 ‘Cire.

across; fruit big, oval almost round, 3.6x4.7 to 4.7x5.7 cm.

(once 8.5 x 7.0 cm.).

Two sterile collections from the Papua Gulf, Kikori R.

NGF 4551, and Oriomo R. NGF 10410, are of very similar leaf

and provisionally can go here too, although fertile collections are

of course essential for final judgement, as is true also of the Kei

Islands collections (above). The field description of NGF 10410

says ‘fruit axillary, globular, somewhat pointed at apex 3 in. diam’

and this is strong circumstantial evidence that the species is

C. peekelii as such big fruits are unique. Both collections are of

a swamp tree with stilt roots. Mr. Womersley tells me this species

is locally common.

12 Gardens’ Bulletin, Singapore — XXII (1967)

C. macrophyllum Scheffer Nat. Tijd. N.I. 32 (2) 405, 1873.

Lauterbach (loc. cit.) says this is probably identical with

C. peekelii. | have seen the type at Leiden (Teysmann, s.n., Herb.

Bog. 7574, Isl. Gebeh), a single huge lanceolate leaf 45 x 10 cm.

quite unlike that of any other Calophyllum I have ever seen,

and certainly difffferent from C. peekelii.

C. euryphyllum Lauterbach, loc. cit. 14

Cited specimens. NEW GUINEA: Sepik River at Lagerberg.

Hollrung 761, sterile.

To this I add NEW GUINEA: Sterile: Sepik River nr. Yellow

River NGF 3920; Sekoli nr. Hollandia BW 8141; Oregon Trail

BW 12032; Meos Waar Isl. 1255; nr. Manokwari BW 4352.

Flowering: Hollandia BW 4797. Fruiting: Hollandia BW 4827.

MOLUCCAS: Sterile: Morotai, Tobelo, Totodokoe bb. 33.771.

Diagnostic characters. Leaf blade broadly ovate, 8x11 to

10 x 16 cm., rounded at base and apex; secondary veins almost

invisible on upper surface, faint and rounded not angular on lower

surface. Flowers (in bud) 1—3—4, in 5 cm. racemes, clustered in

2-3 s in leaf axils; pedicel 1 cm. Fruits globular 28 mm. across.

A species of northern New Guinea;

C. sil Lauterbach loc. cit. 14

Cited specimen. NEW GUINEA SW: South coast by Gelieb

village: Branderhorst 179 fruits, TYPE.

No material seen.

Diagnostic characters. Leaf blade obovate, 3.57 to 5x10 cm.

Few flowered 5 cm. inflorescence. Fruit spherical 10-12 mm..

unripe.

B. Other Species

1. C. vitiense Turrill J. Linn. Soc. Bot. 43, 17, 1915.

A big timber tree described from Fiji which reaches its northern

limit in the Solomons. This species is characterised by its long,

slender, blue-black petiole, lanceolate leaf, racemose inflorescence,

usually with peduncle to 5 cm. and pedicels 2 cm., and ellipsoidal

seed 2.5 cm. long.

Collections seen:

SOLOMONS (BSIP series unless stated otherwise)

Bougainville NGF 2873, Kajewski 2020

‘Choiseul 5271

New Georgia Islands Baga 1377, 2812 Kolombangara 836,

1410, 2099 Rendova 1880 1911 New Georgia 1977, 2154

Vangunu 429, 899, 967, 1260 Gatukai 1249

Santa Ysabel 2186, 2420

Guadalcanal 648, 1120, 1123, Kajewski 2657

Whitmore — Solomon Islands’ Plants 13

SANTA CRUZ

Tevai 1570

Vanikoro 1596, 1706

2. C. cerasiferum, C. paludosum, C. soulattri.

These are the small-tree Calophyllums of the Solomons. They

seldom attain 5 ft. girth and are usually only 2-3 ft. They are

more easily distinguished in forest than herbarium and some

collections I cannot place. In the forest the Kwara’ae can uncan-

nily tell Kaumanu (C. cerasiferum) from Oleole (the others)

although the species often grow together. I have not been able

fully to analyse the bole and bark characters they use.

All three species have small, usually umbellate, inflorescences,

often much reduced in the first two; flower buds are tiny (2-3

mm.), and peduncles slender; fruits are small, globose to subellip-

soidal, and do not exceed 15 mm. long. The leaves are chartaceous

and the secondary veins close.

The species have a wide habitat range from the lowlands to

mountain ridges, and from swamps to dry land. I am unable to

correlate this physiological diversity with morphological variation.

C. kajewskii has a similar range and is equally indivisible; physio-

logical evolution has not yet been matched by morphological.

C. cerasiferum Vesque Epharmosis 2, 10, 1889.

Diagnostic characters. Bole with flying buttresses; exudate

from cut bark clear golden yellow (except BSIP 2388 — opaque

white). Leaves small, ovate-elliptic, 5x2.5 to 10x4 cm. (ratio

2 to 2.5) with midrib slightly raised and broadly channelled on

upper surface and rather clearly cut raised veins prominent on

both surfaces, distant, 25-30 per em. Sometimes in high mountains.

Kwara’ae name: Kaumanu.

The Solomons’ collections are a reasonable match for C.

cerasiferum of Fiji in leaf shape, size and details of venation. The

type, Seemann 49, (isotpye at K!) has most but not all the midribs

raised and channelled above. In both Fijian and Solomons’ material

the inflorescence may be a short raceme with the flowers clustered

at two points along the main axis (Vaisewa 18, BSIP 1497). The

vernacular name in Fiji is Damanu, cognate with the Kwara’ae

name”*.

Closely related to C. congestiflorum A. C. Smith and C.

pauciflorum A. C. Smith of highland New Guinea which, however,

have consistently smaller and rounder leaves with the midrib

above always sunken rather than raised on the upper surface,

and the secondary veins more distant and less clear-cut.

*Dr. B. C. Stone tells me this name and cognate versions are wide

spread through the Pacific for Calophyllum species, e.g. Kamani Kamanu

(Haiwaii), Tamanu (Samoa, Tahiti).

14 Gardens’ Bulletin, Singapore — X XII (1967)

Collections seen, and habitat

SOLOMONS (BSIP series)

Choiseul 5278 (ridgetop 50 ft.) 5399 (hillside 100 ft)

New Georgia Islands Kolombangara 852 (lowland), 2100 (ridge-

top 2700 ft.), New Georgia 2013 (swamp), Vangunu 1207

(crater rim 2400 ft.)

Santa Ysabel 2388 (summit ridge, Mt. Sasari 3600 ft.)

Malaita 70 (littoral swamp)

FIJI Seemann 49 (type), Horne 732 Vaiwesa 18, Bola 11.

In addition the following collections appear intermediate with C.

paludosum:

New Georgia Islands Baga 2837 (lowland), 2912 (valley bottom,

lowland).

Malaita 70 (littoral swamp-forest), 3458 (periodically flooded

lowland).

C. paludosum C. T. White J. Arn. Arb. 31, 98-9, 1950.

Diagnostic characters. Buttresses absent. Exudate from cut bark

opaque cream-yellow (except NGF 12351-clear). Leaves ovate-

lanceolate 7x2 to 10x3 cm. (ratio 3.5 to 3.3), veins clear-cut,

visible on both surfaces close, c. 40 per cm. (50 on BSIP 316),

drying brown. Kwara’ae names: Oleole (Kwai dialect) Kaumanu-

bala (Auki dialect). Distinguished in herbarium from C. cerasiferum

with difficulty, the leaves are on the whole narrower, the veins

closer, less clearly cut and less prominent; and it has not yet

been found in high mountains.

Clearly distinct (pace C. T. White) from C. pulcherrimum Wall.,

the sheets of which at Singapore (labelled by Henderson & Wyatt

Smith, see Gard. Bull. Sing., 15, 1956), all hove smaller, more

ovate leaves with more distant veins, much more delicate twigs

and long racemose inflorescences with bigger flowers, (isotype,

Cuming. L077, 1, abe:

C. neo-ebudicum Guill. is also well distinguished by its open

racemose inflorescences 6 cm. long and big flowers 10 mm. across,

although in leaf it is identical, (isotype, Kajewski 705,! at K).

Kajewski 642 from Santa Cruz Islands, Vanikoro has a small

raceme 3 cm. long, and flower buds of intermediate size: in

morphology as in locality it is intermediate.

Collections seen, and habitat:

BISMARCKS

Mussau A. C. Richardson W 294 (over coral-limstone)

New Ireland NGF 12351 (well drained ground, sea-level).

SOLOMONS (BSIP series)

Shortland 5746 (hillside 100 ft.)

Whitmore — Solomon Islands’ Plants 15

Choiseul 5278 (ridgetop SO ft.)

Rob Roy 5340 (ridgetop 100 ft.)

New Georgia Islands Baga 1330 (lowland ridgetop) Kolomban-

gara 1497 (ridgetop 700 ft.) 2101 (ridgetop 2700 ft.) New

Georgia 192 (swamp) 4759 (hillside 240 ft.) Vangunu 944

(ridgetop) Gatukai 316 (coastal swamp)

Santa Ysabel 2769 (ultrabasic ridge)

Malaita 3469 (ridge 400 ft.), 3549 (lowland hillside)

San Cristobal 4373 (narrow rocky ridge 1500 ft)

C. soulattri Burm. Fl. Ind. 2, 121, 1768; Henderson & Wyatt-

south Gard.” Bull, ‘Sing., 25, 319, 1956.

C. kiong Lauterbach et Schumann. FI. Deutsch. Schutzgeb.

Siidsee 450, 1901; Ltb. Bot. Jahrb. 58, 11, 1923.

Mr. J. S. Womersley has long suspected that C. kiong is the

same as C. soulattri. C. kiong was described from collections made

on Sattelberg 20-30 miles north east of Lae. The locus has un-

fortunately not been revisited despite its relative accessibility,

but there isa common Calophyllum around the Huon gulf, well

represented in Lae herbarium. It is common in the foothills of

the ranges immediately to the north of Lae which adjoin Sattel-

berg. It fits the description of C. kiong, and matches one of our

Solomons’ species, notably in leaf shape, size and venation and

the umbelliform inflorescences. | have examined the sheets at

Singapore determined as C. soulattri by Henderson and Wyatt-

Smith and find them a good match with the species round Lae;

inter alia umbelliform inflorescences are uncommon in Calophy-

llum. The main difference is that the New Guinea-Solomons’

collections sometimes have smaller leaves; this cannot, I consider,

be maintained as a specific difference, especially as there is a

continuous range of variation in leaf size. I therefore propose to

reduce C. kiong to synonymy with C. soulattri.

Diagnostic characters. Buttresses absent. Exudate from cut

bark, opaque, cream-yellow. Big ovate-oblong leaf blade, broadest

near base, 9x4 to 20x7 cm. or rarely 28 x9 cm. (ratio 2.25—

2.7-3.1); drying grey-green; veins faint, very close together. c. 50

(40) per cm. Kwara’ae names: Oleole (Kwai dialect), Kaumanubala

(Auki dialect). Sometimes confusable with C. paludosum; distinct

in the bigger leaves with closer, fainter veins, drying pale grey-

green rather than brown. |

Collections seen, and habitat:

NEW GUINEA

West

Tami R: BW 2712

Mt. Krabo: BW 10752

Gardens’ Bulletin, Singapore — XXII (1967)

East

Lae environs NGF 3293 (coastal), 11919 (Atzera

Range).

Bulolo (300-400 ft.) NGF 7543, 10144, 12239, 17025

Womersley s.n., 5/6/61.

Wau NGF 1463 (ridge 3,500 ft.)

Papua

Oriomo R. NGF 2723, 13196.

Sogeri Plateau (2,000 ft.) Schodde 3131, NGF 2829.

SOLOMONS (BSIP series unless indicated).

Choiseul 3959 (ridge 200 ft.), 5399 (hillside 100 ft.).

New Georgia Islands Baga 1360 (swamp) Gizo 5602 (ridge 200

ft.) Kolombangara 817 (lowland plain) 1485 (ridge 100 ft.)

Rendova 1856, 1857 (ridge 300 ft.) 1912 (ridge 150 ft.) New

Georgia 2041 (riverbank) 2520, 2521, 2523, 2524 (ridge 600—

750 ft.), Waterhouse 309 (damp country).

Santa Ysabel 2703 (ridge).

Guadalcanal 3388 (ridge 200 ft.), 3833 (low ridge).

3. Calophyllum sp. BSIP 424, from New Georgia Islands,

Vangunu. A small tree of lowland forest, unique and distinctive

in its tiny lanceolate leaves only 3 cm. long. I demur from

describing it as a new species with only one collection and the

flowers unknown. [But see the following paragraph which was

added in proof.|

IV Guttiferae, Calophyllum sp. BSIP 424

The note that this represents a new species was based on the

BSIP sheet. In sorting through the set now deposited at SING I

have just noticed that the sheet there of BSIP 424 has much

bigger leaves. In fact the collection is none other than Calophyllum

cerasiferum.

VY. Myrtaceae

New combinations in Eugenia

_ I take the broad view shared by many other botanists that

Syzygium is not generically distinct from Eugenia and make

the following transfers.

Eugenia cincta (Merr. et Perry) Whitmore comb. noy.

Type: Brass 3344.

Syzygium cinctum (Merr. et Perry)

J. Arn. Arb. 23, 272, 1942, basionym.

Eugenia myriadena (Merr. et Perry) Whitmore comb. nov.

Type: Kajewski 2713.

Syzygium myriadenum Merr. et Perry

J. Arn. Arb. 23, 293, 1942, basionym.

Whitmore — Solomon Islands’ Plants | tT

Eugenia onesima (Merr. et Perry) Whitmore comb. nov.

Type: Kajewski 2043. .

Syzygium onesimum Merr. et Perry

J. Arn. Arb. 23, 296, 1942, basionym.

VI. Sapindaceae

A. Pometia J. et G. Forster

The abundant collections we now have of this common

Solomons’ timber tree fully support Jacobs’ contention (Rein-

wardtia 6, 140, 1962) that in the Solomons Pometia is taxonomi-

cally homogeneous. The fertile and some sterile collections all

run down in his key to forma pinnata. Some sterile collections

have a glabrous midrib above and cannot be keyed out, namely

BSIP 5913, 5916, 5918.

The New Guinea foresters, notwithstanding Jacobs’ treatment,

still find it useful to distinguish three species of Pometia (see van

Royen, 1964, ‘Manual of the Forest Trees of Papua & New Guinea’

2, 35-42) which they distinguish on a number of morphological

characters and which differ in their ecology and bole form (hence

commercial value).

On their criteria all the Solomons’ collections are P. pinnata

Forst. except one BSIP 2694, from Santa Ysabel, Tatamba, which

is P. tomentosa T. et B.

I propose that in the Solomons we follow the New Guinea

foresters for this genus, which contains one of our most important

timber species. The group seems comparable to the Buchanania

arborescens group, a widespread polymorphic species with various-

ly distinct local forms; rather than to the ochlospecies Schizomeria

serrata, which cannot be taxonomically subdivided.

Collections examined (all BSIP series).

SOLOMONS

Shortland 5913, 5914, 5915, 5916, 5918, 5920, 5924, 5925, 5927,

5928.

New Georgia Islands Baga 2882, Gizo 5608, Kolombangara 804.

1483.

Choiseul 5667.

Rob Roy 5416.

Santa Ysabel 2694, 3626.

Guadalcanal 1073, 2775.

Malaita 3456.

San Cristobal 4260, 4324.

SANTA CRUZ

Vanikoro 1722 (with edible fruit but forma pinnata nevertheless).

B._ Tristiropsis Radlikofer

Tristiropsis acutangula Radlk. Sitz. Ber. K. (Bayer.) Ak. Wiss.

M.-Ph. KI. (Cl.) Miinch. 20, 248, 1890: listed also in Durand.

Ind. Gen. 1887.

18 Gardens’ Bulletin, Singapore — XXII (1967)

IT. obtusangula Radlk. loc cit.

T. dentata Radlk. Denkschr. K. Ak. Wiss. M.-N. Kl. Wien 89,

572.. 18ts.

This note is prepared after consultation with Dr. Leenhouts

who is revising the whole family Sapindaceae, and is published

now to provide a sound name for an important Solomons’ species.

The nine or so species described in this genus are very difficult

to tell apart and have very slight differences. From the Solomons

are I’. acutangula Radlk. based on a fruiting collection, Guppy

272, from Oima (should be Oema) in the Shortland Islands: and

T. dentata Radlk. based on two sterile collections, one a juvenile,

from Popoco, a small island and village in Kieta Bay, Bougainville,

about fifty miles north of Oema.

We have found a Tristiropsis which is a common tree of the

coral-strand —on Baga (BSIP 4211, 4417), where considerable

numbers have been felled for timber, on Kolombangara in the New

Georgia Islands, and as a relict tree on the coastal coral of north

central Guadalcanal within Honiara town (BSIP 4421). The species

has a patchy distribution, we have not seen the species anywhere

else despite the abundant and highly distinctive seedlings which

make it easy to spot and we know for certain that it does not

grow in northern Malaita where no one has seen it and there is

no local name for it; tree climber Susui went to Malaita and

round the villages to check this for me.

My collections are a good match for Guppy’s from Oema (type

at Kew!) and can be referred to T. acutangula.

Two other species have been described from the Pacific. T.

obtusangula from the Marianas, differing in the obtuse rather

than acute-angled fruits and 7. dentata from Bougainville differing

in having toothed leaflets. Dr. Leenhouts and I examined at Leiden:

(on loan from Paris) the type of the former. It is a good match

on leaf with my collections and has two flattish detached fruits

which appear simply to be young, not yet fully developed. 7.

dentata is almost certainly merely a juvenile plant, for the seed-

lings of my collections BSIP 4211 have serrate leaves.

It is not possible to maintain these three Pacific Island species

as distinct, and as 7. acutangula has the most complete type

specimen it seems the best name to adopt for this important

Solomons’ timber species.

VII. Verbenaceae

Gmelina Linnaeus

G. moluccana (Bl.) Backer in Heyne, Nutt. Pl. 4, 118, 1917.

G. salomonensis Bakh. J. Arn. Arb. 16, 72-3, 1935.

There is a common Gmelina found through the Solomons. It

is a big tree of disturbed lowland-forest and is well known locally

as it is the best canoe timber in the archipelago. We have made

Whitmore — Solomon Islands’ Plants | 19

many collections of this species. There are two fairly distinct

varieties which have different ranges. Thus:

(1) Leaves densely velvety below, rather thick in texture; in-

florescence axes, stems and petioles densely fulvous tomentose

all over: fruit sometimes cylindrical; central and eastern Solomons.

Collections seen (all BSIP series unless indicated):

Santa Ysable 2487, 4072, 2301 Brass 3309.

Guadalcanal 59*, 649.

Malaita 3501.

San Cristobal 4255; Brass 2860*.

(2) Leaves glabrous below except for a few hairs on veins,

rather thin in texture; inflorescence axes, stems and petioles partly

glabrous partly fulvous tomentose but thinly so; fruit always coni-

cal; western Solomons (except 257).

Collections seen (all BSIP series):

San Cristobal 257*.

Shortlands Fauro 3949, 5708.

New Georgia Islands: Baga 1870, 5573 Gizo 5603 Kolombangara

819, 851. Rendova 1854 New Georgia 169*, 3709.

The starred numbers are at BRIS and L; the others (i.e. the

recent part of the BSIP series) at K, L, LAE, US, and SING.

The hairiness and texture of the leaves apparently does not

depend on their maturity, all leaves on every collection are similar

to one another. Nevertheless I prefer not to give these differences

specific status as in their essential parts these taxa are the same.

The hairy leaved eastern form is a good match for G. moluccana

(BI.) Backer, abundantly represented at Leiden and Lae from

New Britain, all over New Guinea and the Moluccas, much of

which was determined by Moldenke, and including Teysmann

1859, 5031 from Ambon cited by Backer (loc. cit.). All the sheets

seen have coriaceous leaves, most of them velvety hairy but a

few glabrous or glabrescent below (e.g. NGF 1854, 4580, 5870,

8213). There is none of the other form is Lae or Leiden. As with

the Solomons’ material there is no suggestion that texture or

degree of hairines changes much on ageing.

G. salomonensis Bakh. was described on the basis of a single

collection, Brass 3309, from Santa Ysabel, Tiratona. Bakhuizen

stated that it is intermediate and possibly a hybrid between G.

moluccana and G. macrophylla (R. Br.) Benth., which is properly

called G. dalrympleana (F. Muell.) H. J. Lam (Verb. Malay. Arch.

223, 1919). I have examined an isotype at Leiden which fits the

20 Gardens’ Bulletin, Singapore — XXII (1967)

hairy-leaved typical G. moluccana of the eastern Solomons.

although the leaves are rather larger, slightly thinner and less hairy

than is usual.

White (J. Arn. Arb. 21, 113, 1950) annotating Walker’s Solo-

mons’ collections also noted how close G. salomonensis is to G.

moluccana. On the other hand G. dalrympleana in Moldenke’s

sense is a very distinct entity as I show below. It has not been

found yet east of mainland New Guinea and none of the Solomons’

collections come near to it, including, in my opinion, the type of

G. salomonensis.

In my opinion G. salomonensis must be considered synonymous

with G. moluccana, which occurs in the archipelago in its typical

form in the eastern islands and in a glabrous form in the western

islands. The glabrous form may be Moldenke’s G. salomonensis

forma glabrescens (Résumé of Verbenaceae etc. 204, 1959) from

Bougainville of which I have been unable to trace description or

material.

Gmelina in New Guinea

The next question is, what is G. dalrympleana? | have seen

material of three closely related published species of Gmelina

from New Guinea. They may be distinguished thus: —

Leaves densely velvety hairy below, without conspicuous glands; inflore-

scence axes stout

Flowers sessile, partially covered by large triangular bracts c. 12 mm.

long; inflorescence a simple spike;

... G. sessilis White et Francis in Lane Poole Rep. For. Resources

Terr. Papua & New Guinea 136, 1925 and more fully in Proc. R.

S. Queensland 38, 257-8, 1927.

Flowers stalked not covered, bracts smaller lanceolate; inflorescence

usually branched paniculate; ...... G. moluccana (BI.) Backer

Leaves glabrous and shiny below with a pair of conspicuous glands

at base of midrib; inflorescence axes slender

.. . G. dalrympleana (F. Muell.) H. J. Lam.

The difference between G. sessilis and G. moluccana is slight

and not absolute; for instance the Lae sheet of NGF 10883 has

simple spikes, and the Leiden sheet branching ones, and both

sheets have big foliaceous bracts partially enclosing the flower.

The position is further complicated by G. papuana Bakh. (J. Arn.

Arb. 10, 71, 1949). I have not been able to find the type and only

cited sheet (Papua, Iawarere, 695, 25.xi.25) of this. Clearly as

Bakhuizen himself states and from his description and photograph

it is very close to G. moluccana. These three species are all close

and more or less intergrading; G. moluccana is the first one pub-

lished. G. dalrympleana by contrast appears to be distinct.

The following collections fit these species; a star indicates the

sheet was determined by Moldenke. L: at Leiden, otherwise at Lae.

Whitmore — Solomon Islands’ Plants | 21

G. sessilis:

NEW GUINEA

East

Dobodura NGF 2005*

Aitape NGF 1221.

Papua

Milne Bay NGF 1345*.

BISMARCKS

New Britain Talasea NGF 10883.

G. moluccana:

(numerous sheets at Lae & Leiden, including many seen by

Moldenke and some by Backer).

G. dalrympleana:

NEW GUINEA

West

Merauke: Van Royen 4891.

Salawati Isl. BW 1343.

Jalan Kp. Keliki Anta 250 (L).

Animanharin Anta 179 (L).

Okaba Branderhorst 23 (L) (cited by Backer 1921).

East

Sepik, Green R. NGF 9375*.

Morobe NGF 2922.

Papua

N., between Ambasi & Devatutu Villages Hoog-

land 3405*. |

Robinson Bay 3422.

Darn Isl. Brass 6319 (L).

Tarara Brass 3539 (L).

Lake Daviumbu Brass 7666 (L).

Oroimo R. NGF 10374, Brass 5753 (L).

Normanby Isl. NGF 8680* (L), Brass 25388.

Milne Bay NGF 1298* (L), Brass 21719*,

YA :

Fergusson Isl. Brass 27281.

Modewa Bay Brass 28910.

MOLUCCAS

Aroe Isl. Buwalda 5431*.

QUEENSLAND

Kajewski 146 (L); Mueller s.n. (Cape York) GL).

eer nerfed

Pas: how page eve i

, sabe Dirt p

ce 4s Ciaaeei oe

—— lls ~

‘a © a.

F ,

a >

slat > f' ’ 5 ' pea

. sv” 4d ‘ie pet

at le | acon vel | balay Cf) £2 vy \

wat EO c ai

2 alutatn. u OF a

si ;

a -

¥ f info > r -

val

ra see

7 ii 4 htt

| 1 P

‘ Tt i+" 3 7 ;

Z ? ‘ ey 7 a

‘ a. ve ey te dy

¢ ewer >» J ij Les tat , ‘

a We 2 ao

i oa F

- ‘ ¢ <a a

yale ee aa AWW? enét

* WA © i» hen

: , <: ;

; iM ; ="

9 PADI Is

a WMULZROCERK

7 iy Gt] AVA; /, Ki

ag? ‘1/7

a te

‘ee ce-uteurete- | a

= >

(ea?

Notes on the Systematy of Solomon Islands’

Plants and some of their New Guinea

Relatives, VIII and IX

F. MARKGRAF

Director, Botanic Gardens, University of Zurich

VIII Apocynaceae

A. Allowoodsonia Markgraf

Allowoodsonia*whitmorei Mef.n.gen. et sp.

Arbor 6-12 m. alta, radicibus tabularibus instructa, cortice

fusco, laevi obtecta. Folia opposita, chartacea, glabra, breviter

(8 mm.) petiolata, elliptica, apice breviter caudata, 13-18 cm.

longa, 5—7 cm. lata; nervi secundarii arcuati, 6-7 in utroque

latere. Inflorescentiae axillares, glabrae, pauciflorae, pedunculo

1-1.5 mm. longo, simplici, in parte superiore bracteis minimis

ornato, a basi ad apicem flore altero post alterum florentes.

Pedicellus gracilis, 1.5 cm. longus, 0.5 mm. latus. Calycis lobi 5,

ovato-lanceolati, usque ad basin liberi, 4 mm. longi, 2 mm. lati,

in marginibus brevissime ciliati, intus in latere tecto uniglandulosi.

Corollae albae tubus extus glaber, 10-12 mm. longus, 3 mm. latus,

a medio ad apicem ad 4 mm. ampliatus, intus longepilosus, fauce

non calloso; lobi 5 sinistrorsum tegentes, lancéolati, paulum

obliqui, margine tecto magis arcuato et basi subauriculato, 15

mm. longi, 4 mm. lati, extus glabri, intus longe-pilosi. Stamina 5,

medio tubo inserta, filamenta brevissima, antherae sagittiformes,

capiti stigmatico agglutinatae, thecae exteriores 4 mm., interiores

] mm. longae, connectivum in apiculum pilosum, 0.5 mm. longum,

faucem attingens prolongatum. Caput. stigmaticum subglobosum,

quinquangulare, apice in appendices 2 erectas, brevissimas, pilosas

attenuatum; stylus 5 mm. longus; ovarium glabrum, biloculare,

apocarpum, | mm. altum, oblongum, disco anulari quinquelobo

circumdatum, ovula in utroque carpello ad 8. Fructus apocarpus,

mericarpia horizontaliter divergentia, cylindrica acuminata, hinc

inde inflata, glabra, coriacea, complura semina gerentia, 12 mm.

crassa, usque ad 28 cm. longa. Semina applanata, oblique oblongo-

lanceolata, 5 cm. longa, 1 cm. lata, glabra, non comata. Embryo

rectus, radicula 6 mm. longa, 1 mm. lata, cotyledones cordato-

lanceolati, 34 mm. longi, 8 mm. lati.

SOLOMONS: Guadalcanal: BSIP 3313, NE at Rere R., c. 3

miles inland, primary forest in valley bottom. Tree H. 20 ft.,

G. 1 ft. Bole straight, bark grey, smooth. Slash: wood soft, white;

* Dedicated 1 to ‘the late Prof. “Dr. R. E. Feisedean Ir. Be ‘St. ‘Tons.

Missouri, who with great success explored the systematics of Apocynaceae.

24 Gardens’ Bulletin, Singapore — XXII (1967)

bark soft. brown, exudate whitish-cream, clear, sticky but free

flowing. Flowers white. Fruits brown, 0.5 in. across, long; two

joined, about 5 in. Fl. 19.xi.63—Z. Lipangeto (holotype of the

genus, in BSIP). |

Shortland: BSIP 5774, SE at Hisie R., valley bottom, well

drained primary forest. Tree, H. 40 ft., G. 3 ft. Bole straight,

buttresses thick, spreading, to 2 ft. Bark surface dark brown,

smooth. Slash: wood hard, yellow, bark soft, brown and yellow.

Exudate opaque, sticky and free flowing. Flowers white. Fruits

brown, 11 in long. Fr. 30.iv.64, Whitmore’s Collectors.

Santa Ysabel: BSIP 2632, Allardyce Hbr, coastal swamp.

Small tree, fruits long and brown. Fr.11.vii.63, J. Sore, W. Masu’u,

Z. Lipangeto.

This genus, belonging to the tribe Echitideae with anthers

enclosed and sticking to the stigma head, represents an Asiatic

counterpart to the tropical-American (and west-African) genus

Malouetia, which has similar fruits and also seeds without a hair

tuft. The large cotyledons call in mind those of Tabernaemon-

tanoideae.

B. Alstonia R. Brown

Alstonia vitiensis Seem. var. whitmorei Mef.n.v.

Arbor usque ad 15 m. alta. Ramuli glabri, teretes. Folia

opposita, glabra; petiolus 2-6 mm. longus, basi intus glandulosus;

lamina tenuiter chartacea, lanceolato-elliptica, apice acuta, basi

cuneata et in petiolum decurrens, 15-25 cm. longa, 5-11 cm.

lata; nervi secundarii ad 16 in utroque latere, conspicui, obliqui,

in ipsum marginem decurrentes. Inflorescentiae foliis vix breviores,

glabrae, multiflorae, e bracteis minimis triangularibus axis prin-

cipalis ramuli oppositi, subpleiochasiales oriundi, bracteis minimis

scariosis ornati; bracteolae duae minimae scariosae calyci appro-

ximatae. Calycis lobi glabri, obtusi, orbiculari-ovati, eglandulosi,

1.5 mm. longi et lati. Corollae albae, odoratissimae, tubus clavi-

formis, 3 mm. longus, infra faucem constrictum 1 mm. latus,

extus glaber, intus infra stamina et in ipsa fauce longe pilosus:

lobi lineares, obtusi, 6 mm. longi, 1 mm. lati, glabri, dextrorsum

tegentes, in basi tegente subauriculati. Antherae breviter ovoideae,

infra faucem subsessiles, 1 mm. longae. Stigma conicum, verru-

colosum, anulo basali angusto circumdatum, antheras attingens.

Stylus 1 mm. longus. Ovarium apocarpum, cylindraceum, glabrum,

1 mm. altum, 0.6 mm. latum, multiovulatum:; discus nullus.

Mericarpia (immatura) ad 40 cm. longa, 3 mm. lata, laevia,

glabra, teretia.

SOLOMONS: Guadalcanal: BSIP 605, Gold Ridge, alt. 2000

ft., ridge forest. Several-stemmed small tree, 30 ft. tall. Flowers

white, highly fragrant. — FI1.13.x.1962 — T. C. Whitmore, (holo-

type of the variety, in BSIP).

Markgraf — Solomon Islands’ Plants | 25

Santa Ysabel: BSIP 2463, nr. Maringe Lagoon, long western

ridge of Mt. Sasari, ridge forest, alt. 2000 ft. Small tree, H. 30 ft..

G. 9 in. Fruits green, unripe, leaves bright green. Fr.28.x.1963

T. C. Whitmore. BSIP 2433, Maringe Lagoon, Molau village.

alt. 2200 ft., secondary forest; ‘tree, H. 40 ft. G. 1 ft. Crown

conical, tiered, the old fruits pendent from every tier; bole

smooth, bark grey; slash: wood yellow, bark coarsely granular:

vivid sulphureous yellow at cambium, paler outwards, with fawn

flecks; no exudate, but cut twigs with sticky white exudate;

flowers yellow in bud, with white lobes when open. — FI.27.x.1963,

T. C. Whitmore.

S. E. Choiseul: BSIP 2985, ultrabasic outcrop across bay from

Ruruvai, sea level, inside edge of mangrove. Tree, H. 15 ft.,

flowers white, scented; white exudate from twigs. FI.27.11.1964,

T. C. Whitmore. BSIP 4015, Easternmost Choiseul, ultrabasic hill

on coast opposite Bembalama Island. Forest with much Casuraina

and thick leaf litter layer. Tree, G. 1 ft. Flowers white. F1.3.111,1964,

T. C. Whitmore.

SANTA CRUZ: Vanikoro: BSIP 1653, E. side Saboe Bay.

ridge forest, alt. c. 500 ft. Tree, H. 50 ft., bole smooth, fawn;

flowers yellow-green in bud; cut twigs exuding white sap. Fl.Liv.

1963, T. C. Whitmore. BSIP 1630, S. coast at Emwa, ridge forest,

alt. c. 1000 ft., soil deep, red sticky clay under a mull-type

humus layer. Small tree to 20 ft., flowers white, fruits green.

F1.8.iv.1963, T. C. Whitmore.

This Solomon’s taxon belongs to the polymorphous Polynesian

group of A. plumosa Lab., mentioned by Monachino in his mono-

graph (Pac. Sc. 3, 143, 1949, point 4, second paragraph), and

especially to his “‘dubious form in the Solomon Islands” (171),

which (172) “in general appearance is markedly like’ his var.

novo-ebudica from the New Hebrides, but also ‘‘to the glabrous-

leaved series from Fiji’’.

C. Carruthersia Seemann

Carruthersia mollis Mef.n.sp.

Frutex alte scandens. Ramuli juveniles teretes, setis brevibus

patulis molliter fusco-velutini. Folia chartacea, supra laxe hirsuta,

subtus molliter fusco-velutina, in marginibus ciliata: petiolus

I-1.5 cm. longus; lamina firme chartacea, ovata, apice acuminata.

basi cordata, ad 9 cm. longa, 5 cm. lata; nervi secundarii utrinsecus

6-8. Inflorescentiae axillares, trichasiales, molliter fusco-velutinas

5 cm. longae et latae, bracteis et bracteolis minimis, supra glabris,

subtus velutinae ornatae. Pedicelli 6-8 mm. longi. Calycis lobi

late ovati, 1 mm. alti et lati, extus breviter setosi, intus glabri

et multiglandulosi. Corollae tubus ruber, 9 mm. longus, 2 mm.

latus, supra calycem paulo ampliatus et inde usque ad faucem

extus breviter setosus, intus longiuscule pilosus; faux pilis longio-

ribus clausus; lobi rosei, dextrorsum tegentes, valde obliqui.

26 Gardens’ Bulletin, Singapore — X XII (1967)

securiformes, pars erecta 7 mm. longa a basi angustiore ad 5

mm. dilatata, pars lateralis obtusa, 3 mm. lata, 3 mm. longa

(cum apice partis erectae, ex qua oritur, 8 mm. longa). Antherae

oblongae, breviter acuminatae, caudis basalibus incurvis, glabrae,

15 mm. longae, 0.5 mm. latae, filamentis 0.5 mm. longis, glabris

1 mm. supra basin tubi insertae. Ovarium glabrum, apocarpum,

carpella subglobosa, 0.3 mm. alta, cum 2 squamis disci obtusis

alternantia. Ovula pauca, biseriata. Stylus 1.5 mm. longus, capite

stigmatico 1.5 mm. longo, fusiformi-incrassato coronatus. Fructus.

desunt.

SOLOMONS: Santa Ysabel: BSIP 2749, Bogotu, forest on

gentle sloping ridge-side west of Parega village; deep, sticky,

orang clay soil overlying ultrabasic rock. Woody climber, flower-

ing at 50 ft., corolla tube red, limbs pink. FI.6.xi.63 T. C. Whit-

more, ( holotype in BSIP).

In most characters corresponding well to C. brassii Merr. et

Perry, and to C. latifolia Gillespie, but both are essentially glabrous,

and have an outside glabrous corolla tube; their filaments are

hairy; they differ slightly too in the measurements of leaves,

flowers and flower parts.

D. Kentrochosia Ltb. et K. Schum. and Kopsia Blume

The vegetative parts and flowers of Kopsia flavida Bl., the fruit

of which is unknown, completely agree with those of Kentrochrosia

monocarpa K. Schum. (cf. Markgraf in Bot. Jahrb. 61,196, 1928).

All the years over no indigenous Kopsia has been discovered in

New Guinea and surrounding countries. Thus it becomes obvious

that both names refer to the same species. As moreover another

species with a minor lateral excrescence of the fruit has now been

found in Borneo (Kopsia lapidilecta v.d. Sleesen in Blumea 10,

137, 1960), the genus Kentrochrosia should best be united with

Kopsia.

E. Ochrosia Jussieu

Ochrosia sciadophylla Mef.n.sp.

Arbor 9-12 m. alta. Ramuli glabri, angulosi. Folia ternatim vel

quaternatim verticillata; petiolus 2-4 cm. longus, intra basin multi-

glandulosus; lamina firme chartacea, glabra, oblongo-elliptica,

apice obtuse acuminata, in petiolum cuneato-decurrens, 12-25

cm. longa, 3.5-6 cm. lata; nervi secundarii arcuati, sed e costa

rectangulariter orti, 18-25 paria, paucis abbreviatis tenuioribus.

interiectis. Inflorescentiae glabrae, axillares, pauciflorae; pedun-

culus communis 2-3 cm. longus, erectus, pedunculi ultimi 2-5

mm. longi floribus subsessilibus glomeratis coronati; bracteae et

bracteolae triangulares minimae. Lobi calycis ovati, obtusi.

glaberrimi, 1.5 mm. longi, 1 mm. lati, intus eglandulosi. Corollae

albae, glabrae, gracilis tubus 2 mm. longus, 1 mm. latus, etiam

Markgraf — Solomon Islands’ Plants ay

intus glaber, lobi glabri, oblongi, 5 mm. longi, | mm. lati, sinis-

trorsum tegentes, obtusi. Antherae infra faucem sessiles, glabrae,

breviter ovatae, | mm. longae. Stigma glabrum, cylindrico-

conicum, collo basali cinctum, antheras vix attingens. Ovarium

apocarpum, biloculare, ovoideum, in utroque loculo biovulatum.

Mericarpia viridia, oblique ovata, 5 cm. longa, 2.5 cm. crassa, in

rostrum apicale | cm. longum recurvum attenuata exocarpium

chartaceum, nitidulum, mesocarpium suberosum, vix evolutum,

endocarpium lignosum, compactum, nonnisi prope marginem in

costulas latiores et inpaucos fibros tenues dissolutum, intus laeve

et nitidulum. Pars basalis laterum carpelli ad 15 mm. longitudinis

et 7 mm. latitudinis concreta; dissepimentum coriaceum, in utroque

eius latere unum semen valde applanatum, oblique ovatum, 2 cm.

longum, 1.5 cm. latum, prope basin ad | cm. erosum, ala 3, mm.

lata cinctum gerens.

SOLOMONS: New Georgia: BSIP 3700, NW at Vaimbu River;

well drained primary forest on hill side, alt. 350 ft. Tree, H. 80 ft.,

G. 4 ft.; bole straight, bark surface light brown, smooth; slash:

wood hard, yellow, bark hard, yellow; white, opaque, sticky, free-

flowing exudate; flowers white, scented; fruits green. FI. and fr.

19.11.1964, Whitmore’s Collectors (holotype in BSIP). BSIP 3182.

NW at Viambu River, hillside 400 ft., well drained primary forest.

Tree, H. 40 ft.. G. 3 ft.; bole straight, buttresses absent, bark

surface dark brown, fissured; slash: wood hard, light brown,

bark hard, fawn inside, brown outside; fruits green, Fr.30.iv.1964,

Cowmeadow’s Collectors. BSIP 3734, N. coast near Hovoro, ridge

top, 564 ft., well drained primary forest. Tree, H. 38 ft., G. 2 ft.;

buttresses thick, steep, spreading, bark surface dark brown,

scaly; slash: wood soft, yellow, bark soft, flecked yellow; flowers

yellow, scented, fruits green. Fr.21.v.1964, Whitmore’s Collectors.

BSIP 2529, Viru R., Viru Hbr., flat land, volcanic rock, alt. 200

ft.. Campnosperma forest, red clay loam. Small understory pole;

G. 2 ft. 13.viii.63, A. W. Cowmeadow & R. Teona. BSIP 1950,

Roviana Lagoon, nr. Nusahope village, small stream in lowland

forest. Tree, H. 30 ft., open, deep crown, flowers white, fruits

green, Fl. & fr.19.1x.1963, T. C. Whitmore.

This taxon belongs to a small group of distantly-nerved species

with cuneate leaf base, like O. ficifolia (Sp. Moore) Mef. in New

Guinea. It is distinguished by its long-petioled and long-bladed,

many-nerved leaves, short inflorescences, by a thick endcarp,

containing only one seed at either side of the false septum.

28 Gardens’ Bulletin, Singapore — X XII (1967)

F. Pagiantha Markgraf

Pagiantha koroana Mef. var. salomonensis Mef.n.yv.

Arbor ad 18 m. alta, radicibus tabularibus instructa, cortice

fusca, squamosa et verrucosa obtecta. Ramuli crassiusculi, glabri,

subquadranguli. Folia opposita, coriacea, glabra, petiolus 5-10

mm. longus, in axilla stipula una cyathiformi 5 mm. longa prae-

ditus; lamina elliptica, obtusa, usque ad 25 cm. longa, 16 cm. lata,

nervi secundarii arcuati, 10-12 in utroque latere. Inflorescentiae

axillares, trichasiales, pedunculi crassiusculi, angulosi, bracteae

et bracteolae triangulares, 3 mm. longae, semiamplectentes, pedi-

celli 5 mm. longi, crassiusculi. Lobi calycis orbiculares, glabri,

minute ciliati intus pluriglandulosi, 6 mm. longi. Corollae albae,

odoratae, tubus crassiusculus, 20 mm. longus, 5 mm. latus, glaber;

lobi in alabastrum globosum, 8 mm. latum inflexi, glabri, sinistror-

sum tegentes, oblique ovati, in basi tegente auriculati, 10 mm.

longi, 7 mm. lati. Stamina in medio tubo subsessilia, antherae

oblongae, 4 mm. longae, | mm. latae, apice breviter apiculatae.

Caput stigmaticum crassum, verrucosum, oblongum, 2 mm. lon-

gum, | mm. latum, apice breviter biapiculatum, antheras non

attingens. Stylus crassus, brevissimus, 0.5 mm. longus. Ovarium

apocarpum, biloculare, 2 mm. altum, 2.5 mm. latum, glabrum,

carnosum, angulosum, multiovulatum. Fructus apocarpi; mericar-

pia (plerumque unum solum maturans) subglobosa, breviter

acuminata, recurvata, fusca, verrucosa, carnosa, usque ad 15 cm.

crassa, pulpa rubra impleta. Semina numerosa, oblonga, longitu-

dinaliter unisulcata et in superficie multirimosa, ad 12 mm. longa

et 6 mm. lata. Testa tenuis, albumen corneum. Embryo rectus,

longitudinalis, sulco oppositus et in latere sulci concavus, radicula

5 mm. longa, | mm. crassa, cotyledones triangulari-ovati, 4 mm.

longi, 3 mm. lati.

SOLOMONS: New Georgia: BSIP 3195, NW at Hehebere R.,

near Hovoro, valley bottom, 300 ft. above sea level, well drained

primary forest. Tree, H. 60 ft. G. 8 ft. Bole straight. Buttresses

thick, steep to 7 ft. Bark surface light brown, scaly and warty.

Slash: wood hard, light brown, bark soft and fawn. Exudate

white, opaque, sticky in free-flowing drops. Flowers white, scented.

Fruits brown, 3 x 1.5 in. ovoid. FI.4.v.64.A.W. Cowmeadow.

(holotype of the variety, in BSIP). BSIP 3129, NW at Vaimbu R..,

well drained primary forest on hillside, alt. 320 ft. Tree, H. 60 ft.,

G. 3 ft. Bole straight. Bark surface light brown, fissured. Slash:

wood hard, light brown, bark hard, fawn. White, opaque, sticky,

free-flowing exudate. Flowers green in bud, unscented. FI. 24.iii.64.

A. W. Cowmeadow. Kolombangara S.W.: BSIP 1193, Tree, up

to 40 ft., overall with widely spreading branches. Flowers white,

sweetly scented. Fruits rarely paired, grey brown. Fl. 26.x1.62,

Womersley and Whitmore.

Markeraf — Solomon Islands’ Plants 29

Santa Ysabel: BSIP 4103, Bogotu Peninsula, near Koloajoa

village. Disturbed forest. Tree, H. 30 ft. Flowers white. Fruits

grey-green, paired. Young fl. and fr. 23.ii1.64, T. C. Whitmore.

BSIP 2183, N.W. coast, Bagi R., 1 mile S. of Allardyce Hbr.

Low-lying, periodically swamped forest near shore. Tree, H. 60

ft.. G. 5 ft. Crown rounded, dense with large leaves. Bole orange-

fawn, coarsely rugose with large lenticels. Low, thick, rounded

buttresses. Slash: wood fine grained, orange, bark granular, fawn.

Copious slightly sticky white, milky latex. Fruits very curious,

some paired, ovate to kidney-shaped, brown warty surface, to

7 in. long. Ultimately dehiscent longitudinally. Seeds in a central

red mass which is probably flora! tissue: the pericarp very very

thick; persistent exuding latex. Fr. 15.x.63, T. C. Whitmore.

The genus Pagiantha is distributed from India to Polynesia,

but in isolated species areas. (cf. Markgraf in Notizbl. Berl.-Dahl

12, 549, 1935 and Bull. Bish. Mus. 141, 129-130, 1936). This new

variety of the Fijian P. koroana differs mainly by being larger in

all vegetative parts, and by pressenting elliptic leaves with rounded

base instead of obovate ones. Moreover, the corolla lobes seem

to remain shorter, and the stigma head does not reach the

anthers. The fruit of the species is here described for the first time.

IX. Gnetaceae

By this collection, our knowledge of the genus.Gnetum in the

Solomon Islands has been considerbly enhanced. As yet it was

unknown that e.g. the climbing: Gnetum latifolium Bl. occurred

there at all. Now its presence is proved by numerous good samples.

It is represented chiefly by the common f. Jatifolium, not by f.

brachypodum Mef. so frequent in the Philippine Islands, and once

by var. laxifrutescens (Elm.) Mgf., whose area previously extended

from the Philippine Islands to the Bismarck archipelago. In this

Solomon Islands collection, there has been found also var.

latifolium f. longipes Mgf., to which, however, no geographical

value can be attributed, as it occurs here and there in the whole

area of the species. |

The numerous samples of the tree Gnetum gnemon L. offer

some interesting features. The typical variety of this species, var.

gnemon, centres in eastern Malesia, from Palawan, Sulu, Celebes

and Sumba eastward, though it is often cultivated in other parts.

At any rate, its area is accompanied by some low-growing and

small-seeded varieties. Four varieties thrive in western Malesia,

from Celebes through Borneo and the Malay Peninsula in to

30 Gardens’ Bulletin, Singapore — X XII (1967)

Assam and Burma; one other, var. silvestre Parl. has the same

range as var. gnemon, from the Philippine Islands and Celebes

through the Moluccas and New Guinea to the Fiji Islands. It

appears from Whitmore’s collections that this variety hybridizes

with var. gnemon in the Solomon Islands. Of course it may do

so elsewhere, but previously we have never had enough material

from a single region to make a comparison.

In order to distinguish the two varieties and the intermediate

specimens, one can find out several graded characters: height

of the plant, length of the fertile part of the inflorescences in

anthesis, the same of female plants with ripe seeds, length of

their internodes, size of ripe seeds, shape of ripe seeds, number

of male flowers in one middle bracteate collar of male plants.

The specimens of the present collection are arranged in a scheme

according to these steps in Table 1 which shows as a first general

result that var. silvestre seems to be the more frequent one;

morever it is often expressively called frequent by the collectors.

As earlier collections have shown too, it proves to be the prevailing

variety in the rain — forests east of New Guinea. None of the

Solomons’ specimens reaches the mean height of true var. gnemon.

Even in those which come nearest to var. gnemon, the characters

taken into consideration hold its lowermost level. The same

predominance of var. silvestre appears also in the intermediate

specimens; most of them incline more to var. silvestre than to

var. gnemon. This trend may be understood in the easiest way

by assuming that var. silvestre, prevailing in quantity, will prevail

also in morphological characters over the interbreeding var.

gnemon. This interpretation of the intermediates as being of hybrid

origin, is favoured too by the fact that the respective characters

appear in them in almost any combination. Only the lengths of

the inflorescences show a certain parallelism to the lengths of

their internodes. Of our scheme, the first three collections, BSIP

1875, 3942, 3448 may still be reckoned under true var. gnemon,

and the last seven under var. silvestre, namely. BSIP 2202, 5319,

2799, 4008, 3435, 5653, 3680.

All the specimens cited in these notes were examined on loan

from the Forest Herbarium, Honiara (BSIP). Replicates are at

K, L, LAE, SING and US; the last two sets are however

incomplete.

TABLE 1

The two common Guetum gnemon varieties in Solomons and their intermediates.

Hybridisation is suspected. See text

5444 2519 1155 2708 5250 826 5635 4079 3061

2 ? 2 2 2 2 7 a 2

2202

©)

seeds

young open fi.

treelet

broken

adult fl. & ripe unripe openfl. ripe ripe ripe adult unripe young

seeds seeds seeds seeds seeds

4.8 shrub 12 treelet under 6 tree treelet tree

4.5

=. 2 = 2.8 = => = =

8 5 3 = 4 broken 4 4

6-7 fl. 2 4-5 3-4 3-4 1-2 2-3 3

ripe 6

16x10 17x8 15x12

Sx8 20x10 15x11 15x10 15x10

pointed pointed pointed = pointed pointed round pointed round

with with

. tip tip

BSIP No. average 1875 3942

v. gnemon

sex O&O mG Q

stage adult open fl. adult

plant ht. m. 8-20 3 9

in flower 2-5 2 =

4 | cm.

= Ee

20 .

» |with ripe 6-8 = 7-8

3 | -seedsicm:

fe)

= internodes |, 6% 1-2 0.5 6-7

cm. 0 2-5

seed size mm. 20-25 = 25 <5

x 10-15

seed shape blunt = blunt

approx. no. 3 80 60 _

fl.

conclusion true v. gnemon

intermediate between v. gnemon and vy. silvestre

3435 5653 3680 average BSIP No.

v. silvestre

oy 2 e S&Q | sex

young ripeseeds fl. adult stage

under tree es) 8-12 plant ht. m.

15 = 25 O° 1.5-2 | in flower

2 23 cm. 5

iS)

= 4 — 2-5 with ripe e

seeds 5

cm. @

oo.

0.1 2-3 1-2 fl. O-1 intenodes | >

ripe 2 mm.

— bili = 8-12 seed size mm.

= round round pointed seed shape

with

tip

25 = = 40 approx. no. of

fl.

true v. silvestre conclusion

os -

- ter

Lae i Wihuene 4 »

viet Jey ee, Oe yasarg

{ "

o t

} a:

+ — - a

j . ‘ ~ ao.

‘ €) ) @) a

‘ . ¢ 7

ie othe eqinan bape ‘“Hpbe ah

>. .8 oe

ze "a

(xs) oat - ot

a a be

wets lids i e

ar ‘

)

. “7

t a T \

ms 7

— ——* ; a a

: !

nr = '

| a yt ; é

' ‘| « )

* ui

be t :& Sxe! -

ne ,

sien ii 1 } ' Z

‘ , ;

= nl atl ;

—+ a ' _ 7 nail

it? al a

1 si baperanen nine \4

> > aes *

Notes on the Systematy of Solomon Islands’ Plants

and some of their New Guinea Relatives, X.

P. VAN ROYEN

Dept. of Forests, Division of Botany, Lae, New Guinea

X. Sapotaceae

The Sapotaceae of the Solomon Islands are in general not very

well known, and material of one of its most interesting genera,

Chelonespermum, was so scrappy that for a long time it remained

a rather doubtful one. In the large collections by T. C. Whitmore

from many parts of the Solomons this genus turns out to be well

represented and seems to be sufficiently distinct to keep it separate

from Burckella.

Apart from the specimens in Chelonespermum, two new species

in Burckella and Palaquium are found, descriptions of which will

be given below. In addition we will add some remarks on

Chelonespermum.

The sheets examined were all at LAE; there are replicates at

_BSIP, K, L, SING and US. The last two series are incomplete.

A. Burckella Pierre

Burckella sorei Royen nsp. Fig. |

Arbor magna. Ramuli subcrassi, foliis, bracteis et flores apice

conferta, dense pallide brunneo-sericei, glabrescentes. Folia apice

ramulorum conferta, pseudo-verticillata, oblongo-elliptica, 10-14

x 4-6 cm., apice obtuse acuminata, basi rotundata vel late cuneata

interdum obliqua, nervi secundarii 17-22 pares, arcuatim conjuncti,

tertiarii reticulati: utrinque glabra. Petiolus 2.2-4 cm. longus,

glaber, supra late canaliculatus. Flores fasciculi; pedicelli 3-6

mm. longi, dense ferrugineo-sericei. Sepala 4, ovata, 2—2.5 x 11.5

mm., subacuta, dorsa subcarinata, extus disperse sericea, apice

ferrugineo-plumulosa, intus glabra. Corolla 8-lobata, glabra sed

apice plumulosa (ultimo glabrata?). Stamina 15, filamenta pilosa,

antherae pilosa apice solis. Ovarium 4-loculare, glabrum. Fructus

obovoideo-ellipsoideum, usque ad 7.5 x 3.8 cm., 1-spermus, glabrus.

Semen usque ad 5 x2.8 cm., cicatrice testam } parte includente

irregulariter bullata.

Tree up to 33 m. with dense crown. Buttresses steep and broad,

up to 2.4 m. Bole fluted. Bark surface dark, dull brown, closely

fissured, in part with elongate, loose scales. Cut inner bark orange-

brown or pink, with copious white exudate. Wood orange-yellow

or pink-brown. Branchlets relatively slender, densely pale brown,

sericeous, ultimately glabrous. Leaves conferted at tip of flushes,

34 Gardens’ Bulletin, Singapore — X XII (1967)

Fig. 1. Burckella sorei.

Whitmore BSIP 2771. A Habit, B Calyx. C Corolla. All sizes in mm.

Van Royen — Solomon Islands’ Plants os)

sometimes seemingly whorled, limb oblong-elliptic, 10-14 by 4-6

cm., short obtusely acuminate at tip, rounded to broadly cuneate

at base and sometimes asymmetric; midrib narrowly crested above,

rounded below, lateral nerves 17-22 on either side of midrib,

inconspicuous above, distinct below, archingly joined, nervation

widely reticulate, hardly visible above, conspicuous below; glabrous

on either side; Petiole 2.2-4 cm., broadly grooved in the apical

part, flat in lower parts, glabrous. Flowers closely bundled at tip

of branchlets; pedicels 3-6 mm. long, densely ferruginous, silky.

Sepals 4, ovate, 2-2.5 by 1-1.5 mm., subacute, slightly crested on

outside, scattered silky hairy on inside, with a plumule of rusty

hairs at tip, glabrous on inside, the 2 inner sepals with membranous

margins. Corolla-tube c. 1 mm. long, glabrous, lobes 8, ovate,

2-3 by 1-1.2 mm., obtuse, plumose at tip (ultimately glabrous?).

Stamens 15, c. 1 mm. long, filaments hairy, anthers hairy at tip

only. Ovary 4-celled, glabrous, style c. 1 mm. long, glabrous.

Fruits obovoid-ellipsoid, up to 7.5 by 3.8 cm., subacute at tip,

glabrous, 1-seeded; pericarp fleshy. Seed up to 5 by 2.8 cm., scar

covering # of entire seed, roughly knobbly.

SOLOMONS: Guadalcanal: BSIP 2771, Rere R. c. 3 miles

inland, T. C. Whitmore, Nov., flowers, Holotype in SING.

Santa Ysabel: BSIP 2477, Garona, a few miles W. of Maringe

Lagoon, T. C. Whitmore Oct., fruits.

A lowland rain-forest species.

The flowers are known in the bud stage only and details parti-

cularly those of the stamens and ovary, are incomplete.

This species, which by its 8 corolla-lobes belongs to Burckella,

could not be matched against any of the known Burckella species,

not even against the very variable Burckella obovata Pierre, from

which it differs by the larger number of lateral nerves coupled

with smaller leaves. The calyx of B. sorei is much smaller than

that of B. obovata, but the flowers being young, this detail might

not be reliable.

The species is named in honour of Mr. J. Sore, a tree-climber

working for the Forestry Department B.S.I.P., and living on

Guadalcanal at Bambasu village in Longgulenggu near the Rere

R. where the type material was collected.

Kwara’ae names: Faigona, Kona.

B. Chelonespermum Hemsley

This genus was described by Hemsley (Ann. Bot. 6, 205, 1892)

based mainly on the peculiar seeds that separate it immediately

from the closely related genus Burckella. Of the four species

described only C. majus had leaves (though fragmentary) and

fruits, while C. fijiense was based on leaves and seeds only. The

Other two species, C. minus and C. unguiculatum are based on

36 Gardens’ Bulletin, Singapore — XXII (1967)

seeds only. In 1959 I described a new species (Nova Guinea N.S.

10, 140) C. banikiense, from Banika Island, in the Russells, of

which at that time no fruits were known. The reason to place this

species in Chelonespermum and implicitly the maintaining of that

genus against Burckella was the number of corolla-lobes in

Chelonespermum. In Burckella this is 8, in Chelonespermum 4,

rarely 5. Additional to this, though the differences are overlapping,

are the number of cells in the ovary, viz. 1-4 in Chelonespermum,

(3-) 4-8 in Burckella.

Though these differences are small, they are not smaller than

is recognised in many other groups in the Sapotaceae, for instance

Madhuca and Burckella 4, Palaquium 6 sepals, though in both

Madhuca, Burckella and Palaquium occasionally 5 sepals are

found.

It is doubtful whether the fruits, and: more in particular the

seeds ultimately can be used as sufficiently different to keep the

genera apart, emphasizing only their close relationship. In Burckella

the scar, representing that part of the seed with which it is attached

to the placenta, is usually smooth, without any appendages, while

those of Chelonespermum have a most intricate system of crests,

knobs and protuberances. However, in Burckella magusun, des-

cribed by myself in 1959 (1.c.) the scar is very irregularly grooved

and knobbly, and the same applies to Burckella sorei described

in this paper.

In C. banikiense, material of which turned up several times in

Whitmore’s collection, the seeds show the same type of scar as in

the two Burckella’s mentioned in the previous paragraph. To

complete the description given in 1959, that of the fruits and seeds

follows here.

C. banikiense. Fig. 2. Amended description. Fruits globose,

narrowed at base, up to 3.5 cm. across, pericarp fleshy-woody,

glabrous; 1-seeded. Testa of seed narrow, brown, glossy, scar

covering about 3 of entire seed, knobbly; seeds ellipsoid, c. 2 by

1.2 by 1 cm. obtuse at either end.

Fig. 2. Chelonespermum banikiense.

Whitmore BSIP 1512. A Fruit. B Seed. All sizes in mm.

Van Royen — Solomon Islands’ Plants 37

SOLOMONS: New Georgia Islands: Kolombangara: BSIP

1512, W. coast, Sandfly Hbr. T. C. Whitmore fruits, Febr. Baga

BSIP 2849, Whitmore’s Collectors, fruits, Jan. BSIP 3008,

Whitmore’s Collectors, fruits, Febr.

Shortland: BSIP 5876, N. at 1 mile west of Kupola R.,

Whitmore’s Collectors, flowers, May.

Kwara’ae names: Faigona, Kona.

C. Palaquium Blanco

Palaquium masuui Royen nsp. Fig 3.

Arbor magna. Ramuli ferrugineo-sericei, glabrescentes; stipulae

lanceolato-subulatae, acutae, dense ferrugineo-pilosae. Folia apice

ramulorum subconferta, obovata, 6-15 x4-7 cm., apice obtuse-

acuminata, basi cuneata, supra glabra basi excepta, subtus glabra;

nervi secundarii utroque latere 11-15, sursum evanescentes, ter-

tiarii transversi. Petiolus 2-4 cm. longus. Pedicellus 2-2.8 cm.

longus, dense ferrugineo-pilosus. Sepala 6, extus ferrugineo-pilosus,

intus pilosus in parte apicali solis. Corolla intus glabra, tubus

extus ferrugineo-sericea, lobi sericei in linea mediana. Stamina

c. 15, filamenta glabra, antherae extus ferrugineo-pilosum. Ovario

6-loculare, hemi-globosum vel late ovoideum, ferrugineo-sericeum,

stylus glaber. Fructus ignotus.

Tree up to 21 m., d.b.h. 45 cm. Buttresses up to 2.4 m. Bark

surface dark brown, smooth, with close, fine superficial fissures.

Cut inner bark red or orange-brown, fibrous, with sticky white

exudate. Sapwood pink or ordnge-fawn, heartwood dark dull

brown or red. Branchlets rusty sericeous, ultimately glabrous.

Stipules lanceolate-subulate, 2-3 mm. long, densely rusty hairy.

Leaves (sub-) crowded at tip of branchlets, limb obovate, 6-14

by 4-7 cm., obtusely acuminate at tip, cuneate at base and decur-

rent along upper side of petiole, midrib shallowly grooved above

and longitudinally crested, prominent and rounded below, lateral

nerves 7—9 on either side of midrib, diminishing until inconspicuous

along margin, prominulous and grooved above, prominent below,

nervation transverse, slender; glabrous except above, greyish hairy

at the extreme base. Petiole 2-4 cm. long, grooved above, rounded

below but crested in basal part, densely rusty or greyish silky,

becoming glabrous. Flowers in 2-4 flowered, axillary clusters or

solitary; pedicels angular, 22.8 cm. long, densely appressed, rusty

hairy. Sepals 6, outer 3 ovate-triangular, 3-4 by 3-3.5 mm., obtuse

or obtusely acuminate, inner 3 more lanceolate-ovate, 3.5-5 by

3-3.5 mm., rounded, crested, all sepals appressed rusty hairy on

Outside, sparsely hairy on inside in apical part only, inner sepals

plumose at tip and glabrous along the membranous margins.

38 Gardens’ Bulletin, Singapore — XXII (1967)

Fig. 3. Palaquium masuui.

Whitmore BSIP 1638. A Habit. B Corolla outside. C Ovary

with part of corolla. D Outer sepal. E Inner sepal. F Anther.

All sizes in mm.

Van Royen — Solomon Islands’ Plants xo

Corolla-tube 2-3 mm. long, lobes 6, elliptic or oblong-elliptic,

5-6 by 2.5-3 mm., rounded or truncate at tip; corolla on outside

appressed rusty hairy on tube and along part of midrib or corolla-

lobes, glabrous on inside. Stamens c. 15, c. 6 mm. long, filaments

c. 4.5 mm. long, glabrous, anthers c. 2.5 mm., connective prolonged,

truncate or bifid at tip, appressed rusty hairy on outside. Ovary

6-celled, hemiglobose or broadly ovoid, 2-3.5 mm. across,

appressedly rusty hairy, style up to 15 mm. long, glabrous. Fruits

not known.

SANTA CRUZ: Vanikoro: BSIP 1638, E. side of Saboe Bay,

T. C. Whitmore April, holotype in SING. BSIP 1814, ridge near

Peou, T. C. Whitmore, May.

A species of primary or old secondary lowland rain forest.

This species is related to Palaquium neo-ebudicum Guillaumin,

from the New Hebrides. It differs from that species by its appressed

pubescence of branchlets, petioles and pedicels, by its smaller

number of lateral nerves, by the pubescence of the sepals which

are hairy on outside and only so on inside in the apical part, while

those of P. neo-ebudicum are glabrous on the outside. Also the

corolla has a pubescence on outside, that of P. neo-ebudicum being

glabrous.

This species is named in honour of Mr. W. Masu’u, a tree-

climber from Totonga Village, Longgolenggu, Guadalcanal, who

worked for the Forestry Department, B.S.I.P., between 1962 and

1964.

ana a aaitine &

: aie) a tee vi bestd wT OA

cae ts eo aly b gnol Tuo tt oF ty olgie yi

> yt '

i ;

“ Pi ;

i , 2. ” ry VW ms Met ’ pay

« ed . fe; #7 m™ ‘ie et ya ne 4 al

bal 4

Taeoes aaa 1 pte 19 a ae Pte ii

1297 ol on Hat baielerit try I

sire bb

a) asic hw nat Yo Railay | “dh me

— i sitenen’ uit qd: ete, hem

7 LHF: dabei, eth ero wl 7 he» acspeatdinttises

7 a wetwng| Lani a: afl) it olteni iG

iy gd? onfA. SBiat ey att no ever |

: . “4 repeat hy - mart .“\ Ye tre ob

* an TR. n'ont iW. er Teen

as daca tubaaa? op ahepertio F

Oo: beth CAOT péawy tel a.

a» &, 7

Pe, aha

at Pay

The Tree-ferns of Malaya

R. E. HoLTTUM

AND

BETTY MOLESwWoRTH ALLEN

introduction (R.E.H.)

Preparation of the account of the tree-fern family (Cyatheaceae)

in Flora Malesiana (Series II, Vol. 1, part 2, 1963) involved a

study of all the species which have been named throughout the

Malayan region, some 350 names in all. This led to the discovery

that a good many species had been re-named, sometimes more than

once, by different authors; in such cases, the earliest name has to

be used. For this reason, the names of some species described in

Ferns of Malaya (1954) have to be changed. An example is

Cyathea kingii, named in 1883 from a Malayan specimen, which is

found to be indistinguishable from Cyathea lurida, originally des-

cribed from a Java specimen in 1828. Two more names were later

given to specimens from Sumatra, but they are not distinct from

C. kingii, the variability of which is shown by more ample collec-

tions made in Malaya. Besides changes made for this reason, two

more are due to the discovery that names used in Ferns of Malaya

were mis-applied; and three species have to be added to the list.

The changes are as follows.

Ferns of Malaya Flora Malesiana

Cyathea brunonis p. 117 = C. moluccana R. Br. ‘p. 143

C. latebrosa var indusiata p 121 = C. hymenodes Mett. p. 89

C. obtusata p. 121 = C. borneensis Copel. p. 110

C. burbidgei p. 124 = C. trichodesma

(Scort.) Copel. p. 150

C. ampla p. 125 — C. polypoda Bak. p 151

C. kingii p. 126 — C. lurida (Bl.) Copel. p. 121.

Misapplications. The true C. burbidgei (Bak.) Copel. and the

true C. ampla Copel. are both species of Sarawak which do not

occur in Malaya, and the names were wrongly applied to Malayan

ferns in Ferns of Malaya. The Malayan specimens misnamed C.

burbidgei were found to be identical with the type-specimen of

C. trichodesma at the British Museum (Natural History); this

species had been overlooked when Ferns of Malaya was being

written. The Malayan specimens misnamed C. ampla are only

distinct from C. polypoda in their broader leaflets, and are now

included in that species (true C. ampla has indusiate sori).

Additional species are C. incisoserrata Copel., C. sumatrana Bak.,

and C. alleniae Holttum.

In Flora Malesiana are new keys for the identification of species.

But as these keys cover 191 species, mostly not in Malaya, they

42 Gardens’ Bulletin, Singapore — XXII (1967)

are unnecessarily complex for local use. A simplified key is there-

fore here provided, covering only the species known to be native in

Malaya.

The final part of the present paper consists of a summary by Mrs.

Allen of her observations, made over many years, on the local

occurrence of the individual species in Malaya. These notes will be

of great value to local field botanists. They will also be of historic

interest, as unfortunately species have been exterminated in some

localities by felling of the forest in which they once grew, and more

areas of forest will suffer the same fate, as more land is taken over

for other use. In addition to this information on local occurrence

of species within Malaya, a summary of their known distribution

outside Malaya is given; this is taken from Flora Malesiana, and is

based on a study of specimens preserved in many herbaria in

various parts of the world.

Key for identification of tree-ferns in Malaya

Note. In this key, the word pinnules, unless otherwise stated,

refers to the larger pinnules of middle pinnae; upper pinnae are

always smaller, and lower ones more or less reduced.

1. Scales at base of stipe dark, shining (except in C. excavata), with paler

fragile edges which are often abraded when fronds are old.

2. Pinnules lobed almost to costa throughout; an indusium always present

(sometimes very small and hidden by sorus); lower surfaces of rachises

rarely very dark; little difference in size between sterile and fertile

pinnules.

3. Pinna-rachis bearing some hairs on lower surface, at least distally:

indusium a little wider than base of sorus, at least on costular side.

4. Pinna-rachis rather densely hairy throughout on lower surface;

indusium very fragile, completely covering young sorus, breaking

at maturity and leaving an irregular shallow cup ..................

1. C. sumatrana.

4. Pinna-rachis hairy towards apex only on lower surface; indusium

rather firm and disc-shaped or only on costular side of

receptacle.

5. Indusium forming a disc round receptacle (sometimes

asymmetric); stipe-scales to 4 mm. wide ...... 2. C. hymenodes.

5. Indusium only on costular side of receptacle; stipe-scales

to 1 or 14 mm. wide.

6. Lower pinnae usually much reduced; larger scales on

costae not setiferous; pinnules to 10 cm. Iong ...............

3. C. borneensis.

6. Lower pinnae not much reduced; larger scales on costae

setiferous; pinnules to 14-15 cm. long ...... 4. C. alleniae.

3. Pinna-rachis lacking hairs on lower surface; indusium either com-

pletely covering young sorus, or quite covered by sorus.

7. Indusium completely covering young sorus, breaking later; no

bullate scales on costae and costules ............ 5. C. excavata.

7. Indusium very small, covered by sporangia; bullate scales

abundant on costae and costules.

8. Pinnules of well-grown plants commonly 23 cm. wide, some-

times to 34 cm., with several basal pairs of segments separately

attached to costa; largest segments deeply lobed near bases;

costules 44-54 mm. apart .................. 6. C. incisoserrata.

8. Pinnules of well-grown plants rarely to 2 cm. wide; at most

one basal segment almost free; segments not deeply lobed;

costules usually 3-34 mm. apart .................. 7. C. latebrosa.

Holttum & Allen — Cyathea in Malaya 43

2.- Pinnules rarely lobed more than 4 way to costa, or if lobed nearly to

costa the fertile segments much narrower than sterile; no indusium;

lower surfaces of rachises always dark, often very dark.

9. Greatly reduced pinnae, separate from the rest, on basal part of

in Pee edie cotta nqdinenarineenny 1 etaae RAT tate 8. C. recommutata.

9. No greatly reduced pinnae at base of stipe.

10. Sterile pinnules lobed almost to costa throughout, bearing

bullate scales on lower surfaces of costules; lobes of fertile

pinnules: greatly :contracted (¢:.2:.1.8.000N011.055.!.. 9. C. lurida.

10. Sterile pinnules not lobed nearly to costa, with no bullate scales

beneath; lobes of fertile pinnules not greatly contracted.

11. Pinnules borne on very short stalks and lobed 4 way to

costa or more; sori on lowest veins not near costule, those

on higher veins progressively nearer to it, the arrangement

thus an inverted V; basal basiscopic vein of each group

often from costa, not from costule ......... 10. C. gigantea.

11. Pinnules borne on stalks 2-4 mm. long, lobed less than

4 way to costa; sori on all veins about equidistant from

costule; basal basiscopic vein of a group attached to costule

above. base coccostule. 24W...8078...600.5 00% 11. C. glabra.

1. Scales at. base of stipe thinner, usually pale, edges not of different texture

but bearing many short oblique dark setae.

12. Most segments of pinnules quite free as tertiary leaflets; sori apparently

indusiate, actually covered by overlapping scales ...... 12. C. tripinnata.

12. Most segments of pinnules not free: sori indusiate in nos 13 & 14, in

other species exindusiate, never covered by overlapping scales.

13. Fronds simply pinnate; pinnae entire or + crenate; indusium con-

spicuous, covering young sorus (disppearing HOCET) CR eres kek et

13. C. moluccana.

13. Fronds with deeply lobed or pinnate pinnae.

14. Pinnae deeply lobed, or almost pinnate near base ...............

14. C. alternans.

14. Pinnae fully pinnate or almost so throughout; no indusia.

15. Pinnules lobed almost to costa throughout; stipe and lower

surfaces glaucous, stipe strongly spiny ....................0..005.

15. C. contaminans.

{5. Pinnules not lobed nearly to costa; lower surfaces not

glaucous, stipe not strongly spiny.

16. Long spreading hairs present on lower surfaces of pinna-

rachis, costules and veins ......... 16. C. trichodesma.*

16. No long spreading hairs on lower surfaces.

17. Largest pinnules with a free segment at base;

pinnules on stalks to 4 mm. or more long; texture

METRO soos sacs Chap ¢2 Gage ett «See on 2 17. C. polypoda.

17. Largest pinnules lacking a free segment at base,

sessile or nearly so; texture thinner.

18. Sori on 3-4 pairs of basal veins of each group

only, not on distal veins, at maturity confluent

18. C. obscura.

18. Sori on almost all veins, not confluent at

NAAM mses cay asset swraian- 19. C. squamulata.

* A plant intermediate between C. trichodesma and C. alternans, pro-

bably a hybrid, has been found near Ampang Reservoir, Kuala Lumpur

(see Flora Malesiana p. 150). .

44 Gardens’ Bulletin, Singapore — XXII (1967)

Notes on the distribution of the Malayan Tree-ferns (B.M.A.).

Cyathea sumatrana Baker

General distribution: Sumatra and the Malay Peninsula, between

500—1500m.

One sterile specimen has been found in Malaya, in 1957 (M.A.

3447). It was growing at an altitude of about 500m in Selangor in

tall forest on the upper side of the Ginting Sempak road, near the

19 9/10 milestone. The fern was in a sandy patch which was in

the middle of a flat, wide but shallow stream. Although it was not

in the water it must frequently be inundated, and perhaps had been

washed down from a higher altitude, for no others were found in

the vicinity, either in the stream-bed or in the forest, where there

was comparatively little floor covering. Some young plants which

appear to be this species were found alongside the Kinta and Ulu

Piah rivers in Perak at about 170m in altitude. No adults were

found although the area was searched, and so they should be

looked for further upstream.

The Selangor specimen had a very short trunk, not more than

30cm tall and the fronds when living, were soft in texture and

bluish-green and dull above. The stipes, rachises and costae were

almost black above but pale below and there were distinct

excavations at the stipe bases.

C. hymenodes Mett. (syn. C. latebrosa var indusiata Holtt.)

General distribution: Sumatra and the Malay Peninsula, 700-

2200m.

This is not a common Malayan fern for so far has been found

only in Pahang and Johore in mountain areas, but of course may

turn up on other hills, especially on the Main Range. It grows in

the shade of tall dark forests often where the ground is sloping and

the soil loose and damp, sometimes by streams but more often well

above them.

It is not uncommon at Fraser’s Hill but is easily overlooked as

it resembles C. latebrosa. One of the places it occurs frequently is

on the forest slopes on the north side, below Victory and Cecily

bungalows, near and below the main contour path. It has also been

found on the quartz ridge at the beginning of the path above

Pahang bungalow, and in the forest about half-way to Pine Tree

Hill. At Cameron Highlands where it seems to be more local, it

has been collected near the junction of the Terla and Telom rivers

at about 1200m, and on Gunong Batu Brinchang where specimens

were collected at an altitude of 2200m; the plant was growing on

steep but wet ground and had a trunk of nearly 250cm in height. An

interesting habitat was in an area of recently cleared forest, where

only a few saplings and shrubs remained; near some well established

young plants of C. excavata were a few adult C. hymenodes in full

sun. One had a trunk of about 1m in height, and the fronds were

shiny and tough. Although it was too early then (1962) to know

whether they were really established, it would be interesting now to

Holttum & Allen— Cyathea in Malaya 45

have further observations. The area is in a peaty ridge near Walker-

burn cottage (now Che Foo) and above Brinchang village, on a

woodcutters’ path a few metres from the road.

In Johore C. hymenodes was collected from the summit of

Gunong Muntahak. Fronds are usually dark green above and the

trunk is slender and not more than about 350cm in height.

C. borneensis Copel. (syn. C. obtusata Rosenst.)

General distribution: from Mergui southwards, and Peninsula

Thailand; Malayan peninsula and Sarawak, in lowland forest to

1200m.

In Malaya this species has been collected from between 100-

1200m from Perak, Pahang and Selangor, but not very much is

known about its distribution and is possibly more widespread than

we know. It is a forest species not tolerating much sunlight and

therefore may have been more common when it was first collected

in Perak last century, for specimens do not seem to have turned up

again until 1957. In an area which many years ago was milled but

not cleared, where there was tall secondary growth mixed amongst

the taller trees, C. borneensis was found to be common. In 1962

this place (on the slopes of Bujang Melaka mountain) was com-

pletely cleared for rural development, but on more remote places

on the mountain it may still exist. It has also been found in several

places in the Tapah Hills Forest Reserve on the foothills of the

Main Range in Perak, and although apparently absent from wide

areas is common enough where it does occur. In Pahang it grows in

more or less undisturbed tall forest below Fraser’s Hill on the north

side at about 1200m, sometimes together with C. hymenodes. In

Selangor it occurs sporadically in the tall forest by the side of the

Sungei Berok above Kepong Forest Research Institute; in the

forest surrounding the Ampang Reservoir, alongside streams, and

in the Ginting Sempak it was once collected by a streamside in the

forest above the road at 300m.

The fern is nearly always near water and always in gloomy

places. When living, the fronds are usually very dark green above

and pale below, and one Pahang specimen had a trunk of nearly

5m in height with a circumferance of only 30cm.

C. alleniae Holttum

General distribution: Malayan Peninsula, between 1100-1200m.

So far known only from the Kuala Terla area in the Cameron

Highlands district of Pahang. The original plant was growing in a

band of forest above the Terla river, near the junction of the Terla

and Telom rivers. No others were found here, but a few hundred

metres further on by the main road towards Blue Valley Estate,

several patches of young plants (some fertile) were found, and

about 16m above the Telom river, under a light forest canopy and

on steeply sloping ground, others were seen. Some had trunks to

2m in height and all had pale green fronds. Several miles away,

46 Gardens’ Bulletin, Singapore — X XII (1967)

near a tributary of the Telom more plants were found, but none

so tall as the type which had a trunk about 4m tall, with small

crowns of fronds from near the base of the trunk. The species

apparently tolerates a great deal of light but not exposure to wind,

favouring forest edges and open places which are sheltered, or in

light forest and not on streamsides, but often growing well above

where the humidity would be more or less constant. Without more

field work it is difficult to be sure of its proper habitat, adapting

itself as it does to disturbed forest. The pinnules break off very

easily but the stipe base does not (in most species of Cyathea this

is easily detached from the trunk by a tug).

C. excavata Holttum

General distribution: Malay Peninsula; again known only from

the Cameron Highlands district of Pahang, from about 1170

to 1900m.

This is a very distinctive species and occurs in many areas around

the hill station environs. It is common in open places alongside

‘streams where the forest has been cleared but where a few small

trees, shrubs and tall grasses line the banks, and seems to be in-

creasing by the sides of shaded streams near bungalows where the

banks undergo periodic cleaning. These two habitats are no doubt

artificial ones, and as Dr. Holttum states this species was originally

found in primary forest, and it is still to be found on the more or

less undisturbed forest slopes of mountains such as Jasar, Ruil and

Brinchang by small streams or on flat wettish ground. It also occurs

in the forests below Cameron Highlands near the Terla and Telom

rivers. Apparently it will grow when away from water for in a

cleared area on the slopes of a peaty ridge (see under C.

hymenodes) C. excavata was quite common, yet there was no

stream near enough to influence it.

It is easily distinguished from C. contaminans witch is the

common tree-fern of the open places at Cameron Highlands, by

its soft fronds and completely smooth stipes.

C. incisoserrata Copel. (syn. Alsophila latebrosa var. ornata

Ridley*)

General distribution: Sarawak and the Malay Peninsula; from

the lowlands to 1400m or more.

This has been found in Penang, Perak, Pahang and Johore,

and is a tall species growing on forest edges or in clearings within

the forest, requiring light and sun. On Penang Hill it was growing

at about 650m altitude, and in Perak was quite common on forest

edges both on the lower slopes of Bujang Melaka and the foothills

behind Gopeng. At Fraser’s Hill it grows commonly on the edges

of tall forest which borders Girdle Road, the trunks being as tall

as C. contaminans with which it grows, but it differs by the more

* See H. N. Ridley: The Ferns of Malay Peninsula —J. Mal. Br. R.

As. Soc. IV, 8: 1926.

Holttum & Allen— Cyathea in Malaya 47

lacy appearance of the fronds and the non-glaucous stipe bases.

In Selangor it is quite conspicuous in sunny places on the forest

edges at Ampang Reservoir: here the fronds are yellowish. In

Johore it was found at Tanjong Kupang (Ridley) and on the

forest edge between Mersing and Endau.

This is a species which is possibly increasing with the changing

face of our forests, for it obviously tolerates much sun and seems

equally happy in damp or dry ground; also plants become fertile

at an early age.

C. latebrosa (Wall. ex Hook.) Copel.

General distribution: Hainan, Indochina and Thailand; Malaya,

Sumatra, Borneo.

In Singapore and Malaya, throughout the lowlands and up to

2000m. This is the species most frequently seen on roadside

ditches where there is shade; on the floor of old rubber estates; on

streamsides in secondary growth; in swamp forests and is one of

the few Cyathea species to be recorded from the bases of limestone

outcrops. In the hill stations it is a common fern in both forest

and rather open places, the highest being at 2000m on Gunong

Batu Brinchang in Pahang. It is usually a small species with a

slender trunk and soft fronds.

C. recommutata Copel.

General distribution: Central and S. Sumatra, Malay Peninsula

& Borneo.

In Malaya it has been found between 600-1500m in Perak on

Maxwell’s Hill; in Pahang at Fraser’s Hill and gn Gunong Tahan;

in one or two places on the mountain ridges in Selangor, and on Mt.

Ophir in Johore. C. recommutata does not seem to be a common

fern, apparently requiring a. special environment. It has been found

on peaty ground, on quartzite ridges and in short mossy forest,

sometimes in association with C. lurida, but always in the shade.

Previously it was collected from Bujang Melaka in Perak, but

does not seem to be there now. It is a small dark tree-fern, easily

distinguished by the very small pinnae on the stipe bases.

C. lurida (Bl.) Copel. (syn. C. kingii (Clarke) Copel.)

General distribution: Sumatra, Malay Peninsula, West Java and

the Philippines (Mindoro only).

This species is found in Malaya between 1250-2220m in Perak

and Pahang, on some of the higher hills. It is most easily seen in

Perak near the summit of Gunong Hijau above Maxwell’s Hill,

and at Cameron Highlands and Fraser’s Hill in Pahang, the highest

altitude recorded being near the summit of Brinchang in mossy

forest. Although a fern of the forest shade it seems to persist for

some years on forest edges in rather open places, and is very

common on the quartzite ridges around Fraser’s Hill. It is nearly

always trunkless, occasionally short ones are seen; this, together

with the long stipes, drooping laminae and contracted fertile

pinnae make it easy to recognise.

48 Gardens’ Bulletin, Singapore — XXII (1967)

C. gigantea (Wall. ex Hook.) Holttum

General distribution: Ceylon and S. India, N. E. India to Burma,

Thailand and Annam, Malaya, Central Sumatra and W. Java.

In Malaya found only on Langkawi Is. (Kedah), Penang and

Upper Perak from sealevel to about 350m, in rather open places.

In Langkawi it was collected from Gunong Raya, and in Penang

on the east and north eastern slopes of the hill, but as much of

this area has now been cleared it may not still be there. In Perak

it was found recently (1960) growing on flat banks of the Plus

river, towards Legap on the outskirts of light forest, and some,

being close to the riverside were quite exposed. Another species

(not fertile) was nearly always growing with it. Neither species was

common. In the field C. gigantea has a similar appearance to C.

glabra; the differences are given in the key.

C. glabra (Bl.) Copel.

General distribution: Sumatra, Malaya, Borneo and W. Java.

In Malaya it occurs in forest from sealevel to 1700m and has been

collected from most states except Penang. On Kedah Peak it grows

in light forest at about 860m; in Perak on the higher parts of

Maxwell’s Hill and near Ipoh on the edges of the rocky lowland

forest bordering the Kinta river at more or less sealevel, and at

270m near a quartz outcrop on sloping dry ground. It does not

seem to be common in the lowland swampy forests here, as it is

further south. In Selangor and Pahang it grows under tall

forest on sloping ground near the tops of ridges on the Main

Range, but not near streams. It is common at both Cameron High-

lands and Fraser’s Hill. In Singapore it still occurs in Mandai

forest but is now rare.

C. tripinnata Copel.

General distribution: Malaya, N. Borneo, W. Java, Amboyna,

Philippines.

In Malaya found only on Pulau Tioman at Bukit Telang and

Sedagong at 270m. This species should be looked for on the

mountains near the east coast of the Peninsula( see Holttum, Ferns

of Malaya p. 120). On Mt. Kinabalu where it is fairly common

locally, it is quite distinctive with its attractive lacy fronds. It is a

tall tree-fern.

C. moluccana R. Br. (syn. C. brunonis (J. Sm.) Wall. ex Hook.)

General distribution: Central Sumatra, Malaya, Lingga, Borneo

(excluding S. & S.W.), South and (?) Central Celebes, Moluccas

(Ceram, Amboyna).

It has been found throughout the Malay Peninsula except from

perhaps Perlis, from sealevel to 1300m. Although a common

lowland and foothill fern of disturbed forest and short secondary

growth, it appears to be absent from wide areas. It is very common

on Penang Hill; on the lower (dryer) slopes of Maxwell’s Hill, on

forest edges; and on the roadsides to both Cameron Highlands and

Fraser’s Hill, becoming rarer about 650m (one young plant was

Holttum & Allen — Cyathea in Malaya 49

found on a ridge near F. H. at 1300m). As it not only tolerates,

but appears to thrive in short second growth, especially when near

streams, it may be on the increase in many areas. C. moluccana is

easily recognised by its simply pinnate fronds; it is nearly always

trunkless, but in old plants in tall foothill forest, trunks may be

seen up to 50cm. This species probably hybridises with C. alternans.

C. alternans (Wall. ex Hook.) Presl

General distribution: Sumatra, Malay Peninsula, Sarawak and

N. Borneo.

In Malaya this species occurs in most States from Penang and

Kelantan to Johore, from the lowlands to 1430m, but is often very

local and thus may be overlooked. It is possible that it is more

common than is known at present. It is certainly not uncommon on

Penang Hill and in parts of Perak, and often grows in association

with C. moluccana on forest edges, but is much rarer than this

species, sometimes only solitary plants being found. In Perak C.

alternans has been seen several times growing together with a

Pleocnemia sp. (which was always infertile). Mature specimens on

Penang Hill at about 650m had trunks 2-3m in height, but this

does not seem to be commonly seen now, for young, trunkless

plants are usually encountered, and these in shaded places in tall

secondary growth. Tall and mature specimens which have also been

seen in Perak, Pahang and Selangor, appear to be quite happy in

full sun, on forest edges, and with one exception were not near

streams, yet young specimens frequently are. Observation for nearly

ten years on two plants suggests that they are slow growing, but

young ones are definitely increasing in undisturbed secondary

growth. At Fraser’s Hill there is a mature plant with a trunk about

45cm high growing on the forest edge, on the lower side of Girdle

Road which is just over 1300m, and a young trunkless one was

seen on the summit of Peninjau near the wireless station. Fronds

vary tremendously in the pinnae shape and lobing, and on the Gap

road in Selangor, in one of the Agricultural plots at about 700m, are

some tall specimens with curiously shaped pinnae.

C. contaminans (Wall. ex Hook.) Copel.

General distribution: Throughout Malaysia; from the Malay

Peninsula northwards to the latitude of Mergui.

In Malaya this species is almost as widespread as C. latebrosa

and certainly the commonest species of the hills. It grows in

abundance on forest edges, on streamsides in cleared valleys and

on mountain slopes in open places, and tolerates a great deal of

sun and exposure, and is not a fern of the forest understory. This

is our tallest species and adapts itself easily to new habitats, and

has fairly recently been recorded from about sealevel, on riversides

in flat areas (near Taiping, behind the town; on the banksides of

the Sungei Raia where it emerges into the Kinta Valley). It would

thus appear to be on the increase in the lowlands. It has been

collected from as high as 2000m on Gunong Batu Brinchang in

Pahang. |

50 Gardens’ Bulletin, Singapore — XXII (1967)

C. trichodesma (Scort.) Copel. (syn. C. burbidgei of Ferns of

Malaya, not (Bak.) ).

General distribution: Central part of the Malay Peninsula,

Sarawak, N. Borneo in lowland and hilly forest (to 1500m in

Borneo).

So far it has been found in Perak, Pahang, Selangor and Negri

Sembilan, up to 430m. It seems rather rare, except in Selangor

where it is common locally. In Perak it grows in rocky forest (near

the edges and thus in a fair amount of light) on the Kampar side of

Bujang Melaka mountain, facing west, yet apparently not on the

Chenderiang side which is of a more northern aspect. In Selangor

it has been found on the edges of light forest below the Gap,

usually in damp places; in the forest above Kepong and Sungei

Buloh and at Ampang Reservoir near streams. This species with

its softly hairy olive-green fronds is easily recognised in the field.

Two adult plants found at Ampang Reservoir may be hybrids

having C. trichodesma as one of the parents. More observation is

needed; the plants were on the lower (left hand side) side of the

contour path past the lake at about 60m above the stream, and on

very steep ground. C. trichodesma and four other species were

growing fairly near.

C. polypoda Bak.

General distribution: Malay Peninsula, Sarawak, N. Borneo and

the Philippines (Panay, Mindanao).

C. polypoda has been found only in Perak and Johore, with an

uncertain record from Pahang, between 1100-1300m and is a species,

so far we know, of restricted distribution. In Perak it is common

where it grows on the upper slopes and summit of Gunong Kledang,

a small mountain near Ipoh, where there were many tall specimens.

Below a forest ridge above Lake Chenderoh, several small

infertile plants were found, but were undoubtedly this species. In

Pahang in the Telom Valley below Cameron Highlands at about

930m. one small infertile plant was seen and the adults should be

looked for on the small isolated peaks around there (below Mt.

Penelope). This is a distinctive species with shiny deep green

fronds and pinnules cut into large segments. Apparently it prefers

a dryish ridge or steeply sloping ground in mid-mountain forest,

but not near streams.

C. obscura (Scort.) Copel.

General distribution: Sumatra and the Malay Peninsula, in forest

between 900-2000m.

This has been collected from Penang, Perak, Pahang and

Selangor, and is a common species of the higher forest slopes of

Maxwell’s Hill, Cameron Highlands and Fraser’s Hill and on other

hills in the Main Range in Selangor. It is a fern of the heavy forest,

Holttum & Allen — Cyathea in Malaya 51

not necessarily near streams, but usually in dampish places, yet it

will survive on forest edges where clearing has exposed the plants.

The fern is a large one with trunks to about 3-4m; fronds are deep

green above (yellowish and contracted when in exposed places)

and rather thick and brittle. The long paraphyses which are amongst

the sporangia in the sori are very easily seen.

C. squamulata (Bl.) Copel.

General distribution: Sumatra, Malaya, W. Java, Borneo and

the Sulu Archipelago.

To be found in lowland forest to at least 300m, but more field

work is needed in Malaya for us to know its present day distri-

bution and habitat. The fern was locally common on the lower part

of Penang Hill, but appears to be rare there now; there is one

record from Perak, growing in light secondary growth by an old

rubber estate towards the base of Maxwell’s Hill; in Selangor it

was found (again one plant) on the edge of secondary growth

where there was a Resam (Gleichenia sp.) thicket In the past it

has been collected quite often in Johore and in Singapore and in

the latter place it is still found in several localities: Nee Soon and

Mandai forests and in light growth near the base of Bukit Timah.

Almost certainly the original habitat of this fern in Malaya was

moist lowland forest (see Holttum, Ferns of Malaya p. 123), but

with so much of this now destroyed, it seems to be, today, more

frequent in dryer and lighter places in secondary growth.

— |

i %

7; a

. is >

i wi

’

‘

4 =

i ut Whe Makes Beye

ep twaly ata teh : a.

ipinees-: vant eR pethcns vel

haste me tw want neds

vos

} iy

* §

ary

‘

: . +p

* ~ fy

fe Te

oy s

hi )

et « b

a? 4 '

rhs

dj AN iA)

, sf Aft - ‘7

4 ; Me

] M ;

AM Pe

yg a: fee

y- ,

* $

ey ¢ 7

of b whl 6

‘

‘ ; a » edt all

~ | sn , * ae 7

= ong i, peewee ,- See

aT ory ‘Meld ye

. a

ats =

Ww - ;

j \

a BA!

chittai a4 ic]

or 8

a A ¥ —_— 7

Y aa Piao

Ke way i; Ty |

iv Pet, Lares

Lip. } hea

Notes on the distribution of Nepenthes

Species in Singapore

SALLY GREEN.*

Species found in Singapore.

N. gracilis Korth, N. rafflesiana Jack and N. ampullaria Jack

are all common: N. hookeriana Lindl. and N. trichocarpa Miq.

both occur rarely and singly. In the past N. reinwardtiana Miq.

and N. phyllamphora Willd. were recorded from the Island. Quite

recently a few plants were found with a superficial resemblance to

N. mirabilis Druce, with which N. phyllamphora is now merged,

but differing from it in important ways, such as the shape, of the

peristome, the venation of the lamina and the lid, and the number

and complexity of the glands under the lid. The Singapore plant

cannot be N. mirabilis, and since it occurs singly it is probably a

hybrid.

Hybrids.

So far as is known, all species of Nepenthes can be crossed,

producing offspring of great vigour and fertility, which can be

used in further crosses; as was done in nineteenth century Europe,

where nepenthes breeding had some of the appeal of orchid

breeding to-day. Many new varieties were formed; some were

beautiful and most sold for high prices. Although gardens hybrids

are produced so readily it appears that there is no proved case of

a natural hybrid occurring, though taxonomists agree that they

probably do. The three possible hybrids in Singapore are N.

hookeriana, N. trichocarpa, and the plant resembling N. mirabilis,

all of which are rare and generally grow singly. N. hookeriana is

widely thought to be the hybrid of N. ampullaria and N.

rafflesiana, having some characters of each, and the right geogra-

phical and local distribution. The nineteenth century horticul-

turalists knew N. hookeriana, and esteemed it highly; they used it

in their crosses, but unfortunately they did not determine its status.

N. trichocarpa has been accepted as a good species with an odd

geographical distribution: Sibolga in N. W. Sumatra, and the

Singapore, S. Johore region (Danser, 1928). In the herbarium of

the Singapore Botanic Garden however Holttum has classed it as

hybrid of N. ampullaria and N. gracilis, and the appearance of the

plant and its occurence, in Singapore, make this very probable.

The third possible hybrid resembles N. trichocarpa even more

than it resembles N. mirabilis; and it is perhaps a cross of N.

trichocarpa with N. gracilis. All of the three possible hybrids are

very vigorous and pleasant to cultivate, producing plenty of good-

sized pitchers under a fairly wide range of external conditions;

unfortunately this is not true of the three good species.

* Present address: c/o Institute of Education, University of Keele,

Staffordshire, England.

54 Gardens’ Bulletin, Singapore — XXII (1967)

Distribution of Nepenthes species in Singapore. (See Map).

Nepenthes has an ancillary food supply in the prey caught and

digested in its pitchers, and it might be expected that it would grow

on peculiarly infertile soil. This is indeed so. Singapore soils are

of volcanic, sedimentary or alluvial origin, and Nepenthes is re-

corded from the first two of these but not from the richer alluvium.

Thus N. gracilis is found on the sedimentary outcrop at Ponggol

Point, but not further inland, and there are many old records

of Nepenthes growing on the granite at Changi, but none from the

alluvial Tampines region just inland from this. At first sight the

Tampines scrubland is very similar to that in which Nepenthes

grows on volcanic and sedimentary soils, and some of the same

plants associated with Nepenthes in such sites, can be found there;

these include Melastoma malabathricum and Dicranopteris linearis,

which are both abundant. Others however are absent; Lycopodium

cernuum, a constant associate of Nepenthes, is most rare, and

Dillenia suffruticosa is represented only by occasional unhealthy-

looking plants. Evidently the region is unsuitable for Nepenthes

only in its soil. The Tanah Merah cliff in the alluvial region, which

bears Nepenthes, appears to be an exception, but in fact it is not

so; the Nepenthes grow on exposed ancient rock below the alluvial

level.

In the sedimentary and volcanic regions all the local Nepenthes

spp. can be found growing together, though their relative frequen-

cies vary. Typical habitats are: —

1. Coastal cliffs. e.g. Tanah Merah, Labrador, Tanjong Gul.

Pulau Tekukor.

There is no record of clearing or burning of the cliff vegetation,

so that it may be primary. The Nepenthes found here are often

very large indeed, flowering and fruiting abundantly, and with

many pitchers. N. rafflesiana predominates.

2. Roadside cuttings.

This is a habitat resembling the coastal cliffs in many ways,

though the cuttings are usually less exposed than the cliffs, and

many of them are of course quite recent. The Nepenthes are often

very numerous, though smaller than the cliff plants. Flowers, fruits

and seedlings are numerous. N. gracilis predominates.

3. Regenerating waste ground.

In Singapore there are many areas which have been cleared,

farmed and abandoned after the soil is exhausted. Exposure to

high insolation, leaching and erosion lowers the fertility still more.

Their history, if they are undisturbed, is of a slow change from

sandy wastes to secondary forests; Holttum (1954), Gilliland

(1958) and others have recorded the stages. It is not known at

what stage in the succession the Nepenthes plants first appear,

but they are not among the pioneers. In the scrub stage

Holttum’s ‘‘Adinandra Belukar” — all the local species are

common climbing in the bushes. All flower and fruit and there

fanpergey ay incr nt

Ter

Ine ni wnePaar eas weirs NC)

Oreos

DOnNqsIq

SUIS UO sayyUedaN Jo

‘puvysy olodesur -

? z 9

SOitw ads

cD MOVIG NI SHIOANTSTA

0 oa

HOMMNUEE be: awrng

’

‘

kl 5 we

ae = ns )

: i

-* - as | | |

- : . |

e ce eo it |

7 pease Seco, : 7

6 = ; Poe - ~ | :

‘eat om

eo, Sar) H |}

Sees.

“4 .

° m aes

e $03

. 9

. ese : Pbe

fees

NUdGUN

SaLIS SHHL

TIddIgs NI

Pooy yous sing

P98 UNTIL

SyIOY ais i vi lit

wnsanniy #0 (777

ANG MIN TAS

ee .

° °

See

e ee

lis

eeete?

i? oie

. HH ee ete

| . eid)

xt \e.@

4 = far el

ee. |

de .

s i

e EB

%e.

. SOREST ESS 2.

° ce eeees ..

e SOIC ESS ' :

e Pee, + ;

e :

i = Pee 2

5 %e fit,

eee r

ea AZ 7

cane eee fi

fa ZN

*"Seeed e j

ee ee

ce,

ee Z

= 2 :

——rs2

56 Gardens’ Bulletin, Singapore — XXII (1967)

are many seedlings. N. gracilis predominates. By the time the suc-

cession has reached forest conditions the relative frequency of the

species has changed; N. rafflesiana is now rare, N. gracilis is less

common and N. ampullaria predominates, This is the situation

where the forest is comparatively open and the trees of no great

height; it can be seen at present around the margins of Pierce and

MacRitchie Reservoirs. With the growth of the trees the height

of the Nepenthes has increased; they can be seen among the

branches 30 ft. or more above the ground. Seedling plants,

chiefly N. ampullaria, can be found, usually along rides where

these are lower than the forest level, so that there are sloping

banks rather like miniature cliffs on which the small plants grow.

In denser forests such as that bordering Mandai Road the relative

frequency of the species changes again; here N. ampullaria is very

plentiful, but the other two species are quite rare. There are no

flowers but N. ampullaria proliferates from shoots on prostrate

stems. Its dense clumps of ground pitchers are extraordinarily

- numerous, but on the few plants of the other species pitchers are

rare. The plants are old; that and the absence of flowers and

seeding suggests that they are the last survivors from an earlier

stage of regeneration. This is the end of the Nepenthes period in

the re-development of forest; the plants are absent from well-

established secondary forest, as they are from primary Dipterocarp

forest.

4. Peat swamp. Nee Soon.

Singapore has one small peat swamp at Nee Soon in which

Nepenthes is plentiful. All the local species grow on the banks of

the broad cutting leading to the swamp, with some hybrids; seed-

lings, particularly of N. ampullaria, are abundant. In the swamp

itself there is no N. ampullaria; N. gracilis is common, small plants

apparently growing on detritus floating on the water, and larger

ones growing from islands formed round the bases of Ploiarium

trees, where N. rafflesiana is also found. The island plants are in

no way stunted — they are in fact very fine. Both species flower

and fruit.

It appears that the ecological needs of the three common

Singapore species of Nepenthes are somewhat different. The sites

on the cliffs, cuttings and waste ground can perhaps be considered

together as a continuum, with the cliffs at one end of the scale

and secondary forest at the other; on such a scale shade, soil

water content, fertility and pH increase together, maximum

temperature, diurnal temperature range and evaporating capacity

of the air decrease; with available water still further reduced on the

cliffs by salt spray. From the distribution it is clear that the more

exposed, less fertile end of the continuum is best suited to

N. rafflesiana, the shadier, moister, more fertile end to N.

oe .

Green — Nepenthes in Singapore 57

ampullaria; while N. gracilis finds optimum conditions between the

extremes. The differences are not great; there is, for instance, a very

fine female plant of N. ampullaria growing on the Tanjong Gul

cliff, and fruiting abundantly; however this is an exception, and its

seedlings do not survive. The Nee Soon peat swamp ranges from

| a cutting in secondary forest through increasingly acid, infertile,

physiologically dry conditions to the wet centre. The Nepenthes

distribution follows a predictable pattern, with all species growing

in the cutting,V. ampullaria predominant at the forest edges, and

N. gracilis alone with one or two plants of N. raffiesiana in the

standing water at the centre. Anderson (1963) in a study of the

coastal peat swamps of Sarawak records a similar distribution.

After zoning the bogs into Phasic Communities 1 — 6, P.C. 6 being

the climax, he found N. ampullaria occassional in P.C.s'1 and 2,

N. gracilis frequent in P.C. 4 and surviving though stunted in P.C. 6,

and N. raffesiana rare in P.C.s 3 and 4 but frequent in P.C. 6. In

peat swamp as well as belukar N. rafflesiana favours the most

infertile conditions, N. ampullaria the least.

Plate 1. N. ampullaria Jack.

| Ground, stem and leaf pitchers. Water Catchment Nature Reserve,

Singapore.

ee pre ei

58 Gardens’ Bulletin, Singapore — XXII (1967)

Some Ecological Factors affecting Distribution.

An exhaustive account of the ecology of the local Nepenthes

cannot be attempted here, but a general description of some of the

important points may help in understanding distribution. All the

Singapore species grow on the same or very nearly the same sites,

only their relative frequency varies; so that their ecological needs

can all be considered together. These are met not only in the

belukar and cliff habitats, which are clearly rather similar, but also

in the peat swamp. In Borneo the same species grow in peat

swamps and also in the heath forests, which have some resemblance

ecologically to the Singapore belukar. The two kinds of habitats

have in common acid, highly infertile soil, which is widely recognis-

ed as being a necessity, perhaps the prime edaphic one, for the

growth of Nepenthes. Indeed in 1914 Barrowcliff proposed using

the plants as an index of soil poverty. Macfarlane (1922) thought

that a well-aerated soil was also essential, and cultured his plants

accordingly; but this can hardly be so for the peat swamp plants,

which prosper as well as the belukar ones. Aeration is evidently

not important. Available water is low in the peat swamps, the

belukar has spells of aridity, and as well, Nepenthes is particularly

abundant on slopes and cliffs, from which the water drains quickly.

Anatomically the plants show signs of xerophytic adaptation, which

fit them to survive on a dry soil, though this may not be essential.

Cultivating Nepenthes, even in a suitable soil, is somewhat

difficult because their growth is much affected by climatic factors.

Holttum (1938) pointed out that the micro-climate changes

during the upward growth of climbing plants; so that adaptive

changes in their responses to external conditions are to be expected.

These are not very evident in Nepenthes until the flowering stage:

although the plants do show very considerable changes in

morphology during their growth, their response to climatic

factors appears to be relatively constant.

The Seed and Seedling.

About 10,000 seeds are produced from one inflorescence of

N. gracilis. The seeds are remarkably light, and are winged, so that

they can be dispersed by the wind over long distances. Each of

the two filamentous wings is hollow, so that the seeds are buoyant;

they can be carried for short distances by rainwater rills. Wastage

is obviously high by either method of dispersal, but the seeds are

so numerous that some must find their way to suitable habitats.

Garrard (1955) estimated the viability of N. gracilis as 8.3 per cent

but counts made by the author gave 79 per cent and 63 per cent;

Stern (1917) mentions differences in fertility in seeds from

different parts which may account for the discrepancy, and probably

viability does not limit distribution. The seeds germinate in the

light, and within limits increasing the light shortens the time

required. This is a fortnight to three weeks after sowing for most

of the seeds, but for others it is very much longer; as late as six

or seven months after sowing it is still possible for seeds to

Green — Nepenthes in Singapore 59

germinate. This variation may have some survival value, since the

seedlings emerge into different climatic conditions, some more

suitable than others. The seedling is very small. From the first it

is adapted to life on infertile soil, since the leaves, though marked-

ly different from those of any adult plant, all bear tiny pitchers.

These are functional with digestive glands in the cavities. The

attractive glands, which are so numerous and so complex in adult

pitchers, are restricted in the primary leaves to the rudimentary

rim under the mouth. Development is slow; a tiny rosette of

leaves is formed, and the plant remains in this form for the next

two or three years, with successive leaves gradually getting larger,

and successive pitchers slowly changing towards their adult form.

The seedling is quickly killed by dessication, either by wind or by

strong sunlight, and this even in damp soil, which perhaps sufficient-

ly explains the absence of Nepenthes from the pioneering plants on

bare ground. The seedling root is a very meagre one, and often

does not appear until after the cotyledons; wind can uproot the

small plants, as well as dry them. If the humidity is high the

seedlings survive even in a seemingly dry soil, though it is not

clear how this is achieved. A bunch of long, tough root hairs

appears outside the seed case even before the tip of the root

shows, but these are not enough for anchorage and they are

doubtfully enough for absorption. Stern (1917) suggested that the

seed coat was important, supplying water by capillary action along

its furrows. Macfarlane (1908) thought that minute glands

scattered all over the surface of the cotyledons and leaves absorbed

moisture directly from the air, though not all authors agree with

him. The problem is intriguing, but to understand distribution it

is perhaps enough to note that the seedlings are adapted for life

where the humidity is high, the air is still, there is some shade, and

not over-much dependence on soil water. In such situations they

are found—at the base of bushes, in nooks on banks and along

forest rides. However, there is another limitation; in shade —

by no means sufficient to cause etiolation — no pitcher is formed,

but only a large lamina. It is unlikely that the seedlings can survive

this deprivation for long.

The mature Plant.

The need for humidity, and a rather limited light tolerance,

dominate the life of the older plant as they do that of the seedling:

though as the plant matures some protective adaptations develop.

The cuticle thickens, the young leaf, exposed in the seedling, is

shielded in the older plant, the rudiment of the pitcher — the most

vulnerable part— is wrapped in hairs. But exposed leaves still

curl’ pitchers wither, actual burnt patches appear. Nevertheless the

plants, once established, survive quite adverse conditions, some-

times even fire. No doubt sheer size plays a part; a patch of strong

sunlight kills the whole of a seedling, but only one or two leaves of a

large plant. Also the root system becomes extensive, and once the

rhizome is formed new growth can come from that. The effect of

light can be seen most clearly if the growth of pitchers is watched.

60 Gardens’ Bulletin, Singapore — XXII (1967)

Where shade is too deep pitchers and tendrils do not develop on

stem or rosette leaves, laminas become large and thin. If exposure

is not high enough for burning, but still too high, both pitchers and {

their laminas are small; some leaves, notably in WN. rafflesiana, i

show twisting of the lamina as well as of the tendril, bringing the |

pitcher away from exposure. The species all react in this way |

until climbing is some way advanced; after that there are differences

which may be related to their slightly different habitats. Singapore |

has among its three common species one that is simple, and two .

that are highly specialised in quite different ways. The simple

species, N. gracilis, normally has a pitcher on every leaf, borne on

the end of a tendril which can coil round supports. The pitchers are

tubular and not very large; the wings are never wide, and may be

absent. On leaves at the base of the plant, the pitcher mouth opens

inwards, towards the stem; on the upper parts it opens outwards.

This twisting round of the upper pitchers appears to be normal in

the development of Nepenthes, since it occurs in most species. In

the simpler species, like N. gracilis, it is a gradual process with

intermediate stages; it involves no change in the shape of the

pitcher beyond a slight narrowing of the base, and no change

either in the response to light. Matters are quite otherwise with

Plate 2. N. gracilis Korth. ;

Growing on the surface of the freshwater peat swamp forest, Nee Soon

area of the Water Catchment Nature Reserve, Singapore.

Green — Nepenthes in Singapore 6]

the more specialised species; not only is the change in direction of

the pitcher opening quite sudden, between one leaf and the next,

but there may be a dramatic change in the shape of the pitchers.

N. rafflesiana is such a species. An altogether stouter plant than

N. gracilis, it has correspondingly larger pitchers. The lower ones

are broad-based with widely reaching wings; usually they sit on the

ground, joined by straight tendrils to their laminas — which may

be two feet or more above their level. The upper pitchers are

wingless, with the base so narrowed and curved that they have been

described as cornucopias. There are clearly biological advantages

| in this form; not only is wind resistance lessened, but so also is

the volume of digestive fluid in the base which has to be supported,

while the trapping area round the mouth remains wide. The two

| types of pitcher respond differently to light. In conditions of under

or oOver-exposure pitchers or tendrils develop on neither; but

where exposure is right, the tendril of the lower pitchers grows

rapidly downwards, carrying the pitcher away from the light —

which is normally of course towards the base of the plant. The

tendril of the upper pitchers grows outwards in an arc, and

normally coils round a support before the pitcher grows too big

for this to be possible. The lower pitchers wither very easily; they

cannot survive even that amount of exposure which, when applied

to the lamina, stimulated their development. The upper pitchers are

Plate 3. N. rafflesiana Jack.

On the sea cliffs of Pulau Terkukor, Singapore.

62 Gardens’ Bulletin, Singapore — XXII (1967)

more resistant, though still less hardy than their laminas. The upper

and lower leaves are adapted to different conditions in fact; as much

in their tropistic responses as in the different shape and hardiness of

their pitchers. Evidently making the best of two worlds is a feature

of Nepenthes organisation; N. raffiesiana — and most other species

too —can catch not only running insects on the ground, but —

with the mouths of the upper pitchers opening invitingly outwards

—flying insects as well, though the catch of flying insects is in

fact small. N. ampullaria is specialised in a different fashion; each

Hy YHjjp

Plate 4. N. rafflesiana Jack.

Ground Pitcher.

Green — Nepenthes in Singapore 63

leaf on the lower part of the growing stem bears a pitcher on a

short tendril which does not coil, but on the upper leaves, although

there is a coiling tendril, the pitcher does not develop. There is no

record of a pitcher of N. ampullaria in which the mouth opens

outwards. As the lower leaves and their pitchers die, they are not

replaced. Instead ground pitchers appear at the base of the plants,

borne on short shoots from prostrate stems and rhizomes. Each

shoot bears a tight rosette of pitchers, in which the rudimentary

laminas and tendrils serve only to bring the pitchers into position.

In forest conditions the ground pitchers cover large areas of

forest floor, half hidden in fallen leaves; they must be a most

efficient trap. Similar pitchers can appear on the brown, seemingly

lifeless stems that climb high in the trees. In effect, N. ampullaria

bears two kinds of leaves, one photosynthetic and climbing, the

other absorbtive, Perhaps this is a biological advantage, since

its distribution is wider than that of the other local species, not only

locally but geographically, and it can grow at higher altitudes. The

ground pitchers are borne in quite dense shade, which is some-

times provided on open ground by the bushy growth of this

remarkably adaptable plant. Many species of Nepenthes, including

all the Singapore ones except N. rafflesiana, can produce clumps of

ground pitchers from the mature plants. The pitchers have the same

form as those from the leaves on the lower part of the stem; as in N.

ampullaria their laminas and tendrils are so reduced as to be

scarcely discernible. These ground pitchers are a remarkable

development; all other leaves of the plants show enlarged laminas

when they are grown in shade, and pitchers are not produced.

Cultivators of Nepenthes in private or botanic gardens are some-

times troubled by a scarcity of pitchers. This can be due, not only

to too much or too little shade, but also to too rich a soil

(Smythies 1963). Such pitcherless plants sometimes flower and

seed normally. Evidently carnivory is not obligatory, at least for

older plants, though it may well be for early stages, since

Nepenthes, growing naturally, is usually excluded from habitats

with rich soils.

The Flower

The inflorescences are terminal, though growth continues

from lateral shoots. The climbing habit has the effect of bringing

the flowers out into the open, to the advantage of both pollination

and seed dispersal. They appear only where they can be fully

exposed, so that they protrude from the tops and sides of trees,

and a foot or more over the tops of bushes; and it seems that no

degree of exposure can be too high. A different micro-climate

indeed from that of preceding stages! A few leaves project along

with the inflorescences, and these are pitcherless — according to

Stern, because of diversion of food to the inflorescence. But this

is to neglect the function of the pitchers in procuring food, and it

64 Gardens’ Bulletin, Singapore — XXII (1967)

is perhaps more probable that the pitchers fail through over-

exposure. Few flowers remain unfertilised, but it is uncertain how

fertilisation is accomplished. The surfaces of the tepals are dotted

with large glands, which secrete so much nectar that it fills the

base of the spoonshaped tepals; nevertheless the flowers are seldom

visited by insects, not even by ants. Because of the dioecious habit,

and the floral structure, a pollinating insect would need to be large ~

and winged. The writer once had the good fortune to watch a bee

absorbed in drinking the nectar, which it did while straddling the

stamen column with its body. Such insects are well-fitted to be

pollinators, but there is no other report of their visiting the flowers.

Pollination is probably mainly by wind. Dr. Pallot, lately of the

Botany Department, University of Singapore, points out that the

pollen grains, though joined together in tetrads, are small enough

for this to be possible.

Conclusion.

Nepenthes distribution in Singapore appears to be rather

narrowly limited to the most infertile areas which can also provide

suitable climatic conditions. The plants are in fact fairly wide-

spread, but this reflects rather the prevalence of man-made waste

ground than any abundance of natural sites, of which only the

cliffs and perhaps Nee Soon swamp give any picture. The

Nepenthes population must have been quite small before culti-

vation began on Singapore Island; and must become so again

as one site after another is taken over for industrial and other

development.

All things considered, it cannot be said that Nepenthes is ideally

adapted to the dryland conditions in which it grows in Singapore,

especially in the early stages. The leisurely growth, the meagre root,

and the readiness with which the small plants and the older

pitchers — dry out, all suggest an origin in a different habitat. It

is tempting to see this in the peat swamps, home of many insecti-

vorous plants; and to suggest that the ancestral Nepenthes invaded

the dry land where it found a suitable soil and tolerable climate.

However, for the Singapore species in the existing lowland swamps

in Sarawak light is insufficient; Anderson describes them as

appearing to climb only where there is an opening in the tree

canopy, and only the climbing plants can reproduce. The ancestral

home of Nepenthes, as so much else concerning these unusual

plants, must remain problematic.

Acknowledgements

I have to thank the late Professor H. B. Gilliland, of the Univer-

sity of Singapore, Mr. B. S. Smythies and Dr. J. A. R. Anderson,

both of the Forestry Department, Sarawak, and Mr. H. M.

Burkill, Director of Singapore Botanic Gardens, for their help, and

my husband, for the photographs.

Green — Nepenthes in Singapore 65

REFERENCES

ANDERSON, J. A. R. The Flora of the Peat Swamp Forests of

Sarawak and Brunei.

Gard. Bull., Sing. 20, 2. 1963.

BARROWCLIFF, M. Malayan Rubber and Coconut Soils.

Trans 3rd. Internat. Cong. Trop. Agric. 2, 1914.

DANSER, B. H. The Nepenthaceae of the Netherlands Indies.

Bull. Jard. bot. Buit. ser. 3, 9, 1928.

GARRARD, A. The Germination and Longevity of Seeds in an

equatorial Climate.

Gard. Bull., Sing. 14, 2. 1955.

GILLILAND, H.B. Plant Communities on Singapore Island.

Gard. Bull., Sing. 17, 1958.

HOLTTUM, R.E. Adinandra Belukar. A succession of Vegeta-

tion from bare ground on Singapore Island.

Mal. Journ. Trop. Geog. 3. 1954.

HOLTTUM, R. E. (through Richards) The Ecology of Tropical

Pteridophytes in Verdoorn’s Manual of Pteridology. The Hague

1938.

MACFARLANE, J. M. WNepenthaceae. Das Pflanzenreich.

Leipzig 1908.

MACFARLANE, J. M. Nepenthaceae. chain's Cyclop. Hort.

New Ed. 4 1922.

RICHARDS P. W. The Tropical Rain Forest. O.U.P. 1952.

STERN, K. Beitrage zur. Kenntnis der Nepenthaceen. Flora,

[Ge 191 7. '

SMYTHIES, B. E. The Distribution and Ecology of Pitcher-

plants (Nepenthes) in Sarawak. Symposium on Ecological Re-

search in Humid Tropics Vegetation, Kuching, Sarawak, July

1963. UNESCO Science Co-operation office for South-east Asia,

1965.

ISAAC HENRY BURKILL 1870-1965

R. E. HoLTTUM

Volume XVII, part 3, of this Bulletin was published on May

18th 1960, to commemorate the 90th birthday of Isaac Henry

Burkill, Director of Gardens, Straits Settelements, from 1912 to

1925. Mr. Burkill died on March 8th 1965, thus having almost

attained the age of 95 years. In 1960 Dr. H. Santapau gave details

of Mr. Burkill’s early life and of his service in India before he

came to Singapore, and Dr. C. X. Furtado and I collaborated in an

account of his work in Singapore. The following is to be regarded

as supplementing these earlier statements.

Mr. Burkill’s early work as assistant in the University Herbarium

at Cambridge had trained him as a field botanist in the careful

recording of much detailed observation, especially regarding the

relations between flowers and insects. His work at Kew had first

introduced him to tropical floras and then to an interest in useful

tropical plants. In India he was officially engaged in collecting

information about useful plants, and during his many travels for

this purpose he took every opportunity to extend to Indian plants

his investigations of floral biology. He also began the detailed

study of wild and cultivated species of Dioscorea (true yams)

which he continued almost to the end of his long life. Thus he

came to Singapore with considerable experience as a field botanist,

with an extensive knowledge of the plants of tropical Asia,

especially those of use to man, and with a particular interest in

yams and in floral biology.

Burkill arrived at Singaore several months after the retirement of

H. N. Ridley, that dynamic and erratic pioneer genius who accom-

plished so much in 23 years, during which period the plantation

rubber industry was established, largely as a result of his experi-

ments with the Hevea trees in Singapore. Ridley was always in a

hurry and often uncritically careless about details. Thus Burkill

found much reorganization necessary in every aspect of the work

of the Botanic Gardens. In his efforts to achieve reorganization he

was hampered by lack of trained assistants, especially during the

period of World War I; he was also hampered by a failure of the

senior civil servants of his day to understand the significance and

needs of the work of the Botanic Gardens. But despite these

difficulties he did achieve important developments which had a

great influence both on the Gardens Department and, through his

published work, on many people throughout the world who have

been concerned with useful tropical plants.

68 Gardens’ Bulletin, Singapore — X XII (1967)

During the period between Ridley’s departure and Burkill’s

arrival, J. W. Anderson, Assistant Curator, prepared and had

printed a catalogue of the plants in cultivation in the Gardens. This

contained too many errors to satisfy Burkill’s critical examination,

but it showed the need for carefully compiled records of the

plants which had been introduced and of their behaviour in cultiva-

tion. Burkill therefore had prepared a complete series of extracts

from the plant introduction books, thus obtaining a list of all names

under which plants had been introduced, with their sources and

the date of each record. He also had herbarium specimens prepared

of plants in cultivation, and himself checked their names, attempting

thus to compile a complete record of the actual state of the

Gardens for comparison with the introduction records. This survey

included all the useful plants which has been introduced into the

Economic Garden (now the site of the Faculties of Arts and

Science of the University of Singapore). He also reorganized the

labelling of plants (especially trees, shrubs and palms) and every

week checked the painted labels before they were put out. He thus

trained Malay youths to have a knowledge of botanical plant names

and also to recognize the plants which bore them and to know

their places in the Gardens. Among these Malays was a very

promising young man named Mohamed Nur.

The herbarium, and field work for collecting new specimens, also

needed reorganization. Ridley’s methods had been casual, and in

some cases resulted in confusions of labelling. Burkill therefore

devised a new system of field labels, based on his Indian experience,

and trained his staff in the importance of labelling all specimens

at the time of collections. Needing a new assistant in the herbarium,

he promoted Mohamed Nur, who subsequently made such a great

contribution to this aspect of the Gardens’ work. Burkill collected

specimens during his travels wherever possible, and also arranged

for other members of the staff to do so, sending duplicates to

Kew, where they were available to Ridley, who was preparing his

Flora of the Malay Peninsula. As the successive volumes of this

Flora appeared, the herbarium was re-arranged, and Burkill took

the opportunity of making a careful survey of the extent of

botanical exploration of the various parts of the Peninsula (Gard.

Bull. 4: 113-202, 1927). He also made careful analyses of the floras

of two restricted localities in which he had himself made extensive

collections, namely Fraser’s Hill (with R. E. Holttum, Gard. Bull.

3: 19-110, 1923) and Taiping (with M. R. Henderson, Gard. Bull.

3: 303-458, 1925); these accounts give an indication of his critically

detailed treatment of his data, and also the wide scope of his

thought. ;

He made a special effort to secure herbarium specimens of wild

and cultivated yams, and also cultivated living plants whenever

possible, to observe their growth both above and below ground.

He cultivated many yams from other parts of the world, notably a

collection from Tahiti. He had collected local names for yams in

Holttum — Burkill, 1870-1965 69

India, and continued this practice in Malaya; he searched a very

wide range of literature for other recorded vernacular names, the

final list being more than 1500. His published commentary on the

list throws light on the migrations of man and on the history

of the use of yams as food; it also points out the value of such

records in linguistic studies.

On the subject of floral biology, Burkill was not able to publish

much, apart from studies of various Malayan orchids in cultivation;

these included the very remarkable Plocoglottis lowii. It was how-

ever clear to anyone accompanying him in the field (as I did on

several occasions during the years 1922-25) how deep was his

interest in such matters and how wide his knowledge.

Throughout his service in Singapore, Burkill was in close contact

with the Forest Department, and from 1918, when Dr.’ F. W.

Foxworthy was appointed to establish a Forest Research service,

this contact became closer, as Foxworthy needed frequently to

consult the Singapore Herbarium. Mr. G. E. S. Cubitt, head of the

Forest Department, hearing of Burkill’s accumulated records of

useful plants and their history in culture in Singapore, and knowing

also his work in connection with the preparation of Watt’s

Dictionary of the Commercial Products of India, suggested to the

Governor of the Straits Settlements that Burkill should spend the

years following his retirement in compiling such a dictionary for

Malaya. This proposal was approved, and in his last two years

of service Burkill spent as much time as he could in collecting data

for the Dictionary in various part of Malaya. In this work he was

assisted by Mohamed Haniff of Penang. They collected especially

information about local medical uses of plants, and in so doing also

collected a large amount of information on local plant-name. The

results of this work were published in Gard. Bull., 6 (2), 1930.

It had been suggested that the Dictionary would be completed

in the three years following Burkill’s retirement. But he spent the

whole of those years searching literature, at Kew and elsewhere,

and accumulated a card-index which he told me contained 36,000

entries; only then did he begin the writing. The work was finally

published in 1935, ten years after its commencement. It is the most

comprehensive account of useful tropical plants, and of the history

_ Of their use by man, that has yet been published. Naturally, it is

now not up to date in many matters, but as a basis of fact about the

past it will always be of great value.

This great work accomplished, Burkill reverted to his interest in

yams, and (in collaboration with Sir David Prain) completed the

text of a great monograph on the oriental species of Dioscorea

(Annals of the Royal Botanic Garden, Calcutta, 14, 1936-38). He

extended his studies to African yams, and devoted much time also

to basic studies on the anatomy and morphology of Dioscorea and

the other genera of the yam family. At the age of 81 he completed

an account of the family Dioscoreaceae for Flora Malesiana. Finally.

at the age of 90, he completed a remarkable survey of the whole

family, treating it from many viewpoints (Journ. Linn. Soc. Bot. 56:

319-412, 1960).

70 Gardens’ Bulletin, Singapore — XXII (1967)

The historical studies on which the Dictionary was based, and

those on the dispersal and uses of yams, led Burkill to a general

interest in the history of food plants and so to a wide reading of

literature on early human history and pre-history. In his usual way

he accumulated a large number of reference slips dealing with

all aspects of this subject, and used them in preparing his Hooker

Lecture to the Linnean Society in 1951 on “Habits of Man and

the origins of the cultivated plants of the Old World’’. He had

thoughts of expanding this study into a book, but was obliged to

give up the idea.

About 1950 Burkill was asked if he would write something on the

history of botany in India, a subject on which he had already a

wide knowledge. He set about the work with his usual thorough-

ness, and went to much trouble in searching for biographical details

from many sources. His last visit to Kew was in connection with

this work, which was interrupted by a serious illness, after which

he received much help from his wife in bringing this final labour

to a conclusion. It was published, as ““Chapters on the History of

Botany in India’’, in the Journal of the Bombay Natural History

Society, from 1953 to 1963, and is now re-published in book form

by the Botanical Survey of India.

ISAAC HENRY BURKILL 1870-1965

A Bibliography

H. M. Burkitt, filius

The following lists are by way of supplementing the notes

published in the Gardens’ Bulletin, Singapore 18 (3), 1960, and

in this issue on the life of I. H. Burkill. His published papers total

over 7,700 printed pages. Burkillian taxa number 2 genera, 7

sections, 233 species, 78 varieties and 72 new combinations. The

extent of his collaboration with his friend and colleague, Sir

David Prain can be seen from these lists. Joint publications amount

to 768 printed papes, the first in 1904 and the last in 1938, and

113 new species and 55 new varieties.

I am indebted to Dr. C. G. G. J. van Steenis, Director, and Mr.

L. Vogelgang, Librarian, of the Rijksherbarium, Leiden, to Dr. R.

E. Holttum of Kew, and to Father H. Santapau, Director of the

Botanical Survey of India, who have helped me with additions and

corrections.

T2 Gardens’ Bulletin, Singapore — XXII (1967)

I. PUBLICATIONS

1892

‘Gloucestershire Rubi. J. Bot., 30, 1892: 376.

1893

‘The Flora of Gloucestershire, Reports of Cheltenham Nat. Sci. Soc., 1893,

57-67. [Being the substance of a lecture.]

Reprinted in Cheltenham Examiner, 10 and 17 May, 1893.

‘Observations on the Flora of the pollard willows near Cambridge.

Proc. Camb. Phil. Soc., 8, 1893: 82-91. (J. C. Willis and I.H.B.)

‘Notes on the plants distributed by the Cambridge dustcarts.

Proc. Camb. Phil. Soc., 8, 1893: 91-94.

1894

Botanical notes from north Cardiganshire.

J. Bot., 1894; 1-7. (1.H.B. and J. C. Willis.)

‘On the fertilisation of some species of Medicago L. in England.

Proc. Camb. Phil. Soc., 8, 1894: 141-52.

1895

‘On some variation in the number of stamens and carpels.

J. Linn. Soc. Lond., Bot., 31, 1895: 216-45.

Flowers and Insects in Great Britain. I.

Ann. Bot., 9, 1895: 227-73. (J. C. Willis and I.H.B.)

1896

‘On a collection of plants from New Britain (Neu Pommern).

Proc. Camb. Phil. Soc., 9, 1896: 91-98.

‘Teratological observations on Parnassia palustris, L.

J. Bot., 1896: 1-4

1897

Fertilisation of spring flowers on the Yorkshire coast.

J. Bot., 1897: 92-99; 138-45; 184-9.

List of the known plants occurring in British Central Africa.

In JOHNSTON: British Central Africa, London, 1897, Bot. App., II:

233-84.

1898

‘Changes in the sex of willows.

Ann. Bot., 12, 1898: 557.

[Abstract of a paper read to the British Association for the Advancement

of Science, Section K, meeting of 1898 — vide 1899.]

Pittosporum spathaceum.

Hook. Ic. Pl., 26 (3), 1898: t.2561.

1899

Changes in the sex of willows.

Rep. 68 Meeting Brit. Ass. Adv. Sci., 1898. 1065-6 (Abstract).

Botany [of Gloucestershire. ]

In BEACON C.W.: Gloucester Court Guide and County Blue Book,

1899: 156-82.

Burkill, f. — Burkill, 1870-1965, a bibliography 73

‘Limacia monilifera.

Hook. Ic. Pl., 26 (4), 1899: t. 2585.

On Pelargonium rapaceum, Jacq.

Ann. Bot., 13, 1899: 181-2.

Acanthaceae.

In THISELTON-DYER: Flora of Tropical Africa, 5: 1-44.

Notes on plants collected in the Aconcagua valleys by Phillip GOSSE.

In FITZGERALD, E.A.: The Highest Andes, London 1899, Botany,

pp. 361-79.

Cupuliferae: Betuleae.

In FORBES, F.B. & W.B. HEMSLEY: An enumeration of all the

plants known from China Proper, Formosa, Hainan, Corea, The Luchu

Archipelago and the Island of Hongkong, together with their dis-

tribution and synonymy.

J. Linn. Soc. Lond., Bot. 26, 1899: 496-500.

Cupuliferae: Coryleae.

In FORBES, F.B. & W.B. HEMSLEY: op. cit.

J. Linn. Soc. Lond., Bot. 26, 1899: 501-5.

Salicaceae.

In FORBES, F.B. & W.B. HEMSLEY: op. cit.

J. Linn. Soc. Lond., Bot. 26, 1899: 526-38.

On some African Labiatae with alternate leaves.

J. Linn. Soc. Lond., Bot. 34, 1899: 268-76.

(.H.B. and C.H. Wright.)

1900

Trifolium pratense var. parviflorum. .

Proc. Camb. Phil. Soc., 11, 1900: 29-31. (Reprinted in J. Bot., 1901::

235-6.)

Dolicholobium acuminatum. .

Hook. Ic. Pl., 27 (2), 1900: t. 2630.

1901

The ovary of Parnassia palustris, Linn.

Ann. Bot., 15, 1901: 187-92.

Ground-nut, or pea-nut (Arachis hypogoea, Linn.).

Kew Bull., 1901: 175-200.

The Flora of Vavau, one of the Tonga Islands.

J. Linn. Soc. Lond., Bot. 35, 1901: 20-65. (With a short account of

its vegetation by C. S. Crosby.)

Report on two botanical collections from Mt. Roraima, Introduction.

Trans. Linn. Soc. Lind., Tl, 6, 1901: 1-17.

The phanerogamic flora of the Clova Mountains in special relation:

to flower biology.

Trans. Bot. Soc. Edin., 66, 1901: 109-25.

(J. C. Willis and I.H.B.)

1902

On the variation of the flower of Ranunculus arvensis.

J. As. Soc. Beng., 71, 1902: 93-120.

Paspalum dilatatum, an American fodder grass.

Kew Bull., 1902: 1-4.

Ginseng in China (Aralia quinquefolia).

Kew Bull. 1902: 4-11.

74 Gardens’ Bulletin, Singapore — XXII (1967)

Aleurites cordata, Steud. Tung Oil or Chinese Wood Oil.

Commercial Circ. No. 2 of 1902-3, Calcutta, 1902. [Issued by the office

of the Reporter on Economic Products to the Government of India.]

Annual Report of the Industrial Section of the Indian Museum for the year

1901-1902; 34 pp., Calcutta.

1903

Irregular casings of the Faggot Worm.

Proc. As. Soc. Beng., 1903: 97-98.

The diffusion process of extracting Sugar from Sugar Cane.

Agric. Ledger 1903, 10 (8): 193-213. Calcutta, 1903.

Flowers and Insects in Great Britain. II. Observations on the natural

orders Dipsacaceae, Plumbaginaceae, Compositae, Umbelliferae, and

Cornaceae, made in the Clova Mountains.

Ann. Bot. 17, 1903: 313-49. (J. C. Willis and I.H.B.)

Flowers and Insects in Great Britain. III. Observations on the most

specialised flowers of the Clova Mountains.

Ann. Bot. 17, 1903: 539-70. (J. C. Willis and I.H.B.)

Notes on anthophilous insect fauna of the Clova Mountains.

Ann. Scot. Nat. His., 1903: 29-37, 99-103.

(J. C. Willis and I.H.B.)

Report and Programme, 1903. Collections for the Indian Museum, Calcutta,

and the Imperial Institute, London. Calcutta, 1903, 34 pp.

Annual Report of the Industrial Section of the Indian Museum for the year

1902-1903; 40 pp., Calcutta.

1904

A note upon the use of Marsdenia in the Rajmahal Hills.

Agric. Ledger 1904, 11 (8): 113-14. Calucutta, 1904.

The Tapioca plant: its history, cultivation and uses. A review of existing

information.

Agric Ledger 1904, 11 (10): 123-48. Calcutta, 1904.

Annatto Dye. A review of existing information.

Agric. Ledger 1904, 11 (12): 177-87. Calcutta, 1904.

Notes on the anthophilous insect fauna of the Clova Mountains. II.

Ann, Scot. Nat. His., 1904: 166-8.

(J. C. Willis and J.H.B.)

A report on cotton in Burma.

Calcutta, 1904. [Issued from the office of the Reporter on Economic

Products to the Government of India.]

On Dioscorea burmanica—a new species from Burma—and two allied

species.

J. As. Soc. Beng. II, 73, 1904: 183-7.

(D. Prain and I.H.B.)

The Central Factory System, with reference especially to countries growing

sugar cane.

Agric. Ledger 1903, 11 (12): 245-60. Calcutta, 1904.

(I1.H.B. and J. Weinberg).

On Dioscorea deltoidea, Wall., Dioscorea quinqueloba, Thunb., and their

allies.

J. As. Soc. Beng., Tl, 73 Suppl., 1904: 1-11.

(D. Prain and I.H.B.)

Burkill, f. — Burkill, 1870-1965, a bibliography 75:

1905

Frankincense, Myrrh and Bdellium.

Note No. 157 of the Reporter on Economic Products to the Government.

of India. Calcutta, 1905. 12 pp.

The collection and composition of the dye-stuff Kamala.

Agric. Ledger 1905, No. 4: 49-70. Calcutta, 1905.

1906

A treatment for hydrophobia.

Ind. For., 32, 1906: 27.

A parasite upon a parasite—a Viscum, apparently V. articulatum, Burm...

on Loranthus vestitus, Wall., on Quercus incana Roxb.

J. As. Soc. Beng., ns., 2, 1906: 299-301.

On Swertia angustifolia, Ham., and its allies.

J. As. Soc. Beng., ns., 2, 1906: 363-81.

Gentianacearum species asiaticus novas descripsit.

J. As. Soc. Beng., ns. 2, 1906: 309-27.

Notes on the pollination of flowers in India. I. The pollination of

Thunbergia grandiflora, Roxb., in Calcutta.

J. As. Soc. Beng., ns. 2, 1906: 511-14

Notes on the pollination of flowers in India, II. The pollination of

Corchorus in Bengal and Assam.

J. As. Soc. Beng., ns. 2, 1906: 515-20.

Notes on the pollination of flowers in India, III.

The mechanism of six flowers of the northwest Himalaya.

J. As. Soc. Beng., ns. 2, 1906: 521-S.

The fertilisation of Pieris.

Nature, July 26, 1906.

Bambarra ground-nut (Voandzeia subterrranea, Thou.)

Kew Bull., 1906: 68-70.

Goa Beans in India.

Agric. Ledger 1906, 13 (4): 51-64. Calcutta, 1906.

Gossypium obtusifolium, Roxb.

Mem. Dep. Agr. India, Bot. ser., 1906, 1 (4): 1-10.

Swertiam novam japonicam ex affinitate Swertiae tetrapterae.

J. As. Soc. Beng., ns. 2, 1906: 329.

(S. Le M. Moore and I.H.B.)

Annual Report of the Industrial Section of the Indian Museum for the:

year 1905-1906; 39 pp., Calcutta.

1907

Alpine Notes from Sikkim.

Kew Bull., 1907: 92-94.

A note on Swertia tongluensis and on a new variety of Swertia purpurascens.

J. As. Soc. Beng., n.s., 3, 1907: 33-35.

On Gentiana coronata, Royle.

J. As. Soc. Beng., n.s., 3, 1907: 149-68.

A fish trap from northern Arakan.

J. As. Soc. Beng., ns., 3, 1907: 441-2.

76 Gardens’ Bulletin, Singapore — XXII (1966)

Notes on the pollination of flowers in India, IV. On Cotton in Behar.

J. As. Soc. Beng., ns. 3, 1907: 517-26.

Anguillicarpus—a new genus of Cruciferae.

J. As. Soc. Beng., NS» 3, 1907: 559-61.

A variety of Ducrosia anethifolia, Boiss., from Baluchistan.

J. As. Soc. Beng, ns. 3, 1907: 563-4.

A note on Impatiens balsamina, Linn., as a dye plant.

J. As. Soc. Beng, ns., 3, 1907: 565-6.

An abnormal branch of the mango (Mangifera indica, Linn.).

J. As. Soc. Beng., n8., 3, 1907: 427-32.

(I.H.B. and G. C. Bose.)

‘On three varieties of Corchorus capsuaris, Linn., which are eaten.

J. As. Soc. Beng., ns. 3, 1907: 633-8.

(1.H.B. and R.S. Finlow.)

Annual Report of the Industrial Section of the Indian Museum for the

year 1906-1907; 22 pp., Calcutta.

1908

‘Notes on the pollination of flowers in India, V. Some autumn observations

in the Sikkim Himalaya.

J. As. Soc. Beng., nS. 4, 1908: 179-95.

“Notes on the pollination of flowers in India, VI. The spring flora in the

Simla Hills.

J. As. Soc. Beng., ns., 4, 1908: 197-231.

“The Indian Doum (Hyphaene) Palm.

J. Bomb. Nat. Hist. Soc. 18, 1908: 929-30.

_A review of information on Burmese peas and beans so far collected in

the office of the Reporter on Economic Products to the Government of

India. Calcutta, 1908.

Dioscorearum novarum descriptiones quaedam.

J. As. Soc. Beng., ns. 4, 1908: 447-57.

(D. Prain and JI.H.B.)

Melchior Treub. [A biographic note on the occasion of his election to

honorary membership of the As. Soc. Beng.]

Proc. As. Soc. Beng., 1908: 1xxxviii—lxxxix.

“Rai Ram Brahma Sanyal Bahadur. [Obituary.]

Proc. As Soc. Beng., 1908: cxiii.

“The Races of Jute.

Agr. Ledger 1907, 14 (6): 41-137. Calcutta, 1908.

(1.H.B. and R.S. Finlow.)

Flowers and Insects in Great Britain, IV. Observations on the less

specialized flowers of the Clova Mountains.

Ann. Bot., 22, 1908: 603-49. (J. C. Willis and I.H.B.)

Annual Report of the Industrial Section of the Indian Museum for the

year 1907-1908; 32 pp., Calcutta.

1909

First notes on Cymbopogon martini, Stapf.

J. As. Soc. Beng., ns. 5, 1909: 89-93.

The drug Astukhudus, nowadays Lavandula dentata, and not Lavandula

stoechas.

J. As. Soc. Beng., n.s., 5, 1909: 67-71.

On Coptis.

J. As. Soc. Beng., ns. 5, 1909: 73-88.

Burkill, f. — Burkill, 1870-1965, a bibliography aT

Dyes’ from Flowers. A summary of our present knowledge regarding the

use of dyes from flowers in India, together with two reports on Thespesia’

lampas and Hibiscus subdariffa by A. G. Perkin.

Agr. Ledger 1908, 15 (2): 7-27, 1909.

A Working list of the flowering plants of Baluchistan.

Calcutta, 1909, 136 pp.

Indian pens, their history, classification, materials used and methods of

manufacture.

Agr. Ledger, 1908-9, 15 (6): 111-30, 1909.

Note on the extension of cultivation of fibre plants in India.

Agr. Res. Inst., Pusa, Bull., 15, 1909: 1-14.

(I.H.B. et al.)

Soy Bean.

Ind. Trade J., July 29, 1909: 136-8.

Annual Report of the Industrial Section of the Indian Museum for the

year 1908-1909; 25 pp., Calcutta.

1910

Fashion in Iron Styles

J. As. Soc. Beng., n.s.,.6, 1910: 1-18.

Notes on the pollination of flowers in India, VII. A few observations made

in the Central Provinces and Berar.

J. As. Soc. Beng., ns. 6, 1910: 101-7.

Notes on the spreading of Croton sparsiflorus, Morung, along the Assam--

Bengal Railway.

Proc. As. Soc. Beng., 1910: ci-cii.

Notes from a journey to Nepal.

Rec. Bot. Surv. Ind., 4, 1910: 59-140 + index, 3 pp.

Bat and Bird Guanos in India.

Agric. Ledger 1911, 17 (4): 1-9, 1910.

Edible Frogs in India.

Agric. Ledger 1911, 17 (2): 11-15, 1910.

Report and Programme of the Indian Museum, Calcutta, for the year

1910. Calcutta, 1910.

Industrial and Agricultural Chemistry. Report of the Board of Scientific

Advice for 1908-09: Government of India. Calcutta, 1910. |

(.H.B. and H. E. Annet.)

Annual Report of the Industrial Section of the Indian Museum tor the

year 1909-1910; 52 pp., Calcutta.

1911

Determinations of the Prickly Pears now wild in India.

Rec. Bot. Surv. Ind., 4 (6), 1911: 287-322.

Swertias chinenses quatuor novas ex herbario- G. Bonati descripsit.

J. As. Soc. Beng., ns. 7, 1911: 81-82.

Further spreading of Croton sparsiflorus (Morung).

¥. As. Soc; Beng:; ns. 7,°1911: 132

78 Gardens’ Bulletin, Singapore — XXII (1967)

The polarity of the bulbils of Dioscorea bulbifera, Linn.

J. As. Soc. Beng., n.s.. 7, 1911: 467-9.

A Lepcha’s medicine bag.

Proc. As. Soc. Beng, 191i: xcix-xc.

Corchorus capsularis var. oocarpus—a new variety of common jute plant.

J. As. Soc. Beng., n.8.. 7, 1911: 465-6.

(I.H.B. and R. S. Finlow.)

Annual Report of the Industrial Section of the Indian Museum for the

year 1910-1911; 21 pp., Calcutta.

1912

Croton sparsiflorus (Morung) from Makum.

Proc. As. Soc. Beng., 1912: cxi.

Abor Expedition.

Kew Bull., 1912: 159. [Being an abstract of a letter.]

‘The inheritance of red colour, and the regularity of self-fertilisation, in

Corchorus capsularis, Linn.—the common jute plant.

Mem. Dept. Agr. in Ind., Bot. ser., 4, 1912: 73-92.

(R. S. Finlow and I.H.B.)

1913

‘The botany of the Abor expedition. A study of the forests of the Abor

Hills, eastern Himalayas.

Rep. 82 Meeting Brit. Ass. Adv. Sci., Dundee, 1912: pp. 683-4.

{Being the abstract of a paper read to the British Association for the

Advancement of Science, Section K, Dundee 1912.]

Dioscoreae Elmerianae. A contribution to our knowledge of the genus

Dioscorea in the Philippine Islands.

Elmer, Leafl. Philip. Bot., 5, 1913: 1589-99.

(D. Prain and J.H.B.)

The Coconut Beetles, Oryctes rhinoceros and Rhyncophorus ferrugineus.

Gard. Bull. S.S., 1, 1913: 176-188.

Clerome gracilis, a butterfly destructive to palms.

Gard. Bull. S.S., 1, 1913: 188-9.

Notes on plants of interest in the Botanic Gardens, Singapore.

Gard. Bull. S.S., 1, 1913: 189-92.

‘The extreme hardness of the seeds of Caesalpinia digyna.

Gard. Bulli \SS.,. 0554013 : 03:

A disease of Agaves.

Gard. Bull. S.S., 1, 1913: 193-4.

Annual Report on the Botanic Gardens, Singapore and Penang, for the

year 19172.

Government Printing Office, Singapore, 1913, 8 pp.

1914

The Sirangoon outbreak (1913) of Brachartona catoxantha.

Gard. Bull. S.S., 1, 1914: 207-8.

Calogramma festiva, Donovan,—a caterpillar destructive to Crinum

asiaticum.

Gard. Bull. S.S., 1, 1914: 209.

Burkill, f. — Burkill, 1870-1965, a bibliography 79

The big Rubber Tree, Singapore.

Gard. Bull. S.S., 1, 1914: 210-11.

Croton sparsiflorus, (Morung), an American invader.

Gard. Bull. S.S., 1, 1914: 235-7.

A spadix in the axil of the spathe of Xanthosoma.

Gard. Bull. S.S., 1, 1914: 244.

Grammatophyllum flowering in January.

Gard. Bull. S.S., 1, 1914: 244.

A synopsis of the Dioscoreas of the Old World, Africa excluded, with

descriptions of new species and of varieties.

J. As. Soc. Beng., ns. 10, 1914: 5-41.

Annual Report on the Botanic Gardens, Singapore and Penang, for the

year 1913.

Government Printing Office, Singapore, 1914, 8 pp.

1915

An abnormality in the coconut palm.

i. Str. Br..R. As. Soc;.68, 19154 15.

The Treatment to which the para-rubber trees of the Botanic Gardens,

Singapore, have been subjected.

Gard. Bull. S.S., 1, 1915: 247-95.

Some notes on philippine yams, and the results of cultivation of three

indian races in 1914.

Gard. Bull. S.S., 1, 1915: 297-304.

Preparation of yams for the table.

Gard. Bull. S.S., 1, 1915: 304—S.

Different parts of the tubers of Dioscorea alata sprout at different rates.

Gard. Bull. S.S., 1, 1915: 306-7.

Three Lepidoptera which attack Dioscoreas in Singapore.

Gard. Bull. S.S., 1, 1915: 308-10.

Fragments of malayan geographic botany.

I. Enumeration of Pahang plants collected by the late A. M. Burn

Murdoch.

Gard. Bull. S.S., 1, 1915: 310-18.

Orchid Notes.

Gard. Bull. S.S., 1, 1915: 318-20.

The Singapore Prickly-Pear.

Gard. Bull. S.S., 1, 1915: 320-1.

A nest of living Drymoglossum piloselloides.

Gard... Bull. S.8.; 1,.1915: 321.

A very destructive flash of lightning.

Gard. Bull. S.S., 1, 1915: 329.

A report on philippine yams. é

Phil. Agr. For., 3, 1915: 205-9.

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1914.

Government Printing Office, Singapore, 1915, 7 pp.

80 Gardens’ Bulletin, Singapore — X XII (1967)

1916

Notes on the pollination of flowers in India, VIII. Miscellanea.

J. As. Soc. Beng., n.s.. 12, 1916: 239-65.

A note on the Terai forests between Gandak and the Teesta.

J. As. Soc. Beng., n.8.. 12, 1916: 267-72.

William Jack’s letters to Nathaniel Wallich, 1819-1821.

J. Str. Br. R. As. Soc., 73, 1916: 147-268.

Selection.

Straits Times Press Ltd., Singapore 1916. [Being the substance of a

lecture given to the Malacca Planters’ Association.]

Orchid Notes.

Gard. Bull. S.S., 1, 1916: 349-53.

A skipper trapped in a flower of Dipladenia harrisii.

Gard. Bull? S.S., ¥, T9162 355.

Locusts in Malacca.

Gard. Bull. S.S., 1, 1916: 355-49.

(P. C. Cowley-Brown and I.H.B.)

Dioscorea asclepiadea, Pr. et Burk.; D. bernouilliana, Pr. et Burk.; D.-

carionis, Pr. et Burk.; D. melastomifolia, Uline, and D. truncata, Miq.

Kew Bull., 1916: 190-6. (D. Prain and J.H.B.)

Annual Report of the Director of Gardens, Straits Settlements, for the year

1915.

Government Printing Office, Singapore, 1916: 7 pp.

1917

Elaeocarpus barnardii, a new species described from Perak.

J. Str. Bronte: As. Secs, Tay 19173. 41.

Notes on Dipterocarps, I. The seedling of Anisoptera costata, Korth.

J. Str. Br. R. As. Soc., 75, 1917: 43-48.

Gordonia.

J. Str. Br. R.-As. Soc, 76,1917: 133-32:

Notes on Dipterocarps, II. The seedling and the seed production in some

species of Shorea.

J. Str. Br. R. As. Soe. FW, 1917: 161k

A note upon the way in which bees settle on flowers of Derris thyrsiflora,

and the injury resulting from their search of honey.

J. Str. Br. R. As. Soc, Ti, V917:. 263-4.

Plethiandra sahebii, a new Melastomacea from Sarawak, described.

J. Str. Br. R. As. Soe, Ti, 1917: 265-9,

Food-Crops of the Malay Peninsula and some thoughts arising out of a

review of them.

Proc. Agr. Conf. Kuala Lumpur, 25-28 April 1917, Kuala Lumpur,

1917: 16 pp.

Reprinted in Agr. Bull. F.M.S., 5, 1917: 406-21.

A report on races of the Greater or Ten-Months Yam,—Dioscorea alata—

cultivated in the Botanic Gardens, Singapore.

Gard. Bull. S.S., 1, 1917: 371-96.

The Lesser Yam,—Dioscorea esculenta.

Gard. Bull. S.S., 1, 1917: 396-9.

Burkill, f. — Burkill, 1870-1965, a bibliography 81

Scolia erratica, Smith, a parasite of the Red Coconut Weevil (Rhynco-

phorus ferrugineus).

Gard. Bull. S.S., 1, 1917: 399-400.

The flowering of the Pigeon Orchid, Dendrobium crumenatum, Lindl.

Gard. Bull. S.S., 1, 1917: 400-S.

Precocious flowering of Melia.

Gard. Buil. S.S,,; 1,1: 191%: 306.

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1916.

Government Printing Office, Singapore, 1917, 8 pp.

1918

The circumstances attending the murder in 1859 of the botanist. James

Motley.

#, Sir. Br, RM. As. Soe. 79,1918 : <37=38,

Notes on Dipterocarps, III. The seedling of Shorea robusta, Roxb., and the

conditions under which it grows in pure forests.

J. Str. Br. R. As. Soc., 79, 1918: 39-44.

A new Dendrobium, D. gracilipes, from the Rhio Archipelago.

J. Str. Br. R. As. Soc., 79, 1918: 45-46.

Begonia haniffii, a small tuberous species of the Islands of Langkawi.

I. ttre Br. SR. AS. 5806079, 1918- 103-4,

Catochrysops pandara, a butterfly destructive to cycads.

Gard. Bull. S.S., 2, 1918: 1-2.

Promecotheca cumingii Baly, another coconut hispid and a pest in Malacca.

Gard. Bull.S.S.,.2,.1918: 3-5.

A beetle which attacks yams.

Gard. Bull. S.S.,°2, 1918: 6.

A report upon the experimental cultivation of the Greater Yam—Dioscorea

alata—in 1917.

Gard. Bull. S.S., 2, 1918: 37-44.

Orchid Notes.

Gard. Bull. S.S., 2, 1918: 44-51.

Propagation of Hevea from stakes.

Gard. Bull. S.S., 2, 1918: 54.

The establishment of the Botanic Gardens, Singapore.

Gard. Buill,.S.S., 2, 1918: 55-72.

The Rozelle—Hibiscus sabdariffa.

Gard. Bull. S.S., 2, 1918: 73.

Some cultivated yams from Africa, and elsewhere.

Gard. Bull. S.S., 2, 1918: 86-92.

The second phase in the history of the Botanic Gardens, Singapore.

Gard. Bull. S.S., 2, 1918: 93-108.

Notes on Cola trees in the Economic Gardens, Singapore.

Gard. Bull. S.S., 2, 1918: 74-86. |

Annual Report of the Director of Gardens, Straits Settlements, for the year

1917.

Government Printing Office, Singapore, 1918: 8 pp.

82 Gardens’ Bulletin, Singapore — XXII (1967)

1919

The Gardens’ Hevea tree No. 1844—H. confusa, Hemsl.

Gard. Bull. S.S., 2, 1919: 113-15.

A progress report on the cultivation of the Greater Yam,

Dioscorea alata, in the Botanic Gardens, Singapore.

Gard. Bull. S.S., 2, 1919: 129-35.

It needs want to make people change their food-habits.

Gard. Bull. S.S., 2, 1919: 135-6.

Lightning and Hevea.

Gard. Bull. S.S., 2, 1919: 145.

The composition of a piece of well-drained Singapore secondary jungle

thirty years old.

Gard. Bull. S.S., 2, 1919: 145-57.

Twin Nutmeg Seeds.

Gard. Bull. S.S., 2, 1919: 158.

Dioscorea alata, The Greater Yam, Race No. SO.

Gard. Bull. S.S., 2, 1919: 158.

Dioscorea kegeliana, Griseb., the “Yam Poule” of the West Indies.

Gard. Bull. S.S., 2, 1919: 158.

Yields of the Lesser Yam and of some african yams.

Gard. Bull. S.S., 2, 1919: 159-65.

Some notes on the pollination of flowers in the Botanic Gardens, Singapore,

and in other parts of the Malay Peninsula.

Gard. Bull. S:,.2; T1919. 165-76.

The genus Gordonia in the Philippine Islands.

Philip J. Sci., 15, 1919: 475-8.

Dioscorea sativa.

Kew Bull., 1919: 339-75. (D. Prain and I.H.B.)

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1918.

Government Printing Office, Singapore, 1919, 7 pp.

1920

Notes on Dipterocarps, IV. On the embryo, seedling and positicn of the

flower in various species.

J. Str. Br. R. As. Soc Ot, 1920: 49-76.

Notes on Dipterocarps, V. The embryo and seedling of Balanocarpus

maximus, King.

J. Sir: Br: R. As: Soc: ee, 1920: ‘3-4.

Staking Yams.

Gard. Bull. S.S., 2, 1920: 301-2.

1921

Forests and their retention of rain water.

Gard. Bull. S.S., 2, 1921: 419-21.

[Being mainly a résumé of ENGLER, A.: Untersuchugen iiber den

Einfluss des Waldes auf den Stand der Gewédsser; Mitteil. Schweiz

Zentralanst. Forst. Versuchsw., 12, 1919: 1-626.]

A note upon plants grown for blue dyes in the north of the Malay

Peninsula.

Gard. Bull. S.S., 2, 1921: 426-9.

Burkill, f. — Burkill, 1870-1965, a bibliography 83:

A melon fly, Bactrocera cucurbitae.

Gard. Bull. S.S., 2, 1921: 432.

The correct botanic names for the White and the Yellow Guinea Yams..

Gard. Bull. S.S., 2, 1921: 438-41.

Hibiscus sabdariffa var. altissima.

Gard. Bull. S.S., 2, 1921: 441.

Orchid Notes.

Gard. Bull. S.S., 2, 1921: 441-4.

| The present state of our knowledge of the vegetation of Malaya.

Sing. Naturalist: 1, 1921: 23-24.

[Being a précis of a paper read to the Singapore Natural History Society

at the Society’s inaugural meeting on 8th August 1921.]

Odoardo Beccari.

J. Str. Br. R. As. Soc., 83, 1921: 166-73.

(I1.H.B. and J. C. Moulton.)

Annual Report of the Director of Gardens, Straits Settlements for the year

1919

Government Printing Office, Singapore, 1921, 5 pp.

Annual Report of the Director of Gardens, Straits Settlements, for the year

1920.

Government Printing Office, Singapore, 1921, 6 pp.

1922

The irregularity of a spider’s feeding.

J, Str. .Br. K.. As. 50e... 86, 1922>. 2/0.

Notes on Dipterocarps, VI. On the genus Pachynocarpus.

J. Str. Br. R. As. Soc., 86,-1922: 271-80

(.H.B. and F. W. Foxworthy.)

Notes on Dipterocarps, VII. On the fruit and germination of Jsoptera

borneensis.

J. Str. Br. R. As. Soc., 86, 1922: 281-4.

Notes on Dipterocarps, VIII. On some large-fruited species and in

particular upon the effects of the pressure of the embryo against the

interior of the fruit wall.

J. Str. Br. R. As. Soc., 86, 1922: 285-91.

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1921.

Government Printing Office, Singapore, 1922, 5 pp.

The fertility of branched coconut palms.

Gard. Bull. S.S., 3, 1923: 1-2.

Branching in Arenga pinnata.

arard. Bull. S.S.,°3, 1929s''2.

A spiny yam from Sumatra.

Gard. Bull. S.S., 3, 1923: 3-4.

Tahitian Yams.

Gard. Bull. S.S., 3, 1923: 4-5.

Yams at the Malaya-Borneo Exhibition.

Gard. Bull. S.S., 3, 1923: 5-8.

84 Gardens’ Bulletin, Singapore — X XII (1966)

The as-yet botanically unexplored parts of the Malay Peninsula.

Gard. Bull. S.S., 3, 1923: 8-10.

Orchid notes.

Gard. Bull. S.S., 3, 1923: 12-18.

Haplochorema sumatranum.

Gard. Bull. S.S., 3, 1923: 18-19.

A record of the occurrence of some ferns in northern Sumatra, being ad-

ditions to Mr. Ridley’s list.

J. Mal. Br. R. As. Soc., 1, 1923: 114-15.

‘Notes on Dipterocarps, IX. On the differences in the seedlings between

Balanocarpus maximus, King, and B. heimii, King.

J. Mal. Br. R. As. Soc., 1, 1923: 218-22.

A botanical reconnaissance upon the main range of the Peninsula at

Fraser’s Hill.

Gard. Bull. S.S., 3, 1923: 19-110.

(.H.B. and R. E. Holttum).

The historical aspect of certain local industries.

Sing. Naturalist, 2, 1923: 34-44.

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1922.

Government Printing Office, Singapore, 1923, 6 pp.

1924

A list of oriental vernacular names of the genus Dioscorea.

Gard. Bull. S.S., 3, 1924: 121-244.

Two hybrid trees of Hevea brasiliensis x H. confusa.

Gard. Bull. S.S., 3, 1924: 257-8.

Varieties of Dioscorea pentaphylla in Malaysia.

Gard. Bull. S.S., 3, 1924: 258-9.

A Chinese belief regarding Phyllocactus hookeri, Walp.

Gard. Bull. S.S., 3, 1924: 280.

Cleome chelidonii, Linn. f., in the Malay Peninsula.

Gard. Bull. S.S., 3, 1924: 280-1.

Stenomeris in the Malay Peninsula.

Gard. Bull. S.S., 3, 1924: 289-90.

The floras of the Malay Peninsula, Borneo and the Philippine Islands.

Gard. Bull. S.S., 3, 1924: 283-9.

Orchid notes.

Gard. Bull. S.S., 3, 1924: 292-6.

Dioscorea piscatorum, or Tuba-ubi, a fish poison.

Gard. Bull. S.S., 3, 1924: 260. (I.LH.B. and R. E. Hottum.)

The Botany of the Abor Expedition.

Rec. Bot. Surv. Ind., 10: 1924: 1-154; 1925: 155-420, Pt. I-X.

Annual Report of the Director of Gardens, Straits Settlements, for the

year 1923.

Government Printing Office, Singapore, 1924: 5 pp.

Burkill, f. — Burkill, 1870-1965, a bibliography 85:

1925

Notes on Dipterocarps, X. On Balanocarpus hemsleyanus, King.

J. Mal. Br. R. As. Soc., 3, 1925: 4-9.

Solanum macrocarpum.

Kew Bull., 1925: 333-41.

The Botanic Gardens, Singapore.

J. Kew Guild, 4, 1925: 313-8.

Insect vision in connection with flower fertilisation.

Sing. Naturalist, 5, 1925: 23-46.

The possibility of adding new species to the known flora of the Malay.

Peninsula.

Sing. Naturalist, 5, 1925: 57-61.

(I1.H.B. and R. E. Holttum.)

The flowering plants of Taiping in the Malay Peninsula.

Gard. Bull. S.S., 3, 1925: 303-458.

(I.H.B. and M. R. Henderson.)

Diagnoses specierum novarum generis Dioscoreae.

Kew Bull., 1925: 58-66. (D. Prain and I.H.B.)

Annual Report of the Director of Gardens, Straits Settlements, for the:

year 1924.

Government Printing Office, Singapore, 1925, 6 pp.

1926

The Botanic Gardens, Singaore. Illustrated Guide.

Waterlow & Sons, London, 1926, 67 pp.

Island occurrence of Ipomoea pes-caprae.

Kew Bull., 1926: 425-6.

Ad Dioscorearum orientalium historiam commentarii.

Kew Bull., 1926: 118-20. (D. Prain and I.H.B.)

Vegetation on lava surfaces of various ages in the crater of Kilauea, Hawaii:.

Proc. Linn. Soc. Lond., 138, 1926: 53-54.

1927

A note relating to the history of The Dell in the Gardens.

Gard. Bull. S.S., 4, 1927: 69-77.

Tulang Daing or Sisek Puyuh, Carallia suffruticosa.

Gard. Bull. S.S., 4, 1927: 77-78.

Botanical collectors, collections and collecting places in the Malay

Peninsula.

Gard. Bull. S.S., 4, 1927: 113-202.

Dioscorea tamarisciflora, Pr. and Burk.

Gard. Bull. S.S., 4, 1927: 86-88. (D. Prain and I.H.B.)

The genus Dioscorea in Siam. ,

Kew Bull., 1927: 225-47. (D. Prain and I.H.B.)

Insect vision.

London Naturalist for 1926, 1927: 3-6.

86 Gardens’ Bulletin, Singapore — XXII (1967)

1928

The main features of the vegetation of Pahang.

Malayan Naturalist, 2, 1928: 11-21.

1929

Anisophyllea gaudichaudiana, Baill., is A. grandis, Benth.

Gard. Bull. S.S., 4, 1929: 426.

1930

The chinese mustards in the Malay Peninsula.

Gard. Bull. S.S., 5, 1930: 99-117.

Cosmos in the East.

Gard. Bull. S.S., 5, 1930: 118-20.

Cedrela in the Malay Peninsula.

Gara! Bull S.S., 5501930: 120-2.

Blainvillea, Cass.

Gard. Bul. SS: 3, 19302 123.

Cricket fighting.

J. Siam Soc. Nat. Hist. Suppl., 8, 1930: 119-20.

‘Cardamoms.

Kew Bull., 1930: 32-37.

Malay Village Medicine.

Gard. Bull. S.S., 6, 1930: 165-321. [Reprinted 1960.]

(I.H.B. and Mohamed Haniff.)

“The medical book of Malayan Medicine”. Translated by Inche Ismail,

Munshi, possibly in Penang, circa 1886.’ Edited with medical notes and

determinations.

Gard. Bull. S.S., 6, 1930: 323-474. [Reprinted 1960.]

(J. G. Gimlette and I.H.B.)

On Dioscorea gibbiflora, Hook. fil.

Gard. Bull. S.S., 5, 1930: 51-57.

(D. Prain and J.H.B.)

1931

An enumeration of the species of Paramignya, Atalantia and Citrus found in

Malaya.

Gard.. Bull. SiS.4\ 593 212-23.

Notes on Gluta in Malaya.

Gard. Bull. S.S., 5, 1931: 224-30.

Herpestis monniera, H.B. and K., as pa-chi-t’ien.

Gard. Bull. S.S.,. Sy. A93L Aisi.

Erythropsis colorata.

Gard. Bull. S845, 193i ¢ 251.

Sir George Watt. [Obituary]

Proc. Linn. Soc. Lond. Sess. 142 (1929-30), 1931: 226-9.

Dioscorea: Section Stenocorea.

Kew Bull., 1931: 88-91. (D. Prain and I.H.B.)

Dioscoreae novae asiaticae.

Kew Bull., 1931: 425-7. (D. Prain and I.H.B.)

;

Burkill, f. — Burkill, 1870-1965, a bibliography 87

1932

Edward Arthur FitzGerald. [Obituary]

Proc. Linn. Soc. Lond. Sess. 143 (1930-31), 1932: 176.

An experiment upon the polarity of the tuber of Tamus communis Linn.

Proc. Linn. Soc. Lond., Sess. 144 (1931-32), 1932: Pt. III; 60-62.

1933

Dioscoreae novae asiaticae.

Kew Bull., 1933: 240-6. (D. Prain and I.H.B.)

1934

Dioscoréacées.

In LECOMTE, H., and H. HUMBERT, Flora générale de I’Indo-Chine,

Paris, 6, 1934: 698-745.

(D. Prain and I.H.B.)

1935

Some changes in plant names.

Kew Bull., 1935: 316-19.

A dictionary of the economic products of the Malay Peninsula.

OanP,, 1935, 2 vol., 2402 pp.

[Second edition in press, Kuala Lumpur, 1966.]

The bracteole in Scilla nutans, Sm., and extra organs in the flower.

J. Bot., 73, 1935: 280-6.

1936

The phyllotaxy of Dioscorea glauca, Muhl.

J. Bot., 74, 1936: 89-101.

The phyllotaxy of Tamus communis, L.

J. Bot., 74, 1936: 153-63.

Dioscorea garrettii, Pr. and Burk.: D. hainanensis, Pr. and Burk.: and

D. scortechinii, Pr. and Burk., var. parviflora.

Kew Bull., 1936: 493-5. (D. Prain and I.H.B.)

An account of the genus Dioscorea in the East, I. The species which twine

to the left.

Ann. R. Bot. Gard. Calc., 14, 1936: 1-210, pl. 1-85.

(D. Prain and I.H.B.)

1937

The Races of Sorghum.

Kew Bull., 1937: 112-19.

[Being a review and phytogeographic analysis of SNOWDEN, J.D.: The

Cultivated Races of Sorghum. London, 1936.]

The life-cycle of Tamus communis L.

J. Bot., 75, 1937: 1-12, 33-43, 65-74,,

Phil. Soc., 9, 1896: 91-98.

Cedric Erroll Carr. [Obituary]

J. Bot., 75, 1937: 143-4.

The development of the tuber of Dioscorea

sansibarensis Pax.

Blumea, Suppl. 1, 1937: 232-7.

88 Gardens’ Bulletin, Singapore — XXII (1967)

1938

‘Illustrated guide to the Trees and Flowers of England and Wales’ by

H. G. Jameson. [Review]

J- Bot., ‘76, 1938> 35.

Alfred Barton Rendle. [Obituary]

J. Bot., 76, 1938: 65-68.

‘Lebensgeschichte der Bliitenpflanzen Mitteleuropas’, vol. 2 (1).

[Review]

J. Bot., 76, 1938: 87-88.

The vision of certain small Ophrys-visiting bees.

J. Bot., 76, 1938: 150-1.

Stephen Troyte Dunn. [Obituary]

J. Bot., 76, 1938: 183-4.

An account of the genus Dioscorea in the East, II. The species which

twine to the right; with addenda to Part I, and a summary.

Ann. R. Bot. Gard. Calc., 14, 1938: 211-528, pl. 86-150, Index i-xx.

[Title page dated 1939.]

(D. Prain and I.H.B.)

The contact of the Portuguese with African food-plants which gave words

such as ‘yam’ to European languages.

Proc. Linn. Soc. Lond., Sess. 150 (1937-38), 1938; pt. If: 84-95, 2 maps.

1939

The trigger mechanism in the germination of the seed of Tamus communis,

J. Bot., 77, 1939: 44-50.

Growth and tensions between the nerves in the leaf-blade of Tamus

communis, L.

J. Bot. 971939) 325-33:

Notes on the genus Dioscorea in the Belgian Congo.

Bull. Jard. Bot. Etat, 15, 1939: 345-92.

Two notes on Dioscoreas in the Congo: (1) the acarodomatia of D.

minutiflora, Engl., and D. smilacifolia, de Wild., and (2) twining in both

directions in D. baya, de Wild.

Proc. Linn. Soc. Lond., 151 Sess. (1938-39), 1939: 57-61.

1940

The distribution of raphides in the leaves of Tamus communis, Linn.

J, Bot., 78, 19403" 17-19.

Slight zygomorphy in Dioscorea sylvatica Ecklon.

J. Bot., 78, 1940: 100-1.

Empis pennipes, Linn., on the flowers of Geranium robertianum.

J. Bot., 78, 1940: 175-6.

Varthema’s Corcopal, at one time said to be papaya, was the fruit of

Garcinia indica.

Proc. Linn. Soc. Lond., 152 Sess. (1939-40), 1940: 147-9.

Burkill, f. — Burkill, 1870-1965, a bibliography 89

1941

Duarte Barbosa’s references to trade at Malacca in Cutch, Costus and

Aleppa Galls.

J. Mal. Br. R. As. Soc., 19, 1941: 120-1.

The make-up of the flower of Ranunculus arvensis, Linn.,—a study in the

evolution of isomerism in phanerogamous flowers.

Proc. Linn. Soc. Lond., 153 Sess. (1940-41), 1941: 161-214.

An ontogenetic analysis of Ranunculus arvensis, Linn.

J: rot., 79, 1941: 121-31.

1942

The terrestrial vegetative buds of Dioscorea alata, Linn.

Ann. Bot. Gard. Calc., 150th Anniversary Volume, 1942: 109-14.

1943

The biogeographic division of the Indo-Australian Archipelago. A history

of the divisions which have been proposed.

Proc. Linn. Soc. Lond., 154 Sess. (1941-42), 1943: 127-38.

A leaf of Tamus communis, Linn. with two petioles as a result of an early

injury.

Proc. Linn. Soc. Lond., 155 Sess. (1942-43), 1943: 31-34.

1944

Biological Flora of the British Isles. Tamus.

J. Bot., 32, 1944: 121-9.

The way in which the leaves of Tamus communis and Dioscoreas attain

their light-fixed position.

Proc. Linn. Soc. Lond., 155 Sess. (1942-43), 1944: 252-77.

The early economic history of the tree, Mesua ferrea (Guttiferae).

Proc. Linn. Soc. Lond., 156 Sess. (1943-44), 1944: 85-91.

David Prain, 1857-1944. [Obituary]

Obit. Not. Fellows R. Soc. 4, 1944: 747-70.

1946

Abnormal gamopetalous flowers of the poppy, Romneya coulteri, and the

way in which its sepals protect the sexual organs.

Proc Linn. Soc. Lond., 157 Sess. (1944-45), 1946: 92-93.

Flies of the family Empididae and other insect-visitors to the flowers of

Tamus communis.

Proc Linn. Soc. Lond., 157 Sess. (1944-45), 1946: 92-102.

On the dispersal of the plants most intimate to Buddhism.

J. Arn. Arb., 27, 1946: 327-39.

[This volume is dedicated to Elmer Drew Merrill on the occasion of his

70th birthday, 15th October, 1946.]

1947

A for the description from life of the african Dioscorea minutiflora,

ngl.

Proc. Linn. Soc. Lond., 159 Sess. (1946-47), 1947: 77-81.

90 Gardens’ Bulletin, Singapore — XXII (1967)

1949

William Farquhar’s drawings of Malacca plants.

Gard. Bull. Sing., 12, 1949: 404-7.

The ontogeny of the stem of the Common Bryony, Tamus communis, Linn.

J. Linn. Soc. Lond., 53, 1949: 313-82.

An interruption in the after ripening of Tamus which alters the range

of temperature controlling germination.

Proc. Linn. Soc. Lond., 1961 Sess. (1948-49), 1949; 151-S.

James Hornell. [Obituary]

Proc. Linn. Soc. Lond., 161 Sess. (1948-49), 1949: 244-7.

1950

A new Dioscorea from Celebes.

Kew Bull., 1950: 259-60.

Dioscoréacées.

In HUMBERT: Flore de Madagascar, fam. 44: 1-78, 14 fig.

(I.H.B. and H. Perrier de la Bathie.) |

1951

Dioscoreaceae.

In VAN STEENIS, C.G.G.J.: Flora Malesiana, I, 4, 1951: 293-335.

The rise and decline of the Greater Yam in the service of man.

The Advancement of Science, 7 (28), 1951: 443-8. [Being the substance

of a paper read to Section H (Anthropology and Archeology) of the

meeting of the British Association for the Advancement of Science

held at Birmingham, September 1950.]

New units in the taxonomy of the madagascan Dioscoreaceae.

Mus. Natl. Hist. Nat. Not. Syst., 14, 1951: 131-7.

(I1.H.B. and H. Perrier de la Bathie.)

1952

Testudinaria as a section of the genus Dioscorea.

J..S. Afr. Bot., 18, 19522 477-91.

Mr. Cecil Ernest Claude Fisher, 1873-1950. [Obituary]

Ind. For. 78, 1952: 50.

1953

Vegetables eaten with rice.

Gard. Bull. Sing., 14, 1953: 17-29.

Habits of man and the origins of the cultivated plants of the Old World.

Proc. Linn. Soc. Lond., 164, Sess. 1951-52), 1953: 12-42.

[Hooker Lecture—Linnean Society of London, 1951.]

Chapters on the history of botany in India.

I. From the beginning to the middle of Wallich’s service.

J. Bomb. Nat. Hist. Soc., 51, 1953: 846-78.

Pepper, Nutmegs and Rubber, I.

Malaya, October 1953: 587-9.

Pepper, Nutmegs and Rubber, II.

Malaya, November 1953: 647-50.

Burkill, f. — Burkill, 1870-1965, a bibliography 91

1954

Aji and Batata as group names within the species Ipomoea batatas.

Ceiba, 4, 1954: 227-40.

1955

William Farquhar’s second book of drawings of Malacca plants.

Gard. Bull. Sing., 14, 1955: 530-3.

1956

Chapters on the history of botany in India.

II. The advances, and in particular the plant collecting, of the , Thirties

and Forties of the 19th century.

J. Bomb. Nat. Soc., 54, 1956: 42-86.

Prof. E. D. Merrill. [Obituary]

Nature, 177, April 14, 1956: 687-8.

1957

Elmer Drew Merrill. [Obituary]

Proc. Linn. Soc. Lond., 168 Sess. (1955-56), 1957: 51-56.

Elmer Drew Merrill, M.S., Hon. D. Sc., Hon. LI, D., Hon. F.R.S.E.

[Obituary]

Year Book R. Soc. Edin. 1955-56, 35-37.

1958

Inhibition of germination of the white mustard by bryony juice.

Proc. Linn. Soc. Lond., 169 Sess. (1956-57), 1958: 62-63.

The relics and offerings from a bronze bust of Tsong-Kha-Pa, founder

of the Lamaist Sect of the Yellow Hats.

J. Mal. Br. R. As. Soc., 31, 1958: 175-6. [Issued July 1965.]

Henry Nicholas Ridley. [Obituary]

Proc. Linn. Soc. Lond., 169 Sess. (1956-57), 1958: 35-38.

John Christopher Willis. [Obituary]

Proc. Linn. Soc. Lond., 169 Sess. (1956-57), 1958: 245-50.

1959

A note on the Gardens Jungle.

Gard. Bull. Sing. 17 (2), 1959: 171.

[An issue to mark the centenary of the establishment of the Singapore

Botanic Gardens.]

1960

Organograph and evolution of Dioscoreaceae, the family of the Yams.

J. Linn. Soc. Lond. Bot., 56, 1960: 319-412.

1961

Chapaters on the history of botany in India.

III. At the middle of the 19th century.

J. Bomb. Nat. Hist. Soc., 58. 1961: 678-706.

92 Gardens’ Bulletin, Singapore — X XII (1967)

1962

Chapters on the history of botany in India.

IV. The Royal Gardens at Kew begin to guide the direction of botany

in India.

J. Bomb. Nat., Hist. Soc., 59. 1962: 335-59.

Chapters on the history of botany in India.

V. The undertaking of two great enterprises, The Flora of the Higher

Plants, and the Forest Service.

J. Bomb. Nat. Hist. Soc., 59. 1962: 747-77.

1963

Chapters on the history of botany in India.

VI. The publication of Hooker’s Flora of British India, and what its

publication released.

J. Bomb. Nat. Hist. Soc., 60. 1963: 49-83.

Chapters on the history of botany in India.

VII. Epilogue.

J. Bomb. Nat. Hist. Soc., 60. 1963: 356-70.

[The seven chapters of this work are reprinted in book form by the

Botanical Survey of India, and published by the Government of India

Press. There is an introduction by Father H. Santapau, Director of the

Botanical Survey of India, and Chapter VII was rewritten by the

author for the book. Pp. XI + 245, maps 2, pl. 4; 25 April 1966.]}

I. BURKILLIAN TAXA

ACANTHACEAE

Afromendoncia floribunda Burkill, sp. nov. in Thiselton-Dyer: Fl. Trop.

Afr. 5 (1899): 6.

Brillantaisia debilis Burkill, sp. nov., l.c.: 39.

leonensis Burkill, sp. nov., Lce.: 41.

nyanzarum Burkill, sp. nov., l.c.: 39.

subulugurica Burkill, sp. nov., l.c.: 42.

Eranthemum huegelii Burkill, sp. nov., Proc. Camb. Phil. Soc. 9, TE

(1896): 96.

Hygropila acutisepala Burkill, sp. nov., in Thiselton-Dyer: Fl. Trop. Afr.

5. (1899): 34.

ciliata Burkill, sp. mov. 1¢.: 35:

gigas Burkill, sp. nov., I.c.: 36.

gracillima Burkill, sp. nov., l.c.: 36.

lindaviana Burkill, sp. nov., l.c.: 509.

linearis Burkill, sp. nov., l.c.: 35.

pilosa Burkill, sp. nov., l.c.: 35.

Thunbergia crispa Burkill, sp. nov., l.c.: 12.

hanningtonii Burkill, sp. nov., l.c.: 19.

laborans Burkill, sp.nov., l.c.: 507.

lathyroides Burkill, sp. nov., l.c.: 24.

mellinocaulis Burkill, sp. nov., l.c.: 23.

sericea Burkill, sp. nov., l.c.: 14.

stellarioides Burkill, sp. nov., l.c.: 26.

Synnema brevitubum Burkill, sp. nov., l.c.: 30.

Burkill, f. — Burkill, 1870-1965, a bibliography 93

ACTINIDIACEAE

Suarauja rufa Burkill, sp. nov. in Kew Bull. 1899: 97.

AMPELIDACEAE

Vitis simplex (Blanco) Burkill, comb. nov. in Kew Bull, 1935. 319.

(= Cissus simplex.)

ANNONACEAE

Fissistigma kingii (Boerl.) Burkill, comb. nov. in Kew Bull. 1935: 317.

( =Melodorum kingii.)

APOCYNACEAE

Alyxia kurzii Burkill, nom. nov. in Kew Bull. 1935. 317. (= Gynopogon

breviflorus Kurz.)

Ervatamia dichotoma (Roxb.) Burkill, comb. nov. in Kew Bull. 1935, 317.

(= Tabernaemontana dichotoma.)

divaricata (Linn.) Burkill, comb. nov. in Rec. Bot. Surv. India 10

(1925): 320. (= Nerium divaricatum.)

sphaerocarpa (Blume) Burkill, comb. nov. in Kew Bull. 1935, 317.

= Tabernaemontana sphaerocarpa.)

ARACEAE

Arisaema listeri Prain ex Burkill, sp. nov. in Rec. Bot. Surv. India 10,

(1925) , 397.

Aralia quinquefolia Dec’ne and Planch var. angustifolia Burkill, var.

nov. in Kew Bull., 1902: 7.

var. elegantior Burkill, var. nov., l.c.: 8.

var. major Burkill, var. nov., l.c.: 7.

var. notoginseng Burkill, var. nov., l.c.: 7.

var. pseudo-ginseng Burkill, var. nov. l.c.: 7.

var. repens Burkill, var. nov., l.c.: 6.

ASCLEPIADACEAE

Hoya subcalva Burkill, sp. nov. in Kew Bull. 1901: 141.

BEGONIACEAE

Begonia foxworthyi Burkill ex Ridley, sp. nov. in FI. Mal. Penins.

i es i) eee

haniffii Burkill, sp. nov. in J. Str. Br. Roy. As. Soc. 79 (1918): 103-4.

tampinica Burkill ex Irmsch. sp. nov. in Mitt. Inst. Bot. Hamburg 8

(1929): 130.

BETULACEAE

Alnus cremastogyne Burkill , sp. nov. in J. Linn. Soc. Lond. Bot. 26

(1899): 499. :

maritima Nutt., var. formosana Burkill, var. nov., l.c.: 500.

Betula albo-sinensis Burkill, sp. nov., l.c.: 497.

utilis D. Don, var. prattii Burkill, var. nov., l.c.: 499.

Carpinus laxiflora Blume, var. fargesii Burkill, var. nov., l.c. 501.

pubescens Burkill, sp. nov., l.c.: 502.

Corylus heterophylla Fisch., var. crista-galli Burkill, var. nov., l.c.: 504.

94 Gardens’ Bulletin, Singapore — XXII (1967)

BOMBACACEAE

Ceiba occidentalis (Sprengel) Burkill, comb. nov. in Kew Bull., 1935: 317.

(= Bombax occidentale.)

BROMELIACEAE

Karatas serra (Grisebach) Burkill, comb. nov. in Kew Bull. 1935: 317.

(= Bromelia serra.)

BURSERACEAE

Garuga pacifica Burkill, sp. nov. in J. Linn. Soc. Lond. Bot. 35 (1901):

30-31.

COMPOSITAE

Acomis lesteri Burkill, sp. nov. in Kew Bull. 1901, 140.

Chrysanthemum umbelliferarum (Boissier) Burkill, comb. nov. in Kew

Bull. 1935: 317. (= Tanacetum umbeiliferum.)

Launaea crassiniana (Boiss.) Burkill, comb. nov. in FI. Pl. Baluchistan

44; 1909. (= Zollikofera crassiniana.)

polyclada (Boiss.) Burkill, comb. nov., l.c., 44. (= Z. polyclada.)

stenocephala (Boiss.) Burkill, comb. nov., l.c., 44. (= Z. stenocephala.)

stocksiana (Boiss.) Burkill, comb. nov., l.c., 44. (= Z. stocksiana.)

Icomum lineare Burkill, sp. nov. in J. Linn. Soc. Lond. Bot. 34, (1899) 270.

salicifolium Burkill, sp. nov., l.c.: 270.

subacaule Burkill, sp. nov., l.c.: 271.

CRUCIFERAE

Anguillicarpus Burkill, gen. nov. in J. As. Soc. Bengal ns. 3, (1907) 59.

bulleri Burkill, sp. nov. l.c.: 60.

CONVOLVULACEAE

Merremia petaloidea (Choisy) Burkill, comb. nov. in Kew Bull. 1935, 318.

(= Ipomoea petaloidea.)

DIOSCOREACEAE

Sec. Campanuliflorae Burkill & Perrier, sec. nov. in Mus. Natl.

Hist. Nat. Notulae Syst. 14 (1951): 132-7.

Sec. Illigerastrum Prain & Burkill, sec. nov. in Kew Bull. 1933: 241.

Sec. Pachycapsa Burkill & Perrier, sec. nov. in Mus. Natl. Hist. Notulae

Syst. 14 (1951): 132-7.

Sec. Paramecocarpa Burkill, sec. nov. in Gard. Bull. S.S., 3 (1924): 123—

(name); Prain and Burkill in Ann. R. Bot. Gard. Calc., 14 (1936): 93-94

— (description).

Sec. Seriflorae Burkill & Perrier, sec. nov. in Mus. Natl. Hist. Nat.

Notulae Syst. 14 (1951): 132-7.

Sec. Shannicorea Prain & Burkill, sec. nov. in J. As. Soc. Bengal

n.s., 10 (1914): 10, and Ann. R. Bot. Gard. Calc., 14 (1936): 100.

Sec. Xylinocapsa Burkill & Perrier, sec. nov. in Mus. Natl. Hist. Nat.

Notulae Syst., 14 (1951): 132-7.

Burkill, f. — Burkill, 1870-1965, a bibliography 95.

Dioscorea acerifolia Diels ex Prain & Burkill, sp. nov. in J. As. Soc.

Bengal 73 (Suppl.) (1905): 7.

aculeata Linn. var. fasciculata Prain & Burkill, var. nov. in J. As. Soc.

Bengal, n.s. 10 (1914): 20.

var. spinosa Prain & Burkill, var. nov. l.c. 10: (1914) 20.

var. tillaefolia (Kunth) Prain & Burkill, comb. nov. in Elmer:

Leaflets Philipp. Bot. 5 (1913): 1594, and J. As. Soc. Bengal,

ns. 10 (1914): 20.

alata Linn. var. tarri Prain & Burkill, var. nov. in J. As. Soc. Bengal,

ns. 10 (1914): 39.

alatipes Burkill & Perrier, sp. nov. in Humbert: Fl. Madag. Fam.

44 (1950): 50— (name); Burkill & Perrier in Not. Syst. 14 (1951):

136 —— (description).

arachnida Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 16:

(1914): 21.

asclepiada Prain & Burkill, sp. nov. in Kew Bull. 1916: 190.

aspera Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns., 4 (1908):

447. -

asteriscus Burkill, sp. nov. in Bull. Jard. Bot. Brux., 15 (1939): 356-7.

bancana Prain & Burkill, sp. nov. in Kew Bull. 1925: 62.

belophylloides Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 4

(1908): 448.

benthamii Prain & Burkill, sp. nov. in l.c., 4 (1908): 448-9.

bernoulliana Prain & Burkill, sp. nov. in Kew Bull. 1916: 192.

bicolor Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 4

(1908): 449.

birmanica Prain & Burkill, sp. nov. in l.c., 73 (1904): 185.

biserialis Prain & Burkill, sp. nov. in Kew Bull. 1925: 58-59.

blumei Prain & Burkill, sp. nov., in J. As. Soc. Bengal, n.s., 10 (1914): 25.

bonatiana Prain & Burkill, sp. nov. in Kew Bull. 1925: 61.

bonii Prain & Burkill, sp. nov., l.c., 1933: 244-S.

brandisii Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 10

(1914): 27.

brevipetiolata Prain & Burkill ex Craib, sp. nov. in Kew Bull. 1912:

407 —(name); Prain & Burkill in J. As. Soc. Bengal, ns., 10

(1914): 38 — (description).

bulbifera Linn., var. birmanica Prain & Burkill, var. nov. in J. As.

Soc. Bengal, n.s., 10: (1914): 26.

var. deltoidea Prain & Burkill, var. nov. l.c.: 26.

var. elongata Prain & Burkill, var. nov. l.c.: 26

var. kacheo Prain & Burkill, var. nov. l.c.: 26.

var. sativa Prain & Burkill, var. nov. l.c.: 26.

var. simbha Prain & Burkill, var. nov. l.c.: 26.

var. suavior Prain & Burkill, var. nov. l.c.: 26.

bullata Prain & Burkill, sp. nov. in Kew Bull. 1925: 60.

calcicola Prain & Burkill, sp. nov., l.c.: 64.

cambodiana Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns., 10

(1914): 12.

carionis Prain & Burkill, sp. nov. in Kew Bull. 1916: 193-4.

chingii Prain & Burkill, sp. nov., l.c. 1931: 425.

clarkei Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 10 (1914): 16.

collinsae Prain & Burkill, sp. nov. in Kew Bull. 1927: 234.

craibiana Prain & Burkill, sp. nov. l.c., 1931: 425-6.

cumingii Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 4

(1908): 449-SO.

‘96 Gardens’ Bulletin, Singapore — X XII (1967) |

Dioscorea, cont.

daunaea Prain & Burkill, sp. nov., l.c.: 450.

depauperata Prain & Burkill, sp nov. in Kew Bull. 1933: 245-6.

dissimulans Prain & Burkill, sp. nov. l.c.: 241-2.

elegans Ridley ex Prain & Burkill, nom. nov.: l.c.: 1925: 65

(= P. papuana Ridley.)

elmeri Prain & Burkill, sp. nov. in Elmer: Leaflets Philipp. Bot. 5

(1913): 1594.

var. dubia Prain & Burkill, var. nov. l.c. 1594, J. As. Soc. Bengal,

n.s., 10: (1914): 24.

enneaneura Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 73

Suppl. (1904): 11.

esculenta (Lour.) Burkill, comb. nov. in Gard. Bull. S.S. 1 (1917) 396.

(=Oncus esculentus.)

esquirolii Prain & Burkill, sp. nov. in Kew Bull. 1931, 426.

filicaulis Prain & Burkill, sp. nov., l.c. 1932: 242-3.

filiformis Prain & Burkill, ex Koorders-Schumacher, nom. nov. in

syst. Verzeichn. d. Herbar Koorders 3 (1914): 20. (=D. myriantha

Kunth.)

flabellifolia Prain & Burkill, sp. nov. in Elmer: Leaflets Philipp. Bot.

5 (1913): 1595.

fordii Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 4

(1908): 450-1.

foxworthyi Prain & Burkill, sp. nov. in Leaflets Philipp. Bot., 5

(1913): 1591— (name); J. As. Soc. Bengal, ns., 10 (1914): 34.

garrettii Prain & Burkill, sp. nov. in Kew Bull. 1936: 493.

gedensis Prain & Burkill, sp. nov., l.c., 1925: 64.

glabra Roxb., var. grisea Prain & Burkill, var. nov. in J. As. Soe.

Bengal, n.s., 10 (1914): 37.

var. hastifolia Prain & Burkill, var. nov. l.c.: 37.

var. longifolia Prain & Burkill, var. nov. l.c.: 38.

var. salicifolia Prain & Burkill, var. nov. l.c.: 37.

var. tenuifolia Prain & Burkill, var. nov. l.c.: 38.

‘gracilipes Prain & Burkill, sp. nov. in Kew Bull. 1925: 63.

grata Prain & Burkill, sp. nov. in Leaflets Philipp. Bot., 5 (1913):

1591 —(name); J. As. Soc. Bengal, ns. 10 (1914): 34.

hainanensis Prain & Burkill, sp. nov. in Kew Bull. 1936: 494.

havilandii Prain & Burkill, sp. nov. J. As. Soc. Bengal, n.s., 10 (1914): 40.

hemicrypta Burkill, sp. nov. in J. Linn. Soc. Lond. Bot. 53 (1949): 377 —

(name); J. S. Afr. Bot. 18 (1952): 187-8 — (description).

hemsleyi Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s.4

(1908): 451-2.

hispida Dennst. var. mollissima Prain & Burkill, var. nov. l.c., 10

(1914): 25.

var. neo-scaphoides Prain & Burkill, var. nov. in Kew Bull. 1927: 237.

var. scaphoides Prain & Burkill, var. nov. l.c.: 237.

inaequifolia Elmer ex Prain & Burkill, sp. nov. in Leaflets Philipp.

Bot, 5 (1913); s1598,

inopinata Prain & Burkill, sp. nov. in Kew Bull. 1927: 245-6.

intempestiva Prain & Burkill, sp. nov. l.c. 1933: 243-4.

japonica Thunb. var. tenuiaxon Prain & Burkill, var. nov. in J. AS.

Soc. Bengal, n.s., 10 (1914): 28.

kalkapershadii Prain & Burkill, sp. nov. in J. As. Soc. Bengal,

n.s., 10 (1914): 24.

Burkill, f. — Burkill, 1870-1965, a_ bibliography oF

Dioscorea, cont.

kamoonensis Kunth, var. delavayi Prain & Burkill, var. nov. l.c: 22.

var. fargesii Prain & Burkill, var. nov., l.c.: 21.

var. henryi Prain & Burkill, var. nov., l.c.: 22.

var. straminea Prain & Burkill, var. nov. l.c.: 21.

keduensis Prain & Burkill sp. nov. in Backer: Handb. Flora van Java, 3:

(1924): 114— (name); Kew Bull. 1925: 59 — (description).

kerrii Prain & Burkill sp. nov. in Craib: Kew Bull. 1912: 407 — (name).

J. As. Soc. Bengal, n.s., 10 (1914): 20— (description).

kratica Prain & Burkill, sp. nov. in Kew Bull. 1927: 241.

lamprocaula Prain & Burkill, sp. nov., l.c. 1932: 245.

lepcharum Prain & Burkill, sp. nov., in J. As. Soc. Bengal, n.s., 10

(1914): 36-37.

var. bhamoica Prain & Burkill, var. nov. l.c. 37.

lineari-cordata Prain & Burkill, sp. nov. in Kew Bull. 1925: 61-62.

listeri Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 6 (1908): 452.

loheri Prain & Burkill sp. nov. in Elmer: Leaflets Philipp. Bot., 5

(1913): 1591—(name); J. As. Soc. Bengal, n.s., 10 (1914): 33-34 —

(description).

madiunensis Prain & Burkill, sp. nov. in Kew Bull. 1925: 63.

martini Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 10 (1914):.

18-19.

melanophyma Prain & Burkill ex Duthie sp. nov. in Strachey, Cat.

Plants Kumaon, p. 186, 1906—(name); J. As. Soc. Bengal, ns. 4

(1908): 452-3 — (description).

membranacea Pierre ex Prain & Burkill, sp. nov. in J. As. Soc. Bengal,

ns. 10 (1914): 13.

merrillii Prain & Burkill, sp. nov. Leaflets Philipp. Bot. 5 (1913): 1958.

morsei Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 4 (1908): 454.

moultonii Prain & Burkill, sp. nov. in Kew Bull. 1925: 62.

nieuwenhuisii Prain & Burkill, sp. nov. l.c.: 65.

nitens Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns., 10 (1914): 18.

nipponica Makino, var. jamesii Prain & Burkill, var. nov. l.c.: 14.

var. rosthornii Prain & Burkill, var. nov. l.c.: 14.

nummularia Lamk., var. glauca Prain & Burkill, var. nov. in Leaflets

Philipp. Bot., 5 (1913): 1599; J. As. Soc. Bengal, n.s., 10 (1914): 35.

oenea Prain & Burkill, sp. nov. in J. As. Soc. Bengal, ns. 10 (1914):

16-17.

oryzetorum Prain & Burikll, sp. nov. in Kew Bull. 1927: 242-3.

var. angustifolia Prain & Burkill, var. nov. l.c.: 243.

var. latifolia Prain & Burkill, var. nov. l.c.: 243.

var. mediifolia Prain & Burkill, var. nov. l.c.: 243.

owenli Prain & Burkill, sp. nov., l.c., 1925: 63-64.

palatanifolia Prain & Burkill, sp. nov. l.c.: 60.

palawana Prain & Burkill, sp. nov., l.c.: 59.

paleata Burkill, sp. nov., in Bull. Jard. Bot. Brux., 15 (1939): 360.

panthaica Prain & Burkill, sp. nov. in J. As. Soc. Bengal, 73 Suppl.

(1904): 6.

‘98 Gardens’ Bulletin, Singapore — XXII (1967)

Dioscorea, cont.

paradoxa Prain & Burkill, sp. nov. in Kew Bull. 1927: 246-7.

pentaphylla Linn., var. cardonii Prain & Burkill, var. nov. in J. As. Soc.

Bengal, n.s., 10 (1914): 23.

var. communis Prain & Burkill, var. nov., l.c.: 23.

var. hortorum Prain & Burkill, var. nov., l.c.: 23.

var. jacquemontii Prain & Burkill, var. nov., l.c.: 23.

var. kussok Prain & Burkill, var. nov., l.c.: 23.

var. linnaei Prain & Burkill, var. nov., l.c.: 23.

var. malaica Prain & Burkill, var. nov., l.c.: 23.

var. rheedei Prain & Burkill, var. nov., 1G.2 aa.

var. siamensis Prain & Burkill, var. nov. in Kew Bull. 1927: 236.

var. simplicifolia Prain & Burkill, var. nov. in J. As. Soc. Bengal,

ns. 10 (1914): 23.

var. suli Prain & Burkill, var. nov., l.c.: 23.

var. thwaitesii Prain & Burkill, var. nov., l.c.: 23.

peperoides Prain & Burkill, sp. nov. in Leaflets Philipp. Bot., 5 (1913):

1597.

var. angulata Prain & Burkill, var. nov. in J. As. Soc. Bengal,

ns.. 10 (1914): 28.

var. sagittifolia Prain & Burkill, var. nov. in Leaflets Philipp. Bot.,

5 (1913): 1591 — (name); J. As. Soc. Bengal, n.s., 10 (1914): 28 —

(description).

persimilis Prain & Burkill, sp. nov., J. As. Soc. Bengal, ns., 4 (1908):

454-5.

petelotii Prain & Burkill, sp. nov. in Kew Bull. 1933: 240-1.

pierrei Prain & Burkill, sp. nov. in J. As. Soc. Bengal, n.s., 10 (1914): 22.

piscatorum Prain & Burkill ex Burkill, sp. nov. in Gard. Bull. S.S. 3

(1924): 123—(mame); Kew Bull. 1925: 61.

platanifolia Prain & Burkill, sp. nov., in Kew Bull. 1925: 60.

platycarpa Prain & Burkill, sp. nov. l.c.: 65.

poilanei Prain & Burkill, sp. nov. l.c. 1933: 240.

porteri Prain & Burkill ex Knuth sp. nov. in Engl. Pflanzenr. IV, 43:

352, 1924; ex Ridley in Fl. Mal. Penins. 4, 318, 1924.

var. andersoni Prain & Burkill ex Ridley, var nov. l.c.: 318.

var. purpureo-venia Prain & Burkill ex Ridley var. nov. l.c.: 318.

potanini Prain & Burkill, sp. nov. in Kew Bull. 1933: 243.

praecox Prain & Burkill sp. nov. in J. As. Soc. Bengal, n.s., 4 (1908):

455.

prazeri Prain & Burkill, sp. nov. in J. As. Soc. Bengal, 73 Suppl.

(1904): 2; lc., ms. 10 (1914): 15.

pseudo-nitens Prain & Burkill sp. nov. in Kew Bull. 1927: 231.

pulvera Prain & Burkill sp. nov. in J. As. Soc. Bengal, n.s., 10 (1914):

31.

pyrifolia Knuth, var. diepenhorstii Prain & Burkill var. nov. in J.

As. Soc. Bengal, n.s., 10 (1914): 33.

var. ferruginea Prain & Burkill var. nov. l.c.: 33.

ridleyi Prain & Burkill sp. nov., l.c.: 12-13.

rockii Prain & Burkill sp. nov. in Kew Bull. 1927: 229-30.

rogersii Prain & Burkill sp. nov. in J. As. Soc. Bengal, ns. 10

(1914); 27.

‘scortechinii Prain & Burkill sp. nov. l.c., ns., 4 (1908): 455-6.

seemanii Prain & Burkill, nom. nov., lec.: 34 (=D. nummularia

Seemann).

seniavinii Prain & Burkill sp. nov. in Kew Bull. 1925: 59.

sexrimata Prain & Burkill, sp. nov. l.c., 1950: 259-60.

sikkimensis Prain & Burkill sp. nov. in J. As. Soc. Bengal, 73 Suppl.

(1904): 3.

Burkill, f. — Burkill, 1870-1965, a bibliography 99

Dioscorea, cont.

simulans Prain & Burkill sp. nov. in Kew Bull. 1931: 427.

sitamiana Prain & Burkill sp. nov. l.c., 1925: 64-65.

soror Prain & Burkill sp. nov. in Leaflets Philipp. Bot., 5 (1913):

1958-9.

var. glauca Prain & Burkill var. nov. l.c.: 1591—(name); J. As.

Soc. Bengal, ns.. 10 (1914): 34.

spicata Roth, var. anamallayana Prain & Burkill, var. nov. in J. As.

Soc. Bengal, n.s., 10 (1914): 29.

var. parvifolia Prain & Burkill, var. nov. l.c.: 29.

stemonoides Prain & Burkill sp. nov. in Kew Bull. 1927: 244.

stenomeriflora Prain & Burkill, sp. nov., 10 (1914): 40-41.

subcalva Prain & Burkill sp. nov. in J. As. Soc. Bengal, n.s., 10 (1914):

18.

sumatrana Prain & Burkill sp. nov. in Kew Bull., 1931: 90-91.

sylvatica Ecklon var. paniculata (Dummer) Burkill, stat. nov. in J. S. Afr.

Bot. 18 -(1952): 189.

var. brevipes (Burtt-Davy) Burkill, stat. nov. l.c.: 189.

var. rehmanni (Baker) Burkill, stat. nov. l.c.: 189.

var. multiflora (Marloth) Burkill, stat. nov. l.c.: 189.

tamarisciflora Prain & Burkill sp. nov. in J. As. Soc. Bengal, n.s., 10

(1914): 22.

tentaculigera Prain & Burkill, sp. nov. l.c.: 15.

trimenii Prain & Burkill sp. nov. l.c.: 29.

trinervia Roxb. ex Prain & Burkill, sp. nov. l.c.: 32-33.

triphylla Linn., var. daemona Prain & Burkill var. nov. l.c.: 26.

var. mollissima (Mlume) Prain & Burkill, stat. nov. in Leaflets Philipp.

var. mollissima (Blume) Prain-& Burkill, stat. nov. in Leaflets Philipp.

Bot. 5 (1913): 1596; J. As. Soc. Bengal, n.s., 10 (1914): 26.

var. reticulata Prain & Burkill, var. nov. in J. As. Soc. Bengal,

ns., 10 (1914): 26.

van vuurenii Prain & Burkill sp. nov. in Kew Bull. 1925: 63.

velutipes Prain & Burkill sp. nov. in J. As. Soc. Bengal, ns., 10

(1914): 19.

vexans Prain & Burkill sp. nov., l.c., ns., 4 (1908): 446.

ae Linn., var. coreana Prain & Burkill, var. nov. l.c., ns. 19 (1914):

Ly.

warburgiana Koorders ex Prain & Burkill sp. nov. lc. ns. 4

(1908): 456-7.

wattii Prain & Burkill sp. nov. in J. As. Soc. Bengal, l.c.: 457.

yokusai Prain & Burkill, sp. nov. l.c., 73 Suppl. (1904): 10.

yunnanensis Prain Burkill, sp. nov. l.c.: 186.

DIPTEROCARPACEAE

Isoptera seminis (de Vries) Burkill, comb. nov. in Kew Bull. 1935: 317.

(= Hopea seminis.)

EUPHORBIACEAE

Euphorbia calabarica Burkill, sp. nov. in Kew Bull. 1901: 133.

FAGACEAE

Quercus lampadaria (Gamble) Burkill, comb. nov. in Kew Bull. 1935:

318. (= Pasania lampadaria.)

maingayi (Schottky) Burkill, comb. nov., l.c.: 318. (= Pasania maingayi.)

100 Gardens’ Bulletin, Singapore — XXII (1967)

FLACOURTIACEAE

Trimeria tropica Burkill, sp. nov. in Kew Bull. 1898: 145.

GENTIANACEAE

Gentiana albicalyx Burkill, sp. nov. in J. As. Soc. Bengal, ns. 2

(1906): 314-S.

amoena C.B. Clarke, var. major Burkill, var. nov. l.c.: 312.

amplicrater Burkill, sp. nov. l.c.: 312.

arethusae Burkill, sp. nov. l.c.: 329.

atkinsonii Burkill, sp. nov. l.c.: 309-10.

bryoides Burkill, sp. nov. l.c.: 316.

crassicaulis Duthie ex Burkill, sp. nov., l.c.: 311-2.

detonsa Rottb., var. lutea Burkill, var. nov., l.c.: 319.

var. ovata-deltoidea Burkill, var. nov., l.c.: 319.

duthei Burkill, sp. nov., l.c.: 318.

intermedia Burkill, sp. nov. l.c.: 313.

lawrencei Burkill, sp. nov. in Gard. Chron. 2 (1905): 307.

lhassica Burkill, sp. nov. in J. As. Soc. Bengal, n.s. 2 (1906): 311.

listeri Burkill, sp. nov. l.c.: 314.

micantiformis Burkill, sp. nov., l.c.: 315-6.

panthaica Burkill, sp. nov., l.c.: 313-4.

pharica Burkill, sp. nov. l.c.: 310.

prainii Burkill, sp. nov., l.c.: 317.

pseudo-humilis Burkill, sp. nov., l.c.: 313.

saginoides Burkill, sp. nov., l.c.: 318.

sorocula Burkill, sp. nov., l.c.: 315.

waltonii Burkill, sp. nov., l.c.: 310-1.

yokusai Burkill, sp. nov., l.c.: 316-7

Parajaeschkea Burkill, gen. nov. in Rec. Bot. Surv. India, 4 (1911): 223.

smithii Burkill, sp. nov. l.c.: 223.

Swertia angustifolia Hamilton ex D. Don, var florida (Wall.) Burkill

stat. nov. in J. As. Soc. Bengal, ns., 2 (1906): 374. (= S. florida.)

var. hamiltoniana Burkill, nom. nov. l.c., 374. (= S. angustifolia Ham.)

var. pulchella Burkill, stat. nov. l.c., 375. (= S. pulchella Ham

et alii.) |

var. wallichiana Burkill, stat. nov. l.c., 374. (= S. angustifolia Wall.

in parte.) |

australis Burkill sp. nov. in J. As. Soc. Benegal, n.s. 7 (1911): 81-82.

bisseti Moore & Burkill, sp. nov. l.c., ns. 2 (1906) 329.

bonatiana Burkill, sp. nov., l.c. n.s. 7 (1911): 81.

carinthiaca Griseb., var. afghanica Burkill, var. nov., l.c.: ns. 2 (1906):

324.

chumbica Burkill, sp. nov. l.c.: 323.

cincta Burkill, sp. nov., lic.: 319.

deltoidea Burkill, sp. nov., l.c.: 324.

duclouxii Burkill, sp. nov., l.c.: ns. 7 (1911): 81.

exacoides Burkill, sp. nov., l.c.: ns. 2 (1906): 321-2.

gamosepala Burkill, sp. nov., l.c.: 324-5.

hickinii Burkill, sp. nov., l.c.: 320-1.

Burkill, f. — Burkill, 1870-1965, .a, bibliography

hispidicalyx Burkill, sp. nov., lc.: 321.

var. major Burkill, var. nov., l.c.: 321.

var. minima Burkill, var. nov. l.c.: 321.

101

lawii (Wright & Arnott) Burkill, comb. nov. l.c.: 379. (= Ophelia lawii.)

lloydioides Burkill, sp. nov. l.c.: ns. 2 (1906): 323.

patens Burkill, sp. nov., l.c.: ns. 7 (1911). 82.

paupera Burkill, sp. nov., l.c., ns. 2 (1906): 322.

purpurascens Wall., var. ramosa Burkill, var. nov., l.c., n.s. 3 (1907): 34.

rosea Burkill, sp. nov., l.c., ns. 7 (1911): 82.

sikkimensis Burkill, sp. nov., l.c., ns. 2 (1906): 322-3.

souliaei Burkill, sp. nov., l.c.: 325-6.

stapfii Burkill, sp. nov., l.c.: 325.

speciosa Wall., var. lacei Burkill, var. nov., l.c.: 326.

subspeciosa Burkill, sp. nov., l.c.: 326.

tongulensis Burkill, sp. nov., l.c.: 319-20.

younghusbandii Burkill sp. nov., l.c.: 325.

yunnanensis Burkill, sp. nov. l.c.: 320.

GESNERIACEAE

Boea hians Burkill,.sp. nov., in Kew Bull..1901: 142.

Paraboea tiumanica Burkill sp. nov. in Ridley: Fl. Mal.

(ig25) 2532:

Trichosporum lampongum (Miquel) Burkill, comb. nov. in

1935: 319. (= Aeschynanthus lamponga.)

nummularium Burkill & S. Moore ex S. Moore, sp. nov.

1899: 173.

GRAMINEAE

Acroceras crassiapiculatum (Merrill) Burkill, comb. nov. in

1935: 316-7. (= Panicum crasiapiculatum.)

HYPOXIDACEAE

Curculigo grandis MS. in Herb. Calc. ex Burkill sp. nov. in

Surv. India, 10 (1925): 386 — (name).

LEGUMINOSAE

Whitfordiodendron pubescens (Craib) Burkill comb. nov. in

1935: 319. (= Padbruggea pubescens.)

LORANTHACEAE

Loranthus pachypus Burkill, sp. nov. in Kew Bull. 1899: 109.

MELASTOMATACEAE

Dissochaeta pentamera Burkill, sp. nov. in Kew Bull. 1906: 5.

Penins. “2

Kew Bull.

im J. pot.

Kew Bull.

Rec. Bot.

Kew Bull.

Memecylon curtisii Burkill & Henderson, nom. nov. in Gard. Bull.

Straits Settlements 2 (1925): 377. (= M. gracilipes Ridl.)

Plethiandra sahebii Burkill sp. nov. in J. Straits Br. R. As. Soc. 77

(1917): 265-9.

Pternandra azurea (de C.) Burkill comb. nov. in Kew Bull. 1935:

319. (= Kibessia azurea.)

102 Gardens’ Bulletin, Singapore — XXII (1967)

MUSACEAE

Musa pruinosa Burkill, sp. nov. in Rec. Bot. Surv. Ind., 10 (1925): 384.

MYRTACEAE

Eugenia crosbyi Burkill, sp. nov. in J. Linn. Soc. Lond., 35 (1901): 38.

prora Burkill, sp. nov. in Kew Bull. 1906: 4.

samoensis Burkill, sp. nov. in J. Linn. Soc. London., 35 (1901: 38.

Rhodamnia uniflora (Ridley) Burkill, stat. nov. in Gard. Bull. Straits

Settlements 3 (1923): 42. (= R. trinervia var. uniflora Ridley.)

OCHNACEAE

Ouratea crocea (Griffith) Burkill, comb. nov. in Kew Bull. 1935: 218.

(= Ochna crocea.)

hookeri (Planchon) Burkill, comb. nov. l.c.: 318. (= Ochna hookeri.)

ORCHIDACEAE

Bulbophyllum oeneum Burkill ex Ridley, sp. nov. in Fl. Mal. Penins.

4 (1924): 63.

Corymborchis longiflora (Hk.f.) Burkill, comb. nov. in Kew Bull. 1935:

317. (= Corymbis longiflora.)

Dendrobium citrino-castaneum Burkill, sp. nov. in Gard. Bull. Straits

Settlements, 3 (1923): 12.

gracilipes Burkill, sp. nov. in J. Straits Br. R. As. Soc., 79 (1918): 45-46.

Gastrochilus subbilobus Burkill sp. nov. in Valeton: Bull. Jard. Bot.

Buitenzorg;, II, 27 (1918): 87; in Ridley: Fl. Mal. Penins., 4 (1924):

250.

Liparis purpureo-viridis Burkill & MHolttum, sp. nov. in Gard. Bull.

Straits Settlements 3 (1923): 77.

Oberonia fungum-olens Burkill, sp. nov. l.c., 3 (1924): 292.

Saccolabium crucicallus Burkill, sp. nov., 3 (1923): 7.

PITTOSPORACEAE

Pittosporum berberdoides Burkill, sp. nov. in Kew Bull. 1899: 96.

pullifolium Burkill, sp. nov., l.c.: 96.

spathaceum Burkill, sp. nov. in Hock. Ic. Pl. t. 2561 (1898).

RUBIACEAE

Dolicholobium acuminatum Burkill, sp. nov. in Hook. Ic. Pl. t. 2630.

(1900).

RUTACEAE

Citrus polyandra (Ridley) Burkill, comb. nov. in Gard. Bull. Straits

Settlements. 5 (1931): 219. (= Atalantia polyandra.)

swinglei Burkill ex Harms, sp. nov. in Engl. & Prantl, Nat. Pfanzenfam.

ed. 2, XIX c. 459, 1931.

Paramignya angulata (Willd.) Burkill, comb. nov. in Gard. Bull. Straits

Settlements, 5 (1931): 214-5. (= Citrus angulata.)

cuspidata (Ridley) Burkill, comb. nov., l.c.: 213. (= Atalantia cuspidata.)

lobata Burkill, nom. nov., l.c.: 214. (= Atalantia hispida Ridley.)

missionis (Oliv.) Burkill, comb. nov., l.c.: 213 (= Atalantia missionis.)

ridleyi Burkill, nom. nov., l.c.: 214. (= P. graffithii Ridley.)

trimera (Qliv.) Burkill, comb. nov., l.c.: 213. (= Atalantia trimera.}

Burkill, f. — Burkill, 1870-1965, a bibliography 103

SALICACEAE

Populus tremula Linn., var. adenopoda (Maxim) Burkill, stat. nov. in J.

Linn. Soc. Lond. Bot. 26 (1899): 537. (= P. adenopoda Maxim.)

Salix daltoniana Anderss., var. franchetiana Burkill, var. nov., l.c.: 528.

fargesii Burkill, sp. nov., l.c.: 528-9.

floccosa Burkill, sp. nov., l.c.: 529.

henryi Burkill, sp. nov., l.c.: 530.

longiflora Anderss., var. albescens Burkill, var. nov., l.c.: 530.

subpycnostachya Burkill, sp. nov., l.c.: 532.

SAPOTACEAE

Planchonella pohlmanniana (Benth. & Hook. f.) Burkill, comb. nov. in

Kew Bull. 1935: 319 (= Sideroxylon pohlmanniana.)

Sideroxylon vitiense (A. Gray) Burkill, comb. nov. in J. Linn. Soc. Lond.

Bot. 35 (1901): 44. (= Sapota vitiense.)

STERCULARIACEAE

Erythropsis colorata (Roxb.) Burkill, comb. nov. in Gard. Bull. Straits

Settlements 5 (1931): 231. (= Sterculia colorata.)

Firmiana lychnophora (Hance) Burkill, comb. nov. in Kew Bull. 1935:

317. (= Sterculia lychnophora.)

Pterospermum proteus Burkill, sp. nov., l.c. 1901: 138.

THEACEAE

Gordonia benguetica Burkill, sp. nov. in Philipp. J. Sci., 5 (1919): 478.

concentricicatrix Burkill, sp. nov. in J. Straits Br. R. As. Soc., 76

(1917): 152-3.

havillandii Burkill, sp. nov., l.c.: 156-7.

lanceifolia Burkill, sp. nov., l.c.: 150-1.

polisana Burkill, sp. nov., in Philipp. J. Sci. 15 (1919): 478.

subclavata Burkill, sp. nov., l.c.: 478.

taipingensis Burkill, sp. nov. in J. Straits Br. R. As. Soc. 76 (1917): 148.

TILIACEAE

Corchorus capsularis Linn., var. oocarpus Burkill & Finlow, var. nov.

in Proc. As. Soc. Bengal, 1910, 101; J. As. Soc. Bengal ns. 7 (1911):

465-6.

Elaeocarpus barnardii Burkill, sp. nov. in J. Straits Br. A. As. Soc.

75 (1917): 41.

tonganus Burkill, sp. nov. in J. Linn. Soc. Lond. Bot. 35 (1901) 29.

UMBELLIFERAE

Ducrosia anethifolia (de C.) Boiss., var. jamiatii Burkill var. nov. in J.

As Soc. Bengal, ns., 3 (1907): 564.

URTICACEAE

Polychroa javanica (Weddell) Burkill, comb. nov. in Kew Bull. 1935: 319.

(= Pellionia javanica.)

104 Gardens’ Bulletin, Singapore — XXII (1967)

VIOLACEAE

Rinorea hookeriana (King) Burkill comb. nov. in Kew Bull. 1935:

319. (= Alsodeia hookeriana.) .

ZINGIBERACEAE

Aframomum macrospermum (Smith) Burkill, comb. nov. in Kew Bull.

1930: 37. (= Amomum macrospermum.)

Alpinia oceanica Burkill, sp. nov. in Proc. Camb. Phil. Soc. IX, II: 93,

1896.

Amomum kepulaga Sprague & Burkill, nom. nov. in Kew Bull. 1930:

35. (= Amomum cardamomum Roxb.)

Elettaria cardamomum Maton, var. minuscula Burkill, var. nov. in Kew

Bull. 1930: 35.

Haplochorema sumatrana Burkill, sp. nov. in Gard. Bull. Straits Settle-

ments 3 (1923): 18-19.

Languas allughas (Roscoe) Burkill, comb. nov. in Kew Bull. 1935:

317. (= Alpinia allughas.)

cannifolia (Ridl.) Burkill, comb. nov., l.c.: 318. (= Alpinia cannifolia.)

conchigera (Griff.) Burkill, comb. nov., l.c.: 1930: 37. (= Alpinia

conchigera.)

elegans (K. Schumann) Burkill, comb. nov., l.c., 1935: 318. (= Alpinia

elegans.)

galanga (Swartz) Burkill, comb. nov., l.c., 1930: 36 (= Alpinia galanga.)

globosa (Horanin) Burkill, comb. nov., in Gard. Bull. Straits Settle-

ments 6 (1929): 80; Kew Bull. 1930: 36. (= Alpinia globosa.)

javanica (Blume) Burkill, comb. nov. in Kew Bull. 1935: 318. (=

Alpinia javanica.)

melanocarpa (Ridl.) Burkill, comb. nov. in Gard. Bull. Straits Settle-

ments 6 (1930): 260; Kew Bull. 1935: 318. (=Alpinia melanocarpa.)

officinarum (Hance) Burkill, comb. nov. in Gard. Bull. Straits Settle-

ments 6 (1929): 81; Kew Bull. 1931: 324. (= Alpina officinaum.)

padacanca (Val.) Burkill, comb. nov. in Kew Bull. 1935: 318. (=Alpinia

padacanca.)

raffiesiana (Wall.) Burkill comb. nov., l.c.: 318 (= Alpinia rafflesiana.)

regia (Val.) Burkill, comb. nov., l.c.: 318. (= Alpinia regia.)

uviformis (Horaninow) Burkill, comb. nov., l.c.: 318. (= Alpinia

uviformis.)

scabra (Benth.) Burkill, comb. nov. in Gard. Bull. Straits Settlements

6 (1930): 260. (=Alpinia scabra )

Phaeomeria heyneana (Valeton) Burkill, comb. nov. in Kew Bull. 1935:

318. (= Nicolaia heyneana.)

Il. TAXA NAMED AFTER I. H. BURKILL

Begonia burkillii Dunn (Begoniaceae) in Kew Bull., 1920, 110.

Begonia burkillii Irmsch. (Begoniaceae) in Mitt. Inst. Bot. Hamburg 8

116, 1929.

Bulbophyllum burkillii Gage (Orchidaceae) in J. As. Soc. Bengal, ms., 2:

343, 1906.

Burkill, f. — Burkill, 1870-1965, a bibliography 105

Burkillanthus Swingle (Rutaceae) in J. Arn. Arb. 20: 255-6, 1939.

Burkillia West & West (Coelastraceae-Chlorococcales) in Ann. R. Bot.

Gard. Calcutta 6: 228, 1907.

Burkillia Ridley (Leguminosae) in Fl. Mal. Penins. 5: 304, 1925.

Calamus burkillianthus Becc. ex Ridley (Palmae) in FI. Mal. Penins., 5:

56, 1925; Furtado in Gard. Bull. S.S. 8: 245, 1935.

Cosmarium burkillii West & West (Desmidiaceae) in Ann. R. Bot. Gard.

Calcutta, 6: 208, 1907.

Dioscorea burkillii Knuth (Dioscoreaceae) in Engl. Pflanzenr. IV, 43: 143,

1924.

Ficus burkillii Ridley ex Burkill & Holttum (Moraceae) in Gard. Bull.

3: 73, 1923; Ridley in Fl. Mal. Penins. 3: 330, 1924.

Homalonema burkilliana Ridley (Araceae) in J. Mal. Br. R. As. Soc., 1:

104-5, 1923.

Galera burkillii Massee (Agaricaceae) in Kew Bull., 1910, 2.

Lithocarpus burkillii A. Camus (Fagaceae) in Bull. Mus. Hist. Nat. Paris

a7, 5: 88, 1933.

eae burkillianus Ridley (Pandanaceae) in J. Mal. Br. R. As. Soc., 1:

06, 1923.

Piper burkillii Ridley (Piperaceae) in J. Straits Br. R. As. Soc., 82:

189, 1920.

Psaliota burkillii Massee (Agaricaceae) in Kew Bull., 1907, 123.

Rubus burkillii Rolfe (Rosaceae) in Kew Bull., 1920, 109.

Schefflera burkillii Merrill (Araliaceae) in J. Straits Br. R. As. Soc., 79:

28-29, 1918.

Sordaria burkillii Massee (Ascomycetes) in Kew Bull., 1913, 105.

Staurastrum burkillii West & West (Desmidiaceae) in Ann. Roy. Bot. Gard.,

Calcutta 6: 216, 1907.

Stemona burkillii Prain (Roxburgiaceae) in J. As. Soc. Bengal, 73: 43, 1904.

Strobilanthus burkillii Dunn (Acanthaceae) in Kew Bull., 1920, 208.

Sewertia burkilliana W. W. Smith (Gentianaceae) in J. As. Soc. Bengal,

ms. 7; 78, 1911.

hig acer burkillii Cowan (Rubiaceae) in Notes Bot. Gard. Edinb. 16:

Xanthidium burkillii West & West (Desmidiaceae) in Ann. R. Bot. Gard.,

6: 210, 1907.

Xanthophyllum burkillii Drummond & Dunn (Polygalaceae) in Kew Bull.,

1920, 245-6. fi

Rana burkillii Annandale, in Rec. Ind. Mus. Calc., 5, 1910. (a Burmese frog).

The following are named after E. M. Burkill, the wife of I. H. Burkill,

who collected fungi at Calcutta and at Singapore, and prepared coloured

drawings. The material was curated by I. H. Burkill.

Annularia burkillae Massee (Agaricaceae) in Kew Bull., 1912, 255.

Entoloma burkillae Massee (Agaricaceae) in Kew Bull., 1914, 74.

H. B. Gilliland, 1911 - 1965

An Appreciation.

Hamish Boyd Gilliland was born on 2nd October 1911 in

Southern Rhodesia, educated at George Watson’s College, Edin-

burgh, and read Botany and Zoology in the University of Edin-

burgh, for the degree of B.Sc. In due course he read for an honours

degree under Professor Sir William Wright Smith, Regius Professor

and Regius Keeper of the Royal Botanic Garden. His interests were

both taxonomic and ecological. Later he studied at the British

Museum (Nat. Hist.), where he improved his knowledge of

systematic botany and generally prepared himself for a career in

southern Africa. Persons who helped and influenced Gilliland at

this stage were John Ramsbottom, then Keeper of the Botanical

Department at the Museum, and George Taylor. For these he

always had special niches in his heart and in his mind.

In the course of field studies in Southern Rhodesia, supported

by a Beit Scholarship, Gilliland visited the University of the

Witwatersrand in 1934, when his obvious interest and ability in

taxonomy and field ecology led to his being invited to serve on

the staff of the Department of Botany, then under the administra-

tion of Professor John Phillips.

In June, 1935 Gilliland took up his appointment and in a

remarkably short time proved himself an excellent lecturer and

demonstrator, a keen guide to post-graduate students and an

indefatigable investigator. Because of the emphasis then laid upon

both taxonomy and pure and applied ecology in the Department,

he was able to make important contributions to these subjects

before the onset of war in 1940.

Shortly before the war he published an important contribution

te the ecology of Manicald, Rhodesia (Journal of South African

Botany, 1938) based upon his studies as a Beit Research Scholar.

Volunteering to serve in the South African Forces as a private,

Gilliland in a short time proved his competence as a soldier and

in due course was commissioned. He served in East and North

Africa and, during the course of the campaign, was gazetted by

the Military Administration to study various ecological and

related matters in British Somaliland. His co-worker for a time

was one of the Department’s and also Gilliland’s own earlier

students, P. E. Glover, later to become known for his ecological

researches in East Africa. )

Gilliland’s own studies in North-east Africa provided inform-

tion which he admirably worked up after the war for a thesis for

the degree of D.Sc., which was awarded by the University of the

Witwatersrand in 1947. Abstracts from this thesis were published

in the Journal of Ecology in 1952.

108 Gardens’ Bulletin, Singapore — XXII (1967)

On his return from active service Gilliland enthusiastically

supported his Professor in the educating and training of large

special classes of ex-servicemen being prepared for careers in

conservation and veld and pasture management and in other

aspects of applied ecology.

In 1948 the Head of his Department was invited to undertake

advisory work abroad and for several years Gilliland served as

Acting Head — a duty which he fulfilled with ability.

Throughout his career in South Africa Gilliland showed himself

to be a capable, keen and pleasant colleague, a friendly and able

guide to students and one who was interested in and prepared

to work for the general good of the University and of his adopted

country, South Africa. Many a student during the periods 1935-

1940 and 1945-1955 was generously and ably helped toward a

career in pure or applied botany.

Although by nature reserved, Gilliland ever showed to those

he worked with a kindliness, a warmth and a generosity of spirit.

He was always obective, no matter how great the subjectivity of

some of those with whom he inevitably had contact at various

times. Critical of all that was ethically unworthy and scientifically

Slipshod, he was the first to encourage sincere effort no matter how

faltering this might be. For these and other qualities his name and

influence will remain cherished by a large number of people in

South Africa and Rhodesia.

Gilliland came to Singapore in September, 1955 as the second

holder of the Chair of Botany in the young University of Malaya.

He had a staff of three lecturers and some temporary assistance

and very cramped quarters and inadequate equipment. The depart-

ment in that year had through its hand some 37-40 science and

medical students. The air was heady with political change, and

expansion was the order of the day. By the end of 1964, the

department had grown in stature to 6 lecturers and 220 students,

in quarters, though very greatly expanded, which still were barely

adaquate. The path was far from smooth, and within a couple of

years of his arrival Gilliland was faced with the necessity of

establishing a new and separately equiped and staffed department

at the new Kuala Lumpur Division of the University of Malaya.

He had in fact two departments of botany on his hands 240 miles

apart, one established and inadequately provided for, the other

with nothing, but the promise of money and the prospect of a

very large number of students. In 1959, when the Department of

Botany at Kuala Lumpur opend its doors he had to operate both

departments with staff commuting to and fro between Singapore

and Kuala Lumpur. This difficult situation was resolved in part

by three of his lecturers and certain subordinate staff from

Singapore opting to transfer permanently to Kuala Lumpur. In

1960 the department in Kuala Lumpur became separate under its

own professor, and in the 1961/62 session the two divisions of

the University of Malaya split, that in Kuala Lumpur retaining

the University name, while that in Singapore, where Gilliland

remained, became the University of Singapore.

J.P. & H.M.B.— Gilliland, 1911-1965 109

These staffing problems were a challenge to Gilliland, and even

though after 1960 he had responsibilities only towards the Singa-

pore department, it was a no easy matter to engage adequate

competent staff. While promising graduates of the university

were the obvious source for future staffing, there was still the

hiatus to be bridged between their going overseas for higher

degrees and their return. Amongst the families of British Service

personnel stationed in Singapore, there has always been an

immensely varied talent, and Gilliland made providential tem-

porary use of dependents with university training in botany to

help out. Visiting research workers and Ph.D. students from

overseas were also called upon till adequate teaching staff could

be recruited from local and expatriate sources. It was not in fact

till 1962 that the Department was functioning with a full comple-

ment of staff.

Singapore has an unusually good health record amongst tro-

pical cities, but the climate can be trying, and after some five

years in Singapore Gilliland began to suffer from an asthmatic

condition. This placed great physical strain on him which made

field work especially arduous. Nevertheless he accepted this as

no excuse for lightening his load of duties, neither towards his

students and teaching, nor towards his own botanical interests.

From May, 1960 till mid 1964 he was Dean of Science, and for

two occasions in the latter half of 1964, he was Acting Vice-

Chancellor during a particularly difficult period for the University

administration which he handled with shrewd competence, charm

and tact.

Though heavily preoccupied with administrative duties, his

botanical interest in ecology and secondary vegetation was

diligently followed. The observations of earlier workers on

Singapore’s secondary vegetation were used as a basis for suc-

cession studies, which, if these areas survive untouched, will

make an interesting starting point for future ecologists to record

the final stages of transition to climax forest. His special botanical

interest was in grasses, an interest which he brought from his

earlier days in South Africa. Shortly prior to his departure from

Singapore he completed the text of ‘Grasses of Malaya’ which has

been accepted for publication as Volume 3 of the revised Flora of

Malaya, He was also keenly interested in conservation and served

as a trustee on the Singapore Nature Reserves Board for 53 years

between 1955 and 1961. He sought to bring out the educational

aspects of conservation so that the common citizen with no special

training in biology, as well as biology students, might learn to

understand and appreciate natural history.

In 1964 he was taken seriously ill with a pneumo-thorax and

had to spend some time in hospital. For many months before

this he had accepted the fact that to regain full health he would

have to move to a drier climate, and it was mixed feelings that he

left Singapore on 3rd February, 1965 for South Africa where he

OO —————

110 Gardens’ Bulletin, Singapore — XXII (1967)

took up an appointment at the Botany Department of the Univer-

sity of Natal at Pietermaritzburg. First indications were hopeful:

his health appeared to improve; he liked the department: he liked

his colleagues, and he enjoyed the stimulating change of students.

Soon after his arrival in Pietermaritzburg, he made an 800 mile

car journey to Johannesburg and back to attend his son’s gra-

duation at Witwatersrand University. The effort of this, the sudden

change to a high elevation climate, and the lack of his physical

reserves of strength brought on a relapse of pneumo-thorax and

shortly after he had an attack of pneumonia which brought his

life to an end on 23rd June, 1965 at the age of 54 years.

Known as Gilly to a wide circle of friends, Hamish Boyd

Gilliland and his wife Rita, will long be remembered in Singapore.

Their university flat was an open house for students, and often

there would be a group come in for a social gathering to meet

some eminent biologist travelling through Singapore, or simply to

have a get-together. Gilliland was an active Rotarian and was a

founder member of the Rotary Club of Singapore-West. He was

dedicated to the cause of international co-operation, goodwill and

service.

A Bibligraphy

1931. Anomalous stem structure in Ruscus aculeatus, Linn. Trans. &

Proc. Bot. Soc. Edin., 30, 284-285: 1931.

1935. A new giant Lobelia from Rhodesian Manicaland. J. Bot. Lond.,

23, 247-8: 1935.

1938. A proposed delimitation of botanical countires for Southern

Rhodesia, J. S. Afr. Bot., 4, 65-71: 1938.

The vegetation of Rhodesian Manicaland. J. S. Afr. Bot., 4,

73-99 (35 P1.): 1938.

Notes on the flora of Rhodesian Manicaland.

I. Bryophyta (with W. R. Sherrin)

IJ. Pteridophyta (with A. H. G. Alston)

III. Gymnosperme.

J. S. Afr. Bot., 4, 143-56 (5 Pl.): 1938.

‘Flora of Tropical Africa, ed. Sir. A. W. Hill, 10 (1).

Arundinella to Agrostis, by E. C. Hubbard. A review. J. S.

Afr. Bot., ©. 312-1938.

‘Conspectus Flore Angolensis, ed. Dr. L. Wiltnich Carisso, I,

Ranunculacee to Malvacee, by A. W. Exell and F. A. Mun-

donca.’ A review. l.c. 4, 31-2: 1938.

‘The Geology and Archeology of the Vaal River Basin, by P. G.

Sohnge, D. J. L. Visser and C. van Riet Lowe.’ A review. lI.c.,

4, 32, 1938.

1939. A note on the Rhodesian Cycad. Trans. Rhod. Sci. Ass., 37,

133: 1939.

1939, ‘The classification of Tropical Woody Vegetation types, by Dr.

J. Burtt Davy. Imp. For. Inst. Pap. 13.’ A review. J. S. Afr.

Bot. 5, 39%« 1939.

1940. The propagation of lawn grass, Cynodon dactylon.

I. S. Afr. Hort. Jour. 2.(2), 7: 1940.

II. l.c. 2 (3), 18-19: 1940.

1947. An approach to the problem of the government of nomadic

peoples; with special reference to Eastern British Somaliland.

S. Afr. Geogr. J., April 1947.

[Thesis for D.Sc. Witwatersrand University.]

1949. Some current problems concerning the understanding of African

vegetation. Geogr. Rev., XL (3), 466-468: 1949.

1950/52. Synopsis of the flora of the Witwatersrand. Witwatersrand Uni-

versity. [Prepared and commenced to issue in multi-copy form

for the students — the first seven parts.]

J.P. & H.M.B.— Gilliland, 1911-1965 111

1951.

152.

1953,

1954.

1955.

1956.

1958.

1959,

1960.

1961.

1962.

1963.

1964.

1965.

Three new Acacias from the Ogaden, Kew Bull. 1951. 139-40.

Notes on Witwatersrand Acacias, vol. 3, no. 3 of Trees in

south Africa, 1951, Tree Society of South Africa, Johannes-

urg.

Africa south of the Sahara, ed. Anne Walsh, South African

Institute for International Affairs: Chapter 4: The vegetation

of Africa, O.U.P.

ioe? eames of eastern British Somaliland. J. Ecol., 40, 91-124:

1952

A student's key to the Monocotyledons of the Witwatersrand.

Witwatersrand Univ. Press.

Experiments on the vegetative propagation of Cynodon species.

Chapter 9 ex Better turf through research: Johannesburg.

The birth of Joubert Park. J. Tvl. Hort. Soc., 2 (6), 154: 1953.

On the history of the plant study upon the Witwatersrand. J. S.

Afr. Bot., 19, 93-104: 1953.

The experimental investigation of grass communties on the Wit-

watersrand University’s Field Research Station, Frankenwald

- at Johannesburg, S. Africa. Proc. 7 Int. Bot. Congr., Stockholm,

1950, p. 241.

rae ae plant records. Afr. Wild Life Mag. 8, 58-60:

1954.

The International Code of Nomenclature for Cultivated Plants

do 2 viort.: Sac. 2G), Zi: 2 0), 255° 21), 277: 1954.

On the phenology of the veld around Johannesburg. I. Mono-

cotyledons, J. S. Afr. Bot., 21, 77-82: 1955.

On classifying vegetation. Inaugural address delivered on

16/11/56. Univ, Mal. Press, Singapore.

Plant communities on Singapore Island. Gard. Bull. Sing. 17 (A),

82-90: 1958.

Common Malayan plants. Univ. Mal. Press, Singapore. [Edited

extracts from Watanabe: Illustrations of useful plants of the

Southern Regions, Botanic Gardens, Shonan: Showa 20

( = Singapore: 1942).]

An Evocative Habitat. Trans. Bot. Soc. Edin. 38, 56-63: 1959.

Regenerating high forest on Singapore Island. Gard. Bull. Sing.,

17 (2), 228-43: 1959. (With M. J. Wantman.)

Progress with vegetation studies in Brunei, North Borneo,

Sarawak and Singapore. Proc. Symp. Humid Trop, Veget.,

Tjiawi, Indonesia, Dec. 1958, 12-14, UNESCO Sci. Coop.

Office S.E. Asia, 1960.

Notes on the classification of vegetation, Joc. cit. 60-68. (With

Muhammed bin Habil.)

A coniferous forest from Borneo. Mal. For. 1960.

On the symmetry of the orchid embryo, Proc. Cent. Bicent.

Congr. Biol,, Singapore, Dec. 2-9, 1958. 277-9, Univ. Mal.

Press. Singapore, 1960.

Conservation and education. Mal. Nat. J. Conservation Issue,

26-28: 1961.

‘The grasses of Burma, Ceylon, India and Pakistan (excluding

Bambusez)’ by N. L. Bor — a review, Gard. Bull. Sing., 18 (3),

330-2: 1961.

A checklist of Malayan grasses. Gard. Bull. Sing. 19 (1), 147-80:

1962.

Geographical distribution of Malayan grasses. J, Trop. Geog.

17, 20: 1963.

Further notes on the grasses of the Malay Peninsula, I. Gard.

Bull. Sing. 20 (4), 313-14: 1964.

Further notes on the grasses of the Malay Peninsula, II. Gard,

Bull. Sing. 21 (2), 209-211: 1965S.

In Press. Flora of Malaya, III. Grasses. Government Printing Office,

Singapore.

J. P. and H.M.B.

uy Gardens’ Bulletin, Singapore — XXII (1967)

H. B. GILLILAND

2nd October 1911 — 23rd June 1965.

Bow

BS KOS SARS OR ee ee tas nd

pee sa De

[Gp DE, SOO nnn ate

Yes ee

WER

WAAAY

TRL

Z Z

Y CLA: V4

Y Yj yh SLR OOP.

VU WU YI

tp Z

Y yy Z

Yi y hg’ ME y Aig

WUE a ee ab is apis PP

Taken on 2nd February 1965,

the day before he left Singapore.

With acknowledgement to the Botany Department, University of Singapore.

Observations on Ancistrocladus tectorius

HSUAN KENG

Department of Botany, University of Singapore

The conserved generic name Ancistrocladus (ancistrus —a

small hook, cladus — a branch; referring to the sympodially hooked

young shoot) was first proposed by N. Wallich in 1832. A full

description was provided by G. A. W. Arnott four years later. It

is a rather small genus comprising about 12 species, disjunctively

distributed in tropical western Africa (with about 3 species) and in

southeastern Asia (with about 9 species). The only Malesian species,

Ancistrocladus tectorius (Lour.) Merr., also known as A.

pinangianus Planch. or A. extensus Planch., occurs in the Malay

Peninsula, S. Sumatra, W. Borneo and their adjacent small islands.

Outside of the Malesian region, this species is found in Burma,

Thailand, The Andamans, Indo-China, S. China and Hainan Island.

For a citation of literature and synonyms of the genus and

of this species, see Steenis, 1948. The same author pointed out that

owing to the deficiency of properly preserved flowering material

which after drying always shinks to brittle remnants) and fruiting

material in herbarium specimens, certain androecial, gynoecial, and

seed characters are not fully understood, and the descriptions given

by various authors are sometimes controversial. The seedling

character is completely unknown.

The following observations are based on the preserved flower

buds collected by Mr. Engkik Soepadmo at the Bogor Botanic

Garden, and the flowers, fruit and seedlings collected by myself

from Kuala Sidili, Johore, Malaysia in October, 1965 and in

April, 1966. Microtome sections were made at 8-10 pu and stained

with a safranin-fast green combination.

Observations

Ancistrocladus tectorius (Lour.) Merr. is a woody simber The

young plants if growing solitarily in open places are, however, more

or less erect as a result of the intertwining of the vigorous young

shoots. The modified young shoots are tendril-like (Plate 1, a). They

consist of 3 or more sympodially arranged segments, each normally

bearing a curved hook at the tip. From the extra-axillary bud of

the basal segment, or sometimes from 2 or 3 buds of the basal

segments of these tendril-like shoots, the dwarf foliage branches

arise. Inflorescences are borne sub-terminally on the dwarf foliage

branches, which in turn, are on the old stems scrambling on or

above the ground. They are flat-topped, composed of profusely

dichotomously branched cymes (Plate 1, b). Flowers are borne on

the cymules. Penduncles are subtended by minute, ovate bracts.

Observations on the major flower parts (perianth, androecium,

gynoecium), flower anatomy, fruit, seed and seedling follow.

114 Gardens’ Bulletin, Singapore — X XII (1967)

Perianth

Calyx and corolla are clearly differentiated (Figs. 1 & 2).

The calyx is 5-lobed; the tube or hypanthium is cup-shaped.

The aestivation of the calyx-lobes in all the literature is described as

imbricate. From a study of the serial sections of young flower buds,

it reveals that they are quincuncially arranged at the base (Fig. 2,

b) with two slightly larger exterior lobes covering the other three,

of which two are completely interior, and the fifth one is inter-

mediate.

Two or three conspicuous glands are found on the waist of

the external surface of some of the calyx-lobes. Usually two

(very rarely three or one) of these cup-shaped glands are pre-

sent on the two exterior calyx-lobes, one is found on the inter-

mediate ones. So far, none has been observed on either of the two

interior calyx-lobes. They are apothecium- or perithecium-like in

the sectional views with a large number of Poon hypha-like

glandular hairs compactly arranged.

The corolla consists of 5 free petals which are papery in texture,

and strongly contorted (Figs. 1 & 2). Moderately thick-walled

secretory cells are distributed in the parenchyma tissues of sepais

and petals, especially abundant on the hypodermal layers of the

abaxial surfaces.

Figure 1. Floral diagram of Ancistrocladus tectorius Merr. (based on the

transverse sections of flower buds).

Hsuan Keng —On Ancistrocladus tectorius 115

tj—~

recur. vein.

BE ae

Figure 2. Flower bud of Ancistrocladus tectorius Merr. in_ sections.

The position of the serial transverse sections aw e are indicated on

the longitudinal section.

Androecium

The number of stamens in a flower of the genus varies from 10

to 5. Based on this character alone, the species of the genus are

arranged in 2 groups (Gilg 1895): only the Ceylonese species

(A. hamatus Gilg) belongs to the 5-stamen group, whereas the

rest, including the Malesian species belong to the 10-stamen group.

From a study of the flower sections of A. tectorius, two types

of androecium are observed: (1) the anthers of ten stamens are

almost equal in size, but the filaments differ in length, with the

antisepalous ones slightly longer than the antipetalous ones, (2)

116 Gardens’ Bulletin, Singapore — XXII (1967)

the ten stamens differ in the size of their anthers and in the length

of their filaments, with those of the antisepalous ones both larger

and longer than those of the antipetalous ones.

The anthers are 2-lobed and 4-loculate, intro-latrorse (Plate 2, i).

Owing to the disintegration of the tissue between adjacent locules of

the anther—lobes, they merge and thus the mature anthers appear

to be 2-loculate (Plate 2, j). The longitudinal lines of dehiscence

are lateral. A short projection (‘awn’) is found on the upper end

of each anther-lobe.

The filaments are comparatively short. They are swollen at the

base and almost joined together into a ring (Fig. 2, c, Plate 2, m),

which is in turn, adnate to the basal portion of the corolla.

A transverse section of a filament is orbicular or quadrangular in

outline. The epidermis is cutinized and the ground tissue is com-

posed of 2-5 layers of parenchyma cells. A single vascular bundle,

often associated with a few thick-walled secretory cells is observed.

The epidermis of the anther consists of a layer of small, elongate

cells with prominent nuclei. The endothecium in a mature anther

is composed of a layer of very large cells with strips of secondary

thickening, underneath which, one or sometimes two layers of

parietal cells line the pollen sac cavity.

Gynoecium

The three styles are articulated at the medial portions (cf. Plate 2,

g, h & k). The upper halves are totally free, erect or recurved,

with the rounded or truncate tips probably serving as stigmas. The

lower halves are partly. connivent and partly joined form a

cone-shaped structure. Only this cone-shaped structure is persistent

and becomes very prominent in fruit, while the upper halves are

deciduous.

The ovary wall is completely fused with the calyx-tube or

hypanthium, therefore the position of the ovary ‘s typically inferior

(Fig. 2; Plate 2, g, h & 1). The ovary is one-loculate, one-ovulate,

with the ovule basal-laterally attached (Plate 2, 1). All the sections

of the flower-buds studied fail to reveal the detailed structure of

the ovules themselves. It suggests that the differentiation of the

tissues in the ovule comes likely at a very late stage of development.

Vascular anatomy of flower

The elongate pedicel contains a cylindric vascular region. In the

receptacle at the level of sepal attachment, one single trace (which

later divides into 3, 5 or 7) diverges from the vascular cylinder and

enters each sepal (Fig. 2a, t,). Above this level, the bundles which

form the vascular cylinder separate and ramify. At first, 5 traces

between the 5 main traces, and another 3 traces inside the main

traces become clear. Each of the 5 traces (those between the main

traces) (t, in Fig. 2b & Plate 2, n) enters into one antisepalous

stamen; and all the 3 traces (those inside the main traces) (t,; in Fig.

2b & Plate 2, n) enter into the gynoecium as dorsal traces of the

carpels and end at the tips of the styles. From the main traces (t; in

Hsuan Keng— On Ancistrocladus tectorius 117

Fig. 2a & Plate 2, n) 5 bundles further diverge; each of them divides

tangentially into 2 traces, the outer one (which further divides into

3, 5 or 7 branches) and the inner one enter a petal and a anti-

petalous stamen respectively. The main traces themselves, however,

travel in a downward direction (forming the so-called “‘recurrent”’

veins) (t, in Fig. 2a & Plate 2, n; also in Fig. 2 & Plate 2, 1) at

low level and then bend abruptly upward and meet at the base

of the ovary-locule where the placenta is located. These traces

further appear on the top of the ovary-locule and finally meet

the dorsal traces at the tip of the styles.

Fruit

The fruit is an indehiscent nut, enveloped by the corky hypan-

thium, and crowned by the shining accrescent calx-lobes (Fig. 3 &

Plate 1, c). The five quincuncially arranged calyx-lobes are un-

equally developed: the two interior ones are usually much smaller

and form the short wings; the two exterior and the fifth sepal

are larger and form the long wings. In the fruiting stage their

aestivation nevertheless, is almost sub-valvate.

The cone-shaped basal portions of the styles are completely fused

and enlarged, constituting the persistent spherical or columnar

structure. The pericarp is corky, adnate to, and inseparable from

the hypanthium tissue.

Seed

The seed is pyramidal with a thin membranceous seedcoat. The

endosperm is ruminate, cerebral-like, starchy and horny (Fig. 4 &

Plate 1, d). This ruminate endosperm was erroneously identified as

the “‘remarkably folded”’ cotyledons (Hutchinson 1959, p. 286).

The embryo is fairly small, erect, medially but generally slightly

obliquely disposed and embedded in endosperm. The cotyledons are

somewhat divergent.

Seedling

The primary root is slender, tapering, with a large number of

short lateral rootlets (Fig. 5), black in colour. The hypocotyl is 3-5

cm. long, suberect, tetragonous and thick. The cotyledons emerge

from the pericarp almost simultaneously with the well-organized

young shoot (Plate 1, e & f). They are narrowly lanceolate (2-3

mm. by 0.5 mm.), sessile and prolonged decurrent at the base,

wither and cease to function at a very early stage and when they are

replaced by the foliage leaves. The transitional leaves (Fig. 5, b)

usually two, are subulate, (3-4 mm. by 0.6—0.8 mm.) sessile and

decurrent. }

The foliage leaves on young seedlings are simple, entire, exsti-

pulate, alternate and spirally arranged (Fig. 5). The lamina of each

is elliptic or lanceolate (1.5-3.5 cm. by 0.6-1.2 cm.), glabrous,

deep green above, with a distinct midrib; the lateral nerves are

inconspicuous. Their petioles are very short (2-3 mm. long) and

strongly decurrent, usually with a large purplish crateriform gland

on the exterior surface near the base.

118 Gardens’ Bulletin, Singapore — XXII (1967)

Figures 3-5. Ancistrocladus tectorius Merr.

3. Winged fruit, in two different views.

4. Sectional view of the seed (semi-diagramatic).

5. A seedling. Sa, a cotyledon; Sb, two transitional leaves; Sc,

a juvenile foliage leaf.

Hsuan Keng -—On Ancistrocladus tectorius 119

3 ep : eS 2 ee

Plate 1. Ancistrocladus tectorius Merr.

(a) Terminal branch bearing tendril-like young shoots; (b) Fruiting

inflorescence; (c) Two winged fruits; (d) Seeds (external and

sectional views); (e) Seedlings in various stages of development;

(f) Detail of e. A young seedling showing the radicle, hypocotyl

and cotyledons (with white background) just emerged from the

pericarp.

120

Gardens’ Bulletin, Singapore — X XII (1967)

Medlidiaddddiiiis

Plate 2. Ancistrocladus tectorius Mert.

(g) and (h) Longitudinal sections of flower buds (cf. fig. 2): (i) and

(j) young and mature anthers in transverse sections; (k) Detail of

h, showing style structures; (/) Enlargement of portion of a

longitudinal section of the flower bud, showing the basi-laterally

attached ovule and the reccurent veins (cf. fig. 2); (m) Transverse

section of a flower bud showing the filaments almost joined

together into a ring (slightly above the level showing in fig. 2, c);

(n) Transverse section of flower bud through the receptacular

region showing the vasculature (slightly below the level showing

in fig. 2, b).

Hsuan Keng — On Ancistrocladus tectorius pal

Phylogenetic consideration — A review

The genus of Ancistrocladus in modern classification systems

is generally considered to represent a monogeneric family, the

Ancistrocladaceae. The phylogenetic position of this family, how-

ever, is not well established. There are very few, if any, angiosper-

ous families like the Ancistrocladaceae which it has been suggested

at one time or another is related to such vastly distinct families as:

Annonaceae, Combretaceae, Dioncophyllaceae, Dipterocarpaceae,

Linaceae, Malpighiaceae, Myristicaceae, Pittosporaceae, Symplo-

caceae, Violaceae (Gilg 1995, Melchior 1964, Steenis 1948).

Current opinions still revolve around the placement of the

Ancistrocladaceae in either the Guttiferales (or Parietales in part,

including Dipterocarpaceae, Dioncophyllaceae and others) or the

Geraniales (including Linaceae and others). Hutchinson (1959)

shares with Planchon’s original view that this family and the

Dipterocarpaceae are closely related and both are classified under

his order Ochnales. At first, Erdtman (1952) stated that pollen

morphological evidence did not conflict with this family being

referred to, or placed near to such families as Cactaceae,

Dipterocarpaceae and Theaceae. He later (1958, cited by Melchior

1964) pointed out the similarities between pollen grains of this

family and the Dinocophyllaceae, a small monogeneric African

family which is grouped under the Guttiferae by Melchior (1964).

Steenis (1948) listed a number of important characters which dis-

tinguish this family from the Dipterocarpaceae; he appears to be

in favour of Miquel’s and Hallier f.’s suggestion that this family

is probably allied to the Linaceae-Hugoniaceae.

No positive conclusion can be drawn without thorough com-

parative morphological study of the above-mentioned taxa. The

present writer, nevertheless, feels that the resemblance of the per-

sistent and accrescent calyx-lobes in Ancistrocladaceae and

Dipterocarpaceae is rather superficial. Similar types of fruit can be

found in other unrelated families (Ridley 1930, pp. 103-112). The

seed and seedling characters apparently indicate no affinities existing

between these two families.

Acknowledgement

The writer wishes to express his thanks Dr. J. M. Machin for

reading the manuscript of this paper, to Mr. E. Soepadmo, for

supplying the preserved flower buds used in the study, and to Mr.

D. Teow for taking the photograph reproduced in this paper.

Literature cited.

ERDTMAN, G. (1952)—Pollen morphology and _ plant taxonomy:

Angiosperms. Stockholm: Almgvist and Wilsell.

GiLG, E. (1895)—Ancistrocladaceae, in A. Engler and E. Prantl, Die

natiirlichen Pflanzen familien III, VI, 6: 274-276.

HUTCHINSON, J. (1959)—The families of flowering plants. vol. 1:

Dicotyledons. 2d ed. Oxford: Clarendon Press.

MELcuHIorR, H. (1964)—A. Engler’s Syllabus der Pflanzenfamilien, ed. 12,

II. Angiospermen. Berlin: Gebriider Bornt.

RIDLEY, H. N. (1930)—The dispersal of plants throughout the world.

Kent: L. Reeve & Co.

STEENIS, C. G. Me + re se leap so lal in Flora Malesiana,

ser -

New records of plant diseases in Sarawak

for the year 1965

G. J. TURNER

Department of Agriculture, Sarawak

Plant diseases recorded for the first time in Sarawak, are given

below. The causal organisms are arranged alphabetically under

their individual hosts. The frequency of occurrence is given,

together with the Commonwealth Mycological Institute Herbarium

serial number, where identification has been performed by the

Institute.

Acacia auriculaeformis A. Cunn. ex Benth. (Black wattle)

Sooty mould Meliola species Locally 115463

common

Damping-off Pythium species 1 record —

Allium ascalonicum L. (Shallot)

Leaf blight Phyllostictina species Common 115456

Amaranthus gangeticus L. (Bayam)

Leaf spot Cercospora species Common 115453

Leaf spot Phyllosticta atriplicis 1 record 115451

Desm.

Ananas comosus Merr. (Pineapple)

Thread blight Marasmiellus scandens 1 record —

(Mass.) Denis & Reid

Horse hair blight Marasmius equicrinus 1 record —

Miill.

Annona muricata L. (Soursop)

Leaf blight Pestalotiopsis glandicola 1 record 114589

(Cast.) Steyaert :

Antigonon leptopus Hook. & Arn. (Coral creeper)

Leaf rot Corticium solani (Prill. Occasional —

& Delacr.) Bourd. &

Galz.

Apium graveolens L. (Celery)

Leaf spot Cercospora apii Fresen. Common 1145884

Leaf smut Entyloma helosciadii 1 record 1145885

Magn.

Argyreia speciosa Sweet (Elephant vine)

Leaf blight Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz.

Artocarpus elastica Reinw. (Terap)

Seedling blight Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz.

Trunk rot Ganoderma lucidum 1 record —

(Leyss. ex Fr.) Karst.

124 Gardens’ Bulletin, Singapore — X XII (1967)

Artocarpus integra Merr. (Jack fruit)

Horse hair blight Marasmius equicrinus

Mill.

Averrhoa carambola L. (Carambola)

Glomerella cingulata

(Stonem.) Spauld. &

Schrenk.

Leaf spot

Baccaurea motleyana Muell. Arg. (Rambai)

Sooty mould

Bambusa species (Bamboo)

Caldariomyces species

Marasmius equicrinus

Mill.

Sooty mould

Horse hair blight

Bidens pilosa L.

Leaf spot Cercospora bidentis

Tharp.

Rust Uromyces bidenticola

Arth.

Bignonia magnifica Bull.

Leaf blight Corticium solani (Prill.

& Delacr.) Bourd. &

Galz.

Camellia sinensis (L.) Kuntze (Tea)

Red root disease Ganoderma pseudofer-

reum (Wakef.) van

Over. & Steinm.

Canavalia ensiformis L. (Jack bean)

Leaf rot Corticium solani (Prill.

& Delacr.) Bourd. &

Galz. f

Capsicum annuum L. Chilli)

On branches Septobasidium species

Carica papaya L. (Papaya)

Leaf spot Cladosporium species

Leaf spot Curvularia intermedia

Boedijn

Leaf spot Mycosphaerella caricae

Syd.

Leaf spot Nigrospora sphaerica

(Sacc.) Mason

Celosia cristata L. (Cockscomb)

Leaf speckle

Leaf spot Physalospora species

Chrysanthemum species

Bud blight

Cladosporium species

Corynespora cassiicola

(Berk. & Curt.) Wei.

Asterina aporosae Hansf.

Cercospora celosiae Syd.

1 record

Occasional

1 record

Locally

common

Locally

Common

1 record

1 vecord

record

1 record

—

1 record

115452a

115459

115466

1154495

115449a

115450e

115450c

115450,

115450d

115473a@

115473b

1145906

114590a

Turner — New Sarawak ‘Plant Diseases, -1965 125

‘Cinnamomum zeylanicum Breyn (Cinnamon)

Leat blight ~ Corticium solani (Prill. 1 record —

{ & Delacr.) Bourd. &

Galz.

| Horse hair blight Marasmius equicrinus 1 record —

Miill.

‘Citrus grandis (L.) Osb. (Pomelo)

Branch die back Botryodiplodia 1 record —

theobromae_ Pat. '

‘Citrus nobilis Lour. (Mandarin)

Thread blight Marasmiellus scandens 1 record

(Mass.) Denis & Reid

‘Clerodendron thomsonae Balf. F.

Flower blight - Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz.

‘Cocos nucifera L. (Coconut)

Leat spot Helminthosporium incur- 1 record 115471

vatum Bern.

‘Coffea arabica L. (Arabian coffee)

Thread blight Marasmiellus scandens 1 record —

(Mass.) Denis & Reid

‘Congea species

| Flower and leaf blight Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz.

‘Crossandra undulifolia Salisb.

White root disease Fomes_ lignosus 1 record --

(Klotzsch) Bres.

Desmodium uncinatum DC.

Leaf rot Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz.

Dioscorea alata L. (Yam)

Leaf spot Pestalotiopsis species 1 record 115470

Dioscorea bulbifera L. (Potato yam)

Leaf blight Corticium solani (Prill. 1 record —

& Delacr.) Bourd. &

Galz. ,

Elaeis guineensis Jacq. (Oil palm)

On dying leaves Botryodiplodia Occasional —

theobromae Pat.

Eleusine indica Gaertn.

Flower mould Cochliobolus nodulosus 1 record 114582a

Luttrell

Stem blight Colletotrichum gramini- 1 record 1145825

cola (Ces.) Wilson

sense

126 Gardens’ Bulletin, Singapore — XXII (1967)

Eugenia malaccensis L. (Malay apple)

On leaves Aschersonia species

Sooty mould Phaeosaccardinula

javanica (Zimm.)

Yamamoto

Eugenia species (Jambu)

Leaf spot Cercospora species

Thread blight Marasmiellus scandens

(Mass.)Denis & Reid

Garcinia mangostana L. (Mangosteen)

Horse hair blight Marasmius equicrinus

Miill.

Hevea brasiliensis Muell. Arg. (Rubber)

Leaf spot andrim Ascochyta heveae Petch

blight.

Imperata cylindrica Beauv. (Lalang)

On leaves Pirostoma_ species

Ipomoea batatas Lam. (Sweet potato)

Leaf spot Cercospora timorensis

Cooke

Ipomoea carnea Jacq.

Sooty mould Meliola malacotricha

Speg.

Ischaemum rugosum Salisb.

False smut Cerebella andropogonis

Ces.

Ixora species

Leaf blight Corticium solani (Prill. &

Delacr.) Bourd. &

Galz.

Jasminum sambac Ait (Jasmine)

Leaf blight Corticium solani (Prill.

& Delacr.) Bourd. &

Galz.

Thread blight Marasmiellus scandens

(Mass.) Denis & Reid

Mangifera indica L. (Mango)

On branches Septobasidium species

Mangifera odorata Griff. (Kwini)

Thread blight Marasmiellus scandens

(Mass.) Denis & Reid

Mucuna benettii F. Muell. (New Guinea creeper)

Leaf blight Corticium solani

(Prill. & Delacr.)

Bourd. & Galz.

Muntingia calabura L. (Japanese cherry)

Thread blight Marasmiellus scandens

(Mass.) Deis & Reid

Denis

—

record

—

record

—

record

1 record

Occasional

1 record

Occasional

1 record

1 secord

1 zecord

114591

115474-

115457

111054

115454.

114585.

111052:

. » i

Turner — New Sarawak Plant Diseases, 1965

Musa textilis Née (Manila hemp)

Stem rot Marasmius stenophyllus

Mont.

Myristica fragrans L. (Nutmeg)

Pink disease Corticium salmonicolor

Berk. & Br.

Nephelium lappaceum L. (Rambutan)

Seedling blight Corticium salmonicolor

Berk. & Br.

Marasmius equicrinus

Mill.

Oidium nephelii Hadi.

Horse hair blight

Powdery mildew

Pachyrrhizus erosus Urban (Yam bean)

Leaf rot Choanephora

cucurbitarum (Berk.

& Rav.) Thaxt.

Leaf spot Myrothecium roridum

Tode ex Fr.

Paspalum scrobiculatum L.

False smut Cerebella andropogonis

Ces.

Psidium cattleyanum (Sabine) Purple Guava

Thread blight Marasmiellus scandens

(Mass.) Denis & Reid

Rhynchospora aurea Valh.

Smut Cintractia spicularum

Rac.

Rosa species (Rose)

Leaf spot Cercospora puderi B.H.

Davis

Saccharum officinarum L. (Sugar cane)

Leaf spot Leptosphaeria sacchari

v. Breda de Haan.

Sesamum indicum .: (Sesame)

Leaf spot Cercospora sesami

Zimm.

Leaf spot Mycosphaerella species

Sesbania aculeata Poir.

Leaf blight Corticium solani (Prill.

& Delacr.) Bourd. &

Galz.

Solanum melongena L. (Brinjal)

Wilt Sclerotium rolfsii Sacc.

Sorghum almum Parodi (Columbus grass)

Leaf spot Phyllachora sorghi

Hohn

Spathodea campanulata Beauv. (African tulip tree)

Leaf blight Corticium solani (Prill.

& Delacr.) Bourd. &

Galz.

Marasmiellus scandens

(Mass.) Denis & Reid

Thread blight

Locally

common

1 1ecord

1 record

1 1ecord

1 record

Occasional

1 record

Occasional

1 record

Common

1 record

127

114577

119468

114575

115464

115455

1145766

114578

114579

114580

128 Gardens’ Bulletin, Singapore — XXII (1967)

Stenochlaena palustris Bedd. (Paku miding, Edible fern) _.

Thread blight Marasmiellus scandens 1 record —

(Mass.) Denis & Reid ©

Horse hair blight Marasmius equicrinus 1 record —

Mill. tye

Tabernaemontana coronaria Willd.

Cercospora tabernaemon- Common 115476

Leaf spot tanae H. & P. Syd.

Pink disease Corticium salmonicolor . 1 record a

Berk & Br.

Thread blight Marasmiellus scandens 1 record —

(Mass.) Denis & Reid

Theobroma cacao L. (Cocoa)

Brown pod Botryodiplodia theobro- locally —

mae Pat. common

Thunbergia grandiflora Roxb.

Leaf blight . Corticium solani (Prill.

& Delacr.) Bourd. &

Galz.

record -=

—

Zea mays L. (Maize)

Leaf blight Corticium solani (Prill. 1 record pele

& Delacr.) Bourd. &

Galz.

record 115477

—

Smut - Ustilago maydis (DC.)

Corda.

Acknowledgements

The writer wishes to thank the Director of Agriculture, Sarawak,

for permission to publish this list, and the Director and staff of the

Commonwealth Mycological Institute without whose help, in

identifying many of the species, this list could not have been

published.

OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

1. Annual Reports.

Many from 1909 onward remain available.

Prices variable.

2. The Agricultural Bulletin of the Malay Peninsula (Series I).

Only Nos. 3, 5, 7, 8 and 9 available at 20 cents each.

_ 3. The Agricultural Bulletin of the Straits and F.MLS. (Series HU).

Vols. 1-10, 1901-1911, monthly issues.

All parts available, except Vols. 1, 2, 5 (7), 6 (11) and 9.

Price: $5 per volume, Cents 50 per part. |

4. The Gardens’ Bulletin, Straits Settiements (Series D.

Vols. 1-11, 1912-1947.

Vols. 1 (1-5) January-May 1912 is issued under title of

Agricultural Bulletin of the Straits & F.M.S.

All parts available, except Vol. 1 No. 2, 10, Vol. 2, Vol.

3 No. 1-3 Vol. 6, Vol. 10 pt. 2 and Vol. 11 pt. 3.

Prices variable.

5. The Gardens’ Bulletin, Singapore (Series IV). |

~< Mols, 42-21, 1949+. *

All parts available keene 13 (1). Prices variable. Issues

are irregular.

Subscription will be entered against deposit.

Available on exchange.

6. Materials for a Flora of the Malay Peninsula, Monocotyledons.

Parts 1, 2 and 3 remain available.

7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H.

Addison.

Price $21.

(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison.

Price $15 (Photo-lithographic reprint).

8. A Revised Flora of Malaya. —

(a) Vol. 1, Orchids, by R. E. Holttum. Price $20.

(b) Vol. 2, Ferns, by R. E. Holttum. Price $20.

9. Wayside Trees of Malaya by E. J. H. ae 2 Vols. . 2

Price $25. ‘

[tems 1-6 can be obtained from the Director, Botanic Gardens, . A

Singapore. .

Items 7-9 can be obtained from the Government Printer, Govern: a

* ment Printing Office, Singapore. a

Prices quoted are in Malayan Dollars

Overseas postage is extra

OS 5 Sa C9 a

mT YEO YEA: ECE YE EEE YE

COMPLIMENTARY

GARDENS’ BULLETIN

SINGAPORE

Vol. XXII, Part II

27th December, 1967

CONTENTS

BENJAMIN C. STONE: Materials for a ee one of Freycinetia

_ (Pandanaceae) I - - - - -

Munir AHMAD ABID: A Revision of Symphorema. Verbenaceae

Betty MOLESworTH ALLEN: Malayan Fern Notes. V_ - -

B. V. SkvortTzov: Notes‘on the Flagellata of Hongkong -

~HarDIAL SINGH: Sclereids in Fagraea_ - - eine es

JAMES SINCLAIR: Notes on Sapotaceae_ - - + -

JAMES SINCLAIR: A note on Myriophyllum - . -

BENJAMIN C. STONE: Studies of Malesian Pandanaceae, I -

PP. S. ASHTON: Taxonomic Notes on Bornean Dipterocarpaceae,

peel

ANON: Burkill’s Dictionary of the Foonomic Products of the

uMalay Peninsula. Edition 2. - - > : 3

Price: $8

Published by Authori.

_-—s* Printed by Lim Bian Han, Governme

. 1967

~D ARBOREF

Wnt

pO RECEIVED

PARRY Leo 1968

PAGES

129 — 152

1d F71

173 — 185

187 — 191

193 —212

213 — 228

229 — 230

231-257

poe ~ SSZ

353

To be purchased at the Botanic Gardens, Singapore

Re HE, a

Zea KE HK <e—.

KE KO FF

Zea

Sse

my ee

Sa KK ORK

Re DK

Zag

Ky 7

Exe

en | a

aha

a ws :

Materials for a Monograph of Freycinetia

(Pandanaceae). I.

BENJAMIN C. STONE

Department of Botany, University of Malaya, Kuala Lumpur.

The following notes represent the results of studies in the

genus Freycinetia, a large and somewhat neglected group of

plants with perhaps as many as two hundred species. Previous

studies in the group have appeared by the present author in the

form of a review of the species of the Solomon Islands (Proc.

Biol. Soc. Wash. 76: 1-8. 1963), a proposed new species from

Morotai, Moluccas Is. (Journ. Arn. Arb. 43: 248-350. 1962),

and a review of the species known from Fiji, Samoa, and Tonga

(Proc. Biol. Soc. Wash., 80: 47-60. 1967). The work here reported

concerns species of Polynesia, Micronesia, Melanesia, and Malay-

sia, and consists chiefly of some shorter studies in particular

species, some nomenclatural adjustments, some reports of recent

collections, and the description of a few new taxa. Because it is

proposed to initiate further publications in this series, paragraphs

or sections are numbered for ready future reference.

CONTENTS.

1.1. The typification of the genus Freycinetia and remarks on the

Hawaiian species.

1.2. Freycinetia kiekie B. C. Stone, nom. nov.

1.3. Freycinetia distigmata B. C. Stone, nom. nov. —

1.4. Freycinetia collected in New Guinea in 1959 by L. J. Brass.

1.5. Freycinetia of Micronesia.

1.6. Preliminary studies on ‘leaf spectrum’ series in Freycinetia.

1.7. A note on Freycinetia boninensis Nakai.

1.1. The typification of the genus Freycinetia and remarks on the

Hawaiian species. ‘

The genus Freycinetia was established by Gaudichaud (Bot. Voy.

Freycinet . . . l! Uranie et Physicienne . . . Atlas, t. 41. 1826), with

three species: F. arborea, F. scandens, and F. radicans. The first

of these is selected as generitype. It is represented by a collection

made by Gaudichaud in Hawaii, and is well figured in the table

referred to above. Freycinetia scandens Gaud., a valid species, is

from Timor (the holotype is Zollinger 647), and F. radicans is from

Rawak, New Guinea (the holotype a Gaudichaud specimen).

130 Gardens’ Bulletin, Singapore — XXII (1967)

Freycinetia arborea Gaud. is the only species occurring in the

Hawaiian Archipelago. However, in Hillebrand’s Flora of the

Hawaiian Islands (1888: 453) the species is described under the

name ‘“‘Freycinetia Arnotti Gaudich.”’ This was proposed by

Gaudichaud in a later publication, of 1843 (Bot. Voy. La Bonite,

Atlas t. 36, 37), and was taken up by some later authors. It is

however nothing more than a slightly differing specimen of the

single widely distributed species in the Hawaiian group. The

alleged distinction, based on the number of stigmas per berry,

is unnatural; stigmatic number varies in this species generally, from

4 to 10 (or more), and the supposedly distinctive ranges (““4—6”

in F. Arnotti, ““6-9” in F. arborea) are mere samplings of the

inclusive range.

Hillebrand was aware that but a single species was to be found

in the Hawaiian flora, but he took up the latter name F. Arnotti

because “‘the older name F. arborea has to be abandoned as in-

appropriate and misleading.’ He was right in saying that the older

name was inappropriate, as the species is a woody climber, as are

all other known species of the genus, but with regard to the choice

of name we are retroactively required to use the earlier name.

Freycinetia arborea Gaud., in Freyc. Voy. Uranie et Physicienne,

Bot., Atlas t. 41. 1826. Solms-Laubach, Linnaea ns. 8: 100.

1878-79. Mann, Enum. Haw. Pits. n. 469. 1866. [Fig I: 1]

F. scandens sensu Hooker & Arnott, Bot. Beechey Voy. 87. 141.

(non Gaudichaud).

F. arnotti Gaud., Bot. Voy. Bonite, Atlas t. 36, 37. 1843.

F. arnotti Gaud., Bot. Voy. Bonite, Atlas t. 36, 37. 1843. Solms-

Laubach, Linnaea n.s. 8: 95. 1878-79. Hillebrand, Fl. Haw. Is.

453. 1888.

Holotype: HAWAIIAN ISLANDS: “In insulis Sandwich’

(Gaudichaud). PARIS; and isotype in the Delessert Herbarium

(GENEVA). The holotype of F. Arnotti is also marked simply

“In insulis Sandwich’’ (Gaudichaud).

Occurrence: A common species throughout the six major islands

of the Hawaiian Archipelago (Kauai, Oahu, Molokai, Maui, Lanai,

and Hawaii). It does not occur on the dry barren island of

Kahoolawe, nor on the tiny crescentic rim Molokini. It apparently

does not occur on the island of Niihau (neither Forbes, in his

Enumeration of Niihau Plants of 1913, nor St. John, in his paper on

Niihau plants in 1959, records the species). It is not known from

the Leeward Islands of the Hawaiian group (Nihoa, Necker, Laysan,

Lisiansky, Midway and Kure).

Habitat: Freycinetia arborea is most abundant in the lower and

middle forest zones (of Rock, 1913) or Vegetation Zones B, C, and

D (of Ripperton and Hosaka, 1942), where it may be found climb-

ing by its clinging roots on forest trees, or sometimes with the

loose stems falling and intermingling to form dense clumps of what

appears to be terrestrial growth especially on steep slopes. It

roots in old lava soils, either rocky or in rich loam. The largest

Stone — on Freycinetia 131

plants are found in moderate shade, but they tolerate windswept

exposed conditions on cliffs as well. In general, Freycinetia arborea

can be considered a rather common species between about 300

meters and about 1500 meters altitude.

Some recent collections of F. arborea:

HAWAIIAN ISLANDS: Kauai: Summit of Mt. Haupu, S.E.

side at 700 m alt., Dec. 1956, Stone 1645 (BISHOP, K, US); Oahu:

Puulupe, Kawailoa, Feb. 1937, Degener, Topping, Martinez &

Salucop 11070 (US): Waimano, Oct. 1935, Degener, Park &

Shigeura 10140 (US); Piko trail, Mokuleia, Yuncker 3316 (US);

Punaluu, Castle Trail, Stone 1166 (BISHOP); Manoa Cliff Trail,

Stone 3109 (BISHOP): Pupukea, Sept. 1958, Stone s.n. (BISHOP);

Maui: Puu Mukui, Sept. 1916, Hitchcock 14777 (US); without

locality, April 1911, Curran 83 (US); Hawaii: Volcano Road, July

1924, Setchell sn. (US); Honaunau Mountains, Sept. 1916, Hitch-

cock 14561 (US). |

An illustration of the berry and seed is provided (Fig. I).

1.2. Freycinetia kiekie B. C. Stone, nom. nov.

F. monticola F. B. H. Brown, Bishop Mus. Bull. 84: 29. pl. 19.

1931; not F. monticola Rendle, Journ. Linn. Soc. Bot. 45: 259.

1921.

Type locality: French Oceania: Marquesas Islands: Hivaoa

Island, Feani, F. Brown 870 (BISHOP).

This endemic species is also reported from Uapou Island on the

basis of Quayle 1135. It is quite distinct from Rendle’s species;

rather it is related to the above Hawaiian F. arborea. The new

name is taken from the Marquesan version of the widespread

Polynesian name for this genus. Subject to the normal changes (as

in glottal stops) the same word is used throughout Polynesia. In

Hawaii, it is ‘ie’ie.

1.3. Freycinetia distigmata B. C. Stone, nom. nov.

F. klossii Ridley in Kew Bull. 1926: 92. 1926; not F. Klossii

Ridley in Trans. Linn. Soc. Bot. 9: 236. 1916.

Type locality: Sumatra: Mentawi Island; Siberut, Boden-Kloss

14547.

Ridley inadvertently used the same epithet twice in the genus in

honor of Boden-Kloss. The species first so named, and to which the

name continues to apply, is from New Guinea. The Sumatran

Species requires a new name, given above in reference to its

generally 2-stigmatic carpels.

1.4. Freycinetia collected in New Guinea in 1959 by L. J. Brass.

During a year spent at the U. S. National Herbarium (Smith-

’ sonian Institution, Washington D.C.), I had the opportunity of

studying the collections of Freycinetia made by Leonard J. Brass in

1959, during the Fifth Archbold Expedition, sponsored by the

American Museum of Natural History with the aid of a grant from

the National Science Foundation. As is well known, the series of

132 Gardens’ Bulletin, Singapore — XXII (1967)

Archbold Expenditions have been invaluable in increasing our

knowledge of New Guinea, and Mr. Brass’s collections are generally

exemplary in quality. In fact, a large number of species of

Pandanaceae are based on Brass specimens.

The following account details collection data of species collected,

and includes the descriptions of two proposed new species of

Freycinetia based on Mr. Brass’s material. The text may be con-

sidered as a continuation of the papers by Merrill and Perry on

previous Pandanaceae from New Guinea, which appeared in Journ.

Arn. Arb. 20: 139-186. 1339; and 21: 163-175. 1940. I am grateful

to Dr. Lily M. Perry for her kind assistance during a visit to the

Harvard University Herbaria in 1961, and also to Dr. Richard A.

Howard, Director of the Arnold Arboretum.

Section Freycinetia !

In the interest of convenience there are included here a few

nomenclatural adjustments.

1. Freycinetia funicularis (Savigny) Merr., Interpret. Rumph.

Herb. Amb. 83. 1917; Merr. & Perry, Journ. Arn. Arb. 20: 155.

1939; 21: 168. 1940. [Fig. I: 2.]

Freycinetia strobilacea Bl. in Rumphia 1: 156. t. 39. 1835;

Schnizlein, Icon. 1: t. 74, f. 3-5. 1846; Solms-Laubach, Ann.

Jard. Bot. Buitenz. 3: 99. 1883; Warb. in Pflanzenr. 3 (IV. 9):

34. 1900; Martelli, Webbia 3: 315. 1910.

Freycinetia cyrtocarpa Kanehira, Bot. Mag. Tokyo 55: 295. f.

13. 1941. Syn. nov.

Type locality: Amboina. Recorded from the Solomon Islands

by Merrill & Perry. The type of F. strobilacea is the Rumphian

plate, on which is based the correct name taken up by Merrill.

The type of Kanehira’s species is from ‘‘Netherlands New Guinea

(Boemi, about 40 km inland from Geelvink Bay, at about 300 m alt.,

Kanehira & Hatusima 12680)’.Although this species may be dis-

tinct, (we are less than certain of all details of the plant pictured by

Rumphius) it seems to be synonymous. The few differences are

the smaller, nearly smooth leaves, the unarmed bracts, and the

slightly curving syncarps; but these do not carry much significance.

NEW GUINEA: Papua: Northern District, near Koreaf, Tuji

subdistrict, 23 September 1954, Hoogland 4794 (A): near Budi

Barracks, “‘urur’” in Onjob language, 25 August 1954, Hoogland

4552 (A, US); Morobe District, Sattelberg, 1000 m alt., 8 Nov.

1935, Clemens 832 (A). Finschhaven Peninscula, in beach woods,

19 October 1936, Clemens 4286 (A).

1 Article 22 of the International Code of Botanical Nomenclature

requires that the name of the section that includes the type species of

a genus repeat that name unaltered as its epithet. Consequently Sect.

Freycinetia must be used for the section including F. arborea Gaud.,

rather than the earlier proposed Sect. Pleiostigma Warb. in Pflanzenr. 3

(IV. 9): 28. 1900. This section is characterized by berries with mostly

5-10 stigmas.

--——-«

Stone — on Freycinetia 133

Both Hoogland and Clemens note that the bracts are “‘deep red’”’

or “flame color’. Solomon Islands specimens collected by L. J.

Brass were, according to Brass’s field data, characterized by “‘pink

bracts.”” These specimens have also the much less conspicuoulsy

dentate vegetative bracts which Kanehira mentions for his F.

cyrtocarpa.

2. Freycinetia salamauensis Merr. & Perry, Journ. Arn. Arb.

20: 152. pl. 1, f. 7. 1939. [Fig. I: 3.]

Type locality: North-eastern New Guinea, Morobe District,

Sattelberg, Salamaua (Clemens 28, A).

This species appears to be very close to F. biroi Warb. The

authors distinguished it by its shorter common peduncle, longer

pedicels, and the hispidulous angles and apex of the pedicels. It

is one of the species which is intermediate between the generic

sections, having for the most part berries with 3 stigmas; the range

is from 2 to 6 stigmas per berry.

NEW GUINEA: Eastern Highlands District, Kassam Gap, 1460

m alt., 28 Oct. 1959, climbing to middle spaces in Castanopsis-oak

forest; leaves glaucous on both sides; ripening fruits orange-red,

Brass 32285 (US, and to be distrib.).

3, Freycinetia sogerensis Rendle, Journ. Bot. 61. Suppl. 58.

1923.

Type locality: ‘““New Guinea, Sogere, 2,500 ft.’ (Forbes 75).

NEW GUINEA: Eastern Highlands District, Purosa, Okapa

area, 150 m alt., common root-climber on exposed tree -trunks in

rainforest; ripe fruiting heads orange, soft, ovoid, avg. 3.5 cm long,

2.5 cm broad, 2 October 1959, Brass 31851 (US, and distrib.).

4, Freycinetia stenodonta Merr. & Perry, Journ. Arn. Arb. 20:

151. 1939.

Type locality: Papua, Central Division, Ononge Road, Dieni

(Briss 3838).

NEW GUINEA: Morobe District, Gurakor, 640 m alt., climbing

to 3 m in oak-dipterocarp forest, flat leaves appressed to supporting

trunk, sterile, 6 May 1959, Brass 29452 (US); Sattelberg, 3200

ft. alt., Clemens 862i-bis (A).

A. readily recognizable species, with marked vegetative charac-

ters. The syncarps of the Clemens specimen are shortly oblong, and

they are noted as “‘red, fleshy.”

5. Freycinetia perryana B. C. Stone, sp. nov. (Sect. Freycinetia).

[Fig. I: 4.].

Scandens; caule lignoso subrobusto ad 1 cm crasso; foliis

ensiformibus basi imbricatis 39-47 cm longis, 16-18 mm latis,

apice anguste acuminatis, basi vix angustatis, amplexicaulibus.

margine basim et apicem versus et costa media in parte superiore

serrato-dentatis; auriculis scarioso-membranaceis, circiter 8-9 cm

longis, 8-9 mm latis, margine integerrimis, in parte superiore de-

currentibus (vel rotundatis?) adnatis, demum transverse fractis:

inflorescentia terminalia, spathis caducis non visis, pedicellis 5—6.5

cm longis ad 4 mm crassis, ubique denticulis crassis deltoideis

134 Gardens’ Bulletin, Singapore — XXII (1967)

erectis 0.3 -0.6 mm longis et latis armatis, pedicello apicem versus

subdense scabridis; syncarpiis 5, cylindraceis, circiter 8 cm longis,

et 1.7 cm latis; baccis circiter 9 mm longis et 3 mm latis, infra

4/5 carnosis apice coriaceis truncatis angulosis, areola stigmatica

pallide brunnea nitente annulos cinctos kermesinos, stigmatibus

vulgo 2 vel 4 rarius 5; fissuris stigmaticis angustis conspicuis;

seminibus anguste semi-ellipsoideis usque ad 1.5-1.7 mm longis,

fusco-rubris, rhaphide cum cellulis raphidophoris argenteis niten-

tibus, strophiolo nullo.

Holotype: NEW GUINEA: Eastern Highlands District, Mt.

Wilhelm, eastern slopes, 277Q m alt., 29 July 1959, L. J. Brass

30765 (US, and isotypes for distrib.).

This species is similar to Freycinetia trachypoda Merr. & Perry

and also to F. tafaensis Merr. & Perry; differing however from the

former in having smaller leaves, coarsely scabrid (rather than his-

pidulous) pedicels, and more numerous syncarps; and from the

latter in having somewhat longer leaves with apically narrowed,

entire auricles (rather than truncate, spinulose auricles), longer

more coarsely scabrid pedicels (to 6.5 cm long, rather than only

1.5 cm long), and larger more numerous syncarps per inflorescence.

It is a pleasure to dedicate the species to Dr. Lily M. Perry,

long an associate and colleague of Dr. E. D. Merrill, long associated

with the Arnold Arboretum, and well-known for numerous studies

on Melanesian plants.

Section Oligostigma Warb.

6. Freycinetia angustissima Ridley, Journ. Bot. 24: 359. 1886;

Martelli, Webbia 3: 309; 1910; Rendle, Journ. Bot. 61. Suppl. 57.

1923; Kanehira, Bot. Tokyo 55: 251. f. 1. 1941; Merr. & Perry,

Journ. Arn. Arb. 21: 163. 1940.

Freycinetia stenophylla Warb. in K. Schum. & Lauterb. FI.

Deutsch. Schiitzeb. i. d. Sitidsee, Nachtr. 53. 1905; Martelli,

Webbia 3: 315. 1910; Journ. Arn. Arb. 10: 137. 1929; White,

Journ. Arn. Arb. 10: 201. 1929; Merr. & Perry, Journ. Arn.

Arb. 20: 141. 1939.

F. sp. Solms-Laubach, Jard. Bot. Buitenz. Bull. 3: 101. 1883.

Freycinetia polyclada Merr. & Perry, Journ. Arn. Arb. 20: 141.

p.. 1, 1. 3. rose.

Type locality: New Guinea, Sogere, 2000 ft. (Forbes).

The type of Freycinetia stenophylla was from the Torricelli

Mountains, Kaiser-Wilhelmsland, 600 m alt. (Schlechter 14525);

and that of F. polyclada was from North-eastern New Guinea,

Central Division, Mount Tafa, 2400 m alt. (Brass 4961). The latter

species was synonymized by Merrill and Perry in their second

paper of 1940 on New Guinea Pandanaceae (Journ. Arn. Arb. 21).

NEW GUINEA: Eastern Highlands District, Mt. Otto, south

slopes, 2200 m alt., common in disturbed forest, climbing to 7 m

not much branched; fruit-heads usually solitary, sometimes 2 or 3,

ovoid or subglobose, orange, about 2.5 cm in diameter when ripe;

14 Aug. 1959, Brass 31050 (US and distrib.); same loc., 2680

m alt., plentiful in mossy beech forest, 7 August 1959, Brass

Stone — on Freycinetia 135

30874 (US, unicate). Papua: Boridi, in forest, fruit to 2 cm long,

16 September 1935, Carr 13182 (A, BM). Morobe District;

Sattelberg, 12 May 1938, Clemens 8221 (A).

7. Freycinetia archboldiana Merr. & Perry, Journ. Arn. Arb.

20: 147. pl. 1, f. 10. 1939.

Freycinetia inouei Kanehira, Bot. Mag. Tokyo 55: 254. f. 4.

1941. Syn. nov.

Type locality: Papua, Central Division, Bella Vista (Brass

5458).

The type of Freycinetia inouei Kaneh. was from Dallmann in

then Netherlands New Guinea (Kanehira & Hatusima 12163).

Kanehira stated that the species is distinguished from F. arch-

boldiana by its ‘“‘much thicker syncarp and more numerous drupes’”’

(sic; i.e. berries), but these differences seem more due to individual

variation than to a species difference. A comparison of an’ isotype

(A) with the type of F. archboldiana (also at A) failed to reveal any

further differences of significance.

Kanehira mentions that the leaves are rigid, erect, and tinged

purplish when fresh. The seeds show a conspicuous raphe but lack

a strophiole.

8. Freycinetia arfakiana Martelli, Webbia 3: 173. 1910.

Type locality: “‘Netherlands New Guinea, Mt. Arfak, Putat’’

(Beccari 821).

NEW GUINEA: Eastern Highlands District; Arau, 1400 m alt.,

frequent in castanopsis-oak forest, climbing to several meters and

sparingly branched, bracts white with pale green tips, 14 Oct.

159, Brass 32070 (US and distrib.).

This specimen is referred here with some hesitation; the leaves

are mostly 25-29 cm long and 44.5 cm broad (shorter than those

of Beccari’s type by 10-15 cm), entire, with auricles 5-6 cm long

and 8-9 mm broad, these entire and deciduous; syncarps terminal,

narrowly cylindric, on smooth pedicels 2-3 cm long, about 3 mm

thick, four in number, each syncarp to 3 cm long and 9 mm thick;

berries with mostly 2-4 stigmas but very immature.

9. Freycinetia brassii Merr. & Perry, Journ. Arn. Arb. 20:

145. 1939. [Fig. I: 6.]

Type locality: Papua, Fly River, 528 Mile Camp (Brass 6651).

NEW GUINEA: Morobe District; Gurakor, 640 m alt., scramb-

ling to 10 meters in rainforest, 5 May 1959, Brass 29431 (US,

unicate).

This specimen matches the original description well except that

the inflorescence is lateral (i.e. pseudo-lateral). Since this is a

variable character, (sometimes both terminal and ‘lateral’ inflores-

cences occur On the same plant), and the other features correspond

so closely, there is no apparent reason for keeping it distinct. It

should be remarked that Freycinetia lacinulata Kanehira is

scarcely to be distinguished from F. brassii. Kanehira stated that

the differences lie in the ellipsoid, rather than oblong-cylindric

136 Gardens’ Bulletin, Singapore — XXII (1967)

syncarps of the former, and the brownish to orange-red color of

the berries; these differences are however unimportant. Brass’s

notes mention that the syncarps of his no. 6657 (cited by Merrill

and Perry under F. brassii) were orange-red; while those of his no.

6927 were brown.

10. Freycinetia crucigera Kanehira, Bot. Mag. Tokyo 55: 252.

1941.

Type locality: ““Netherlands New Guinea; Dallmann, 600 m alt.”

(Kanehira & Hatusima 12286).

NEW GUINEA: Eastern Highlands District; Mt. Michael, north-

eastern slopes 200 Om alt., climbing to 2 meters in castanopsis-

oak-nothofagus forest; fruits immature; seeds purple; 9 September

1959, Brass 31471 (US); Purosa, Okapa area, 1950 m alt., frequent

in mixed rainforest, climbing to several meters, leaves thin and

crinkled, flowering bracts green, fruits unripe, 20 September 1959,

Brass 31593 (US).

The first specimen cited fits the original description and figure

very well, except for the terminal infructescence; and the second

specimen bears smaller leaves, but of the same proportions, and

the infructescence is either lateral or terminal, indicating this

character’s variability. The principal difference between this species

and Freycinetia beccarii seems to be the relatively broader leaves

and the seeds without strophiole, of the present species.

11. Freycinetia fibrosa Martelli, Journ. Arn. Arb. 10: 138. 1929;

Merr. & Perry, ibid. 21: 164. 1940.

F. sogerensis sensu Merr. & Perry, loc. cit. 20: 147. 1939, non

Rendle 1923.

Type locality: Papua; Gulf District, Vailala River, Hokoro, 100

m alt. (Brass 1048).

NEW GUINEA: Eastern Highlands District; Mt. Elandora,

Kratke Mts., 2130 m alt., climbing on forest undergrowth, fruits

immature; 18 Oct. 1959, Brass & J. D. Collins 32164 (US); Purosa,

Okapa Area, 1950 m alt., climbing to 10 meters in rainforest,

common, flowering bracts thin, greenish-white; 21 September 1959,

Brass 31632 (US).

13. Freycinetia kanehirae B. C. Stone, nom. nov.

Freycinetia platyphylla Kanehira, Bot. Mag. Tokyo 55: 261. f.

10. 1941; not F. platyphylla Merrill, Philipp. Journ. Sci. Bot.

13: 267, 19 ia.

Type locality: ‘Netherlands New Guinea: Dallman-Patema,

600 m alt.;”’ (Kanehira & Hatusima 12285).

The required new name commemorates Professor Ryozo

Kanehira, formerly of the Department of Agriculture of the Kyushu

Imperial University in Fukuoka, Japan, a student of the

Pandanaceae and authority on the flora of Micronesia. There is

also a Pandanus kanehirae Martelli from Palau. Merrill’s species

is from the Philippines.

Stone — on Freycinetia 137

Freycinetia kanehirae is charcterized by its very large leaves

(120 cm long, 7-8 cm broad), its large cylindric syncarps of

numerous slender berries, and its smooth fruit-pedicels. It is

reminiscent of F. tessellata Merr. & Perry of the Solomon Islands,

but differs in its smooth rather than scabrid pedicels; and of F.

ponapensis Martelli of Ponape (Caroline Islands), but differs in

the obtuse leaf tips and much longer syncarps.

14. Freycinetia radicans Gaudich., in Freycinet, Voy. autour

du Monde...sur les corvettes 1’Uranie et le Physicienne... Bot.

432; et Atlas Bot. t. 43. 1826; emend. Warb. in Pflanzenr. 3 (IV. 9):

34. 1900. [Fig. I: 5.]

Type locality: Is. Rawak, near West Irian (West New Guinea).

The type is Gaudichaud’s collection.

NEW GUINEA: Morobe District; Kaindi, 2250 malt.

occasional in Nothofagus forest, scandent to 20 meters or more;

10 May 1959, Brass 29666 (US).

This species is imperfectly known; but the Brass collection cor-

responds with what little is known.

15. Freycinetia spinellosa Kanehira, Bot. Mag. Tokyo 55: 265.

f. 12. 1941.

Type locality: ‘‘Netherlands New Guinea: Dallmann (Kanehira

& Hatusima 12507).

NEW GUINEA: North-western New Guinea: Sorong, Kp.

Baroe, in forest, nom. vern. “riligino”, 28 July 1948, Djamhari

389 (A).

A species notable for its long narrow leaves and spinulose-

margined auricles.

16. Freycinetia tafaensis Merr. & Perry, Journ. Arn. Arb. 20:

146. 1939.

Type locality: Papua: Central Division, Mt. Tafa, 2400 m alt.;

(Brass 5001).

NEW GUINEA: Eastern Highlands District, Mt. Elandora,.

Kratke Mts., 2530 m alt., scrambling to 1.5 meters in scrubby

growth of summit, flower bracts orange; 18 Oct. 1959, Brass and

Collins 32144 (US). Morobe District, Mt. Kaindi, 2200 m alt.,

frequent in Nothofagus forest, climbing to 12 meters, branches

spreading, flower bracts orange-yellow; 23 May 1959, Brass 29733.

(US). :

17. Freycinetia elegantula B. C. Stone, sp. nov. (Sect.

Oligostigma). [Fig. I: 7.]

Liana nana ad arbores adpressa, caulis lignosis, 2-6 mm diame-

tro, laevis, in sicco rugulosis; ramis angustis, radicibus fibrosis

emittens; folia minima, ovata, 15-22 mm longa, 5—9 mm lata, acuta,

longitudine nervosa (nervis utrinque circiter 8) margine et in costa

dorsale spinulosa vel fimbriatulosa (spinulis 1-1.3 vel 2 mm longis

et 1-2 per mm); auriculis caducis minutis inconspicuis amplectenti-

bus; inflorescentia lateralia, caule breve (8-10 mm longo) bracteato

138 Gardens’ Bulletin, Singapore — XXII (1967)

terminans; bracteis caulae basin versus deltoideis acuminatis mar-

gine integerrimis; bracteis caulae apicem versus foliaceis; bracteis

floraleis viridis margine integerrimis; pedicellis 3, apicem versus

minute et persparse breve-spinulosis vel sublaevibus, 5-7 mm

longis; syncarpiis 3, subglobosis, ad 12 mm diametro; baccis

carnosis, circiter 4 mm longis, 1.5-3 mm latis, apice rotundatis;

stigmatibus 1-2; seminibus anguste semi-ovatis, circiter 0.7 mm

longis, strophiolo nullo (?).

Holotype: NEW GUINEA: Eastern Highlands District, Arau,

1400 m alt., 8 October 1959, L. J. Brass 31945 (US, and isotypes

for distrib.).

An extraordinary, tiny and elegant species related to Freycinetia

beccarii but with smaller yet proportionately broader leaves with

conspicuously spinulose margins and costa.

From tne available material, which is without flowering or

fruiting spikes, I judge this species to be the same or very nearly the

same as that described in English only in 1939 by A. D. E. Elmer

under the name Freycinetia brevifolia (Leafl. Philipp. Bet. 10:

3771. 1939). This name is invalid, since it is without a Latin dia-

gnosis and printed after 1935; it is also a later homonym, since

there is the earlier valid name Freycinetia brevifolia Martelli

(Webbia 3: 186. 1910). Elmer’s sole collection was from Camp

Lambi, Nueva Ecija Province, Luzon. The leaves are described

as 25 mm long, 20 mm broad, ovate- elliptic, and “margine ser-

ratuli-spinulosis, supra distincte nervatis”. This corresponds very

closely to the above description of Freycinetia elegantula except in

the greater width. It is possible that but a single species is con-

cerned, occurring both in New Guinea and the Philippines.

FIGURE I

Details of features of some species of Freycinetia.

1. Freycinetia arborea. Berry in profile and top view, x 5; seed x 5S,

and enlarged x 20. (From Stone 3109).

2. Freycinetia funicularis. Staminate plant; stamens x 5; male inflorescence

x 4; leaf apex x 4+. (From Hoogland 4552).

3. Freycinetia salamauensis. Berry in profile and top view x 5; seed

enlarged x 20; portion of cellular structure of strophiole, greatly

enlarged. The embryo is 0.2 mm thick; endosperm dark pink; seed

walls crimson, 0.05 mm thick; raphe and strophiole translucent.

(From Brass 32285).

4. Freycinetia perryana. Berry in profile and top view x 5; seed, two views,

x 5. (From holotype, Brass 30765).

3. 7 nerd radicans. Berry in profile and top view x 5. (From Brass

9666).

6. Freycinetia brassii. Berry in profile and top view x 5; leaf apex x 4.

(From Brass 29431).

7. Freycinetia elegantula. Berry in profile and top view x 5; seed x 5; leaf

x 1; and infructescence x 1. (From holotype, Brass 31945).

139

Stone — on Freycinetia

)

FIGURE I

‘

See caption on page 10

140 | Gardens’ Bulletin, Singapore — XXII (1967)

1.5. The genus Freycinetia in Micronesia.

Four species have been recorded from Micronesia (Caroline and

Marianne Is.), but the number is here reduced to three, based on

the reduction of Kanehira’s proposed Freycinetia almonoguiensis

to synonymy with F. villalobosii Martelli emend. St. John.

Freycinetia ponapensis Martelli is very closely similar to F.

mariannensis but after long consideration I here maintain

their specific difference. However, Kanehira’s proposed F.

carolinensis of Palau is identical with F. mariannensis (not with

F. ponapensis, aS Kanehira himself later decided). Hosokawa des-

cribed the Ponape plants as F. mariannensis var. microsyncarpia,

while reducing F. ponapensis to the synonymy of F. mariannensis:

but he indicated (Journ. Jap. Bot. 13: 191. 1937) that both his

newly described and the typical varieties occurred in Ponape;

while only the typical var. (mariannensis) was eslewhere in Micron-

sia. His discussion of the leaf tesselations, said to be distinctive

as a taxonomic feature by both Merrill and Kanehira, discloses that

it is too variable a feature for any taxonomic use. Hosokawa did

not, however, investigate the auricles, leaf spectrum, seeds, or

pedicels, all characteristic structures of major taxonomic import-

ance. The following key will show some of the specific character

correlations of the taxa here regarded as species:

Key to Micronesian species of Freycinetia

1. Stigmas mostly 1-3 per berry (Sect. Oligostigma); leaves thick

coriaceous, mostly 3-5 cm broad, the apex acuminate with some

degree of negative or concave curvature; auricles rounded gradu-

ally at apex, unarmed; syncarps stout cylindric.

2. Pedicels of syncarps sparsely scabrid at their distal ends; seeds

nearly straight, about 1.3 mm long, the raphe scarcely or not

exceeding the embryo; syncarps mostly 6-7 cm long.

Freycinetia ponapensis

2. Pedicels glabrous, smooth; seeds slightly lunulate, about 1.7 mm

long, the raphe slightly exceeding the embryo; syncarps usually

8-9 cm long. Freycinetia mariannensis

1. Stigmas mostly 3-10 per berry (Sect. Freycinetia); leaves rather thinly

brittle coriaceous, mostly 1-3 cm broad, the apex tapered, not

acuminate, to a narrow swordlike point; auricles abruptly rounded

at apex and there with a few prickles; syncarps slender cylindric.

Freycinetia villalobosii

Sect. Oligostigma

1. Freycinetia mariannensis Merr., Philipp. Journ. Sci. Bot. 9:

48. 1941; Kanehira, Bot. Mag. Tokyo, 49: 185. 1935; Journ. Dept.

Agric. Kyushu Imper. Univ. (Enum. Micron. Pl.) 4 (6): 259. 1935;

Bot. Mag. Tokyo 51: 906, f. 62-63. 1937; Hosokawa, Bull.

Biogeogr. Soc. Jap. 5 (2): 132: 1934; Glassman, Journ. Arn. Arb.

29: 178, in obs. 1948. [Fig. If and III: 1-4, Pl. I and IL]

Freycinetia carolinensis Kanehira, Bot. Mag. Tokyo 49: 185.

f. 17. 1935; Journ. Dept. Agric. Kyushu Imper. Univ. (Enum.

Micron. Pl.) 4 (6): 259. 1935; Bot. Mag. Tokyo 51: 906.

in synon. 1937.

141

Stone — on Freycinetia

FIGURE II.

_— nw

0 oN

SEA

524

OOO

Bee

Bag

ae .

40”

a on

g.aaq

Fao

a o§

igh

a. fon

ww ,

GC.

a x

Yo .

S38

from Palau, Kanehira 2359).

Freycinetia mariannensis. 1-9:

(prophyll) to first f

in profile x 5. 12:

Gardens’ Bulletin, Singapore — XXII (1967)

142

FIGURE III

See caption on page 143

Stone — on Freycinetia 143

Type locality: Guam, Marianas Is.

Distribution: Marianas Islands, Caroline Islands (Palau, Truk).

GUAM: North of Yona, Rodin 577 (US); 1.5 miles north-west

of Yona, Rodin 759 (US); 2 miles S. of North-West Field,

Steere 12 (US); between Agana and North-West Field, Steere 15

(US); Mt. Alifan, Bryan 1207 (BISHOP, US); Mt. Almagosa,

Hosaka 3173 (BISHOP, US); Talofofo Valley, Stone 3970, 3971

(BISHOP, GUAM, L, US); Agat hills, Stone 4217 (BISHOP,

GUAM, L, US); Fena River, Stone 4491, 4492 (BISHOP, GUAM,

L, US); Guam, without locality, Costenoble (US, isosyntype);

Moore 291 (US).

ROTA: Sabana, high plateau, in forest, Stone 5206a, b

(GUAM); Feb. 1941, Tuyama s.n. (TOKYO).

SAIPAN: April 1930, Shizuwo Momose (TOKYO). (For

previous collections see also Kanehira, Enum. Micron. Pl. 1935).

The seeds are slightly lunulate, or sometimes straight, about

1.6-1.7 mm long, and 0.25-0.37 mm broad, the raphe moderately

slender, white, with scattered shining raphidophorous cells, exceed-

ing the seed proper slightly at both ends, about 0.1 mm broad; no

strophiole is present.

Kanehira originally reduced his Freycinetia carolinensis to F.

ponapensis; however, the glabrous pedicels serve to associate it

rather with F. mariannensis under this revised treatment.

2. Freycinetia ponapensis Martelli in Kanehira, in Bot. Mag.

Tokyo 48: 129. 1934; Kanehira, Journ. Dept. Agric.

Kyushu Imper. Univ. (Enum. Micron. Pl.) 4 (6): 259.

1935; Hosokawa, Trans. Nat. Hist. Soc. Formosa 33: 18.

1943; Glassman, Bishop Mus. Bull. 209: 111. 1952. [Fig.

Ii: 5-9, Pl. WII.]

FIGURE III.

1-4. Freycinetia mariannensis. Floral details.

1: small part of a pistillate spike, showing two flowers, each consisting

of a carpel with 2 papillate stigmas, and a few (3-5) staminodes at

base. RC=raphidophorous cells. Magnified x 10.

2: apex of a carpel, showing the papillae of the stigma, small epidermal

cells, and the interior RC. Divisions of scale=0. 1 mm.

3: small part of a staminate spike, showing stamens; note RC in

filaments. Magnification x 10.

4: pollen grains to same scale as 2. (All from Stone 5206a-b).

5-9. Freycinetia ponapensis. Floral details and berry.

5: cellular detail from filament margin; to same scale as fig. 2.

6: stamens with pistillodes at base, and top view of one pistillode.

Magnification x 10.

7: berry in profile and top view, with seed, all x 10.

8: seed, showing raphe, x 20.

9: epidermis of seed, greatly enlarged. (All from Stone 5479 and

5480).

144 Gardens’ Bulletin, Singapore — XXII (1967)

F. mariannensis var. microsyncarpia Hosokawa, Journ. Jap.

Bot. 13: 191. 1937; Kudoa 5: 79. 1937.

Type locality: Ponape; Patapat (Ledermann 13245).

Distribution: Ponape and Kusaie.

Specimens examined: PONAPE: Mt. Nanalaut, near summit,

June 1957, Stone 2034 (BISHOP): Headlands of Pilen-kiepw

River, 400 m alt., March 1957, Stone 1728 (BISHOP): Mt.

Nginani, summit, 2550 ft. alt. Feb. 1965, Stone 5479, 5480

(BISHOP, GUAM, UNIV. MALAYA); without locality, Jan.

1915, July 1915, Koidzumi s.n. (TOKYO).

KUSAIE: Innemu River, 130 m alt., Dec. 1945, St. John 21449

(BISHOP); Limes, near Lele, 100 m alt., May 1957, Stone 1903

(BISHOP): Jan. 1941, Tuyama (TOKYO).

The St. John specimen and Stone 5479 are both staminate

plants, thus permitting the following description:

Bracts of male inflorescence yellow at base, apically scarlet;

spadices 3, on pedicels 2.5—-3 cm long and c. 3 mm thick, smooth,

cylindric, about 6 cm long and 1 cm thick, the stamens numerous

and densely congested, with long slender filaments 3.5-4 mm

long; white anthers about 0.7 mm long.

The seeds of this species appear to differ from those of F.

mariannensis; they are ellipsoid, nearly straight, about 1.2-1.3 mm

long, 0.35-0.39 mm broad, raphe 0.1 mm broad, white, barely or

not exceeding the embryo, strophiole lacking.

The differences in the seeds, and in the pedical scabridity, serve

to differentiate these species, which had been considered identical

by Hosokawa. It should be noted that the male inflorescence:

bracts of all specimens were yellow.

Sect. Freycinetia

3. Freycinetia villalobosii Martelli in Kanehira, Bot. Mag.

Tokyo 48: 128, as Villalobosensis. 1934; Kanehira, Bot. Mag.

Tokyo 49: 195. 1935; Journ. Dept. Agric. Kyushu Imper. Univ.

(Enum. Micron. Pl.) 4 (6): 259. 1935. Nom. emend. H. St. John,

Philipp. Journ. Sci. 88 (3): 403. 1960 [Fig. IV: 4—6; PI. IV.]

Freycinetia almonoguiensis Kanehira, Bot. Mag. Tokyo 49:

186. f. 18. 1935; Journ. Dept. Agric. Kyushu Imper. Univ.

(Enum. Micron. Pl.) 4 (6): 259. 1935.

Type locality: Palau Is., Babeldaob Is., in forest at 200-300

m alt., ““Urwaldahnl. Mittelwald 10-15 m h. mit viel Fiederpal-

men, viel Unterholz; . . . Hochkeltternd, Kolben griin, B. glanzend

dunkel griin, Rind grau,” Feb. 1914 (Ledermann 14323).

The type of Kanehira’s proposed species was from “Palau,

Almonogui; coll. Feb. 1933, MNisida 2775”. Almonogui (also:

spelled Arumonogui) is the headland to the north of the large

bay or inlet at the mouth of the Gaspan R. in Babeldaob (Babel-

thuap) Island, on the west coast toward the southern end. It is

probably not very far from the locality where Ledermann obtained

the specimen upon which Martelli based the species description.

Stone — on Freycinetia 145

FIGURE IV.

1-3. Freycinetia boninensis. 1— outline of syncarp and part of pedicel,

natural size. 2— part of leaf base, showing part of blade and part of

auricular sheath, natural size. 3— berry, x 5. (From isotype).

4-6. Freycinetia villalobosii. 4- base of leaf, showing auricular sheath,

natural size. 5— top view of berry, x 5. 6— profile of berry, with

seed to scale (but immature), x 5. (From Hosokawa 6902).

146 Gardens’ Bulletin, Singapore — XXII (1967)

Distribution: Endemic to Palau Islands, West Caroline Is.

In his original description, Martelli stated: “I named this plant

in honour of the Spanish navigator Arrecipes Ruy Lopez de

Villalobos, who discovered the groups of the Islands in 1543 and

died at Amboina.’’ Somehow he attached the Latin place-name

ending to this man’s name, and published it as Villalobosensis.

The necessary correction of orthography was made recently by

St. John.

The only apparent differences which might serve to distinguish

Freycinetia almonoguiensis are the larger syncarps (7.5 cm vs.

10 cm); and an alleged slight difference in leaf-tip shape. Not

mentioned by Kanehira is the pedicel scabridity or lack of it; in

his figure, the pedicels appear smooth and glabrous. In F. villa-

lobosii, there is a minute scabridity distally on the pedicels. One .

specimen studied (Takamatsu 1548, cited below) does have glabr-

ous pedicels. Certainly the syncarp sizes are too similar for any

significant difference; the leaf tip shape does not seem different at

all; and the pedicel scabridity is a somewhat variable character.

All told, it seems difficult to consider the available specimens

as representing more than one species, and thus I have united

them; a viewpoint in concordance with that of T. Hosokawa

(in herb.).

On the other hand, Freycinetia villalobosii needs further com-

parison with certain species of the Philippines (the nearest Malay-

sian land area, using the term Malaysian in the sense of van Steenis).

Martelli mentioned the similarity of such Philippine species as

Freycinetia negrosensis, F. superba, and some of Elmer’s later,

unvalidated species. It remains to be determined whether the Palau

form is an endemic species (as does appear likely) or if it may be

a Micronesian outlier of a Philippine species.

Specimens examined: PALAU: Babeldaob Island; Almonogui,

Feb. 1933, §. Nisida 2775 (TOKYO, isotype); Aimiriik (Aimelik),

July 1933, Kanehira 2366 (TOKYO); Garamiscan, or Almiokan,

River, swamp-forest, Fosberg 25729 (US); Mt. Luisalumonogui

(=Almonogui), Hosokawa 6902 (A, BISHOP, TAIWAN); near

Ngaldok Lake, in thickets, Hosokawa 9290 (TAIWAN); Gara-

sumao, Takamatsu 1548 (BISHOP); Banks of Garamiscan River,

observed only, Jan. 1963, Stone; Garamiscan, Sept. 1937, Tuyama,

(TOKYO).

1.6. Preliminary studies on ‘leaf spectrum’ series in Freycinetia.

[Fig. II.]

As pointed out by Melville (Proc. Linn. Soc. Lond. 164: 173- |

181. 1953) a sequence of leaves in ontogeny, follows a characteristic

pattern, which may be termed the leaf spectrum. As an example,

Stone — on Freycinetia 147

selected leaves from Freycinetia mariannensis are shown in Fig. II,

beginning with the first foliar organ produced on the lateral

shoot emergent immediately below an infructescence. This has

on occasion been called a “‘vegetative bract’’. It differs, as do also

to a lessening extent the subsequent organs (Fig II. 2-9), from

fully developed foliage leaves in several major features; it is un-

armed (as are nos. 2-5); it is much shorter (as are nos. 2-9); it

lacks the characteristic auricles (as do nos. 2-6, although in nos.

5 and 6 they are partially developed); it has a central, ventral

longitudinal channel, the result of its being pressed closely against

the stem which terminates in an infructescence, and this channel

is thickly ridged on both sides; its midrib is inconspicuous (this is

also true of no. 2); and it is considerably thicker, especially at the

base, than foliage leaves. The spectrum shown here would be con-

tinued by an organ which, although with a blade not much longer

than no. 9, would have to be considered a mature foliage leaf.

Since in the Pandanaceae there is also a continuous series of

organs from foliage leaves to inflorescence bracts, it may be neces-

sary to include the latter in the leaf spectrum. Only a few species,

in both Pandanus and Freycinetia, produce what may be termed

a lateral inflorescence. This is a morphological structure deserving

much more intensive study. In some Freycinetiae, both these

‘lateral’ and the more common ‘terminal’ inflorescences are pro-

duced by the same plant. In ‘lateral’ inflorescences the foliage leaf

stage is omitted, the transition being from the ‘“‘vegetative tract’

directly to the “‘inflorescence bract’’s

A study of leaf spectra of various species is planned as living

plants are encountered in the field.

1.7. A note on Freycinetia boninensis Nakai. [Fig. IV: 1.3.]

Previously published accounts of this species failed to give a

clear idea of the auricles, which have proved to be necessary in

the taxonomy of the genus. During a short study visit at the

Department of Botany, University of Tokyo, I was enabled,

through the courtesy of Prof. Hiroshi Hara and Dr. Y.:Kimura, to

study material of this species. The following notes and accompany-

ing figures partly clarify the morphology of this species.

Freycinetia boninensis (Nakai) Nakai, Journ. Jap. Bot. 11: 153.

1935.

Type locality: Bonin Is.

First described as a variety of F. formosana, this species is very

similar to Formosan material, but a critical review of both species

is still required. The following notes add to available information

concerning the Bonin Is. plant:

148 Gardens’ Bulletin, Singapore — XXII (1967)

Leaves upwards of a meter in length; at base stoutly toothed on

the margins; the apex caudate-flagellate, toothed; but intervening

middle margins unarmed. Syncarps 3 or more per inflorescence on

minutely scabridulous peduncles, the scabridity on all the surfaces

and along the whole length of the peduncle. Bracts bright red-

orange. Stigmas 4-8, or rarely 9-10, per berry.

The syncarps reach a length of 9.5 cm, but perfectly mature

ones have not been seen; the diameter is about 1.8 cm.

Figure IV: 1-3 show auricles, syncarp, and berry.

Literature Cited

ForBES, CHARLES N. 1913. Notes on the Flora of Kahoolawe and

Molokini; (and) An Enumeration of Niihau Plants. Bishop

Museum, Occas. Pap. 5 (3).

St. JOHN, HAROLD. 1959. Botanical Novelties on the Island of Niihau.

Hawaiian Islands. Pacific Plant Studies 25. Pacific Science

13: 156-190.

RIPPERTON, J. C., and E. Y. Hosaka. 1942. Vegetation Zones of Hawaii.

Rock, JOSEPH F. 1913. The Indigenous Trees of the Hawaiian Islands.

Honolulu, privately printed.

Stone — on Freycinetia 149

Plate I.

Freycinetia mariannensis. A fruiting specimen, in riverine forest along

the Ugum River, Guam. (Photo: B. C. Stone).

150 Gardens’ Bulletin, Singapore — XXII (1967)

Plate II.

Freycinetia mariannensis. A specimen with young spikes to show bracts

and leaf apex; from Rota is., Stone 5206a.

Stone — on Freycinetia 151

Plate III.

Freycinetia ponapensis Martelli, in mossy forest at the summit of Mt.

Nginani, Ponape, alt. 2550 ft.; ripe syncarps red. (Stone 5480).

152 Gardens’ Bulletin, Singapore — XXII (1967)

Plate IV.

Freycinetia villalobosii Martelli emend. St. John. Palau (Hosokawa 9290).

A Revision of Symphorema. Verbenaceae.

MuNIR AHMAD ABID *

University of Sind, Hyderabad, West Pakistan

INTRODUCTION

The genus Symphorema was founded by Roxburgh in 1798 on

S. involucratum, a species described from Coromandel, the eastern

coast of southern India. The generic description is brief, but it is

accompanied with a coloured plate (t. 186) and a detailed descrip-

tion of the species. The plant is stated to flower “‘in February,

March and April’.

In 1805 Jussieu established the genus Analectis which has

been declared to be congeneric with Symphorema. Wallich (Cat.

1828 p. 47, No. 1734) placed the genus “‘Analectis Juss, apud

Vahl’ (1810) between Congea and Symphorema, a statement

recorded by Meisner (1843 p. 200) on the authority of Wallich,

since Meisner was not able to examine the specimen. It is not sure

whether Wallich considered the genus as published by Jussieu

(1805), or merely Vahl’s use of the name in connection with his new

species Analectis speciosa (1810). Steudal (1840) listed the genus

as of uncertain position. Vahl’s binomial is found listed in

Moldenke’s Résumé (1959 p. 234) as a synonym of S. involucratum

Roxb. It is desirable to know the locality and the collector of the

specimen.

Blanco in his Flora Filipinas (1845) 284 described a philippine

plant as a species of Lauraceae, namely Litsaea luzonica, which

Llanos (1858) referred to the genus Symphorema. In 1863

Turczaninow established Sczegleewia on Cuming’s No. 648 from

the Philippines, with S. luconiensis as the type. This genus was

recognized by Bentham and Hooker (1876) to be also congeneric

with Symphorema Roxb., though they made no reference to

Blanco’s species.

Presently the genus consists of only three species: S$. involucratum

Roxb., §. polyandrum Wight and S. luzonicum (Blanco) F-Villar.

The types of the first two species were from India, and the type of

the third one was from the Philippines. Those described from India

are closely allied in their floral characters and in the shape of their

leaves and in having stellate tomentum on their leaves and

branchlets.

The basionym of S$. luzonicum (Blanco) F-Villar is Litsaea

luzonica Blanco (1845) based on a specimen from Arayat in the

island of Luzon, Philippines. Llanos and others like Fernandez-

Villar (1880), Perkins (1904) and Merrill (1906) have recognized

* Colombo Plan Fellow, Botanic Gardens, Singapore, 1965.

154 Gardens’ Bulletin, Singapore — X XII (1967)

that it is a true Symphorema. However, before Blanco’s species was

made the basionym of the new combination under Symphorema,

Sczegleewia luconiensis Turcz. (1863) and Symphorema glabrum

Hassk., (1965) were published, apparently based on two different

collections of Cuming. This Philippine species is the only one of

this genus which has glabrous and entire leaves and glabrous twigs.

The leaves are oblong-elliptic and superficially resemble those of

the species of some genera of Lauraceae. Hence, it was regarded

by Blanco as a lauraceous species.

Distribution

Of the two species which occur in India, S. involucratum has

the widest distribution, occurring alsc in Burma and northern

regions of Thailand, while the other S$. polyandrum, is restricted

to India only. These two are confined to the area lying between

70-105° East longitude and 7—27° North latitude (See Maps 2 & 3).

S. luzonicum is apparently limited to an area in Luzon and a few

neighbouring islands lying southwest of Luzon, between 120—124°

East longitude and 11-19° North latitude. But there is a collection

from the Tanimbar Islands lying to the south of the Moluccas,

between 130-135° East longitude and 5-10° South latitude (See

Map 4). The collector has not given any clue of its being cultivated

or wild.

Involucral Bracts in Symphoremeae

Inflorescence in Symphoremeae consists of cymes disposed on

axillary and terminal panicles. Each cyme consists of a peduncle

and a cluster of flowers surrounded by involucral bracts. These

bracts are generally much longer than the floral calyces, the only

exception being in Sphenodesme eryciboides Kurz where bracts are

comparatively smaller Fig. 1-C). In Symphorema and Sphenodesme

these bracts are always free to the base, while in Congea they are

either free or united to form a cup at the base. Bearing these

general characters of the flower-heads in mind, it would be easy

to understand the many variations discussed below.

The structure of the involucral-heads in Symphorema and

Sphenodesme shows that these two genera are closer to each other

than either is to Congea. In both these genera the bracts are always

6, subequal, and free to the base. But, as shown in my paper on

Sphenodesme (Gard. Bull. Sing. Vol. XXI, Part III), the flower-

head consists of two dichotomous branchlets with main axis ending

in a flower. Each of these branchlets ends in a bract with one

axillary flower and bears two lateral bracts each with a flower. One

can often see that these two main bracts are the largest and Dest

developed. These are regarded by Wight (1850 p. 215) as true bracts

while the other four as subsidiary ones or “‘bracteoles’’. Since these

“‘bracteoles” are often almost as large as the real bracts, it is not

easy to distinguish between them without a careful examination.

- =—

Munir — Symphorema 155

160°E

EQUATOR

° e

g0°E q0°e 100°e No € 160°C

Map. 1. Distribution of Genus Symphorema (————— ).

160°E

Cancer,

EQUATOR

gore q0°e 100° 110°E

160°E

Map 2. Distribution of S. involucratum (——— ).

156 Gardens’ Bulletin, Singapore — XXII (1967)

Ws Ta 1 60°e

ial

Tr. of

Concer.

EQUATOR

g0°e q0°E 100° 10°E 20°E 130°€ HOPE 150° 160°E

ee Se Sean ee eee eere

Map 3. Distribution of S. polyandrum (———— ).

asi NOE. ed 1 60°E

30°N

a ee a ee ee eee ee Lit oe eS OR SS 2 ieee

Cancer,

20°N

2 C

° ’

AJ e

Ion ° 4

()

EQUATOR

Yew.

10°S 7 os

20°s

g0°¢ q0°e 100°%e 110°E

Map 4. Distribution of S. luzonicum (————).

Munir — Symphorema 157

However, that Wight’s view is correct is readily seen in the

species of Congea. Here the involucral bracts show different stages

of development not present in the other two genera. The bracteolar

head consists of not more than 4 distinct members nor less than 3

to a cyme. Moreover, the bracts may be entirely free or united

at base to form a cup surrounding the flowers in the head. But

Congea forbesii and its var. ridleyana show an intermediate stage

in the reduction of the bracts from 6 to 4. In both these taxa

there are six involucral bracts, all free to the base, of which two

are so reduced that they have been often overlooked (Fig. 1-D).

3cm

JURAIMI1 DEL.

Fig. 1. Variation in Involucral Bracts of Symphoremeae.

A, Symphorema. B, Sphenodesme. C, Sphenodesme eryciboides.

D, Congea forbesii. E, Congea griffithiana. F, Congea chinensis.

G, Congea tomentosa. H, Congea connata.

158 Gardens’ Bulletin, Singapore — XXII (1967)

In C. griffithiana and C. rockii these two reduced bracts are

absent and the four present are large and free to the base (Fig. 1-E).

The number of bracts to a cyme in C. chinensis and its variety

is also four, but they are united below into a cup (Fig. 1-F), but

in C. pedicellata, C. vestita and C. tomentosa, two of the four

bracts are connate half-way to make them sub 4; otherwise the

bracts are free (Fig. 1-G). In C. siamensis, C. connata and C.

velutina the bracts are united to form a definite cup at the base, but,

while in S. siamensis they are 3 or sub 4, in C. connata and C.

velutina they are always three (Fig. 1-H).

Systematic Position of Symphorema and allied genera

Roxburgh established the genus Symphorema in 1798 as a mem-

ber of the group “Octandria Monogynia’’. Subsequently, in 1819,

the same author published (posthumously) the genus Congea under

“Didynamia Angiospermia, Natural Order Vitices Juss.” In Rox-

burgh’s manuscript itself this second genus was named Roscoea,

a name, owing to the delay in publishing the manuscript became

unusable from 1804, when it was taken up for another taxon

(see introduction of my paper on Congea, Gard. Bull. Sing. Vol.

XXI, Part II[). Endlicher (1836) placed Symphorema and Congea

(with Sphenodesme as a synonym) under the tribe Aegiphyleae

Endl. Almost simultaneously Meisner (1836) created a new tribe

Symphoremeae exclusively for Symphorema and Congea with

Sphenodesme included in the latter as its synonym. Presl (1844)

published two monotypic genera Calochlamys and Viticastrum and

placed them under the tribe Lantaneae Endl.

Schauer (1847), who made a careful revision of Verbenaceae

and recognized Sphenodesme Jack as distinct from Congea Roxb.

and Calochlamys capitata Presl and Viticastrum racemosum Presl

as species of Congea and Sphenodesme respectively, divided the

family into three tribes and many subtribes. The tribes are as

follows: Verbeneae, Viteae (or Viticeae) and Avicennieae. In this

classification Symphoremeae Meisner is made a subtribe of Viticeae

for three genera: Symphorema, Congea and Sphenodesme. In

1850 Wight who had previously suggested (1840) that the two

genera Symphorema and Congea with Sphenodesme as the synonym

of the latter, should form a distinct family, examined the status of

these tribes of Schauer. After a detailed analysis of these divisions

he concluded that Symphoremeae cannot be placed as a subtribe of

Viticeae, being quite distinct from the latter though in many

respects allied to Avicenneae. He therefore retained the group as

a tribe by itself between Viticeae and Avicenneae, and stressed

some characteristics which, in my opinion, would justify raising

Symphoremeae and Avicenneae to the status of subfamilies, or

even of families, if smaller families are desirable. Bentham and

Hooker (1876) included in Verbenaceae the anamalous genus

Phryma Linn. which Schauer had made the type of Phrymaceae

Schauer (DC. Prodr. XI, 1847 p. 520), and subdivided the family

Munir — Symphorema 159

thus constituted into 8 tribes as follows: Phrymaeae, Stilbeae,

Chloantheae, Verbeneae, Viticeae, Caryopterideae, Symphoremeae

and Avicenneae.

Briquet (1897) who, like Schauer, had in 1895 treated Phryma

as forming a distinct family, did not include the genus under

Verbenaceae, but raised the remaining seven tribes of Bentham

and Hooker into higher groups (probably subfamilies) as:

Stilboideae, Chloanthoideae, Verbenoideae, Viticoideae, Caryop-

teridoideae, Symphoremoideae and Avicennioideae. In addition

he subdivided Verbenoideae, Chloanthoideae and Viticoideae into

6, 3 and 4 tribes (?) respectively, with their names ending in eae.

Lam (1919 & 1921) who dealt with Indonesian Verbenaceae

regarded Briquet’s names ending in oideae and eae as tribes and

subtribes respectively.

Moldenke (1959) like Schauer and Briquet retained Phrymaceae

as valid; but in addition separated Stilboideae, Symphoremoideae

and Avicennioideas as distinct families viz: Stilbaceae, Symphore-

maceae and Avicenniaceae. The remaining three groups Verbe-

noideae, Viticoideae and Caryopteroideae Moldenke retained as

subfamilies of Verbenaceae, but added a new subfamily Nyctan-

thoidea consisting of MNyctanthes and Dimetra, which were

generally regarded as members of Oleaceae.

Without going into the propriety either of including the two

oleaceous genera in Verbenaceae or of separating Stillbaceae from

it, there seems to be justifiable reason either to retain

Symphoremoideae and Avicennioideae as two subfamilies or, if

microfamilies are desirable, to follow Moldenke who raised them

to the rank of two distinct families.

Moldenke characterises these two families as follows: placenta

central, free; ovules apically attached, pendulous, orthotropous;

while the other two, Stilbaceae and Verbenaceae as follows:

placenta axial, carpels 2 or 4, each bearing 2 ovules, but one

carpel often more or less aborted, the carpel-edges turning back

from the middle of the ovary to the midrib of each carpel, making

false partitions; ovules basally or laterally attached, anatropus or

hemi-anatropous, apotropous. The distinctions between the first

two families are as follows:

Avicenniaceae

Ovary incompletely 4-celled; cotyledons folded; growth

in diameter of trunks and stems brought about by concentric

layers of mestome rings; branches, branchlets, and twigs

commonly terete, prominently nodose and articulated; typic-

ally saline lagoon shrubs or trees.

Symphoremaceae

Ovary 2-celled to the middle; cotyledons not folded; trunk

and stem growth normal; branches, branchlets, and twigs

commonly more or less tetragonal, not articulate; typically

woody vines (climbers) of non saline situations.

160 Gardens’ Bulletin, Singapore — XXII (1967)

ACKNOWLEDGEMENT

It is my pleasant duty to record my gratitude to the following:

to the Government of Singapore for extending my Fellowship

which enabled me to complete the research in the genera of the

Symphoremeae group of Verbenaceae; to Mr. H. M. Burkill,

Director of Gardens, Singapore, and to Dr. Chew Wee-Lek, Keeper

of the Herbarium, for putting at my disposal the facilities of the

Singapore Botanic Gardens; to Dr. C. X. Furtado for his general

guidance; to Che Juraimi bin Samsuri for making the drawings

published here; to the typist who patiently typed the manuscript.

My thanks are due to the Directors and Curators of the institu-

tions mentioned below for the loan of herbarium specimens:

1. Arnold Arboretum, Harvard University, 22 Divinity

Avenue, Cambridge 38, Massachusetts, U.S.A. (A).

2. Botanisches Garten und Museum, Belin-Dahlem, Germany.

(B).

3. Forest Herbarium, Bangkok, Thailand. (BKF).

British Museum of Natural History, London, S.W. 7,

England. (BM).

Botanic Gardens, Brisbane, Queensland, Australia. (BRI).

Botanic Gardens, Calcutta, India. (CAL).

Forest Research Institute, Dehra Dun, India. (DD.)

Royal Botanic Garden, Edinburgh, Scotland. (E).

Royal Botanic Gardens, Kew, Richmond, Surrey, England.

(K).

10. Forest Research Institute, Kepong, Selangor. (KEP).

11. Ruijksherbarium, Schelpenkade 6, Leiden, Holland. (L).

12. New York Botanical Garden, Bronx Park, New York,

U.S.A. N.Y).

13. National Herbarium & Museum, Manila, Philippines.

(PHN).

14. Herbarium, Department of Botany, University of

California, Berkeley 4, U.S.A. (UC).

tre. ae SN

Symphorema Roxb.

Symphorema Roxb., Corom. Pl. II (1798) 46 t. 186 & FI. Ind. II

(1832) 262; Endl., Gen. Pl. (1836) 638, no. 3716; Meisner, Gen.

Pl. I (1836) 292 & II Comment. (1843) 200; Walp., Repert. Bot.

IV (1844) 116; Schauer in DC, Prodr. XI (1847) 621; Wight, Ic.

Ind. Or. IV, 3 (1849) 13 & Ill. Ind. Bot. II (1850) 216; Pfeiffer,

Nomencl. Bot. II, 2 (1874) 1326; Benth & Hook., Gen. Pl. II, 2

(1876) 1159 p.p. (excl. Decadontia Griff.); Kurz, For. Fl. Burma,

II (1877) 253 p.p. (altera parte Sphenodesme); Clarke in Hook.

f., Fl. Br. Ind. IV (1885) 599; Trim., Fl. Ceyl. III (1895) 362;

Briq. in Engl. & Prantl, Pflanzenf. IV, 3a (1879) 179 p.p. (excl.

Decadontia Griff.); Cooke, Fl. Bomb. II (1908) 434; Lam,

Munir — Symphorema 161

Malay. Verb. (1919) 329 & Bull. Jard. Bot. Buitenz. III, Ser.

Ill (1921) 99; Haines, Bot. Bihar & Orissa, III] (1922) 724;

Gamble, Fl. Madras II, 6 (1924) 1103; Dop in FI. Gén. Indoch.

IV (1936) 896; Mold., Résumé Geogr. distr. & Syn. (1959) 234,

343, 351 & 403.

Analectis Juss. in Jaume St. Hil. Expos. fam. II (1805) 362;

Vahl in Danske Selsk. Skrivt. VI (1810) 90.

““Litsaea’’ sec Blanco, FI. Philip. Ed. 2 (1845) 284 & Ed. 3, U (1878)

162.

Sczegleewia Turcz. in Bull. Soc. Imp. Nat. Mosc. 36, If (1863) 212.

TYPE SPECIES: S. involucratum Roxb.

Scandent shrubs. Stem almost cyclindrical sometimes faintly

quadrangular in the older parts. Leaves opposite, simple, petioled,

entire or irregularly crenate-toothed or serrate, reticulate, unicos-

tate, pubescent-tomentose or completely glabrous. /nflorescence

paniculate, consisting of pedunculate 7-flowered cymes with an

involucre of 6 accrescent bracis; flower sessile. Calyx 3-8 lobed,

almost obovoid, accresent in fruit. Corolla white; tube cylindrical,

glabrous, or with a villous ring in the throat; lobes 6-16, subequal,

narrowly oblong. Stamens 6-18, exserted; filaments filiform,

glabrous; anther oblong-elliptic, dorsifixed, cells parallel. Ovary

obovoid, 2 or imperfectly 4-celled, 4-ovuled, the ovules pendulous,

but only one fertile; style exserted, filiform, glabrous; stigma dis-

tinctly 2-fid. Fruit capsular, dry, included in the accrescent calyx,

l-seeded. Seed erect; cotyledons fleshy.

Distribution: India, Burma, Thailand, Philippines and Tanimbar

Islands (south of the Moluccas).

Under Symphorema, Bentham et Hooker (1876) and Briquet

(1897) have given as synonym Decadontia Griff. which was based

on D. coerulescens Griff., a synonym of Sphenodesme griffithiana

Wight. These authors, though included Sczegleewia Turcz. as a

synonym of Symphorema, overlooked Litsaea luzonica Blanco, and

so made no refrence to the literature where the latter was published.

Key to the Species

1 (a) Leaves oblong-elliptic, entire, glabrous; calyx 3-4-lobed, with

dense, long, appressed hairs within; corolla tube densely villous

in the throat; ovary often covered sparingly with stellate-hairs

_in the upper half, glandular at apex; stigma lobes peltate:

o_o) SR eee terre on ee 7! Ain ee ete 2 a S. luzonicum

(b) Leaves ovate or sub-orbicular, crenate to almost dentate, stellately

pubsecent or tomentose beneath; calyx 6-8-lobed, glabrous in

the tube; corolla glabrous all over; ovary glabrous; stigma lobes

Uo” Si ee ee | z

2 (a) Leaves pubescent beneath; involucral bracts elliptic, entire, papy-

raceous, deciduously pubescent; calyx-lobes 6, obtuse, glabrous

within; corolla 6-8-lobed; stamens usually 6-8, rarely 9, hardly

longer than the petal lobes; ovary not glandular at apex: (India,

Burma and Thailand) ............. RI ee S. involucratum

(b) Leaves densely tomentose beneath; involucral bracts obovate,

usually entire, sometimes coarsely toothed, densely tomentose;

calyx lobes 6-8, acute, tomentose on both sides; corolla lobes

12-16; stamens 12-18, much exserted; ovary slightly glandular

Po ee Pe ree ot: eee S. polyandrum

162 Gardens’ Bulletin, Singapore — X XII (1967)

DESCRIPTION OF SPECIES

1. Symphorema luzonicum (Blanco) F-Vill. in Blanco, Fl. Philip.

Ed. 3, IV, Novis Append. (1880) 162; Perk., Frag. Fl. Philip.

(1904) 3; Merr. in Philip. Journ. Sc. I Suppl. I (1906) 112, FI.

Manila (1912) 400, Sp. Blancoanae (1918) 335 & Enum.

Philip. Pl. III (1923) 406; Mold., Résumé Geogr. distr. & Syn.

(1959) 185, 236, 243 & 350. — Fig. 2.

co)

t » 4

GY, Viz

‘a\

JURAIMI DEL.

Fig. 2. Spmphorema luzonicum (Llanos 69).

A, Fertile twig. B, Flower. C, Flower vertically cut open. D,

Cyme showing flowers and bracts.

Munir — Symphorema 163

S. cumingianum Briq. in Engl. and Prantl, Pflanzenf. 1V, 3a

(1897) 180: nom. tantum.

S. glabrum Hassk. in Flora XLVIII (1865) 402; Lam. Verb.

Malay. (1919) 330 & in Bull. ard. Bot. Buitenz. Ser. 3, IIL (1921)

99.

“§. involucratum Spreng” sec. Llanos in Mem. Acad. Cienc.

Madrid III, 4 (1858) 507 reprinted in Blanco FI. Filip Ed. 3, IV,

(1880) 105; Mold., Résumé Geogr. distr. and Syn. [sub

Symphorensa] (1959) 351: pro syn.

S. luzoniense (Turcz.) Benth. and Hook. ex Vidal, Sinop.

Atlas (1883) 36 t. 75, fig. F; ibid, Phan. Cuming. Philip. (1885)

135 & Rev. Fl. Vasc. Filip. (1886) 212.

Litsaea luzonica Blanco, Fl. Philip. Ed. 2 (1845) 284. & Ed.

3, If (1878) 162. basionym.

Sczegleewia luconiensis Turcz. in Bull. Soc. Imp. Nat. Mosc.

XXXVI, 2 (1863) 212: basionym of S. luzoniense.

Balibai Blanco in FI. Philip. (1837) 406: nom. vern.

Neotype: Blanos 69, in Herb. A.

S. luzonicum is an unique species of this genus not only because

it is found in the Philippine and possibly also in the Tanimbar

Islands, south of the Moluccas, but also because its leaves are

glabrous and entire; calyx 3-4 lobed with dense, long, appressed

hairs within; corolla tube densely villous in throat; stigma lobes

peltate.

Branchlets almost cylindrical or faintly quadrangular in the older

parts, glabrous excepting the grey-pubescent apical parts. Leaves

coriaceous, oblong-elliptic, obtuse or obtusely acuminate at apex,

sub-truncate to rounded at base, entire, glabrous, more or less

nitid above, dull below, up to 24 cm long, 10 cm abroad; main

Jateral nerves 4 pairs, the lower two pair very prominent and long:

petioles 5-10 mm long, glabrescent. /nflorescence grey pubescent.

Cymes 7-flowered; involucral bracts broadly obovate-elliptic, nar-

row towards base, obtusely rounded at apex, softly pubescent,

accrescent, up to 3.5 cm long, 1.6 cm broad; peduncles thick,

pubescent, 2. 5-5 cm long. Calyx + tubular, 7-9 mm long, slightly

accrescent, almost urceolate in fruit, distinctly 3-4 lobed, some-

what 2-lipped, pubescent or minutely tomentose without, hirsute

with appressedly distinct long hairs within; lobes subequal, 1-3

mm long. Corolla 6-7 lobed, glabrous in the tube, with a villous

band in throat, sparingly puberulent without, 6.5-9 mm long;

lobes oblong, softly pubescent without, glabrous within, 3-5 mm

long. Stamens 8-16, inserted in the villous corolla throat, much

exserted; filaments filiform, up to 12 mm long, often united into

pairs at base; anthers oblong-elliptic, glabrous. Ovary almost

obovoid, imperfectly 4-celled, glandular at apex, provided in the

upper half with deciduous hairs, glabrous below, 2. 5-3 mm long;

style long exserted, filiform, up to 1.5 cm long, glabrous, bilobed at

apex; lobes rarely armed with a few hairs; stigma peltate,

glandular. Fruit almost globose, glabrous, enclosed in the per-

sistent, accescent calyx, 4-6 mm in diameter.

164 Gardens’ Bulletin, Singapore — XXII (1967)

PHILIPPINES: Luzon, /locos, Pasuquin (Felix sub For. Bur. No.

30,914: SING); Burgos (Ramos sub Bur. Sc. No. 27,122: A

& BRI); Mt. Quebrada (Edano sub Philip. Nat. Herb. No.

17,797: L; 17,784: -A); Mt. Pico de Loro (Edano sub Philip.

Nat. Herb. No. 17,960: A). Cagayan, Alcala (Clemens sub Bur.

Sc. No. 17,534: B, BM & SING). Pangasinan, Bolinao (Clemens

18,176: SING). Nueva Ecija, Ecija (Villavicencio sub For. Bur.

No. 23,654: A); Zambales, Subig (Merrill 2,075: NY). Bataan.

Mt. Mariveles, Lamao River (Borden sub For. Bur. No. 2,549:

NY & UC; Meyer sub For. Bur. No. 2,516: NY & SING;

Williams 707: A & NY; Barnes 343: BM & NY; Whitford 2:

NY; Pascual sub For. Bur. No. 28,766: UC). Protacio, Brandt's

Garden (Steiner 1.011: PNH, Prob. cultivated). Pampanga,

Arayat (Merrill 1,421: A & NY). Tayabas, Attmonan (Gregory

81: K); Lucena (Merrill 2,416: NY). Laguna. Calauan (Cuming

648: A, isotype of Sczegleewia luconiensis Turcz & BM):

Los Banos (Merrill spec. Blanco 467: A, BM, K, L & NY;

Elmer 9,418: E; Tamesis sub For. Bur. No. 11,924: BM: Steiner

1,742: L, PNH & SING, Prob. cultivated); Mt. Makiling (Ilagan

sub Phil. Nat. Herb. No. 35.488: PNH & SING; Mabanag sub

Phil. Nat. Herb. No. 9,601: L, SING & UC; Banaga sub Phil.

Nat. Herb No. 33.398: A & L; Susara sub Phil. Nat. Herb. No.

37,313: BM & PNH; Baker 3.042: BM & SING; Sinclair

9,470: SING; Amihan sub Phil. Nat. Herb. No. 40,320: PNH:;

Sulit sub Phil. Nat. Herb. No. 8,320: PNH; Ebron sub Phil.

Nat. Herb. No. 34,236: PNH: Abadilla sub Phil. Nat. Herb.

No. 35,389: PNH; Holman 53: A). Rizal, Manila, San Mateo

(Vidal 501 & 848: A, K & L); Montalban (Topping sub Bur. Sc.

No. 5,228: A; Loher s.n.: UC; Elmer 17,419: A, BM, L, PNH

& UC; 17,429: A, BM, K, L & UC; 17,467: A, BM, L, PNH

& UC); Antipolo (Ahern 40: A; Merrill 1,934: A; 1,334: NY);

Paningtingan (Loher 13,444: UC); Bosoboso (Ahern sub Bur.

Sc. No. 1,172: NY; 4,454: CAL; 4,455: CAL & K; 4,456:

CAL). loc. incert. (Ahern sub For. Bur. No. 2,654: NY & SING;

Llanos 69: A, neotype), Albay, loc. incert (Cuming s.n.: L,

holotype of S. glabrum Hassk.; BM & K, isotypes). Mindoro,

Paluan (Merrill 954: A, NY & SING). Busuanga (Weber 1,546:

A, E & PNH).

INDONESIA: Tanimbar Island, Jamdena, Ranarmoje River,

Norkese (Borssum 3,283: L & SING).

Apparently there are two collections of Cuming; one bears

No. 648 from Calauan in Laguna Province of Luzon which formed

the basis of Sczegleewia luconiensis Turcz. and the other is un-

numbered which formed the basis of Symphorema glabrum Hassk.

What appears to be the duplicate of the latter is in Kew and is

recorded as being from the “‘Province of Albay” in Luzon. How-

ever, the No. “‘648, Luzon’’ is later added in a different handwriting.

Since No. 648 is from the Laguna Province, this Albay specimen

cannot be its duplicate. Apparently Turczaninow spelt the epithet

of his species with c—cedilla, which is equivalent to z in Latin

as shown in the combination made by Vidal.

)

Munir — Symphorema 165

As there is no Blanco type available Llanos 69 is taken as the

neotype, for it is on this specimen that Blanco’s Litsaea luzonica

(1845) was correctly interpreted by Llanos in 1878.

Since Bentham and Hooker (1876) included Sczegleewia Turcz.

as a synonym of Symphorema and quoted Cuming No. 648, Vidal

followed the old custom and attributed the new combination S.

luzoniense (Turcz.) to Bentham and Hooker; but under the Code,

Vidal has to be ackonwledged as the actual author of the combina-:

tion.

F-Villar states that the species is very common in Central Luzon

and he saw a living plant on the bank of river of Mount San

Francisco and at San Mateo in the Manila Province.

Since Lam (1919 & 1921) did not know Litsaea luzonica Bianco

(1845) and admitted erroneously 1835 as the date of publication of

Hasskarl’s $. glabrum, he took the latter name as the correct one

for S. luzonicum (Blanco) F-Vill. However, as shown above in the

synonymy, Hasskarl’s species was published only in 1865, twenty

years after Blanco’s species and as such the latter must take

precedence.

Merrill (1923) credits the species for Palawan but the only

specimen I found is from Paluan in the Mindoro island, collected

by Merrill lkimself.

Moldenke (1959) erroneously gives Symphorema (Symphorensa)

involucratum as having been published by Sprengel for the

Philippine plants, though the latter merely recorded Roxburgh’s

species of India. Similarly Moldenke retains S. grossum Kurz as

a good species of the genus, but as shown in my previous paper it 1s

a later synonym of Sphenodesme eryciboides Kurz.

Borssum 3,283 from Tanimbar islands, south of Moluccas, is

the only record of this species or even of the genus collected so

far outside the Philippines. Since the collector has not given any

indication of its being cultivated or wild, a further investigation

is necessary to confirm this unusual distribution.

2. Symphorema involucratum Roxb. Corom. Pl. I (1798) 46 t. 186

& Fl. Ind. II (1832) 262; Spreng., Syst. Veg. IL (1825) 208;

Wight, Ic Ind. Or. IJ, 1 (1840) 5 t. 362; Walp., Repert. Bot. IV

(1844) 116; Voigt, Cat. Pl. Calc. (1845) 470; Schauer in DC.,

Prodr. XI (1847) 621 exclud. descript. et Wall. No. 1740; Dalz.

& Gibs., Bomb. FI. (1861) 199; Thwaites, Enum. Pl. Zeylan.

(1861) 242; Kurz, For. Fl. Burma, II (1877) 254; Clarke in

Hook. f., Fl. Br. Ind. IV (1885) 599; Trim., Fl. Ceyl. III (1895)

363; Brig. in Engl. & Prantl, Pflanzenf. [V 3a (1897) 180; Cooke,

Fl. Bomb. II (1908) 434, Craib in Kew Bull. (1911) 445; Haines,

Bot. Bihar and Orissa III (1922) 724; Gamble, Fl. Madras II, 6

(1924) 1104; Dop in FI. Gén Indoch. IV (1936) 898 fig. 93/2

to 5; Fletcher in Kew Bull. (1938) 441; Mold., Résumé Geogr.

distr. & Syn. (1959) 164, 166, 167, 178 & 234. — Fig. 3.

166 Gardens’ Bulletin, Singapore — X XI1 (1967)

JURAIMI DEL.

Fig. 3. Symphorema involucratum (A Roxburgh s.n.; B-E Wight 2,586).

A, Part of an inflorescence. B, Flowering twig. C, Cyme with its

bracts and flowers. D, Flower. E, Ibid. longitudinally cut open

to show ihe internal organs.

Analectis speciosa Vahl in Danske Selsk. Skrivt. VI (1810) 90

(fide Moldenke).

This species is easily recognized by its leaves being pubescent

beneath; involucral bracts elliptic, entire, papyraceous; calyx-lobes

6, glabrous within; corolla 6-8 lobed; stamens usually 6-8. hardly

longer than the petal lobes and ovary non-glandular.

Branchlets cylindrical, stellately sub-tomentose when young,

later pubescent. Leaves ovate or almost elliptic, sub acute at apex,

usually cordulate at base, sometimes almost rounded, irregularly

crenate-serrate or toothed, puberulent above when young, later

almost glabrous, densely and stellately pubescent beneath, 10 by 6

Munir — Symphorema 167

cm on fertile twigs; main lateral nerves 4—S pairs; petioles usually

5-15 mm long, pubescent. Cymes 7-flowered; involucral bracts 6,

elliptic, entire, papyraceous, deciduously pubescent, 1-1.7 by

0.2-0.5 cm at anthesis of the flower, later accrescent, up to 3.5 by

1.5 cm; peduncles up to 7 cm long, stellately pubescent. Calyx

tubular, 6-lobed, slightly accrescent, 4-5 mm long, stellately to-

mentose without, glabrous within; lobes + 1 mm long, obtuse,

glabrous within. Corolla 6-8 lobed, white, glabrous except at few

villous hairs outside at the tip of its lobes, 6-8 mm long; lobes

linear oblong, obtuse, 2-3 mm long; tube 4-5 mm long. Stamens

6-8, rarely more, hardly longe rthan the corolla lobes; filaments

filiform, glabrous; anthers oblong-elliptic. Ovary obovoid, glabrous,

non-glandular; style exserted, filiform; stigma distinctly 2-lobed,

lobes porrect, cylindrical. Fruit subglobose, glabrous, 4-6 mm in

diameter, enclosed in accrescent, papyraceous, persistent calyx.

INDIA: Maharashtra, Khandala (Herb. Blatter No. 6,112: K); loc.

incert. (Dalzell s.n.: DD). Mysore, North Kanara; near Yellapore

(Fernandes 220: A; Talbot 52: K & s.n.: DD).Konkan (Stocks &

Laws s.n.: A, BM, CAL & L). Kerala, Travancore (Bourdillon

537; CAL). Madras, coromandel (Roxburgh s.n.: E, holotype);

Naggur hills (Wight 2.586 & 2,587: E; Cleghorn s.n.: CAL & E);

loc. incert. (Wight 909: E & NY; 2,303: A, CAL, K & L);

Shevaroy hills (Perrottet 487: CAL). Andhra Pradesh, Cuddapoh

hills (Beddome 6,520 & 6,522: BM: 6,521 & 6,523: SING; 39:

CAL; s.n.: CAL); Godavari (Beddome 6,519: BM & SING):

Vishakhapatnam, at Ragupalien (Barber 1,573: K). Bihar &

Orissa, Ganjam Distr., Mahendragiri (Fischer & Gage 83: CAL);

Barkuda (Carter 1,507: CAL); Valiki (Gamble 13,745: CAL):

Kuaduli (Haines 4,944: K); Puri (Haines 2,542: DD); Seeta-

koond! (Madden 663: E). Madhya Pradesh. Chanda Distr.

(Duthie 9,687: DD); Jabalpore (Leg.? s.n.: BM). Nagaland

(Beddome s.n.: BM).

CEYLON: Trincomali (Glenie sub Thwaites C.P. No. 3,645: BM,

probably cultivated).

BURMA: Shan State, Mong Noi, near Salween, alt. 3,000 ft.

(Robertson 316: K); Taunggyi (Khalil s.n.: CAL); loc. incert.

(Collect 435: CAL). Magwe, by the Kinmondaing (Rogers 910:

CAL & E). Minbu, Chichaung, Mon Chaung (Parkinson 15,733:

DD & E); Mezali (Parkinson 15,750: DD & E); Nwamadaung

hills (Aubert & Gage s.n.: CAL). Pegu, loc. incert. (Brandis

881: CAL; Kurz 1,040; 2,392 & 2,399: CAL). West Central

Burma, loc. incert. (Kingdon-Ward 21,796: BM). Tharrawaddy,

Myaung Waingale (Lace 2,794: CAL, E & K):; Taungnyo

Reserve, alt. 100 m. (Rogers 274: CAL).

168 Gardens’ Bulletin, Singapore — X XII (1967)

THAILAND: Northern regions, Muang Lamphun, (Larsen &

Hansen 861: A; Smitinand 4,169: BKF); Muang Phrae, at

Hooey-Kamin (Kerr 991: K); Muang Fang, alt. 700 m. (Kerr

5,234: E); Chawn Fong, alt. 270 m. (Kerr 2,932: E & K).

Roxburgh’s type specimen was apparently collected by a Telugu

man “‘Moodu”’ (= Muthu ?), under vernacular name “‘ceroodiddu”’

which Roxburgh rendered as “‘Suroodoo’.

Thwaites (Enum. Pl. Zeylaniae, 1861 p. 242) records S. involu-

cratum Roxb. from Trincomalee, Ceylon, which was brought to

him by one Glenie, but so far no one has found it indigenous

there. The specimen was probably from a cultivated plant.

Moldenke who has recorded it as wild in Ceylon, has apparently

followed Thwaites.

Dop (1936) included Congea paniculata Wall. as synonym of

S. involucratum Roxb. However, this binominal was a nomen

nudum for Sphenodesme involucrata (Presl) Robinson vat. pani-

culata (Clarke) Munir (see my paper on Sphenodesme in Gard.

Bull, Sing. Vol. 21 Part IID.

3. Symphorema polyandrum (polyandra) Wight, Ic. Ind. Or. IL.

1 (1840) 5 t. 363 & ILI. Ind. Bot. II (1850) 217 t. 173/b-—7: Voight,

Cat. Pl. Calc. (1845) 470; Schauer in DC., Prodr. XI (1847)

621; Clarke in Hook. f., Fl. Br. Ind. IV (1885) 599; Briq. in

Engl. & Prantl, Pflanzenf. [V 3a (1897) 180; Cook, Fl. Bomb.

II (1908) 434; Haines, Bot. Bihar and Orissa III (1922) 728;

Gamble, Fl. Madras II, 6 (1924) 1104; Mold., Résumé Geogr.

distr. & Syn. (1959) 164 & 222. — Fig. 4. |

S. involucratum Roxb. sensu Wall. Cat, (1828) No. 1740; Schauer

in DC., Prodr. XI (1847) 621 pro parte non typica.

Allied closely to S$. involucratum, but is distinguished easily by

its leaves being densely tomentose beneath; involucral bracts

obovate, usually entire, sometimes coarsely toothed, densely to-

mentose; calyx lobes 6-8, tomentose on both sides; corolla-lobes

12-16; stamens 12-18, much exserted; ovary slightly glandular at

apex.

Branchlets almost cylindrical, thickly and stellately tomentose

when young, later pubescent. Leaves suborbicular or broadly ovate,

obtuse or sub-acute, almost rounded at base, irregularly crenate or

dentate, densely tomentose on both sides when young, later re-

motely hirtellous and scabrous above, up to 15.5 cm long, 12 cm

broad; main lateral nerves 4-5 pairs; petiole thick, 1—2.5 cm long,

tomentose. Cymes 7-flowered; involucral bracts 6, obovate, usually

entire sometimes coarsely toothed, densely tomentose, accrescent,

up to 4 by 2.5 cm in fruit; peduncles thick, densely tomentose,

2.5-4.5 cm long. Calyx campanulate infundibuliform at anthesis,

somewhat urceolate in fruit, 6-8 lobed, stellately tomentose outside,

Munir — Symphorema 169

glabrous in the tube, accrescent, 1-1.3 cm long; lobes somewhat

triangular, acute, tomentose on both sides, 1.5-3 mm long. Corolla

white, glabrous, usually 12-16 lobed, rarely less, up to 1.8 cm long;

lobes lanceolate, acute, up to 8 mm long; tube almost as long as

the calyx. Stamens 12-18, exserted; filaments filiform; lower parts

of the filaments often connate in pairs; anthers oblong-elliptic.

Ovary obovoid or somewhat sub-globose, glabrous, scarcely glan-

dular at apex; style long exserted, filiform; stigma distinctly bilobed;

lobes cylindrical. Fruit 8-10 mm in diameter, sub-globose, glabrous.

enclosed in the persistent accrescent calyx.

Say

ges

NYG QD }\

OARS

JURAMI DEL.

Fig. 4. Symphorema polyandrum (A-D Wight 2,304; E-G Talbot 362;

H-I Haines 109).

A, Parts of an inflorescence. B, Flower. C, Vertically dissected

flower. D, Stellate hairs from twigs. E, Non-fertile twig with

leaves. F, Fertile branchlet showing its dichotomous divisions. G,

Cyme. H, Fruit enclosed in the accrescent calyx. I, Fruit without

the calyx.

170 Gardens’ Bulletin, Singapore — X XII (1967)

INDIA: Uttar Pradesh, N. Chanda (Dowitt 1: DD; Ramrao 1,370:

DD); Raipura (Marten s.n., DD). Bihar & Orissa, Singbhum

(Haines 109: CAL & K); Chota Nagpur (Gamble 9,149: K):

Mayurbhanj, at Dhobinisole (Econ. Pl. Surv. No. 677: DD).

Madhya Pradesh, [Prob. from Balaghat] (Wight 2,304: K,

prob. holotype; A, CAL & L). Andhra Pradesh, Godavari

(Leg. ? s.n.: CAL; Beddome 6,516: BM; s.n.: SING); Kurnool

hills (Gamble 10, 932: CAL; Beddome 8,181: K; 6,512: BM);

Cuddapah hills (Beddome 40: CAL, 6,513: SING; 6,514 &

6.515: BM; s.n.: CAL & SING). Madras, Pondicherry (Perrottet

410: L & K; 326 & 530: K). Maharashtra, Belgaum near Gokak

(Ritchie 925: E; s.n.: NY). Mysore, N. Kanara (Talbot 362: (E).

CULTIVATED: Hort. Bot. Calc. (Wallich 1,740: BM; Kurz 56

& s.n.: CAL; Leg. ? [Prob. K.d. No. 6,007]: CAL; s.n.: CAL,

E & L).

In 1828 Wallich distributed under his Cat. No. 1,740 as S.

involucratum Roxb. specimens from a plant cultivated in the

Botanic Gardens, Calcutta. Schauer who had with him a specimen

of this collection, used it to give a detailed description under

Roxburgh’s species, but stated that the specimen differs in many

respects from the description of Roxburgh’s type; but failed to

identify with S. polyandrum Wight of which he had seen only

a plate but no specimen. This error was noted by Clarke (1885)

who made the correct reduction.

Munir — Symphorema 17!

Index to Collectors’ Numbers

Collectors’ names are in alphabetical order and their collections are

in numerical order. The number in brackets refers to the number given to

each taxon.

ABADILLA PNH 35,389 (1). KERR 991, 2,932 & 5,234 (2).

AHERN 40, Bur. Sc. No. 1,172, 4,454, KHALIL s.n, (2).

> 4,456 & For. Bur. No. 2,654 KINGDOM-WARD 21.796 (2).

(1). :

AMIHAN PNH 40,320 (1). oe oe (3); 1,040, 2,392 &

AUBERT & GAGE s.n. (2). LACE 2,794 (2)

BAKER 3,042 (1). LARSEN & HANSEN 861 (2).

BANAGA PNH 33,398 (1). LLANOS 69 (1).

BARBER 1,573 (2). LoHER 13,444 & s.n. (1).

BARNES 343 (1).

BEDDOME 39 (2); 40 (3); 6,512 MABANAG PNH 9, 601 (1). .

6,513, 6,514, 6,515, 6,516 & s.n. MADDEN 663 (2).

(3); 6,519, 6,520, 6,521, 6,522, MARTEN s.n. (3),

6,523 & s.n. (2); 8,181 (3). MERRILL 954, 1,334, 1,421, 1,934,

BLATTER 6,112 (2). 2,075 & 2,416 (1); sub. spec.

BORDEN For. Bur. No. 2,549 (1). Blanco No. 467 (1).

BORSSUM 3,283 (1). MEYER For. Bur. No. 2,516 (1).

BOURDILLON 537 (2). MILL s.n. (3).

BRANDIS 881 (2). PARKINSON 15,733 & 15,750 (2).

CARTER 1,507 (2). PascuaL For. Bur, No. 28,766 (1).

CLEGHORN s.n. (2.) PERROTTET 326 & 410 (3); 487 (2);

CLEMENS Bur. Sc. No. 17,534 & 530 (3).

18,176 (1).

RaMos Bur. Sc. No. 27,122 (1).

RAMRAO 1,370 (3).

RITCHIE 925 & s.n. (3).

ROBERTSON 316 (2).

COLLET 435 (2).

CUMING 648 & s.n. (1).

DALZELL s.n. (2),

DowirTr 1 (3). RoGERS 274 & 910 (2).

DUTHIE 9,687 (2). ROXBURGH s.n. (2).

EBRON PNH 34,236 (1). SINCLAIR 9,470 (1).

Econ. Pl. Surv. No. 677 (3). SMITINAND 4,169 (2).

EDANO PNH 17,748, 17,797 & STEINER 1,011 & 1,742 (1).

stacy og 17,419, 17,429 & or ag ee) aie

17,467 (1). ine SuLIT PNH 8,320 (1).

SuSARA PNH 37,313 (1).

TALBor 52 & s.n, (2); 362 (3).

TAMESIS For. Bur. No. 11,924 (1).

THWAITES C.P. No. 3,645 (2).

GAMBLE 9,149 & 10,932 (3); 13,745 Toppinc Bur. Sc. 5,228 (1).

FELIX For Bur. No. 3,914 (1).

FERNANDES 220 (2),

FISCHER & GAGE 83 (2).

(2).

GLENIE sub Thwaites C.P. No. VIDAL 501, & 848 (1).

3,645 (2). VILLAVICENCIO For, Bur. No

GreEGorY 81 (1). 23,654 (1).

WALLICH 1,740 (3).

Gaeecubnie, Go Mexetal wacten e546, (1).

Hotman 53 (1). a el},

WIGHT 909 & 2,303 (2); 2,304 (3);

LLAGAN sub Phil. Nat. Herb. No. 2,586 & 2,587 (2).

35,488 (1). WILLIAMS 707 (1).

Malayan Fern Notes. V

BETTY MOLESWORTH ALLEN*

Dryopteris hirtipes (Bl.) O. Kuntze, Rev. Gen. Pl. 2; 813. 1891.

Basinyum: Aspidium hirtipes Blume, En. Pl. Jav. 148. 1828.

Synonyms: Lastrea hirtipes (Bl.) Moore in Beddome, Ferns of

S. India, tab. xcvi. 1863-64.

Nephrodium hirtipes (Bl.) Hooker in Hk. & Bak. Syn. Fil.

261: 1883.

Malaya. Slopes of Gunong Palas, Cameron Highlands, Pahang.

23. 4. 63, ca. 1870 m. Coll. no. 4946.

This is a widespread species found from south China through

Burma and north India to Ceylon, Borneo and Indonesia, and to

some Polynesian islands. It does not appear to have been recorded

in Malaya previously, but occurs in Siam where it bears a local

name, Ta-ku-khu’i-do.

Description of living plants: Rootstock stout but fiat, with tufted

fronds. Scales to about 20 mm. long by 7 mm. wide at the base,

tapered, midbrown with a dark base; margins entire. Scales become

sparse on the upper part of the stipe, and also on the rachis where

some are dark, almost black. Stipe approximately 1/3—4 the length

of the lamina; base black for about 3 cm. then green (stramineous

when dry), the base is usually densely scaly for about 5 cm. Lamina

simply pinnate and to about 54 x 30 cm. or more, widest at the

basal or penultimate pair of pinnae. Rachis green, grooved above

and usually rounded below. About 12—15 subopposite to alternate

pairs of pinnae, the basal ones on stalks 1—-2.5 mm. long, gradually

becoming sessile about the middle pinnae and adnate or slightly

decurrent in the upper ones; apex of lamina lobed. Pinnae

commonly 14 x 2.5 to 30 x 3.5 cm., ascending and widest just

above the base; shallowly lobed for 1—1.5 mm.; lobes falcate

especially when dry, entire with a blunt tip. Acroscopic basal lobe

overlapping the rachis, but not the basioscopic one. Colour deep

green and shiny above, but pale below, texture firm, thick and

brittle, and pinnae glabrous except on the veins. Veins obscure,

becoming visible when dry, ending just short of the margins; 4—5

veinlets in each group, sometimes 6 on the basioscopic side; very

narrow hairlike brown scales scattered along the veins. Sori when

immature, pale cream in colour with a whitish indusium which is

persistent when mature or dry. Sori vary from one either side of

the costule, to several uneven rows which are always nearer the

costae than to the pinnae margins. ,Indusia reniform, maturing

medium-brown in colour.

*Authors present address:

c/o Bardays Bank D.F.O.,

Gibraltar.

174 Gardens’ Bulletin, Singapore — XXII (1967)

The first plants found were growing on steeply sloping ground

amongst tea bushes (Camellia sinensis Kze.) on the Sungei Palas

Tea Estate. These had very small infertile fronds not more than

15 cm. long, but had very large thick rootstocks, and were pro-

bably old plants which were frequently cut back by the weeding

carried out around the tea bushes, so they produced no larger

fronds. The edge of the estate was a few metres from here, and

was bordered by a steep and narrow forest-clad gully. In the

secondary growth bordering the forest I found five more plants of

thesDryopteris growing where the soil was loose (see plate 1).

They were larger and were partly in shade, and their roots were

protected by other growth. Only one fertile frond was found and

this was immature. Associated plants noted besides grasses and

small herbs were. Thelypteris uliginosa (Kze.) Ching, T. brunnea

(Wall.) Ching, T. beddomei (Bak.) Ching (which was very

common), Athyrium japonicum (Thbg.) Copel., Cyclosorus ecallosis

Holtt., tall Cyathea contaminans (Hk.) Copel., and the introduced

Passiflora edulis Sims, which was scrambling all over plants at

the forest edge. Just inside the forest in dark wet places, were seven

more plants of D. hirtipes; these were smaller but were fertile, the

sori being mature. No more were found in the remainder of this

narrow strip of forest. On all the fertile fronds the indusia showed

clearly, even when past maturity.

D. hirtipes is the only simply pinnate Dryopteris so far found in

Malaya, the other three having much more deeply divided fronds.

Through the courtesy of the Director of the Botanic Gardens in

Singapore I was able to compare the Cameron Highland material

with speciments from other countries, and they were almost identi-

cal with some from Mt. Kinabalu, and very close to a sheet

labelled D. atrata (Wall.) Ching, from the Khasi Hills in India,

except that on my specimens the basal scales are larger and paler.

In Beddome’s description (1892) the stipe and rachis are densely

scaly.

Dryopteris hirtipes (Blume) O. Ktze. var. exinvolucrata (C. B.

Clarke) B. Allen comb. nov.

Synonym: Nephrodium hirtipes (Bl.) Hooker var. exinvolucrata

C. B. Clarke, Journ. Linn. Soc. Bot. xxv: 93. 1889.

Malaya: In clearing in forest at filter tanks, Cameron Highlands,

Pahang. 1-5-1963, ca. 1,700 m. Coll. no. 4980.

Differs from the species in having the basal scales very dark

brown, almost black, and narrower (less than 3 mm. wide at the

base), and being quite black at the base. Pinnae are slightly smaller

and narrower the widest measured being 2 cm. wide by 10 cm.

long, and the fertile fronds are slightly contracted. Lobing is very

shallow and the apex of the lobes is frequently toothed. Fronds

are about the same length so that the pinnae are wider apart and

the acroscopic lobe does not overlap the rachis. Sori are naked,

there being no sign of indusia at any stage.

Allen — Malayan Fern Notes 175

This variety was collected near a stream in a small open space

surrounded by tall forest, and just above where the water filter

tanks are (previously the trout hatcheries). This is about two miles

from where D. hirtipes was found. The ground was flat and rocky

and had been cleared of forest before the war and periodically cut

since, but a few Cyathea contaminans grew on the stream edges

and gave slight shade. Here several plants of the variety were

growing amongst plants of similar height and about 1-2 metres

from the stream bank. Two other ferns were growing with them;

they were small specimens of Athyrium amplissimum (Bak.) Holltt.,

and a Cyclosorus, possibly C. megaphyllus (Mett.) Ching. The

Dryopteris bore fertile fronds at all stages of maturity, and the

indusia was clearly missing.

That the species and the variety should turn up together is in-

teresting. Also that the two localities, although not far apart and

both in disturbed areas, are quite different from each other. They

are large enough ferns not to be easily overlooked, and surround-

ing areas and similar places were searched but no more of either

was found, so appear to be very restricted in their distribution.

This suggests to me that they may be comparatively new arrivals

in Cameron Highlands as is Thelypteris beddomei. (This fern was

first recorded for Malaya in 1949 when I collected it from

Maxwell’s Hill in Perak, where tea had been grown about 50 years

ago. The only other Malayan locality I know of, is at Camerons

where tea is commonly grown today.) Dryopteris hirtipes was not

growing near water, but on sloping ground both in the open and in

the shade. The variety, on the other hand was near a stream on

flat ground which well may be flooded, but was not found in real

shade. |

I saw neither in Camerons when I was there in 1948, although

I collected ferns from the exact area of the variety. Nor did Dr.

Holttum record them during his collecting, prior to 1949 (after that

it was virtually closed for 11 years). Christensen and Holttum

(1934) reported both kinds from Mt. Kinabalu area and stated

(about the Lobang specimens) ‘‘H. 25557 has conspicuous indusia,

but no indusia can be found on the Tenompok specimens, which

thus agree with the Himalayan-Chinese Phegopteris scottii Bedd.”’

I believe that Dryopteris scottii (Bedd.) Ching (1933a), syn.

Phegopteris scottii is identical with var. exinvolucrata, and thus

would become a synonym of it. The illustration in Beddome (1892)

shows almost elliptic pinnae, but he states in the text that it is

“perhaps an abnormal form of Lastrea cuspidata or hirtipes’’. Dr.

Holttum writes (in a letter to me) that the differences between D.

hirtipes and D. scottii as given by Ching, agree with my descrip-

tion of var. exinvolucrata. He points out however, that after com-

paring many specimens of D. hirtipes and the variety, he finds that

the toothing on the edges of the pinna-lobes is variable and does

not seem to be a good distinctive character.

176 Gardens’ Bulletin, Singapore — X XII (1967)

Dryopteris atrata is a species very closely related to D. hirtipes

and in fact was cited as a synonym by van Alderwerelt van

Rosenburgh (1908) and Backer and Posthumus (1939) but con-

sidered a variety by C. Christensen (1931) and retained as a distinct

species by Ching (1938). He states that the true D. Airtipes occurs

in south India and Siam, but not in north east India and S. China.

_Microsorium superficiale (Bl.) Ching, Bull. Fan Mem. Inst. Biol.

4:299. 1933.

Basinym: Polypodium superficiale Blume Flor. Jav., 11; 136.

1828-29.

Synonyms: P. normale Don, var. sumatranum Bk. Journ. Bot.

XVIII, 215.

P. nigricans v.A.v.R. Bull. Btz., Vii: 24. 1912.

Pleopeltis superficialis (Bl.) Bedd. Handb. 350. 1885.

PI. nigricans (v.A.v.R.) v.A.v.R. Bull. Btz. XX:22. 1915.

Pl. rupestris var. nigricans v.A.v.R. VIL: 24. 1912.

Distribution: From Japan and India to Indonesia.

Malaya: Slopes of Gunong Batu Brinchang, Cameron Highlands

district; Pahang. altitude, 1960 m. upwards for at least 200 m.

Both epiphytic and terrestial. Coll. nos. 4024 and 4685. There

appear to be no previous Malayan records.

Description. Rhizome long-creeping, dark brown, about 4 mm.

wide when dry. Rhizome scales conspicuously iridescent, cell walls

dark brown and forming a clear network, edges irregularly toothed,

Scales spreading, overlapping, gradually tapered from a wide base

to a pointed apex; apices often breaking off on old scales. Fronds

well spaced on rhizome and stipes from 4 to 10 cm long or more,

glabrous when mature, usually paler than the lamina. Lamina

varying greatly in size according to habitat; mature fronds from

15-30 cm. long or more, by 3—5 cm. wide with tapered base and

apex, usually widest just below the middle; edges slightly thickened,

entire, often sinuate when dry. Fronds deep green and shiny above

and paler below and glabrous. Veins indistinct but visible by trans-

mitted light, especially the more or less parallel side veins. Sori

numerous, in 3-5 irregular rows between the midrib and margin

(or in 1-2 irregular to fairly regular rows between the slightly

parallel veins. Sori round, but varying in size from 1.5—2.5 cm.

in diameter.

In general appearance this fern resembles M. normale (Don)

Ching, but does not have the curious scales of this species, but

copious shiny scales which are most conspicuous. M. superficiale

is in fact closer to M. sarawakense (Bak.) Ching, and at a distance

may look like a large form of it.

I have found M. superficiale only in one (large) area, on the

slopes of Batu Brinchang, in both disturbed and more or less

intact rain forest. It seems to favour the outskirts of tall wet

jungle, on both living and dead tree trunks, on horizontal logs,

and on one occasion on the trunk of a Cyathea sp. Where it is

Allen — Malayan Fern Notes 177

common, it seems to have become established on the ground

around the host trees, where it produces large fronds; these I

have never found fertile. In no cases were the ferns growing

very near streams.

There was tall second growth, with a small rotan, probably

Calamus sp., rather effectively sealing this area from the road.

The ground was flat and damp with patches of Athyrium dilatatum

near the terrestial M. superficiale which was very common here.

It was also epiphytic on the trunks of small trees and on the

remains of trunks of larger cut trees, and the long thin rhizomes

frequently reached the top of a stump, twisting itself around,

making a conspicuous bunch of rhizomes and fronds, which hung

away from the trunk.

Other ferns noted in this area included Cyathea contaminans

(on the outside edge), C. latebrosa, Cyclosorus ecallosis, C. stipel-

latus, Bolbitis simplicifolia, and occasional Athyrium subintegrum;

these were terrestial. Epiphytic ferns were Asplenium tenerum, A.

nidus, Elaphoglossum callifolium, Grammitis hirtella var. imajor,

G. reinwardtii and one G. holttumii (very low on a branch of a

sapling). Xiphopteris hieronymusii and Ctenopteris obliquata otten

grew with the Grammitis spp. on the mossy trunks.

Other places where the fern grew near the above habitat were

on trees in tall forest on sloping ground and some others were

seen in full sun on stumps; the fronds of the latter were small

and pale green. It was so common here on the eastern slope of the

mountain that I feel sure it must grow in similar places on other

parts of the Main Range, and may even have been collected pre-

viously for it is stated to occur in Malaya by both Beddome end

van Alderwerelt van Rosenburgh although no herbarium material

appears to exist.

Specimens of Microsorium superficiale have been distributed to

the following institutions: Royal Botanic Gardens, Kew; Smith-

sonian Institution; Chicago Natural History Museum; Gray

Herbarium; Singapore Botanic Gardens; Sarawak Museum, and

Swedish Museum (Nat. Hist.).

Tectaria decurrens (Pres!) Copeland, Phil, Journ. Sci. 2C: 412 1907.

Synonyms: Aspidium decurrens Presl; A. mamillosum C. Chr.:;

A. pteropus Kze.;

Nephrodium decurrens (Presl) Bak.; Sagena decurrens (Presl)

Houlst.;

S. mamillosa (C. Chr.) Moore; S. pteropus (Kze.) Beddcme.

van Alderwerelt van Rosenburgh calls the Malayan form var.

mamillosum as this seems to have the sori deeply impressed, but

it is doubtful whether it is really sufficiently distinct to maintain

the variety. ?

Known collections in Malaya: Batu Kurau, Perak. 1911; coll.

G.C. Matthew. Foothills of the Main Range behind Gopeng.

1960; coll. B. Molesworth Allen nos. 4486, 4555.

178 Gardens’ Bulletin, Singapore — X XII (1967)

Distribution: widespread from south China, N. India to

Polynesia.

Holttum (1954) does not include Yectaria decurrens as a

Malayan fern for the existence of Matthew’s specimen was not

then known. Admittedly Ridley (1921) does record it from Pahang,

Trengganu and Perak, but it appeared there was no material in

herbaria from these States. As I will show, it is a fern that is

easily mistaken for two other Malayan Tectaria which grow in

similar places. So, Dr. Holttum who was fully aware of this,

being unable to find any collection from Malaya rightly omitted

it from the Flora. I think now that Ridley did collect it himself

from near Bujang Melaka mountain in Perak, and I believe that

T. decurrens was far more common before the lowland jungle

was greatly milled or destroyed for cultivation, for it prefers tall

wet jungle, usually near streams.

It does seem strange that no other herbarium material trom

Malaya is known; Matthew’s collection only came to light recently

when Dr. Holttum was kindly examining Tectaria for me at Kew.

Description of the Malayan plants: Stock stout, built-up sith

supporting rootlets; scales dull, medium-brown, stiff, entire and

tapered, commonly 102 mm. Stipes medium-brown to 50 or 60

cm long, commonly about 45 cm. long, and winged to within 3-4

cm. of the base, then ceasing abruptly. Wing fairly even but usually

widest near the lamina, where it may be over 3 cm. (across both

wings) on sterile fronds. Scales are present only on the base of the

stipe, the underside being globrous for the remainder (it is always

scaly in T. vasta). Fronds erect, glabrous medium-green and shiny

above, drying olive brown. Lamina, simple when very young;

adult deeply lobed with 3-4 pairs of lobes and a basal pair

which often have a lobe each. Apex large, usually with a pai of

small lobes at the base. Lamina length commonly about 45—55

cm. long and about as wide (across the lobes) as long. Fertile

fronds slightly contracted; the upper 16 cm. of the wing of tlie

stipe bearing scattered sori. Sori similar to that of T. crenata on a

receptacle, very deeply impressed so that the upper surface of the

lamina is strongly papilliform. Sori placed on free terminal veins

within aeroles in two, more or less even rows, between the costules.

Indusium large, reniform and persistent.

When I first saw T. decurrens in Malaya (and I was familar

with it in Borneo) in July 1959, it was sterile, and I noted that

it was possibly young T. vasta. Later, however, in August I found

a fertile plant in the same area and not expecting T. decurrens,

[ was careless enough to pass it off as T.crenata, but did not

collect it then, only adding that the stipe was winged to its base

as in T. vasta. I mention this to illustrate how easy it is to

mistake this fern, and if records are not backed by herbarium

material, they should be treated with caution.

Allen — Malayan Fern Notes 179

-Tectaria decurrens was growing on the floor of tall mixed

jungle, which lies in the foothills of the Main Range between

Gopeng and Jor Camp. The forest here has remained more or

less intact as it is a water catchment reserve. Next to a large

Pteridys sp., it was the commonest fern within an area of about

30 square metres, at an altitude of about 100 m., and higher up

in the continuation of the forest, it became rare and was not seen

above 300 m. The ground where it was common was generally

sloping, but with a flat area just above a rocky stream. The soil

was black and sticky, the undergrowth fairly sparse, and in a

fewplaces some trees had fallen and exposed a few Tectaria to

direct sun. These seemed to be surviving when I last saw them

(1963), but were continually fertile whereas the others in shaded

places, in what is probably their natural habitat, produced new

growth in December and January, and sori from January to April.

During August — September only sterile plants, or those with

very old sori were noted.

Associated floor plants included Tectaria maingayi which was

very common, a very large form of Pteridys which approached

acutissima, as well as normal P. australis (which was not very

common here), Tacca cristata and several small gingers, Globba

auranticea and G. variabilis, and large ones, Achasma sp. and

Hornstedtia sp.; Forrestia sp., Curculigo latifolia, Angiopteris

evecta, and some dwart palms including Pinanga (probably

disticha), and some aroids Anadendrum sp. and Schismatoglottis

sp. in the dryer parts, also Tectaria singaporeana on sloping

ground. Juvenile Lomariopsis cochinchinensis was frequent at tree

bases, and taller growth in this area included Cyathea latebrosa,

Baccaurea lanceolata and B. brevipes. Growing in and on the

sides of the stream were noted, Saraca thaipingensis, Salmalia

valetonii and a huge Annonaceous tree in the stream with large

cauliferous flowers borne low on the trunk, and Laportea stimulans;

on wet rocks were Bolbitis diversifolia which was very common.

B. heteroclita, Egenolfia appendiculata and a possible hybrid

between two of these; Microsorium pteropus, Abacopteris meniscii-

carpa, Trichomanes maximum and one Athyrium simplicivenium

(which commonly occured higher up), Stauranthera grandiflora,

and a Begonia sp. which was very common. On large rocks by

the stream were Polypodium papillosum and Phymatodes nigres-

cens. Lianes were common over the stream, one was a Vitis sp.

others were Calamus spp.

The only other place I have seen the fern in Malaya was at

Klian Intan in Upper Perak. It was growing on the floor of tall

rather similar forest at about 300 m. There was scant undergrowth

on flat ground near a stream. Again, the Tectaria was found in

only one small area, but as I did not collect it then (intending

to return later), I can only make a note of it in here.

180 Gardens’ Bulletin, Singapore — XXII (1967)

Cyclosorus papilio (Hope) Ching, Bull. Fan Mem. Inst. Biol.

8:214. 1938 syn. Nephrodium papilio Hope, Journ. Bombay

Nat. Hist. Soc. 12: 625 t. 12.1899.

Distribution: N. India, Ceylon.

In 1958 I found C. papilio on the borders of Pahang and Perak,

by the side of a small stream under tall trees, growing alongside

some large gingers (Achasma macrocheilos). This was just below

Ringlet at about 1,000 m. It was identified by Dr. Holttum. Later

I found it in several areas lower down, on the same part of the

Main Range, where in places it is quite common. The second

area was all within the Tapah Hills Forest Reserve, from about

670-1,000 m. in altitude. Coll. nos. 3913, 3986, 4312, 4449, 4949.

In the past C. papilio was included under Nephrodium molle

var. major Bedd. (as was C. sumatranus (v.A.v.R.) Ching). Then

the specimens from north India and Ceylon were removed by Hope

and called N. papilio (see Holttum (1954) p. 275).

Extreme forms of the Malayan species do appear to be quite

distinct, especially in the remarkable auricles on the stipes (always

about 5 or 6 pairs) which have such enlarged and pointed acros-

copic lobes that they resemble the Malayan Dragontail butterfly

(Lamproptera megas virescens), thus the specific name is most

apt. There are oblong glands on the underside of the lamina,

sometimes copious, which I have not seen on C. sumatranus.

There are, however, specimens which I consider are intermediate

between the latter and C. papilio. I have found C. sumatranus

growing together with C. papilio at about 700 m. which is a high

altitude for this normally lowland fern.

I have not yet seen any other specimens of C. papilio with which

to compare mine. If this species has already been collected in

Malaya, it will most probably be found under C. sumatranus, for

there appear to be no other records of it.

I am grateful to the Director of the Royal Botanic Gardens,

Kew, for assistance in checking specimens and to Dr. R. E.

Holttum who has spent much of his valuable time on my behalf

checking my identifications, and especially for the information

on the status of Dryopteris hirtipes and the var. exinvolucrata.

Literature Cited

BACKER, C. A. & Posthumus, O., 1939: Varenflora voor Java.

BEDDOME, R. H. 1892: Handbook to the Ferns of British India.

CHING, ., 1933: Bull. Dept. Biol. Sunyatsen Univ. no. 6. 3.

CHING, .. 1938: Bull. Fan Memorial Institute, Botany vol. 8.

CHRISTENSEN, C., 1931: Contributions to the U.S. National Herbarium, 26.

CHRISTENSEN, C. & HoLtrum, R. E., 1934: Gardens Bull. Straits Settle-

menis,.J ...pt,: 3.

Ho.ttum, R. E. 1954: Flora of Malaya. Vol. 2. Ferns of Malaya.

RIpLey, H. N., 1926: Malayan Branch, Royal Asiatic Soc. 4, pt. 1, p. 79.

VAN ALDERWERELT VAN ROSENBURGH, C.R.W.K., 1908: Malayan Ferns

Handbook.

Allen — Malayan Fern Notes 181]

1. Dryopteris hirtipes on forest edge, Cameron Highlands. Pahang.

Gardens’ Bulletin, Singapore — X XII (1967)

182

SSS

RGC—LQ@0AEEGUR

nchang. Pahang.

;

le from the slopes of Batu Br

1Cid

icrosorium superf

Allen — Malayan Fern Notes 183

3. Tectaria decurrens growing in forest near Gopeng. Perak.

184 Gardens’ Bulletin, Singapore — X XII (1967)

4. Tectaria decurrens; a fertile frond showing the deeply immersed sori.

oes

Allen — Malayan Fern Notes

}

) SI

yed

olidpd s

HNAOSO]

Dy)

Notes on the Flagellata of Hongkong.

B. V. SKVORTZOV

(Sao Paulo, Brazil)

1. Species of the genus Chlorogonium Ehr. (Chlorophyceae,

Volvocales, Chlamydomonadaceae) from polluted waters in

Hong Kong.

Notulae ad floram Flagellatarum Hongkongensis 1. Species generi

Chlorogonium Ehr., Chlorophyceae, Volvocales, Chlamydo-

monadaceaae in Hongkong aquae impurae inventae cum 8 fig.

Abstract

In the present note the author describes 7 species of the genus

Chlorogonium Ehr. from polluted waters of Hong Kong, recorded from

materials received in January 1966 and studied in the Botanical Institute

of Sao Paulo, Brazil. The collection from Hong Kong has been received

through the kindness of Professor Thomas B. Widdowson of the Botanical

Department of Hong Kong University. Samples of mud cultured in

glass vessels were collected in Hong Kong by Mr. S. T. Chan of the

Botanical Department of the University.

Preface.

Green flagellata of the genus Chlorogonium Ehr. form a typical

algal component of polluted waters. The genus was proposed by

Ehrenberg in 1830 to cover such forms. Cells of Chlorogonium

are almost fusiform in shape and have 2 flagella at the anterior

part of the cell. The chloroplast is entire, massive, green and not

Jaminate in the specimens from Hong Kong, According to the

species, the chloroplast contains one, two, or several pyrenoids

or may lack them entirely. The nucleus is almost centrally located.

Some species have an eyespot, while others lack one. Asexual

reproduction is by transverse division of the protoplast and by the

production of zoospores. Sexual reporduction is by fusion in pairs

of equal sized (isogamous) biflagellate gametes. |

Seven species of Chlorogonium are recorded from the cultures

of polluted mud collected near Hong Kong. One has been identified

as Chlorogonium euchlorum Ehr., but the remaining appear to be

new to science and are so designated.

Key to the Species.

A. A single pyrenoid present.

a. Eyespot present, cell fusiform |... 1. Chlorogonium chanii

Sp. nov.

b. Eyespot lacking, cell fusiform

1. Pyrenoid central. 2. Chlorogonium astigmatae

sp. nov.

2. Pyrenoid posterior (rarely 2) 3. Chlorogonium subtropicale

sp. nov.

188 Gardens’ Bulletin, Singapore — X XII (1967)

B. Two pyrenoids present.

a. Eyespot lacking ... 4. Chlorogonium widdow-

soni sp. nov.

b. Eyespot present, apical and very

small it

5. Chlorogonium aberdeenii

sp. nov.

C. Numerous pyrenoids.

a. Eyespot lacking 6. Chlorogonium stentoni

Sp. nov.

b. Eyespot present ... 7. Chlorogonium euchlorum

Ehr.

1. Chlorogonium chanii sp. nov. [Fig. I: 1.]

Cellula brevi vel longi-fusiformis cum apicibus attenuatis et

cum flagella 4-plo cellulae paulo longiore; chromatophoris

parietalis; stigma elongatum prope vacuolae parte anteriore cel-

lulae; pyrenoide centralis prope nucleus; cellulae 9-15 x 4».

Differt a Chlorogonium fusiforme Matv. in stigmate fere apicale.

Dedicavi hance species in honourem Do. S. T. Chan collectoris

algarum Hongkongensis. Hab. Hong Kong, in orysetis cum aqua

impura, Ig. S. T. Chan, 12th August, 1965.

2. Chlorogonium astigmatae sp. nov. [Fig. I: 2.]

Cellula brevi-fusiformis cum apicibus brevi acutis, lateribus.

plusminusve rotundatis; flagella 22—plo cellulae longioribus; stig-.

mate nullo; pyrenoide et nucleus centrales; vacuola parte anteriore

prope flagellis; cellula 9-14 x 4-5 yw. Differt a Chlorogonium

obliquum Skv. 1946, Ch. uinutum Sky. 1957, Ch. leiostractum

Str. et Ch. tetragamum Bohi in stigmate nullun. Hab. Hong Kong,

in rivularis et orysetis cum aquae impurae, lg. S. T. Chan, 12th

August, 1965.

3. Chlorogonium subtropicale sp. nov. [Fig. I: 3 & 4.]

Cellulae fusiformis cum apicibus attenuatis et acutis; flagella 2.

apice, 4%-plo cellulae longioribus; vacuola_ contractilis pone;

nucleus centralis; pyrenoide 1 rarior 2, inter nucleus et parte

posteriore cellulae; cellulae 18-30 x 7.4 u. Differt per totem species

in 1-2 pyrenoidibus. Hab. Hong Kong, in rivularis montanis cum

aqua impura, Ig. S. T. Chan, 12th August, 1965.

4. Chiorogonium widdowsonii sp. nov. [Fig. I: 5.]

Cellula perfecte lanceolata vel spatulata cum parte posteriore

late cuneata; flagella 2%-plo cellulae longiora; nucleus centralis;

stigmate nullo; pyrenoide 2; cellulae 14-18 x 7.4 yw. Differt a

Chlorogonium aculeatum (Korsch.) Pascher, Ch. acuminatum Sky.

1957, Ch. elongatum Dang., in stigmate nullo. Dedicavi hanc species

in honourem Dom. Prof. Thomas B. Widdowson, Botan. Depart.

Hong Kong University. Hab. Hong Kong, in rivularis montalis curn

aquae impurae, lg. S. T. Chan, 12th August, 1965.

5. Chlorogonium aberdeenii sp. nov. [Fig I: 6.]

Cellula fusiformis cum apicibus attenuatis et acutis, parte

mediane cellulae cum lateribus conversis; flagella 2, 4-plo cellulae

paulo brevioribus; stigma minor fere apice; nucleus centralis;

vaculoles contractiles nonnihil; cauda hyalina sine chromato-

phoris. Differt a Chlorogonium elongatum Dang. in stigmate mintae

Skvortzov — Hong Kong Flagellata 189

apicale et in margines cellulae protractis. Hab. Hong Kong, prope

Aberdeen, in rivularis montanis cum aqua impura, lg. S. T. Chan,

12th August, 1965.

6. Chlorogonium stentoni sp. nov. [Fig. I: 7.]

Cellulae elongato-fusiformis cum apicibus elongatis et protractis;

flagella 4~-5-plo cellulae brevior: stigma nullo; nucleus fere

centralis; pyrenoide 5-10; cellulae 37-74 x 9-5 u. Differt a

Chlorogonium euchlorum Ehr. (see Pascher 1927, p. 314-315, fig.

281-282) in cellulis astigmatis. Dedico hanc species in memoriam

Dom Prof. O. H. Stentoni Universitatis Hongkongeusis Hab.

Hong Kong, Aberdeen, in rivularis montains cum aquae impurae,

Ig. S. T. Chan, 12th August, 1965.

7. Chlorogonium euchlorum Ehr. [Fig. I: 8.]

Cells lancelate and acute at both ends. Flagella 2, about 4

the length of the cell. Chromatophores in the form of a green

plate close to the periplast. Stigma large, long, in the anterior part

of the cell. Pyrenoids 6. Nucleus central and large. Contractile

vacuoles small and numerous. Hab. Hong Kong, in a polluted

stream, collected by S. T. Chan, 19th August, 1965. Reported

from Europe and N. America.

Acknowledgments.

I wish to express my gratitude to the Foundation of Science

(Fundaso de ampera a Pesquisa, Sao Paulo) which is supporting

my research work, to the Director of the Botanical Institute of Sao

Paulo, Dr. Alcides Ribeiro Teixeira and the leaders of the Crypto-

gamic Section of the Botanical Institute, Dr. Oswaldo Fidalgo,

Dr. Joao Salvador Furtado, Dr. Carlos Eduardo de Mattos

Bicudo and the secretaries of the section Aparecida de Castle

Miccoli and Mallene Ocana Orlane for help in the work. I am

indebted to Prof. Thomas B. Widdowson and Mr. S. T. Chan of

the Department of Botany, Hong Kong University, for collecting

and sending to Sao Paulo, Brazil the algological collection for

my studies. I wish to express my gratitude to the Director, Botanic

Gardens, Singapore, for permission to publish these notes in the

Gardens’ Bulletin, Singapore.

Literature.

Dedusenko-Stzegoleva, N. T., A. M. Matvienko & L. A. Schkor-

vatov. 1959. Chlorophyta. Volvocineae. Key of freshwater algae

of U.S.S.R. Vol. 8. Moscow 1959.

Pascher A. 1927. Volvocales—Die Siisswasserflora Deutschland

Oesterreich und der Schweiz.

Skvortzov, B. 1946. Species novae et minus cognitae Algarum,

Flagellatarum, et Phycomycetarum Asiae, Africae, Americae et

Japonae nec non Ceylon anno 1931-1945 descripto et illustrato

per tab. 1-18. |

Proc. Harbin Soc. Nat. Hist. and Ethnogr. 2. Botany, Harbin

1946.

1957. New and rare flagellatae from Manchuria, Eastern Asia.

Philipp. J. Sc. 86. Manila.

oS)

eo” ee i

Figure I

Chlorogonium

chanii sp. nov.

astigmatae sp. nov.

subtropica sp. nov.

widdowsonii sp. nov.

aberdeenii sp. nov.

stentoni sp. nov.

euchlorum Ehr.

Sclereids in Fagraea

( Loganiaceae )

HARDIAL SINGH

Botanic Gardens, Singapore

INTRODUCTION

Caspary (1865) was one of the early workers to realize the

taxonomic value of sclereids in distinguishing species within the

genus. Van Tiegham (1891) used this character when he observed

sclereids in the mesophyll of a number of species of Mouriria and

Memecylon which led him to group both the genera in the sub-tribe

‘‘Mouririées” under the subfamily ‘“‘Melastomacées”. Since Van

Tiegham in his work included neither the author-names of the

species of Mouriria studied nor citations to specific herbarium

specimens, it led Foster (1946) to do a complete re-examination of

the foliar sclereids of Mouriria, utilising an extensive series of

herbarium collections and he saw that throughout the sixty-nine

species investigated, the sclereids were restricted in position to the

ends of the veinlets, regardless of their form. He thus considered

the presence of terminal foliar sclereids as an important generic

character of Mouriria which could be utilized in the identification

of ‘“‘sterile’ and doubtful materials. Since then, many workers

Morley (1953), Rao (1957), Barua and Wight (1958), Tomlinson

(1959) and Carlquist (1961) have used sclereid characters to assess

the taxonomic positions of certain angiosperms.

In the family Loganiaceae, Metcalfe and Chalk (1950) have

recorded the presence of ‘sclerenchymatous idioblasts’ in the

mesophyll of Anthocleista, Fagraea, Potalia and Strychnos. More

recent work by Rao (1965) on the genus Fagraea showed that

the size and arrangement of the sclereids was different in each of

three species examined. Preliminary studies on foliar sclereids in

Fagraea revealed great morphological variation in the structure

of the sclereid and it was this that led the writer to the present

investigation of all the species of Fagraea with a view to study

the anatomy of sclereids and to ascertain their systematic value in

this genus.

Materials and Methods:—

The genus Fagraea though distinctly centered in Malaysia is

found distributed from Ceylon and the Malabar coast through SE

continental Asia to South China, Hainan and the southern penin-

sula of Formosa, in the Northern Territory and NE Queensland,

‘

194 Gardens’ Bulletin, Singapore — X XII (1967)

and in the Pacific from the Marianas to the Marquesas and the

Tubuai Islands and New Caledonia in the southwest. (Plate 1).

The materials for the present investigation were derived partly

from fresh and partly from herbarium specimens. Fresh material

was available for four of the species. Of these, F. auriculata, F.

fragrans and F. ridleyi are growing in the Botanic Gardens, while

material of I’. crenulata was collected from trees growing at Farrer

Road and in the campus of the University of Singapore. Material

for the rest of the species was taken from herbarium sheets at the

Singapore Botanic Gardens Herbarium and material for ten of the

species, was very kindly sent from the herbarium at Leiden by Dr.

P. W. Leenhouts (see below for details).

The following specimens were used in the study: —

1. F. acuminatissima Merr., Ngadiman SFN. 36671 at

Pengkalan Raja, Pontian, Johore, Malaya. June 1939.

SING.

2. F. annulata Hiern., Versteeg G. 1237. Between Zandroort

and Sabang, W. New Guinea. June 1907. LEIDEN.

3. F. auriculata Jack, Fresh specimens, Singapore Botanic

Gardens.*

4. F. beteriana Benth., Barrett J.H. N.G.F. 4231 at Rouna

Gorge, Sogeri Central District Papua. August 195!

SING.

5. F. blumii G. Don, Wray L, 1992. SING.

6. F. bodenii Wernh., Carr C.E. 14132 at Alola, Papua.

January 1936. SING.

7. F. calcarea Henders, Henderson M.R. 25036 at Bukit

Chintamani, Pahang, Malaya. October 1931, SING.

8. F. carnosa Jack, Sinclair J. 7948 at Belara Forest Reserve,

Trengganu, Malaya. July 1953. SING.

9. F. carstensensis Wernh., van Royen P. et Sleumer H. 7755,

Tamran Mts., W. New Guinea, November 1961.

LEIDEN.

10. F. ceilenica Thunb., Burkill H.M. 2421 at Fraser’s Hill,

Pahang, Malaya. August 1960. SING.

11. F. crenulata Clarke, Fresh specimens. Singapore. *

12. F. curtisii K & G, Curtis 1676 at Tanjong Tirie Kedah,

Malaya. September 1890. SING.

13. F. elliptica Roxb., Henderson M.R. 19658 at Sungei Keteh,

Kelantan, Malaya. October 1927. SING.

Hardial Singh — Sclereids in Fagraea 195 ’

14.

1S.

16.

US.

18.

19.

20.

at.

22;

y 2%

24.

25s

26.

p45 %

28.

29.

30.

bl,

eymae Baker, Versteeg Chr. BW. 3070, Wissel Lakes,

West New Guinea. September 1955. LEIDEN.

fastigiata Bl. L. 908. 127-208, Java ? 18—. LEIDEN.

fragrans Roxb., Fresh specimens, Singapore Botanic

Gardens. *

. gardenioides Ridl., Burkill H.M. 2384 at Fraser’s Hill,

Pahang, Malaya. August 1960. SING.

gitingensis Elm., Lam. H.J. 3264, P. Karakelang, Talaud

Islands, Celebes. April 1926. LEIDEN.

. gracilipes A. Gray, Kostermans A. 371 at Manokwari,

Momi, New Guinea. September 1948. SING.

. involucrata Merr., Sinclair J. 8996 at Mt. Kinabalu,

Borneo. June 1957. SING.

. longiflora Merr., Conklin H.C. et Buwala PNH. 80440

Mt. Province, Luzon, Philippines. April 1963. LEIDEN.

. macroscypha Baker, Ashton P.S. BRUN. 5203 at Kuala

Belatong, Brunei. February 1959. SING.

. racemosa Jack, Nur Md. 25168 at Bukit Sagu, Pahang,

Malaya. October 1931. SING.

. resinosa Leenh., Asah v. BRUN 3083 at Bukit Belatong,

Brunei. June 1958. Sing.

. ridleyi K & G, Fresh specimens. Singapore Botanic

Gardens.*

. salticola Leenh., Saunders J.C. 777, Eastern Highlands

District, Terr. of New Guinea. August 1957. LEIDEN.

. tacapala Leenh., Eyma P.J. 2473 at W. Ceram, Moluccas.

December 1937. SING.

truncata Bl., de Vriese W.H. et Teijsmann J.E. L. 908.

127-616 Celebes 1859-60. LEIDEN.

tubulosa Bl., Ridley H.N. 9738 at Bujong Malacca,

Perak, Malaya. 1898. SING.

umbelliflora Gilg & Bened., Ledermann 9614, Sepik

Region, E. New Buinea. LEIDEN.

woodiana F v M, Womersley J.S. NGF. 15313, Morobe

District, E. New Guinea. September 1962. LEIDEN.

(*) A herbarium specimen was made for the Singapore

Herbarium.

196 Gardens’ Bulletin, Singapore — XXII (1967)

The procedures outlined previously by Foster (1955) and Arnott

(1959) were followed to obtain the leaf clearings and macerations.

The leaf clearings were used to study the density and mode of dis-

tribution of the sclereids and the macerations were used for the

study of the form, shape and size of the individual sclereid.

Transverse and paradermal sections were cut to study the struc-

ture of the leaf and the positional relationship of the sclereids,

and these were stained with Safranin and Fast Green, while clear-

ings and macerations were stained with Safranin alone.

To obtain the length of the sclereid, the two distant points

of the arms of the sclereid oriented in opposite directions were

measured; while for the width, the widest portion of the central

axis was measured. An average of ten such measurements was

taken. Sclereid density was assessed by visual observations.

Observation and Discussion

Of the thirty-one species investigated, sclereids were present

in the mesophyll tissues of all the species except Fagraea annulata.

However a transverse section did show the presence of four or five

sclereids in the midrib region of the leaf, which was also confirmed

with leaf clearings. In the remaining thirty species, a single type

of sclereid was seen in eleven of them, while sixteen others had two

types of sclereids and three species were found to have three types

of sclereids. (see Table 1).

The sclereids found in the various species could be grouped

together based on their morphology into types like the ASTER,

COLUMNAR, T-SHAPED and the DENDROID form.

1. Aster type: —

This type of sclereid displayed a central body with radiating

arms and a central lumen. The arms of the sclereid were branched

or unbranched, with or without spicules and had either short thick

arms with a lamellated cell wall or long, slender arms having a

homogenous cell wall.

For the convenience of description, the aster type sclereids can

be further classified into different sub-types: —

Aster Type A has few, short arms which are rarely branching.

Here the cell wall is thick and lamellated but the central lumen is

not prominent. This type of sclereid is seen in F. acuminatissima,

F. carnosa, F. crenulata, F. fragrans, F. gitingensis F. involucrata

and F. racemosa, (Figs. 1-12).

Aster Type B has a greater number of arms that are commonly

branching. The cell wall is thick and lamellated and the central

lumen is again not prominent. Such sclereids are seen in F. blumii,

F. bodenii, F. calcarea, F. carnosa, F. eymae, F. gardenioides, F.

resinosa, F. ridleyi and F. tubulosa, (Figs. 13-26, 69, 70, 74, 75).

Hardial Singh — Sclereids in Fagraea 197

Aster Type C has unbranched, short, radiating, slender arms.

The cell wall is homogenous. The relatively large central axis

encloses a very large central lumen which extends into the tips of

the arms. This type of sclereid is seen in F. curtisii, F. longiflora

and F. umberlliflora, (Figs. 27-31).

Aster Type D has a relatively small central axis with extremely

long, slender arms. Relative to the size of the central axis, the

lumen which it encloses is prominent and extends to the tips of the

arms. The cell wall is thin and homogenous. Occasional branching

of the arms does occur. This type of sclereid is seen in F.

acuminatissima, F. beteriana, F. calcarea, F. carstensensis, F.

ceilenica, F. fastigiata, F. gracilipes, F. tacapala, F. truncata,

F. salticola and F. woodiana, (Figs. 32-49, 68, 71, 81).

Thus, the aster type sclereid with its radiating arms was seen

in twenty-seven species and this group was further divided based

on size and mode of branching into four sub-types. The smallest

and largest sclereids were seen in this group. The small sclereids

were seen in F. carnosa, F. curtisii, F. fragrans, F. resinosa and

F. umbelliflora while the larger ones were seen in F.. acuminatissima,

F. calcarea, F. fastigiata and F. woodiana, (see Table 1).

2. Columnar type: —

This type of sclereid was very prominent being at times very

long and wide. Here the body of the sclereid varied from being

short to extremely long. In these sclereids the cell wall is thick and

lamellated while the central lumen is prominent. This columnar type

of sclereid could be further divided into Subtype A where the arms

were short and restricted to the very ends of the sclereids, and the

cell body had an extremely prominent columnar form. Such

sclereids were seen in F. auriculata, F. macroscypha and F. resinosa

(Figs. 50-54, 72, 85). In a transverse section of the leaf

of F. resinosa, the pillar-like sclereids can be seen as vertical

columns connecting the upper to the lower epidermis, (Fig. 86).

Sub-type B has arms which arise more or less from the entire

length of the cell body and hence a prominent pillar-like form is

not seen. Also these sclereids are much smaller than those placed

in sub-type A. Such sclereids are seen only in F. ridleyi, (Figs. 55-

56, 74).

3. ‘T’-shaped type:—

In these sclereids, the cell wall lying next to the upper epidermis

develops a number of outgrowths that grow in between the tan-

gential walls of the epidermal or hypodermal cells. The vertical arm

of the ‘T’-shaped sclereid extends for varying distances into the

mesophyll tissue and occasionally it produces a number of branches.

The cell wall of the sclereids could be thick and lamellated or thin

and homogenous. The central lumen is usually prominent. ‘T’-

shaped sclereids are found in F. acuminatissima, F. auriculata,

F. beteriana, F. blumii, F. bodenii, F. carnosa, F. fastigiata F.

198 Gardens’ Bulletin, Singapore — XXII (1967)

gardenioides, F. gracilipes, F. macroscypha, F. ridleyi, F. salticola,

F. tacapala, truncata and F. umbelliflora, (Figs: 57-63, 73).

This type of sclereid though seen in sixteen of the species

was not very prominent since relatively few such sclereids were

seen in the leaves. In species where other types of sclereids

were found besides the ‘T’-shaped form, the latter exhibited

characters common to the other types of sclereids with regard to the

nature of the arms, cell wall and lumen.

4. Dendroid type: —

This type of sclereid has no typical shape nor a definite pattern

of branching. The branches of the sclereids appear as tube-like

filaments with a rather uniform thickness along their length. This

type of sclereid is seen in only three species, namely F. elliptica,

F. involucrata and F. racemosa, (Figs. 64-67, 76).

Foster (1946) working on Mouriria, recognised four types of

sclereids, namely Parenchymatous, Stellate, Columnar and Filiform.

Though the parenchymatous and filiform types are not seen in

Fagraea, the aster type which is alike the stellate type and the

columnar type are present. In Mouriria, the majority of the species

showed conspicuously branched polymorphic sclereids as also seen

in the genus Fagraea and in marked contrast to this, Rao (1957)

working on Memecylon, recognised three major types of sclereids

of which the filiform type predominated in eighty-two of the ninety-

five species investigated. Foster (1946) did record in one species —

Mormriria cauliflora, the presence of two types of sclereids namely

columnar and parenchymatous. In Fagraea however, this pheno-

menon of having more than a single type of sclereid in a species

appears to be very common and in some species like F. acuminatis-

sima, F. carnosa and F. ridleyi, three different types of sclereids are

present, (see Table 1). Also Rao (1957) did observe in Memecylon

cuneatum, thick columnar sclereids alternating with fusiform

branched sclereids.

In most of the species the sclereids are rather crowded but

in the following, F. beteriana, F. crenulata, F. curtisii, F. fragrans,

F. gardenioides, F. gitingensis and F. involucrata they are sparsely

distributed, (Figs: 77-80, 83). There appears to be no correlation

between the sclereid density and the thickness of the leaves. Though

in some thick leaves like F. bodenii and F. ridleyi the sclereids

are densely distributed (Fig. 84) there are also those species with

thick leaves having sparsely distributed sclereids as seen in F.

curtisii, F. gardenioides and F. involucrata. Again in certain species

with thin leaves the sclereids were densely distributed as seen in

F. ceilenica, F. fastigiata and F. umbelliflora (Fig. 82). As has

been revealed by the present studies, unless more studies are made

with other angiosperm species, one cannot make any definite in-

ference regarding the relationship between thickness of leaves and

sclereid density.

Hardial Singh — Sclereids in Fagraea 199

Distribution of sclereids is predominantly diffuse except in

F. beteriana, F. crenulata, F. fragrans, F. gardenioides, F. gitin-

gensis and F. involucrata where the sclereids were seen in both the

diffuse and terminal positions. This predominantly diffuse con-

dition is alike that found in Memecylon (Rao, 1957) but contrasts

to that seen in Mouriria (Foster, 1946) where the sclereids were

observed to be mainly in a terminal position. Foster (1955) working

on Boronella observed that though the sclereids were predominantly

terminal in B. crassifolia and B. verticillata, they were predomin-

antly diffuse in B. pancheri and B. francii though some collections

of B. pancheri showed a predominantly terminal postition of the

sclereids.

Leenhouts (1962) revised the genus Fagraea and recognised

thirty-one species distinguishing them by using vegetative and floral

characters like the form of leaves, being crenulate or entire, rounded

or acute, with or without auricles, the inflorescence being axillary

or terminal, the flowers being small or large and with a tubular

or funnel shaped corolla tube. Leenhouts work has been used as

the basis for names of the species investigated. Leenhouts divided

the genus into three subgroups but the sclereid types of the different

species placed by him within the same subgroup vary tremendouslv.

In the section Cyrtophyllum which includes F. elliptica, F. fragrans

and F. umbelliflora, F. elliptica has a single type of sclereid which

is dendroid in form. F. fragrans has also a single type of sclereid

which is aster shaped while F. umbelliflora has two types of

sclereids, one being aster shaped while the other is the “T’-shaped

type. Also the aster type seen in F. fragrans is unlike that found in

F. umbelliflora. It is also observed that the only three species having

dendroid type sclereids, namely F. elliptica, F. racemosa and F.

involucrata, have been each placed in a different section. It there-

fore appears that the sclereid characters of the different species

of this genus do not seem to be helpful in further substantiating

the taxonomic classification of Leenhouts based on the criteria

already mentioned.

Summary:

Sclereids were seen to be present amongst the mesophyll tissues:

of the leaf in thirty of the species of Fagraea investigated. Due

to the great variation in form of the sclereids, these were classified

into four distinct types namely, aster, columnar, “I’-shaped and

dendroid forms. Presence of more than a single type of sclereid in a

species was commonly seen and some species like F’. acuminatis-

sima, F. carnosa and F. ridleyi had three different types of sclereids.

Most of the species had densely distributed sclereids though no

correllation was seen between the sclereid density and the thickness

of the leaf. Distribution of the sclereids was predominently diffuse

and only in a few species was the terminal postion seen together

with the diffuse sclereids. The sclereid characters do not appear

to be helpful in substantiating the taxonomic classification of

Leenhouts.

200 Gardens’ Bulletin, Singapore — XXII (1967)

The writer is grateful to Mr. H. M. Burkill, Director Botanic

Gardens, for his encouragement, Dr. A. N. Rao, Department of

Botany, University of Singapore, for his constant guidance and the

use of the university facilities, Mr. Raymond Tay for assistance in

the practical work, Mr. Douglas Teow for the photographs and

Mr. Juraimi bin Samsuri for help in the drawings.

Literature Cited:

Arnott, H. J., Leaf Clearings, Turtox News, 1959, 37, 192.

BaRuA, P. K. and WIGHT, W., Phytomorphology, 1958, 8, 257.

CARLQUIST, S., Comparative Plant Anatomy, Holt, Reinhert and Winston,

New York, 1961.

Caspary, R., Mig. Ann. Musei Bot. Legd. Bat., 1865, 2, 241.

Foster A. S., J. Arn. Arboretum, 1946, 27, 253.

Foster, A. S., J. Arn. Arboretum, 1955, 36, 189.

LEENHOUTS, P. W., Flora Malesiana, 1962, 6, 293.

METCALFE, C. R., and Chalk, L., Anatomy of the Dicotyledons, Clarendon

Press, Oxford, 1950.

Rao, T. A., Phytomorphology, 1957, 7, 306.

Rao., A. N., Curr. Sci., 1965, 34, 509.

TOMLINSON, P. B., New Phytol., 1959, 58, 253.

TIEGHAM, P. van, Ann. Sci. Nat. vii. Bot., 1891, 13, 23.

Ad | 431 | 304

TABLE I

SCLEREID

Species a

P thickness ;

Types | Length Width |Positional R| Density

. acuminatissima 512 us Aa 256 52 ps i eee N2

Ad 426 35 pu

T 454 u 3

. annulata 128 pu dee — — — a

. auriculata 611 u Cc 302 52 Sg N2

T 530 ph 50 Be

. beteriana 398 Ad 226 pu 32 p ‘.. 23 oe N3

T 220 28 us

. bodenii Sll pu Ab 355 pu 35 um 1. 355% Nl

T 284 u 25 p

wy 255 ph 19 u

F. carstensensis 327 p Ad 372 p 38 pb 3, 4 N2

F. calcarea 284 pu Ab 369 uo 36 pu 3, 4 N2

Hardial Singh — Sclereids in Fagraea 201

TABLE 1— continued

SCLEREID

; Leaf

grees thickness

Types | Length | Width |Positional R| Density

9. F. carnosa 412 u Aa 142 u 32 p Ly ay oat & N2

Ab | 196 x | 60 pu

Oe re ee

10. F. ceilenica 199 Ad | 309 | 22 p 2,374 | NI

11. F. crenulata 341 u Aa 239 29 3, 4 N3

12. F. curtisii 525 Ac 173 pu 38 pu Zs SF N3

14. F. eymae 497 u Ab 338 u 35 > S N2

15. F. fastigiata 241 pu Ad | 830 4 38 3 Nl

T8600 9 | 35 p

16. F. fragrans 256 pu Aa 173 p 35 us 3, 4 N3

17. F. gardenioides rare |. Ab 993 gp | 4fin | 1/5. 3,-4 N3

fT. (969 we | 35 »

18. F. gitingensis 340 u Aa 255 41 u 2, 34 N3

19. F. gracilipes 327 p Ad 312 uw 16 p I, 4 3, 4 Nl

T | 21 | 16 »

20. F. involucrata 426 Aa | 227» | 324 * | Na

D | 3584 | 16 4

21. F. longiflora 284 u Ac 269 u 43 po 2, 3, 4 N2

22. F. macroscypha 341 e 213 26 u Lf) 38 N2

T 324m | 29-p

32 a, 3, 4 N2

23. +F. racemosa 241 pu Aa | 200 u ph

| . G 426 pu Dp |

24. F. resinosa 511 Ae | eit) Ste KS Ne

| C 350 wp | 42 p

25. F. ridleyi 525 A200, | S442 | 1.43.4) Ni

| + 315 p 22 po

26. F. salticola 600 pu Aus |6308 ary | 40 me -) 34] ND

| Po |\,2845 | 32m

27. F, tacapala 284 Ey i a

| i 213 w | 16

| 28. F. truncata 369 Ad | 332 | 22h | ' 3,4 | N2

| rT 426 ps 16 p

29. F. tubulosa 256 u Ab | 244 4 49 u 2, 3, 4 N2

30. F. umbellifiora 213 p Ac 184 16 u pas, @ Nl

|. T*, 210 p 32 p

31. FB. woodiana 200 » Ad 437 p 42 u 3, 4 N2

N1 = Dense

N2 = Numerous

N3 = Few

Gardens’ Bulletin, Singapore — XXII (1967)

202

WoLynogs

Hardial Singh — Sclereids in Fagraea 203

13 |

15 17

a, 20

Plate II. “

Sclereid forms: Fig. 1—F. carnosa, Figs. 2-4—F. crenulata, Figs.

S-7—F. fragrans, Figs. 8-9—F. gitingensis, Figs. 10-11—F.

involucrata, Fig. 12—F. racemosa, Figs. 13-15—F. blumii, Fig.

16—F. bodenii, Fig. 17—F. calcarea, Fig. 18—F. carnosa, Figs.

19-20 F. eymae. Camera lucida drawings, magnification x 100.

204 Gardens’ Bulletin, Singapore — XXII (1967)

30 52

Plate III.

Sclereid forms: Figs, 21-22—F. gardenioides, Fig. 23—F. resinosa,

Fig. 24—F. ridleyi, Figs. 25-26—F. tubulosa, Fig. 2 —F. curtisii,

Figs. 28-29—F. longiflora, Fig. 32—F. acuminatissima, Figs. 33-34

—F. beteriana, Fig. 35—F. calcarea, Figs. 36-37—F. carstensensis.

Camera lucida drawings, magnification x 100.

Hardial Singh — Sclereids in Fagraea 205

AA 5 42

Plate IV.

Sclereid forms: Figs. 38-39—F. ceilenica, Figs. 40—F. fastigiata, Figs.

41-42—F. gracilipes, Fig. 43—-F. tacapala, Figs. 44-45—F. truncata,

Figs. 46-47—F. salticola. Camera lucida drawings, magnification

x 100.

206 Gardens’ Bulletin, Singapore — XXII (1967)

ee a ee

ebay gh

Plate V.

Sclereid forms: Figs. 48-49—F. woodiana, Fig. 50—F. auriculata, Figs.

51-52—F. macroscypha, Figs. 53-54—F. resinosa, Figs. 55-56—F.

ridleyi. Camera lucida drawings, magnification x 100.

Hardial Singh — Sclereids in Fagraea 207

66 67

Plate VI.

Sclereid forms: Figs. 57-58—F. acuminatissima, Fig. 5)—F. auriculata,

Fig. 60—F. blumii, Fig. 61—F. bodenii, Fig. 62—F. carnosa, Fig.

63—F. gracilipes, Fig. 64—F. elliptica, Fig. 65—F. involucrata,

Figs. 66-67—F. resinosa. Camera lucida drawings, magnification

x 100.

208 Gardens’ Bulletin, Singapore — XXII (1967)

Plate VII.

Sclereid forms: Fig. 68—F. tacapala x 105, Fig. 69—bodenii x 61,

Fig. 70—F. tubulosa x 136, Fig. 71—F. gracilipes x 71, Fig.

72—F. macroscypha x 92, Fig. 73—F. acuminatissima x 65.

Hardial Singh — Sclereids in Fagraea 209

| Plate VIII.

| Sclereid forms: Fig. 74—F. ridleyi x 109, Fig. 75—F. calcarea x

95, Fig. 76—F. elliptica x 97.

210 Gardens’ Bulletin, Singapore — XXII (1967)

Plate IX.

Leaf clearings showing sclereid densities. Fig. 77—F. crenulata x 67, |

Fig. 78—F. fragrans x 71. Fig. 79—F. beteriana x 71, Fig. 80— |

F. gardenioides x 67, Fig. 81—F. gracilipes x 63, Fig. 82—F.

ceilenica x 80.

Hardial Singh — Sclereids in Fagraea 21k

Plate X.

Transverse sections of leaves showing sclereid densities. Fig. 83—F.

fragrans x 105, Fig. 84—F. ridleyi x 189, Fig. 85—F. auriculata

; he

212 Gardens’ Bulletin, Singapore — XXII (1967)

Upper

epidermis

Palisade

Spondy

mesophyll

Lower

e pidermis

Plate XI.

Fig. 86—F. resinosa—T.S. leaf showing columnar scalereids x 214,

Fig. 87—Diagram indicating the positional relations of sclereids.

Notes on Sapotaceae

(Written July 1965)

JAMES SINCLAIR

Botanic Gardens, Singapore.

It was through the misidentification by Leiden for a certain

Malayan species of Sapotaceae and the rather unusual circum-

stances connected with it that the following notes came to be

written. In trying to identify it myself later, I had to consult the

Precursores for this family to some extent and as a result noticed

a number of minor details with which I cannot agree. I have,

however, to acknowledge the good work that has been accom-

plished by the authors of these Precursores.

I. A Misunderstood Species of Sapotaceae

On 21st May 1953 I collected at Tanjong Gul, Singapore, a

sterile Sapotaceae, Sinclair S.F.N. 39640, and also numbered Sinclair

7446 in my own series. It was growing by the sea-shore at the foot

of the grassy wooded sea-cliffs. Incidentally Tanjong Gul is the

station for a number of rare plants in Singapore such as Vatica

wallichii, Casearia capitellata, Adinobotrys erianthus, Rhizophora

stylosa, Diospyros styraciformis, Urceola torulosa, Ipomoea digitata,

I. illustris, Knema corticosa (globularia), Ficus consociata var.

murtoni, F. xylophylla, Athyrium prescottianum and Dipteris conju-

gata. Haji Mohamed Nur, herbarium assistant at that time, made

various suggestions as to its identity. The sheets bear the names

Payena, Planchonella obovata and Madhuca sericea but I deleted

them all in turn, not being satisfied. It was eventually sent on

loan to Leiden with the rest of the Sapotaceae when they were

being revised for Flora Malesiana and returned in due course with

the remarks cf. Myristicaceae. Dr. P. van Royen had also seen

it as his initials and the date 8-4-61 were inscribed on the sheet.

As it happened I had revised and published the Malayan species of

Myristicaceae by 1958 and would not have mistaken the present

plant with its white sap for one of the Myristicaceae. Perhaps the

person who wrote cf. Myristicaceae on the sheet was not looking

at collectors’ names but was more concerned with the appearance

of the specimen itself.

On looking among the named Sapotaceae then back from loan,

I noticed that it resembled two sheets, both named Payena grandi-

flora Ridley by van Royen——(1) Ridley 6497 from Sungei

Morai, Singapore and (2) Ridley 11371 from? Sumatra, but culti-

vated in the Botanic Gardens, Singapore.

I picked out these two sheets and my own sterile specimen and

sent them again to Leiden so that van Royen could see them side by

side. I wrote on the species cover “Dr. v. Royen —— Please

return these three mounted sheets to Singapore. The two specimens

214 Gardens’ Bulletin, Singapore — XXII (1967)

collected in Singapore near the sea-shore are obviously the same

as the Sumatran one. I think you overlooked them and did not

see the connection. I cannot at present check if you have kept

Payena grandiflora as a good species since our copy of your

publication has gone to the binders and they usually keep such

things for a long time. Can you please re-examine and annotate?”’

J. Sinclair.

This material was returned later with other Sapotaceae but

without any comments or further annotations. I do not think van

Royen saw it at all. It was about that time he left Leiden to take

up an appointment with the Forestry Department at Lae. In fact

I can recall receiving a letter from someone at Leiden informing

me of his departure and of the fact that he had not been able :

to examine a duplicate of Palaquium beccarianum which I had sent :

to him for checking. Our own specimen of this was sent to him

later at Lae and I only came across it the other day in the her-

barium when writing this article. But returning to the subject of the

unidentified plant I had nothing further to expect until the 3rd of

November 1963 when it eventually flowered at Tanjong Gul.

The Tanjong Gul specimens are indeed Payena grandiflora

Ridley and Ridley cites the following specimens with his descrip-

tion (1) Ridley 6497 (SING) Sungei Morai, Singapore; (2)

Goodenough 1268 (SING) Panchur, Malacca and (3) Ridley

113711 (SING) cultivated in Botanic Gardens, Singapore, believed

to have come from Sumatra. The last mentioned specimen actu-

ally bears the number //37/ on the label of the Singapore sheet

and not /13711 as stated in the publication. Ridley says that the

species is described mainly from the tree in the Botanic Gardens, :

a ——s =

i.e. Ridley 11371 and adds that this fine species has been confused

with Payena maingayi by King and Gamble in the Materials. But

there are four sepals in its flowers and not five as stated by him. /

In the revision of Payena in Blumea 9, 1 (1958) 89-138, the

author A.C. van Bruggen has sunk Payena grandiflora Ridley under

P. maingayi Cl. On page 107 he states that on account of Ridley’s

erroneous remark about the sepals (Ridley loc. cit. 29) “Sepals |

ovate, subobtuse, nearly half an inch long 5,...’’ Lam transferred

P. grandiflora to the genus Diploknema. In our opinion it is

evident that this species is quite identical with P. maingayi. I must

point out now that van Bruggen annotated most of our specimens

of Payena while van Royen, who wrote up Madhuca, annotated

most of the latter. There are, however, occasional sheets of Payena !

which van Royen named. Lam was in a way nearer to the truth

when he regarded it as a good species and also when he removed |

it from Payena; the sepals are, indeed, half an inch long. One |

of the syntypes, Goodenough 1268, is actually Payena maingayi,

but the other two are one species, namely Payena grandiflora of

Ridley which was based, as pointed out above, mainly on Ridley ;

11371. It was not the custom then to choose a holotype and so I |

have now to make Ridley 11371 the lectotype of Payena grandi- |

flora and to exclude Goodenough 1268. |

Sinclair — On Sapotaceae 215

I was not, however, satisfied that Payena was the correct genus

and after investigation found that the specimens really belong to

Madhuca chiefly because of the structure of the flower and the

number of its parts. I cannot call the species Madhuca grandiflora

because a Madhuca grandiflora Fletcher already exists. A new

name must therefore be chosen so I have adopted the epithet

Madhuca decipiens J. Sinclair.

Madhuca decipiens J. Sinclair, nom. nov.

Basionym: Payena grandiflora Ridley in J. Roy. As. Soc.

Str. Br. 61 (1912) 28 excl. spec. Goodenough 1268; Ridley, FI.

Mal. Pen. 2 (1923) 262 excl. Goodenough 1268 = P. maingayi Cl.

Isonym: Diploknema grandiflora (Ridley) H.J. Lam in Bull.

Jard. Bot. Btzg 7, 1 and 2 (1925) 185 excl. Goodenough 1268 et

in Bull. Jard. Bot. Btzg 8, 4 (1927) 463.

Shrub (6-10 ft) 2-3 m high with copious white sap. Leaves

dark green and glossy above with a paler midrib, pale yellowish

green and dull beneath, drying greyish brown to dark brown

above and pale yellow beneath, becoming greyish when old. Sepals

4, greyish tomentulose outside, glabrous inside and on the margins

of the two inner ones as well, the outer two ovate, 1 cm long and 8

mm broad, the inner two narrower, 1 cm long and 7 mm broad and

distinct from the outer in having a carinate midrib on the outside.

Corolla white, thin in texture, 1 cm long in bud and 1.5 cm long

when mature (i.e. it protrudes 5 mm beyond the calyx when mature)

split down 4—way into the 9-12 acuminate lobes, tomentulose

except for the free parts of the lobes. Stamens 26-31 and in three

rows, the very short filaments only 0.75 mm long, densely hirsute,

the anthers 2 mm long, their slender appendages 1.5 mm long, both

sparsely covered with 0.5—-1 mm long whitish hairs. Ovary densely

tomentose with 10 loculi and one ovule in each loculus; style 1 cm

long, its basal third with short adpressed, greyish white hairs,

the rest glabrous; pedicels 1.5—2.5 cm long and 1.5-2 mm thick,

minutely pubescent.

JOHORE : Gunong Pulai, Md Nur & Kiah S.F.N. 7785

(SING) not “Md Mutkiah’” as stated by

Lam l.c. 186.

SINGAPORE : Sungei Morai, Ridley 6497 (SING); Tanjong

Gul, Sinclair 7446, also numbered Sinclair

S.F.N. 39640 (SING) sterile and Sinclair

10761 (A, E, G, K, L, LAE, SING) in

flower.

(CULTIVATED : Cultivated in Bot. Gard. Sing. probably

from Sumatra, Ridley 11371 (SING)

quoted as //3711.

TYPE MATERIAL: Lectotype of Payena grandiflora Ridley =

Ridley 11371 (SING).

The above description is taken from Sinclair 10761 and duplicates

of it are being distributed. Three shrubs were seen in all. Lam is

right} when he says that the Johore specimen cited above is not

Payena maingayi but probably belongs to the present, i.e. Ridley’s

species. He, however, did not read the handwriting on the label

correctly, see above under JOHORE. I at first thought that this

Species might be confined to the sea-shore as both Tanjong Gul

216 Gardens’ Bulletin, Singapore — XXII (1967)

and Sungei Morai are sea-shore localities. Apparently this is not

the case as Gunong Pulai is a mountain. The present species is.

very different from Payena maingayi, the latter a tall forest tree,

reaching at least 70 feet high and which I have seen in Bukit Timah

Nature Reserve, Singapore. The leaves of P. maingayi are much

thinner in texture, dry blackish above, and have a dark rusty, scaly,

powdery pubescence on the under-surface like those of Chrysophyl-

lum cainito. Those of the Madhuca are much paler beneath, a

pale yellow which becomes greyish when old and tends to disap-

pear; furthermore they are finely reticulate on both surfaces with

longer and stouter petioles. There is a number of differences also in

the flowers, especially in the larger number of petals, stamens and

loculi. But it will be seen without dissecting them that those of the

Madhuca are slightly larger with stouter pedicels and more coria-

ceouls sepals.

I was not so interested, however, in comparing this Madhuca

with P. maingayi as in trying to find out its proper place among the

other Madhuca species. It soon became apparent that it did resem-

ble one other species and this investigation led to further un-

expected discoveries as I had then to consult rather closely the:

Precursores on Payena and Madhuca. From this consultation L

found also that there were some minor points in which I cannot

quite agree with all that has been written in the Precursores. It

seems best to discuss the relationship of Madhuca decipiens under

the next heading since some name changes result.

II. On the Relationship of Madhuca decipiens; A New

Combination in Madhuca and Madhuca dubardii

H. J. Lam redefined

I soon found a near relative to the above species in Payena

selangorica King and Gamble, the type being Ridley 7387 (SING)

from Bukit Kutu, Selangor, Malaya (young fruiting material). In

fact this is a miniature edition of Madhuca decipiens, a very

similar but more elegant species. The rather similar leaves are

smaller and their petioles are also reduced proportionately in

length and thickness. Ridley 7387 is the only authentic specimen

in the Singapore herbarium named P. selangorica, the only one

quoted under the original description by King and Gamble in the

Materials, page 175 and also the only one quoted in the Pre-

cursores, page 131. But if P. selangorica is so like Madhuca de-

cipiens then it must surely be a Madhuca and this can indeed be

seen from a comparison of the actual specimens. Van Bruggen in

the Precursores, page 133 states that its status is still doubtful and

that according to the present material it could belong to Madhuca

or Ganua as well,...etc. A search was then made among our

Madhuca material to see if any other specimens of it could be

located under some other name. Four sheets were found and the

specimens were in flower. From these I am able to confirm that it is

a Madhuca. Two of these sheets are named Madhuca dubardii

H. J. Lam var. dubardii, namely Strong 11115 and Strugnell

11116 and the other two Abdul Rahman 10527 and Symington

Se ee ee ee

Sinclair — On Sapotaceae 217

24097 Madhuca dubardii var. lanceolata H. J. Lam. The Abdul

Rahman specimen 10527 (SING) is the type of the var. lanceolata.

In my opinion, however, these specimens are all very uniform and

represent only one species which should not have been divided

into two varieties. The leaves are narrowly elliptic and acute at

the apex; those with the apex damaged tend to be obovate and

obtuse at the apex. I find the same thing applies in my specimens

of Madhuca decipiens. A Penang specimen, Paul s.n., named by

van Royen and quoted by him in the Precursores, page 105 under

M. dubardii var. dubardii is, however, not that species but

Palaquium curtisii (K. & G.) H. J. Lam instead. The leaves are

slightly juvenile and match those of a similar specimen of P.

curtisii from Penang wrongly named P. gutta (Hooker f.) Baillon

by van Royen. This is Curtis s.n. date 25th October, 1900.

The type of M. dubardii H.J. Lam (1925) was based on

material from Suan Lamba, Sabah (British North Borneo) namely

Agama 538 (BO holotype, K). Suan Lamba is not in Sarawak as

is stated by van Royen. I then got together all the named and

unnamed sheets of this species in our herbarium from all the Sabah

localities which I could find. Two of these are quoted under

dubardii by van Royen himself in the Precursores, page 105,

namely Cuadra A2148 and Puasa 10076. The former bears the

determinavit slip of van Royen who saw this sheet; the latter, the

SING duplicate, was not sent to him as it was discovered later

among unmounted material, but he has seen its K and L duplicates.

I found two other sheets of dubardii namely Castro 3793 and

Sales 4311 which had been wrongly named and quoted as Payena

acuminata var. acuminata. by van Bruggen on page 103 of his

Precursores on Payena. Having made a careful comparison of the

Sabah material of M. dubardii with that of the Malay Peninsula I

find that they represent two distinct species. The Malay Peninsula

material is not different from Payena selangorica but the latter is

a Madhuca and a new combination is necessary.

Madhuca selangorica (King & Gamble) J. Sinclair, comb. nov.

Basionym: Payena selangorica King & Gamble, Mat. Fl. Mal.

Pen. 17 in J. As. Soc. Bengal 74, 2 Extra Nr. (1906) 175; idem

Addenda-Corrigenda (1908) 873; Ridley, Fl. Mal. Pen. 2 (1923)

264; Lam in Bull. Jard. Bot. Btzg 7, 1 and 2 (1925) 132 et l.c. 8, 4

(1927) 439; van Bruggen in Blumea 9, 1 (1958) 131 f. 11.

Synonyms: Madhuca dubardii (non H. J. Lam 1925) H.J. Lam

in Bull. Jard. Bot. Btzg 8, 4 (1927) 444 f. 18; van Royen in

Blumea 10, 1 (1960) 104 quoad spec. selangorenses tantum.

M. dubardii var. lanceolata H. J. Lam in Bull. Jard. Bot. Btzg

8, 4 (1927) 446 f. 18; van Royen in Blumea 10, 1 (1960) 105

syn. nov.

Sepals 7 mm long and 5 mm broad, medium brown tomentulose

to tomentose outside, rugose when dry, the inner two with glabrous

margins and carinate on the outside like those of M. decipiens.

‘Corolla 5 mm long and split down ?—way into 8 lobes, the latter

acute at the apex, adpressed sericeous on the outside and at the

218 Gardens’ Bulletin, Singapore — X XII (1967)

base inside. Anthers 16, hirsute with an acute apex (not produced

into a slender process as in decipiens but the flowers examined

were in a Slightly younger stage). Ovary tomentose with 8 loculi

and a 5 mm long style; pedicels sulcate, 2 cm long and 1 mm thick.

SELANGOR : Bukit Kutu, Ridley 7387 (SING); Ayer

Hitam Reserve, Kajang, Abdul Rahman

10527 (SING); Bukit Tunggal Reserve,

Strong 11115 (SING); Ampang Reserve,

Selangor, Strugnell 11116 (SING); Sungei

Lalang, Kajang, Symington 24097 (SING).

DISTRIBUTION : Selangor in Malaya. Van Royen in Blumea

l.c. 105 mentions a specimen from West

Coast, Sumatra. This is not in Herb.

Sing. and I have not seen it at the time

of writing.

TYPE MATERIAL: Payena selangorica K. & G., Ridley 7387

(SING holotype). M. dubardii var. lan-

ceolata H.J. Lam, Abdul Rahman 10527

(SING holotype).

As pointed out M. selangorica is smaller in all its parts than

M. decipiens. The colour of the under-surface of the leaves is of

a paler yellow in decipiens, that in selangorica being more rusty.

Later on when the scales tend to fall off the colour changes to

an ashy grey in both. There are 9-12 corolla lobes in decipiens as

against 8 in selangorica. The lobes are acuminate at the apex

in the former and acute in the latter. The anthers are more numer-

ous also, 26-31 as against 16 in selangorica. In decipiens they are

produced at the apex into a slender filiform process; in selangorica

they are simply acute and not produced. The anthers were not

quite mature in the flowers of selangorica which I examined, but

I do not think they will increase very much more at the apex.

Madhuca dubardii H.J. Lam in Bull. Jard. Bot. Btzg 7, 1 and 2

(1925) 162 f. 43 [non M. dubardii Lam et vars Lam l.c. (1927) 444

f. 18=M._ selangorica (K. & G.) Sinclair]; van Royen in Blumea

10, 1 (1960) 104 quoad spec. borneenses tantum.

Sepals 5 mm long and 4 mm broad, pale brown-tomentulose

when dry, the two inner ones with a groove on the outside in

place of the mid-vein. Corolla 4 mm long and split down #—-way

into about 9 lobes, the latter obtuse at the apex, pubescent at the

base inside but not outside and not on the free lobes. Anthers

16-19 with a few scattered hairs, acuminate at the apex, some

with a slender process. Ovary densely tomentose with 10-11 loculi;

style 4 mm long; pedicels sulcate, 1.5—-1.8 cm long and 1 mm thick.

SABAH (BRITISH

NORTH BORNEO): Suan Lamba, Agama 538 (BO, K); Balach-

ing River, Sandakan, P. Castro 3793

(SING); Sekong River valley, Sales 431]

(SING); the remainder Kinabatangan :—

Kori timber camp, Austin Cuadra A2148

(SING); Supu Forest Reserve, Puasa

10076 (SING); Daramakud timber camp,

Kadir bin Abdul SAN 16864 (SING).

DISTRIBUTION : So far confined to Sabah.

TYPE MATERIAL: Madhuca dubardii H. J. Lam, Agama 538

(BO holotype, K) not seen by me at the

time of writing.

Sinclair — On Sapotaceae

219

I hereby exclude all the Malay Peninsula, Selangor specimens

of M. dubardii and M. dubardii var. lanceolata from this species.

They are, as has been pointed out M. selangorica. The differences

between M. selangorica and M. dubardii are here expressed in

tabular form.

Leaves

Petioles

Flowers

Pedicels

Calyx

Corolla

Anthers

Madhuca selangorica

Mostly narrowly elliptic,

less often obovate; up-

per surface finely re-

ticulate with raised veins

and reticulations, mostly

glossy when dry; lower

surface when young a

rusty brown colour due

to scales, the veins very

faint

2-2.3 cm long, proportion-

ately longer than in

dubardii depending on

the size of the blade

Larger, 7 mm long and 5

mm broad, medium

brown, tomentulose to

tomentose

2 cm long

7 mm long and 5 mm

broad, rugose, the inner

two with a carinate mid-

vein on the back (out-

side)

5 mm long, densely pubes-

cent at the base inside

and adpressed-sericeous

outside except towards

the apices of the lobes;

lobes acute at the apex

2 mm long, hairy, acute

at the apex but not pro-

duced into a filiform

appendage

8 loculi observed (pro-

bably 10 also)

Madhuca dubardii

Mostly obovate, less often

elliptic; broader at the

middle; upper surface

smooth and dull without

the veins and: reticula-

tions being visible or if

present then very faint,

the veins sunk; lower

surface paler and more

yellowish, the veins

more prominent

1-1.5 cm long, the blade-

petiole ratio propottion-

ately less than in selan-

gorica

Smaller, 5 mm long and 4

mm broad, paler brown,

tomeritulose, the tomen-

tum less

1.5-1.8 cm long

5 mm long and 4 mm

broad, smooth and not

rugose, the inner two

with a sulcate mid-vein

on the back

4 mm long, pubescent only

at the base, glabrous

outside and inside higher

up; lobes obtuse at the

apex

2.5 mm long, much less

hairy, the apex acumin-

ate or sometimes pro-

duced into a slender ap-

pendage

10-11 loculi

220 Gardens’ Bulletin, Singapore — XXI1 (1967)

III. On the Identity of Payena ridleyi Gandoger

It was, when trying to identify my specimens of Madhuca de-

cipiens from Tanjong Gul, that I noticed a doubtful species Payena

ridleyi Gandoger at the end of A.C. van Bruggen’s Precursores on

Payena in Blumea 9, 1 (1958) 133. The type of that species is there

stated to be Ridley 6698, Singapore, and van Bruggen adds: —“‘Not

having seen the type specimen we can not refer it to one of the

known species; moreover, the description is quite insufficient. There

is a possibility that it is identical with P. maingayi (1882) according

to the remark that the leaves are twice as long as those of P.

costata=P. lucida.’’ As I did not then know the identity of my

Tanjong Gul specimens there was the possibility that they might be

identical with Gandoger’s species if that could be found. By some

kind of intuition and remembering that I had once written

Madhuca sericea on the sterile sheet, Sinclair S.F.N. 39640 and

scored it out later, I went straight to the genus cover of Madhuca

sericea in the herbarium and found the missing Ridley’s number

there. It had been identified and labelled as Madhuca sericea (Miq.)

Lam by van Royen who revised Madhuca. I do not know whether

van Bruggen, the author of Payena in the Precursores, saw it or

not. However, the collector is not Ridley himself but Mat, his

collector. All inscriptions on the label are in Ridley’s handwriting.

Mat is Ahmad bin Hassan M.B.E. who is still alive and probably

nearly ninety years old. He collected this specimen, “‘a big tree’,

according to the label, on 15th May 1894, some 71 years ago at

Chan Chu Kang, Singapore. Sometimes we find on labels of plants

collected in the past in Malaya the names of native collectors

instead of those of the botanists with whom they worked, e.g.

Alvins = Cantley’s collector, Haniff = Curtis, and Mohamed

Nur or Haniff = I. H. Burkill’s collectors. The specimen in Singa-

pore will be the isotype of Payena ridleyi Gandoger, the holotype

will be a duplicate of this at Lyons. I have thus to record this

species aS a new synonym of Madhuca sericea.

Madhuca sericea (Miquel) H.J. Lam in Bull. Jard. Bot. Btzg 7,

1 and 2 (1925) 163 and 264 f. 44; Ridley, Fl. Mal. Pen. 5 (1925)

Suppl. 319; Lam, l.c. 8, 4 (1927) 446; van Royen in Blumea

10, 1 (1960) 70.

Basionym: Payena? sericea Miq. Fl. Ind. Bat. 2 (1859) 1039.

Synonym: Payena ridleyi Gandoger in Bull. Soc. Bot. France

65 (1918) 56; van Bruggen in Blumea 9, 1 (1958) 133

syn. nov.

SINGAPORE : Chan Chu Kang, Mat 6698 (LY, SING).

TYPE MATERIAL: Payena sericea Miq., Horsfield s.n. (BO).

Payena ridleyi Gandoger (Ridley) Mat

6698 (LY holotype, SING).

“4

Re a

a th SE

eee ae

—

en eee ee

Sinclair — On Sapotaceae 221

IV. On Other Missing Specimens

There are other cases of missing specimens which could not be

located. The reason again seems to be that mentioned above where

it was not realized that the collections of certain botanists could

also be that of their native collectors.

In the Precursores for Ganua by J. van den Assem in Blumea

7, 2 (1953) 380 is stated:—“‘Of some specimens from British

North Borneo (A postal 22, Wood 1261, 1889) and from Sarawak

({Garaman 2311, 2789), annotations by Lam were found in the

Rijksherbarium. Unfortunately, however, the specimens themselves

could not be traced anymore, which is the more regrettable since

they would mean new localities. Merrill... etc.”

Garaman was a collector of Haviland and the Singapore speci-

men labelled Haviland 2311 was sent to Leiden for revision and

determined personally by van Royen as Payena havilandii King

and Gamble. P. havilandii is a synonym of P. obscura Burck, but

it is certainly not that species. It is probably quite correctly deter-

mined as Ganua monticola by Lam. It is very similar to a sterile

specimen, Anderson SAR 2683 wrongly named Payena lucida by

van Royen and quoted by van Bruggen. Both have the under-sur-

face of the leaves very pale when dry. The locality of the Haviland

or Garaman specimen 23/1] now required is Sk near Kuching,

Sarawak, 10th June 1893. Tree. Pedicels pink, calyx yellowish.

V. Some Notes on Payena species

When trying to solve the identity of the Tanjong Gul species

mentioned previously I had to look through the other Payena

species in the Singapore Herbarium. I sometimes found that the

species in certain covers did not always form a uniform series.

I extracted those which appeared wrongly named and after a little

manipulation and rearrangement got a clear picture. The changes

are only minor ones but I feel they ought to be recorded.

1. Payena acuminata (Blume) Pierre in Bull. Mens. Soc. Linn.

Paris (1885) 528; van Bruggen in Blumea 9, 1 (1958) 100 f. 2.

Var. acuminata

MALAY PENINSULA Pananc: Sungei Mai Estates, Kadim &

Mahmood Nos. 15 (K, L, SING)

and 32 (A, BO, K, L, SING).

This species has not been recorded previously from Pahang

so the above collections appear to be the first records. They were

collected in March 1959 after the revision of Payena appeared

in Blumea.

222 Gardens’ Bulletin, Singapore — X XII (1967)

I must point out that a sheet Corner S.F.N. 30276 from Kema-

man, Trengganu 1s identical with Moysey & Kiah S.F.N. 33749

from Ulu Brang, Trengganu. One can see at a glance that they are

the same, yet the first has been incorrectly identified and quoted

by van Bruggen as Payena lucida when it should have been P.

acuminata var. acuminata.

2. Payena lucida (G. Don) A. DC. Prodr. 8 (1844) 197; van

Bruggen in Blumea 9, 1 (1958) 111.

I have added to our collections of Payena lucida another sheet,

Derry 937 from Malacca, wrongly named and quoted as Madhuca

sericea by van Royen. I spotted it as out of place among the

specimens of M. sericea by its very obtuse, almost orbicular

sepals in contrast to the sharply acute ones of this Madhuca. It

was in young flower-bud with as yet very short pedicels, but in

P. lucida these seem to lengthen rapidly as the flowers mature.

There are many specimens of this very distinct species in Herb.

Sing. and they form a very uniform series throughout the covers

marked ‘‘Malay Peninsula” in our collection. I am afraid I cannot

say this about the specimens named lucida from the Malay

Islands, especially Borneo. In fact I do not see any specimens

either from Borneo or Sumatra in our collection which I can

identify with lucida and it may be that lucida is not found in

Borneo at all. I have sorted out a series from Borneo which looks

more or less uniform but differs from Jucida in the finer and more

closely spaced nerves and in the line of interarching of the nerves

much nearer to the margin of the leaf than in that species. I discuss

this series here below under 3. Payena sp. but I do not know for

certain to which species it belongs.

3. Payena sp.

BRUNEI: Ulu Ropan, Belalong watershed, Ashton

BRUN 5245 (SING).

WEST BORNEO: Melawi Tyjatit, B. Tengkujung, 5bb26347

(SING).

EAST AND NORTH-EAST

BORNEO: Peak of Balikpapan, Beoul, Kostermans

7331 (SING); Loa Djanan, west of

Samarinda, Kostermans 6747 (SING); Loa

Haur, west of Samarinda, Kostermans

Nos. 6941 (SING) and 9899 (SING);

Pleihari, S. Alang, bb14197 (SING);

Pasir S. Ongka, bb25638 (SING); Bulun-

gan, bb26158 (SING).

P. NUNUKAN: Southern part, Kostermans 9204 (SING).

SABAH: Tawau Residency:—Merotai Besar, Tawau,

Aban Gibot SAN 31292 (SING).

Sandakan Residencv:—Bettotan, Sanda-

kan, Puasa 4558 (SING).

Sinclair — On Sapotaceae 223

West Coast Residency:—Kundasan, 10 miles.

west of Ranau, Wood & Kapis SAN

16392 (SING); west of Tenompok Pass by

roadside, Smythies SAR 10951 (SING);

Tenompok, Clemens Nos. 27974; 28178;

28538 and 28688 (all SING); Dallas,

Clemens 26715 (SING).

Interior Residency:—Kaingaran, 10 miles

from Tambunan, Wood & Charington SAN

17028 (SING); Bukit Tenom, Wood &

Wyatt-Smith A4392 (SING).

The above are the specimens I have sorted out as probably

consisting of one species which is not Payena lucida. One of them,

bb14197, is named Payena endertii by van Bruggen and a fruiting

twig of it is figured in Blumea 9, 1 (1958) 125. The fruits.

are glabrous, black, very hard and ovoid. Most of our specimens

are in flower. The only other one in fruit is Aban Gibot SAN 31292

(immature fruit). Lam in 1925 described P. endertii from Sumatra

with two syntypes, For. Bur. Lab. E1051 and For. Bur. Lab. T.B.

452 but in 1/927 he included some Bornean specimens in this

species. Consequently van Bruggen has chosen E/J05/ from Sumatra

as the lectotype. The only Sumatran specimen that we have here

is one of the syntypes 7.B. 452 and it is in flower. From it I am

not entirely convinced that it is the same species as bb/4197 and

the rest of the Bornean specimens. Unfortunately there is no other

authentic material of P. endertii known from Sumatra except

these two syntypes and they are not in fruit. That means we do

not know what the fruit of the Sumatran specimens is really like.

The specimen of 7.B. 452 is very like P. lowiana Pierre. The type

of P. lowiana came from Perak in the Malay Peninsula but this

species is also distributed in Sumatra and Borneo. Its fruits are

glabrous, hard and black but more elongate than those of 14197.

Young fruits, however, would look more like those of bb/4197.

Could the Sumatran material of P. endertii be P. lowiana? The

Bornean specimens which IJ have sorted out are very like Payena

lanceolata Ridley var. lanceolata except for their ovoid fruits and

smaller flowers. If the Bornean species is not lanceolata and not

endertii, could it be an undescribed species? This is a matter for

the authors of the Precursores and perhaps they may wish to

re-examine the material.

Of the above sorted material the following in my opinion are

wrongly named P. lucida by van Bruggen: — 6626/58; Clemens

Nos. 27974 and 28688; Kostermans 6941 and Wood & Wyatt-

Smith A4392. The following are wrongly named P. leerii:—

Kostermans Nos. 6747 and 7331 by van Royen and bb26347 by

van Bruggen. Clemens 26715 is wrongly named obscura by van

Bruggen and cannot be that because the veins of the leaf are

almost at right angles to the midrib. The following were returned

without identification slips: — bb25638; Clemens Nos. 28178 end

28538 and Puasa 4558. Some of the Clemens’ and Kostermans’

numbers had already been determined as P. endertii before they

224 Gardens’ Bulletin, Singapore — XXII (1967)

were sent to Leiden either by their collectors or by those who

prepared the typed labels as P. endertii is part of the typed script

there. The remainder of the specimens quoted by me were not

sent to Leiden since they were only acquired after the publication

of the Precursores.

4. Payena lowiana Pierre in Bull. Mens. Soc. Linn. Paris (1885)

525; van Bruggen in Blumea 9, 1 (1958) 126.

BRUNEI : Kuala Belalong, Temburong, Ashton BRUN

462 (SING).

EAST AND NORTH-EAST

BORNEO: Central Kutei, Belajan River, near Long

Bleh, Kostermans 10222 (SING).

SABAH: Mostyn Estate, Lahad Datu, Wood & Wyatt-

Smith A4299 (SING).

This species has not been recorded previously from either

Brunei or Sabah.

The flowers of 7.B. 452, one of the syntypes of P. endertii from

Sumatra, are very similar to those of P. lowiana. Both have rusty

obtuse sepals. Once more, as has been suggested above, could the

Sumatran material of P. endertii be the same as P. lowiana?

5. Payena lanceolata Ridley in J. Roy. As. Soc. Str. Br. 79 (1918)

93; van Bruggen in Blumea 9, 1 (1958) 128.

var. lanceolata

A fruiting specimen from Sumatra without locality collected by

Burck s.n. is this species rather than P. leerii as quoted in Blumea

l.c. page 123. The leaves are too narrow for leerii and the fruit

agrees with that of lanceolata.

6. Payena leerii (T. & B.) Kurz in J. As. Soc. Bengal 40, 2 (1871)

69; van Bruggen in Blumea 9, 1 (1958) 121 f. 8.

Basionym: Azoala leerii T. & B. in Nat. Tijdschr. Ned. Ind. 6

(1854) 116.

Synonyms: Isonandra microphylla de Vriese in Nat. Tijdschr.

Ned. Ind. 21 (1860) 312 —— syn. nov. Payena microphylla (De

Vriese) Pierre in Bull. Mens. Soc. Linn. Paris (1885) 531; van

Bruggen in Blumea 9, 1 (1958) 120 f£.7 —— syn. nov.

When I remove the following Singapore Island specimens named

and quoted as Payena obscura by van Bruggen from their genus

cover and place them with P. leerii I then get a very uniform selec-

tion of the two species and can see at a glance most of their differ-

ences: —Negadiman S.F. Nos. 35908; 36431 and 36456; Ridley 9203

all from Bukit Timah Nature Reserve and Ridley 4957 from Bajau.

The specimens of leerii from Singapore Island are correctly named

8 ee eee As 8 Oe | Re ee ae ee eh

Sinclair — On Sapotaceae 225

and it was noticed that they are all from Bukit Timah Nature

Reserve. In fact all the specimens from Bukit Timah are leerii and

not obscura. The latter is not known from Singapore Island, yet

both species are found in other parts of the Peninsula. It did seem

rather strange having specimens named leerii and obscura which

looked alike and yet both coming from a small area like Bukit

Timah Nature Reserve. It seemed equally strange when Ngadiman

S.F.N. 35800 is quoted twice in the Precursores, once on page 118

as obscura and again on page 123 as leerii. In the index of collec-

tors’ numbers it is given correctly as leerii.

Payena obscura has larger and more coriaceous leaves than leerit.

Its flowers and fruits are larger also, the sepals being more pubes-

cent and of a rusty colour. Juvenile leaves of leerii may be as

large as those of obscura but they are not so thick and leathery.

This may be seen in a specimen Price s.n. from Pulau Rimau,

Sumatra. Another very similar sterile specimen with slightly larger

leaves, Daud 10857 from Pulau Lavau, Sumatra is cited as lucida

but I think it should be leerii.

Looking through the covers of leerii one at first gets the impres-

sion that the flowers are slightly variable in size but this is actually

due to their age more than anything else. This is well illustrated by

looking at Smythies SAR 7804 from Berakas Forest Reserve,

Brunei. The material has quite young flowers. They have increased

considerably in very similar material, Anderson SAR 2163 from

the same locality. Here most of the flowers have the developing

Ovaries now visible, protruding some distance beyond the sepals,

but these flowers are still not so large as those of obscura. Berakas

Forest Reserve covers a very small area. The Anderson material

from Berakas is named P. microphylla. There is another sheet of

microphylla, bb28096 from East Borneo in the Singapore her-

barium. Other specimens exactly like these two are named leerii.

I can see no difference between microphylla and leerii and even van

Bruggen himself states that the two are nearly related. I have ac-

cordingly made a formal reduction above.

7. Payena pseudoterminalis H. J. Lam in Bull. Jard. Bot. Btzg

7, 1 and 2 (1925) 260 et l.c. 8, 4 (1927) 439 f. 17; van Bruggen in

Blumea 9, 1 (1958) 119.

A.C. van Bruggen states on page 119 of his Precursores that

Payena pseudoterminalis is a close relative of P. leerii, a species

which shows almost the same type of inflorescences; generally the

tertiary nerves of the leaf are much fainter or completely invisible

in the first-mentioned species. But under P. leerii in the key on

page 98, he says “tertiary nerves of leaves faint or hardly visible

below’.

226 Gardens’ Bulletin, Singapore — X XII (1967)

There is an isotype of P. pseudoterminalis, namely bb7190 from

Sumatra in the Singapore herbarium. This specimen is large enough

but has been badly dried without pressure; its leaves are folded

double and not one of them has been laid out flat. The Leiden

holotype seems to have been more carefully prepared. At least

one gets this impression from Lam’s drawing, figure 17.

Although P. pseudoterminalis is close to leerii its leaves are

much narrower and have a fine pubescence. In fact they are of the

same shape as those of lanceolata. Could this species be synony-

mous with lanceolata? I am not able to prove this for lack of

better material.

There are three sheets from Indragiri, Sumatra which are

obviously similar, the leaves having dried black. The flowers are

in a young stage, all rusty-tomentulose. In fact these three are

not different from two other sheets also from Indragiri with

sterile specimens, Curtis 3631 and Curtis s.n. which have been, in

my opinion, correctly named leerii. Yet the first three sheets have

been named differently. Two of them, 5527452 and bb27500

are quoted as obscura and the other Buwalda 6430 as pseudotermin-

alis. They are not pseudoterminalis as they do not have narrow

elliptic leaves. I do not understand how these specimens, all from

the same locality and so similar, should bear three different

identifications.

8. Payena obscura Burck in Ann. Jard. Bot. Btzg 5 (1886) 60;

van Bruggen in Blumea 9, 1 (1958) 117.

Synonyms: P. havilandii King & Gamble, Mat. Fl. Mal. Pen.

17, J. As. Soc. Bengal 74, 2 (1960) Extra Nr. 169. P. longipedi-

cellata Brace ex King & Gamble, Mat. Fl. Mal. Pen. Lc.

(1906) 169; van Bruggen in Blumea 9, 1 (1958) 115 f.6 ——

syn. nov.

Haviland 3035 figured on page 116 of Blumea 9, 1 (1958) as

Payena longipedicellata is one of two syntypes of that species.

The other syntype King 2940 is now the lectotype. The Singapore

duplicate of Haviland 3035 is not really different from P. havilandii

(of which there are five sheets in Singapore, these five making up

two syntypes) except that its pedicels are longer. Its flowers are

older than those of havilandii and will naturally tend to have

longer pedicels just as in lucida where the pedicels lengthen con-

siderably as the flowers mature. The leaves of Haviland 3035 are

not different from those of havilandii or from the other specimens

of obscura from Penang and Sarawak. Payena longipedicellata

should never have been created and I have no hesitation in reducing

it to obscura.

I must point out that Clemens 26329 from Dallas named lucida

by van Bruggen is obscura. This specimen has pedicels about the

same length as those of Haviland 3035.

Sinclair — On Sapotaceae 227

POSTSCRIPT

(Written April 1966)

A postscript to this paper is necessary to explain why I have

made no reference to Dr. Charles Baehni’s ‘“‘Mémoires sur les

Sapotacées, III Inventaire des genres’, Boissiera, vol. 2 (1965).

This contribution had not yet appeared when I wrote my notes in

July 1965. In fact it was not received in Singapore until 19th

March 1966 and I did not see it before mid-April. My notes were

sent to the Editor of the Gardens’ Bulletin Singapore in July 1965

but are still with the press at the time of writing this postscript.

Their aim as pointed out in the first paragraph of the text was to

draw attention to some minor details in the Precursores on

Sapotaceae, stating where I did not agree with the authors on

Payena and to find a name for a member of the same family col-

lected at Tanjong Gul, Singapore. There was no intention there

to criticise Baehni’s work, then non-existent or at least not avail-

able to the public. This aim still stands for I do not intend making

any alterations to what I have already written, not-with-standing

the profound changes that Baehni has made in the system of

classification of the Sapotaceae. I must, however, point out that

1 cannot agree with several changes that he proposes and I am

inclined at present to maintain Payena as a distinct genus. He jas

included it in Madhuca.

I laid out a series of specimens from the genera he has included in

Madhuca and another of the plants he calls /sonandra in order to

test them for uniformity.. Many of the Madhuca species in the

sense of the Precursores and of the authors of Sapotaceae for Flora

Malesiana now go into his /sonandra. These two lots which I laid

out are, alas, far from uniform and I can never agree that they

each represent one genus. Each bundle is most mixed! It is

quite ridiculous putting Palaquium obovatum in Isonandra and

Palaquium obtusifolium with very similar leaves and venation

in Madhuca. Payena leeriét and P. lancifolia are so very similar;

in fact I have difficulty in separating them when sterile. They both

have a long narrow scar on the seed extending from one end to the

other of its length. It is true that his drawings of these two species

also show similar scars, but he places P. leerii in Madhuca and

P. lancifolia in Isonandra. He says in the key that the scar is narrow

and long in Madhuca and narrow and short in Jsonandra, but here

as pointed out above both of these Payena species have the scars

long and narrow. Most of his so-called /sonandra species are

illustrated with short scars and on page 189 fig. 137 of his publica-

tion he depicts the well-known Palaquium obovatum (his Isonandra

228 Gardens’ Bulletin, Singapore — XXII (1967)

obovata) with a short obovate scar extending three quarters way

along the length of the seed. I checked on this species which I

pass every day and collected from the tree 22 seeds. I found that

every one of these has the scar extending along the entire length of

the seed almost from end to end and in no case did the scar ter-

minate three quarters way up as shown by Baehni.

It seems to me that he has laid too much stress on this character

of the seed for distinguishing genera and has been carried away with

it. Leaf characters are not used at all. The authors of Flora

Malesiana Precursores often use leaf characters for the separation

of species and have not neglected them for identifying genera. I

feel that in this family much more attention should be paid to leaf

characters.

Most of the Payena species mentioned here by me have rather

similar leaves and appear to be a uniform lot. Madhuca sensu van

Royen is a large genus and certain species may have to be removed

from it. But the great majority of them form a very uniform series

all with a distinct type of leaf which has a reddish tinge, the midrib

glossy and smooth, the veins very distinct beneath and the twigs

pale and angled towards the apex. Some of the species are so close

that I have difficulty in distinguishing them. If they are so close

surely then they must all belong to the same genus whatever name

future systematists may give it. Most of these species belong to van

Royen’s groups 2 and 3 on pages 4-6 of the Precursores on

Madhuca, thus: —M. aristulata, burckiana, cuneata, elmeri, gla-

brescens, korthalsii, laurifolia, longistyla, malaccensis, mindanaen-

sis, montana, ovata, penangiana, pedicellata, pubicalyx, sepilokensis,

sericea, spectabilis, woodii and others which are not in the Singa-

pore herbarium and which I have not yet seen.

In the genus Ganua there is a group of species with fine, close,

slender veins in the leaves. To this lot belong G. curtisii, the type of

the genus, G. motleyana and several others. In another group the

leaves have long petioles, the veins very prominent on the lower

surface, there raised and arising at a wide angle, (often 90°) with

bold sweeping curves, and the reticulations beneath equally pro-

minent. Here belong G. hirtiflora, kingiana and prolixa etc., but

there are similar species in a few other genera. I do think that

some of them should be removed and placed in this group because

of their rather different vegetative characters. Some of these species

were also formerly placed in a genus Dasyaulus. They should be

studied more carefully in relation to each other.

A Note on Myriophylium

By JAMES SINCLAIR

Botanic Gardens, Singapore.

On looking through the material of Limnophila in the Singapore

Herbarium | came across several sheets of a plant from Kedah and

Perlis wrongly named Limnophila heterophylla. These actually

belong to a Myriophyllum which I have identified as M. tubercula-

tum Roxb. The numbers concerned are Corner S.F.N. 37985; Curtis

2102 and Henderson S. F. Nos. 22907 and 22908. The two Curtis

sheets had originally been named Myriophyllum intermédium but

this name is deleted by C. X. Furtado and Limnophila heterophylla

substituted. The rest of the material was named by M. R.

Henderson.

Under the genus Myriophyllum itself there is not a lot of

material in this herbarium. In fact the only record from the Malay

Peninsula is a sheet, J. A. Baker s.n. 30th October 1939 from

near Alor Star, Kedah. This is named M. tuberculatum and is

stated, on the label, to be the first definite record for the Malay

Peninsula. The Curtis material, however, was collected in Novem-

ber 1889, the two Henderson sheets 18th November 1929 and

Corner’s on 14th November 1941.

Ridley in Fl. Mal. Pen. 1 (1922) 692 states ‘““Myriophyllumi inter-

medium DC. Malacca, Griffith, probably on Mt. Ophir. No

specimen occurs in Griffith’s collections at Kew, nor is it mentioned

in the F.B.. Mt. Ophir is a most improbable locality for this

pondweed”’.

M. R. Henderson in Gard. Bull. Sing. 7, 2. (1933) 103 in a paper

entitled Additions to the Flora of the Malay Peninsula has

*“Myriophyllum sp. Johore: Bukit Tiga, Sungai Sedili, Corner sine

num. Common by edge of river from Kuala Bohol to a short way

below Bukit Tiga. Does not grow in the salt water of the mangrove

reaches. Malay name: Rumput Ekor Kuching. Unfortunately only

the submersed stems and leaves, without flowers or fruit have so far

been found, so that it is impossible to say to what species it may

belong. This is the first definite record of the family from the

Malay Peninsula’.

At first I could not find the above specimen. It was not filed

under Myriophyllum so I came to the conclusion that it had been

removed from that genus and put elsewhere by someone who

disagreed with the identification. A search was made under other

likely genera and I soon located it also under Limnophila. It had

been put there by Corner who had later deleted the word Myriophyl-

lum and added Limnophila ?sessiliflora. These alterations are in

ink of a different and fresher shade than that of his original hand-

writing. The specimen was collected in March 1932. It is certainly a

Limnophila and not a Myriophyllum, and unfortunately, as pointed

out by Henderson, is sterile. The bushy tassels of leaves resembling

a cat’s tail (ekor kuching) are covered with, and somewhat hidden

230 Gardens’ Bulletin, Singapore — XXII (1967)

by an epiphytic green alga which is present. There are two other

specimens of Limnophila like it but these are also sterile and

unnamed at the time of writing. One is from Kedah and the other

from Kota Tinggi. These may or may not be sessiliflora. I am

unable to identify them because of my lack of knowledge about

Limnophila in the vegetative condition. I have no doubt that they

could be identified by a person with experience in cultivating them.

They could easily be grown in tanks and this would make an

interesting study. Myriophyllum tuberculatum certainly resembles

Limnophila heterophylla and sessiliflora vegetatively and for this

reason there has been confusion over them in the Singapore Her-

barium. Some of the more important references in the literature are

now given for M. tuberculatum with the citation of specimens in the

Singapore Herbarium.

Myriophyllum tuberculatum Roxb. Hort. Beng. (1814) 12

nomen nudum; FI. Ind. 1 (1820) 471; 1 (1832) 451 and (1874)

151; A.DC. Prodr. 3 (1828) 69; Miq. Fl. Ind. Bat. 1 (1) 4 (1855)

635; C.B. Cl. in Hk. f. Fl. Br. Ind. 2 (1878) 432; Peterson in

Engl. & Prantl, Pflfam. 3, 7 (1893) 234 f£.104 A-C; Schindler

in Engl. Pflreich 23 (4, 225) (1905) 96 f£.28 A-C; Sinclair in

Bull. Bot. Soc. Beng. 9, 2 (1955) 94.

MALAY PENINSULA PERLIS: Kangar, Henderson S.F. Nos. 22907

(SING) and 22908 (SING).

KEDAH: Kuah, Langkawi, Corner S.F.N.

37985 (SING); Langkawi in paddy

fields, Curtis 2102 (SING); near

Alor Star, J.A. Baker s.n. 30th

October 1939 (SING).

MOLUCCAS HALMAHEIRA: Pajahi Road, District Weda, G.A.L.

de Haan 1788 (A, BO, K, L, P,

SING) only the SING specimen

seen by me.

Most of the species in this genus are confined to Australia. Some

are from New Zealand, Europe, North and South America, India

and the Mascarene Islands, while M. spicatum is of world-wide

distribution. M. tuberculatum is an East Bengal species and I have

collected it at Cox’s Bazar. I am surprised that it has not, so far,

been recorded from Siam where siamense and tetrandrum grow.

Neither has it been seen in Indo-China where there are five species,

namely bonii, intermedium, siamense, spicatum and tetrandrum.

See M.-L. Tardieu-Blot in Flore du Camboge du Laos et du

Vietnam 4 (1965) 124. M. tuberculatum is nearest to tetrandrum

but has larger, minutely serrate, ovate bracteoles and a larger

fruit which is tuberculate and not rugose-granulate. M. tetrandrum

was first recorded in Malesia from Kendari, Celebes where it was

collected by Beccari. See van Steenis in Webbia 8 (1952) 435. In

Java, according to Backer and Bakhuizen f., Flora Java, 1 (1963)

266, the species are M. brasiliense (naturalized and locally abun-

dant) and M. dicoccum (Lake Burnih, Madura). The latter was

originally described from North Australia. In New Guinea there

are probably several species. There is one resembling verticillatum

from the Wissel Lake region, represented in the Singapore

Herbarium by Eyma 4733.

Studies of Maiesian Pandanaceae, I

Polymorphism in Pandanus odoratissimus

L.f. of Asia

BENJAMIN C. STONE

School of Biological Sciences, Botany Unit, University of Malaya

In Ridley’s Flora of the Malay Peninsula, vol. 5, (1925) the

common pandan of beaches and coasts of Malaya is described

under the name of Pandanus fascicularis Lamarck, and placed in

the section Keura. That section, since it includes the type species.

of the genus, must now be called section Pandanus, and P.

fascicularis is considered a synonym of P. odoratissimus Lf...

which is the type species of the genus. P. odoratissimus has had

a history of considerable confusion. It has an acknowledged list

of many synonyms, but it has also been frequently misapplied.

‘to quite different species. It has at one time or another been

considered as equivalent to P. tectorius, with now one and now

the other name considered as a synonym. It has been used as a

kind of receptacle for undeterminable collections and for sterile

cultivars. The result of this historical confusion has been the

obscuring of the real specific characters and the true distribution

of the species.

Recently a considerable number of species have been described

from Malaya and adjacent or nearby areas in South-east Asia

by Dr. H. St. John in Pacific Science (1959-65). In attempting

to write up the Malayan Pandanaceae, it of course became

necessary to undertake a study of these species. To do so, not

only have the holotype specimens in the Herbarium of the Botanic

Gardens, Singapore, been examined, but travel to some of the

type localities was made to study the living plants. Some types

were Seen at the Arnold Arboretum. In some cases I have not

seen the types, but the descriptions are so full, and the illustrations

so good, that little doubt as to the character of the plant was left.

The conclusion I have reached is that only one species of

section Pandanus is wild in Malaya, but that it is represented by

several forms, most of which are so minor in distinctiveness, and

so clearly united by transitional forms, that it is impossible to

regard them as separate species. Since this view appears to be so

greatly at odds with the treatment in Pacific Science, it has seemed

necessary, as well as valuable, to discuss the situation at length,

and especially to set forth all the reasons available for the

reduction of so many species to Pandanus odoratissimus L..

Taxonomists often list new synonyms with a remark or two such

as “‘indistinguishable”’ or “differing onlv in the somewhat larger

232 Gardens’ Bulletin, Singapore — XXII (1967)

leaves”, and not rarely give no remark at all, assuming evidently

that none is required. On the contrary, it seems to me that listing

a mame as a new synonym requires explanation. Therefore I

present in this paper the data which have convinced me of the

need to bring the indicated species to synonymy.

Important Taxonomic Characters. The characters which are used

in the discrimination of species in the whole genus are not all

of use in the present case. For example, the highly important

characters of leaf-tip shape and of armament of ventral pleats are of

no value among the forms considered here, since they are identical

(ali leaves have long, narrowly attenuate, flagellate tips, and un-

armed ventral pleats). The habit and habitat characters differ

slightly, but far less than in other sections of the genus. The

presence of a racemose or spicate female inflorescence likewise

is of great importance, but in all our forms the cephalium is

solitary on the peduncle. The fruits are syncarpous, with phalanges

of several fused carpels, and none of the forms considered here

has simple drupes. The stigmatic structure is of the same type

throughout in our forms, as is evidenced by their inclusion in

section Pandanus.

The remaining characters, which have been considered import-

ant in the original diagnoses of the species and in my own

experience are as follows: (1) shape and dimensions of the

cephalium; (2) length of the peduncle; (3) number and range, and

median number (major carpel number) of carpels per phalange;

(4) dimensions of phalanges; (5) presence or absence of fleshy

mesocarp shoulders around middle of phalange; (6) color of pha-

langes; (7) presence or absence of lateral sutures in phalanges;

(8) depth of apical sinuses between carpel apices; (9) position of

endocarp within the phalange; (10) dimensions of the leaves;

(11) number of veins in leaf (as counted near leaf midpoint);

(12) color, size, and spacing of leaf prickles; (13) direction of

midrib prickles.

The following remarks on the above characters precede a com-

parison of them in the species concerned.

1. Shape and dimensions of the cephalium. It should be super-

flous to note that measurements of the character are useful only

if full-grown cephalia are available. In addition it must be noted

that (a) fruit-size decreases somewhat as the plants grow older;

very old trees often produce cephalia as much as 40 per cent

smaller than young, vigorous plants. Furthermore, adverse environ-

mental conditions produce a size reduction; (6) the distinction

between globose, subglobose, and ellipsoid cephalia is a very lax

one; smaller cephalia are often concomitantly more globose.

2. Length of peduncle. This character appears to vary in much

the same manner as cephalium size.

3. Carpel number. This is an important character, which I

have discussed in another paper (1967). Briefly, it has been found

that by plotting data from one cephalium in graphic form, the

Stone — On Pandanaceae 233

carpels-per-phalange number on the horizontal axis and the number

of phalanges on the vertical axis, a curve is obtained which typi-

cally begins at a relatively small carpel number, quickly rises to

a peak, then more gradually descends to a relatively large number

(Figure 5). Thus, of a cephalium bearing sixty phalanges, it may

be found that three phalanges were composed of four carpels; 23

phalanges were composed of five carpels; 20 phalanges of six

carpels; 8 phalanges of seven carpels; and 6 phalanges of eight

carpels each. In this case, the “‘range’’ of carpel number would

be 4-8, and the ‘major carpel number’ would be 5. Ostensibly,

cephalia showing different major carpel numbers are different in

a taxonomic sense; but, as pointed out in the paper already

referred to, the taxonomic function of this difference is often over-

estimated.

4. Dimensions of the phalanges. As pointed out above in

sect. 1, size is partly a function of age and of environmental

conditions, as well as of hereditary causes. Especially important

is the dictum that all measurements be taken on ripe, full-grown

material. For the sake of comparibility, phalanges from the same

part of the cephalium should be utilized; basal (peduncular

region) phalanges are invariably shorter than average, as well as

being curved, and have a greater than normal number of carpels.

5. Shoulders. The pulpy, firm expansion of the midregion of

the phalange is conveniently termed shoulders. These are often

eaten by insects on fruits still on the tree, and rather quickly

“disappear” on dried or weathered phalanges. In my experience,

the presence of shoulders on phalanges is a major character of

Pandanus odoratissimus, and is correlated with other major cha-

racters such as the white color of the leaf prickles, the twin

glaucous strips on the underside of the leaves, etc. Shoulders are

either absent, or of a different shape and color, or are not corre-

lated with other characters, in similar but distinct species.

6. Phalange color. Clearly, a good color terminology is required

here; also it is important that fruits be fully ripe. In our species,

the entire phalange becomes some shade of red-orange; but the

last part to turn this color is the summit (carpel apices) of the

phalange. The shoulders are often a deeper red than the rest of

the phalange; and the extreme base may be yellowish,

7. Lateral sutures. By this is meant deep divisions between

the carpels, visible in side view. Generally such sutures are absent

in Our species, though an occasional one or two occur. Phalanges

with marked, deep, and regular sutures between carpels could

indicate a different species.

8. Depth of apical sinuses between carpel apices. This character

is a fairly important one, as it contributes largely to the overall

impression one obtains when seeing a cephalium, or even loose

phalanges, for the first time. Furthermore, it often is of specific

importance. However, unless the depths concerned differ by more

than just a few millimeters, they do not seem to merit overmuch

234 Gardens’ Bulletin, Singapore — XXII (1967)

attention at the species level of distinctiveness. It must be pointed

out that phalanges from the peduncular end of the cephalium will

generally have shallower sinuses than phalanges from the midre-

gion or apex of the same cephalium.

9. Endocarp position. This too can be a significant character,

as for example in discriminating such species as Pandanus dubius

and P. compressus, or in the circumscription of P. basilocularis.

Nevertheless, different positions may be attained in different ways;

in some cases, extra growth of the apex of the phalange; in other

‘cases, extra growth of the basal fibers; and both of these types of

cases may be found to occur on one and the same plant at

different times. More important for our discussion is the fact that

since growth of the phalange continues at the base until the full

length of the phalange is attained, an immature phalange will

generally have an apparently sub-basal endocarp.

10. Leaf dimensions. These vary from several causes. Pandans

have several categories of leaves: (a) prophylls and preliminary

foliar bracts; (6) normal foliage leaves; (c) transitional leaves

merging with inflorescence bracts; (d) seedling leaves; (e) juvenile

leaves. It is generally useless to compare an organ of type (a) from

specimen 1 with an organ of any other type from plant 2.

Obviously, comparable structures must be compared. Besides this,

older plants produce smaller leaves than younger plants. In

general it may be stated that normal foliage leaves of unbranched

juveniles may be up to four or five times as long, and half again to

three times as wide, as leaves of adult plants.

11. Number of main longitudinal veins per leaf. The same

cautions mentioned in sect. 10 should be heeded. Naturally,

smaller leaves will have fewer longitudinal veins, even on the same

individual.

12. Prickles. The underlying features of the prickles of the

species under consideration is their white or very pale color. This

character at once distinguishes these plants in the field. It is in

marked distinction to the dark or merely pale green prickles of

many other species, including the ones of the Eastern Pacific often

lumped under the name P. tectorius. These white prickles are also

relatively long, i.e. on the leaf bases of mature leaves they may

exceed 6-7 mm. in length. Their spacing may be expressed as a

range of distances between successive prickles at given positions

(e.g., marginal prickles near leaf-base 6-12 mm. apart); or in terms

of how many occur per unit (e.g., marginal prickles near base

3-4 per 10 cm.).

13. Direction of midrib prickles. In our species, the midrib

prickles of the basal portion of the leaf are always retrorsely

curved. This quickly distinguishes these plants from some related

species and from some unrelated species (as well as from

Cyperaceae).

Stone — On Pandanaceae 235

Before proceeding with a comparative analysis of species, it is

necessary to fix the character of Pandanus odoratissimus. This is

no easy matter, since the description is very brief, and worst of

all, it is based on the male plant. The type locality is Ceylon, so

it may be expected that a collection made there of both male and

female plants would establish the identity of the species. The fruit

is needed, because nearly all subsequent descriptions emphasise the

fruits. Luckily this has been done by Dr. St. John and we are

therefore in a position to make the comparisons needed with

Ceylon material.

Comparative analysis

The essential characters are set out in Table 1. In summary,.

it may be stated:

(1) Size range for cephalia is from 8.8 x 8.8 cm. in P. globosus

and 9 x 8.5 cm. in P. rubricoloratus to 23 x 20 cm. in P. hueensis.

Average size appears then to be about 16.8 x 14 cm. The only

gap seems to be for P. hueensis and P. vietnamensis.

(2) Peduncular length varies from 9 to 40 cm., with the average:

length near 25 cm.

(3) Carpel number. Since the original data is not expressed in

the form outlined earlier, only ranges can be given. These rarely

cover more than 5 carpels difference between minimum and maxi-

mum in any one cephalium. The smallest number given is 4 (in 2

species); the largest number is 14 (in 2 species). The general

range is thus 4-14, with 5-12 being far more common. A gap:

seems to exist between species with the “‘major carpel number” at

5, 6. or 7, and those at 9 or 11, but this seems uncorrelated with

any other significant feature.

(4) Phalange size varies more or less continuously between 3.5

and 7 cm. long and 1.6 and 4.6 cm. wide. Phalange color is always

some shade of red or red-orange, if fully ripe; if too young, yellow

or green.

(5) Number of phalanges per cephalium ranges from 26 to 143,

with figures between 60 and 90 apparently most common.

(6) None of these forms have regular, deep lateral sutures; most

have none; a few have occasional shallow sutures.

(7) All forms have fleshy shoulders except those which are

clearly immature. It is believed that these too would exhibit

shoulders at full ripeness.

(8) Leaves range in length from 0.8 to 2.4 (or 3?) meters, and’

in width from 2.8 to 6.3 cm. All have pale or white prickles; many

have dark or brownish tips on the prickles. The number of longi-

tudinal veins in each half varies from 21 to 45 (in the known

examples).

(9) Ecology. All forms are beach, coastal, or swamp species, near

the sea. Geographically, they range from the Indian Ocean through

S.E. Asia to Malesia.

236 Gardens’ Bulletin, Singapore — X XII (1967)

In conclusion, it appears from this analysis that 9 species des-

cribed from Malaya, one from Anamba Is., 10 from Vietnam, one

from Hongkong, and four from the Maldive Is., can all be con-

sidered synonymous. Several are considered to be at least tem-

porarily recognizable as forms. There follows a list of the synonyms

and discussion of the forms and species maintained.

Description of Pandanus odoratissimus.

Pandanus odoratissimus Linnaeus fil., Suppl. Pl. 64, 1781. St.

John, Taxon 12 (5): 201-204, 1963.

P. verus Rumph., Herb. Amb. 4: 139. t. 74. 1743; Kurz, J.

Bot. Brit. & For. 5: 125, 1867, in part.

Keura odorifera Forsskal, Fl. Aegypt.-Arab. 172, 1775;

Pandanus odorifera (Forssk.) Kuntze, Rev. Gen. 2: 737, 1891.

(Nomen dubium).

P. fascicularis Lamarck, Encycl. 1: 372, 1783.

P. leucanthus Hasskarl, F1. Bot. Ztg. 25 (2), Beibl. 14, 1842.

Hasskarlia leucacantha Walpers, Ann. 1: 753.

Note: this is by no means all the older synonyms, but a full

account of these is impossible until their exact identities have been

set forth, since earlier authors, including Warburg, confounded the

Indian-Malaysian species with the Polynesian-Melanesian species,

and Warburg gave to this very broadly conceived species the name

Pandanus tectorius Solander ex Parkinson, a name which is now

agreed not to have been validly published (see Airy Shaw, Taxon

11 (7): 223, 1962) and therefore dates only from Warburg’s publi-

cation in 1900. Neither his taxonomic nor his nomenclatural con-

clusions can be supported now; and the name P. odoratissimus L.,

though later than Keura odorifera Forsskal, will be the type of

the genus Pandanus, since it has been shown by St. John to be

better established. Forsskal’s name must remain dubious, as it

lacks a type specimen and was based only on part of a male

flower. The species of the Pacific are distinctive enough to merit

retention, though the number of binomials existing now is certainly

too large; but even if a very widespread specific concept is adopted,

it seems clear at this stage that P. odoratissimus is easily discri-

minated, both on vegetative and fruit characters, and its presently

known range does not extend east of Malesia (in the sense of van

Steenis in “‘Flora Malesiana’’). The boundary is not certainly

known but seems to be in the general vicinity of Wallace’s Line.

The Pacific species (or one of them) however appears to overlap

the above area, and occurs at least in eastern Malesia; it is this

occurrence which has helped to foster the confusion of the species.

‘The plant illustrated in Rumphius’ Tab. 81 seems to me to be an

example of such overlapping. What name, or names, may be used

for this plant and its Pacific relatives is a problem that can safely

be deferred for the present.

Stone — On Pandanaceae 237

NEW SYNONYMS: MALAYAN TYPE LOCALITIES:

. albibracteatus St. John, Pac. Sci. 17: 10. 1963.

. ambiglaucus St. John, ibid. 17: 13. 1963.

. carnosus St. John, ibid. 17: 20. 1963.

. globosus St. John, ibid. 17: 21. 1963.

. inclinatus St. John, ibid. 17: 24. 1963.

. incrassatus St. John, ibid. 17: 26. 1963.

. obtusus St. John, ibid. 17: 28. 1963.

. rubricoloratus St. John, ibid. 17: 32. 1963.

. Boryi Gaud. Voy. Bonite, t. 22, f. 15. 1841; em. St. John, thick

17: 18. 1963.

Cows 2 Ses Soe ae

ANAMBA ISLANDS:

P. Hendersonii St. John, ibid. 19: 116. 1965.

VIET NAM:

. Integriapicis St. John, ibid. 16: 91. 1962.

. Phamhoangii St. John, ibid. 16: 96. 1962.

. projectens St. John, ibid. 16: 100. 1962.

. reversispiralis St. John, ibid. 16: 101. 1962.

semiorbicularis St. John, ibid. 16: 106. 1962.

. Smitinandii St. John, ibid. 16: 109. 1962.

. subcarnosus St. John, ibid. 16: 111. 1962.

. Subulatus St. John, ibid. 16: 114. 1962.

. vietnamensis St. John, ibid. 16: 116. 1962.

a Bile- la-la-la-la ~~

HONG KONG:

P. remotus St. John, ibid. 16: 70. 1962.

MALDIVE ISLANDS:

. adduensis St. John, ibid. 15: 328. 1961.

. Hartmannii St. John, ibid. 15: 333. 1961.

. Karikayo St. John, ibid. 15: 335. 1961.

. maldivecus St. John, ibid. 15: 339. 1961.

a Bias SE, a

238 Gardens’ Bulletin, Singapore — XXII (1967)

DESCRIPTION: (Figs. 1-6).

Habit. Coarsely branched trees with a tendency to candela-

briform branching, the secondary branches sympodial by

development of a lateral vegetative bud at the base of the

terminal inflorescence; branching di- or _ trichotomous, or

irregular. Main stem rarely more than 20 cm. thick, erect or

more or less decumbent, usually thickest a little above the

extreme base; later branches of the same or lesser diameter

because of lack of cambium; leaves of three- to five-year-old

vigorous juveniles often four or five times longer and twice or

three times broader than leaves of senescent, much-branched

adult plants. Main stem and not rarely upper stem and lower

branches producing prop-roots up to 10 cm. thick, but usually

more slender. Adult plant seldom over 10 m. tall; suckering from

base occasional. Occurrence in groves not uncommon.

Stems green in growing stages towards apex, earliest parts

white, soon discoloring on exposure, but covered by leaves;

exposed part usually turned pale buff or gray-brown, grossly

ringed by leaf-scars, these as much as 3-4 cm. apart on rapidly-

grown parts, usually 1 cm. apart or a little less. Uppermost

ultimate branches as slender as 3 cm. diameter in old trees.

Base of branch at attachment to stem dorsoventrally flattened

and somewhat clasping. Stems and branches usually more or

less prickly because of blunt or acute adventitious roots with

arrested growth, up to about Icm. long.

Leaves spiralled, in three rows (tristichies), clasping at base

around the stem, the extreme edges just meeting there or slightly

overlapping. Leaf margins of the clasping portion gradually

thinner toward the edge, whitish or pale green. Blade ensiform,

nearly parallel-sided, toward the apex very gradually reduced

(in foliage leaves), more abruptly reduced in prophylls. Prophylls

deltoid (approaching isosceles form), entire, the first one channelled

on both sides, the remainder channelled ventrally, the series

merging gradually to foliage-leaf form, with the last few prophylls

(usually around the 9th) developing marginal pickles toward the

apex; early foliage-leaves short, rather bluntly acute, marginal

prickles developed; later ones with dorsal midrib prickles

developed, and apices increasingly prolonged; adult foliage-leaves

with the apex gradually narrowed into an elongated triquete

flagella, the margins and dorsal midrib armed with prickles.

Largest leaves as much as 3 m. long and 9 cm. broad, but

usually less, 1-2 m. long and 4-7 cm. broad; length : width ratio

commonly between 20:1 and 30:1. Blade M-shaped in section.

Apex of youngest leaves stiffly erect, but older leaves drooping

from near the tip or the midpoint or even lower. Dorsal surface

glaucous on each side of the midrib; ventral surface also glaucous,

but less so. Prickles white, or the extreme tip dark, commonly

3-5 mm. long, often longer; in some forms up to 10 mm. long.

slender, very sharp, very slightly curved; prickles of margins all

forwardly directed; prickles of midrib forwardly directed in the

Stone — On Pandanaceae 239

distal half (approximately) of the leaf, but the prickles of the

proximal part retrorse. Spacing of the prickles rather irregular,

but distally the prickles always smaller and more crowded,

proximally always larger and farther apart; commonly (near the

leaf base) the prickles 1-3 cm. apart. (Some forms probably for

genetic reasons unarmed). Parallel veins mostly 40-160, depending

on age of leaf, age of tree, etc., visible but not prominent along

most of the blade but less or not so toward the base. Transverse

lines not at all evident, unless under microscope. Colored

pigmentation absent, i.e. leaves never reddish, purplish, etc.;

white- or yellow-striped variegated forms may exist.

Female inflorescence borne only on female trees, terminal,

pedunculate, the peduncle 10-30 (rarely more) cm. long, trigonal,

bracteate, the lower bracts leaflike, upper bracts whitish-yellow,

successively shorter, more or less navicular, the midrib and

margins with reduced prickles, texture sub-fleshy, the last bracts

much reduced to linear-lanceolate shape and much shorter than

the syncarp; these bracts all soon withering and more or less

persistent but at last fragmenting and falling. Inflorescence a

cephalium, globose to ellipsoid, up to about 30X20 cm., slightly

rounded-trigonal, mostly 15-20 cm. x 12-18 cm.; composed of a

variable number (26-143, usually about 50-70) of carpellate

phalanges, these in turn composed of mostly 4-10 concentrically

arranged fused carpels (phalanges of only 1-3 carpels or of 11-14

carpels rarely also occur, the former only near the cephalium

apex, the latter only near the cephalium base). Carpels each

terminating in a single U- or V-shaped stigma; carpel apex

truncate, low-rounded,.or low-pyramidal, sometimes somewhat

acute. Phalanges laterally mostly smooth, rarely with 1 or 2

intercarpellary grooves; apically with slight grooves between the

carpel tips. At maturity the phalanges red-orange to vermilion,

sometimes the carpel tips still greenish; median part of phalange

rather abruptly expanded into fleshy shoulders, these not evident

in dried or insect-damaged or animal-gnawed specimens, but

always obvious in fresh fully ripe fruit. Phalanges mostly 3-8

cm. long, 2.5-4.5 cm. wide, pentagonal or hexagonal but some-

times slightly exceeding these measurements; phalanges from

cephalium base (peduncular end) always curved,. shorter, and

with more carpels, than those from lateral or apical part of

cephalium; phalanges from apex of cephalium usually longest

and with fewest carpels, the carpel tips more expanded and

sometimes more acute. Young phalanges greenish-white, later

yellow, finally orange then reddish to vermilion. Phalanges free

of each other but tightly crowded; eventually falling from the

cephalium receptacle, apical ones first, leaving a pentagonal or

hexagonal scar. Cephalium when mature more or less drooping;

when young usually suberect. Phalange externally covered by a

pericarp; within this the upper mesocarp is lacunose, each carpel

apex with a chamber traversed around the periphery by longitudinal

fibers and crossed by thin plates of pithy tissues (aereiichyma);

lower mesocarp fibrous, the fibers traversing a pulpy tissue;

240 Gardens’ Bulletin, Singapore — X XII (1967)

endocarp central, osseous, reddish-brown, unitary, each carpel

represented by one ovular chamber, this fusiform, a few fibers

traversing the endocarp, some central in the mesocarp and leading

to the stigma, these entering the ovular chamber along one side;

upper part of ovular chamber filled with soft white endosperm;

embryo basal; germinating downward. Outermost seeds often

not developing.

Male inflorescence only on male trees, pedunculate, bracteate,

the bracts similar in color and form to those of the female, but

the inflorescence a raceme of spikes (an odd number, usually

5, 7, 9, or 11), each in the axil of a bract, each consisting of

a fleshy central axis from which emerge numerous lateral phalanges

of stamens about 13 mm. long, bearing about 19-23 stamens on

short filaments 0.5—-2 mm. long, with anthers 2—3.5 mm. long,

narrowly sagittate, with an apiculum 0.5 mm. long; the basal

part of the phalange is about 3 mm. long up to the first filament.

The phalange, and the receptacle and major axis as well, are

all clearly seen to be fasciculate. The overall length of the whole

inflorescence varies from about 30 to 60 cm. long; the bracts

subtending the spikes are mostly less than 30 cm. long, the

last ones only about 6 cm. long. The inflorescence is ephemeral

and decays very rapidly after maturing, usually within a couple

of days.

Citation of specimens. No attempt is made here to cite all

specimens of Pandanus odoratissimus, since it has been collected

many, many times; and indeed many specimens are not worth

the paper they are mounted on, for lack of representative parts,

poor labelling, and so on. The specimens cited here are simply

those of the herbaria at the Singapore Botanic Gardens and at

the University of Malaya. The main purpose in citing these is

to mark precisely those which are in fact this species to distinguish

them from the very similar plants (often in villages) which have

recently been called P. spurius Miq., and those which represent

the Pacific-Melanesian group which has been called P. tectorius,

such as Lam 3285.

MALAYA. Singapore: Changi, Dec. 1892, Ridley (2). Bajau,

9 May 1892, J. S. G. (c‘). Passu Panjan, 1901, Ridley (¢).

Same locality, 1909, Ridley 14157 (c"). Kranji, mangrove swamps,

10 April 1966, Stone 6237, 6238 (2), KLU.— Johore: Tanjong

Bungah, 1894, Ridley 6284 (3). Tana Panti, 22 April 1890,

Ridley 4594 (co). Kukul, May 1909, Ridley 14169 (¢). Tana

Panti, 1892, Ridley (2). Jason Bay, 15 June 1934, Corner (@ ).

3 miles west of Kuala Sendai, 7 June 1922, Goodall (c‘). Johore

Bahru, west side mangrove area, 21 Nov. 1965, Stone 6106

(2, KLU). Same locality, 12 April 1966, Stone 6106-A (2°,

obs. ¢, KLU). Mersing, 6 Jan. 1959, Paine ( 2 ).— Malacca:

Near Malacca, 1892, Ridley (3). ‘‘Sea-shore’, 1 May 1895,

Maheiros (sterile). Negri Sembilan: Kuala Pilah, kampong

pandan, 26 Feb. 1936, Corner SFN 30762 (c‘). — Pahang:

Kuala Pahang, 17 May 1890, Ridley (3); 10 May 1890, Ridley

Stone — On Pandanaceae 241

(92). — Trengganu: Kuala Trengganu, 9 May 1925, Holttum

SFN 15169 (o). Cherating River, 25 Aug. 1884, Ridley (sterile).

— Perak: Dindings, Pangkor, March 1896, Ridley (co). Pulau

Lalang, 30 Dec. 1925, Sermund (o'). Langkawi: April 1892,

Curtis (co). Penang: Batu Ferringhi, Tanjong Bungah, 10 Feb.

1966, Stone 6171, (2); 2 May 1966, Stone 6350 (2) KLU.

{SING unless otherwise noted].

SUMATRA. Atjeh, Troemon, 27 Aug. 1941, Asdat 107. (@).

PULAU BANGKEI: 30 June 1936, Liitjeharms 5134. (9).

NORTH BORNEO: SABAH: Sibubu River, Mengalong Forest

Reserve, 5 miles S. W. of Sipitang, 30 July 1954, Wood & Wyatt-

Smith SAN ‘A’ 4567 (¢ ).

Some Forms of the species:

Pandanus odoratissimus L.f£. forma vietnamensis (St. John) B. C.

Stone, comb. et stat. nov.

P. vietnamensis St. John, Pac. Sci. 16: 116. f. 93. 1962.

Holotype: Vietnam, 6 km e. of Mui Ne, Phan Thiet Province,

top of beach, 9 Feb. 1960, St. John 26,342 (BISH).

- This seems to be distinctive in its unusually long prickles;

those on the leaf base up to 11 mm. long; in the large cephalium,

27 cm. long, 19 cm. thick, with 106 phalanges; and in the

stout phalanges of 5-9, usually 6-7 carpels, about 6 cm. long

and 4 cm. thick.

Pandanus odoratissimus. L.f£. forma karikayo (St. John) B. C.

Stone, comb. et stat. nov.

P. Karikayo St. John, Pac. Sci. 15; 335. f. 16. 1961.

Fr. adduensis ‘St. John, ibid’ FS: °328.°r 12: 1967.

P. Hartmannii St. John, ibid. 15; 333. f. 15. 1961.

Holotype: Maldive Islands: Fadiffolu Atoll, Kuredu Islet,

Shore, 25 Sept. 1957, W. D. Hartmann | (A).

This form may be tenable because of the larger number of

carpels per phalange, usually from 9 to 14.

Pandanus odoratissimus L.f. forma rubricoloratus (St. John) B. C.

Stone, comb. et stat. nov.

P. rubricoloratus St. John, Pac. Sci. 17: 32, f. 163. 1963.

Holotype: Malaya: Pahang; Pulau Tioman, Telok Paya,

sealevel, 19 May 1927, M. R. Henderson 18,444 (SING).

Rather distinctive because of its small globose cephalium only

9X85 cm., with 34 phalanges, each about 3.6x2.5 cm., the

stigmas somewhat elevated.

Pandanus odoratissimus L.f£. forma hueensis (St. John) B. C. Stone,

comb. et stat. nov.

Pandanus hueensis St. John, Pac. Sci. 16: 88. f. 60. 1961.

242 Gardens’ Bulletin, Singapore — XXII (1967)

Holotype: Vietnam: Thai Duong Ha, Hue, thicket at edge of

brackish marsh by inner beach, sandy soil, 2 m. alt., 3 April, 1960,

H. St. John 26,343 (BISH).

This seems to have certain characters permitting it to be ranked,

at least for the time being, as a forma, although its relationship

to P. odoratissimus is certainly very close. The important features

are these: the short prickles, which even at the leaf base are

only 2-4 mm. long (but white! ); the large cephalium, 28 X20 cm.,

with 143 phalanges; the large, rather narrow phalanges, about

7.4-8 cm. long and 2.8-4.6 cm. thick, with 4-9 carpels; and the high

position of the endocarp, in the upper third of the phalange.

Summary and post-script.

Pandanus odoratissimus L.f. (Sect. Pandanus), the type species

of the genus, was described from a male flowering specimen from

Ceylon. A neotype collection has been made by Prof. H. St. John

(Pac. Sci., in sched.) to establish the identity of the female plant,

particularly the fruit: cf. Fig. 6. This is a coastal species, and the

phalanges are distributed by ocean-currents. It is to be expected

then that it should have a fairly wide distribution, in the manner

of most coastal plants with oceanic dispersal, e.g. lpomoea

pes-caprae, Canavalia rosea, Hernandia sonora, Heritiera litoralis,

etc. Nonetheless a very large number of species have been

described, especially in the last few years, on the basis of

specimens which are agreed to be close relatives (at least) of

P. odoratissimus. By means of an analysis of the taxonomic

criteria involved, and a tabular comparison of the alleged dis-

tinguishing features, it is belived to be evident that only one

species can be distinguished, but it is clearly rather polymorphic,

and hence several formae may be recognized. Although there are

still no doubt further synonyms among the relevant binomials

in existence, it is already possible to confirm that the broad

concept of Warburg (1900) who considered that only one species

ranged from the Indian Ocean to the limits of Polynesia, is

erroneous; and that instead, there are undoubtedly a few distinct

species in the Pacific Islands, though certainly fewer than the

available binomials in existence would indicate. One of the

Pacific species has borne the name Pandanus tectorius, which

however is considered now to date validly only from its publica-

tion by Warburg in 1900, and not from Parkinson, 1773. Con-

sequently another name must be found for the species (singular

or plural) of Tahiti; several are available, including Pandanus

douglasii Gaudich. (1841) which has already been used for the

Hawaiian plants (manifestly of only one species) which certainly

also occurs in Tahiti and no doubt elsewhere in Polynesia.

How many species of this general ‘Pacific’ alliance can finally

be discriminated, it is too early to tell; but it is certain that none

penetrates into the Indian Ocean, at least not to the Maldives or

beyond, and apparently throughout that area and in Malaysia

(Malesia) the vicariant representative species is P. odoratissimus.

Nonetheless, a member of this ‘tectorius’ group does overlap into

the area of P. odoratissimus, evidently in the general region of

Stone — On Pandanaceae 243

Wallacea, i.e. in parts of the Malayan-Indonesian Archipelago,

particularly in the vicinity of Celebes-Amboina-Halmaheira, as

also perhaps on the eastern coast of the Philippines.

Pandanus odoratissimus owes a number of its synonyms and

in general much of the confusion surrounding the species is also

due to the ephemeral nature of one of the major characteristics

of the species, namely the abrupt fleshy dark red shoulders of

the mature phalanges. Immature fruits, even when seen in the

field, lack these; dried fruits, even when collected fully mature,

also lack them (due to shrinkage); and many field-collected fruits

lack them from depredations of ants or other insects. Con-

sequently, this character, so effective in defining P. odoratissimus,

has either been missed or ignored by most previous authorities.

The other major character is the nature of the large, white spines

of the leaves. By itself, this feature of armature must have seemed

of secondary importance to earlier botanists; yet in conjunction

with the fleshy shoulders, it is the hallmark of the species. Neither

character is found in the Pacific Islands beyond Malaysia. Instead,

these plants — whether of one, or as I rather believe, a few

closely related species — have no shoulders on the phalanges,

or if present, these are orange or vermilion, merely slightly

bulging and never abrupt or dark red, and persisting after drying,

i.e. not merely consisting of pulpy exocarp; and further, the

leaves bear shorter, green spines, rarely as much as 4 mm. long,

in contrast to those of P. odoratissimus, which may reach a

length of 8 or 10 mm. Other distinguishing features will no doubt

be found to correlate with these two obvious ones.

The western limit of P. odoratissimus appears to be the Maldive

Islands of the Indian Ocean, although I suspect that P. balfourii

Martelli of the Seychelle Islands is merely a form; but nothing

like the species has ever been recorded from Mauritius, Madagascar,

or the South African coast. Rendle’s P. rabaiensis needs reviewing

in the light of the above discussion also: this was from Mombasa

— Dar-es-Salaam.

The eastern limit of P. odoratissimus seems to be along Wallace’s

Line, approximately. Field investigations in this region are

critically needed. The status of P. coronatus Martelli (Philippines)

must be reviewed. The distribution clearly extends northward into

the Ryukyu Islands (cf. P. tectorius var. liukiuensis Warb., and

var. ferreus Y. Kimura). No doubt also Warburg’s P. tectorius

var. sinensis pertains to P. odoratissimus. Exploration on these

borders is a major desideratum, for only until these borders

are fully known can the questions relating to distribution be

approached.

REFERENCES

RIDLEY, H. N. 1925. The Flora of the Malay Peninsula. Vol. V. Mono-

cotyledones (concluded), Gymnospermae, General Indices. Pandanus,

pp. 72-81. L. Reeve & Co., London.

St. JOHN, H. 1959-67. Revision of the Genus Pandanus. Pacific Science,

14-21, (continuing).

STONE, B. C. 1967. On the use of Carpel Number as a Taxonomic Criterion

in distinguishing species of Pandanus. Amer. Journ. Bot. 54 (in press).

—~

lon

—

“

co)

ea)

ah)

1S)

244

« YSIPPol,,

qupiu wo

10 o]ed,,

sopyorid “ws

St x ECT

oyM sourds

“UD ¢°9 — Bg

we c=.

UMOIG

“Ysippol

sdi} surds

mE cos

oe — LT

s[reyop

oulds

-UIPrM

x YSU]

2SOAROT

dues

dAOISUBUT

a10desuls

durems

JOVEM YSoIJ

‘fuer

‘o10desulsg

yoraq

vos Suvudg

"1099

“AV TCIO'T

Cy — 6¢/

9p — vb/

¢¢/

SoA

Jey

“JI aod

suloA Jo Ft /

UMOUY aDW

SOA

(91n}e WUT 3nq)

SoA

jou IO

slop[noys

AYsoy WIAA

YSIMOTIOA

uMOdAG—Uaad8

(VE = 02

x 09 — SS)

“WU Pe

— 8C X €9

ouou

"WU Of — 87

* 99 = 09

ouou

pol-osue1o

yuid—uaass

~— [| Ajeser | ‘ww Op — CZ

ouou ‘nsn oo = SF

jou

Iojoo +

Jo juasoid sosurejeyd Jo

ba le odeys + ozIg

jesoeT ‘

i /

toE—)9

"yd ¢/

‘yd 19/

uimniyeydes od

sosuvyeyd

JO "ON

/osuvjeyd

Jod sjodies

Jo “ON

"(WD ZZ

‘pod) od

udYyM pol

(prosd}j]9)

oLIpuryAs

[ews

(Wd pT x 97)

"WO Op ‘pod

prosdijjo

"wd OT xX IZ

‘Wd /7 ‘pod

esoqgo]sqns

"WO CLT X €7

9]ounped =

pod ‘(qse]

ye yuopuod

sdieouAs ]]e)

odeys + ozis

dievsuhg

SONVONVd dO SdIOddS GHSOdOUd $7 AO SHYNLVAA OILSONOVIG AHL AO NOSIYVdWOO — I ATAVL

> ‘pned iuog ‘d

** UYOL 1S SNonDpsiquiv *g

uYyOf 3S Snjvajovagiqw “_

so1sedg

245,

Stone — On Pandanaceae

sdij uMOIg

YIM ovTGM dureMs

[[e sourds ysryoviq :

‘Wd Lp — PP ‘Tue Or — 8¢/

xX WLT—s' Tl ‘s10desuls ou SoA ouou

‘sy neing

-ooulog

«poddn Seqe[9)

uUMOIG,, 1SDOI

sopyporid SUO}SOWT]

"ud ¢'¢ |UOTMeYSsULT g¢/ 91} BUT

xX ‘dd 06—I8 neng ou A4aa ynq ‘ou ouou

poddn uejuENyy

-Ysipper ynq “sueyed

gjed =sourds asD]]1A

wd 7S — PP Huery Sp/

x “al p’7—9' | ‘aIodesuls ou SoA ouou

S[Te}op

outd aed ou JO nee

ra _, 1092 J, sod art ous Jo yuosoid

x YISuey Ayyeoo'T suloA Jo FF / | gion 1M soinjns

:SOARO] UMOUY ADIN 4sen Ut [er19}eT

pol-osuvlo

YSiuaa¢s

"WU Eh — €7

SO ="'09

“WU 8Z — 77

AGt ie

oSuUvIO-Pol

uaads AjjAvd

"ULL CE — PZ

| Peace by

IOJOS +

sosuvjeyd jo

odeyg + 92ZI¢

"ud OF ‘pod

19 — vb/ esoqo]sqns

8I-peiqnop "Wd OC—ET X

Ajorel ."wd €7 — OT

(ZI) (6—L) (9) a UYOL "IG Snjouroul

"Wd 6 ‘pod

97/ asoqos

9—G | “Wd 8°8 x 8's

‘T "* UYOL “1g Snsogojs “gq

oesoqo]sqns

ao | ‘WICT X 81

OI —9 | a | uYOL 1S SNSOUdDI *g

winiyeydes Jod gjounpod =

sosuvjeyd pod :(qsv]

JO ‘ON 78 Juopued

/osuepeyd sdieous |[v) ope

Jod sjadievs odeys + 9ozIs

JOON diesusg

panujju0d —SONVGNVd HO SHIOddS GHSOdOUd $7 AO SHAUNLVAA OLLSONOVIG AHL AO NOSIAVdWOO — I ATaVL

Gardens’ Bulletin, Singapore — XXII (1967)

246

poddn

-UMOIG

+ ofed

sopyorid

‘Ud € — BZ

ae 99

YsIppor 3nq

ged sopyorid

‘Wo ye x

poddi-mMo1q

“au1yM soutds

‘WD Z'p

ze | ohm “al

s[rejop

oulds

“YIpIM

xX Ysusy

:SOABY]

L7/

uvWIOL] Neng a ia

o10desulsg

upjupny 6¢ a IZ/ adnjouiu1

-sueyed ou ynq ‘ou

fuer ct — vE/

‘s10desuig ou soa

feet I

‘Joos "JI tod scseihon

‘AqI]VOIO7] suIoA Jo tf / | AYSOL UITAA

UMOUY aD

ou0Uu

osuou

ouou

jou

Io Juosoid

soins

[e19}e'T]

pol

pod be/ "Wd +¢] “pod

“UU {7

SS OCR OE b—9 "WI 6°83 x6

(pelsp) 8¢/ "WO+€Z “pad

WU BE —LZ

x $$—0S OIl—8 | “WO ?7ITXZI ‘9

o8ULIO-poz

udads rs/ ‘wd ¢7 ‘pod

"UIUI OF —EZ

a Gs OV 8—S | “WO SPL XxSsT

tuntyjeydas Jad | sajounpod—

sosurjeyd pod :(qsvy]

OJOS -+ ;

JO "ON ye JUuspued

pee eel SBS /osuvyeyd sdieouAs |v)

aS 'S Jod sjodies odeys + ozs

JO ‘ON diesuks

UYOL "}S SJD40]0914GN4 “I

a, UYOL IS SNSsnjgGO "g

"* UYOLr 1g SmIDssvsdUI “gd

so1sedg

penunuoI—~SONVANWd AO SHIOddS GHSOdOUd $7 AO SAUNLVAA OLLSONOVIC AHL AO NOSIYVdNOO — I STAVL

247

Stone — On Pandanaceae

| Sopyorud

jo —_ ploaAsp

jsOwye

dry ‘sopyorid

ged ase]

‘US SPs CV

HEB PBL

peddnumoig

udyjO OTM

sourds jjous

"Wd ¢°¢ — LP

Ro OTe

YSIMOTIOA

sopyorid

‘WD Cp

x Ure’ l—Ss6 0

s[reyop

oulds

IpIM

x YSU]

:SOABOT

panuyjuoI—SONVANVWd AO SdIOdadS GAsSOdOuUd

WIPUJOIA,

WIeU}OI A

SUO}SUOH

"[O99

‘Ay[LOOT

é/

ou SOA quou

inal (Atuoredde)

ou ou ouou

MOTeYs 1nqG

. # souo [erjied

SoA sad | IayJO JO Moj B

Jey jou

: jou 10

nagar smpmones:| Bamaea

AYSOY YAN

UMOUY ADIN [es1ojeT

osuvio

ugass vL/

"uUILU OF — IT

x 1€—LZ ane o

pues

o3uvio €vt/

‘WLU Op — QZ

xX 08 — PL |6—) L—9(—S)

osuPvio

mMopjad +0s/

"ul OF — OT

—9rx7p | OL (6—8) ()

uiniyeydas sod

OJOS ++ pre

sosuepeyd JO | / s3urjeyd

odeyg + 2ZI¢ Jod sjadievo

JO ‘ON

"Wd O¢ ‘pod

PrAo-or diya

"Wd OZ X 8Z

"ld QZ “pod

prosdiqj9

“UID OZ

— 81 X €@

gjounped =

pod :(jse]

18 yUSpued

sdiesuAs |e)

odeys + 9oZIs

dieouks

UuYOL 31S S1IIdD14sa]UI “J

uyOL “1g SiSuaany *d

uyor IS Snjowad

so1sodg

S$? JO SAUNLVAA OLLSONDVIG AHL SO NOSIYVdNOD — I aTAVL

Gardens’ Bulletin, Singapore — XXII (1967)

248

poddnumoiq

“OUTYM

sopyorid

‘Wd 9 — 8'P

x “Ul €*I—L'0

poddn

-uMOIq Inq

YSIMOTIOA

sopyorid

"UID ¢

x “U9C'T

poddnuMmoiq

usyjO o}IGM

sopyorid

oe Pe

gers 8 a ae 8

s[tejop

ouldg

“YIpIM

xX YIsusy

7SOARO'T

a ee er

souo

é MOT[VYS AIOA

WU} A SoA SOA MO} IO suUOU

é/ o1N} BUT

WUT A ou ynq ‘ou ouou

‘s[sy] MNOnRy

wosj ‘studs

syiodo

ose ‘f 1S ih on} BWI

WeujyorA, ou yng ‘ou ouou

yey jou

"1099 "JI tod - ae g Jo juasoid

“AV[VIOT SUIOA Jo Ft / k sa, : somnjns

UMOUY aD YSU UTM [e19je' 7]

asuvAO

"WU BE — EZ

at: | Pee 14

TEEBs

“WU Oh — CC

A Biro Ve

IOJOS +

sosuvjeyd jo

odevyg + 2ZzIg¢

16 — 7S/

L—S

78/

6—9

tunijeydas sod

/esuepeyd

JO "ON

/osuejpeyd

Jod sjadies

JO ‘ON

"WdS7—0Z ‘pod

Sa 4 ame 9 |

*-9L—.CI

‘wd + Q] ‘pod

“USS PLXS BT

"ud ST ‘pod

Wd COI — 6

5 SoGE EE

ojounpod ==

pod :(ase]

}v JuNpuod

sdeouAs |e)

odvys + ozs

dievsudks

UYOF “1G S1jD41dsisdadad *_

"* wyor 3G suajoafoud *g

UYOLF “IG WSuvoYwuDY, “I

sa1sedg

panuyuom—SNVANVWd AO SHIOddS GHSOdOUd

St dO SAUNLVAA OLLSONOVIG AHL SO NOSIYVdWOO — I ATAVL

249

ovIgM

sopyoud

"Ud p

TESOL

OUT

sopyorid

Df L-6

a eal

<,lO]OO

-MPIS,,

sopyorid

"8 Vy ese ee

aS SP

x ‘Wd [—8'0

s[Te}Op

ourldg

Stone — On Pandanaceae

o1N}B WUT

' é/ Apystys

WIeUIOIA OU | ynq ‘ou

é/

WUT A, ou SOA

sounp él oIn}e LUT

WPUSIA ou ynq ‘ou

é oIn} eww

WPUJIIA Sod ynq ‘ou

JTeY

. ; }Ou JO

[O90 JI tod sropjnoys

‘AU[COO'T SUIOA JO 3 / knsou 1

UMOUY aD 4Soy UTM

ouoUu

ouou

Moy JO ouOU

SAOQP SB SUBS

o8ueIO

“uu ¢¢ — gT

SB) | Sree

osuerso

‘WU Tp — CZ

xX 6b — SP

YSIMO]TOA

uaads

Or te LC

x tv — OV

osurio

> — u00I18

“WU Ep — PT

xX 0S — SL

jou

Io juosoid

soinyns

jeiojey

JojOo +

soduvyeyd jo

odeyg + 92I§

(F=).0b<=9

ZL/

6-8) Ti = 9

98/

(OI)

(¢—2). 19)

wnijeydes Jod

sosuvjeyd

JO “ON

/osuejeyd

Jod sjadies

JO ON

"Wd O7-S | “pod

“UD O]

4 et 3

uyOL "3S SMsvjngns *g

"UID OZ “pod

osuevIO

‘Wd pT X OT

“ql

** UYOL 1G Snsousvoqns *q

‘WOC TT X py | °° wyor aT HPUubUuljIUs ‘d

‘pod

"WS CPT X LT | UYOL 3G Svjnobgojwas ‘d

‘T

gyounped =

pod :(qse]

ye Juopuod

sdiesus Ie)

odeys + ozs

divsusg

so1sadg

penujuoI—SQNVANVWd AO SaIOddS GaSOdOUd $7 JO SAUNLVAA DASONOVIG AHJ JO NOSIAVdNOO — IAaTavL

poddnumoiq

sopyord

"wud €°9 X “UW C*]

poddnumoiq

‘gjed

sopyorid

D.C —-p

Sul) ) mae

“uuu TT

0} soutds

JOMOT OJIYM

sopyorid

aD os

Raat |

Gardens’ Bulletin, Singapore — XXII (1967)

s[rejop

duldg

-UIPiM

x YISsUd]

:SOARO'T

250

a te a eh i a

spuvys]

OAIDILIL ou

Js} equieuy tp — 8¢/

VISUOPUT ou sok

é/

WU A ou SOA

Jyey |

Ou JO

satel ahded sJopynogs

‘AyPeAO'T sulerto.we./ || itePihed

UMOUY ajDIV AYsey WIAA

ouoUu

ouou

duou

jou

IO juosoId

soinjns

]e19}e'T]

‘WH [yp — ve

xX 6¥—S'P E Lis 6 é

OL/ "Wd +07 ‘pod

“UU

Ob—v7 x (0S (Ol — 8)

a7 (Sk. — Gg) TT ah.) UT OX er

YSIMO]JOA

pad-asUuvso 901/ "Wd O¢ ‘ped

+ yoazeOs

“WU 7p — ST (L — 9) | po1—joy1v9s

ADP By 6S) "Ogi x Le

wuniyeydes Jad ajounpod =

r sosuvreyd pod :(jsey

soaueyijo | J270N, | ae 1uapuad

ade I - a /osueryeyd sdivouAs |[e)

4S Ss Jod sjadies odvys + ozs

JO'"ON diesudg

"* UYor “Ig sysuanppy “gq

** WYOL "IG mUOSdapuaz] “_

UYOL "1S SisuauDUjalA “gq

sorsedg

panuyjuoo—SONVAGNVd AO SHIDAdS GHSOdOUd $7 JO SAUYNLVA OLLSONDVIG AHL HO NOSTYVdNOO — I AIAVL

251

iS)

sopyorid

+ poner plosdijo

~yAeD,, ; 09/ peolq

wd¢ — FP spur]s] Wd PE — ZZ

XO Lt sATP[PW ou é ouou | x 7¢ — Bp | 9 ‘Nsn “6 —¢ | “WOTT X PT

<«<o]ed,,

“UID Z°C «Po IY4sIg,,

xX ‘w ({E—) spuvys] wo pe —Z

SC Eat oATP[PW ou é suou | xX HH —B8eE vI—Ol | “MO €T X BI

sapyorid ao |

«Sd Yep,, spurs] (Mo]TeYys Moy Wd p — 3°Z prosdiyo

"MID "5 x UT SAIP[VIAL ou i AidA JO) oUOU Xx Sp—Pp (pI —) ZI ‘wd pI X OZ

sjreyop winijeydes od ojounpod =

suds : aed jou JO }0U Jojoo -+ soduvjeyd pod :(jse]

“YPM Raa pe nee sJopjnoys stag eoaeata jo ged! ye USpusd

x Yisue AypesoT =| Sula JO FF/ | soinjns ade ae /asuereyd | sdavouds ][e)

:SOARO UMOUY BID i [e1oyeT WS + IS | tod sjodies odeys + ozIs

" | JO ‘ON diesusg ©

panuijuoI—SANVGNVd AO SHIOddS GHSOdOUd $7 AO SAUNLVAA OJLSONOVIG AHL AO NOSIAVdWOD — 1 ATAVL

SNIAAIPIVIAW “dT

uYyor “Ig odvynDy “g

** WYOr IS NuuDUAD]T “gd

so1sedg

a2, Gardens’ Bulletin, Singapore — XXII (1967)

Figure 1

Pandanus odoratissimus L.f. A fruiting female tree about 8 m. tall, on

the sandy beach at Tanjong Bungah, north coast of Penang (Stone 6/71), |

near the type locality of P. albibracteatus. :

Stone — On Pandanaceae 253

Figure 2

Pandanus odoratissimus L.f. A young plant in the Singapore Botanic

Gardens.

Above, view of the whole plant. Note the numerous small leafy

branches near the base at left.

Below, midportions of several leaves showing the ventral surfaces.

Note the slender, elongated, white marginal prickles.

254 Gardens’ Bulletin, Singapore — X XII (1967)

Figure 3

Mature cephalium of Pandanus odoratissimus L.f. (Stone 6171 from

Penang). The color is red-orange or vermilion; a few carpel tips are still

green.

Stone — On Pandaraceae 255

Figure 4

| Cephalia of Pandanus odoratissimus L.f. (Stone 6171 from Penang).

| Above, an unripe young cephalium.

| Below, a ripe cephalium cut in half.

256 Gardens’ Bulletin, Singapore — XXII (1967)

|40 6106 6171

tag. A

(51) ZN (115)

CEYLON P , 6350

peti ind ¥y ’ \ (sa)

! AA

i s g *

/ os he 5 \

oe eared Wek

/ : \\

Vs i : \ om:

vad ¥ x ‘a

O e ewe eet = 4 x “7

1 2 3 4 5 6 7 8 9 10 11

Figure 5

‘Carpel numbers in Pandanus cdoratissimus L.f.

The vertical axis shows number of phalanges; the horizontal axis shows

number of carpels per phalange. Whole-peohalium samples only are used.

The numbers above the sample line refer to collections. The number in

parentheses below the sample line indicate the total number of phalanges

borne by the cephalium. No. 6/06 is from Johore; no. 6/7] is from Penang;

and no. 6350 is from Penang. Major carpel numbers are: for 6/06 6;

for 6171, 5; and for 6350, 7. The total range is 2-10. Individual ranges are:

for 6106, 3—9; for 6171, 2-10; for 6350, 5-9. It should be added that two

additional cephalia, not shown graphically here, from the same individual

as 6106, gave the following data: (a) total phalanges 40; range 59; major

carpel number 7. (b) total phalanges 47; range 4-8; major carpel number 7.

The Ceylon sample (St. John 24212) shows a major carpel number 8; the

range is 5-11; total phalanges 59.

Note.—Strictly, these graphs should be histograms, but in order to con-

serve space, lines have been used so that several samples may be super-

imposed. The variation between each carpel number is of course not

continuous.

Stone — On Pandanaceae 257

Figure 6

Phalanges of Pandanus odoratissimus L.f.

Left: profile, top view, and sectional view of St John 24212 from Ceylon.

Centre: profile, top view, and sectional view of St. John 24218 from

Ceylon.

Right: profile, top view, and sectional view of Stone 6350 from Penang

(Natural size). :

Taxonomic Notes on Bornean

Dipterocarpaceae, III

P. S. ASHTON

Forest Department, Kuching, Sarawak*

This paper succeeds two others (this journal, XIX (1962)

253-319; id.. XX (1963) 229-284) in which new species of

Dipterocarpaceae from Brunei and Malaysian Borneo were

described, others reduced to synonymy and reassessments were

made of the infrageneric classification of some genera. The first

two papers were published as precurors to my Manual of the

Dipterocarp trees of Brunei State (O.U.P., 1964); the present

similarly precedes a forthcoming supplement to the Brunei Manual

to include additional information on the same species, and ali

further species, that occur in Sarawak. Furthermore, since the

Brunei manual was sent for publication, more complete material

has been obtained of several taxa there referred to separate and

undescribed taxa.

This paper is the last in the present series to be devoted solely

to problems in Bornean Dipterocarpaceae. Subsequent papers in

this series will directly relate to an eventual revision of the family

for Flora Malesiana. .

This work has been greatly assisted both by the inspiring leader-

ship and continued encouragement of B. E. Smythies, Conservator

of Forests in Sarawak during the initial stages of this work, and by

Dr. J. A. R. Anderson, who has given invaluable guidance

throughout.

The plates are the work of Mr. Ahmad Akip, a promising young

artist at present attached to the Sarawak Forest Department.

SUMMARY

Anisoptera Korth., sect. Anisopterae: A. reticulata Ashton is described.

Dipterocarpus Gaertn.f.: D. cuspidatus Ashton is_ described.

Dryobalanops Gaertn. f.: D. abnormis Sloot. is reduced to synonymy

with D. oblongifolius Dyer.

Vatica L.: Retinodendropis aspera Heim is reduced to V. sarawakensis

Heim.

The Bornean species previously referred to V. odorata Griff. is now

identified as V. mindanensis Foxw., which is considered to be a subspecies

of V. odorata. V. aerea Sloot. is reduced to the same subspecies.

5 subspecies are recognised in V. oblongifolia Hook. f.

7 new species are described: V. badiifolia, V. brunigi, V. compressa,

V. congesta, V. globosa, V. rotata, and V. rynchocarpa.

Hopea Roxb.: H. laxa Sym. is reduced to H. pachycarpa (Heim) Sym.

*Present address: Botany Department, University of Abardeen, Scotland.

260 Gardens’ Bulletin, Singapore — XXII (1967)

12 species are described: H. andersoni with subspecies, H. depressinerva,

and H. megacarpa in sect. Hopeae, subject. Hopeae; H. aequalis, H.

bullatifolia, H. centipeda, H. enicosanthoides, H. pterygota and H.

tenuinervula in subsect. Pierreae (Heim) Ashton; H. altocollina, H.

kerangasensis and H. longirostrata in sect. Dryobalanoides Miq., subsect.

Dryobalanoides; and H. mesuoides in subject. Sphaerocarpae (Heim) Ashton.

Shorea Roxb. ex Gaertn. f.: Two new subsections are described:

Polyandrae in sect. Richetioides Heim, and Sagittatae in sect. Muticae

Brandis.

S. meadiana Syrn. is reduced to scrobiculata Burck.

2 subspecies are recognised in S. agami Ashton.

S. grandiflora Brandis is recognised as a subspecies of S. hemsleyana

King.

S. leptoclados Sym. is reduced to S. johorensis Foxw.

S. rugosa Heim var uliginosa (Foxw.) Sym. is reinstated as a species.

17 new species are described: S. brunnescens and S. lunduensis in sect.

Shoreae, subsect. Shoreae: S. cordata in sect. Anthoshoreae; Pierre; S.

alutacea, S. bakoensis, S. cuspidata, S. iliasii, S. macrobalanos, and S.

mujongensis in sect. Richetioides Heim subsect. Richetioides; S. polyandra

in subsect, Polyandrae; S. praestans and S. rotundifolia in sect. Pachycarpae

Heim; S. pubistyla and S. carapae in sect. Brachypterae Heim Brandis sub-

sect. Brachypterae; S. foraminigera and §S. pallidifolia in sect. Muticae

subsect. Muticae; and S. sagittatae in subsect. Sagittatae.

ANISOPTERA

Anisoptera reticulata Ashton sp. nov Piate 1.

Species floribus ut in sectione sed foliis ramulisque glabris

intercostis subreticulatis.

Foliae ramulique glabri. Ramuli apices versus c. 2 mm. diam.

teretes ruguli fusci; internodis -1.5 cm. longis; cicatricibus

stipularibus brevibus horizontalibus. Gemmae —2 x 2 mm. ovoideae

subacutae. Stipulae ignotae. Lamina 4.5—13 x 2.2-5.5 cm. elliptico-

obovata coriacea basim versus late cuneata, apice in acumen

—-5 mm. longum breve abrupte terminata; costis lateralibus

utrinsecus 9-14 curvatis subtus prominentibus angulo 60—65°

exorientibus; intercostis subreticulatis; costa media supra applanata

vel subdepressa subtus prominenti. Petiolus 15-35 mm. longus,

c. 2 mm. diam. teres prominente geniculatus apicem versus

tumescens, in sicco niger. Racemi —6 cm. longi, terminales vel

axillares breves teretes plus minus persistente aureo-pubescentes

stipitis basi in fructu —3 mm. diam. semel ramosi. Alabastrum

-8 x3 mm. fusiforme. Lobi calycis anguste deltoidei inaequales

valvati. Petala lorata cremia partibus in alabastro expositis

pubescentibus. Stamina c. 35 subaequalia, filamentis brevibus

compressis antheris oblongis attenuatis aristis filiformibus.

Stylopodium oblongum obtusum pubescente. Pedicellus in fructu

—4 mm. longus —3 mm. diam. crassus prominens. Calyx in fructu

sparsim caduce aureopubescens, tubo —2 cm. diam. subgloboso;

lobis longioribus 2, -133 cm. lorato-spatulatis obtusis basim

versus —9 mm. latis attenuatis; lobis brevioribus 3, —20X3 mm.

lineare-lanceolatis acutis. Nux apicem versus conferte persistente

alutaceo-pubescens stylopodio -4 mm. longo -3 mm. diam.

oblongo obtuso pubescenti coronata.

Ashton — Bornean Dipterocarpaceae 261

Collections —Sabah: SAN. 12864 (Wood) SAN. 15172 (Wood)

(Holotypus in Herb. Kew) Ulu Sipitang; Brunei: Hotta 12988,

Bt. Teraja; Sarawak: S. 1503 (Drahman) Lawas.

This distinctive species resembles A. marginata Korth. of the

swamps but differs however in the c. 35, not c. 25, stamens, larger

fruits, fewer nerves, subreticulate intercostals, and lustrous entirely

glabrous thickly coriaceous lamina. It appears to be rare and

found in Mixed Dipterocarp forest on leached yellow sandy soils.

DIPTEROCARPUS

Dipterocarpus cuspidatus Ashton sp. nov. Plate 2.

D. eurhynchi Migq. affinis sed petiole longiore lamina cuspidata

alis tubi nucis latioribus differt.

Gemma conferte persistente longe alutaceo-tomentosa; stipulae

externe (intus glabrae) ramuli petioli costa media supra nervi

laminarum subtus sparsim longe subpersistente fusco-tomentosi.

Ramuli apices versus c. 1 mm. diam. graciles teretes multo ramosi

griseo-brunnei; internodis —2 cm. longis, cicatricibus stipularibus

gracilibus obscuris. Gemmae -7X3 mm. oblongae. Stipulae

-18x5 mm. lineares caducae. Lamina 6-11 x 2-4 cm. anguste

elliptico-ovata applanata in statu expanso vix plicata; basi acuta

vel rare cuneata; apice in acumen —2 cm. longum gracile prominente

cuspidatum attenuata; costis lateralibus utrinsecus 8-9 gracilibus

sed subtus prominentibus angulo 35-45° exorientibus; intercostis

conferte scalariformibus; costa media gracili subtus prominent.

Petiolus 12-18 mm. longus c. 1 mm. diam. gracilis geniculatus

apicem versus tumescens. Flores ignoti. Racemi —5 cm. longi stipitis

basi in fructu c. 2 mm. diam. terminales vel axilares teretes graciles

sparsim persistente longi fusco-tomentosi semel ramosi. Pedicellus

in fructu —5 mm. longus c. 2 mm. diam. prominens. Calyx in fructu

glaber; lobis longioribus 2, -8x1.8 cm. spatulatis obtusis vel

subacutis basim versus 4-6 mm. latis attenuatis; lobis brevioribus

3, -9x4 mm. oblongis obtusis subrevolutis; tubo —2 cm. longo

-1.8 cm. diam. alis prominentibus 5 chartaceis subundulatis basim

versus attenuatis sed apicibus —9 mm. latis subauriculatis. Nux

~—3 x 1.6 cm. ovoidea supra tubum maxime extrusa.

Collections Sarawak: S. 15138 (Ilias Paie), S. 22061, 22064

(Ashton) Segan F. R.; S. 22058 (Ashton) Ulu Segan: S. 15821

(Ilias Paie) Labang F. R., Bintulu (Holotypus in Herb. Kew).

The leaves of this species resemble in shape the sapling leaves of

D. eurhynchus Migq., thus differing from the leaves of the type

Specimen of that species only in their smaller average size.

The relatively longer petiole and’ cuspidate acumen serve to

distinguish this species at once from mature D. eurhynchus, with

which it certainly appears to be closely allied, as well as the fruit

with its broader, distally subauriculate, aliform processes on the

tube. The species appears to be a local endemic of central northern

Sarawak, where it is found on low hills in Mixed Dipterocarp

forest on arenaceous soils.

262 Gardens’ Bulletin, Singapore — XXII (1967)

DRYOBALANOPS

The identity of Dryobalanops abnormis Sloot.

Dryobalanops oblongifolia Dyer (J. Bot. 12 (1874) 100) was

originally based on two collections, Beccari 2533 from G. Matang,

Sarawak, and Beccari 3734 from Bintulu; the latter was later cited

as type of D. kayanensis Becc., a synonym of D. lanceolata, so

that Beccari 2533 becomes Jectotype. Brandis (J. Linn. Soc. Bot. 31

(1895) 51) first described the fruits, from Beccari 2993, also froni

Matang.

In his revision of the genus Dryobalanops Van Slooten (Bull.

Jard. Bot. Btzg. 12 (1932) 22) correctly reduced Baillonodendron

malayanum Heim (Bull. Soc. Linn. Paris, 2 (1891) 867) and

Dryobalanops beccariana Ridl. (Fl. Mal. Pen. I (1922) 211) to

D. oblongifolius Dyer. Nevertheless his description of D.

Oblongifolius fruit is clearly based on Malayan and Sumatran

material, and his illustration shows Malayan fruit, and leaves that

in their shape and their revolute bases appear characteristic of

D. lanceolata Burck and not of this species.

Van Slooten founded D. abnormis (Bull. Jard. Bot. Btzg. 16

(1940) 449) on 9 numbers from West and South Indonesian Borneo,

two of which, bb. 29641 and 30213, from near Tjatit, Melawi,

Kapuas, bear fruit; I choose the latter (at BO) as lectotype. He

states that D. abnormis differs from D. oblongifolius ‘among other

characters ... by its larger rigid leaves, its lenticellate and

differently shaped calyx tubes, and its nuts, which are lenticellose

and smooth’; elsewhere he states that the calyx tube is constricted

medially and longer in D. abnormis than in D. oblongifolius:

furthermore the sepals in D. abnormis are deltoid and prominent,

whereas in D. oblongifolius they are reduced almost to an

undulating ring. Though these comparisons are, with the exception

of the length and construction of the calyx tube which depends on

the ripeness of the fruit, valid between D. abnormis and Malayan

material of D. oblongifolius, the Beccari type and his fruiting

collection collected nearby match D. abnormis, so that D. abnormis

must be considered a synomym.

The lamina dimensions quoted by Van Slooten under the two

names overlap and those quoted for Bornean D. abnormis are well

within the range of Malayan and Sumatran D. oblongifolius

specimens now at my disposal. I do not on present evidence

consider that the Malayan and Sumatran specimens represents a

separate taxon.

VATICA

The identity of Retinodendropsis Heim.

This genus was described (Compt. Rend. Ass. Franc. Pau 1892

(1893) 470) from a single number, Lowe s.n., consisting of two

shoots bearing somewhat fragmentary leafy twigs, inflorescences,

and part of a fruit, in the Kew herbarium; the single species,

R. aspera Heim, was published in the same paper.

Ashton — Bornean Dipterocarpaceae 263

Heim described the anatomy of the paris available to him in

considerable detail. He concluded that the material represented a

dipterocarp, whose fruit recalled those of the genus Retinodendron

Korth. (synonymous with the type section of Vatica, see Ashton,

Gard. Bull. Sing. 20 (1963) 264), though with a thinner pericarp

than usual and with ascending, not reflexed, sepals. He noted that

in other characters it resembled his own Vatica sarawakensis,

founded on Beccari 3018 from Sarawak, which is a flowering

collection. Heim, whose generic concepts were extremely narrow

(see Ashton, loc. cit.) considered that as the material was inadequate

to confirm its position in Retinodendron a new genus should be

created for it!

The specimen is without doubt one of V. sarawakensis Heim

bearing immature fruit. In this species the fruit calyx lobes are at

first ascending, as in the specimen, and become reflexted as the

fruit ripens.

The identity of Vatica mindanensis Foxw. and Vatica aerea Sloot.

Vatica mindanensis was founded on Elmer 13680 with young

fruit, and 13398 and 13359 with flowers from Mr. Urdaneta,

Cabadbaran, Agusan Province, Mindanao.

Vatica sorsogonensis was later described from Elmer 16840,

from Mt. Bulusan, Irosin, Sorsogon, Luzon and clearly represents

a fruiting specimen of the same taxon. The complete material of

V. mindanensis is found to be identical with V. odorata (Griff)

Sym. as it occurs m Borneo. Furthermore V. odorata in Borneo

occurs typically between 350-1,300 m. altitude, as also does V..

mindanensis in the Philippines.

V. mindanensis differs from the type and other mainland material

of V. odorata in having a somewhat longer, more or less geniculate,

petiole, and lamina drying greyish, not yellow-brown. The ranges

in petiole length overlap, and I consider that V. mindanensis

should be considered as a subspecies of V. odorata.

Bornean material has been separately described by Van Slooten:

under the name Vatica aerea (Bull. Jard. Bot. Btzg. 3, 17 (1942)

133) based on bb. 26438 (Budding), a fruiting collection from

Tjatit B. Tengkoejoeng, Melawi; this decision is based on my

examination of the isotype in the Leiden herbarium; the Holotype

is at Bogor.

V. odorata (Griff.) Sym. ss. odorata.

Petiole 6-18 mm. long, not geniculate; lamina drying yellow-

brown.

Type: Griffith s.n., Mergui (K).

Distribution —Indo-China, Thailand, Peninsular Burma, Malaya,

West Borneo.

ssp. mindanensis (Foxw.) Ashton., stat. nov.

Lectotype: Elmer 13398, Mt. Urdaneta, Mindanao (K).

V. mindanensis Foxw., Leafl. Philip. Bot. 6 (1913) 1957.

iat,

264 Gardens’ Bulletin, Singapore — XXII (1967)

V. sorsogonensis Foxw., Philip. J. Sc. Bot. 13 (1918) 196.

196.

V. aerea Sloot., Bull. Jard. Bot. Btzg. 3, 17 (1942) 241.

Petiolus 15-20 mm. longus, plus minus geniculatus, lamina

delapsa sicco grisea.

Distribution—Borneo, Philippines.

Intraspecific variation in V. oblongifolia Hook. f. Plate 3-7.

Lowe s.n. (K), the fruiting specimen upon which this species

is based, is clearly distinguishable from other species in the type

section of Vatica both by its stout compressed twigs, narrowly

oblong-obovate thickly coriaceous leaves (blue-green when fresh),

and vinous tomentum. Nevertheless 4 distinct taxa, sharing these

characters but each differing in lesser criteria, including the

number and appearance of the nerves, length of the petiole, size

of lamina and twigs, and size and shape of the fruit sepals, exist.

Each of these 4 taxa have a well defined geographical and

ecological distribution and I have not seen two sharing a common

habitat. In view of their close similarity, however, I perfer on

present evidence to regard them as subspecies, though intermediate

forms have not as yet been found.

Subspecies oblongifolia——Ramuli c. 4X2 mm. diam. crassi.

Lamina 10-31 x4.5-10 cm. obovata vel oblonga basi obtusa

acumine brevi lato; costis lateralibus utrinsecus 10-18 subtus

prominentibus. Petiolus 2.5-5 cm. longus. Lobi calycis in fructu

—3 x2 mm. deltoidei acuti.

Collections—Lowe, s.n. Borneo (Holotypus in herb Kew).

Sarawak: S. 21550, 21560 (Ashton) Batu Empar, Santubong;

S. 15907 (Brain) Nyabau F.R., S. 9618 (Ariffin) Lundu; S. 9605

(Ariffin) G. Pueh F.R.; S. 8644 (Brunig) Similajau F.R., S. 7980

(Bojeng) G. Gading; S. 12783 (Hj. Bujang) G. Santubong; S. 13145,

13251 (Smythies) S. Bena, Kapit; S. 22293 (Suib) Bt. Raya, Kapit;

S. 12816 (Asah) Semengoh F.R.; Kep. 358 (Omar) Bt. Siol; S. 13305

(Ilias) G. Periggi; S. 21371 (Othman) Bt. Lambir. Brunei: Brun.

5683 (Ashton) K. Belalong; Brun. 2625 (Ashton) Andulau F:R.;

Kep. 48222 (Wyatt-Smith) Badas-Labi road; Kep. 80112 (Wyatt-

Smith) Peradayan F.R. Sabah: SAN. 16424 (Wood and Charring-

ton) Timbang Batu, Kudat; SAN. 16660 (Wood) Ulu Moyah

Sipitang; SAN. 23857 (Jad Binting) Pulau Sepanggor, Jesselton;

SAN. 32014 (Mikil) Halogilat, Beaufort; SAN. 20702 (Meijer)

Bt. Ataidon, Ranau.

Distribution Throughout Sarawak to western Sabah. Found

on shallow leached loams derived from both granite, sandstone,

and shale.

Subspecies selakoensis Ashton, ssp. nov. Ramuli c. 2X1 mm.

graciles. Lamina 6.5—22 x 2.5-6.5 cm. anguste elliptica basi cuneata

apice in acumen -—1.5 cm. longum angustum attenuata, costis

lateralibus utrinsecus 11-18 gracilibus subtus vix elevatis. Petiolus

1.5-2.5 cm. longus. Lobi calycis in fructu —4 x 3 mm. oblongi obtusi.

Ashton — Bornean Dipterocarpaceae 265

Collections —Sarawak: S. 18466 (Chai), S. 13340 (Ilias) (Holo-

typus in herb Kew), S. 12619 (Smythies), S. 9639 (Ariffin), Sinclair

and Kadim 10387, G. Gading F.R.; S. 13171, 13174 (Smythies) G.

Berumput.

Distribution—Found on shallow leached raw-humus_ bearing

soil above 600 m. alt. on the upper slopes and summits of the

granodiorite mountains of W. Sarawak. Ssp. oblongifolia and ssp.

multinervosa occur on the same mountains but at lower altitudes.

Subspecies crassilobata Ashton, ssp. nov. Ramuli c. 4X2 mm.

crassi. Lamina 11.5—21 x 4.5-10.5 cm. Jate oblonga basi obtusa vel

cordata apice in acumen breve latum attenuata; costis lateralibus

utrinsecus 16-23 crassi subtus prominentibus. Petiolus 1.8-2.8 cm.

longus. Lobi calycis in fructu -15x12 mm. obovati obtusi vel

emarginati subrotati.

Collections——Brunei: Brun. 614 (Holotypus in Herb. Kew), 615

(Ashton), S. 2246, 5601 (Anderson), Andulau F.R.; Brun. 5157

(Ashton) Berakas F.R.; Hotta 13002, Bt. Puan. Sarawak: Fuchs

21371, Bt. Pesu, Ulu Semut, Tubau; s.n. (Anderson) Bt. Gaharu.

Distribution.—Local, in Sarawak and Brunei, on strongly leached

very deep pale yellow sandy soils derived from poorly consolidated

soft sandstones and clays.

Subspecies multinervosa Ashton ssp. nov. Ramuli c. 4X2 mm.

crassi. Lamina 14-31 x4-8.5 cm. anguste obovata basi anguste

obtusa vel cuneata apice in acumen —1.5 cm. longum gracile

attenuata; costis lateralibus utrinsecus 18-27 subtus prominentis-

sime elevatis intercostis prominentibus. Petiolus 1.5—2.5 cm. longus.

Lobi calycis in fructu -13 x 8 mm. oblongi obtusi reflexi.

Collections Sarawak: S. 22239, 22240, 22295 (Suib) Bt. Raya,

Kapit; S. 17739 (Ashton) Ulu Belaga; S. 12623 (Smythies) G.

Gading. Brunei: S. 5786 (Wood, Smythies, Ashton) Bt. Biang.

Sabah: SAN. 26077 (Muin Chai) Takun, Lahad Datu; SAN. 18475

(Martyn) Ulu Kelumpang, Tawau; SAN. 26029 (Meijer and Muin

Chai) Mt. Silam, Lahad Datu; SAN. 30052( Aban) Beradaya F.R.,

Tawau; SAN. 29835 (Muin Chai) Kelumpang, Tawau; SAN. 19713

(Meijer) (Holotypus in herb. Kew), SAN. 22509 (Singh) Sepilok

F.R.; SAN. 34083 (Aban) Ayer Panas, Tawau.

Distribution —Throughout Sarawak, Brunei and Sabah, on

relatively fertile clay rich latosolic soils.

Subspecies eiliptifolia Ashton ssp. nov. Ramuli c. 3X2 mm.

Lamina 12-—20-3.5—-7 cm. elliptico vel cbovata basi late cuneata

apice in acumen —1 cm. longum breve attenuata; costis lateralibus

utrinsecus 14-16 gracilibus subtus prominentibus. Petiolus 10—14

mm. longus brevis. Lobi calycis in fructu —-8 x 4 mm. oblongi obtusi

revoluti. |

Collections —Brunei: Brun. 925 (Ashton) Andulau hills at

Bt. Kukub (Holotypus in Herb. Kew). Sarawak: S. 23225 (Sibat),

Bt. ju, Arip, Balingian.

266 Gardens’ Bulletin, Singapore — XXII (1967)

Distribution—Known only from these two collections from

Mixed Dipterocarp forest, the one from leached yellow sands

overlying the Seria formation, the other from similar soils on the

Arip rhyolite.

Vatica badiifolia Ashton. Plate 8.

V. bantamensis Sensu Ashton, Man. Dipt. Brun. (1964), 67,

non Burck.

V. bantamensis (Hassk.) Burck affinis, sed lamina delapsa in

sicco badia costis lateralibus utrinsecus 6-10 costa media supra

prominente elevata tomento roseo-brunneo differt.

Ramuli apices versus racemi gemmae stipulae petiolique plus

minus conferte caduce pulverulente fulvo-tomentosi. Ramuli apices

versus —3 mm. diam. teretes vel costati dein levescentes pallide

roseo-brunnescentes; internodis 1.5—2 cm. longis. Gemima —1.5 x |

mm. ovoidea subacuta. Stipula —5 x 1.5 mm. hastata acuta. Lamina

7.5—15 x 3-6.5 cm. coriacea plus minus elliptica; basi plus minus

cuneata, apice in acumen —5 mm. longum deltoideum attenuata;

costis lateralibus utrinsecus 9-12 curvatis subtus prominentibus

teretibus supra paullum elevatis. Petiolus 2—3.5 cm. longus —3 mm.

apicem versus incrassatus geniculatus sicco fulvescens vel

nigrescens. Lamina delapsa sicco badius. Racemi —8 cm. longi

stipitis basi in fructu —-2 mm. diam., terminales vel axillares semel

vel bis ramosi. Alabastrum —1.3 cm. longum; calyx conferte breviter

cremio-pubescens; flores aliter ut in Vaticis aliis. Pedicellus in

fructu —5 mm. longus —1 mm. diam. gracilis. Lobi calycis in fructu

caduce badio-tomentosi ad basim liberi; longioribus 2, —8 x 2.5 cm.

chartaceis oblongo-spatulatis basi -8 mm. latis consirictis;

brevioribus 3, -30x8 mm. hastatis acutis basi -4 mm. latis

constrictis. Nux —8x 8 mm. globosa breviter plane badio-pubescens

stylo —2 mm. longo lineari, coronata.

Collections —Sarawak: S. 15857 (Ilias and Johnson) Similajau,

Labang F.R. (Holotypus in herb. Kew); S. 22080 (Ashton) Nyabau

reservoir. Brunei: S. 2115, 2120 (Smythies) Bt. Teraja; Kep. 47025

(Smith) ‘Brunei’; Kep. 48135 (Flemmich) Balai Rotan; Kep. 39650

(Durant) Labi F.R.

This species, found on yellow sandy loams in Mixed Dipterocarp

forest from the Belait valley to the lower Rejang in northern

coastal Borneo, bears a close superficial resemblance to V.

bantamensis (Hassk.) Burck of West Java; the two can be

differentiated as follows:

V. badiifolia V. bantamensis

Lamina drying chestnut- Lamina drying grey-green.

brown.

Nerves 6-10 pairs, stout. Nerves 8-11 pairs, slender.

Midrib prominently raised Midrib more or less applanate

above. above.

Petiole 2.2-3.5 cm. long. Petiole 1-2 cm. long.

Tomentum pink-brown. Tomentum buff.

Ashton — Bornean Dipterocarpaceae 267

Vatica brunigi Ashton sp. nov. Plate 9..

V. maingayi Dyer affinis sed lamina petiolo ramulisque conferte

breviter ochraceo-tomentosa costis lateralibus subtus haud

prominentibus differt.

Ramuli apices versus gemmae stipuli lamina subtus petiolique

breviter alutaceo-scabrido-tomentosi, Jamina supra fugace floccosa

tomentosa. Ramuli apices versus c. 2 mm. diam. graciles teretes;

internodis —1.5 cm. longis brevibus. Gemmae -—3 x 2 nim. ovoideo-

conicae obtusae. Stipulae -5 x2 mm. loratae obtusae. Lamina

6-12 x 2.5-6.5 cm. ovato-elliptica vel obovata basi cuneaia vel

obtusa apice in acumen —1 cm. longum attenuata: costis lateralibus.

utrinsecus 9-12 gracilibus subtus paullum elevatis angulo. 65°—80°

exorientibus; costa media supra vix elevata subtus prominenti;

intercostis scalariformibus supra obscuris subtus paullum elevatis.

Petiolus 8-15 mm. longus c. 1.5 mm. diam. Racemi —12 cm. Jongi.

stipitis basi in fructu —2 mm. diam.; teretes vel angulati conferte

breviter persistente ochraceo-scabrido-tomentosi terminales_ vel

apices ramulorum versus axillares semel (axillares) vel bis

(terminales) ramosi; ramulis -—6 cm. longis; flores distichos

gerentibus; bracteolis —3 x 1 mm. ellipticis sparsim pubescentibus.

Alabastrum -8x2 mm. ellipsoideum. Calyx conferte breviter

ochraceo-griseo-pubescens; petala cremia anguste oblonga obtusa

externe sparsim pubescentia intus glabra; aristae breves erectae;.

stylum columellare quam ovarium longiore; flores aliter ut in

Vaticis aliis. Pedicellus calyxque in fructu sparsim alutaceo-

pubescens, nux scabrido-alutaceo-pubescens. Pedicellus —3 mm.

longus —1 mm. diam., gracilis. Lobi calycis longiores 2, -6.5 x 1.5

cm. lorati vel spathulati obtusi; breviores 3, -15x6 mm. ovati

acuti paullum recurvati. Nux —9 x 7 mm. ovoidea subacuta.

Collections—Brunei: S. 1011, 12352 (Brunig) Ulu _Ingei.

Sarawak: S. 17045 (Ilias Paie) (Holotypus in Herb. Kew) Sabal

F.R.; S. 6758, 17451 (Brunig), S. 17885, 17871 (Ilias Paie), S. 17300:

(Ilias Paie and Paul Chai) Bako National Park; S. 18389 (Ashton)

Tubau; S. 6437, 7132 (Brunig) G. Pueh F.R.; S. 5969 (Brunig)

Sempadi F.R.; S. 8256 (Yakup), S. 1484 (Dan) Marudi F.R.; S.

6524 (Yakup) Kampong Mienki, Ulu Sebuyau, Simanggang:

S. 8937 (Yakup) Setapok F.R.

At once distinguished by the densely shortly evenly yellowish

buff lamina undersurface, this further addition to Section Sunaptea

(Griff.) Burck is found throughout Sarawak north-eastward to the

Belait basin of Brunei. It is named in honour of Dr. E. F. W. O.

Brunig, who is contributing so much to our knowledge of the

floristics and ecology of Heath forest in Borneo.

This species was not known to me from Brunei when I compiled

the Manual of Dipterocarp trees for that State.

Vatica compressa Ashton sp. nov. | Plate 10.

V. maritimae Sloot. affinis sed ramulo compresso, lamina inter-

costis remotioribus, petiolo 15-23 mm. longo glabrescenti differt.

Novelli breviter plane pallide roseo-pubescentes stipulae intus

exceptae, gemmae stipulaeque externe persistente pubescentes

268 Gardens’ Bulletin, Singapore — XXII (1967)

partes aliter glabri. Ramuli apices versus c. 3X1 mm. compressi

pallide fusci leves; internodis —3 cm. longis; cicatricibus stipularibus

primo c. 2 mm. longis pallidis horizontalibus vel descensis

prominentibus. Gemmae c. 3 x3 mm. ovoideae acutae compressae.

Stipulae -4-1.2 cm. oblongae obtusae subcaducae. Lamina

11-19 x 5-8.5 cm. ovato-elliptica coriacea basi obtusa apice in

acumen —1 cm. longum attenuata; costis lateralibus utrinsecus

10-15 gracilibus subtus paullum elevatis haud prominentibus supra

vix elevatis curvatis angulo c. 50° exorientibus, secundartis brevibus

obscuris; intercostis scalariformibus; costa media subtus prominenti

supra applanata vel elevata. Petiolus 15-23 mm. longus c. 2 mm.

diam. sicco nigrescens. Racemi-7 cm. longi, basim versus — mm.

diam. paullum compressi sparsim_ pallide roseo-brunneo-

pubescentes, irregulariter semel vel bis ramosi. Alabrastrum

~9x4 mm. lanceolatum. Calyx conferte pallide fulvo-pubescens;

corolla cremia partibus in alabastro expositis conferte pubescenti-

bus; flores aliter ut in Vaticis aliis. Pedicellus in fructu -8 mm.

longus, —1 mm. diam. gracilis pubescens. Calyx in fructu puberulens

vel glabrescens; lobis longioribus 2, —6.5 x 1 cm. loratis plus minus

obtusis basim versus c. 3 mm. latis gradatim attenuatis; lobis

brevioribus 3, —-20 x6 mm. deltoideis acutis revolutis. Nux —5 mm.

diam. breviter ovoidea conferte breviter alutaceo-pubescens, stylo

—3 mm. longo glabro coronata.

Collections.—Sarawak: SFN. 35618 (Daud and Tachun) Hutan

Setapok (Holotypus in herb. Kepong); S. 309 (Mead), SA. 604

(Igon) Mile 6 F.R., Kuching; S. 6101, 5984 (Brunig) Sempadi F.R.:;

S. 6604 (Brunig) Bako N.P.; S. 13275 (Anderson) S. Klauh,

Simangang. |

A species which is locally frequent on poorly drained sandy

podsols in Heath forest in Sarawak west of the Batang Lupar.

Vatica congesta Ashton sp. nov. Plate 11.

V. coriaceae Ashton affinis sed petiolo 1.2-2.5 cm. longo,

racemis brevibus fasciculatis conferte congestis differt.

Ramuli apices versus gemmae stipulae petiolique conferte

breviter pallide ochraceo-fusco-scabrido-tomentosi, lamina subtus

costa media basim versus supra sparsim eadem. Ramuli apices

versus c. 3 mm. diam. crassi primo angulati rugosi dein fuscen-

scentes chartaceo-exfoliantes striati; internodis 1-3 cm. longis.

Gemmae —43 mm. late ovoideo-conicae acutae. Stipulae -8 x3

mm. anguste deltoideae acutae. Lamina 8-22 x 3.5-8 cm. oblongo-

elliptica vel obovata coriacea, basi obtusa, margine revoluto, apice

subacuminato vel obtuso vel retuso; costis lateralibus utrinsecus

7-12 supra paullum elevatis subtus crassis prominentibus, angulo

40°—55° exorientibus; intercostis subtus prominentibus, supra

paullum elevatis reticulatis; costa media supra vix elevatis subtus

crassa striata prominenti. Petiolus 1.2-2.5 cm. longus 2-3 mm.

diam. crassus sicco rugescens. Inflorescentiae paniculatae —3 cm.

longae, stipitis basi in fructu —2 mm. diam., breves axillares vel

breviter ochraceo-scabrido-tomentosae semel ramosae; ramulis

—1.8 cm. longis flores distichos gerentibus. Alabastrum —8 x 3 mm.

Ashton — Bornean Dipterocarpaceae 269

Calyx conferte breviter ocharceo-tomentosis, lobis deltoideis acutis

in alabastro expansis; petala lorata conferte pubescentia; aristae

breves bifurcatae; stylum breve sericeum; flores aliter ut in Vaticis

aliis. Pedicellus in fructu —4 mm. longus gracilis. Calyx glaber;

lobis longioribus 2, -12 2.2 cm. loratis obtusis prominente 3-

costatis; lobis brevioribus 3, -4.5 x 1.5 cm. anguste ovatis attenuatis

acutis recurvatis. Nux —12 x 8 mm. ellipsoidea glaber apice in stylo

—6 mm. longo gracili attentuata.

Collections Sarawak: S. 9347A (Holotypus in herb. Kew),

9347 (Yakup), S. 15606, 15713 (Galau), S. 15021, 15045, 14936

(Rosli), Sinclair and Kadim 10219, Semengoh F.R.; S. 15148 (Ilias:

Paie) Segan F.R., Bintulu.

S. 15148 differs from the Semengoh collections in having 7-8.

instead of 10-13, pairs of nerves in the lamina, but shares the

large fruit calyx and the short ramiflorous inflorescences, by which

this species is so clearly distinguissed from V. coriacea Ashton and

V. teysmanniana Burck (as well as in other leaf character). I

therefore have no hesitation in considering it to represent the

same species.

V. congesta has been collected in Mixed Dipterocarp forest on

leached yellow soils rich in clay, on low hills.

Vatica globosa Ashton sp. nov. Plate 12.

Species fructu ut in V. oblongifolia Hook. f. sed tomento rufo

haud vinoso lamina chartacea subcaudata petiolo brevi ramulo

tereti differt.

Ramuli gemmae petiolique persistente rufo-sericei. Ramuli apices

versus c. 2 mm. diam. teretes levescentes pallide brunnescentes;

internodis -—3 cm. longis; cicatricibus stipularibus brevibus

horizontalibus obscuris. Gemmae 2x2 mm. ovoideae acutae.

Stipulae ignotae. Lamina 7-18 x 3-6.5 cm. obovata subchartacea,

basi anguste cuneata, apice in acumen —2 cm. longum cuspidatum

attenuata; costis lateralibus utrinsecus 13-16 gracilibus subtus

prominentibus supra elevatis angulo 35°-60° exorientibus,

secundaris multis brevibus; intercostis gracilibus reticulatis; costa

media supra subtusque prominenti. Petiolus 7-15 mm. longus.

c. 2 mm. diam. supra sulcatus. Racemi —3 cm. longi, stipitis basi

in fructu —1 mm. diam. terminales vel —3 axillares congesti costati

vel teretes persistente rufo-sericei plus minus semel ramosi:

bracteolis fugacis. Alabastrum -5 x2 mm. lanceolatum; calyx

externe conferte rufo-pubescens intus glaber; corolla cremia

partibus in alabastro expositis rufo-pubescentibus; aristae conicae

quam antheras 4-plo longioribus; flores aliter ut in Vaticis aliis.

Fructus omnibus partibus persistente rufo-sericeis. Pedicellus in

fructu —2 mm. longus c. 2 mm. diam. Lobi calycis -4x3 mm.

aequales oblongi obtusi reflexi. Nux -20 mm. diam. globosi obtusi

suturis 3.

Collections —Sarawak: S. 18091 (Ashton) Ulu Stirau, Labang

(Holotypus in herb. Kew); S. 14485 (Brain) Nyabau F.R.; Haviland

and Hose 3159, Niah; Haviland and Hose 3148, near Kuching:

S. 23751 (Jugah) Bt. Iju, Arip.

270 Gardens’ Bulletin, Singapore — XXII (1967)

This species differs from V. oblongifolia Hook. f. both in the

rufous, not vinous tomentum, the terete twigs, and the texturally

thinner lamina with subcaudate acumen and short petiole. Apart

from the single record Haviland and Hose 3148 it is known only

from central Sarawak between the Kemena and Niah drainages,

where it is locally frequent in Mixed Dipterocarp forest.

Vatica rotata Ashton sp. nov. Plate 13.

V.. dulitensis Sym. affinis sed lamina late elliptica costis lateralibus

utrinsecus 9-11 vix elevatis intercostis conferte reticulatis petiolo

8-12 mm. longo differt.

Ramuli gemmae petioli costa media subtus persistente conferte

vinoso-sericei. Ramuli apices versus c. | mm. diam. graciles multum

ramosi teretes griseo-fuscescentes leves. Gemmae c. 21 mm.

ovoideae subacutae. Stipulae ignotae fugaces. Lamina 5.5—10 x 3-5

cm. late elliptica vel ovata coriacea, basi obtusa, apice in acumen

-8 mm. longum attenuata; costis lateralibus utrinsecus 9-11

curvatis vix elevatis angulo 65°—80° exorientibus; intercostis

gracilibus conferte reticulatis; costa media supra obscura depressa

basi excepta subtus prominentibus. Petiolus 8-12 mm. longus

c. 1 mm. diam. gracilis. Flores ignoti. Fructus racemique omnibus

partibus persistente plane vinoso-sericei. Racemi —1.5 cm. longi,

stipitis basi in fructu c. 1 mm. diam., terminales vel axillares

graciles teretes semel ramosi. Pedicellus in fructu —-9 mm. longus

c. 1 mm. diam. longus gracilis. Lobi calycis in fructu -7 X4 mm.

aequales suborbiculares revoluti rotati. Nux c. 4 mm. diam.

subglobosa obtusa suturis 3 obscuris.

Collection Sarawak: S. 19594 (Ashton) Ulu Kenyana, Mukah

{Holotypus in herb. Kew).

A member of the type section. Though only known from this

single collection this species, whose fruit is very similar to that of

V. vinosa Ashton and V. dulitensis Sym., but whose leaf recalls

that of V. borneensis Burck in sect. Sunapteae, is quite distinct

from other described species. It was collected in Heath forest.

Vatica rynchocarpa Ashton sp. nov Plate 14.

Lamina ut in V. venulosam BI. sed costis lateralibus utrinsecus

10-14 vix elevatis lobi calycis in fructu inaequales longiores 2

ascendentes spatulati.

Ramuli apices versus racemi gemmae stipulae petiolique breviter

sparsim griseo-puberulentes, dein glabrescentes. Ramuli apices

versus —1 mm. diam. graciles teretes dein levescentes pallide

griseo- brunnescentes; internodis —8 mm. longis brevibus. Gemma

-1.5x1 mm. conica subacuta. Stipula —2.5 mm. longa anguste

hastata acuta fugax. Lamina 5-8.5 x 1.3-2.5 cm. anguste elliptica

vel lanceolata chartacea; basi obtusa apice in acumen -1 cm.

longum angustum attenuata; costis lateralibus utrinsecus 10-14

gracilibus marginem versus curvatis subtus vix elevatis angulo

75°-85° exorientibus remotis, nervis secundariis brevibus; costa

media gracili subtus elevata supra paullum depressa. Petiolus

—4 mm. longus —1 mm. diam. brevis gracilis sicco nigrescens. Flores

Ashton — Bornean Dipterocarpaceae 271

ignoti. Racemi —3 cm. longi stipitis basi in fructu -1 mm. diam.

terminales vel axillares semel ramosi pallide brunneo-pubescentes.

Pedicellus in fructu -7 mm. longus —1 mm. diam. gracilis. Calyx in

fructu glaber, lobis basi liberis, longioribus 2, -6.2*1.4 cm.

chartaceis spathulatis angustis obtusis basim versus —2.5 mm. latis

gradatim attenuatis; brevioribus 3, —15 x 2.5 mm. hastatis acutis.

Nux -18X8 mm. ovoidea glaber stylo -4 mm. longo lineare

Collections Sarawak: SA. 514 (Egon) N. Temulan, Rejang

(Holotypus in herb. Kepong); S. 16568 (Ashton) Ulu Sinrok,

Simalajau F.R.; S. 22047 (Ashton) N. Sepulau, Segan; S. 12107

(Ashton) N. Amau, Mujong. Brunei: Brun. 3354, Ulu Senuko.

This well-defined species is now fairly well known, and is locally

common along river banks in Sarawak North-East of the Rejang.

HOPEA

The identity of Hopea laxa Sym.

This species, based on Richards 2361, with flowers, from G. Dulit

(Holotype at Kew) was described by Symington (Gard. Bull. S. S.

8 (1934) 33) at a time when only the type of Hopea pachycarpa

(Heim) Sym., Haviland 2730 with fruit, and one other, at that

time unidentified, specimen (S. 512, Merit, Sarawak) has been

collected. The type of the latter has, on the evidence of material

at present available, smaller leaves with fewer nerves than the

average for the species, while in Richards 2361 the reverse is true.

There is no doubt, however, that the two are conspecific, Heim’s

name being the older.

Hopea aequalis Ashton sp. nov. Plate 15.

H. nervosae King affinis, sed lamina basi obtusa costis lateralibus

utrinsecus 16-20, lobi calycis in fructu breves subaequales diifert.

Costa media supra puberulens, partes aliter glabrae. Ramuli

apices versus c. 2 mm. diam. teretes leves pallide fusci: internodis

—4 cm. longis; cicatricibus stipularibus brevibus horizontalibus

obscuris. Gemma c. 2 1 mm. ellpsoidea obtusa. Stipulae ignotae.

Lamina 13-25 x5.5-8 cm. lanceolata chartacea basim versus

obtusa apice in acumen —1 cm. longum attenuata; costis lateralibus

utrinsecus 16-20 gracilibus subtus prominentibus angulo 65°—70°

exorientibus; intercostis conferte scalariformibus gracillimis a

nervis diagonale exorentibus; costa media supra subdepressa subtus

prominenti gracili. Petiolus 15-18 mm. longus c. 2 mm. diam.

sicco niger. Flores ignoti. Racemi —9 cm. longi, stipitis basi in

fructu c. 1 mm. dam., axillares teretes glabri non vel semel ramosi.

fructu c. 1 mm. diam., axillares teretes glabri haud vel semel ramosi.

ovati subacuti saccati incrassati. Nux —23 x15 mm. ovoidea acuta

saepe resinifera. j

Collection —Sarawak: S. 15881 (Hias Paie) Nvyabau F.R.

(Holotypus in Herb. Kew). Sabah: SAN. 38797 (Meijer) Mile 804,

Labuk Road trace, Sandakan.

Zhe Gardens’ Bulletin, Singapore — X XII (1967)

Though founded on but two collections the fruit and leaf

characters of this species are sufficiently striking to distinguish

it at once from its nearest allies, whch appear to be H. nervosa

King and H. sublanceolata Sym.

Hopea altocollina Ashton sp. nov. Plate 16.

H. mengerawan Miq. affinis, sed lamina breviori Janceolata

prominente revoluta acumine caudato differt.

Omnes partes glabrae. Ramuli apices versus c. 2 mm. diam.

teretes fusci lenticellis parvis pallidis punctati; internodis —2 cm.

longis; cicatricibus stipularibus brevibus obscuris. Gemma-l X I

mm. subglobosa parva. Stipulae ignotae. Lamina 7—10 x 3-4.5 cm.

lanceolata prominente revoluta basim versus obtusa apice in

acumen —1 cm. longum caudatum attenuata; costis ‘dryoblanoideis’,

costis lateralibus utrinsecus c. 16 obscuris; costis secundariis

brevibus; costa media gracili supra subtusque vix elevata. Petiolus

10-13 mm. longus c. 1 mm. diam. subgeniculatus apicem versus

subtumescens gracilis sicco niger. Flores ignoti. Racemi —4 cm.

longi, stipitis basi in fructu c. 1 mm. diam. terminales vel —2

axillares teretes glabri semel ramosi. Fructus glaber. Pedicellus in

fructu —2 mm. longus -1 mm. diam. Lobi calycis in fructu

longiores 2, -4.5 0.8 cm. spatulati obtusi basim versus —3 mm.

lati attenuati, partibus basalibus -6X3 mm. anguste ovatis

incrassatis saccatis; breviores 3, -4x4 mm. suborbiculares obtusi.

Nux -7X5 mm. ovoidea apice in stylo — 2 mm. longo gracili

attenuata.

Collections Sarawak: Brun. 1030 (Ashton) Bt. Antu, Ulu

Sembayang, Limbang (Holotypus in Herb. Kew); S. 22461 (Sibat)

Bt. Mersing, Anap; S. 19618 (Ashton) Carapa Pila, Ulu Mujong;

S. 21267 (Ashton) base of Batu Kapal, Mujong; S. 22705, 22706,

22714 (Murthy and Ashton) G. Dulit.

The attenuate apex of th enut suggests that these floral ovary

lacks a stylopodium. This species most resembles H. mengerawan

Mig. of E. Borneo, Sumatra, and Malaya; it differs however in

the powdery flaky not ruggedly fissured bark surface, and in the

revolute margin of the relatively shorter lamina with its caudate

acumen.

H. altocollina is confined to clay rich soils at these altitudinal

limits of the Mixed Dipterocarp forests, and has been observed at

between 900-1,400 m. It is sometimes an enormous tree, —60 m.

tall, -4 m. girth and with tall convex thin spreading plank

buttresses.

Hopea andersoni Ashton sp. nov. Plate 17.

H. sangal Korth. affinis sed staminibus 15 lamina lanceolato-

falcata inaequalis glabra, costis lateralibus remotioribus subtus

minus prominentibus differt.

Foliae ramulique glabri. Ramuli apices versus c. 1 mm. diam.

teretes leves fusci; internodis —3 cm. longis; cicatricibus stipularibus

brevibus obscuris. Gemma —1 X 1 mm. subglobosa. Stipulae ignotae.

Lamina 5—14X2-6 cm. lanceolato-falcata vel elliptica coriacea.

Ashton — Bornean Dipterocarpaceae 273

basim versus adaxiale obtusa abaxiale cuneata inaequalis, apice in

acumen —2 cm. longum gracile saepe falcatum attenuata; costis

lateralibus utrinsecus 9-12 gracilibus subtus vix elevatis curvatis

angulo 65°-75° exorientibus saepe domatiis prominentibus

pustularibus axillaribus; intercostis scalariformibus; costa media

supra vix elevata subtus prominenti. Petiolus 5-10 mm. longus

c. 2 mm. diam. sicco niger. Racemi —12 cm. longi, stipitis basi in

fructu-1 mm. diam., terminales vel axillares teretes saepe fasciculati

conferte plane plus minus persistente pallide griseo-puberulentes;

semel ramosi flores —9 secundis gerentibus; bracteolis -2 mm. longis

linearibus puberulentibus fugacis. Alabastrum -3x*2 mm.

ellipsoideum. Calyx lobis ovatis acutis subaequalibus externe

conferte pubescentibus intus sparsius eisdem. Petala lanceolata

partibus in alabastro expositis conferte pubescentibus. Stamina 15

verticillis 3 inaequalibus; filamentis basim versus compressis

attenuatis apicem versus filiformibus; antheris late ellipsoideis;

aristis gracilibus quam antheras 2-—24-plo longioribus. Ovarium

stylopodiumque cylindricum conferte papillosum puberulente, parte

media subconstricta, stylopodium quam _ ovarium paullum

angustiore; stylo brevi columellari. Pedicellus in fructu —3 mm.

longus c. 1 mm. diam. gracilis. Calyx in fructu basim versus

puberulens aliter glabrescens; lobis longioribus 2, -6x2 cm.

oblongis obtusis basim versus 3 mm. latis attenuatis; partibus

basalibus -4x3 mm. ellipticis subincrassatis saccatis; lobis

brevioribus 3, -4X3 mm. ovatis obtusis saccatis. Nux -85 mm.

ovoidea glabra stylopodio parvo truncato styloque c. | mm. longo

mucronata coronata.

Holotypus: S. 11096 (Anderson) Kuching road, Bau (in Herb.

Kew).

Two subspecies are recognised:—

Subsp. andersoni subsp. nov.

Lamina delapsa in sicco rufescens. Cortex nigrescens altis fissis.

Collections —Sarawak: S. 12238 (Anderson), S. 10038 (Zen),

Seburan, Bau; S. 11096, 4639 (Anderson) G. Benarat, Baram:

S. 10334, 10352, 10353 (Brunig) Bt. Serapah, Serian road; S. 22865,

22876 (Chai and Lee) Bt. Tabai, Bau; S. 16182 (Chai and Lee)

Bt. Kolong, Bau.

Subsp. basalticola subsp. nov.

Lamina delapsa in sicco rava. Cortex fulvescens tenue exfolians.

Collections —Sabah: SAN. 28649 (Meijer and Baker) Ulu

Balong, Tawau; SAN. 23278 (Meijer and Lampangi) Bt. Tingka,

Kinabatangan. Sarawak: S. 25005 (Holotypus in Herb. Kew),

S. 25040 (Sibat) Bt. Mersing, Anap; S. 23481 (Othman and Suib)

Long Kelaby, Ulu Dapoi, Tinjar; S. 22736 (Murthy and Ashton)

G. Dulit.

This species is named after Dr. J. A. R. Anderson, in honour

of his work on the peat swamp and limestone vegetation of Borneo.

274 Gardens’ Bulletin, Singapore — XXII (1967)

The two subspecies, though only recognisable with experience

in the herbarium, are very distinctive in the field owing to their

characteristic bark configurations. The type subspecies is an

obligatory calcicole, whereas ssp. basalticola appears to be confined

to basic intrusive rocks and calcareous shales.

Hopea bullatifolia Ashton sp. nov. Plate 18.

Hic species sect. typici subsect. Pierreae sedis; lamina bullata

subtus badio-pubescenti facile distinguitur.

Gemma stipula externe (intus glaber) petiolus costa media

subtusque conferte persistente plane badio-pubescens; lamina

costaeque laterales subtus costa media supra sparsim eidem. Ramuli

apices versus c. 2 mm. diam. teretes dein leves pallide fusci;

internodis —3 cm. longis; cicatricibus stipularibus brevibus obscuris.

Gemma —1 x1 mm. parva subglobosa. Stipulae —-4 mm. lineares

caducae. Lamina 16—34 x 4.5—9 cm. oblonga inter costas prominente

bullata, basim versus cordata subaequalis, apice in acumen —1 cm.

longum gracile gradatim attenuata; costis lateralibus utrinsecus.

17-26 crassis subtus prominentibus supra depressis angulo 35°-45°

exorientibus basi excepta prope marginem saepe conjunctis; inter-

costis scalariformibus subtus prominentibus supra depressis; costa

media subtus supraque prominenti. Petiolus 3-6 cm. longus 2-5

mm. diam. brevis crassus. Flores racemique ignoti. Fructus glaber.

Lobi longiores calycis in fructu —8 x 1.5 cm. spatulati obtusi basim

versus —4 mm. lati attenuati, partibus basalibus —8 x3 mm. ovatis

incrassatis saccatis; lobi breviores 3, —15 mm. longi lanceolati

graciles basim versus ut in lobilongiores, nucem obscurantes..

Nux c. 10X7 mm. ovoidea apiculata.

Collections.—Sarawak: S. 522 (Mashor) S. Danaw, Rejang

(Holotypus in herb. Kew); S. 17730 (Ashton) Ulu Belaga.

A small smooth-barked tree, endemic to Central Sarawak where

it is found in scattered groups in the understory of Mixed

Dipterocarp forest. It most resembles H. enicosanthoides (also

described in this paper) but differs in the somewhat smaller leaf

with pubescent undersurface and petiole, builate between the

intercostals, drying dark chocolate-brown not pale rust-brown.

Hopea centipeda Ashton sp. nov. Plate 19.

—H. acuminata (non Merr.) Ashton, Man. Dipt. Brun. (1964) 94.

—H. acuminatae Merr. similis, sed ovario stylopodioque pyri-

formi, costis lateralibus lamina domatiis pubescentibus ornatis.

differt.

Ramuli apices versus gemmae stipulae petiolus et costis latera-

libus subtus plus minus conferte breviter brunneo-tomentosi.

Ramuli apices versus —1 mm. diam. horizontale ramosi dein

griseo-brunnescentes levescentes glabrescentes lenticellis cremiis

parvis orbicularibus punctati; internodis 1-1.5 cm. longi. Gemma

~1.5 mm. longa subglobosa. Stipula -3 mm. longa linearis. Lamina

5.5—-9 x 1.5-3.5 cm. lanceolata basi inaequali cuneata apice in

—

Ashton — Bornean Dipterocarpaceae 275:

acumen —1.5 cm. longum caudatum attenuata; costis lateralibus.

utrinsecus 7-9 subtus prominentibus paullum curvatis angulo

30°-40° exorientibus domatiis axillaribus pilosis prominentibus;

intercostis gracilibus subscalariformibus; costa media _ subtus

prominenti tereta supra angusta depressa. Petiolus 4-7 mm. longus

—1 mm. diam. gracilis. Racemi —4 cm. longi, stipitis basi in fructu

-0.5 mm. diam. axillares semel ramosi puberulentes. Alabastrum

—6X2 mm. lanceolatum. Calyx sparisim pubescens, lobis ovatis

acutis externis 2 quam interni 3 paullum longioribus. Petala

cremia basim versus roseo-suffusa. Stamina 15, verticillis 3

inaequalibus; aristis quam antheras subglobosis c. 3-plo longioribus.

Ovarium stylopodiumque pyriforme glabro; stylo stylopodium

aequanti. Pedicellus in fructu —1 mm. longus. Calyx in fructu

glabrescens, lobis longioribus 2, —2.8 x 0.6 cm. spatulatis obtusis

basim versus —1.5 mm. latis attenuatis partibus basalibus —2.5 x 2

mm.; brevioribus 3, —-4 mm. longis, basim versus ut in lobis

longioribus. Nux -4x3 mm. ovoidea glabra apice acuta.

Collections.—Brunei: S. 5750 (Wood, Smythies, Ashton) K.

Belalong; Brun 726 (Ashton) K. Sekurop, Temburong. Sarawak:

S. 18440 (Ashton) Ulu Tubau, Bintulu; S. 12111 (Ashton) N. Amau,

Mujong; S. 19645 (Ashton) N. Sebatang, Mujong; S. 23012, 23011,

23010 (Othman), S. 22682, 22653, 22654, 22655, 23342 (Holotypus

in herb. Kew) (Murthy and Ashton) Ulu Dapoi, Tinjar.

I had previously compared this species, owing to its obvious

leaf similarities, with H. acuminata Merr. of the Phlippines.

The latter, however, is hard wooded, with flaky bark, only 10

stamens, and a truncate stylopodium.

Hopea depressinerva Ashton sp. nov. Plate 20.

H. sangal Korth. affinis sed lamina coriaceori costis lateralibus

utrinsecus 5—8 sine domatiis differt.

Novelli glabrescentes. Ramuli apices versus 1-2 mm. diam.

teretes leves rufo-brunnei; internodis —3 cm. longis; cicatricibus

stipularibus brevibus obscuris. Gemma —1 x 1 mm. minuta. Stipula

ignota. Lamina 5-13 2-5 cm. lanceolata vel anguste elliptica

coriacea basi cuneata apice in acumen —1.5 cm. longum gracile

attenuata; costis lateralibus utrinsecus 6-8 subtus gracilibus elevatis

supra paullum depressa, angulo 40°—50° exorientibus; intercostis

gracilibus scalariformibus. Petiolus 11-13 mm. longus c. 1 mm.

diam. teres sicco nigrescens. Racemi -—7 cm. longi stipitis basi

—l1 mm. diam. teretes terminales vel axillares conferte persistente

alutaceo-pubescentes; semel ramosi ramulis —1.5 cm. longis flores

—5 secundos gerentibus; bracteolfs fugacis ignotis. Alabastrum

—3x2 mm. ellipsoideum. Calyx conferte alutaceo-sericea lobis

ovatis acutis, externis 2 quam internis 3 paullum longioribus

angustioribus. Petala externe rosea, intus flavo-rubra elliptica

obtusa partibus in alabastro expositis conferte pubescentibus.

Stamina 10 aequalia; aristis quam antheras c. 2-plo longioribus

276 Gardens’ Bulletin, Singapore — XXII (1967)

filiformibus gracilibus. Ovarium stylopodiumque glabrum cylind-

ricum truncatum stylo brevi coronatum. Fructus maturus ignotus;

Icbis calycis inaequalibus ut in flores.

Collections —Sarawak: S. 15439 (Holotypus in herb. Kew),

15516 (Anderson), Bt. Gebong, Lundu; S. 395 (Ariffin) G. Pueh

F.R.

This taxon is undoubtedly closely allied to H. sangal Korth.,

but the appearance of the leaf is quite distinct, in many ways

recalling that of immature Shorea maxwelliana King.

Hopea enicosanthoides Ashton sp. nov. Plate 21.

H. polyalthioides Sym. affinis, sed lamina maiori costis lateralibus

utrinsecus 16—30 intercostis conferte scalariformibus differt.

Ramuli gemmae stipulae costa media supra basim versus

petiolusque caduce pallide badio-pubescentes vel glabri. Ramuli

c. 3 mm. diam. teretes vel subcompressi leves dein fusci sub

cicatricibus petiolorum prominente costati; internodis —5 cm.

longis; cicatricibus stipularibus brevibus obscuris. Gemma —2 x 1

mm. conica acuta. Stipula -8 mm. longa linearis subpersistens.

Lamina (16) 27-46 x(5) 8-15 cm. maxima oblonga coriacea facie

Superiore inter costis lateralibus prominente convexa, basim versus

inaequalis cordata, apice in acumen ~—2.5 cm. longum angustum

attenuata; costi lateralibus utrinsecus 16-30 gracilibus subtus pro-

minentibus supra depressis angulo 45°—50° exorientibus; costa media

gracili supra elevata subtus crassa prominenti; intercostis conferte

scalariformibus gracilimis. Petiolus 5-8 mm. longus c. 3 mm. diam.

glaber. Racemi —12 cm. longi stipitis basi in fructu c. 1 mm. diam.

brevis crassus sicco niger. Flores ignoti. Racemus fructusque

—2 axillares saepe ramiflori teretes laxi semel ramosi; ramulis

—3 cm. longi ascendentibus —5 flores gerentibus; bracteolis c. 2 mm.

longis linearibus subpersistentibus. Peticellus in fructu c. 1 mm.

longus, brevis. Lobi calycis in fructu longiores 2, -13 x 3 cm. plus

minus late spatulati obtusi basim versus —6 mm. latus attentuati,

partibus basalibus -12 x10 mm. ovatis saccatis; lobi breviores 2,

—2 mm. longi lanceolati acuti, partibus basalibus saccatis vix

incrassatis, nucem obscurantibus. Nux -10x6 mm. ovoidea

stylopodio —2 mm. longo filiformi attenuata.

Collections Sarawak: S. 18116 (Ashton) Ulu Labang, Bintulu

(Holotypus in herb. Kew); S. 18304 (Ashton) Ulu Sinrok, Similajau

F.R.; S. 13200 (Symthies) S. Bena, Baleh; S. 1467, 1470 (Esmit)

Miri; s.n. (Anderson) S. Sebakai, Bintulu.

This species, which appears to be widespread though local on

alluvium along slow moving non-tidal waterways, is at once

distinguished by its huge pendant lustrous leaves, evocative of

some of the larger-leaved Enicosantha (Anonaceae); they are

conspicuously magenta on first opening. H. enicosanthoides suffered

conspicuously as a result of the unprecedented floods of 1962-63;

in the Bintulu drainage in particular a very high proportion of the

population died.

Ashton — Bornean Dipterocarpaceae 277

Hopea kerangasensis Ashton sp. nov. Pilate 22..

Hic species laminis parvis late ovatis costis lateralibus domatiis

axillaribus prominenti pubescentibus lobi calycis in fructu ovatis.

parvis subaequalibus facile distinguitur.

Ramuli apices versus petioli costa media supra domatiae

gemmaeque breviter plane plus minus persistente pallide flavido-

brunneo-tomentosi. Ramuli apices versus c. | mm. diam. multo

ramosi teretes dein levescentes brunnescentes internodis 3-8 mm.

longis brevibus. Gemma —1 x 1 mm. minuta globosa. Stipula fugax

ignota. Lamina 1.5-4.5x1-3 cm. parva ovata chartacea basi

cuneata apice in acumen —1 cm. longum caudatum attenuata; costis

lateralibus utrinsecus c. 6 dryobalanoidiis obscuris; costa media

supra paullum depressa subtus gracili prominenti, domatiis

utrinsecus —6 magnis pallide fulvo-pubescentibus. Petiolus 3-5 mm.

longus c. 1 mm. diam. gracilis. Racemi —12 cm. longi stipitis basi

in fructu —0.5 mm. diam. axillares parvi teretes sparsim alutaceo-

puberulentes; semel ramosi ramulis —4 mm. longis flores —3 distichos.

gerentibus; bracteolis minutis linearibus fugacibus. Alabastrum

c. 1.51 mm. ovoideum. Sepala ovata acuminata glabra; interna

3 quam externa 2 breviora apicem versus angustiora media parte

latiora. Petala lanceolata partibus in alabastro expositis puberu-

lentibus. Stamina 15 verticillis 3 inaequalibus, antheris subglobosis,

aristis quam antheras 2-3-plo longioribus gracilibus. Ovarium

ovoideum glabrum; stylo ovarium aequanti columellari attenuata.

Fructus ex integro glabra. Peticellus in fructu —1 mm. longus

brevis. Lobi calycis in fructu -6 x5 mm. subaequales ovati acuti

saccati incrassati externis 3 quam internis 2 brevioribus angustior-

ibus incrassatioribus. Nux -85 mm. ovoidea subacuta.

Collections Sarawak: S. 15530 (Anderson et al.) S. Serayan,

Lundu; S. 15608 (Galau), S. 8944 (Holotypus in herb. Kew), 8912,

8912a (Yakup), S. 21299 (Ashton), S. 14926 (Rosli), Whitmore 446,

Semengoh F.R.; S. 9454, 9306 (Bojeng), S. 8932 (Yakup), S. 2274

(Abg. Muas) Setapok F.R.; S. 9669 (Browne) Munggu Unjam F.R.;

Haviland sh Sadong, coal mine hill.

Haviland 857 was identified by Brandis as H. vesquei Heim

but the present species, which is locally common in Mixed

Dipterocarp forest West of the Batang Lupar, is at once recognised

from other members of subsection Dryobalanoides by its very

small leaves with prominent domatia, as well as by its short

subequal calyx lobes.

Hopea longirostrata Ashton sp. nov. Plate 23.

H. beccarianae Burck, H. dryobalanoides Mig. affinis sed

omnibus partibus glabris pedicello c. 5 mm. longissimo differt.

Omnes partes glabri. Ramuli apices versus c. 2 mm. diam. multe

ramosi teretes fusci leves; internodis —1.5 cm. longis, cicatricibus

stipularibus brevibus obscuris. Gemma —2x2 mm. subglobosa

obtusa. Stipula ignota caduca. Lamina 7-9 x 3-5 cm. ovato-elliptica

coriacea basi obtusa apice in acumen —1.5 cm. longum subcaudatum

278 Gardens’ Bulletin, Singapore — X XII (1967)

attentuata; costis ‘dryobalanoidiis’; costis lateralibus utrinsecus

c. 12 obscuris haud elevatis; secundariis longis subaequalibus; costa

media gracili supra prominenti subtus acuta. Petiolus 7-10 mm.

longus c. 1 mm. diam. geniculatus sicco nigrescens. Flores ignoti.

Racemus fructusque glaber. Racemi -4 cm. longi stipitis basi in

fructu c. 1 mm. diam. terminales vel —3 axillares semel ramosi;

ramulis —5 flores gerentibus. Pedicellus c. 5 mm. longus c. 1 mm.

diam. Lobi calycis in fructu longiores 2, —24x6 mm. spatulati

basim versus —3 mm. lati attenuati, partibus basalibus in media

parte tuberculatis incrassatis; lobi breviores -15 mm. longi lineares

vel spatulati obtusi vel acuti basim versus ut in lobi longiores.

Nux -—6 x 4 mm. ovoidea stylo —2 mm. longo attenuata.

Collections.—Sarawak: S. 18191 (Ashton) Ulu Tubau, Bintulu;

S. 17742 (Ashton) Ulu Belaga, Rejang (Holotypus in herb Kew).

This is a further species in the difficult subsection Dryobalanoides.

The leaves closely resemble those of several others, including

H. dryobalanoides, H. beccariana, and particularly H. latifolia

Sym. with which it also shares an overall smooth bark surface.

The completely glabrous parts however distinguish it from H.

beccariana and H. latifolia, while the inordinately long fruit pedicel

and centrally tuberculate basal portion to the calyx lobes are

characteristic and unique in the genus.

H. longirostrata is only known by two collections, though in the

area where I collected it the species was fairly common. It may

well be more widespread as it is easy to overlook when sterile.

Hopea megacarpa Ashton sp. nov. | Plate 24.

H. semicuneatae Sym. affinis sed costis domatiis cbscuris lobis

calycis in fructu —10 cm. longis basim versus subauriculatis differt.

Ramuli apices versus, costi laminae subtus petiolique sparsim

caduce puberulentes, gemmae persistente eaedem. Ramuli apices

versus c. 1 mm. diam. graciles teretes leves dein rufo-brunnescentes;

internodis —3 cm. longi; cicatricibus stipularibus parvis obscuris.

Gemma —1 x1 mm. minuta. Stipula —2 mm. longa lineare-falcata

caduca. Lamina 6-12 1.5-5 cm. plus minus anguste elliptica

chartacea undulata basi cuneata apice in acumen —2 cm. longum

prominenti attenuata; costis lateralibus utringsecus 6-7 gracilibus

subtus elevatis angulo 25°-40° exorientibus; intercostis gracilibus

conferte scalariformibus; costa media subtus prominenti tereti,

supra applanata vel paullum elevata. Petiolus c. 6 mm. longus

c. 1 mm. diam. brevis supra striatus sicco nigrescens. Racemi

—3 cm. longi stipitis basi in fructu —1 mm. diam. axillares teretes

glabri laxi semel ramosi, ramulis flores —3 gerentibus:; bracteis

bracteolisque -2 mm. longis_ linearibus subpersistentibus.

Alabastrum -4x3 mm. subglobosum. Lobi calycis fimbriati,

externi 2 ovati, interni 3 suborbiculares submucronati. Petala

pallide rosea elliptico-oblonga obtusa partibus in alabastro

expositis sparsim puberulentibus. Stamina 15, antheris oblongis:

aristis quam antheras c. 3-plo longioribus gracilibus filiformibus.

Ovarium stylopodiumque cylindricale subtruncatum glabrum stylo

brevi. Fructus ex integro glaber. Pedicellus in fructu -3 mm. longus

Ashton — Bornean Dipterocarpaceae 279

diamque. Lobi calycis in fructu longiores 2, -10 x 1.3 cm. anguste

spatulati subacuti basim versus c. 5 mm. lati attenuati partibus

basalibus —9 x 15 mm. subauriculatis saccatis; breviores 3, —20 x 9

mm. ovati basim versus ut in lobi longiores. Nux -12*10 mm.

ovoidea apice minute truncata mucronata calyce obcura.

Collections —Sarawak: SFN. 35625 (Daud and Tachun) (Holo-

typus in Herb. Kew), SA. 639 (Egon) N. Pelagus; S. 14743

(Anderson) Bt. Raya, Pelagus.

This species, only known at present from a very limited area

on the middle Rejang, is at once distinguished by its fruit. though

the leaves in many ways resemble the widespread H. semicuneata

Sym.

Hopea mesuoides Ashton sp. nov. Plate 25.

H. subalata(non Sym.) Man. Dipt. Brun. (1964) 110.

H. subalatae. Sym. affinis, sed lamina maiore coriaceore

attenuata margine revoluta, lobis calycis in fructu suborbicularibus

aequalibus differt.

Gemma _ stipulaque breviter persistente pallide alutaceo-

tomentosa, aliter glaber. Ramuli apices versus —1.5 mm. diam.

graciles leves teretes badii; internodis 1.5—-3.5 cm. longis. Gemma

—1 mm. longa subglobosa. Stipula —2.5 x 1 mm. hastata acuta fugax.

Lamina 8-14X2.5-5 cm. ovata chartacea basi obtusa apice in

acumen —1.5 mm. caudatum gradatim attenuata margine paullum

revoluto; costis lateralibus utrinsecus c. 11 subdryobalanoidiis

subtus paullum elevatis curvatis angulo 55°-75° exorientibus;

costis intermediis brevibus; intercostis reticulatis obscuris; costa

media subtus prominenti supra anguste depressa. Petiolus 7-10 mm.

longus —1 mm. diam. gracilis sicco nigrescens. Racemi —4 cm.

longi, stipitis basi in fructu —0.7 mm. diam. axillares breves glabri

semel ramosi, bracteolis —1 mm. longis deltoideis glabris sub-

persistentibus. Alabastrum parvum late ovoideum. Calyx externe

sparsim pubescens, intus glaber, lobis externis 2 suborbicularibus

obtusis vel acutis internis 3 suborbicularibus plus minus breviter

mucronatis. Corolla purpurea; petalis oblongis obtusis partibus

in alabastro expositis pubescentibus. Stamina 15, aristis quam

antheras 2-3-plo longioribus gracillimis. Ovarium stylopodiumque

glabrum apice papillosa excepta cylindricum truncatum ovarium

quam stylopodium paullum latiore; stylo brevi glabro. Pedicellus

in fructu c. 1 mm. longum brevis in calyce impressus. Lobi calycis

in fructu —14*12 mm. subaequales ovati chartacei imbricati

saccati ad nucem appressi apices subacutas versus plus minus

erosi. Nux -1414 mm. subglobosa glaber stylopodio truncato

persistenti coronata calyce obscurata.

Collections —Sarawak: S. 15551’ (Ilias) Segan F.R. (Holotypus

in herb. Kew); S. 11247, s.n., Marudi F.R.; S. 25028 (Sibat) Bt.

Mersing, Anap; s.n. (Yakup) Marudi F.R. Brunei: Brun. 91

(Ashton) Mile 3, K. Abang Road; Brun. 879 (Ashton) Bt. Bedawan;

Brun. 5674 (Asah) K. Belalong; Kep. 35532, 35536 (Symington)

280 Gardens’ Bulletin, Singapore — X XII (1967)

Pangkalan Ran; Kep. 34451 (Flemmich) Bt. Rampayoh, Labi

Hills; Kep. 39639, 48203 (Flemmich) Labi Hills F.R.; Kep. 28657

(Smith) Bt. Kerita, Ladan Hills.

At the time of writing my Brunei Manual only immature fruit

were known from Borneo; these appeared to resemble those of

the Malayan species exactly though there were differences in the

leaf. Mature fruit (in S. 15551) however have confirmed that this

taxon should be considered as a separate species.

Hopea pterygota Ashton sp. nov. Plate 26.

H. tenuinervulae Ashton affinis sed lobi calycis in fructu late

auriculati lamina intercostis remotioribus.

Novelli primo conferte alutaceo-pubescentes dein glabrescentes

gemmis stipulis exceptis. Ramuli apices versus c. 2 mm. diam.

fusci dein levescentes glabrescentes; internodis -3 cm. longis;

cicatricibus stipularibus brevibus obscuris. Gemma —2x1 mm.

ovoidea acuta. Stipula -4 mm. longa linearis caduca. Lamina

12-28 x 5-9 cm. oblongo-lanceolata vel oblanceolata, basi obtusa

vel subcordata inaequalis, apice in acumen —2 cm. longum sub-

caudatum attenuata, margine subrevoluto, costis lateralibus

utrinsecus 12-21 gracilibus subtus prominentibus supra depressis

angulo 50°-60° exorientibus curvatis; intercostis gracilibus

scalariformibus; costa media supra prominenti subtus prominenti-

ore. Petiolus 3-8 mm. longus c. 2 mm. diam. brevis crassus. Racemi

—8 cm. longi stipitis basi in fructu —1 mm. diam. teretes glabri

graciles saepe fasciculati axillares saepe ramiflori semel ramosi;

ramulis flores —8 secundos gerentibus. Alabastrum -3 x2 mm.

ellipsoideum. Calyx glaber, sepalis externis 2, lanceolatis .

acuminatis; internis 3, suborbicularibus quam externis 2 paullum

brevioribus breviter mucronatis. Petala lineara partibus in alabastro

expositis pubescentibus. Stamina 15 verticillis 3 inaequalibus,

filamentis basim versus compressis apicibus filiformibus attenuatis;

antheris subglobosis; aristis gracilibus quam antheros 3—4 longio-

ribus. Ovarium anguste ovoideum glabrum stylopodio fusiformi

quam ovarium paullum breviori. Fructus glaber. Pedicellus in

fructu —1 mm. longus —1 mm. diam. brevis. Lobi calycis in fructu

longiores 2, -10 x 1.5 cm. spatulati chartacei basim versus —7 mm.

lati attenuati, partibus basalibus -20 mm. latis auriculatis in media

parte incrassatis; breviores 2, —-3 cm. longi subacuti attenuati basim

versus ut in lobis longioribus. Nux —7 x 5 mm. ovoidea acuta calyce

obscura.

Collections —Sarawak: s.n. (Anderson) G. Lesong, Simanggang;

S. 15206 (Smythies) Bt. Gaharu (Holotypus in Herb. Kew); s.n.

(Tready) Marudi; S. 4355 (Dan) Lambir Hills; S. 25022 (Sibat)

Anap P.F.; S. 22314 (Suib) Bt. Alet. N. Bah, Rejang; S. 23245,

23612 (Sibat) Bt. Iju, Arip.

This species is most obviously distinguished from its allies

H. tenuinervula Ashton and H. philippinensis Dyer by the broadly

auriculate bases to the fruit calyx lobes. In the field it is

distinguished by its smooth, not flaky, bark surface and sofe

white wood.

Ashton — Bornean Dipterocarpaceae 281

H. pterygota is found in Heath forest and sandy ridges in Mixed

Dipterocarp forest; though local it is often a very common

medium-sized tree where it occurs, as in Marudi F.R. where

many trees border the Limbang road.

Hopea tenuivervula Ashton sp. nov. Plate 27.

—H. philippinensis (non Dyer) Ashton, Man. Dipt. Brun. (1964)

107.

H. philippinensis Dyer similis, sed lamina maiore costis laterali-

bus subtus utrinsecus 12-16 differt.

Novelli conferte pallide flavido-brunneo-tomentosi, ramuli

apices versus gemmae stipulae petiolique persistente tomentosi,

costae laminae subtus subpersistente eaedem; aliter tomento caduco.

Ramuli apices versus —-2 mm. diam. dein levescentes glabrescentes

rufo-brunnescentes; internodis 1.5—2.5 cm. longis. Gemma —3 x 1.5

mm. lanceolata. Stipula —5 mm. longa linearis subpersistens.

Lamina 10-18 x 3-5.5 cm. anguste ovata vel lanceolata basi obtusa

inaequalis apice in acumen —1 cm. longum attenuata; margine

paullum revoluto; costis lateralibus utrinsecus 12-16 gracilibus

subtus prominentibus angulo 50°-60° exorientibus_ curvatis:

intercostis gracilibus conferte scalariformibus; costa media tereta

ex integro elevata. Petiolus 3-7 mm. longus —2.5 mm. diam. brevis

crassus. Racemi —8 cm. longi stipitis basi in fructu —1 mm. diam.

—2-axillares rare terminales teretes laxi glabri; semel ramosi ramulis

—3 cm. longis flores secundos —4 gerentibus; bracteolis —1 mm.

longis deltoidiis glabris subpersistentibus. Alabastrum -4 mm.

longum ellipsoideum subsessile. Calyx fimbriatus aliter glaber, lobis

externis 2 anguste ovatis prominente acuminatis, lobis internis 3,

suborbicularibus vel late ovatis mucronatis. Petala oblongo-

lanceolata acuta partibus in alabastro expositis conferte tomentosis

aliter glabra. Stamina 15; antheris oblongo-ellipsoideis, aristis quam

antheras c. 2-plo longioribus gracilibus. Ovarium ovoideum

glabrum; stylopodio ovarium aequali fusiformi glabro stigma

obscura. Fructus ex integro glaber. Pedicellus in fructu —1 mm.

longus brevis. Lobi calycis in fructu longiores 2, —8.5 1.7 cm.

spatulati anguste obtusi basim versus —2 mm. lati attenuati partibus

basalibus —6 x 4 mm. anguste ovatis prominente saccatis incrassatis:

lobi breviores 3, —1.3 cm. longi subaequales acuti partibus basalibus

~7 mm. longis paullum auriculatis. Nux -12*8 mm. ovoidea

stylopodio brevi attenuata.

Collections —Sabah: SAN. 38708 (Mikil) Gaya Island. Brunei:

Brun. 619, 3290, 2624 (Ashton) Andulau F.R.; S. 1187 (Brunig)

Bt. Teraja; Sarawak: S. 14812 (Rosli), S. 11091, 213 (Abg. Muas),

near tree 1154 (Rosli and Asah) Semengoh F.R.:; S. 9869

(Anderson) Rantau Panjang, Sibu; S. 12614 (Symthies) Mt.

Meroyong, Sempadi F.R.; S. 9616 (Ariffin) Lundu (Holotypus in

herb. Kew); Haviland 3167 (Haviland and Hose) near Kuching.

This species, found on leached yellow sandy soils in Mixed

Dipterocarp forest, and on deep white sand podsols in Heath

forest closely resembles H. philippinensis Dyer, but differs, among

other characters, in the larger lamina with more abundant nerves.

282 Gardens’ Bulletin, Singapore — XXII (1967)

SHOREA

The identity of Shorea meadiana Sym.

Symington (Gard. Bull. S.S. 10 (1939) 366) based this species

on Kep. 49845 (Symington) Bt. Goh F.R. Kuantan, Pahang, a

flowering collection. The species is well known in Malaya, and

the many collections now extant show no constant differences from

the also well collected S$. scrobiculata Burck of Borneo, whose

synonymy I have discussed before (Gard. Bull. Sing. 20 (1963) 272).

Symington rightly describes S$. meadiana as having 28-32 stamens,

while Burck describes his species as having 20-30. I have, however

found up to 32 in Bornean collections, though none lower than 24.

I consequently reduce S$. meadiana to S. scrobiculata.

Collections examined.—Sabah: s.n. (Wyatt-Smith) 5 miles from

Dalas, 16.8.54; s.n. (Wyatt-Smith) 5 miles from Dalas, 18.8.54;

SAN. 15012 (Wood) Mt. Silam; SAN. 29590 (Aban) Balong,

Tawau; SAN. 26895 (Bakar) Bt. Serudong, Tawau; SAN. 25848,

26895 (Meijer) Tawau river F.R.; SAN. 44226 (Pereira) J. Pahudia,

Tanjong F.R., Tawau; SAN. 47230 (Meijer) J. Ayer Panas, Tawau:

SAN. 25426 (Meijer) Bt. Mesasan, Beluran; SAN. 23135, 26589

(Meijer) Lubuk Buaya, Kinabatangan; Brunei: S. 5572 (Anderson),

Brun. 3271, 623, 52 (Ashton), SAN. 17530 (Smythies et al.)

Andulau F.R.; Brun. 2606 (Ladi) K. Temburong Machang; S. 2117

(Smythies) Bt. Teraja; S. 5723, 5760 (Smythies et al.) K. Beialong;

Brun. 740 (Ashton) K. Sekurop, Ulu Temburong; Brun. 632

(Ashton) Bt. Puan; S. 1952 (?) Bt. Telugong. Sarawak: Kep. 80491

(Wyatt-Smith) S. Dalam F.R.; s.n. (Anderson) Bt. Gondol, Bako

N.P.; SFN. 10254 (Sinclair and Kadim) G. Gaharu, Serian; SA. 462

(Majid) Bayor, Lundu, S. 1817 (Hj. Sulong) Bt. Tegarai, Limbang;

S. 9441 (Ariffin) Lundu; S. 9923 (Brunig) Bako N.P. (fr. of Vatica

sp.); Jacobs. 5544, Becc. P.B. 2538, Matang; Becc. P.B. 2889,

Sarawak; S. 263, 269 (Sipudin) Nyabau F.R.; S. 1252 (Ariffin)

Batang Ayer Betumbok F.R.; S. 2792 (Brunig) Selang F.R.;

Malaya: Kep. 50604 (Ismail) S. Nal, Kep. 65760 (Nik Yusoff)

Ulu Lebir, Kep. 50507, 50508 (Browne) S. Chalil, Kep. 37976,

37844 (Symington) Tualang Gelam, Kep. 37974 (Symington), Kep.

32820 (Walton) Lata Mak Mik Nol, Kep. 33425 (S.F.O., Kelantan)

Bt. P. Lelah, Kep. 33290 (Walton) Pulai Chondong, Kep. 52904,

52902 (Browne), Kep. 38371 (Mohd. Alias) Chebang Tongkat F.R..,

Kelantan; Kep. 53417 (Ahmad) Kelantan road, mile 37, Besut,

Trengganu; Kep. 80850 (Kochummen) 34 miles from Jerangan

police station, Dungun, Kep. 17224 (Debab), Kep. 94918 (Kochum-

men and Yong), Kep. 49840, 43128 (Symington), Kep. 48042,

47138 (D.F.O., Kuantan), Baloh F.R., Kep. 31751, 31752, 31753,

31761, 31762 (Awan) Jeram, Kep. 49845 (Symington) Dt. Goh

F.R.) Kuantan, Pahang; Kep. 34704 (S.F.O., (Kedah) Relau

F.R., Kedah; Kep. 51554 (Walton), Kep. 45753, 45754, 45755

(Abd. Taib) Kep. 12973 (Durant) Chikus F.R.; Kep. 29904,

29916, 29942, 29877 (Symington) Bubu F.R.; Kep. 40608

(Symington) Manong, Kep. 33771, 32162, 28969, 33702 (Arnot):

Kep. 33685 (Pendak), Kep. 28975 (Tachun and Arnot), Kep. 25442,

Ashton — Bornean Dipterocarpaceae 283

21405, 21406, 25885, 25415, 24632, 24519, 24478 (Symington),

Kep. 11964 (?) Keledang Saiong F.R., Kep. 24632, 24467, 24470

Tg. Tualang F.R., Perak; Kep. 68950 (Aziz Budin) Forest Research

Institute, Kepong, Selangor.

Shorea brunnescens Ashton sp. nov. Plate 28.

S. ciliatae King affinis, sed staminis 40-62 lamina coriaciore

intercostis subtus prominentus elevatis.

Gemma conferte breviter persistente alutaceo-pubescens, novellis

caduce eisdem, lamina glaber. Ramuli apices versus c. 1 mm. diam.

teretes leves nigrescentes; internodis 1-3.5 cm. longis; cicatricibus

stipularibus brevibus horizontalibus obscuris. Gemma —2.5 x 2 mm.

parva ovoidea acuta. Stipulae ignotae. Lamina 6—12 x 2.5-6 cm. late

ovata vel lanceolata coriacea basi cuneata, apice in acumen --1 cm.

longum angustum attenuata; costis lateralibus utrinsecus 9-11,

gracillimis subtus vix elevatis curvatis angulo 45°—60° exorientibus;

intercostis confertis subreticulatis evidentibus vix elevatis; costa

media supra obscura depressa subtus prominentibus sulcatis.

Petiolus 1-1.5 cm. longus 1-2 mm. diam. sicco nigrescens. Racemi

—~9 cm. longi stipitis basi in fructu c. 1 mm. diam. terminales vel

axillares angulati sparsim breviter pallide alutaceo-pubescens semel

ramosi; ramulis —1.5 cm. longis floribus secundis bracteolis ignotis.

Alabastrum -4x2 mm. anguste ellipsoideum Calyx externe

conferte pubescens intus glaber, lobis ovatis acutis internis 2 quam

externis 3 paullum brevioribus latioribus. Corolla albida petalis

lanceolatis partibus in alabatsro expositis breviter pubescentibus.

Stamina 40-62, filamentis glabris basim versus compressis apicibus

filiformibus attenuatis; antheris elliptico-oblongis glabris loculis

internis 2 quam externis 2 paullum minoribus; aristis quam antheros

brevioribus setis longis apicalibus 2 alter glabris. Ovarium stylo-

podiumque pyriformi conferte pubescenti styio brevi columellari

glabro coronatum. Fructus maturus ignotus. Calyx in fructu dein

griseo-sericeus; lobis aliformibus exterioribus 3 quam aliis 2

longioribus. Nux ovoidea conferte breviter pubescens.

Collections —Sarawak: S. 13179 (Smythies) G. Berumput;

S. 15220, 15218 (Holotypus in Herb. Kew) (Smythies) Bt. Gaharu;

Jacobs 5544, G. Matang; S. 19373 (Ashton) Bt. Naoung, Muput

Kanan, Anap; S. 18194 (Ashton) Ulu Tubau; S. 7995, 10179

(Bojeng) G. Gading; S. 14910, 15019, 14904, 14988, 14928 (Rosli).

tree no. 87, 111 (Asah), S. 362 (Omar), S. 45 (Ahmad Kabir),

Sinclair and Kadim 10192, S. 15714 (Galau), S. 15773 (Jugah),

S. 14931 (Anderson), S. 11057 (Ghazalli), S. 8943, 8945 (Yakup),

S. 15172, 15763 (Rosli and Jugah) Semengoh F.R.

This species differs from S. ciliata King of Malaya principally in

the larger number of stamens, though small but consistent differ-

ences in lamina characters are also noticeable. These two species

are therefore allied iti the same way as S. sumatrana (V. Sl. ex

Foxw.) Sym. of Sumatra and Malaya is to S. seminis (De. Vr.)

Sloot of Borneo and the Philippines, S. /umutensis Sym. of Malaya

is to S. inappendiculata Burck of Borneo, and S. astylosa Foxw. of

the Philippines is to $. domatiosa Ashton of Borneo. I have chosen

284 Gardens’ Bulletin, Singapore — XXII (1967)

to consider these as pairs of species rather than as geographical

subspecies of single species partially as the range in the number

of stamens in each taxon of a pair does not overlap, partially

as there are also other, though less clearly defined, differences and

partially also in order to avoid upsetting presently accepted

nomenclature in the absence of experimental or other conclusive

evidence. It appears that a change in the mean numbex of stamens

in separated parts of a population of species in section Shoreae

is commonly one of the earliest evolutionary divergences to occur.

In S. foxworthyi Sym. for example the number varies between

33-41 in Malaya and between 32-34 in Borneo; I do not feel this

difference is sufficient to merit being given taxonomic status.

The ‘instability’ of this character is not altogether surprising in

view of their centrifugal origin.

Though S$. brunnescens is common in Semengoh F.R. at low

altitudes, elsewhere it is confined to hills, being found typically

along ridges between 600—1,000 m., in this resembling its Malayan

counterpart. The present species is known to me up to the present

only from Sarawak, where it is widespread west of the Baram

drainage.

abetiee ned yelte dip tet ne - tee ogi at oe le

Shorea lunduensis Ashton sp. nov. Plate 29.

S. collinae Ridl. affinis sed lamina minus coriaceori lobis calycis

in fructu inaequalibus longioribus differt.

Ramuli gemmaeque breviter plane fugace pallide alutaceo-

pubescentes, laminae glabrae. Ramuli apices versus c. 3X2 mm.

diam. primo compressi dein nigrescentes nitentes; internodis

1-3.5 cm. longis; cicatricibus stipularibus c. 3 mm. lengis pallidis

prominentibus ascensis. Gemma —5 x3 mm. ovoidea acuta niger.

Stipulae ignotae. Lamina 14-24x6-15 cm. magna ovata vel

elliptica basi late cuneata vel subcordata, apice in acumen —1 cm.

Icngo attentuata; costis lateralibus utrinsecus 11-15 supra obscuris

subtus prominentibus angulo 40°-55° exorientibus basi excepta

intercostis sinuatis conferte scalariformibus; costa media supra

vix elevata subtus prominenti. Petiolus 2—3.5 cm. longus c. 2 mm.

diam. crassus sicco nigrescens. Racemi —12 cm. longis stipitis basi

in fructu —2 mm. diam. terminales vel axillares subteretes conferte

breviter fasciculate alutaceo-pubescentes semel vel bis ramosi;

ramulis —3 cm. longis; bracteolis -4 x 2 mm. ovatis conferte breviter

pubescentibus fugacibus. Flores secundi. Alabastrum —10 x3 mm.

anguste ovoideum calyce expansa. Calyx partibus in alabastro

expositis sericeis, lobis anguste ovatis subacutis internis 2 quam

exterioribus 3 minoribus latioribus. Corclla cremia, petalis :

linearibus partibus in alabastro expositis conferte pubescentibus.

Stamina 47-52 filamentis compressis attenuatis glabris; antheris

oblongis glabris apicem versus attenuatis; aristis quam antheras

longioribus conferte setosis. Ovarium stylopodiumque pyriforme

conferte pubescente stylo columellari coronatum. Fructus maturus

ignotus, novello sparsim breviter pubescens; lobis calycis externis

3, inaequalibus quam internis 2 longioribus spatlats subacutis.

Nux apiculo —1 mm. longo attenuata.

Ashton — Bornean Dipterocarpaceae 285

Collections —Sarawak: S. 15396 (Anderson et alia) G. Lundu

(Holotypus in Herb. Kew); S.A. 587 (Baharol) S. 10172, 7984, 7968

(Bojeng), s.n. (Anderson), S. 59 (Browne), Gunong Gading F.R.;

S. 15502 (Anderson et al.) Bt. Gebong; S. 15482 (Anderson) Bt.

Seburan, Bau.

Belonging to the type section and subsection and differing from

S. collina Ridl. of eastern Malaya principally in the aliform fruit

calyx, this species appears to be confined to Mixed Dipterocarp

forest in West Sarawak.

Shorea agami Ashton; infraspecific variation.

Two distinct forms of this species (Ashton, Gard. Bull.. Sing. 19

(1962) 270), based solely on the size of the lamina, are discernable,

and appear to merit subspecific status.

subsp. agami. (Ashton, id. 271, Pl. 9)

Lamina 10-15 x 6—10 cm. late ovata.

Collections (others than those cited in the original description).

Sarawak: S. 10077 (Yong) Baram; S. 1454 (Gusni) Miri; S. 1936,

1462 (Dan) Riam road, Miri; S. 1430 (Keram) Bakam, Miri;

S.-1489 (Suib) Limbang road, Marudi; S. 1757 (Drahman) Bt.

Meringgit, Lawas.

subsp. diminuta Ashton subsp. nov. Plate 30.

Lamina 4.5—10 x 2.5-4 cm. anguste cvata.

Collections.—Sarawak: S. 25027 (Sibat) Bt. Mersing, Anap

(Holotypus in herb. Kew); S. 22335 (Suib) Bt. Leran, Niah;

S. 17771 (Ashton), S. 23961 (Wright), S. 23860 (Jugah) Bt. Raya,

Kapit; S. 15227 (Jugah) S. Sabal Tapang, Serian.

This latter subspecies is also widespread in western Kalimantan;

the ranges of the two subspecies appear not to overlap.

Shorea cordata Ashton sp. nov. Plate 31.

S. hypochrae Hance affinis sed lamina maiore tect lepidota

basim versus cordata.

Ramuli apices versus petioli conferte, lamina costis lateralibus

subtus sparsim caduce pubescentes, gemmae stipulaeque per-

sistente pubescentes. Ramuli apices versus c. 3X2 mm. diam.,

paullum compressi fuscescentes terescentes levescentes vel ruguli;

internodis —5 cm. longis; cicatricibus stipularibus horizontalibus

latis pallidis prominentibus. Gemma -7x4 mm. ellipscidea

compressa acuta. Stipula -12x6 mm. elliptica obtusa. Lamina

8-15 x 5.5-10.5 cm. oblonga vel obovata, basi cordata vel rare

acuta, apice obtusa vel retusa vel in acumen —8 mm. longum abrupte

attenuata; costis lateralibus utrinsecus 15-18 subtus prominentibus

apicem versus angulo 30°-40° basim versus angulo -100°

exorientibus; intercostis gracilibus conferte scalariformibus; costa

media supra depressa subtus prominenti. Petiolus 13-25 mm.

longus c. 2 mm. diam. teres saepe rugulosus sicco nigrescens.

286 Gardens’ Bulletin, Singapore — X Xf1 (1967)

Racemi —12 cm. longi stipitis basi in fructu —2 mm. diam.

terminales vel axillares laxi compressi vel costati persistente

alutaceo-puberulentes semel ramosi; ramulis —5 cm. longis —5 flores

plus minus secundis gerentibus; bracteis -10 x3 mm. lanceolatis

acutis puberulentibus fugacibus; bracteolis -10 x5 mm. ellipticis

obtusis puberulentibus subpersistentibus. Alabastrum —105 mm.

fusiforme. Calyx externe sericeus, intus glaber; lobis anguste

deltoideo-lanceolatis obtusis exterioribus 3 quam interioribus 2

paullum maioribus. Petala oblongo-lanceolata externe sericea intus

glabra. Stamina 15 verticillis 3 inaequalibus vel externis 16

vestigialibus; filamentis angustis compressis attenuatis; antheris

elliptico-oblongis; aristis quam antheras -6-plo longoribus

filiformibus atenuatis. Ovarium ovoideum glabrum, stylo columel-

lari quam ovarium 2-plo longiore attenuato basim versus sericeo

apicem versus obscure trifurcato. Fructus glaber. Pedicellus in

fructu obscurus in calycem decurrens. Lobi calycis longiores 3,

-13x2.5 cm. lorati obtusi basim versus —8 mm. lati attenuati,

partibus basalibus -12 x 12 mm. orbicularibus incrassatis saccatis;

lobi breviores 2, -6.5x0.8 cm. angusti lanceolati acuti, basim

versus ut in lobi longiores. Nux -14x10 mm. globosa calyce

obscurata.

Collections —Sarawak: S. 13183 (Smythies) G. Berumput; SA.

678 (Egon) outside Semengoh F.R. (Holotypus in Herb. Kew);

S. 7988, 7986, 10177 (Bojeng) G. Gading; S. 19306 (Ashton)

Ulu Mayeng, Kakus.

Apparently allied to S. hypochra Hance of Malaya, Thailand,

Cambodia and Cochin China, S. cordata differs mainly in the larger

cordate glabrescent lamina lacking the characteristic pale yellowish

lepidote undersurface of the former species; in this it resembles

immature specimens of S. hypochra. S. cordata is a rare tree,

found to date in Central and West Sarawak; it can be of immense

size.

Shorea polyandra Ashton sp. nov. Plate 32.

Ramuli petioli lamina subtusque persistente rufo-lepidoti,

gemmae stipulaeque persistente breviter rufo-puberulentes. Ramuli

apices versus c. | mm. diam. graciles teretes griseo-fuscescentes

leves vel minute striati; internodis —2 cm. longis; cicatricibus

stipularibus brevibus horizontalibus obscuris. Gemma —2 x1 mm.

parva ellipsoidea subacuta. Stipula —5 x 2 mm. lanceolata subacuta

fugax. Lamina 8-13 x 3-5 cm. lanceolata chartacea undulata basi

cuneata apice in acumen —2 cm. longum gracile gradatim attenuata;

costis lateralibus utrinsecus 11-14 gracilibus subtus elevatus angulo

40°-60° exorientibus; intercostis scalariformibus gracilibus; costa

media gracili supra applanata subtus prominenti. Petiolus 14—20

mm. longus c. 1 mm. diam. gracilis. Racemi —6 cm. longi, stipitis

basi in fructu —2 mm. diam., terminales vel axillares teretes

conferte rufo-pubescentes semel ramosi; ramulis —1 cm. longis flores

—3 plus minus distichos gerentibus; bracteolis -4 x 3 mm. ellipticis

subacutis rufo-pubescentibus caducis. Alabastrum —5 x 4 mm. late

ovoideum. Calyx partibus in alabastro expositis seriuces, lobis

suborbicularibus subaequalibus. Petalae oblongae obtusae partibus

Ashton — Bornean Dipterocarpaceae 287

in alabastro expositis conferte pubescentibus. Stamina 102-107

plurima subaequalia; filamentis basim versus compressis apicem

versus aitenuatis; antheris quam filamentos —2-plo longioribus

anguste oblongis loculis 2; aristis filiformibus attenuatis quam

antheris paullum brevioribus conferte setosis. Ovarium ovoideum

conferte pubescente stylo brevi late columellari trifurcato

pubescenti apice excepto attenuatum. Pedicellus in fructu -1 mm.

longus —2 mm. diam. brevis. Calyx in fructu basim versus sparsim

rufo-pubescens glabrescens; lobis longioribus 3, -8x1.4 cm.

spatulatis obtusis vel subacutis basim versus —8 mm. latis attentuatis

partibus basalibus -10x8 mm. tuberculatis saccatis incassatis;

brevioribus 2, —5 x0.5 cm. linearibus acutis basim versus ut in

lobis longioribus. Nux —30 x 13 mm. anguste ovoidea acuta conferte

rufo-pubescens.

Collections Sarawak: S. 10171 (Holotypus in herb. Kew) 10174,

10189, 7977, 7979 (Bojeng) G. Gading F.R. Sabah: SAN. A3959,

A3465, 15265 (Wood) Kalabakan, Tawau; SAN. 16478 (Wood)

Quoin Hill, Tawau; SAN. 16342 (Wood and Kapis) Pangi, Tenom.

Kalimantan: Kostermans 13302, Mt. Medadam, N. of Sengkulirang;

b.b. 21976 (Zwaan), 21974, 21975 (Henar) Djalepat, P. Laut;

b.b. 31167, 27186 (Chatihoe) Seblimbingan, P. Laut; b.b. 31559

(Kadiri), 12345 (Zwaan), 12901 (Hildebrand) Sei Paring, P. Laut;

b.b. 25158 (Soeli), b.b. 26404, 26405, 26406 (Lammers) Tg. Miri,

P. Laut; b.b. 12239 (Ramli) Sei Taip, P. Laut; b.b. 23888 (Henar)

Seratah, P. Laut; b.b. 20165 (Mahboel) Batoeng, Kandangan;

b.b. 20455 (Nandika) Petarikan, Kota Waringin; b.b. 23349, 23350,

23351 (Dachlan) Sakadoea, Martapoera; b.b. 23353 (Dachlan) Taha

Doea, Martapoera; b.b. 14629 (Pohan) Katak, E. Kutei; b.b. 17557

(Atjil) Sotek, Balikpapan; b.b. 15758, 15777, 15779 (Hamid)

Sebulu, W. Kutei; b.b. 26050 (Soepomo) Mendan, W. Kutei:;

b.b. 23080, 23081 (Dijen) G. Karap, W. Kutei; b.b. 23072 (Dijen)

Lebak Njur, W. Kutei; b.b. 22937 (Nandika) Benua Baru, W.

Kutei; b.b. 2049 (de Jong) Pait, W. Borneo; b.b. 22882, 22610

(Walinjan) Bt. Rompin, Sintang; b.b. 25100, 25101, 25099

(Budding) Tjatit B. Tengkoejoeng, Malawi.

This striking species is in fruit very similar to S$. gidbosa

Brandis and S. faguetiana Heim, but the flowers are so different

from other numbers of Section Richetioides that it justifies the

creation of a new infrageneric group to contain it. The anthers,

though shaped as in Section Anthoshoreae are yet 2-locular,

a character unique to Section Richetioides. In wood anatomy too

Gottwald considers it to be typical of the latter section (in litt.),

though Murthy tells me that there are small differences, as follows.

Character Typical sect. Richetioides S. polyandra

Silica Absent ; Absent, but presence of

crystals in rays very

much like silica distri-

bution.

Calcium oxalate crystals Present but not Abundant

abundant

Rays canals Fairly common Rare, though present.

Chambered crystals in Frequent, alawys Rate.

paranchyma present

288 Gardens’ Bulletin, Singapore — XXII (1967)

I therefore consider that it merits the creation of a new sub-

section within Section Richetioides rather than a separate section

altogether. The description of Section Richetiodes thus needs

further amplification.

Section Richetioides Heim.

Type: S. faguetiana Heim.

Anthers with 2 pollen sacs; appendage to connective erect,

filiform, more or less scabrous or setose. Style shorter than ovary.

Stipules and bracts minute, fugaceous; lamina nervation usually

more or less reticulate, pellucid, midrib raised or sunken above,

evident.

Subsection 1. Richetioides.

Flowers usually small, bud fusiform; stamens 15 (rarely 10),

in 3 verticils; filaments broad at base, frequently gibbous, tapering

more or less abruptly medially, filiform below anther; anthers

broadly oblong to subglobose; appendage to connective longer

than anther, more or less scabrous towards apex; ovary with

stylopodium, shortly tomentose, with a slender columnar style.

Subsection 2. Polyandrae Ashton, subsect. nov.

Type: S. polyandra Ashton

Alabastrum -5X4 mm. magnum late ovoideum. Staminal 8,

sine verticillis; filamentis basim versus compressis apicem versus

gradatim atenuatis; antheris anguste oblongis; aristis quam antheras

paullum brevioribus conferte setosis. Ovarium ovoideum sine

stylopodio conferte longe pubescente stylo brevi lato prominenti

trifurcato.

Shorea alutacea Ashton sp. nov. Plate 33.

S. gibbosae Brandis, S. faguetianae Heim affinis sed lamina

maiore costis lateralibus utrinsecus 14-19 petiolo 6-10 mm. longo

breviore costa media subtus petiolo ramulique conferte persistente

plane pallide ochraceo-pubescentibus.

Ramuli stipulae gemmae petioli costae mediaeque breviter con-

ferte persistente alutaceo-pubescentes, lamina subtus sparsim

eaedem. Ramuli apices versus c. 2 mm. diam. teretes; internodis

1-4.5 cm. longis; cicatricibus stipularibus c. 1.5 mm. longis

horizontalibus. Gemma c. 2 x 1.5 mm. ovoidea acuta parva. Stipula

—8 x3 mm. anguste deltoidea acuta magna. Lamina 15—22 x 6-8

cm. ovata vel lanceolata undulata chartacea basi cordata apice in

acumen -—1 cm. longum attenuata; costis lateralibus utrinsecus

14-19 curvatis gracilibus supra anguste depressis subtus pro-

minentibus basim versus angulo -—100° apicem versus angulo

60°-70° exorientibus; intercostis subscalariformis — gracilibus

obscuris; costa media supra depressa subtus prominenti tereti.

Petiolus 6-10 mm. longus c. 2 mm. diam. sicco rugosus. Racemi

—15 cm. longi, stipitis basi in fructu —2 mm. diam. terminales vel

axillares teretes recti conferte breviter pallide alutaceo-pubescentes

semel vel bis ramosi; ramulis —2.5 cm. longis; bracteolis —-4 x 3 mm.

Ashton — Bornean Dipterocarpaceae 289

ovatis acutis conferte breviter alutaceo-pubescentibus fugacibus.

Flores secundi. Alabastrum -6x3 mm. ellipsoideun. magnum.

Calyx partibus in alabastro expositis conferte pubescentibus, lobis

late ovatis subaequalibus internis 2 breviter acuminatis quam

externis 3 marginem versus characeoribus basim versus angustiori-

bus. Petala lanceolata badia partibus in albastro expositis pube-

scentibus, statu aperto basim versus imbricatis apicem versus

contortis. Stamina 15, verticillis 3 inaequabibus; filamentis basim

versus latis compressis attenuatis apicem versus filiform:bus;

antheris late oblongis loculis 2; aristis quam antheros 3—4-plo

longioribus gracilibus glabris. Ovarium stylopodiumque pyriforme

conferte pubescente stylo columellari glabro attenuatum; stylo

stylopodium aequanti. Pedicellus calyxque in fructu sparsim

breviter caduce pubescens. Pedicellus c. 1 mm. longus brevis.

Lobi calycis longiores 3, -8x1.8 cm. spatulati basim versus.

—4 mm. lati attenuati, partibus basalibus —-7 x 7 mm. ovatis saccatis

incrassatis; lobis brevioribus 2, -61.2 cm. alter ut in lobis

longioribus. Nux —2.2 x1 cm. anguste ovoidea conferte breviter

plane alutaceo-tomentosa apiculo brevi attenuata.

Collections —Sarawak: S. 7976, 7975, 10170 (Holotypus in Herb..

Kew) (Bojeng) Gunong Gading F.R.

A member of section Richetioides Heim and allied to S.

faguetiana Heim and S. gibbosa Brandis among others, the lamina

differences quoted in the preceding diagnosis serve at once to

distinguish it.

S. alutacea is known at present only on the granodiorite Gading

mountains of West Sarawak, where it is scattered below 400 m. alt.

Shorea bakoensis Ashton sp. nov. Plate 34.

S. laxa Sloot. affinis, sed lamina oblongo-lanceolata basi obtusa,

petiolo 10-12 mm. longo c. 2 mm. diam., lamina petiolo ramuloque

glabro differt.

Laminae petioli ramulique glaber. Ramuli apices versus c. 2 mm.

diam. primo leves pallide fusci lenticellis nigris punctati dein

fuscentes exfoliati; internodis c. 1.5 cm. longis; cicatricibus.

stipularibus brevibus horizontalibus. Stipula gemmaque ignota.

Lamina 13-18 x5—6 cm. anguste oblonga vel lanceolata coriacea

basi obtusa apice in acumen —1 cm. longum angustum attenuata;

costis lateralibus utrinsecus 9-10 curvatis subtus prominentibus

angulo 55°—65° exorientibus; intercostis gracilibus subtus vix

elevatis subscalariformibus; costa supra applanata subtus pro-

minenti. Petiolus 10-12 mm. longus c. 2 mm. diam. brevis crassus

sicco niger. Flores racemique ignoti. Peticellus in fructu brevis.

obscurus. Lobi calycis -4x3 mm. subaequales oblongi obtusi

incrassati saccati patentes externe sparsim persistente alutaceo-

sericel. Nux —25x14 mm. ellipsoideo-cylindrica conferte plane

persistente alutaceo-pubescens apice in stylum -1 mm. longum

attenuata.

Collections Sarawak: S. 17502 (Brunig) Bako National Park

(Holotypus in Herb. Kew).

290 Gardens’ Bulletin, Singapore — XXII (1967)

This rare species is only known from a few localities in W.

Sarawak. It somewhat resembles S. laxa Sloot. and S$. obovoidea

V. SI., but differs in the glabrous leaf wth obtuse base and

relatively shorter petole, as well as in the ellpsoid — cylindric, not

obovoid, nut.

Shorea Cuspidata sp. nov. Plate 35.

S. gibbosa Brandis, S. faguetana Heim. affinis, sed lamina

5-9 X 2-6 cm. minori glabra costis lateralibus utrinsecus 5—7.

Gemma stipulaque breviter persistente pubescens, ramulus caduce

pubescens, lamina glaber. Ramuli apices versus c. 1 mm. diam.

graciles teretes fusci minute striati; internodis 1-2.5 cm. longis.

Gemmae c. 1 mm. longae parvae ovoideae. Stipulae parvae fugaces.

Lamina 5-9 x 2-6 cm. late ovata subcoriacea basi cuneata apice in

acumen —1.5 cm. longum subcaudatum attenuata; costis lateralibus

utrinsecus 5—7 gracilibus subtus vix elevatis curvatis angulo 40°—50°

exorientibus; intercostis reticulatis obscuris; costa media gracilt

supra applanata vel paullum elevata subtus prominenti. Petiolus

7-11 mm. longus c. 1 mm. diam. gracilis sicco niger. Racemi —9

cm. longi, stpitis basi in fructu —1.5 mm. diam., terminales vel

axillares teretes graciles conferte plane obrevite: alutaceo-

pubescentes semel ramosi; ramulis —1.5 cm. longis flores secundos

gerentibus. Alabastra —3 x 2 mm. parva ellipsoidea. Calyx partibus

in alabastro expositis pubescentibus sepalis late ovatis parvis

subaequalibus acutis, internis 2 quam externis 3 latioribus marginem

versus chartaceoribus. Petala pallide lutea lanceolata prominente

contorta partibus in alabastro expositis conferte pubescentibus

basibus connatis. Stamina 15, verticillis 3 inaequalibus; filamentis

basim versus compressis apicem versus filiformibus attenuatis;

antheris loculis 2 ellipsoideis; aristis quam antheras c. 14-plo

longioribus gracilibus dimidiis apicalibus pubescentibus. Ovarium

stylopodiumque pyriforme sericeum stylo brevi columellari glabro

coronatum. Pedicellus calyxque in fructu’ breviter sparsim

pubescens. Pedicellus c. 1 mm. longus brevis. Lobi calycis longiores

3, -5x1.5 cm. late spatulati obtusi basim versus —3 mm. lati

attenuati; partibus basalibus -5 mm. latis saccatis incrassatis; lobis

brevioribus 2, -4x1 cm. aliter ut in lobis longioribus. Nux

2.5 x 1.5 cm. obovoidea conferte breviter plane alutaceo-pubescens

in apice mucronata attenuata.

Collections Sarawak: S. 15258 (Holotypus in Herb. Kew),

15715 (Galau), S. 9345, 9345A (Yacup), S. 11056 (Ghazalli),

S. 15179 (Rosli and Jugah), S. 14999, 15718 (Rosli), Tree No. 603

(Asah), S. 10033 (Zen) Semengoh F.R.; S. 15211 (Smythies) Bt.

Gaharu; S. 9607, 9609, 9615 (Ariffin) G. Tamin, Lundu; S. 6513

(Yakup) Gunong Tabut, Ulu Sebuyau; S. 9483, 9487, 9487A (Saidi)

Santubong; S. 364 (Ariffin) Bt. Sebandar, Lundu; S. 10289, 10289A

(Ardzi) Bako N.P.

This species is allied to S. faguetiana Heim and S. gibbosa

Brandis in section Richetioides Heim, but differs in the fewer

nerves, glabrous leaf and smaller size of all parts. Furthermore

Ashton — Bornean Dipterocarpaceae 291

the bark surface configuration appears to be unique in the section:

at first smooth, chocolate-brown with grey lichen-mottling;

becoming regularly surface-fissured, fissures -8 mm. broad, —4 mm.

deep, V-section, anastomosing, separated by short sharp zig-zag

ridges; ridges flaking evenly in small oblong pieces leaving an

overall smooth tawny prominently scroll-marked surface beneath;

in old trees further flaking irregularly. This surface-fissure con-

figuration evokes that of Shorea biawak Ashton of section Shoreae

subsection Barbatae Sym. ex Ashton (see my Manual of the

Dipterocarp Trees of Brunei State, p. 129, pl. 37), but in that

species the flaking is chunky and bears no relation to the surface

fissuring.

S. cuspidata formerly bore the code-letter names Shorea H,,

Shorea K,, and Shorea Z;. It is found on yellow sandy soils in

Mixed Dipterocarp forest. It is widespread in Sarawak west of the

Lupar, and is frequently the most abundant representative of the

section.

Shorea iliasii Ashton sp. nov. Plate 36.

Species lamina ut in S. faguetiana Heim, sed 19-25 9-12 cm.

maiore costis lateralibus utrinsecus 12-14 petiolo 22-30 mm. longo

c. 3 mm. diam. differt.

Novelli caduce griseo-sericei. Ramuli apices versus c. 2 mm.

diam. teretes griseo-fusci saepe chartaceo-exfoliati; internodis

—4 cm. longis; cicatricibus stipularibus brevibus horizontalibus.

Gemma stipulaque ignota. Lamina 19-25 x 9-12 cm. obiongo-ovata,

basi obtusa sed breviter decurrenti subaequali, apice in acumen

—8 mm. longum breve attenuata; costis lateralibus utrinsecus 12-14

curvatis subtus prominentibus angulo 50°—75° exorientibus; inter-

costis remote scalariformibus; costa media supra applanata vel

paullum elevata subtus prominenti. Petiolus 22-30 mm. longus

c. 3 mm. diam. crassus teres sicco nigrescens. Racemi —10 cm.

longi, stipitis basi in fructu c. 3 mm. diam., teretes caduce griseo-

sericei. Alabastrum —4 x 2 mm. parvum fusiforme. Calyx partibus

in alabastro expositis conferte puberulentibus, lobis externis 3

Ovatis acutis internis 2 quam externis minoribus breviter fimbriatis

basim versus angustioribus. Petala cremia lineara obtusa partibus

in alabastro expositis conferte pubescentibus. Stamina 15 verticillis

2 inaequalibus, internis 5 quam aliis paullum longioribus, filamentis

basim versus latis sub antheris subglobosis filiformibus attenuatis;

aristis quam antheras c. 14-plo longioribus apices versus sericeis.

Ovarium anguste ovoideum pubescente stylo brevi glabro

attenuatum. Calyx basim versus sparsim griseo-sericeus, aliter

glaber; lobis longioribus 3, -8x2.2 cm. spatulatis obtusis vel

subacutis basim versus —-4 mm. latis attenuatis; partibus basalibus

—6x5 mm. saccatis incrassatis prominenti tuberculatis; lobis

brevioribus 2, —5.5x1.2 cm. aliter ut in lobis longioribus. Nux

-10x8 mm. ovoidea acuta conferte persistente griseo-sericea.

Collections Sarawak: S. 15102 (ilias) Ulu Segan, Bintulu

(Holotypus in herb. Kew); SA. 310 (Spurway) Long Sangga,

Baram; S. 1455 (Brunig) Ulu Skroh, Kapit; S. 15842 (Ilias and

292 Gardens’ Bulletin, Singapore — XXII (1967)

Johnson) Similajau F.R., Labang; S. 22063, 22041 (Ashton) N.

Sepulau, Segan; S. 22258, 22259 (Suib) Bt. Raya; S. 22304 (Suib)

Bt. Wong, Pelagus.

Differing from S. faguetiana Heim in the larger size of all parts

and the greater number of lamina nerves, this species is known

so far from the Lower Rejang to the Kemena drainage in Central

Sarawak.

Shorea macrobalanos Ashton sp. nov. Plate 37.

S. longiflorae (Brandis) Sym. affinis, sed lamina 19-37 x 8-15 cm.

latiore nervis angulis 45°-65° exorientibus basi excepta petiolo

1.8-3.7 cm. longo differt.

Gemma stipulaque persistente alutaceo-pubescens aliter glaber.

Ramuli apices versus c. 4 mm. diam. primo teretes dein verrucosi

pallide alutacei; internodis —7 cm. longis; cicatricibus stipularibus

c. 3 mm. longis obscuris ascendentibus. Gemma —2 x 2 mm. ovoidea

acuta. Stipula ignota. Lamina 19-37 x 8-15 cm. oblonga coriacea,

margine revoluto, basi cordata, apice in acumen brevi abrupte

attenuata vel obtusa; nervis utrinsecus 12-16 subtus prominentibus

angulo 45—°65° exorientibus basi excepta; intercostis gravilibus

remote subreticulatis; costa media supra applanata_ subtus

prominenti. Petiolus 1.8-3.8 cm. longus c. 4 mm. diam. sicco

nigrescens. Flores ignoti. Racemi —32 cm. iongi, basi stipitis in

fructu —S5 mm. diam., longi terminales vel axiilares paullum

compressi glabri pallide fusci rugescentes semel vel bis ramosi.

Fructus sessilis glaber; lobis calycis -8 x8 mm. subaequalibus

Ovatis saepe marginem versus undulatis expansis. Receptaculum

—1 cm. diam. -8 mm. longum magnum. Nux —5 x 2.5 cm. magna

oblonga breviter apiculata.

Collections Sarawak: S. 13192 (Smythies) G. Berumput (Holo-

typus in Herb. Kew); S. 10324 (Brunig), S. 11726 (Hj. Bujang)

Semengoh F.R.; S. 13712 (Ilias Paie) G. Pueh.

Undoubtedly closely allied to Shorea longiflora (Brandis Sym. of

Section Richetioides Heim, this West Sarawak species differs in

the relatively longer petiole, relatively broader lamina and larger

parts generally, and in the much larger size the tree attains.

Shorea mujongensis sp. nov. Plate 38.

S. gibbosae Brandis affinis sed lamina maiore subtus glabra

margine prominente revoluto petiolo giabrescenti differt.

Ramuli apices versus gemmae petiolique breviter alutaceo-

pubescentes dein glabrescentes. Ramuli apices versus c. 2 mm.

diam. teretes ruguli lenticellis pallidis minutis punctati; internodis

—3 cm. longis; cicatricibus stipularibus brevibus obscuris. Gemma

—2 x 1mm. conica. Stipula ignota. Lamina 7-14 x 2.8-5.5 cm. ovata

margine revoluto basi obtusa vel late cuneata, apice in acumen

-8 mm. longum attenuata; costis lateralibus utrinsecus 9-13

gracilibus subtus prominentibus curvatis angulo c. 50° exorientibus;

intercostis scalariformibus obscuris haud elevatis; costa media

Supra applanata subtus prominente tereta. Petiolus 10-16 mm.

Ashton — Bornean Dipterocarpaceae 293

longus c. 2 mm. diam. teres sicco nigrescens. Flores ignoti. Racemi

—6 cm. longi, stipitis basi -2 mm. diam. semel ramosi teretes rugull

breviter plane alutaceo-pubescentes. Pedicellus in fructu —2 mm.

longus -—1 mm. diam. puberulens. Calyx in fructu’ sparsim

puberulens dein glabrescens; lobis longioribus 3, --7 1.5 cm.

spatulatis obtusis basim versus c. 4 mm. latis attenuatis partibus

basalibus -6 x 5 mm. tuberculatis incrassatis; lobis brevioribus 2,

—4.5x0.7 cm. aliter ut in lobis longioribus. Nux —2.2 x0.7 cm.

anguste ovoidea conferte breviter pallide alutaceo-pubescens.

Collections—Sarawak: S. 19038 (Ashton) Base of Bt. Temedu,

Hose Mountains (Holotypus in Herb. Kew); S. 19993, 19603

(Ashton) Carapa Pila.

At present this magnificent tree is known only from the

inaccessible woods on the middle slopes of the Hose mountains.

The lustrous leaves make its pale crown easily identifiable in

aerial photographs.

Shorea hemsleyana King ex Foxw., Mal. For. Rec. 10 (1932) 167:

Notes on infraspecific variation.

Symington (Gard. Bull. S.S. 7 (1933) 129) revised the nomen-

clature and synonymy of this Malayan species. He reduced Shorea

grandiflora Brandis (J. Linn. Soc. Bot. 31 (1895) 93) to it, based

on Haviland 2121, with flowers and fruits, from the Kuching area

of Sarawak. Until recently this remained the only Bornean

collection; but now that more material has been collected from

Borneo it has become apparent that the type specimen of S.

grandiflora bears leaves and twigs larger than they are on mature

trees of this species in Borneo, inferring that it was collected

from an immature specimen. These parts are considerably smaller

than is normal for the species in Malaya, and justify the Bornean

form’s resurrection as a separate subspecies:

ssp. hemsleyana

Lamina 14-35 x 6-15 cm. Petiolus 1.4—2 cm. longus. Racemus

—25 cm. longus.

Collections examined.—Malaya: Kunstler 6670, 7562, Larut

Perak, Scortechini 1653, Perak (Syntypes); Curtis 2512, 3739,

Province Wellesley; Kep. 16814, Lumut F.R.; Dindings; Kep

75983, 80220, Kepong plantations; Kep. 16826, Bruas F.R.,

Dindings; Kep. 12018, Parit F.R., Tronoh, Perak.

ssp. grandiflora (Brandis) Ashton, stat. nov.

—S. grandiflora Brandis, J. Linn. Soc. Bot. 31 (1859) 93.

Omnes partes ut in ssp. hemsleyana sed lamina 10-23 x 4-11

cm., petiolo 6-12 mm. longo, racemis —4 cm. longis differt.

Collections Sarawak: Haviland 2121; near Kuching (Holotypus

in herb. Kew); S. 18860 (Ashton), S. 12612 (Smythies) Bt. Mero-

yong, Sempadi F.R., Lundu; S. 6503 (Yakup) G. Menuku, Sebuyau

(fruit excepted); S. 21585 (Ashton) G. Matang; S. 22333 (Suib)

Bt. Leran, Niah; S. 6236 (Brunig) G. Pueh F.R.

294 Gardens’ Bulletin, Singapore — XXII (1967)

The identity of Shorea johorensis Foxw.

This species was described (Mal. For. Rec. 10 (1932) 236) from

a single specimen, Kep. 5992 (Bain) Gunong Panti, Johore, now

in the Kew herbarium. It bears somewhat immature fruits.

Symington (Mal. For. Rec. 16 (1943) 72) regarded it as a dubious

species, tentatively synonymous with S. singkawang Miq., notwith-

standing that species has vestigial fruit calyx lobes, whereas the

fruit of Kep. 5992 is prominently winged.

The material of S. singkawang now available shows that the

fruit of Kep. 5992 is definitely not of this species; though it is

detached, Foxworthy’s photograph of the fresh material with

attached fruits is clear enough to be unequivocal. Kep. 5992 is

however identical with Kep. 35758 and 35759, from the same

locality, cited by Symington in his original description of S.

leptoclados (Gard. Bull. S.S. 10 (1939) 376). The Holotype of

S. leptoclados, Kep. 30533 (Smith) Batu Apoi, Brunei in the

Kepong herbarium, differs, as does most, but not in fact all,

Bornean material, from Malayan material in possessing 3-6 pairs

of pale scale-like domatia in the axils of the first 3-6 pairs of

basal nerves. This, the only difference, is not only smail but

inconstant, and I feel compelled, therefore to reduce S. leptoclados

to Foxworthy’s species.

Shorea uliginosa Foxw. reinstated.

This species was described (Mal. For. Rec. 10 (1932) 210) from

Kep. 7938 (Foxworthy) Bukit Cheraka F.R., Selangor, a flowering

collection, represented both at Kepong and Kew. Symington (Gard.

Bull. S.S. 10 (1939) 372) reduced it to a variety of S. rugosa Heim,

stating that ‘S. uliginosa has slightly larger leaves with relatively

longer petioles, and smaller, less hairy fruits’. He concludes ‘I think

a reasonable and satisfactory way of treating the problem is to

consider S. uliginosa Foxw. as a variety of S. rugosa Heim’.

Symington noted that the two varieties might occur in different

habitats; they do, Foxworthy’s species being confined to peat

swamps and Heim’s to leached sandy soils. Though the flowers,

inflorescences, tomentum and bark appear similar in the two taxa

there are other differences besides those mentioned by Symington.

The lamina of uliginosa is chartaceous and prominently concave,

whereas that of rugosa is applanate and thickly coriaceous. The

habit of uliginosa with its oblong diffuse crown with large flopping

emi-pendant leaves is quite different from the dense hemispherical

crown of rugosa. I consequently consider that these two taxa are

specifically distinct, and reinstate S. uliginosa Foxw.

Shorea carapae Ashton sp. nov. Plate 39.

S. coriaceae Burck affinis sed lamina 13-18 7-10 cm. costis

lateralibus utrinsecus 11-13 subtus crassis prominentibus petiolo

breviter conferte plane persistente alutaceo-sericeo differt.

Ramuli gemmae stipuli petiolique persistente conferte plane

alutaceo-sericei, costis subtus costa media supra sparsim eisdem.

Ramuli apices versus c. 3x2 mm. primo compressi rugosi dein

Ashton — Bornean Dipterocarpaceae 295

terescentes levescentes pallide griseo-fusci; internodis —6 cm. longis;

cicatricibus stipularibus prominentibus prope amplexicaulibus

descensis. Gemmae c. 7X2 mm. lanceolatae caudacae. Lamina

13-18 x 7-10 cm. late ovato-elliptica coriacea basi obtusa vel

cordata apice in acumen breve latum attenuata; costis lateralibus

utrinsecus 11-13 crassis prominentibus angulo 50°-70° exorienti-

bus; internodis gracilibus conferte scalariformibus haud elevatis;

costa media supra applanata vel paullum depressa subtus pro-

minenti. Petiolus 23-35 mm. longus c. 2 mm. diam. teres.

Collections —Sarawak: SK. 168 (Brunig) Batu Skalap, Merurong

Plateau; S. 3864 (Pickles) S. Julan, Usun Apau; S. 13991

(Ashton) Ulu Teniai, Hose mountains. Sabah: SAN. 39323

(Meijer) Tavai plateau, Ulu Karamuak. Indonesian Kalimantan:

Kostermans 13095, Mr. Palimasan, near Tabang, Ulu Belajan,

W. Kutei (Holotypus in herb. Kew).

Differing from S. coriacea Burck, to which it appears to be most

closely allied, in its larger leaf with pubescent petiole and stouter

more prominent nerves. This species is characteristic of ‘carapa’ —

pole forest under poor drainage conditions—on high level

plateaux, usually between 700—1,300 m. alt., and has been observed

on sandstone, dacite and also ultrabasic rocks; SAN. 39323, which

is the record from the latter, is however atypical in having an

unusually short petiole.

The bark morphology suggests that this species belongs to sec.

Brachyterae Heim.

Shorea foraminifera Ashton sp. nov. Plate 46.

S. teysmannianae Dyer ex Brandis affinis sed lamina late ovata,

costis lateralibus utrinsecus 8-9 subtus prominentibus domatiis

magnis axillaribus costa media subtus tereti differt.

Novelli lamina stipulaque excepta fugace puberulentes. Ramuli

apices versus 1—2 mm. diam. primo paullum compressi dein teretes

fusci leves; internodis -3.5 cm. longis; cicatricibus stipularibus

brevibus subhorizontalibus. Gemma —8 x 5 mm. ovoidea compressa

subacuta. Stipula -10 x5 mm. oblongo-elliptica obtusa caduca.

Lamina 6—9 x 4-7 cm. late ovata coriacea, basi cordata vel obtusa,

apice in acumen —5 mm. longum paullum falcatum abrupte

attenuata; costis lateralibus utrinsecus 8-9 crassis subtus pro-

minentibus curvatis domatiis axillaribus prominentibus foraminiferis.

angulo 55°-70° exorientibus basi excepta; intercostis obscuris

conferte scalariformibus; costa media supra applanata subtus

crassa tereti prominenti. Petiolus 10-15 mm. longus c. 2 mm. diam.

Flores racemique ignoti. Pedicellus in fructu c. 1 mm. longus

c. 1 mm. diam. brevis. Calyx in fructu basim versus alutaceo-

puberulens apicem versus sparsim idem; lobis longioribus 3,

-7X1.2 cm. spatulatis subacutis basim versus -4 mm. latis

attenuatis partibus basalibus —11 x 5 mm. ovatis incrassatis saccatis;

lobis brevioribus 2, —4.5 x 0.4 cm. loratis acutis basim versus ut in

lobis longioribus. Nux —15 x 10 mm. ovoidea persistente alutaceo-

pubescens breviter apiculata.

296 Gardens’ Bulletin, Singapore — XXII (1967)

Collections —Kep. 35719, Andulau F.R.; S. 1916 (Smythies)

(Holotypus in herb. Kew), SAN. 17531 (Wood, Smythies, and

Ashton) Brun. 5541 (Ashton), Andulau F.R.; Brun. 243 (Ashton)

Ulu Belait Boundary rentis; s.n., Hasan, enumeration survey,

Baram-Rawa, Ulu Belait, 9/2/53.

This species I had previously regarded as a variety of S.

teysmanniana Dyer ex Brandis (Manual of the Dipterocarp Trees

of Brunei State (1964) 225, where the two trees are compared).

It is so far only known from Brunei, where it is sometimes very

common, and is found in fresh water swamp forest unlike S.

teysmanniana which is confined to peat.

Shorea pallidifolia Ashton sp. nov. Plate 41.

Species lamina ut in S. elliptica Burck sed 13-22x7-13 cm.

maiori tomento ochraceo scabrido basi subcordata costis lateralibus

utrinsecus 12-19.

Ramuli gemmae stipulae costae laminarum subtus_ petiolique

persistente pallide flavo-brunneo-scabrido-tomentosi, lamina subtus

brevissime conferte persistente ochraceo-scabrido-pubescens.

Ramuli apices versus c. 6X3 mm. diam. crassi primo compressi

costati dein terescentes; internodis 1-4 cm. longis; cicatricibus

stipularibus -—2 mm. longis subhorizontalibus prominentibus.

Gemma -10X8 mm. late ovoideo-conica compressa. Stipula

-15X7 mm. oblonga obtusa caduca. Lamina 13-22 x 7-13 cm. late

oblonga vel ovata vel obovaia coriacea, basi obtusa vel subcordata,

apice in acumen breve latum abrupte attenuata; costis lateralibus

utrinsecus 12-19 curvatis supra obscuris subtus prominentibus

basim versus angulo —110° exorientibus apicem versus angulo

45°-—70° exorientibus; intercosti gracilibus conferte scalariformibus

subtus vix elevatis; costa media supra obscura depressa subtus

prominenti. Petiolus 2-3.5 cm. longus c. 4 mm. diam. crassus sicco

rugosus. Racemi —15 cm. longi, stipitis basi in fructu -4 mm. diam.,

paullum compressi angulati axillares velrare terminales semel vel bis

ramosi; ramulis —3 cm. longis -6 flores distichos gerentibus;

bracteolis -4x3 mm. ellipticis obtusis caducis externe breviter

plane pubescentibus intus glabris. Alabastrum -63 mm.

ellipsoideum. Lobi calycis partibus in alabastro expositis pube-

scentibus, extoriores 3 deltoidei acuti internis 2 ovati quam externi

3 minores marginem versus tenuores. Petala elliptica partibus in

alabastro expositis pubescentibus. Stamina ignota. Ovarium

ovoideum glabrum, stylo filiformi glabro quam ovarium paullum

longiore. Pedicellus calyxque in fructu breviter sparsim plane

pubescens. Pedicellus c. 1 mm. longus c. 1 mm. diam. parvus.

Lobi calycis longiores 3, -7 x 1.5 cm. late spatulati obtusi basim

versus —7 mm. lati attenuati, partibus basalibus —10 x 8 mm. late

ovatis paullum incrassatis; lobi breviores 2, -4 x 0.3 mm. lineares

basim versus ut in lobi longiores. Nux —12 x 9 mm. ovoidea breviter

apiculata conferte breviter pallide alutaceo-pubescens.

Collections.—Sarawak: S. 16054 (Rashid) Bako National Park

(Holotypus in Herb. Kew); Kep. 79314 (Wyatt-Smith) Munggu

Unjam F.R.; S. 7452 (Brunig) Selang F.R.; S. 9315 (Bujang)

Ashton — Bornean Dipterocarpaceae 297

Setapok F.R.; S. 5957, 6161 (Brunig) Sempadi F.R.; S 13357

(Abang Muas) Balai Ringin P.F.

As the stamens are unknown this species cannot be placed in a

section with conviction, though it appears to be most closely

allied to S. elliptica Burck in Section Rubella Ashton; it differs

in the larger lamina with more abundant nerves and ocherous.

scabrid not cream tomentose undersurface like S. elliptica. It is

confined to the region of West Sarawak, where it is found in Heath

forest.

Shorea praestans Ashton sp. nov. Plate 42.

S. stenoptera Burck, S. macrophylla (De Vr.) Ashton affinis sed

lamina 24-35 x 11-20 cm. maiore basi cordata stipulis 4-6.5 cm.

longis glabris maioribus differt.

Partes ex integro glabri. Ramuli apices versus c. 5X3 mm.

crassi compressi leves; internodis 3—4.5 cm. longis; cicatricibus

stipularibus_ gracilibus evidentibus amplexicaulibus. Gemma

-25 x7 mm. faicato lanceolata compressa acuta. Stipulae —11 x5

cm. grandissimae ellipticae subacutae subpersistentes. Lamina

24-35 x 11-18 cm. grandis plus minus late oblonga coriacea basi

cordata apice obtusa vel breviter late acuminata; costis lateralibus

utrinsecus 11-13 supra obscuris subtus prominentibus angulo

45°—70° exorientibus basi excepta; intercostis remote scalariformi-

bus gracilibus subtus evidentibus; costa media supra applanata

subtus prominenti. Petiolus 4-6.5 cm. longus c. 5 mm. diam.

crassus sicco nigrescens rugescens. Flores racemique ignoti. Fructus.

ex integro glaber. Pedicellus in fructu c. 5 mm. longus c. 5 mm.

diam. crassus. Calyx lobis longioribus 3, —15 x3 cm. spatulatis

obtusis basim versus c. 1.5 cm. latis attenuatis partibus basalibus

-1.5x 1.8 cm. late ellipticis saccatis; lobis brevioribus 2, —10 x 1.5

cm. loratis subacutis basim versus ut in lobis longioribus. Nux

—2x2 cm. ovoidea glabra stylopodio —-4 mm. longo filiform

attenuata.

Collections —Sarawak: s.n., 18.6.58 (Anderson), s.n. (Brunig)

(Holotypus in herb. Kew), S. 22074, 22075 (Ashton) Bintulu

reservoir, Nyabau F.R.; S. 19598 (Ashton) Ulu Kenyana, Mukah.

This is a local species, known so far in Central Sarawak; though

this small tree of Heath forest is closely allied to §. stenoptera

Burck the lamina is considerably larger, the base cordate and the

petiole relatively longer. Though only immature fruit have been

collected they appear to be more similar to those of S. beccariana

Burck than S. stenoptera, with relatively small nut and 3 long

aliform calyx lobes.

Shorea pubistyla Ashton sp. nov. ) Plate 43.

S. scaberrimae Burck S. fallaci Meijer affinis sed lamina latiore

basi cordata petiolo racemoque longiore lobis calycis in fructu

longioribus aliformibus differt.

298 Gardens’ Bulletin, Singapore — XXII (1967)

Ramuli gemmae costa media subtus petiolique conferte per-

sistente scabrido-fulvo-tomentosi; costae laminae costa media

supra stipulae externe breviore scabrido-tomentosi stipulae intus

caduce plane pubescentes. Ramuli apices versus c. 7 mm. diam.

crassi primo prominente costati dein terescentes; internodis 1-3

cm. longis; cicatricibus stipularibus c. 2 mm. longus horizontalibus

obscuris. Gemmae -1.3 mm. longae -1.3 mm. diam. ovoideo-

compressae. Stipulae -2.5x0.7 cm. lanceolatae subcaudatae

caducae. Lamina 11-19x7-12 cm. late oblonga vel obovata

coriacea, basi obtusa vel cordata, apice subretusa vel breviter

acuminata; costis lateralibus utrinsecus 14-16 supra obscure

depressis subtus prominentibus basim versus angulo -110°

exorientibus apicem versus angulo —45° exorientibus; intercostis

remote scalariformibus subtus prominentibus. Petiolus 2.5-4 cm.

longus c. 3 mm. diam. crassus. Racemi —20 cm. longi, stipitis basi

in fructu —3 mm. diam., costati conferte breviter fulvo-scabrido-

tomentosi laxi axillares seme! ramosi; ramulis -6 cm. longis —12

flores secundis gerentibus; bracteolis -8x5 mm. oblongo-ovatis

subacutis breviter plane pubescentibus caducis. Alabastrum —7 x 3

mm. lonceolatum. Lobi calycis conferte tomentosi exterioribus 3

Ovatis acutis interioribus 2 ovatis marginem versus fimbriatis quam

exterioribus tenuioribus basim versus angustioribus. Petala externe

rosea intus coccinea elliptico-lanceolata partibus in alabastro

expositis pubescentibus. Stamina 15 internis 5 quam externis 10

paullum longioribus; filamentis basim versus latis compressis

apicem versus filiformibus abrupte attenuatis; antheris late

ellipsoideis; aristis gracilibus glabris quam antheris 2-3-plo

longioribus. Ovarium ovoideum conferte pubescente stylopodio

quam ovarium paullum longiore crasso columellari conferte

pubescenti stylo brevi glabro. Pedicellus in fructu —3 mm. longus,

—3 mm. diam. crassus. Calyx in fructu basim versus sparsim caduce

pubescens; lobis longioribus 3, -14x2.5 cm. spatuiatis obtusis

basim versus —8 mm. latis attenuatis; partibus basalibus —1.3 x 1.4

cm. late ovatis saccatis incrassatis; lobis brevioribus 2, —12 x 1.2

cm. loratis obtusis aliter ut in lobis longiorbus. Nux —3 x 1.8 cm.

anguste ovoideis conferte breviter plane pallide cremio-pubescens

apice in apiculo —-4 cm. longo attenuata.

Collections——S. 14930 (Anderson) (Holotypus in herb. Kew)

S. 15000, 14938 (Rosli), S. 15182 Galau), tree no. 614, 51 (Asah),

FA. 1301 (Carroll) Semengoh F.R.; S. 6227 (Brunig) Sempadi

F.R.; Sinclair and Kadim 10258, Gunong Gaharu, Serian; S. 16473

(Ashton) Ulu Kelawit, Tatau; S. 19575 (Ashton) Bt. Iju, Arip,

Balingian; Hotta 15720, 219, Minah camp, Kakus; S. 2118

(Smythies) Bt. Teraja.

This species is identical in floral characters to S. scaberrima

Burck, but differs ii the longer inflorescence and fruit sepals,

smaller nut, and larger coriaceous leaves with cordate bases and

longer petioles.

It is found in Mixed Dipterocarp forest on leached clay rich

soils, both overlying shales and rhyolite, in many localities west

of the Belait drainage in Brunei and throughout Sarawak.

Ashton — Bornean Dipterocarpaceae 299

Shorea rotundifolia Ashton sp. nov. Plate 44.

S. amplexicaulis Ashton affinis sed laminis alternatis remotis

orbicularibus; stipulis -62 cm. grandibus subpersistentibus;

petiolis 3-4 cm. longis.

Ramuli apices versus petioli stipulaeque pruinosi aliter glabri.

Ramuli apices versus c. 3X2 mm. diam. teretes vel paullum

compressi atro-brunnei leves dein pallide brunnescentes; internodis

-6 cm. longis; cicatricibus stipularibus amplexicaulibus pro-

minentibus. Gemma —20x8 mm. grandis lanceolata compressa.

Stipula -6 x 2 cm. lanceolato-falcata obtusa subpersistens. Lamina

9-21 x 8-14 cm. alternata late ovata vel suborbicularis coriacea

plus minus lustra basi obtusa vel cordata apice in acumen —1 cm.

longum cuspidatum attenuata; costis lateralibus utrinescus 9-11

curvatis subtus prominentibus basim versus angulo -115° apicem

versus angulo c. 45° exorientibus; intercostis remote scalariformi-

bus; costa media supra applanata subtus prominenti tereti. Petiolus

3-4 cm. longus, c. 3 mm. diam. longissimus sicco nigrescens.

Flores inflorescentiaeque ignoti. Fructus ex integro glaber.

Pedicellus in fructu c. 6 mm. longus c. 3 mm. diam. Lobi longiores:

calycis in fructu 3, c. 132.5 cm. spatulati obtusi basim versus.

c. 6 mm. lati attenuati partibus basalibus c. 15 x 18 mm. saccatis;

lobis breviores 2, —8 x 0.7 cm. anguste lanceolati subacuti basim

versus ut in lobis longioribus. Nux c. 2.5 X 1.8 cm. ovoidea glabra in:

stylopodio —7 mm. longo attenuata.

Collections —S. 9470 (Smythies) S. Iran, Pelagus (Holotypus in

herb. Kew); S. 22051, 22053 (Ashton) Ulu Segan, Bintulu.

This local species of Central Sarawak is locally frequent on

sandstone knolls and ridges below 500 m. alt. There it is frequently

found actually in association with its close ally S$. amplexicaulis,

but I have found no evidence of hybridisation in the forest.

Shorea sagittata Ashton sp. nov. Plate 45.

S. ferrugineae Dyer ex Brandis affinis sed lamina subius raniulis-

que conferte breviter persistente roseo-fusco-scabrido-tomentosis.

costis lateralibus utrinsecus 19-25 differt.

Lamina supra plane caduce cremio-pubescens, folia aliter

ramulique conferte breviter persistente roseo-fusco-scabrido-.

tomentosi. Ramuli apices versus c. 2 mm. diam. teretes costati dein

rufescentes verruculescentes; internodis —3 cm. longis; cicatricibus.

stipularibus brevibus ascensis. Gemmae —7X7 mm. late ovoideae..

Stipulae —10 x 8 mm. ovatae subacutae saccatae caducae. Lamina

7—14.5 x 3-6 cm. anguste oblonga vel lanceolata, basi obtusa, apice

in acumen —1.5 cm. longum gracile attenuata; costis lateralibus

utrinsecus 19-25 gracilibus subtus prominentibus basim versus

angulo 70°—80° apicem versus angulo —50° exorientibus; intercostis

gracilibus conferte scalariformibus; costa media supra depressa

subtus prominenti. Petiolus 9-17 mm. longus 2-3 mm. diam.

persistente roseo-fusco-scabrido-tomentosus. Flores ignoti. Racemi

—6 cm. longi, stipitis basi in fructu c. 2 mm. diam., conferte

persistente roseo-fusco-scabrido-tomentosi semel ramosi. Calyx in

300 Gardens’ Bulletin, Singapore — XXII (1967)

fructu sparsim puberulens; lobis longioribus 3, —11 x 2.5 cm. lorato-

spatulatis obtusis basim versus —8 mm. latis vix attenuatis pro-

minente sagittato-auriculatis in media parte saccatis incrassatis,

lobis longioribus 2, -20x 3 mm. linearibus acutis basim versus

haud auriculatis. Nux -15 <8 mm. ellipsoidea breviter apiculata

apicem versus puberulens.

Collection—S. 15882 (llias) Nyabau F.R.; FA. 566

(Wright) Ulu Gedong, Sadong; S. 18316 (Ashton) Ulu Sinrok,

Similajau F.R.; FA. 420 (Wright) Baleh; S. 7608, 7609 (Brunig)

Bungoh range; Hotta 15714, Minah camp, Kakus; S. 23616 (Sibat)

(Holotypus in herb. Kew), S. 23720 (Jugah) Bt. Iju, Arip; S. 22727

(Murthy and Ashton) G. Dullit.

With close affinities to S. ferruginea Brandis, S. sagittata is in

leaf characters almost indistinguishable from Shorea ovalis (Korth)

BI. ssp. ovalis of southern Kalimantan, Sumatra and Malaya,

though the auriculate fruit calyx is entirely different from that

species. The similarity to S. ferruginea extends to the bark

morphology, which is overall smooth, later patchily shallowly

cracking and flaking to leave a secondary smooth surface.

Recently flowering material of S. ferruginea, SA. 622, has come

to light; the flowers are typical of section Muticae Brandis.

With the addition of the present species, and with the systematic

position of S. ferruginea now confirmed, it is all the more clear

that S. ferruginea Brandis, S$. macroptera Dyer, S. slooteni Wood

ex Ashton, S. myrionerva Ashton, S. acuta Ashton and S. sagittata

Ashton form a well defined natural group within section Muticae

Brandis characteristed by their auriculate fruits and very different

bark morphology from other species. Whitmore (Gard. Bull. Sing.

19 (1962) 358) has already anticipated this in his bark studies.

T therefore propose to designate this unit as a new subsection.

Shorea sect. Muticae Brandis, subsect. Auriculatae Ashton subsect.

nov.

Type species: S. macroptera Dyer.

Flores ut in Sect. Muticae Brandis sed lobi calycis in fructu

basim versus auriculatae.

6 species: Malaya (1), Sumatra (1), Borneo (6).

On the basis of the bark morphology Whitmore places S.

macroptera in the same group as the species of Section Pachycarpae,

calling the group as a whole the Kawang subgroup. The bark of

these species characteristically remains smooth, and of Whitmore’s

Bark Manifestation Smoth C, at least until maturity.

The definition of this manifestation is as follows (quoted from

Whitmore):

‘Periderms: 1—2 present together. Inner bark structure: expansion

tissue of tangentially elongate stone cells and parenchyma; phloem

proliferation tissue usually widespread, of parenchyma and large

circular stone cells. Outer bark structure: thin rhytidome layers

to 1-2 mm. thick. Sloughing: apparently absent. Surface: either

finely sculptured rugose, rugulose, or not sculptured, smooth with

small lenticels.’ At maturity, unlike numbers of sect. Pachycarpae,

members of this subsection may become shallowly fissured,

Ashton — Bornean Dipterocarpaceae 301

exhibiting bark manifestation shallowly fissured C. This manifesta-

tion differs from that of the characteristic manifestation as follows:

‘Expansion tissue in uniformly distributed fingers, — uniseriate files

of tangentially elongate stone cells; fissures not penetrating into

inner bark.’

Shorea sect. Muticae Brandis, subsect. Muticae.

Lobi calycis in fructu haud auriculata.

Bark surface early becoming fissured, of Bark Manifestation

shallow fissured A, sometimes becoming flaky on the butt in old

trees. This manifestation is defined by Whitmore as follows:—

‘Periderms: multiple structure, thickness variable (0.03-0.20

mm.), phellems black and usually leathery, phelloderm either

apparently absent, or 0.25 mm. thick; several, close (0.5-1.0 mm.

apart, rarely more); not parallel to cambium for more than

1-3 cm. on T.S. Inner bark structure: pseudocortex present;

expansion at first by fingers and wedges, then by wedges only.

Outer bark structure: essentially a superficial skin, 3 mm. or less,

usually dipping in at, and bounding the fissures. Sloughing: on

poles usually absent; in crown absent or as layer thick, brittle,.

thin, scales, adherent before sloughing. Surface closely, more or

less regularly, V-section fissured; fissures penetrating inner bark;.

ridges smooth, sometimes micro-lichen flecked; surface firm, dark

coloured, at a periderm; bole smooth-fissured, becoming coarsely

fissured in old trees’.

302 Gardens’ Bulletin, Singapore — XXII (1967)

RS,

SPS Ses

SLO

renee

IKE S

‘ \)

Srass

wale

SSS

ee Ree

SOR

oat!

. Wis

wee

N RT SPRY

SS ‘I

NWP

S72

Gee

“He

Sa.

ZA E

PAL

EY)

eK)

SF @aho

gOS =

Plate 1. Anisoptera reticulata.

1, Fruiting branchlet; 2, minor, 3, major flower sepal; 4, petal;

5, androecium and gynoecium; 6, posterior, 7, lateral view of stamens;

3, fruit. From SAN. 12864..1, 8 =.x 4; 2-5 = x.10; 6, 7.= x 20.

Ashton — Bornean Dipterocarpaceae eS

——

ere

{Voge ee

EY .

—

Plate 2. Dipterocarpus cuspidatus. ‘

1, Fruiting branchlet; 2-4 fruit; 5, apex of nut. From S. 15821.

1-4 =x 4; 5 =x 1.

304 Gardens’ Bulletin, Singapore — XXII (1967)

m KA

mY TSS)

Plate 3. Vatica oblongifolia ssp. oblongifolia.

1, Leafy twig; 2, 4-9, fruit; 3, surface configuration of pericarp. From

S.t2516.° 1, 2, So a, DS

Ashton — Bornean Dipterocarpaceae 305

—

ay.

~~ OF ve

eee Sans yi

ASS

Plate 4. Vatica oblongifolia ssp. selakoensis.

1, Fruiting twig; 2, fallen leaf; 3, fruit. From S. ins lo <4;

ase mi.

306 Gardens’ Bulletin, Singapore — XXII (1967)

LC Ae

SANS

Plate 5. Vatica oglongifolia ssp. crassilobata.

1, branchlet with young fruit; 2, fruit; 3, fruit calyx. 1 from Brun..

614, 2, 3 from S. 2246. 1 = x 1/3, 2 = x4,3=xh

307

Ashton — Bornean Dipterocarpaceae

ae Ny

Cdbbe

igs

A Ms

)

. =

se Sa\ .S

“EO |

ae rom

ae STAY fay 8

Se "rm

( VED

: < AK =~

Wy ml PS LS

WL AOS ERS

NS <—_- ~§ .

Ae )

— > bos

} D}

taéA ‘

5 Z

= L

ze oO

_—

ie)

"ie

gn?

2 ka

5 2 &

= Se

= 2

—wWw

Es |

: _—

Qo x

3 DE

o «x

= a"

fm n

5 oe

Ss op in

og &

Seo

we —

as)

oes SS oS os Se, Ge a XK A

26029. 1,2 — x 4;

308 Gardens’ Bulletin, Singapore — XXII (1967)

Plate 7. Vatica oblongifolia ssp. elliptifolia.

1, Flowering branchlet (x 4, from S. 23225); 2, fruit (x 2, from

Brun 925).

309

x 4,

Ashton — Bornean Dipterocarpaceae

(E.

ey!

= : | ) “a,

LAH

Ie.

1, Fruiting branchlet; 2, fruit; 3, ripe nut. From S. 15857. 1

Plate 8. Vatica badiifolia.

pO et 5 ay. ee eB

Gardens’ Bulletin, Singapore — XXII (1967)

310

4, b aS

SP “WC .

PY) RY “A

™ =, — . Poss

, =

XY SS OSE \\

S \\

Sane

ox x3

ae (ig We

cs “0

CNG ae SD

Ni :

Plate 9. Vatica brunigi.

11 from

x 10:

1, fruiting branchiet; 2, fallen leaf; 3, flowering inflorescence; 4, 6,

major, 5, minor floral sepals; 7, petal; 8, androecium and gynoecium;

S. 17045; 2-10 from S. 17885. 1,3 = x 1/3; 2 = x 4; 4-8, 10

9, lateral view of stamens; 10, gynoecium; 11, ripe nut.

So = EO. At = ee

Ashton — Bornean Dipterocarpaceae 31]

of sos

—-

Plate 10. Vatica compressa.

1, flowering branchlet; 2, fallen ‘leaf; 3-5, flower buds; 6, major,

7, minor floral sepals; 8, petal; 9, androecium and gynoecium; 10,

lateral, 11, posterior view of stamen; 12 gynoecium; 13, 14, young;

15, mature fruit. 1-12 from SA. 604; 13-15, from SFN. 35618.

L2—s 3 — x 3; 6-9, 121 1G bl —-«x SS; 6-1 = x i.

312 Gardens’ Bulletin, Singapore — XXII (1967)

was

{7S

wri Ts

J JL

rt>

yr)

Ze ZF \ : \

et eS

Sot

PO, Vg

oy,

reg

waegi

== en on

MES

TIL Pig ae

=i n%8

Gers [SZ

Gee

of *

Ula

Ea,

~——ee,

Plate 11. Vatica congesta.

1, flowering twig; 2, enlargement of young twig; 3, flower bud;

4, major, 5, 6, minor floral sepals; 7, petal; 8, androecium and

gynoecium; 9, gynoecium; 10, fruit; 11, ripe out. 1-9 from S. 9347;

10, 11° from S8.*9347A.«1 = x*& 452 => x 333 = x 45 49

=i; 20 <2 B/D EeS uit

Ashton — Bornean Lupterocarpaceae

2 The

\ (|

Plate 12. Vatica globosa.

1, flowering branchlet; 2, flower bud; 3, minorg; 4, major floral sepal

5, petal; 6, androecium and gynoécium; 7, lateral view of stamens

8, gynoecium; 9,

11-13, fruits; 10, surface configuration of pericarp.

1-8 from Haviland and Hose 3159; 9-13 from S 18091. 1 = x

2 = x 4;:3-8 = x 15; 9-13 = x 1

313

Gardens’ Bulletin, Singapore — XXII (1967)

314

SUNS

a 9

ESS

ho. N

—

t; OSS

i. c

a a

1, fruiting branchlet; 2, fallen leaf; 3, fruit. 1,2 = x 4;3 = x 3.

Plate 13. Vatica rotata.

315

Ashton — Bornean Dipterocarpaceae

Wass Y

Plate 14. Vatica rynchocarpa.

1, branchlet with young, and 2, mature, fruit; 3—5, fallen leaf; 6, 7,

ripe nut. From Brun. 3354. 1-5 = x 4; 6, 7,

x2;

Gardens’ Bulletin, Singapore — XXII (1967)

316

Z Hf)

Ziyg) r

ly “ids, Wy,

~ Cif»

‘\

Plate 15. Hopea aequalis.

lee)

(or)

—~y

be |

7p)

°o

——

5 =]

vac

ron)

dia

»<

—

ee]

13}

i=)

fas}

bea

ao)

fee

317

Ashton — Bornean Dipterocarpaceae

ioe

Mlicak ae? /

ell 3

= <A

La OT

hy Ls

Ae ZN

e =i ie a

1, fruiting branchlet (x 4); 2, fruit (x 2). From Brun. 1030.

Plate 16. Hopea altocollina.

318 Gardens’ Bulletin, Singapore — XXII (1967)

——_=

= pet

LN,

{{) i

( \ f A 4

Plate 17. Hopea andersoni.

1, fruiting twig of ssp. basalticola; 2-14, ssp. andersoni. 2-4, fallem

leaves; 5, flowering raceme; 6, outer, 7-8, inner floral sepals; 9, petal

and stamens; 10, stamens; 11, gynoecium; 12-14, fruit 1 from S. 25005;

2-14 from $,- 10353. 1-5, 12-14, = x 4569 = = 40, tf = £2

Ashton — Bornean Dipterocarpaceae 319

wy)

My Js

>> =

”

ix,

CRY LEGS

4 he,

ane:

swe,

Jioaeks

= =

> yo

eae

ae 2 a

1 ee oS

—

——

ae es

—_

ans

——y

=a.

a a eT

Saas

——

———

—

Plate 18. Hopea bullatifolia.

1, leafy branchlet (x 4); 2, fruit (x 2/3). From S. 522.

320 Gardens’ Bulletin, Singapore — XXII (1967)

Plate 19. Hopea centipeda.

1, fruiting branchlet; 2, outer, 3, inner, floral sepals; 4, 5 petals;

6, androecium and gynoecium; 7, stamens; 8, gynoecium; 9, fruit;

10, ripe nut. 1, 9, 10 from S. 18440, 2-8 from S. 23342. 1 = 1/3;

2, 3, 6, 8 =-x0l0; 4, S, 10%! a5; 7 i eee SZ

Ashton — Bornean Dipterocarpaceae 321

Plate 20. Hopea depressinerva.

1, flowering branchlets; 2, outer, 3, inner floral sepals; 4, petal;

5, petal with stamens; 6, androecium and gynoecium; 7, frontal; 8,

lateral view of stamens. From S. 15439. 1 = x 4; 2-6 = x 10;

1, = 2

422 Gardens’ Bulletin, Singapore — XXII (1967)

Plate 21. Hopea enicosanthoides.

1, fruiting branchlet; 2 fruiting inflorescences; 3, fruit. From S.

18116. (x 4).

Ashton — Bornean Dipterocarpaceae oad

Plate 22. Hopea kerangasensis.

1, flowering twig; 2, fallen leaf; 3, outer, 4, inner, floral sepals;

6, petal with stamens; 7, stamens; 5, gynoecium; 8, fruit 1-7 from S.

me, ete S, 8944, 1 =e 45.2 — x 2; 34,6 = x 8; 5

1 =z 2 8 = x 5.

324 Gardens’ Bulletin, Singapore — XXII (1967)

eam

—

; 4

Plate 23. Hopea longirostrata.

1, fruiting twig; 2 = fruit; 3 = nut. From S. 17742. 1 = & #4;

3 =x 2. -

em — — ——

Ashton — Bornean Dipterocarpaceae

Plate 24. Hopea megacarpa.

1, flowering and fruiting twigs; 2, flower bud; 3, outer, 4, 5, inner,

floral sepals; 6, petal; 7, androecium and gynoecium;

Fruiting twig from SA. 639, otherwise from S. 14743

2 oe aoe 5-57 x 20;°6 = x 15: = *

325

8, stamens.

- 1 = x 4;

Sa y i, VA

Wie VwenWY) I-77

: EF |B

oF SS

Plate 25. Hopea mesuoides.

1, flowering branchlet; 2, leaf of young tree; 3, inner, 4, outer floral

sepals; 5, ium and gynoecium; 6, stamen; 7, gynoecium; 8, 9,

fruit. 1, 3-7 from S. 11247; 2, 3, 9 trem 3. 19551. 1, 2=— = iyo:

3,4,6,7 = x 10,5 = 2 73.6,9—:27%

Ashton — Bornean Dipterocarpaceae

Plate 26. Hopea pterygota.

aw ) V Wa LEZ Ls

i Ip = re

AD x f Wb Say

: st { Wha age

ie /j aN 53 ak

Pa Wy fl > 7

\\ "/ Za j ;

\ 4 off Az F 4

Oy \ ' Y Ak pe

Sh) | Seer ‘i

PAW \ \ { } Nid

= Whe Wy WI] SA fh

=E=Yr, fo \ eK ) y Gx iy _ Soy

IZ y i Ag ’ x, Za)

VW exere FT XA

4 i] }

= Vu

) Ny,

f My

/ i

327

1, fruiting branchlet; 2, outer, 3, inner floral sepals; 4, petal and

stamens; 5, lateral view of stamens; 6, gynoecium. From S. 15206.

1 x

== x 10; 5-6 = x 20.

328 Gardens’ Bulletin, Singapore — XXII (1967)

Aa \ (Lz <ff

Plate 27. Hopea tenuinervula.

1, leafy branchlet, 2, 3, fruit. From S. 9616. 1,2 = x4;3 =x 1.

Ashton — Bornean Dipterocarpaceae 329

ZL:

Z.——

Plate 28. Shorea brunnescens.

1, flowering twig; 2, detail of lamina nervation below; 3, outer,

4, inner, flower sepals; 5, petal, 6, ovary and gynoecium; 7, 8, stamens;

5 ee ome = pm heer, 2 = = 13,7 = x 1, 5, 6,9 = x 10:

cy oS oe

330 Gardens’ Bulletin, Singapore — XXII (1967) ;

Plate 29. Shorea lunduensis.

1, flowering branchlet; 2, flower bud; 3, 4, outer, 5, inner sepals;

6, petal; 7, androecium and gynoecium; 8, stamens, frontal view; 9,

stigma; 10, gynoecium. From S. 15396. 1 = x 4;2 = x 4; 3-5 = x 15;

6,77 ="x' 1073 =x

Ashton — Bornean Dipterocarpaceue 331

Plate 30. Shorea agami ssp. diminuta.

1, leafy twig; 2, fallen leaf; 3, fruit. From S. 25027. 1,3 = x 4;

332 Gardens’. Bulletin, Singapore — XXII (1967)

> Ip JY LZ (ZB

DT bp a

| EV Fike

Fe JZ

UZ,

Plate 31. Shorea cordata.

1, flowering branchlet; 2, detail of young twig; 3, outer, 4, inner,

floral sepals; 5, petal; 6, stamens and gynoecium; 7, posterior view of

stamen. From SA. 678. 1 = & #% 2:3 3; 3-6 = % 10,7 =a T.

Ashton — Bornean Dipterocarpaceae 333

adizig

P a\~

vo iy

Plate 32. Shorea polyandra.

1, flowering branchlet; 2, flower bud; 3, outer, 4 inner floral sepal;

5, petal; 6, androecium and gynoecium; 7, lateral, 8, frontal view of

outer, 9 same of inner, stamens; 10, gynoecium. From S. 10171.

1 = x4;2 = x 5; 3-6, 10, = x 10; 7-9 = x 30.

334 Gardens’ Bulletin, Singapore — XXII (1967)

(

Hil

Hl }! )

SoS ts:

Bi eR SISOS

me }

Po ee oe en arn a ec Sn ae ee

——

——.

Plate 33. Shorea alutacea.

1, flowering branchlet; 2, fallen leaf; 3, flower bud; 4, opened flower;

5, outer, 6, inner floral sepal; 7, androecium and gynoecium; 8, stigma;

9, stamen, lateral view; 10, gynoecium; 11, fruit; 12, ripe nut. 1-10

from S. 7975; 11, 12 from’S., 10170. 1, 2,.11, 12 = x4; 3, 4 ='x 5;

S—/, 10 = X 10; 8. =X 405.9 = X15.

335

Ashton — Bornean Dipterocarpaceae

0 y

o Wi

é rp

Leg

Fini)

1, leafy branchlet; (x 4); 2, fruit (x 2).

Plate 34. Shorea bakoensis.

336 Gardens’ Bulletin, Singapore — XXII (1967)

Plate 35. Shorea cuspidata.

1, fruiting branchlet; 2, detail of young twig; 3, fallen leaf; 4,

flowering raceme; 5, flower bud; 6 inner, 7, outer floral sepal; 8,

corolla; 9, androecium and gynoecium; 10, lateral; 11, posterior view of

stamen; 12, gynoecium; 13, 14, fruit. 1-3, 13, 14 from S. 15258, 4-12

from tree no, 603. 1, 3, 4, 13, 14 = & 4; 2 = x 2; 3,8 = = 8; G, 7,

10, 11 = x 25; 9B, 12 = &% 15.

Ashton — Bornean Dipterocarpaceae rv sof

— Ly

soe

SARS

see Se

RSD

—

~~ rl

SoS

ney

‘a

——

Lea

$2 PZ

—l=—

= -

=e.

= =e

Plate 36. Shorea iliasii.

1, fruiting branchlet; 2 fruit; 3, ripe nut. From S. 15102. 1, 2 = 1/3;

3=k 4.

meme 6

338 Gardens’ Bulletin, Singapore — XXII (1967)

)

mals °,

Drop

Plate 37. Shorea macrobalanos.

1, fruiting branchlet; 2 young, 3 mature fruit. From S. 13712. 1,

= x42 = x1.

Ashton — Bornean Dipterocarpaceae 339

QMS

Plate 38. Shorea mujongensis.

1, fruiting twig; 2, fruits; 3, ripe nut; from S. 19038. 1, 2 = x 4;

3 = x 2.

340 Gardens’ Bulletin, Singapore — XXII (1967)

Plate 39. Shorea carapae.

1, leafy twig; 2, young fruit. 1 from S. 13991; 2 from SAN. 39323;

ly, Z.=% 5.

Ashton — Bornean Dipterocarpaceae 341

Plate 40. Shorea foraminifera.

1, leafy branchlet; 2-6, fallen leaves; 7, 8, fruit; 9, detail of outer

surface f" fruit calyx lobe. 1 from Brun. 243; 2-9 from S. 1916.

1-8 = x3;9 =x 1.

342 Gardens’ Bulletin, Singapore — XXII (1967)

LY)

\\ ty HA

WENA ASOD

BEN

KAPEAE)

eal

—— =

FSS —

aoe,

evan

S =

See

CSNY

——

SSS

zs a——~

SST

——

Plate 41. Shorea pallidifolia.

1, fruiting branchlet; 2, fallen leaf; 3, detail of young twig; 4, fruit;

5, ripe nut; 6, 7, germinating seedlings; 8, detail of seedling stem.

From. '8:' 16034: °F 52 5"6; * 7" A/S, os Ge

ped : —

™ eee PR MN by

CIEE Algae ey (7

eran) erreees eRe oe ="

—

young fruit. 1, 3 from S. 19598;

8 at

S ae

8 ‘a

S o

ine =

8 =|

: .

S ay

~ -

5 uN »%

Sy S itl

a = 2G

Y a's

S ‘oOo

5 x g Pe

XA am

Y So

NI 8 5

é San

m5 § 25

| 4 2 ¢

= 2

S ae

E E

N Ay

344 Gardens’ Bulletin, Singapore — XXII (1967)

RSS

Iohe >

Naae: xD

<= Se

, b

Wy, UY

IY CPL DORE FG

CY A es Pe bn

ie:

if,

SS a

say

LW t ryt

ieshicaee

Vii WES

‘ DIST oa f

Plate 43. Shorea pubistyla.

1, flowering branchlet; 2, outer, 3, inner, floral sepals; 4, petal;

5, androecium and gynoecium; 6, lateral view of stamen; 7, gynoecium;

8, fruit; 9, ripe nut. 1-7 from S. 14930; 8, 9 from FA. 1301.

1, 9= x 1/3; 2, 3, Se =z 15,585, 7 Se Sse

Ashton — Bornean Dipterocarpaceae

D \ .

4 \ | \

a | 4 NW A

4 e \\ te aa )

= = ——TT/ [

SOF UL 2 SS

Leer

[

Plate 44. Shorea rotundifolia.

Tae branch; 2 young fruit. 1 from S. 22051; 2 from S. 9470.

x ;

345

346 Gardens’ Bulletin, Singapore — XXII (1967)

}

9. & ry) }

)

Ny H |

tai at

iene, y

a 4y

i SS |

Plate 45. Shorea sagittata.

1, leafy branchlet; 2, detail of twig; 3, detail of leaf undersurface;

4, fruit; from S, 15882. 1 = x 4; 2,3 = x 4;4 = x 2/3. :

347

Ashton — Bornean Dipterocarpaceae

‘aqny xA[e> 9y} JO odeys ul UONeIeA oy} pue divoiied aje]J90NUa] oY} s}OU ‘(ULE

‘6

ZS96I'S) Attpesoy odAy oy} wor dn poysid 10Aq snyofisuojqo sdouvjpgodiq JO syinig

348 Gardens’ Bulletin, Singapore — XXII (1967)

Bole of Hopea altocollina sp. nov.; Hose mountains, Sarawak. April 1964.

Ashton — Bornean Dipterocarpaceae 349

Bole of Hopea tenuivervula sp. nov.; Semengoh arboretum, Kuching.

November 1963.

350 Gardens’ Bulletin, Singapore — XXII (1967)

Bole of Shorea cuspidata sp. nov.; Semengoh arboretum, Kuching.

November 1963.

Ashton — Bornean Dipterocarpaceae 351

Bole of a giant Shorea mujongensis sp. nov.: on the Mujong-Balui divide,

Rejang, Sarawak. April 1964.

352

w®&

Bole of Shorea

Gardens’ Bulletin, Singapore — XXII (1967)

sagittata sp. nov.; Similajau F.R.; Bintulu. March 1963.

Reprinting of Burkill’s Dictionary of the

Economic Products of the Malay Peninsula

This work, published in 1935, quickly became a standard

reference book for scientist and administrator alike concerned

with utilisation of natural resources of Malaya. Its usefulness is

acknowledged generally throughout the Tropics and wherever tro-

pical plant products are studied. After being unavailable for the

past 15 years, except at an exorbitant price for the very few

second-hand copies coming onto the market, the Federation of

Malaysia Government is doing an extremely useful service to

tropical agri-horticulture and forestry in reprinting this book. A

number of amendments and corrections on the original edition

were prepared by the author before he died in 1965, and these

have been incorporated in the new edition.

The work in two volumes was published near the end of

1966 at a price of M$59 for both volumes. Inquiries should

be directed to the Publications Officer, Ministry of Agriculture

and Co-operatives, Federation of Malaya, Swettenham Road,

Kuala Lumpur, Malaysia.

;

’ : ¥

“e

: :

A * ‘ .

44 :

* * -

OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

1. Annual Reports.

Many from 1909 onward remain available.

Prices variable.

2. The Agricultural Bulletin of the Malay Peninsula (Series I).

Only Nos. 3, 5, 7, 8 and 9 available at 20 cents each.

3. The Agricultural Bulletin of the Straits and F.M.S. (Series I).

Vols. 1-10, 1901-1911, monthly issues.

All parts available, except Vols. 1, 2, 5 (7), 6 (11) and 9.

Price: $5 per volume, Cents 50 per part.

4. The Gardens’ Bulletin, Straits Settlements (Series III).

Vols. 1-11, 1912-1947. 3

Vols. 1 (1-5) January-May 1912 is issued under title a

Agricultural Bulletin of the Straits & F.M.S.

All parts available, except Vol. 1 No. 2, 10, Vol. 2, Vol.

3 No. 1-3 Vol. 6, Vol. 10 pt. 2 and Vol. 11 pt. 3.

Prices variable.

5. The Gardens’ Bulletin, Singapore (Series ve

Vols. 12-21, 1949 —*

All parts available Sou 13 (1). Prices variable. Issues

are irregular. |

Subscription will be entered against deposit.

Available on exchange.

6. Materials for a Flora of the Malay Peninsula, Monocotyledons.

Parts 1, 2 and 3 remain available.

7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. Bs

Addison.

Price $21. |

(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison.

Price $15 (Photo-lithographic reprint). )

8. A Revised Flora of Malaya. 7 |

(a) Vol. 1, Orchids, by R. E. Holttum. Price $20.

(b) Vol. 2, Ferns, by R. E. Holttum. Price $20.

9. Wayside Trees of Malaya by E. J. H. Corner, 2 Vols

Price $25. .

Items 1-6 can be obtained from the Director, Botanic Gardens,

Singapore.

Items 7-9 can be obtained from the Government Printer, Govern-

ment Printing Office, Singapore.

Prices quoted are in Singapore Dollars

Overseas postage is extra

0223—500—12/67

f Aa a 2 “} j

‘ Y =~

a 4 hoes, é *;

1 i s a ’

a, |

THE

| ‘

GARDENS’ BULLETIN

SINGAPORE

Vol. XXII, Part III 29th June, 1968

CONTENTS

PAGE

ae ‘SoFPADMO: A revision of the genus Quercus _ - - 355

BURKILL et al.: Some studies of Malayan Agarophytic and

Alginophytic seaweeds : - - . - 429

KOSTERMANS : New species of Dipterocarpaceae, etc. - - 443

SKvortzov: On some species of Euglena’ - Shasta 447

SKVORTZOV: On a new species of the genus Collodictyon - — 451

-SKvorTzov: New genera of primitive green Flagellata, etc. - 455

HARDIAL SINGH: Sclereids in Raphidophora and Scindapsus - 461

ALPHONSO: “Gardening in Hot Countries”, a review - - 469

To be purchased at the Botanic Gardens, Singapore

Price: $9 Fa ge

RECEIVED

OCT 29 1968

LIBRARY

wt ra

ee Un,

oe nie Published by Authority

; Bie . Printed by Lim Bian Han, Government Printer, Singapore

| 1968

oS ny

ae ter

LONER IEEE YEO ELEOIIOLOPILEVE

OS OEE YEO YE IERIE Pe \ DFE DBE 2FAnwW FE DES

c< ie ee >

A Revision of the Genus Quercus L. Subgen.

Cyclobalanopsis (Oersted) Schneider* in Malesia

E. SOEPADMO

(Herbarium Bogoriense, Bogor, Java, Indonesia)

SUMMARY

Quercus L. is here compared with and regarded as a separate genus

from Lithocarpus Bl., differing from the latter by its inflorescence, flower

and pollen characters. The genus is subdivided into two subgenera, viz.

subgen. Quercus (=subgen. Eu-Quercus A. Camus) and Cyclobalanopsis

(Oersted) Schneider. The cupule in the former is beset with imbricate

scales, and the male flowers are solitary along the rachis. In the subgen.

Cyclobalanopsis the cupule is lamellate, and the male flowers are in

4-1-flowered dichasial clusters. The generic splits in Quercus proposed by

Oersted (1867, 1871), Schottky (1912), and Schwarz (1936 a, b) are not

accepted.

Quercus subgen. Cyclobalanopsis extends from Japan (Kanto Prov.,

C. Honshu) south- and westwards to Korea, China, Formosa, Indo-China

(Vietnam, Laos, Cambodia), Thailand, Burma, N.E. India, and Western

Malesia, with Indo-China as the centre. In Malesia the distribution does

not extend further East than Borneo nor further North than Palawan.

The centre of distribution in Malesia is in Borneo, from where 17 of the

19 species have been recorded.

Q. kerangasensis and kinabaluensis are described for the first time.

Q. valdinervosa is a new species based on Q. mespilifolia Wall. ex A.DC.

var. borneensis Heine.

The characters in the group, in particular the nature of the cupule

in various genera, are extensively discussed. The questions on the identity

of several species are explained. Keys, synonyms, and descriptions are

given.

INTRODUCTION

This paper is the first contribution towards a revision of the

Fagaceae in Malesia, represented by five of the seven genera

currently recognized in the family. These five genera are Casta-

nopsis Spach, Lithocarpus Bl., Nothofagus Bl., Quercus L. and

Trigonobalanus Forman. Of these five genera no less than 200

species have been attributed to Malesia.

Quercus is the largest and the most widely distributed genus.

In 1938, A. Camus monographed the genus, recognizing no less

than 500 species. She followed Schneider (1906) by dividing the

genus into two different subgenera, viz. Eu-Quercus and Cycloba-

lanopsis; in the former the cupule is armored with imbricate

scales, whilst in the latter the cupule is lamellate. In her mono-

graph, and also in 1948, A. Camus credited 17 species to Malesia.

Of these only 11 are here acknowledged. In April 1966 I described

five new species, and in the present paper I have added two and

have proposed a new name which make the total number of

species known from Malesia amount to 19. According to

Schneider’s circumscription all of these species belong to the

subgen. Cyclobalanopsis.

*In April 1966, I erroneously referred to this subgenus under the

name Quercus L. subgen. Cyclobalanopsis (Oersted) A. Camus.

356 Gardens’ Bulletin, Singapore — XXII (1968)

The major point of interest in the genus Quercus lies in the

controversy over its clear difference from the genus Lithocarpus,

and also over the question on the generic splits as proposed by

Oersted (1867, 1871), Schottky (1912), and Schwarz (1936 a, b).

A. DeCandolle (1863, 1964), Miquel (1863), Oudemans (1865),

Baillon (1877), Bentham & Hooker f. (1880), Wenzig (1886),

Hooker f. (1888), Koorders & Valeton (1904), O. von Seemen

(1906), Brandis (1906), Hosseus (1911), Merrill (1908, 1921, 1923),

Burkill (1935), Corner (1939, 1940), and Backer & Bakhuizen van

den Brink Jr. (1965) were of the opinion that the genera Quercus

(incl. Cyclobalanopsis, Macrobalanus, and Erythrobalanus) and

Lithocarpus (incl. Pasania, Cyclobalanus, and Synaedrys) should

be regarded as a single genus, namely Quercus, as these genera

possess a similar type of cupule.

The opposite view was introduced by Oersted in 1867, in

which Quercus was treated as a distinct genus, differing from

Lithocarpus by its inflorescence and flower characters. This view

was accepted by Prantl (1887, 1894), Schneider (1906), Dalla Torre

& Harms (1900-1907), Schottky (1912), Gamble (1915), Koidzumi

(1916), Rehder & Wilson (1916), Rehder (1919, 1929), Ridley

(1924), Markgraf (1924), Henderson (1930), Lemée (1932, 1934,

1939), Schwarz (1936 a, b), Camus (1938, 1954), Barnett (1943,

1944), Hjelmqvist (1948), Brett (1964), Forman (1964), Melchior

(1964), and many others.

Before deciding which of these two opposite views should be

adopted here, I have carried out a comparative study on the

morphology of the inflorescence, flower, cupule, and pollen grains,

and also on the anatomical structure of the cupule and the fruit

wall of both genera. The main conclusions should be drawn here

are as follows:

(1) I fully agree with Oersted and others in separating Litho-

carpus from Quercus. These two genera may be distinguished

from one another by the following characters:

Quercus Lithocarpus

a. Inflorescence unisexual. ... Inflorescence usually androgy-

nous or mixed.

b. Male inflorescence flexuous, ... Male inflorescence rigid, thick

pendent. or slender, erect.

c. Male flower normally with ... Male flower normally with 12

6 stamens and without a stamens, and always with a

pistillode. pistillode.

d. Anthers basifixed, c. 0.5-1 ... Anthers dorsifixed, c. 0.20—-0.25

mm long. mm long.

e. Female flower mostly with- ... Female flower with 12 stamin-

out staminodes. odes.

f. Styles flattened, bearing ... Styles terete, bearing minute,

broad stigmatic surface, terminal, punctiform _ stig-

or cylindrical and groov- mas.

ed with capitate stigmas.

uese

— we ee ee ee

Soepadmo — Quercus subgen. Cyclobalanopsis 357

Quercus Lithocarpus

g. Pollen grains’. prolate- ... Pollen grains prolate-perpro-

spheroidal, c. 30 X 20 u, late, c. 18 x 10 yp, always

mostly 3-colpate, rarely 3-colporate; exine relative-

3-colporate; exine more ly smooth (seen under light-

or less scabrate. microscope).

h. Cupule normally solitary ... Cupule solitary or aggregated

along the rachis. in dichasial clusters along

the rachis.

(2) In its general form and also in its anatomical structure,

the cupule of Quercus is exactly the same as that of Lithocarpus.

The similarity even extends to the type of external emergences

(scales or lamellae) covering the cupule. The cupule of these two

genera represents a distinct type and differs from the other type

recognized in the Fagaceae, viz. Fagus-Castanea-type, by being

Open, cup- or saucer-shaped or completely covering the solitary

fruit, indehiscent, not spiny but covered with imbricate scales or

concentric lamellae. I do not think, however, that the characters

of the cupule alone are sufficient to merge Lithocarpus with

Quercus.

(3) Three different types of anatomical structure of the fruit wall

in the Fagaceae are distinguished, viz. Quercoid-type (Quercus,

Castanea, Castanopsis, Lithocarpus), Fagoid-type (Fagus, Trigono-

balanus), and Nothofagoid-type (Nothofagus). The descriptions

of these anatomical structures are given on p. 369 and figured in

Plates 3-5.

(4) With respect to the generic splits in Quercus, it may be

pointed out here that Oersted (1867) was the first who suggested

splitting the genus into two smaller genera, viz. Quercus and

Cyclobalanopsis. Amongst the later authors who accepted this

subdivision are: Schottky (1912), Markgraf (1924), Kudo & Masa-

mune (1930), Nakai (1939), Hjelmqvist (1948), Li (1953, 1963),

and Brett (1964). In 1936 Schwarz even proposed subdividing the

genus into four genera, namely Quercus s.s., Macrobalanus,

Erythrobalanus, and Cyclobalanopsis. No subsequent author has

accepted this suggestion.

The characters used by the above mentioned authors, to distin-

guish the two or four smaller genera, are that of the cupule,

abortive ovules, male inflorescence and flower, and the styles.

In the present study however it was found that except for the

presence of a lamellate cupule in Cyclobalanopsis, all the differ-

ing characters are too inconsistent to be regarded as generically

important. There are too many intermediate characters which

obscure a clear distinction between those smaller genera. There-

fore, I am of the opinion that it is not worthwhile to split the

genus Quercus into two or four smaller genera. Amongst the

previous authors who have accepted this idea are: Gamble (1915),

Koidzumi (1916), Rehder & Wilson (1916), Ridley (1924), Trelease

(1924), Handel-Mazzetti (1929), A. Camus (1938), Lemée (1939),

Rehder (1940), Miiller (1942 a, b), Barnett (1943, 1944), Hutchin-

son (1959), Forman (1964), Ohwi (1965), and many others.

Gardens’ Bulletin, Singapore — XXII (1968)

ACKNOWLEDGEMENTS

I would like to express my deep gratitute to the Authority

of the Colombo-Plan Scheme, and to the British Council for the

scholarship which enabled me to carry out the present work.

To Dr. J. Barker, the Secretary of the Gordon Wigan Fund of

the University of Cambridge, England, and to Professor C.G.G.J.

van Steenis, the Director of the Flora Malesiana Foundation, Lei-

den, the Netherlands, I would like to thank them for the financial

support which made my visits to the Botanisches Museum in

Berlin-Dahlem and to the Rijksherbarium, Leiden, in Summer

1965, possible.

I am very grateful to the Directors of the following Herbaria

for the loan of specimens or for their hospitality given to me

during my visits to their Institutions:

Botanisches Museum, Berlin-Dahlem, Germany (B).

British Museum of Natural History, London, England (BM).

Herbarium Bogoriense, Bogor, Indonesia (BO).

Central National Herbarium, Botanical Survey of India, How-

rah, Calcutta, India (CAL).

Herbarium, Botany School, the University of Cambridge, Eng-

land (CGE).

Royal Botanical Gardens, Edinburgh, Scotland (E).

Herbarium Universitatis Florentinae, Istituto Botanico, Firenza,

Italy (FI).

Royal Botanical Gardens, Kew, England (K).

Rijksherbarium, Leiden, the Netherlands. (L).

Museum National d’Histoire Naturelle, Laboratoire de Phanero-

gamie, Paris, France (P).

Sarawak Forest Department, Kuching, Sarawak, Malaysia

(SARF).

Botanic Gardens, Singapore (SING).

Botanical Museum & Herbarium, Utrecht, the Netherlands (U).

In particular, I am very much indebted to Professor E.J.H.

Corner, F.R.S., Professor of Tropical Botany, at the University

of Cambridge, England, for his advice and guidance during my

four years research work in the University.

My gratitude also extends to the following for their invaluable

advice and information: Dr. J.A.R. Anderson (Kuching, Sarawak),

Mr. H.M. Burkill and Dr. Chew Wee Lek (Singapore), Dr. D.M.

Churchill (Monash University, Australia), Mrs. C. Dikson (former-

ly Botany School, Cambridge, England), Mr. L.L. Forman (Kew),

Dr. M. Jacobs (Leiden), Dr. S.M. Walters (Botany School, Cam-

bridge, England), and Dr. F. White (Oxford).

Finally I would like to thank also to Dr. P.S. Ashton (formerly

Kuching, Sarawak), Mr. K. Kartawinata (formerly Department of

Botany, the University of Singapore), Prof. Dr. A.J.G.H. Koster-

mans and Mr. N. Wirawan (Bogor, Indonesia) for their assistance

to collect material.

Soepadmo — Quercus subgen. Cyclobalanopsis 359

CHARACTERS

Habit: in Malesia species of Quercus are trees of medium to

large size. Under favourable conditions an individual tree may

reach c. 40 m in height and 1.20 m in diameter. In each species

there is a general tendency for the plant to attain a smaller size

at a higher elevation. This is probably due to the more favourable

ecological conditions in the lowland forests. The ecological varia-

tion also extends to the size, texture and indumentum of the leaves.

In Q. elmeri, gemelliflora, lineata, lowii, subsericea, and valdi-

nervosa, for example, the leaves of the plant growing at lower

altitude are larger, thinner and less pubescent than those of a

tree inhabiting a higher altitude. The trunk is usually straight,

very rarely stunted. In most species buttresses are absent, but in

Q. argentata, elmeri, gemelliflora, percoriacea, pseudo-verticillata,

and treubiana, it may reach c. 1-3 m in height. The crown is

usually dense, spreading, and makes up about one third or

one half of the total height of the tree.

Bark: in the nineteen species from Malesia, three different types

of bark may be distinguished, viz. the smooth and lenticellate bark

(Q. argentata, kerangasensis, kinabaluensis, nivea, percoriacea, and

valdinervosa), the finely fissured bark (Q. chrysotricha, lowii, and

subsericea), and the scaly bark which peels off profusely into

rectangular pieces (Q. oidocarpa, pseudo-verticillata, and treu-

biana).The bark in Q.gemellifiora and linedta may be smooth and

lenticellate or finely fissured. In all species the outer bark is

usually thin and hard, and pale yellow to greyish-brown in colour;

the inner bark is sometimes as thick as 2.5 cm, brittle, fibrous,

ridged on its inner surface, and pale yellow to reddish-brown in

colour. The tannin content may be up to 20 per cent of the dry

weight.

Wood: from the available information it seems to be impossi-

ble to distinguish species of Quercus by the wood characters alone.

According to Cutler (1964), the wood of some species of Quercus

is almost indistinguishable from that of Lithocarpus and Trigono-

balanus. The wood of these three genera differs from that of Fagus,

Nothofagus, Castanea and -Castanopsis by its dendritic distribution

of the solitary vessels, uni- to multi-seriate rays, concentric bands

of parenchyma, bordered pitting in the wall of vessels and ray-

cells, and by its frequency of the libriform fibres. Cutler also men-

tioned however that the genera in the Fagaceae are not always

clearly defined from one another by the wood characters alone.

There are many overlapping characters between some species, not

only of the same genus but also of different genera. In the Malesian

species the wood is usually pale yellow to yellowish-brown in

colour, moderately hard and heavy, but not durable, splitting

very easily, and difficult to work with. The growth rings are indis-

tinct, and the sapwood is not sharply defined from the heart-

wood.

360 Gardens’ Bulletin, Singapore — XXII (1968)

Twigs: in the Malesian species the new shoots are as a rule

conferted. Young twigs are densely covered with yellowish-brown,

either simple or stellate-adpressed hairs. Older twigs are glabrous

and densely or sparsely lenticellate. Though the twigs are terete,

the pith is always star-shaped with 4-5 lobes in cross-section.

Vegetative buds: in most of the Malesian species the vegetative

buds are not characteristic for a particular species, but usually

small, c. 0.2-1 by 0.2-0.5 cm, ovoid-globose or ovoid-conical, with

the scales ovate-acute and densely simple or stellate tomentellous.

The arrangement of the scales is as a rule imbricate, but in

‘-Q. valdinervosa it is sometimes in four symmetrical rows. In Q.

subsericea and treubiana, the vegetative buds are elongated, ovoid-

ellipsoid, and the scales are linear-acute.

Stipules: in Malesia the stipules are extrapetiolar, caducous,

linear-acute, c. 0.2-1 by 0.1-0.2 cm, with simple or stellate tomen-

tum or with woolly or stiff pubescence. These characters are

however not specific.

Leaves: the simple leaves are spirally arranged, usually crow-

ded near the tip of the twig, or occasionally in a pseudo-whorled

arrangement (Q. pseudo-verticillata); venation pinnate, with the

‘midrib and lateral nerves usually flattened or impressed on the

‘upper surface and more or less prominent on the lower surface.

‘The petiole, whether it is long and slender or short and thick, is

always thickened at the base; glabrous or more commonly densely

or sparsely tomentose or densely stiff or woolly pubescent. In

‘QO. chrysotricha, merrillii, pseudo-verticillata, and steenisii, the

‘petiole is c. (0.1)-0.3-0.5-(1.0) cm long, whilst in the other fifteen

‘species it is slender, c. (1.0}-2-3-(4.5) cm long. The adaxial side

of the petiole in Q. lowii, nivea, oidocarpa, percoriacea, and sub-

sericea, is deeply sulcate at least in the upper part near the leaf-

‘base. In the other species the adaxial side of the petiole is either

flattened or rounded. The petiole of the mature leaf may be

completely glabrous (Q. argentata, oidocarpa, percoriacea, and

valdinervosa), densely stellate tomentose (Q. gaharuensis, lowii,

-and_ nivea), densely sericeous (Q. chrysotricha, gemelliflora, merril-

lii, pseudo-verticillata, subsericea, and sumatrana), or densely stiff

or woolly pubescent (Q. elmeri, kerangasensis, kinabaluensis,

lineata, steenisii, and treubiana). In a combination with the other

characters of the leaf the petiole morphology is very useful for

identification.

In Malesia the leaf-blade is either elliptic- to lanceolate-oblong

(Q. argentata, elmeri, gaharuensis, gemelliflora, lineata, nivea, oido-

carpa, pseudo-verticillata, sumatrana, valdinervosa, and occasion-

ally also in subsericea), or ovate-elliptic to ovate-lanceolate (Q.

chrysotricha, kerangasensis, kinabaluensis, lowii, merrillii, perco-

riacea, steenisii, and occasionally also in subsericea), or rarely

linear-lanceolate (Q. treubiana). The leaf varies from 2 to 24 cm in

length and from 1 to 9 cm in width. On account of this character,

three different species groups may be distinguished in Malesia,

namely Q. argentata, nivea, oidocarpa, percoriacea, pseudo-verticil-

lata, sumatrana, and occasionally also subsericea, which have

Soepadmo — Quercus subgen. Cyclobalanopsis 36h

relatively large leaves, c. 10-17 by 3-5 cm; Q. elmeri, gemelli-

flora, kerangasensis, kinabaluensis, lineata, lowii, steenisii, and.

usually subsericea, which have medium sized leaves, c. 5-10 by 2-4

cm; and Q. chrysotricha, merrillii and treubiana, which have small

leaves, c. 2-4 by 1.5-3 cm. There is however some overlapping,.

so that the size of the leaf should be used in conjunction with the:

other characters. The base of the leaf may be rounded, sub-

cordate or auriculate (Q. lowii, percoriacea, pseudo-verticillata and.

steenisii), or attenuate-rounded or attenuate-acute in the other

species. The margin is either entire as in Q. argentata, chrysotricha,

kerangasensis, kinabaluensis, nivea, and pseudo-verticillata, or

entire in the basal half and distantly serrulate in the apical half.

The apex of the leaf is rounded, bluntly acute or rounded emar-

ginate in QO. chrysotricha, merrillii, pseudo-verticillata, and steeni-

sii, Of more commonly shortly acute, acuminate to caudate. The

texture of the leaf in Q. kinabaluensis, nivea, percoriacea, pseudo-:

verticillata, steenisii, and valdinervosa is thickly coriaceous, whilst

in Q. argentata, chrysotricha, elmeri, gaharuensis, gemelliflora,.

kerangasensis, lineata, merrillii, oidocarpa, and subsericea is thinly’

coriaceous. In Q. sumatrana and treubiana, the texture is thinly’

chartaceous.

Except for Q. chrysotricha, kinabaluensis, lowii, and subsericea,,.

the midrib and lateral nerves are strongly prominent beneath. On.

the upper surface, these nerves are either slightly raised (Q. elmeri,.

gaharuensis, gemelliflora,: kinabaluensis, lineata, steenisii and.

treubiana), or flattened to impressed in the other species. The num--

ber of lateral nerves varies considerably in the species, but fairly

consistent numbers may be found in three species groups, viz..

Q. argentata, lineata, oidocarpa, pseudo-verticillata, and valdiner-

vosa which have c. 10-20 pairs; Q. chrysotricha, kinabaluensis,

lowii, and merrillit which have c. 4-6 pairs, and the rest of the

species which have c. 8-15 pairs. The angles between the lateral

nerves and the midrib vary from 30° to 75°, with the majority of

the species falling between 45° and 60°. In Q. oidocarpa and

pseudo-verticillata the angle is about 30°-45°, whilst in Q. argen-

tata, chrysotricha, elmeri, gemelliflora, kerangasensis, nivea, and.

valdinervosa is c. 60°-70°. The reticulation between the lateral

nerves is generally parallel, very rarely more or less anastomosing

(Q. lowii), and it is either distinct beneath (Q. gaharuensis, lineata,.

oidocarpa, percoriacea, steenisii, and valdinervosa) or more com--.

monly obscure. In Q. lineata, oidocarpa and steenisii the reticula--

tion is parallel and dense, whilst in the other species it is lax.

The type and density of the pubescence on the lower surface of

the leaf is in some species rather consistent. This character how-

ever should be used with a great caution. In Q. argentata, chryso-

tricha, gemelliflora, kerangasensis, and valdinervosa, the mature.

362 Gardens’ Bulletin, Singapore — XXII (1968)

leaves are completely glabrous on both surfaces, whilst in Q. gaha-

ruensis, lowii, nivea, oidocarpa, percoriacea, and subsericea, the

lower surface of the adult leaf is densely or sparsely adpressed

tomentose. In Q. elmeri, kinabaluensis, lineata, merrillii, pseudo-

verticillata, steenisii, sumatrana, and treubiana, the lower leaf

surface is densely or sparsely stiff or woolly pubescent. On account

of this character Q. lineata may be distinguished from the othet

species by its dense layer of straight, adpressed, simple hairs on

the lower surface of the leaf, and Q. steenisii differs from the other

species by its thick layer of yellowish-brown, woolly hairs which

are caducous.

Inflorescene (Fig. 1, f. 1-3): Similar to that of the other

anemophilous genera in Fagaceae (Fagus, Nothofagus), the inflore-

scence in the genus Quercus is either male or female. In Quercus,

however, the inflorescence is catkin-like (condensed raceme of

cymes), whilst in Fagus and Nothofagus it is simply a 1-many-

flowered, somewhat modified dichasium. In the Malesian species

the male inflorescence is as a rule compound, i.e. the flowers are

in 3-4-flowered dichasial clusters, and very rarely simple, i.e.

when the flowers are solitary along the catkin-axis; it is either

solitary in the axil of a lower leaf (Q. gaharuensis, gemelliflora,

kinabaluensis, lineata, lowii, steenisii, subsericea, treubiana, and

valdinervosa) or in a paniculate clusters on the lateral or subter-

minal new shoot (Q. argentata, kerangasensis, nivea, and oido-

carpa). The rachis may be very thin and slender (Q. lowii), or

more commonly rather thick; in both cases it is either stellate

tomentellous or sericeous. The female inflorescence is always sim-

ple, solitary in the axil of a higher leaf, unbranched and densely

minutely tomentose.

Flowers (Fig. 1, f. 4-7): in each dichasial cluster, the flowers

are always protected by ovate-acute, caducous bracts. Male flower:

the perianth is densely minutely simple tomentose, 4—6-lobed, with

the lobes connate at base; stamens 4—6, rarely up to 9 (Q. valdiner-

vosa); filaments slender, filiform, glabrous or minutely puberulous

at base; anthers c. 0.5-1 mm long, globular reniform, basifixed,

longitudinally dehiscent, 2—-locular, 4-lobed; ovary usually absent

or abortive and represented by a cluster of stiff-simple hairs

(Q. gaharuensis, lineata, lowii, steenisii and treubiana). Female

flowers: the ovary is obtusely triangular, 3—-locular, each locule

contains 2 ovules; perianth (4)-6-(9)-lobed; staminodes absent in

most species but 5—7 in Q. gemelliflora, nivea, steenisii, and valdi-

nervosa; styles 3-(6), recurved and free or rarely connate (Q. nivea);

stigmas always broadly capitate.

Pollen grains: from the study of no less than 100 species of

Fagaceae, belonging to seven different genera, it appears that three

different types of pollen grains exist,* namely Quercus-type (Fagus,

Quercus, Trigonobalanus), Castanea-type (Castanea, Castanopsis,

Lithocarpus) and Nothofagus-type.

* The characters used here are the size, general appearance of the

pollen, the structure of the exine, and the number of apertures and pores.

Soepadmo — Quercus subgen. Cyclobalanopsis 363

my ye,

ape

yr 00

ian

XK @

@M 99 o000D

“

Ch= CUPULE

Ee de fe A = PRIMARY FLOWER 2 é ‘ooconmaty 26467

O = MALE FLOWER , pl = eee ee

B= Secondary FLOWER nga Saye

@,r= RACH/S Be 94

Fig. 1:

1 = diagram of a male inflorescence.

2 = diagram of a female inflorescence.

3 = diagram of a paniculate cluster of male inflorescences.

4 = 3-flowered male dichasium.

5 = diagram of figure 4.

6 = a pistillate flower with the perianth removed.

7 = diagram of a female flower.

364 Gardens’ Bulletin, Singapore — XXII (1968)

Except for the pollen type in the genus Fagus therefore this gene-

ric grouping concurs with that based on the inflorescence and

floral characters. In the Quercus-group the pollen is prolate-

spheroidal to suboblate, 3— or rarely 4~angular in polar view;

usually with 3 apertures only, or with 3 apertures and 3 pores as

well; average size c. 30 x 20 yp. The exine c. 1-2.5 » thick, more or

less scabrate. Furrows elongate and the pores are simple or in

Trigonobalanus doichangensis it is striate in equatorial optical

‘section; circular or more or less obtuse-triangular in outline. In

the Castanea-group the pollen is prolate-perprolate to ellipsoid,

cc. 18 x 10 »; always with 3 apertures and 3 pores. The exine is

«. 0.4-1 » thick, and relatively smooth. The furrows are narrow

-and usually with pointed ends. The pores are always simple, well-

‘defined and equatorially elongated. In Nothofagus, the pollen is

‘distinctly oblate to peroblate (flattened) in equatorial view, and

-4-9-angular in polar view. Polar diameter ranges from 9 to 16 n,

equatorial diameter from 20 to 65 pw. The exine c. 0.7-1.6 » in

‘thickness, and distinctly echinate. The furrows are simple, meri-

‘dionally elongated, and usually with rounded ends.

The pollen of Quercus differs from that of Fagus by its smaller

‘size and its finer reticulation; from that of Trigonobalanus by its

‘thinner exine, simpler and sometimes vestigial pores, and to

‘some extent also by its more circular outline in polar view. The

‘pollen in Q. lowii has very well-developed pores and apertures,

‘whilst that of Q. lineata, steenisii, and valdinervosa have vestigial

‘pores and well-defined apertures. In the other Malesian species

‘investigated the pollen has no pore. From this brief survey it seems

‘that pollen morphology supports the separation of Lithocarpus

‘from Quercus, Nothofagus from Fagus, but it does not support

ithe generic splits proposed by various workers in Castanopsis,

Lithocarpus, Nothofagus and Quercus. Pollen morphology also

‘supports Forman’s idea to place Quercus and Trigonobalanus in

ithe subfamily Quercoideae, and Castanea, Castanopsis and Litho-

«carpus in the subfamily Castaneoideae.

Cupule: similar to the other genera of Fagaceae, the most char-

cacteristic feature of the genus Quercus, is the presence of cupule

‘enclosing the basal part of the fruit. In the earlier works, this

‘cupule has been interpreted as an involucre of bracts fused to-

gether with only their tips appearing as scales, lamellae or spines;

of as an axial structure formed by an intercalary outgrowth of the

‘meristematic tissue below the flower bearing modified leaves

(scales or spines). In 1878 Eichler suggested a rather different

interpretation, in which the cupule was regarded as a structure

‘built up by the modification of the four bracteoles in the dichasium,

and the scales or spines as simply external emergences. In the

most recent works (Berridge, 1914, Langdon 1939, 1947, and

‘Brett 1964) the cupule is considered as an axial structure formed

iby the modified axes of the next higher order flowers in the

‘dichasium, and the scales, lamellae or spines as external appen-

dages.

Soepadmo — Quercus subgen. Cyclobalanopsis 365

Development of the cupule: in the earlier stages of the pistillate

flower development, the cupule primodium in Quercus, appears.

as a collar subtending the flower, and it is situated between the

flower and the bracts. The growth of the cupule is at first faster

than that of the flower, so that the partly developed fruit is

completely enclosed by the cupule, except for the perianthodium

and the styles. The faster growth of the cupule in the earlier stages.

gives rise to the ring-like marks on the perianthodium. Soon after

fertilization, the elongation of the cupule slows down or stops before

the fruit is fully developed. At maturity therefore the cupule usually

covers not more than a quarter to two thirds of the fruit. As a

rule the cupule in Quercus encloses one fruit only, but in Q. lowii

it has been observed that in some specimens the cupule contains.

more than one fruit, usually two. There is, however, a clear

evidence that this abnormality is caused by injury during the

development of the fruit and cupule. A similar condition has been

reported by Hjelmqvist in 1948 in Q. acuta from Japan, which

induced him to conclude that the cupule in Quercus subgen. Cyclo-

balanopsis must have a different origin from that of subgenus.

Quercus. Another abnormality which should be mentioned here

is that, while in most species the cupules are solitary along the

rachis, in a few specimens of Q. gemelliflora from Java two or

three cupules are grouped together in a dichasial cluster as normal-

ly occur in the genus Lithocarpus, It is not certain, however, whe-

ther this abnormality is a product of a teratological growth or that

it is a genuine morphological feature.

Anatomy of the cupule: this investigation was performed mainly

to find out whether the distribution of the vascular bundles in the

cupule gives any clue towards a better understanding on the nature

and probable evolutionary lines of the cupule in Fagaceae. The

cupules were cut by razorblades in three different planes, viz.

longitudinal sections following the axis of the catkin, longitudinal

sections across the axis, and cross-sections through the cupules.

The 0.5 to 2 mm thick sections then were treated with lactic acid

and heated for about five minutes. By this simple method clear

sections showing the distribution of the vascular supply were:

obtained.

In a series of longitudinal sections following the axis of the

catkin (Fig. 2, f. 1), the vascular tissue at first appears as a cen-

tral cylinder (M) originating from the vascular tissue supplying the

rachis. Outside this main vascular cylinder, several smaller traces.

may be seen. These smaller traces supply the bracts and bracteoles,

whilst the main vascular cylinder is leading to the flower and

cupule. At the level above the insertion of the bract (qa), i.e. from.

the lower part of the main vascular column, arise several ascendant

lateral traces which are orientated and branched off towards the

cupule. At the level of the insertion of. the flower on the cupule,.

the main vascular cylinder is further subdivided into several

separate bundles which supply the ovary, perianth, staminodes

(if present) and the styles. In a series of longitudinal sections across

the rachis (Fig. 2, f. 2) the organization of the vascular tissue is

366 Gardens’ Bulletin, Singapore — XXII (1968 }

exactly the same. In a cross-section through the basal part of the

cupule (or peduncle), the vascular cylinder appears as a triangular

ring of loose vascular traces (Fig. 2, f. 3). At the level of the

insertion of the bracteoles (b) this vascular column becomes

laterally elongated (Fig. 2, f. 4). Above this level, i.e. at the basal

part of the cupule proper, the main vascular cylinder is subdivided

into a central tract (M1) which directly supplies the flower, and

numerous scattered traces which give rise to the vascularization

of the cupule (Fig. 2, f. 5). At the region of the insertion of the

fruits (Fig. 2, f. 6) the central vascular tract undergoes a further

reorganization and is divided into at least 13 smaller groups of

vascular traces. One of these vascular traces supplies the placental

tissue, and the others are distributed in the periphery of the fruit

wall and leading to the perianth, staminodes and styles. Further

up from this level, where the fruit wall is completely free from

the cupule, the separation of the central vascular tract is more pro-

minent. In the fruit wall the vascular traces situated at the corners

of the locules are clustered in threes, whilst those placed between

the corners of the locules are solitary (Fig. 2, f. 7). At this level it

may be noticed that the traces distributed in the cupule send off

more smaller branches from which the vascular supply of the

lamellae covering the cupule originates. It may be mentioned here

that the pattern of vascular distribution in the cupule of Quercus

just described is exactly the same as that of a solitary cupule in

the genus Lithocarpus.

The nature of the cupule: The cup- or saucer-shaped, solitary

cupule in Quercus is here regarded as an axial structure formed by

the intercalary outgrowth of the meristematic tissue below the

flower, bearing scales or lamellae as external emergences.

The evolutionary interpretation of the cupule: in 1878 Eichler

assumed that the cupule in Quercus must have been derived from

that of Fagus-Castanea-type, through a complete loss of the two

lateral flowers in the originally 3-flowered dichasium, followed by

a complete fusion of the four modified bracteoles. Prantl (1887),

Celakovsky (1887, 1890, 1893), Palibin (1909), Lendner (1916),

Troll (1926), Hjelmqvist (1948), on the other hand, regarded the

cupule in Quercus as a type derived from a dichasial cluster of 3

cupules in the genus Lithocarpus, by means of complete reduction

of the two lateral flowers and their cupules. Recently Brett (1964)

suggested a different interpretation in which the cupule in Quercus

subgen. Cyclobalanopsis (considered by him as a distinct genus):

is regarded as derived from a cupule type occurring in Castanopsis,

through a similar process of reduction to that of Castanopsis

fissa-group, and that of the subgenus Quercus from a 3-flowered

dichasial cluster of cupules in Lithocarpus through a similar way

of reduction as has been proposed by Prantl and others. In his.

recent paper, Forman (1966, in press) considers the cupule in

Quercus (incl. Cyclobalanopsis) as derived from a 3-lobed, 1-flow-

ered cupule like that in Trigonobalanus doichangensis, by a com-

plete fusion of the cupule lobes.

Soepadmo — Quercus subgen.

Fig. 2:

3-7

Cyclobalanopsis 367

a longitudinal section through the rachis of a young cupule.

a longitudinal section across the rachis of a young cupule.

= cross-sections of a young

figure 2.

cupule through the planes shown in

368 Gardens’ Bulletin, Singapore — XXII (1968)

I have pointed out elsewhere that from the earlier stages of

development, there is no indication that the cupule in Quercus

consists of separate lobes or that the cupule normally encloses

more than one fruit. I also mentioned that the anatomical feature

of the cupule in Quercus is exactly the same as that of a solitary

cupule in Lithocarpus. Moreover, in Lithocarpus there is a

gradual reduction from a dichasial cluster of cupules to a solitary

condition by a complete abortion of the higher order flowers and

their cupules. Therefore I conclude that the cupule in Quercus

is closely allied to that of Lithocarpus, and it must have been

derived from one of the types occurring in the latter. The cupule

in these two genera represents a type distinct from that of the other

genera in Fagaceae. The cupule in Quercus and Lithocarpus I con-

sider as a flower-cupule, but that of Castanea, Castanopsis, Fagus,

Nothofagus, and Trigonobalanus as a dichasium-cupule.*

The taxonomic significance of the cupule: in Malesia the charac-

ters of the cupule are very useful for identification. The cupule is

either saucer-shaped (Q. elmeri, percoriacea, steenisii, subsericea

and sumatrana) or deeply cup-shaped with rounded or attenuate

base (in the other species). In Q. argentata, gaharuensis, gemelli-

flora, lineata, nivea, oidocarpa, pseudo-verticillata, treubiana, and

valdinervosa, the cupule is c. 1.5-3.5 cm deep and 2-4 cm in

diameter, whilst in Q. chrysotricha, kerangasensis, kinabaluensis,

lowii and merrillii the cupule is c. 0.7-1.2 cm deep and 1-1.5 cm

in diameter. In the other species the cupule is c. 0.5-1.2 cm deep

and 2-2.5 cm in diameter. The rim of the cupule is c. 0.5-4 mm

thick in Q. elmeri, gemelliflora, kerangasensis, kinabaluensis,

lineata, steenisii, treubiana and valdinervosa, and thinner than 0.5

mm in the other species. The lamellae are c. 8-12 in Q. argentata,

lineata, oidocarpa, pseudo-verticillata, and treubiana, and c. 5-8

in the other species. The lower lamellae are usually denticulate,

whilst the upper ones are entire. The margin of the lamellae may

be free or sometimes slightly connate. In most species the outer

surface of the cupule is densely puberulous or stellate-tomentose

and early glabrescent; the inner surface is densely covered with

thick layer of silky to dark-brown, stiff simple hairs.

Fruit: according to Trelease (1924), Camus (1938), and Schwarz

(1936 a, b) the maturity of the fruit in the genus Quercus may be

annual (subgen. Cyclobalanopsis and subgen. Quercus sect. Macro-

balanus, Mesobalanus and Lepidobalanus) or biennial (subgen.

Quercus sect. Erythrobalanus, Protobalanus, and Cerris). In Male-

sia the data on this particular character are too scanty for a con-

clusion,

*(See Forman 1966, in press).

Soepadmo — Quercus subgen. Cyclobalanopsis 369

In the Malesian species the fruit is ovoid-globose, ovoid-

conical or ovoid-cylindrical. The young fruit is usually densely

sericeous, and gradually becomes glabrous towards maturity. In

Q. argentata, chrysotricha, gaharuensis, gemelliflora, lineata, mer-

rillii, sumatrana and valdinervosa the apex of the fruit is distinctly

attenuate-acute, whilst in the other species it is either attenuate-

rounded or depressed. The base of the fruit in most species is

convex, in Q. sumatrana it is concave, and in Q. elmeri, oidocarpa

and: pseudo-verticillata it is either convex or flat. The fruit-wall

is c. 1-2 mm thick, and usually woolly tomentose inside. The

anatomical structure of the fruit-wall in Quercus is the same as

that of Castanea, Castanopsis and Lithocarpus, i.e. composed of at

least five different layers of cells. These layers are, from outside

inwards: 1), the outer epidermal layer consisting of a single row

of rectangular cells; 2). the pallisade layer which consists of several

rows of radially elongated, thick-walled cells; 3). the outer paren-

chymatous layer which is usually very much thicker than the other

layers and composed of more or less isodiametric, thin-walled

cells, and containing the vascular tissue of the fruit-wall and the

other floral parts; 4). the inner parenchymatous layer which has

smaller cells than that of the former; 5). the inner epidermal layer

consisting of a row of flattened, rectangular, thin-walled cells

(Fig. 3 & 4). The anatomical structure of the fruit-wall just

described differs from that of the other genera in Fagaceae by the

position and type of the supporting layer, i.e. the layer which is

composed of thick-walled cells. In Nothofagus (Fig. 5, f. 1-2)

this supporting layer is situated near the centre of the fruit-wall,

and consists of several rows of tranversely elongated thick-walled

cells. In Fagus and Trigonobalanus, the supporting tissue is situat-

ed directly beneath the outer epidermal layer, and is composed of

several rows of more or less isodiametric, polygonal, thick-walled

cells (Fig 5, f. 3-4). In the examined specimens, the cotyledons

are flat-convex, and fill the entire space of the surviving locule.

The abortive ovules are apical, and the germination is hypogeal

as in Castanea, Castanopsis and Lithocarpus.

Chromosomes number: chromosome counts in various species of

Quercus subgen. Quercus have been made by several cytologists

such as Ghimpu (1929), Sax (1930), Sugira (1931), Yamazaki

(1936), and Duffield (1940). The diploid number reported is 24,

similar to that of Castanea (Jaynes, 1962), and Fagus (Jaretzky,

1930). In the subgen. Cyclobalanopsis on the contrary no chromo-

some count has been made. The only information so far available

is that given by Mr. Kwiton Jong (formerly Dept. of Botany,

University of Malaya, Kuala Lumpur). He made some preliminary

counts in several species of Castanopsis, Lithocarpus, Quercus

{subgen. Cyclobalanopsis) and Trigonobalanus, and found that the

diploid number in the first three’ genera is also 24, whilst that of

Trigonobalanus verticillata is not yet fully understood. For

comparison, it may be recalled here that in 1965, Armstrong &

Wylie reported a diploid number of 26 in the four species of

Nothofagus (fusca, truncata, solandri, and menziesii) from New

Zealand.

370

epidermal layer.

Gardens’ Bulletin, Singapore — XXII (1968)

8 A oe: .

Jord) Scope ot a

<> <= a”. \ Oar, {9} +4 He

f i } "a Ws ) =

BES ae aid sean OU

SSS SAO COHN tek

SS RET |

Kas SoA S2@0;8 SFr sag te. Co Hilit =

0.5 mm.

PERTH TOO EY

SS Raemengnoneasne:

i=—— Wad VeEg® OU ab en@ Br nn 60;

‘=a Sates Seance OG) BA tT WH et nN

Sea Ogracalset ert Pagtecoasys.::

|= eiitreNesileprstetsseaue’:

EEO OO Eg

: ¥

ee \

’ i

Fig. 3: cross-sections of the fruit-wall.

1 = Q. oidocarpa;

outer epidermal layer; = pallisade layer; c = outer paren-

chymatous layer; d = inner parenchymatous layer; e = inner

2 = Q. suber; 3 = Castanopsis chrysophylla; a =

aT!

Soepadmo — Quercus subgen. Cyclobalanopsis

Se Q

iigeereger

oe {2

. e=ee Oe Os

=o®

Sas

Bre

pas

See

Castanea sativa.

on,

‘&

ogc

ee J

cross-sections of the fruit-wall.

1 = L. philippinensis; 2 = Castanopsis javanica; 3

Fig. 4:

oi2 Gardens’ Bulletin, Singapore — XXII (1968)

POO RST AIHA

ar BO i Me

KOOL JOR vita

oo® JS) i

f Soy} h lin

Ocal

ay 1h

WoO, Sy

‘750 (8 ‘eo

@ i

20 '\ a = 4

ony, | Y Pr

wal

HRD LH) Nk

be DESK WON H\

Ge |

Bear |

BVena'|

\\ “gil LA oe

(5@O)( Ri V'seeen(|(

ICOS\ ME arexes:\;

Ral, eOne/

] ! \

\ i

Fig. 5: cross-sections of the fruit-wall.

1 = Nothofagus resinosa; 2 = Nothofagus antarctica; 3 = Trigono-

balanus verticillata; 4 = Fagus sylvatica.

Soepadmo — Quercus subgen. Cyclobalanopsis 373

TAXONOMY

The systematic position of the genus Quercus: on account of the

inflorescence and stigma characters, Oersted (1867) subdivided the

family Fagaceae into two subfamilies, viz. Quercinae and Cas-

taninae. He included Quercus, Cyclobalanopsis and Fagus (incl.

Nothofagus) in Quercinae, and Castanea, Castanopsis, Pasania and

Cyclobalanus (incl. Lithocarpus) in the subfam. Castaninae. He

distinguished the subfam. Castaninae from Quercinae by its rigid

and erect male inflorescence and terminal punctiform stigmas.

In 1871, Oersted revised his classification and excluded Fagus

and Nothofagus (in this paper Nothofagus was regarded as a

separate genus) from the subfam. Quercinae, and placed them in

a separate subfamily, Fagineae. In this new classification the sub-

fam. Quercinae was maintained to include Quercus and Cycloba-

lanopsis. In the subfam. Castaninae, the genus Castanopsis was

reduced to a subgeneric rank under Castanea, and Pasania was

kept separate from Cyclobalanus (incl. Lithocarpus). Oersted dis-

tinguished the subfam. Castaninae from Fagineae and Quercinae

by its terminal punctiform stigmas, and Fagineae from Quercinae

by its plicate cotyledons and epigeal germination. Amongst later

authors who basically accepted Oersted’s subdivision of the family

Fagaceae are Schwarz (1936 a, b), Melchior (1964), and Forman

(1964). However, these later three workers added more characters

which were not mentioned by Oersted or they used different

combinations of characters to distinguish one subfamily from the

others.

In 1894, Prantl introduced a rather different classification, in

which he placed Quercus (incl. Cyclobalanopsis), Castanea (incl.

Castanopsis) and Pasania (incl. Cyclobalanus and Lithocarpus) in

the subfam. Castaneae, and included Fagus and Nothofagus in the

subfam. Fageae. He distinguished these two subfamilies as follows:

Subfam. Fageae: flowers arranged in dichasial clusters, rarely

solitary; fruit triangular; cotyledons plicate. :

Subfam. Castaneae: flowers arranged in dichasial clusters or

solitary along the catkin-axis; fruit rounded; cotyledons

not plicate.

This classification was adopted by Dalla Torre & Harms (1907),

Schottky (1912), and by Trelease (1924). A rather similar type of

classification was proposed by Brett in 1964, though this author

based his system mainly on the cupule characters.

In 1964, Forman suggested another type of classification which

seems to represent a compromised scheme between Oersted’s

(1871) and Prantl’s arrangements. Forman agreed with Oersted in

subdividing the family Fagaceae into three different subfamilies,

_ ae

374 Gardens’ Bulletin, Singapore — XXII (1968)

but used the inflorescence and floral characters to distinguish

subfam. Fagoideae from Castaneoideae and Quercoideae as was

done by Prantl. Forman distinguished subfam. Fagoideae from the

other subfamilies by the inflorescence which consists of a 1-many-

flowered, axillary dichasial cluster, in contrast to that of subfam.

Castaneoideae and Quercoideae which is catkin-like, and com-

posed of dichasial clusters of flowers or solitary flowers. Subfamily

Castaneoideae was distinguished from Quercoideae by the charac-

ters of the styles, stigmas, male inflorescence and flower, anthers,

and the cupule. Trigonobalanus and Quercus were placed in the

subfam. Quercoideae.

From the preceding discussion it is clear that the main difference

between Oersted’s (1871) and Prantl’s classifications hingers on

the question whether the genus Quercus (incl. Cyclobalanopsis)

should be regarded as representing a distinct subfamily or should it

be included in the same subfamily with Castanea, Castanopsis, and

Lithocarpus (incl. Pasania and Cyclobalanus). It is also evident

that though at the subfamily level Forman’s classification is essen-

tially similar to that of Oersted, at the generic level, except for

the status of Castanopsis, it agrees better with Prantl’s system.

Before deciding which of these three different classifications

should be adopted here, it is necessary to discuss the generic

affinity between Quercus and the other genera in Fagaceae. I have

indicated elsewhere that Quercus possess many important consist-

ent characters, viz. that of the inflorescence, flower, pollen grains,

which distinguish this genus not only from Lithocarpus (s.1.) but

also from Castanea and Castanopsis. The difference between

Quercus and Castanea-Castanopsis ate further strengthened by the

characters of the cupule. In Quercus the cupule is always cup- or

saucer-shaped, not lobed nor spiny; it encloses one fruit only, and

is here regarded as flower cupule. In the genera Castanea and

Castanopsis the cupule is a dichasium cupule, and in most

species it is neither cup- nor saucer-shaped, but completely covering

the fruits and eventually splits either into a definite number of

lobes or irregularly; it encloses one to many fruits, and in the

majority of the species it is either spiny or tubercled. In the earlier

works, however, there has been some confusion with regard to the

generic limit between Quercus and Castanopsis.

Species of Quercus are readily distinguished from those of Fagus

and Nothofagus by their catkin-like inflorescence, cup- or saucer-

shaped, not lobed cupule enclosing one fruit only, rounded ovary,

flat-convex cotyledons, hypogeal germination, anatomical structure

of the fruit-wall and the wood, and pollen characters. Taking all

these characters into consideration, the genus Quercus should be

placed in a different subfamily from Fagus and Nothofagus.

Soepadmo — Quercus subgen. Cyclobalanopsis 375

The affinity between Quercus and Trigonobalanus has been dis-

cussed in detail by Forman (1964). Amongst the characters men-

tioned by him which indicate the close relationship between these

two genera are the presence of the catkin-like inflorescence, capitate

stigmas, male flower with 6 stamens, anthers which are c. 0.5—-1 mm

long and more or less basifixed, and the absence of pistillode in

the male flower. It is interesting to note here that the conclusion

reached by Forman which was based on the characters just men-

tioned was subsequently supported by Cutler, who in 1964 investi-

gated the wood anatomy of these two genera. Cutler, however, also

stressed that the genera of Fagaceae are not always clearly defined

from one another by the wood characters alone. In his classifica-

tion, Forman regarded the lobed cupule and the triangular fruit in

Trigonobalanus, and also the presence of androgynous and rigid

and much branched male inflorescence in 7. verticilldta as ancient

characters persisting in the subfam. Quercoideae rather than as

evidence for a closed generic affinity between Trigonobalanus and

the genera included in the subfam. Fagoideae or Castaneoideae. 1

have mentioned elsewhere that except for the detailed structure of

the pores in the pollen of T. doichangensis, as for the pollen,

Trigonobalanus agrees better with Quercus or Fagus than with the

other genera of Fagaceae. As far as the characters of the flower are

concerned therefore Forman’s classification seems to be well-found-

ed. As for the cupule and fruit characters, however, Trigonobalanus

has nothing in common with Quercus, but it possess many charac-

ters in common with the subfam. Fagoideae. These characters.

are: the lobed cupule enclosing one to many fruits, trigonous and

almost winged fruits, epigeal germination, and probably also the

plicate cotyledons. Melchior (1964) included Trigonobalanus in

subfam. Fagoideae, but his argument is not very convincing, as he

indiscriminately mixed up the characters of the inflorescence,

flower, cupule and fruit, which in combination are not always clear-

ly defined in the subfamilies recognized by him. Great credit,

however, should be given to Professor Melchior for suggesting the

alternative systematic position of this enigmatic new genus.

Other characters which seem to contradict the disposition of

Trigonobalanus in subfam. Quercoideae ate the presence of andro-

gynous and much-branched, erect, male inflorescences in 7. verti-

cillata. These characters which are usually associated with entomo-

phily, persist in Fagaceae only in the Castaneoideae. Taking all the

known characters into consideration, however, it is clear that

placing Trigonobalanus in the subfam. Castaneoideae is out of

question. It remains to be settled in the future, therefore, whether

the genus Trigonobalanus should be placed in Fagoideae or Quer-

coideae.

Conclusion: on account of the inflorescence and floral characters,

I agree with Forman (1964) to recognize in the family Fagaceae

three subfamilies, viz. Castaneoideae (Castanea, Castanopsis, and

Lithocarpus s.1.), Fagoideae (Fagus and Nothofagus), and Quer-

coideae (Quercus incl. Cyclobalanopsis, Macrobalanus and Erythro-

balanus). A reservation should be made with regard to the position

of 1vigonobalanus until this genus is better understood.

376 Gardens’ Bulletin, Singapore — XXII (1968)

NOTE ON THE IDENTITY OF SEVERAL MALESIAN

SPECIES |

Q. oidocarpa Korth., Q. brevistyla A. Camus, and Q. gaharuensis

Soepadmo: the type of Q. oidocarpa, preserved in the Rijksher-

barium, Leiden, was collected by Korthals in 1835 from the vicinity

of Mt. Malintang in Central Sumatra. In 1888, Hooker f. cited

several specimens from Malaya and Borneo under this species,

which were accepted by most of the subsequent authors such as

King (1889), Brandis (1906), Gamble (1915), Ridley (1924), Merrill

(1921), and A. Camus (1938). In the present study, however, it was

found that these specimens are quite different from Korthals’

species, and they agree very well. with Q. gaharuensis I described in

April 1966. In 1933, A. Camus described a new species, viz.

Q. brevistyla, based on specimens collected by Syed Ali (Kep.

22574) from Fraser’s Hill, Malaya. She apparently did not see

Korthals’ specimens; I found that Syed Ali’s specimens match very

well those of Korthals. Therefore. I reduce Q. brevistyla to Q.

oidocarpa.

Q. semiserrata Roxb.

In his account on the Indo-Malayan species of Quercus and

Castanopsis, King (1889) erroneously reduced Q. horsfieldii Miquel

(1856) from Bangka (Horsfield 11) and Sumatra (Diepenhorst 42)

to Q. semiserrata Roxb. After comparing these two specimens with

those of Q. semiserrata, I came to the conclusion that the Suma-

tran plants do not belong here, but should be included in Q.

gemelliflora Bl. In 1904, Koorders & Valeton accepted King’s

reduction and credited several specimens from Java collected by

Koorders to Q. semiserrata. I place Koorders’ specimens in

Q. oidocarpa.

Q. lowii King

King’s original description of this species was based on the

specimens collected by Low (s.n., in Herbarium Calcutta) from Mt.

Kinabalu in North Borneo. King (1889) indicated that this species

may be distinguished by its coriaceous leaves which are narrowly

ovate, bluntly acuminate, remotely serrate in the apical half, and

with the base either rounded or slightly cordate. The lower surface

of the leaf was described as minutely cinereous-tomentose, and

the cupule as cup-shaped, c. 1.5 cm in diameter and 0.75 cm in

height, minutely tomentose and covering c. 4 part of the ovoid-

cylindrical fruit. In King’s illustrations (l.c., Plate 21 B, f. 4-6)

the leaf is figured as sharply and deeply serrate in the upper half.

In the actual specimen, however, the leaf margin is not sharply

and deeply serrate as in King’s figures, but it is either shallowly,

remotely serrate in the apical half or almost entire. King’s exag-

gerated figures on the leaf margin have given rise to some doubt to

most later botanists who tried to identify specimens of Quercus

recently collected from Mt. Kinabalu. One of these even thought

that Q. lowii must be a very rare species. From the specimens

examined, however, I found that this species is not very rare as

Soepadmo — Quercus subgen. Cyclobalanopsis 377

thas been assumed, but is very common in Mt. Kinabalu, especially

at 1000-2000 m altitude. In the specimens studied, it appears

that there are two forms, which differ from one another by their

Jeaf pubescence. In one form the leaf is densely, minutely stellate-

tomentose beneath, and almost completely glabrous in the other.

‘The pubescent form is represented by several specimens (including

the type), e.g. Clemens 10936, 32448, Meijer SAN. 24121, 28808,

Chew & Corner RSNB. 4363, 4500, 4662, and Sow Kep. 71649.

Except for Sow’s specimen, all of these are from c. 1500-2500 m

altitude. The majority of the examined specimens belong to the

glabrous form, and most of them are from lower altitude. Clemens

30967, 32933, and 40231 are intermediates, with the leaf being

neither glabrous nor densely, minutely stellate-tomentose, but

-sparsely stellate-tomentose. These three specimens were collected

mainly between 1200 and 1500 m altitude. The cupules and fruits

of the recently collected specimens (e.g. SAN. 28808 and RSNB.

4500) agree very well with King’s description and illustration.

~Therefore, I consider the glabrous and pubescent forms as con-

-specific, representing King’s Q. lowii. This species is closely allied

-to Q. subsericea A. Camus, which also occurs in Mt. Kinabalu, but

.Q. lowii differs from the latter by its thick-coriaceous leaves which

-have rounded to cordate base, sharply acuminate apex, more or

“less net-shaped reticulation, and by its cup-shaped, densely

-tomentose cupule.

-Q. mespilifolia Wall. ex A.DC. var. borneensis Heine

Q. mespilifolia was first mentioned as a nomen nudum in

Wallich’s Catalogue (no. 2766, 1828). It extends from S.W. China

(Yunnan) to the Northern parts of Burma, Laos and Vietnam

(Tonkin). In 1935, A. Camus renamed this species Q. mespili-

folioides, as she was of the opinion that Wallich’s name was

invalidly published. In 1864, however, A. DeCandolle validated

Wallich’s name, so that Q. mespilifolioides A. Camus is a super-

-fluous name for Wallich’s species. In 1953, Heine described Q.

mespilifolia var. borneensis, based on Clemens 31238, 31317, and

31459, from Mt. Kinabalu. Heine distinguished the new variety

by its densely ferrugineous-pubescent leaves which are serrate in

;the apical +4 part. Recent collections from North Borneo and

Sarawak added more information about this new variety, in which

female inflorescences and fruits were gathered, unknown to Heine.

After comparing the type specimens of Q. mespilifolia from the

“Upper Burma and some other specimens credited by King (1889)

‘to this species with the Bornean materials, I have come to the

.conclusion that the Bornean plant represent a distinct species. It

differs from Q. mespilifolia by the characters of the cupule, fruit,

and leaf. The specific epithet borneensis, however, is preoccupied

by Q. borneensis Merr. (1922), the basionym of Lithocarpus bor-

-neensis (Merr.) Rehder. I recognize this species as Q. valdinervosa

-to indicate the strong nervation of the leaf. Clemens 31317 is

chosen as the lectotype, and Anderson S. 4547 as the paratype

«(fruits).

378 Gardens’ Bulletin, Singapore — XXII (1968)

Q. gemelliflora BI.

This species was first described and figured by Blume (1823)

based on mixed specimens consisting of a leafy twig which belong

to Quercus and fruits represent a species of Lithocarpus (cf.

Blume’s illustration in Verh. Bat. Genootsch. 9, f. 6, 1823). In the

type specimen at Leiden, I could not trace any fruit attached, and

I discovered that the leaves are exactly the same as those of Q. tur-

binata BI. (1825). Therefore, I reduce Q. turbinata to the synonymy

of Q. gemelliflora, and choose the remaining sterile specimen

(Blume s.n.) preserved at Leiden as the type.

In 1847, Endlicher erroneously renamed Q. turbinata Bl. as

Q. merkusii, as he thought that the name turbinata had been used

by Roxburgh (1832) for a different species from India, apparently

not realizing that this was a later homonym. Therefore, Q. merkusii

is a superfluous name for Q. turbinata, and accordingly it beeomes

another synonym of Q. gemelliflora. Roxburgh’s Q. turbinata was

eventually reduced by King (1889) to Q. thomsonii Mig. which

in 1919 was transferred to the genus Lithocarpus by Rehder. In

1915, Gamble described under Q. turbinata, a new variety crassila-

mellata, based on Wray 1532 from Johore, Malaya. A Camus

(1931) raised this variety to a specific rank. I reduce this variety to

Q. gemelliflora Bl.

Q. hendersoniana A. Camus

This species was published by A. Camus in 1932, based on

Henderson 18053 from Cameron Highlands, Pahang, Malaya. I

reduce this species to Q. lineata BI.

The other species of Quercus from Malesia which are not

described in this paper are included in the genus Lithocarpus BI.

GEOGRAPHICAL DISTRIBUTION

The distribution of Quercus subgen. Quercus extends from

southern Scandinavia throughout Europe to northern Africa

(Morocco, Algeria and Tunisia), Lebanon, Turkey, and spreads

eastwards to the Caucasus ranges, northern Syria, Iraq, Persia,

Afghanistan, Pakistan, India, Burma, Thailand, Laos and then

northwards to Hainan, China, Formosa, eastern Manchuria, Korea,

Japan and Sakhalin Isl. In America the distribution ranges from

the south-eastern parts of Canada, southwards and westwards

to the United States, Mexico, Cuba, Guatamala, Honduras,

Nicaragua, Costa Rica, Panama and the Columbian Andes (Map I).

Species of Quercus subgen. Quercus may be found up to c. 4300 m

altitude, and the boundary of the distribution lies as far north as

latitude 62° N. In Asia the centre of speciation of this subgenus

is in China, from where not less than 40 species have been describ-

ed. This subgenus is not represented in Malesia.

Soepadmo — Quercus subgen. Cyclobalanopsis 379

Subgenus Cyclobalanopsis is confined to the south-eastern and

eastern parts of Asia, viz. from Japan (Kanto Prov., Honshu) to

Korea, China, Formosa, Indo-China, Thailand, N. Burma, N.E.

India, and southwards to western Malesia (Malaya, Singapore,

Sumatra, Anambas Isl., Palawan Isl., Borneo and Java), with a

centre of speciation in Indo-China. In Malesia (Map II) the centre

of distribution is in Borneo from where 17 of the 19 known species

have been recorded. Species of this subgenus may be found up to

3500 m altitude, more commonly at about 1000 to 1500 m. The

northernmost boundary of the distribution lies at latitude c. 35° N.

(Central Japan) and the southernmost limit is in Java at c. latitude

10° S.

Quercus L. subgen. Cyclobalanopsis (Oersted) Schneider

Quercus subgen. Cyclobalanopsis (Oersted) Schneider, Ill.

Handb. Laubh. /: 210. 1906; Nakai in Bot. Mag. Tokyo 29: 56.

1915; Koidzumi in Bot. Mag. Tokyo 30: 199. 1916; A. Camus, Les

Chénes J; 158. 1938; Rehder, Man. Cult. Trees & Shrubs, 2nd.

ed.: 155. 1940. — Quercus sect. Cyclobalanus Endlicher, Gen. Pl.,

‘Suppl. 4, 2: 28. 1847, p.p.; A.DC. in Ann. Sc. nat. Bot., Sér. IV,

18: 55. 1862, p.p.; Prodr. 16, 2: 91. 1864, p.p.; Oudemans, Annot.

Cupul. Jav.: 22. 1865, p.p. — Quercus sect. Gyrolecana Blume,

Mus. Bot. Lugd. Bat. 7: 299. 1850, p.p.; Miquel, Fl. Ind. Bat. J, 1:

855. 1856. — Cyclobalanopsis Oersted in Vidensk. Medd. naturh.

For. Kjobn. 8: 77. 1867; in Kong. Dansk. Vidensk. Selsk. Skr.

5, 9: 69. 1871; Schottky in Bot. Jahrb. 47: 645. 1912; Hayata,

Icon. Pl. Form. 3: 185. 1913; Markgraf in Bot. Jahrb. 59: 59. 1924;

Kudo & Masamune in J. Soc. Trop Agr. 2: 148. 1930; in Trans.

Nat. Hist. Soc. Form. 20: 161. 1930; Schwarz in Notizbl. Bot. Gart.

Berlin-Dahlem /3; 5-8. 1936; in Fedde’s Repert. Sonderbeiheft D.:

31-35. 1936; Nakai in J. Jap. Botany /5: 201. 1939; Li in Bull.

Torrey Bot. Club 80: 323. 1953; Wood. Fl. Taiwan: 99. 1963. —

Quercus sect. Cyclobalanopsis (Oersted) Benth. & Hook. f., Gen.

Pl. 3: 408. 1880; Hooker f., Fl. Brit. Ind. 5; 603. 1888; King Ann.

Roy. Bot. Gard. Calc. 2: 27. 1889; Prantl in Engler & Prantl, Nat.

Pfl. fam. 3, 1: 55. 1894; Koorders & Valeton, Bijdr. Booms. Java

10: 15. 1904; Brandis, Ind. Trees: 627. 1906; Gamble in J. As.

Soc. Bengal 75: 408. 1915; Barnett in Trans. & Proc. Bot. Soc.

Edinburgh 34: 164. 1994. — Quercus subgen. Cyclotheca Nakai,

in Bot. Mag. Tokyo 29: 57. 1915.

Type species: Q. velutina Lindl. ex Wall., non Lamarck.

Trees of medium to large sizé. Buttresses absent or up to 1-3

m in height, spreading. Bark smooth and lenticellate or finely

fissured or scaly and peeling off profusely into rectangular pieces,

pale grey to greenish-brown in colour; inner bark sometimes up to

2.5 cm in thickness, brittle, fibrous, ridged on the inner side, pale

380 Gardens’ Bulletin, Singapore — XXII (1968 y

yellow to reddish-brown. Wood pale yellow to pale brown, growth

rings indistinct, heart-wood not well defined from the sapwood,.

hard but not durable, splitting very easily; vessel distribution

denritic rarely solitary; rays uni- to multi-seriate, homogeneous;

pitting of the vessel bordered; tannin content little to c. 20 per cent.

of dry weight. Innovations densely simple- or stellate-tomentose or

densely brownish, stiff pubescent. Branchlets glabrous, smooth or

shallowly fissured or markedly warty lenticellate. Buds ovoid-

globose or ovoid-conical, rarely ovoid-ellipsoid; scales tomentose

or densely brownish, woolly pubescent. Stipules extrapetiolar,

linear-acute, densely tomentose or woolly pubescent, caducous.

Petiole always thickened at its base, terete or flattened or sulcate

on the adaxial side, densely tomentose or woolly pubescent or

glabrous. Leaves spirally arranged or rarely pseudo-whorled, pen-

ninerved; midrib and lateral nerves flattened or impressed or

slightly raised above, more or less prominent beneath; margin

entire or remotely minutely serrate in the apical half; glabrous or

densely or sparsely simple or stellate-tomentose or densely or

sparsely stiff or woolly pubescent at least on the lower surface.

Male inflorescence solitary in the axil of a lower leaf or in paniculate

clusters on the lateral or subterminal new shoots, flexuous, pendent,

compound or simple, unbranched or much branched. Male flower

in 3-4-flowered dichasial clusters or solitary along the rachis.

Bracts ovate-linear, acute, densely tomentose, caducous. Perianth

(4)-6-lobed, the lobes connate at base, densely tomentose; stamens

(4)-6—(9), filaments slender, filiform, glabrous or tomentose at base;

anthers c. 0.5-1 mm long, basifixed, 2-locular, 4-lobed, longitu-

dinaliy dehiscent; pistillode normally absent, sometimes present

but reduced to a cluster of stiff simple hairs. Pollen grains prolate,

prolate-spheroidal or very rarely suboblate, 3-colpate, 3-colporate

or 3-colporoidate; polar diameter c. 22-33 », equatorial diameter

c. 15-30 uw; exine c. 1-1.5 p» thick, more or less scabrate; inner wall

(endexine) thinner or thicker or as thick as the outer wall

(ektexine); furrows elongate, tapering towards both ends; pores

circular or elliptic. Female inflorescence solitary in the axil of a

higher leaf, erect, densely woolly pubescent, few- to many-flowered.

Female flowers always solitary along the rachis, sessile; bracts

linear-acute, densely pubescent, caducous; perianth (4)-6—-(9)-lobed;

staminodes 0 or 5-7; styles 3-4-{6), cylindrical, glabrous or

pubescent at base, free and recurved or connate at base; stigmas

broadly capitate, glabrous; locules as many as styles; ovules 2 in

each locule. Cupule cup- or saucer-shaped, obconical or obovoid-

globose, lamellate, densely tomentose outside, densely silvery

pubescent inside; lamellae c. 5-11, denticulate and free at the rim

or more or less smooth and connate, thin or thick. Fruit ovoid-

conical, ovoid-globose or ovoid-cylindrical; apex rounded, atten-

uate-acute or abruptly depressed, umbonate; perianthodium (umbo)

ringed, well-developed; base rounded, truncate or concave; gla-

brous and shining or densely tomentose. Cotyledons flat-convex,

filling the entire space of the surviving locule; germination hypo-

geal; abortive ovules apical; radicle vertical.

Soepadmo — Quercus subgen. Cyclobalanopsis 381

KEY TO THE SPECIES, BASED ON FLOWERING OR

FRUITING SPECIMENS

1. Rim of the cupule thinner than 0.5 mm, lamellae with more or less

free margin; female inflorescence usually many-flowered.

2. Cupule deeply cup-shaped, covering c. 4} part of the fruit, c.

(1)-2-3-(3.5) cm deep and 2-4 cm in diameter.

3. Cupule c. 2—3.5 cm deep and 2.5—4 cm in diameter. Fruit c. 3-5 cm

long and 2-3 cm in diameter. Leaves pseudo-whorled, base

cordate to auriculate; petiole c. 0.5—-1 cm long .....................085

Be derits Mur ORE Feats CULE Leathe Peale chs obnce 1. @Q. pseudo-verticillata.

3. Cupule c. (1)—1.5—2-(2.5) cm deep and c. (1)—-1.5-2-(2.5) cm in

diameter. Fruit c. 2-3-(3.5) cm long and c. 1.5-2-(3) cm in

diameter. Leaves not pseudo-whorled, base not cordate nor auri-

culate; petiole c. 1.5-5 cm long.

4. Cupule tapering towards the base. Leaves densely stellate-

tomentellous, glaucous or silvery beneath; margin entire.

5. Fruit elongate conical or ovoid-globose. Styles free and

recurved. Staminodes O. Leaves silvery beneath; petiole not

deeply sulcate on the adaxial side ............ 2. QO. argentata.

5. Fruit obovoid-globose. Styles connate, not recurved. Stami-

nodes 5S—7. Leaves glaucous beneath; petiole deeply sulcate

Oe Rta Ie... ee acces evap iceen. 3. @Q. nivea.

4. Cupule rounded or truncate at base. Leaves glabrous or densely

brownish stiff-pubescent or sparsely stellate-tomentose, neither

glaucous nor silvery beneath; margin remotely minutely serrate

in the apical half.

6. Cupule c. 2-2.5 cm in depth and in diameter; lamellae 6-8.

Fruit ovoid-conical, c. 2-3 cm in length and c. 2 cm in

@iameter;> base? conver suds Anite. 4. Q. gaharuensis.

6. Cupule c. 3-3.5 cm in depth and in diameter; lamellae c.

9-11. Fruit ovoid-globose or ovoid-cylindrical, c. 2.5—3.5

cm in length and c. 2-3 cm in diameter; base flat or convex

Fe attra oe ee ee RAT rs ubisom satint ce.9 3 5. @Q. oidocarpa.

2. Cupule shallowly cup-shaped or saucer-shaped, covering c. 3-4 part

of the fruit, c. 0.5-1.2 cm in depth and c. (0.7)-1-1.8-(2.5) cm in

diameter.

7. Leaves elliptic-lanceolate or ovate-lanceolate, c. (3)—-10-—15-(24)

by (1)—3-5-(9) cm; petiole c. 1-3 cm long... -

8. Leaves elliptic-lanceolate, chartaceous, base attenuate-acute.

9. Cupule covering c. 1/6-1/5 part of the fruit, base of the

fruit concave. Petiole terete or flattened on the adaxial side

fo ECE ie (OMNES 2 ae NR a 6. Q. sumatrana.

9. Cupule covering c. 4-4 part of the fruit; base of the fruit

convex. Petiole sulcate on the adaxial side Re Tae IRE SBE

ania s MAEN ST atelier date Ne heed ab stesy ch dalpn? 00> Se, dake 7. @Q. subsericea.

8. Leaves ovate-elliptic or ovate-lanceolate, thick-coriaceous; base

rounded or cordate.

10. Cupule covering c. $+ part of the fruit; fruit ovoid-

cylindrical. Leaves c. (3)—5—-10-(14) by 2-5.5 cm, lower

surface not glaucous; lateral nerves c. 5-8 pairs ...............

10. Cupule covering c. 1/6-1/5 part of the ovoid—globose

fruit. Leaves c. (10}+15—18-(20) by 5.9 cm, glaucous beneath;

7. Leaves obovate or elliptic-obovate, c. 1.5-5.5 by 0.7-3.5 cm;

petiole c. 0.1-0.6 cm long.

11. Leaves glabrous’ or with sparse pubescence beneath,

upper surface shining, margin entire, apex rounded or

truncate-emarginate; lateral nerves c. 4-5 pairs .........

ribo cnag SERRE? deeds 3 epee deud. LES oh 10. Q. chrysotricha.

11. Leaves with dense pubescence on both surfaces, upper

surface dull, margin remotely serrulate in the apical

half, apex bluntly acute; lateral nerves c. 5-8 pairs

1g peed Shey beep epee SEA ent pA ee ee Yel eenen 11. Q. merrillii.

382 _ Gardens’ Bulletin, Singapore — XXII (1968)

1. Rim of the cupule c. 0.5-4 mm in thickness, lamellae more or less

connate at the rim; female inflorescence normally few-flowered.

12. Cupule c. 0.7-1.2 cm deep, c. 1-2 cm in diameter; lamellae c. 5-8.

Fruit c. 1-2-(3) by 1.2-1.5 cm. Leaves with entire margin.

13. Leaves elliptic-lanceolate, glabrous, glaucous beneath, base

attenuate-Aciileas ihe occt. pert fends doce marks » 12. @. kerangasensis.

13. Leaves ovate-elliptic densely yellowish-brown pubescent, base

rounded -or acute! ..24)s0ndol:...4.. 282.2: 13. Q. kinabaluensis.

12. Cupule c. 1.5—-2 cm deep and c. (1)-2-2.5 cm in diameter; lamellae

c. (5)-8-10-(12). Fruit c. 2-5.5 by 1-2 cm. Leaves remotely serrulate

in the apical half.

14. Old leaves glabrous. Fruit c. 2—5.5 cm long.

15. Leaves with strong and dense reticulation; lateral nerves

c. 10-15 pairs. Stamens 5-9; styles connate at base; stami-

Ie I cea eS ae 14. Q. valdinervosa.

15. Leaves with obscure and lax reticulation; lateral nerves

c. 7-10 pairs. Stamens 3-6; styles free and recurved; stami-

nodes, 6-Outic:tasredcnee weeps. 15. Q. gemelliflora.

14. Old leaves with dense pubescence beneath. Fruit c. 1-3 cm long.

16. Leaves with strong and dense reticulation.

17. Cupule deeply cup-shaped, obconical, c. 1-1.2 cm in

depth and c. 2-2.5 cm in diameter; lamellae 8-10.

Fruit cylindrical-conical, c. 2-3 by 1-2 cm ...............

dbs Fad « shee eaedars idatataban, an 9S Sa geet RE 16. Q. lineata.

17. Cupule flattened cup-shaped or saucer-shaped, c. 0.7-1

cm in depth and c. 1.2-2.5 cm in diameter; lamellae

7-8. Fruit globose or ovoid, c. 1-2 cm in length and

aiaMeper. . A tag . 41). dsus Podoods «oh - Bagleae 17. Q. steenisii.

16. Leaves with obscure and lax reticulation.

18. Cupule cup-shaped, base rounded, c. 1.5—-2 cm deep and

c. 1.5-2 cm in diameter; lamellae 10-12. Fruit with

convex base. Leaves linear-lanceolate .....................

WANE I dare! Mine per Mn de ee Pe ARM 18. Q. treubiana.

18. Cupule saucer-shaped, base flat, c. 0.5-0.7 cm deep and

c. 2-2.5 cm in diameter; lamellae 5-7. Fruit with flat

base. Leaves elliptic-oblong ............... 19. Q. elmeri.

KEY TO THE SPECIES, BASED ON STERILE SPECIMENS

1. Petiole c. (0.1)-0.3-0.5-(1) cm long. Leaves with rounded or bluntly

acute apex.

2. Leaves c. 6-17 by 3-6.5 cm, base rounded, subcordate or auriculate.

3. Leaves elliptic-oblong or oblanceolate-oblong, c. 7-17 by 3-6.5

cm, margin entire; lower surface sparsely minutely pubescent;

reticulation fine, dense, parallel ......... 1. Q. pseudo-verticillata.

3. Leaves ovate-elliptic or ovate-orbicular, c. 3-8 by 2-5 cm, margin

remotely serrulate in the apical half, lower surface with dense

layer of brownish woolly pubescence; reticulation distinct, pro-

minent on the lower surface ...............:..0ce008 17. Q. steenisii.

2. Leaves c. 1.5-5 by 0.7-3.5 cm, base attenuate-acute or attenuate-

rounded.

4. Leaves glabrous on both surfaces, margin entire, apex rounded

or truncate; lateral nerves c. 4—S5 pairs ......... 10. Q. chrysotricha.

4. Leaves with dense pubescence on both surfaces, at least on the

midrib and lateral nerves, margin remotely serrulate in the apical

half, apex bluntly acute, rarely rounded; lateral nerves c. 5-8

PANS |. ecccvecesscisctiesctldddddintiledgehitA ea. 11. Q. merrillii.

Soepadmo — Quercus subgen. Cyclobalanopsis 383

1. Petiole c. (1)-2-3-(4.5) cm long. Leaves with acute, acuminate or

caudate apex.

5. Leaves elliptic-lanceolate, lanceolate-oblong or ovate-elliptic, c. (5)-10-

17-(24) by (2)-3-5-(9) cm.

6. Leaf margin entire, recurved, reticulation obscure on both sur-

faces.

7. Lower surface of the leaf glaucous. Branchlets sparsely stellate-

tomentose,. shallowly fissured ........................ 3. Q. nivea.

7. Lower surface of the leaf silvery. Branchlets glabrous densely

and prominently lenticellate ..................... 2. @Q. argentata.

6. Leaves remotely serrulate in the apical half, reticulation distinct,

at least on the lower surface.

8. Old leaf glabrous.

9. Leaves elliptic-oblong, lateral nerves c. 10-15 pairs; reticula-

tion ‘strong and dense =:,...06.5...0...... 14. Q. valdinervosa.

9. Leaves elliptic-lanceolate, lateral nerves c. 7-10 pairs;

8. Old leaves not glabrous.

10. Leaves ovate-elliptic, elliptic-oblong or oblong-lanceolate,

c. (5)-10—20-(22) by (2.5)-S-7-(9) cm.

11. Leaves ovate-elliptic, lower surface glaucous ............

Be hes ie | Den tes : tL BW GAT Oe ee ER ae 9. @Q. percoriacea.

11. Leaves elliptic-oblong or oblong-lanceolate, lower sur-

face not glaucous.

12. Leaves densely brownish, stiff-pubescent beneath;

petiole and midrib not sulcate on the adaxial side

eben) et eet eee CP eee, re 4. Q. gaharuensis.

12. Leaves sparsely stellate-tomentose; petiole and

midrib deeply sulcate on the adaxial side ............

MOOR ie See 5. @Q. oidocarpa.

10. Leaves elliptic-lanceolate, c. (1.5)—-5—13-(24) by 2.5-5 cm.

13. Leaves glaucous beneath, reticulation fine, obscure ...

Noa Oho dais « DENG Roe ese zias eels dis 5 6. Q. sumatrana.

13. Leave not glaucous beneath; reticulation strong, dis-

tinct on the lower surface.

14. Leaves with dense, simple, adpressed pubescence

beneath; lateral nerves c. 10-20 pairs, dense;

reticulation distinct beneath ......... 16. Q. lineata.

14. Leaves with dense woolly, rufous-pubescence

beneath; lateral nerves c. 5S—12 pairs, lax; reticula-

tion, obseute dnd fax ..:1f.0.016:.2..48 19. Q. elmeri.

5. Leaves linear-lanceolate or ovate-elliptic, c. 5-10 by 2-4 cm.

15. Leaves linear-lanceolate, with dense pubescence beneath.

5 MOS AS DAE EEE ae eee eee 18. Q. treubiana.

15. Leaves ovate-elliptic, with dense or sparse stellate-tomentum

| or completely glabrous beneath.

| 16. Leaves with attenuate-acute or attenuate-rounded base,

glabrous or sparsely stellate-tomentose beneath.

17. Leaves glabrous, glaucous beneath; petiole c. 0.8-1.2

cm long; branchlets with dense warty lenticels ......

] Q. kerangasensis.

17. Leaves sparsely stellate-tomentose beneath, not glau-

cous; petiole c. 1-2.5 cm long; branchlets sparsely

MMIMUTe-lETMICCIIAIC .........2....06000+. 7. @Q. subsericea.

16. Leaves with rounded or cordate base, Jower surface with

dense stellate-tomentum or simple-pubescence.

18. Leaf-margin remotely serrulate in the apical half;

| lower surface with dense stellate-tomentum or glab-

rous; reticulation more or less anastomosing .........

IR th Se ke a OR SS 8. QO. lowii.

18. Leaf-margin entire; lower surface with dense yellowish

simple-pubescence; reticulation parallel ..................

feb ieh eves: eérocg tO. ATER Shades, 13. Q. kinabaluensis.

384 Gardens’ Bulletin, Singapore — XXII (1968)

1. Quereus pseudo-verticillata Soepadmo, Pl. 1, Map III.

Q. pseudo-verticillata Soepadmo in Gard. Bull. Singapore 21,

3: 380, f. 1. 1966. — Type: Chew & Corner RSNB. 4434, Bem-

bangan River, Mt. Kinabalu, Borneo, alt. c. 1650 m, Febr. fr.

(Holotype: K; isotype L).

Tree c. 30 m tall, trunk c. 90 cm in diameter. Buttresses up to

2 m tall. Bark scaly, peeling off profusely into small, rectangular

pieces. Branchlets glabrous, lenticellate. Leaves pseudo-whorled,

thick-coriaceous, elliptic-lanceolate or oblanceolate-oblong, c. 7-17

by 3-6.5 cm; margin entire, base cordate to auriculate, apex

rounded or bluntly acute; upper surface glabrous, lower surface

with sparse minute simple-pubescence. Midrib and lateral nerves

strongly prominent beneath, flattened to slightly impressed above,

especially in the apical half. Lateral nerves c. 8-15 pairs, forming

c. 30°-45° angle with the midrib, parallel, arcuating; reticulation

obscure, parallel, dense. Petiole c. 0.5—1 cm long and 0.2 cm thick,

tomentose, glabrescent, flattened on the adaxial side. Buds ovoid-

globose, c. 0.2-0.3 cm in length and c. 0.2 cm in diameter. Inflore-

scences unknown. Cupule cup-shaped, obconical-globose, base

pointed, covering c. 44 part of the fruit; c. 2-3.5 cm deep and

2.5-4 cm in diameter; outside brownish-sericeous, glabrescent;

inside with dense brownish, stiff-pubescence; lamellae c. 10-12,

thin, free at the rims, dentate, especially the lower ones. Fruit

cylindrical-globose, c. 3-5 cm in length and 2-3 cm in diameter,

sericeous, apex rounded or depressed, umbo conical, ringed, base

convex or flat.

Ecology: in montane rain forest at c. 1650 m alt. Fruiting in

October-February. ;

Distribution: so far only known from Mt. Kinabalu, in North

Borneo.

2. Quercus argentata Korth., Fig. 6, Map IV.

Q. argentaia Korthals in Verh. Nat. Gesch. Bot.: 215, t. 47, f.

1-17, 1844; Blume, Mus. Bot. Lugd. Bat. J: 299. 1850; Miquel, FI.

Ind. Bat. J, 1: 858. 1856; in Ann. Mus. Bot. Lugd. Bat. J: 115.

1863; A. DeCandolle, Prodr. 1/6, 2: 91. 1864; Scheffer in Nat.

Tijdschr. Ned. Ind. 32: 417. 1872; Wenzig in Jahrb. K6n. bot. Gart.

& Mus. Berlin 4: 230. 1886; King in Ann. Roy. Bot. Gard. Calc.

2: 30, t. 24 A. 1889; von Seemen in Bull. Dép. Agr. Ind. Néerl.

I: 3. 1906; Gamble in J. As. Soc. Bengal 75: 408. 1915, p.p.

Merrill in J. Str. Br. Roy. As. Soc., Sp. No.: 211. 1921; Ridley, FI.

Mal. Pen. 3: 373. 1924, p.p.; Moore in J. Bot. 63, Suppl.: 114.

1925; A. Camis, Les Chénes: 7 >-3'i 91938. tad; t 23

1934. — Cyclobalanopsis argentata (Korth.) Oersted in Vidensk.

Medd. naturh. For. Kjobn. 8: 79, t. I-II, f. 5-8. 1867; in Liebe-

man’s Chénes Amér. Trop.: 20. 4&89. — Lithocarpus argentata

(Korth.) Merrill in Contr. Arn. Arb. 8: 42. 1934. — Type:

Korthals s.n., in forest, Malintang, Sumatra, fruiting (Holotype: L).

Q. pinanga Blume, Mus. Bot. Lugd. Bat. J: 303. 1850. — Type:

Blume s.n., Java, st., (L); syn. nov.

6 Be

v lay ye

= Qa oO

a gS &

oo 0

ee)

<< Si

= a,

a a

~~ —

Q |

: - iia

, ee i ee

3 Ea

om 5

a SS

Ss 0 go

aor

EO __

North

Kinabalu

Mt.

Borneo.

8 i oa

paRes La .

Po a nel ge

Soepadmo — Quercus subgen. Cyclobalanopsis 385

Bs sgh.

Fig. 6: Quercus argentata: 1, 2, 4, 5 after Smythies S. 12165; 3 after

Korthals s.n. (Type); 6-7 after Jacobs 5114.

386 Gardens’ Bulletin, Singapore — XXII (1968)

Q. wilhelminae von Seemen, 1.c.: 9-10, p.p., excl. fructu; Merrill

in J. Str. Br. Roy. As. Soc., Sp. No.: 216. 1921. — Type: Hallier

2628, Amai Ambit, Sarawak, Borneo (Holotype: L); syn. nov.

Tree c. 7-37 m tall, trunk c. 20-100 cm in diameter. Buttresses

up to 1.2 m tall. Bark smooth, lenticellate, pale grey, sometimes

with horizontal cracks; inner bark c. 2 cm thick, pale brown,

brittle. Branchlets glabrous, densely warty lenticellate, greyish-

brown. Buds ovoid-globose, c. 2-3 by 2-2.5 mm, stellate- or

simple-tomentose, glabrescent. Stipules linear-acute, with dense

simple, stiff-pubescence, c. 3-5 by 1-1.5 mm, caducous. Petiole

slender, glabrous flattened on the adaxial side, c. 1.5-3 cm long

and c. 1-1.5 mm thick. Leaves coriaceous, elliptic- or lanceolate-

oblong, c. 8-22 by 3-7 cm; lower surface with dense silvery

stellate-tomentum, upper surface shining, glabrous; base attenuate-

rounded or attenuate-acute, slightly asymmetrical, margin entire

and slightly undulate, apex c. 0.5-1 cm acuminate. Midrib and

lateral nerves strongly prominent beneath, impressed above, gla-

brous; lateral nerves c. 10-17 pairs, forming c. 60°-75° with the

midrib, parallel, arcuating near the margin; reticulation fine,

obscure. Male inflorescence in a paniculate cluster of 3 or 4 on

a lateral new shoot, c. 5-10 cm long. Bract ovate-acute, with

dense simple-tomentum, c. 1-1.5 by 0.5 cm, caducous. Male

flowers in 3-flowered dichasial clusters or solitary along the rachis;

perianth 4-6-lobed, the lobes connate at base; stamens normally

6, filament hairy at base, pistillode absent. Female inflorescence

many-flowered, slender, c. 2-3 cm long, with dense simple-

tomentum; perianth 4-6-lobed, densely tomentose outside; stami-

nodes absent; styles 3-4, hairy at base, free and slightly recurved.

Bracts linear-acute, c. 0.5-1 mm long. Immature cupule turbinate

or ovoid-conical, flattened or rounded at the top, attenuate at

base; lamellae c. 8-10, thin, dentate, with dense brownish tomen-

tum, completely covering the fruit except for the umbo. Mature

cupule cup-shaped, obconical or obovoid, c. 1.5—2 cm deep and

c. 1-1.5 cm in diameter, with dense simple-pubescence outside;

lamellae c. 8-10, free and denticulate at the rim. Fruit elongated

conical or ovoid-globose, attenuating towards the acute apex and

rounded base, c. 3—-3.5 cm long and 1.5 cm in diameter.

Ecology: in lowland to montane forests at c. 0-2700 m alt.

Flowering between July and September; fruiting between

October and May.

Distribution: Malaya, Singapore, Sumatra, Bangka, Anambas

Isl., Borneo, and Java.

Notes: The majority of the examined specimens come from

Borneo and Sumatra.

Soepadmo — Quercus subgen. Cyclobalanopsis 387

3. Quercus nivea King, Fig. 7, Map III.

Quercus nivea King in Ann. Roy. Bot. Gard. Calc. 2: 31, Pl.

24B, 1889; Koidzumi in Bot. Mag. Tokyo 30: 201. 1916; Merrill

in J. Str. Br. Roy. As. Soc., Spec. No.: 214. 1921; A. Camus, Les

Chénes J: 313. 1938; Atl. J: t. 23, f. 10-16. 1934. — Lecto-type:

Beccari P.B. 2551, fr., fl., Sarawak (FI).

Tree c 25 m tall, trunk c. 40 cm in diameter. Buttresses c. 2 m

tall, 10-15 cm thick. Bark pale grey, rough; inner bark chocolate-

brown, fibrous. Branchlets with dense brownish, stellate-tomentum,

glabrescent; lenticels sparse, splitting longitudinally into shallow

furrows. Stipules linear-acute, with stellate-tomentum outside, c.

2-3 by 0.5-1 mm, caducous. Petiole densely stellate-tomentose

when young, soon becomes glabrous, c. 2-4.5 cm long, deeply

sulcate on the adaxial side, especially near the leaf-base. Leaves

thick-coriaceous, elliptic-oblong or lanceolate-oblong, c. 6-15 by

2.5—5.5 cm; upper surface shining, glabrous, lower surface glaucous,

with dense stellate-tomentum; base rounded or abruptly acute,

sometimes asymmetrical, margin entire, slightly undulate near the

acuminate or caudate apex, acument c. 0.5—1 cm long. Midrib and

lateral nerves prominent beneath, impressed or flattened above.

Reticulation obscure on both surfaces. Lateral nerves c. 6-12

pairs, parallel, arcuating near the margin, forming c. 60°-70° angle

with the midrib. Male inflorescence in a paniculate cluster on the

lateral new shoot, c. 5 cm long, densely stellate-tomentose. Male

flowers in 3-flowered dichasial clusters or solitary along the rachis;

perianth 4-6-lobed, rather thick-coriaceous; stamens 4-6; filaments

slender, hairy at base; pistillode absent. Female inflorescence

solitary in the axil of a higher leaf; bracts ovate-acute, caducous.

Female flowers: perianth 5—6-lobed, with dense stellate tomentum;

staminodes developed considerably but remain not functional;

style 3-4, short, connate at base. Young cupule obconical-turbinate,.

base attenuate, lamellae thin, with dense stellate-tomentum. Mature

cupule cup-shaped, obconical, tapering towards the base, c. 1-1.7

cm deep and 1.5-2 cm in diameter; lamellae c. 7. Fruit obovoid-

globose, c. 2 cm long and in diameter, apex depressed, base convex.

Ecology: in high heath forest, at c. 1000 m alt. Flowering in

January.

Distribution: endemic in Sarawak (Mts. Gaharu and Pueh).

Notes: This species is closely allied to Q. argentata, but differs

from the latter by its thick-coriaceous leaves which are glaucous

and densely stellate-tomentose beneath, fewer lateral nerves, deeply

sulcate petiole, and by its shallowly furrowed, not warty lenticellate

branchlets. The lamellae in Q. nivea are more or less connate,

whilst in Q. argentata, they are free and denticulate.

Gardens’ Bulletin, Singapore — XXII (1968)

388

——— ee

5 CI

5.1964,

Fig. 7: Quercus nivea

1, 3 after Beccari P.B. 2551 (Type); 2 after King, l.c. 1889.

Soepadmo — Quercus subgen. Cyclobalanopsis 389

4. Quercus gaharuensis Soepadmo, Map V.

Quercus gaharuensis Soepadmo in Gard. Bull. Singapore 2/, 3:

384, f. 3. 1966. — Type: Anderson et alia S. 15534, Serian, lower

slope of Mt. Gaharu, Sarawak, alt. c. 100 m, October fr. (Holo-

type: K*; isotypes: BO, L, SAN, SAR).

Quercus oidocarpa (non Korth.) Hooker f., Fl. Brit. Ind. 5: 603.

1888; King in Ann. Roy. Bot. Gard. Cal. 2: 28, pl. 21A,

1889, p.p., quoad specim. ex Mal. Pen.; Brandis, Ind. Trees, 3rd.

ed.: 629. 1906, p.p., quoad specim. ex Mal. Pen.; Gamble in J.

As. Soc. Bengal 75: 409. 1915, p.p., quoad specim. ex Mal. Pen.;

Merrill in J. Str. Br. Roy. As. Soc., Spec. No.: 214. 1921, p.p.,

excl.. Beccari 2919; in Contr. Arm. Arb. 8: 42. 1934. p.p.; A.

Camus, Les Chénes J: 213. 1938, p.p., excl. Korthals s.n.; Atl. 1,

t. 7, f. 10-15. 1934.

Tree c. 10-30 m tall, trunk c. 30-100 cm in diameter. Buttresses

short, spreading, up to 0.7 m tall. Bark smooth, mottled hooped,

lenticels scattered in longitudinal rows. /nnovation with dense

stellate-tomentum. Branchlets glabrous, lenticellate. Buds ovoid-

globose, c. 2 by 2 mm. Stipules linear-acute, c. 5 by 1 mm,

caducous. Petiole slender, terete or flattened on the adaxial side,

_ with dense stellate-tomentum, glabrescent, c. 1-3.5 cm long. Leaves

elliptic-lanceolate or elliptic-oblong, c. 5-23 by 2-9 cm, thin

coriaceous, base attenuate-acute or rounded, slightly asymmetrical,

margin entire or remotely serrulate in the apical part, apex rounded

or bluntly acute; upper surface glabrous, lower surface with dense

stellate-pubescence, glabrescent. Midrib and lateral nerves pro-

minent beneath, slightly so above; reticulation distinctly visible

beneath; lateral nerves c. 8-15 pairs, parallel, arcuating towards

the margin, forming c. 45°-60° angle with the midrib. Male

inflorescence c. 1.5-3.5 cm long; bracts ovate-acute, c. 1-1.5 by

1 mm, caducous. Male flowers in 3-flowered dichasial clusters;

perianth 4—6-lobed; stamens 4—6; filament hairy at base, c. 1-2 mm

long; anthers c. 0.5 by 0.5 mm; pistillode reduced to a cluster of

stiff hairs. Female inflorescence c. 1—-1.5 cm long, carrying c. 2-5

solitary, sessile flowers; bracts ovate-linear, c. 1-2 by 0.5-1 mm.

Female flower: perianth 5—6-lobed; staminodes absent; styles 3-4,

slender, cylindrical, hairy at base, c. 1-2 mm long. Young cupule

ovoid-globose, c. 1.5—2 cm in length and in diameter, covering the

entire fruit; apex rounded or truncate, base attenuate-rounded,

densely sericeous outside and densely brownish-stiff-pubescent

inside. Mature cupule deeply cup-shaped, c. 2—2.5 cm in depth

and in diameter; base rounded, enclosing c. 4-4 part of the fruit;

lamellae c. 6-8, rim thin, slightly recurved and denticulate. Young

fruit ovoid or depressed conical, c. 1-1.5 cm in length and in

diameter, densely sericeous, glabrescent; apex rounded or abruptly

truncate. Ripe fruit ovoid-conical, c. 2-3 cm long and 2 cm in

diameter, apex acute or rounded, base convex.

Ecology: in lowland mixed Dipterocarps to submontane forests,

at c. 100-1400 m alt. Fruiting in September-February.

Distribution: Malaya (Penang, Perak), Borneo (Sarawak),

Sumatra (Karolands, Painan, Asahan Forest Reserve).

Recently Mr. Forman informed me that the holotype of this species was

redeposited at Kew.

| ee aa i

390 Gardens’ Builetin, Singapore — XXII (1968)

5. Quercus oidocarpa Korth., Fig. 8, Map VI.

Quercus oidocarpa Korthals in Verh. Nat. Gesch. Bot.: 216,

t. 47, f. 18, 1844; Blume, Mus. Bot. Lugd. Bat. /: 302. 1850;

Miquel, Fl. Ind. Bat. 7, 1: 856, 1856; in Ann. Mus. Bot. Lugd.

Bat. 7: 115. 1863; A. DeCandolle. Prodr. /6, 2: 99. 1864; Wenzig

in Jahrb. K6n. bot. Gart. & Mus. Berlin 4: 233. 1886; King in

Ann. Roy. Bot. Gard. Calc. 2: 28. 1889, p.p., quoad Korthals s.n.;

A. Camus, Les Chénes J: 213. 1938, p.p., quoad Korthals sn. —

Cyclobalanopsis oidocarpa (Korth.) Oersted in Vidensk. Medd.

naturh. For. Kjobn. 8: 78. 1867; Schottky in Bot. Jahrb. 47: 151.

1912. — Type: Korthals s.n., in forest, Malintang, Sumatra, fr.

(Holotype: L).

Quercus brevistyla A. Camus in Bull. Soc. Bot. France 80:

353. 1933; Lzes Chénes 7: 276. 1938; Atl. t. 17. 1934. — Type:

Syed Alie Kep. 22574, Fraser’s Hill, Pahang, Malaya, Febr. fr.

(Holotype: KEP; isotypes: P, SING); syn. nov.

Quercus semiserrata (non Roxb.) Koorders & Valeton, Bijdr.

Booms. Java /0: 25. 1904, p.p., quoad Koorders 1488 & 1533;

Koorders, Exk. Fl. Java 2: 16. 1912. p.p.

Tree c. 25-30 m tall, trunk c. 50 cm in diameter. Bark grey,

scaly and peeling off profusely into-rectangular pieces. Branchlets

with dense stiff-pubescence, glabrescent, lenticellate. Buds ovoid-

globose, c. 3-5 by 2-3 mm. Petiole c. 1.5—3.5 cm long, shallowly

furrowed on the adaxial side, especially near the leaf-base,

glabrous. Leaves oblong-lanceolate or elliptic-oblong, c. 7-17 by

3-7 cm, bas2 acute or rounded or subcordate, margin remotely

serulate in the apical part, apex acute or 0.5—1.5 cm acuminate:

upper surface glabrous, lower surface with spare stellate-

tomentum. Midrib and lateral nerves prominent beneath,

impressed above; lateral nerves c. 9-13 pairs, parallel, forming

ce. 45° angle with the midrib; reticulation distinct beneath,

parallel, dense. Male inflorescence on a new shooi in the axil of a

higher leaf, c. 5-7 cm long, with dense stiff-pubescence; bracts

Ovate-acute, c. 1.5-2 by 1I-1.5 mm, glabrous, caducous. Male

flowers in 3-4-flowered dichasial clusters or solitary along the

rachis; perianth membranaceous, glabrous, 4-6-lobed; stamens

4-7; filaments slender, glabrous, c. 2 mm long; anthers c. 0.7 mm

long: pistillode absent. Female inflorescence c. 3-7 cm _ long.

subglabrous, in the axil of a higher leaf, carrying c. 3-7 flowers;

bracts ovate-acute. Female flowers: perianth 4—6-lobed; staminodes

absent; styles 34, free, recurved, c. 2 mm long, hairy at base.

Cupule dzeply cup-shaped, c. 3—3.5 cm in depth and c. 3—3.5 cm in

diameter; base attenuate-rounded; lamellae thin, c. 9-11. the lower

ones denticulate, the other entire, yellowish-brown tomentose. Fruit

ovoid-globose or ovoid-cylindrical, c. 2.5-3.5 cm in length and c.

2-3 cm in diameter, apex rounded or abruptly depressed, base

rounded or flat.

Ecology: in lowland to submontane forests, at c. 700-1500 m

alt. Flowering between March and April; fruiting between

Apri'—February.

Distribution: Malaya, Sumatra, and Java.

Soepadmo — Quercus subgen. Cyclobalanopsis 391

Fig. 8: Quercus oidocarpa

1-4 after Symington Kep. 31071; 5 after Poore 1315, 6 after Syed

Ali Kep. 22574; 7-8 after Md. Nur s.n.; 9 after Poore 1348; 10 after

Poore 1315. :

— oO ee ee

392 Gardens’ Bulletin, Singapore — XXII (1968)

6. Quercus sumatrana Soepadmo, Map VII.

Quercus sumatrana Soepadmo in Gard. Bull. Singapore 2/, 3:

387, f. 4, 1966. — Type: Achmad 1501, Tapah, N. Sumatra, Nov.

‘fr. (Holotype: BO; isotype: L).

Tree c. 35 m tall, trunk up to c. 130 cm in diameter. Buttresses

-c. 1-2.5 m tall. Bark rough, grey. Young branchlets smooth, with

-dense brownish, stiff-pubescence; older branchlets glabrous, lenti-

cellate. Buds ovoid-globose, with dense pubescence, glabrescent,

¢. 2-3 by 1.5-2 mm. Petiole slender, c. 1-2.5 cm long, terete or

‘flattened on the adaxial side, with sparse pubescence. Leaves thin-

chartaceous, elliptic-lanceolate, c. 8-24 by 2-8 cm, base attenuate-

acute, margin entire-undulate or remotely serrulate in the upper

part; apex sharply 0.5-1.5 cm acuminate; lower surface with

sparse stiff-simple-pubescence, glaucous, upper surface glabrous.

-Midrib and lateral nerves prominent beneath, slightly so above;

lateral nerves c. 8-13 pairs, rarely opposite, forming c. 45°-60°

-angle with the midrib, parallel, arcuating near the margin; reticu-

lation parallel, obscure on both surfaces. Inflorescences not known.

“Young cupule saucer-shaped, covering the whole friut except for

the protruding umbo which is topped by 3, recurved, c. 2 mm

long styles; lamellae 3-5, thin, tomentose, rim recurved and

denticulate. Mature cupule shallowly cup-shaped, c. 0.7-1 cm in

depth and c. 2—2.5 cm in diameter, covering c. 1/6—-1/5 part of the

‘fruit, base attenuate-rounded, with dense tomentum on both sur-

faces; lamellae free at the rim. Mature fruit ovoid-conical, c. 1.8-2

cm in length and c. 1.5—2 cm in diameter, densely sericeous, apex

acute, base concave.

_ Ecology: in lowland to submontane forests, at c. 10-1300 m

alt. Fruiting between April and December. |

Distribution: Sumatra, Borneo.

.7. Quercus subsericea A. Camus Fig. 9, Map VIII.

. Quercus subsericea A. Camus in Bull. Soc. Bot. France 80: 354.

(1933; Les Chénes /: 366. 1938; Atl. J, t. 33, f. 21-26. 1934; Atl.

3, t. 347, f. 1-7. 1939. — Quercus sericea Scheffer in Nat. Tijdschr.

Ned. Ind. 37: 361. 1870; 1.c. 32: 416. 1873; King in Ann. Roy. Bot.

Gard. Calc. 2: 63, pl. 57 B. 1889; Merrill in J. Str. Br. Roy. As.

Scc., Spec. No.: 215. 1921; non Willdenow (1805). — Cycloba-

-lanopsis sericea (Scheff.) Schottky in Bot. Jahrb. 47: 656. 1912. —

-Synaedrys sericea (Scheff.) Koidzumi in Bot. Mag. Tokyo 30:

192. 1916. — Type: Teijsmann HB. 21080, Bangka, fr. (Holotype:

BO; isotype: K).

Quercus lineata (non Bl.) S. Moore in J. Bot. 63, Suppl.: 114.

1925, p.p., quoad specim. ex Forbes 572.

Quercus oidocarpa (non Korth.) Merrill, 1.c.: 214. 1921, p.p.,

‘qucad specim. ex Beccari 2919.

Tree c. 6-15 cm tall, trunk c. 10-40 cm in diameter. Bark finely

fissured or scaly, thin, greyish-brown; inner bark fibrous, brownish,

‘c. 2.5 cm thick. Innovations with dense stiff-simple-pubescence or

adpressed stellate-tomentum, glabrescent. Older branchlets sparsely

Soepadmo — Quercus subgen. Cyclobalanopsis 393

Fig 9: Quercus subsericea

1 after Teijsmann H.B. 7638; 2 after Symington Kep. 37644; 3 after-

SAN. 25119; 4 after SAN. 28276.

ees ees

394 Gardens’ Bulletin, Singapore — XXII (1968 }

minute-lenticellate. Stipules linear-acute, with dense stiff-pubes-

cence, c. 0.5—0.7 by 0.1-0.2 cm, caducous. Petiole slender, c. 1-2.5

cm long, furrowed on the adaxial side. Leaves thin-coriaceous,

elliptic-lanceolate, rarely ovate, c. 5-16 by 1—5 cm, base acute or

roundish, margin entire or remotely serrulate near the sharply

acute or 0.5-1.5 cm acuminate apex; lower surface with sparse

stellate-tomentum, upper surface shining, glabrous. Midrib and

lateral nerves prominent beneath, impressed above; lateral nerves

c. 6-12 pairs, forming c. 45°-60° angle with the midrib, parallel,

arcuating towards the margin; reticulation obscure, especially on

the upper surface. Male inflorescence c. 2-5 cm long, solitary in

the axil of a lower leaf or in paniculate clusters on the lateral new

shoots, with dense stellate-tomentum. Bracts ovate-acute, tomen-

tose outside, c. 1-2 by 1 mm, caducous. Male flowers solitary along

the rachis or in 3-flowered dichasial clusters; perianth tomentose,

4-6-lobed; stamens 4-6; filaments slender, c. 1-2 mm _ long;

anthers c. 1 by 1 mm; pistillode absent. Female inflorescence

solitary in the axil of a higher leaf, c. 0.5-1.5 cm long, with dense

tomentum, carrying 2-5 flowers; bracts ovate-acute, c. 0.5-0.7 by

0.3-0.5 mm. Female flowers: ovary ovoid-cylindrical, c. 0.5 mm

long, rounded-triangular; perianth 5—6-lobed; staminodes 0; styles

3-4, c. | mm long, tomentose at base. Young cupule ovoid-

globose or obconical, lamellae 4-5, the 2-3 lower ones denticulate,

the other entire. Mature cupule cup-shaped, obconical, covering

c. +4 part of the fruit, c. 0.5-1.2 cm in depth and c. 0.7—-1 cm in

diameter, base rounded or attenuate, lamellae with dense tomentum

outside. Fruit ovoid-conical or ovoid-globose, c. 1-2.5 cm in

length and c. 1-1.5 cm in diameter; densely sericeous, base obtuse

or convex, apex acute or abruptly depressed, umbonate.

Ecology: in lowland Dipterocarps to submontane forests, at

c. 500-1500 m alt. Fruiting in June-March.

Distribution: Malaya, Sumatra, Bangka, Borneo, Java.

8. Quercus lowii King, Fig. 10, Map IX.

Quercus lowii King in Ann. Roy. Bot. Gard. Calc. 2: 28, pl. 21B,

1889: Koidzumi in Bot. Mag. Tokyo 30: 201. 1916; Merrill in J.

Str. Br. Roy. As. Soc., Spec. No.: 214. 1921; A. Camus, Les

Chénes /: 366. 1938; Atl. J, t. 33, f. 27-19. 1934. — Cyclobala-

nopsis lowii (King) Schottky in Bot. Jahrb. 47: 653. 1912. — Type:

Low s.n., fr., Mt. Kinabalu, Borneo (CALC).

Tree c. 10-20 m tall, trunk c. 30-40 cm in diameter. Branchlets

dark-grey, shallowly fissured, with dense brownish, stellate-tomen-

tum; older branchlets lenticellate, glabrous. Buds ovoid- globose,

densely tomentose. Stipules linear-acute, c. 2-4 by 0.3-0.5 cm, with

dense tomentum, caducous. Petiole c. 1-2.5 cm long, sulcate on the

adaxial side, with dense simple- or stellate-tomentum. Leaves thick-

coriaceous, ovate-elliptic or ovate-lanceolate, c. (3)-5—10-(14) by

2-5.5 cm, base rounded or cordate, slightly asymmetrical, margin

entire or remotely serrulate in the apical half, apex sharply acute

or 0.5—2 cm acuminate; lower surface with dense, brownish stellate-

tomentum or almost glabrous, upper surface glabrous and shining

Soepadmo — Quercus subgen. Cyclobalanopsis 395

£3 1968 .

—————

Fig. 10: Quercus lowii

1-2 after SAN. 28808; 3 after Clemens 40533; 4-7 after Chew &

Corner RSNB. 4500. 4

— es

396 Gardens’ Bulletin, Singapore — XXII (1968)

or with sparse stellate-tomentum, especially on the midrib. Midrib

and lateral nerves impressed above, prominent or flattened beneath;

teticulation fine, more or less anastomosing, distinct beneath.

Lateral nerves c. 5-8 pairs, forming c. 50°-60° angle with the

midrib, parallel, arcuating towards the margin. Male inflorescence

‘slender, c. 5-10 cm long, with dense stellate-tomentum, unbranched

or rarely branched, solitary in the axil of a lower leaf; bracts

ovate-acute, c. 1-2 by 0.5 mm, with dense stellate-tomentum

outside, glabrescent, caducous. Male flowers solitary or in 3-flower-

ed dichasial clusters along the rachis; perianth 4—-6-lobed, with

simple or stellate-tomentum, glabrescent; stamens 4-6; anthers c.

1 by 0.5-1 mm; filaments c. 1 mm long; pistillode reduced to a

cluster of woolly simple hairs. Female inflorescence c. 1-2 cm

long, with 2-5 solitary female flowers; solitary in the axil of a

‘higher leaf; bracts ovate-acute, with dense simple or stellate-

tomentum, c. 1 mm. Female flowers: perianth 4-7-lobed, with

-dense tomentum; staminode 0; styles 3-6, c. 1 mm long, recurved.

Young cupule obconical, c. 0.8-1 cm deep and 0.7-1.2 cm in

-diameter; lamellae thin, c. 5-7, the lower ones denticulate, the

Others entire, with dense stellate-tomentum. Mature cupule cup-

‘shaped, base rounded, c. 1.3-1.8 cm in diameter and 0.8-1 cm in

depth, covering c. 4-1/3 part of the fruit. Fruit ovoid-cylindrical,

apex rounded or acute, c. 1.5—-2 cm in length and 1.3—-1.5 cm in

diameter, with dense stellate-tomentum, glabrescent; base convex.

Ecology: in lowland to montane forests, at c. 0-2500 m alt.,

more commonly between c. 1000-1500 m alt. Fruiting between

July and March.

Distribution: endemic in Borneo; common in Mt. Kinabalu.

‘9. Quercus percoriacea Soepadmo, Map VI.

Quercus percoriacea Soepadmo in Gard. Bull. Singapore 2/, 3:

382, f. 2. 1966. — Type: Anderson S. 20226, Sarawak, Bario,

‘Ulu Baram, on path to Pa’ Ukad, at alt. c. 1100 m, July fr.

(Holotype: K; isotypes: L, SAN, SAR, SING).

Tree with trunk c. 50 cm in diameter. Buttresses up to 1 m tall.

‘Bark smooth, grey, hoop-marked. Branchlets grey, glabrous, lenti-

‘cellate. Buds ovoid-globose, c. 0.3 by 0.2 cm, with dense tomentum.

Peiiole c. 2-3.5 cm long, glabrous, terete or slightly grooved on

‘the adaxial side. Leaves thick-coriaceous, ovate-elliptic, c. (10)-

15-18-(20) by 5-9 cm, base rounded, margin incurved, remotely

‘serrulate in the apical half; apex acute or 1-1.5 cm acuminate,

‘lower surface with sparse stellate-tomentum, glaucous, upper sur-

face shining, glabrous. Midrib and lateral nerves prominent

beneath, impressed and obscure above. Lateral nerves c. 8-10 pairs,

forming c. 45°-60° angle with the midrib, parallel, arcuating

towards the margin; reticulation parallel, prominent beneath. Male

cand female inflorescence not known. Young infructescence c.

2-3 cm long, sericeous, glabrescent, lenticellate, with 1-4 solitary

young fruits. Young cupule obovoid, sericeous, c. 0-7-1 cm deep,

Soepadmo — Quercus subgen. Cyclobalanopsis 397

0.8-1 cm in diameter; base attenuate; lamellae c. 4-8, the lower

ones denticulate, the others entire. Young fruits ovoid-globose,.

attenuate towards the conical, ringed umbo, sericeous; styles 3,.

recurved, sericeous. Mature cupule flattened cup-shaped, c. 0.7-1

cm in depth and c. 2 cm in diameter, covering c. }—# part of

the fruit, with dense tomentum, base rounded; lamellae c. 6-8,

thin, more or less free at the rim. Mature fruit ovoid-globose, c. 2

cm in length and in diameter, with dense tomentum, apex round-

ed, umbonate, base convex.

Ecology: in primary heath forest on terrace sands, at alt. c.

1200 m. Fruiting in June-July.

Distribution: so far only known from Sarawak.

10. Quercus chrysotricha A. Camus, Fig. 11 — la-c, Map X.

Quercus chrysotricha A. Camus, Les Chénes 3, Expl. Pl.: 50;

Atl. t. 347, f£. 8-14. 1948; Text: 1212. 1954. — Quercus arbutifolia

(non Hickel & A. Camus) Warburg in Kew Bull.: 19. 1936. —

Type: Richards 1885, Sarawak, Mt. Dulit, alt. c. 1300 m, Sept. fr.

_(Holotype: K; isotype: L).

Tree c. 19 m tall, trunk c. 30 cm in diameter. Bark smooth or

finely fissured. Young branchlets with dense, brownish, simple-

pubescence; older branchlets glabrous, lenticellate. Petiole c. 1.0—

0.5 cm long, flattened on the adaxial side, with dense pubescence,

glabrescent. Leaves thin-coriaceous, obovate-elliptic, base atten-

uate-rounded or attenuate-acute, slightly asymmetrical, margin

entire, apex rounded or truncate-emarginate; c. 1-5.5 by 1.5-3.5

cm; upper surface shining, glabrous, lower surface glabrous or with

sparse pubescence. Midrib and lateral nerves slightly prominent.

beneath, flattened or slightly impressed above; reticulation obscure

on both surfaces. Lateral nerves c. 4-5 pairs, forming c. 60°-70°

angle with the midrib, parallel, arcuating towards the margin.

Cupule cup-shaped, obconical, c. 1-1.5 cm in depth and in dia-

meter, base attenuate-acute, covering c. }-1 part of the fruit;.

lamellae thin, c. 6-7, with dense tomentum outside, denticulate.

Fruit ovoid-conical, c. 1.5—2 cm in length and c. 1-1.5 cm in dia-

meter, with dense tomentum, glabrescent; apex attenuate-acute.,,.

base convex.

Ecology: in submontane forests, at c. 1200-1300 m alt. Fruiting

in September.

Distribution: endemic in Borneo (Mt. Dulit, Sarawak).

11. Quercus merrillii von Seemen, Fig. 11 — 2a-c, Map X.

Quercus merrillii von Seemen in Fedde’s Rep. 5: 21. 1908;

Merrill in Philip. J. Sc. Bot. 3:°329. 1908; Enum. Philip. Fl. Pl.

2: 28. 1923; Koidzumi in Bot. Mag. Tokyo 30; 201. 1916; A.

Camus, Les Chénes /: 210. 1938; Atl. J, t. 7, f. 6-9. 1934. —

Cyclobalanopsis merrillii (v. Seemen) Schottky in Bot. Jahrb. 47:

649. 1912. — Type: Foxworthy 566, Palawan Isl., fr. April 1906

(Holotype: K).

Gardens’ Bulletin, Singapore — XXH (1968 }

398

IN )

Quercus chrysotricha, after Richards 1885 (Type).

Quercus merrilli, after Curran 3858.

2a-c

Fig. 11. la-c

Soepadmo — Quercus subgen. Cyclobalanopsis 399

Small tree. Young branchlets with dense brownish, stiff-pubes-

cence; older branchlets glabrous, lenticellate. Stipules linear-acute,

with dense pubescence outside, c. 0.5-1 by 0.1 cm. caducous.

Petiole c. 0.3-0.6 cm long, flattened on the adaxial side, with dense

pubescence, glabrescent. Leaves thin-coriaceous, obovate or elliptic-

ovate, c. 1.5—5 by 0.7—-2.5 cm, base attenuate-acute, margin remotely

serrulate in the apical half, apex bluntly acute; with dense pubes-

cence on both surfaces, glabrescent. Midrib and lateral nerves

prominent beneath, flattened or impressed above; reticulation

obscure on both surfaces. Lateral nerves c. 5-8 pairs, parallel,

forming c. 50°-60° angle with the midrib, arcuating towards the

margin. Inflorescences unknown. Young infructescence c. 1-2 cm

long, with dense tomentum, glabrescent, lenticellate, with 1-2

solitary young fruits. Young cupule obovoid or obconical, c. 0.5

cm in depth and in diameter, with dense tomentum; styles 3, c.

1-2 mm long, with sparse tomentum at base. Mature cupule cup-

shaped, obconical, covering c. }-1 part of the fruit, c. 1 cm deep

and 1-1.5 cm in diameter; lamellae c. 7-8, thin, denticulate, with

dense tomentum outside. Fruit ovoid or ovoid-cylindrical, c. 2—2.5

cm long and 1-1.5 cm in diameter, shining, glabrous; apex acute,

_ base convex. |

Ecology: in lowland forest, at c. 100-500 m alt. Fruiting between

April and August.

Distribution: Borneo and Palawan Isl.

12. Quercus kerangasensis Soepadmo, spec. nov., Fig. 12, Map XI.

Arbor c. 20-30 m alta, trunco c. 25-50 cm lata; cortice laevi vel

scabro, grisei-lenticellato. Ramuli laeves vel dense lenticellati

annulati, grisseo-brunnei, dense et rigide pubescentes, glabrecentes.

Alabastra c. 2-4 x 3-4 mm, ovoideo-globosa, dense pubescens.

Stipulae lineares acutae, c. 2-3 x 0.5-1 mm, dense pubescentes,

glabrescentes. Petiolus gracilis, c. 8-12 x 0.5-1 mm, ad basim

incrassatus, supra applanatus vel subsulcatus. Lamina c. 4-11 x

2-4 cm, elliptico-lanceolata, coriacea, subtus glabra, glaucescens,

supra nitida glabra; costa nervisque lateralibus subtus prominen-

tibus, supra vix elevatis; nervis lateralibus c. 7-10, angulo 60°—70°

exorientibus, ascendentibus, marginem versus arcuatis; nervis reti-

culatis invisibilibus.

Inflorescentia mascula c. 1-2 cm longa, axillaris semiterminalis,

dense pallide brunneo-pubescens. Bracteae c. 1 X 1 mm, ovatae

acutae, dense et rigide brunneo-pubescentes; cymulis trifloris.

Perianthium 5—6-lobatum, extus pubescens; stamina 5-6; filamentis

c. 1-2 cm longis, glabris filiformibus; antheris c. 0.5 x 0.5 mm.

Inflorescentia feminea ignota.

Cupula matura obconica cupuliformis, c. 0.8-1.2 cm alta, 1.5-2

cm lata, dense brunneo-sericea; lamellae c. 5—7, crasse, marginem

crenato vel integro. Nux matura c. 2-3 cm longa, 1.2—1.5 cm lata,

Ovoideo-conica vel obovoideo-cylindrica, dense sericea, glabrescens;

apice rotundato vel acuto, ad basim rotundato; hilum convexum.

Type: Brunig S. 1065, Badas Forest Reserve, Sarawak, alt. c.

10 m, September fr. (Holotype: K; isotypes: L, SAR).

400

Gardens’ Bulletin, Singapore — XXII (1968)

Fig. 12: Quercus kerangasensis

1-2 after Brunig 4661 (2 = male inflorescences); 3-4 after Brunig 1065

(Type)

Soepadmo — Quercus subgen. Cyclobalanopsis 401

Tree c. 20-30 m tall, trunk c. 25-50 cm in diameter. Buttresses

small, fluted. Bark smooth or rough, lenticellate, grey. Young

branchlets with dense stiff-pubescence, smooth or horizontally

crasked; older branchlets glabrous, greyish-brown, with dense

warty lenticels. Buds ovoid-globose, with dense yellowish-brown

pubescence, c. 3-6 by 3-4 mm. Stipules linear-acute, c. 2-3 by

0.5—1 mm, caducous. Petioles slender, c. 0.8-1.2 cm long and 0.5-1

mm thick, with dense pubescence, glabrescent, thickened at base,

flattened or shallowly furrowed on the adaxial side. Leaves elliptic-

lanceolate, coriaceous, c. 4-11 by 2-4 cm, upper surface shining,

glabrous, lower surface pale glaucous, glabrous except the midrib;

margin entire, base attenuate-acute, apex acute or 0.5-l1 cm

acuminate. Midrib and lateral nerves prominent beneath, slightly

so above. Lateral nerves c. 7-10 pairs, ascendant, subparallel,

arcuating towards the margin, forming c. 60°—70° angle with the

midrib; reticulation obscure on both surfaces. Male inflorescence

in paniculate clusters on the lateral or subterminal new shoots, with

dense yellowish-brown pubescence. c. 1-2 cm long. Bracts mem-

branous, ovate-acute, with dense stiff-pubescence outside, c.

1 by 1 mm. Male flowers in 3-flowered dichasial clusters; perianth

membranous, 5-6-lobed; stamens 5-6; filaments glabrous, fili-

form, c. 1-2 mm long; pistillode absent. Female inflorescence not

known. Mature cupule cup-shaped, obconical, c. 0.8-1.2 cm in

depth and 1.5—2 cm in diameter, base attenuate, rim c. 1-1.5 mm

thick, with dense brownish tomentum on both surfaces; lamellae

c. 5-7, margin entire or denticulate. Mature fruit ovoid-conical or

ovoid-cylindrical, c. 2-3 cm long and 1.2—1.5 cm in diameter, dense-

ly sericeous, glabrescent, shining; apex rounded or acute, base

convex.

Ecology: in primary heath forests, at c. 10-100 m alt. Flower-

ing in June, fruiting in September.

Distribution: endemic in Borneo (Brunei, Sarawak).

Notes: This species is named after the local name of heath

forest, viz. “kerangas” forest, from where the specimens were

mainly collected. It may be recognized by its elliptic-lanceolate

leaves with entire margin and pale glaucous lower surface, and

by its small fruit and cupule.

13. Quercus kinabaluensis Soepadmo, spec. nov., Fig. 13, Map XI.

Arbor c. 10-40 m alta, trunco c. 20-30 cm lato: cortice laevi.

Ramuli sparsim lenticellati, dense et ridgide brunneolo-pubescen-

tes, glabrescentes. Alabastra, c. 4-10 x 3-4 mm, ovoideo-globosa

vel ovoideo-conica. Stipulae c. 5-10 x 0.5-1 mm, lineares acutae,

dense et rigide pubescentes, caducae. Petiolus c. 1-1.5 cm longus,

1-1.5 mm crassus, dense brunneolo-pubescens, glabrescens, ad

basim incrassatus, supra applanatus. Lamina c. 5-10 x 2-5 cm,

ovato-elliptica, crasse coriacea,’ dense flavo-brunneolo-pubescens,

glabrescens; margine integro, basi asymmetrico rotundato vel

acuto; apice acuto acuminato, 0.5-1.5 cm longo; costa nervisque

lateralibus subtus vix elevatis, supra applanatis vel subsulcatis;

nervis lateralibus c. 6-8, ascendentibus, marginem versus arcuatis,

angulo 45°-60° exorientibus; nervulis reticulatis invisibilibus.

402

Gardens’ Bulletin, Singapore — XXII (1968)

Fig. 13: Quercus kinabaluensis

1-4 after Chew & Corner RSNB. 4451; 5-6 after Chew & Corner

RSNB. 7148 (2 = a 3-flowered male dichasium; 4 = female

inflorescence).

Soepadmo — Quercus subgen. Cyclobalanopsis 403

Inflorescentia mascula gracilis flexuosa, c. 50-110 x 1 mm, dense

brunneo-pubescens; cymulis tri vel uni-floris. Bracteae c. 2-3 x

1-2 mm, ovatae acutae, tenue, extus dense pubescentes. Perian-

thium 4~7-lobatum, extus dense brunneo-pubescens, glabrescens;

stamina 4-7; filamentia c. 2 mm, glabris; antheris c. | x 1 mm.

Inflorescentia feminea c. 10-30 x 2-3 mm, dense et rigide

brunneo-pubescens, 1—3-flores gerens. Flores femini solitarii. Brac-

teae c. 1-2 x 0.5—1 mm, dense pubescentes, glabrescentes. Perian-

thium 5-6-lobatum, crasse coriaceum, extus dense _ brunneo-

pubescens; staminodia nil; stili 3-5, recurvi, ad basim versus hirti,

c. 2-3 x 0.5—1 mm.

Cupula immatura obconica cupuliformis, c. 0.7—1 cm alta, 1—1.5

cm lata, margine crassa, basi attenuati; lamellae 6-8, dense pubes-

centes, denticulo vel integro. Nux c. 1—-1.2 cm alta, 0.8-1 cm lata,

oOvoideo-conica, dense brunneo-sericea, glabrescens; apice rotun-

dato vel attenuato, basi convexo.

Type: Chew & Corner RSNB. 7148, Mt. Kinabalu, Mentaki

ridges, alt. c. 2600 m, March fr. (Holoytpe: K; isotypes: L).

Tree c. 10-40 m ttall, trunk c. 20-30 cm in diameter. Bark

‘smooth, brownish with white patches. Innovations with dense

brownish, stiff-pubescence, glabrescent. Branchlets glabrous, with

sparse lenticels. Buds ovoid-globose or ovoid-conical, c. 0.4-1 by

0.3-0.4 cm. Stipules linear-acute, c. 5-10 by 0.5-1 mm, caducous,

Petiole c. 1-1.5 cm long and 1-1.5 mm thick, with dense brownish,

stiff-pubescence, glabrescent, slightly thicken at base, flattened on

the adaxial side. Leaves c. 5-10 by 2-5 cm, ovate-elliptic, thick-

coriaceous, with dense yellowish-brown pubescence on both sur-

faces, glabrescent; margin entire, base asymmetrical, rounded or

acute, apex acute or 0.5-1.5 cm acuminate. Midrib, and lateral

nerves slightly prominent beneath, flattened or impressed above.

Lateral nerves c. 6-8 pairs, ascendant, arcuating towards the mar-

gin, forming a c. 45°-60° angle with the midrib; reticulation

obscure on both surfaces. Male inflorescence in paniculate clusters

on a lateral new shoots, with dense yellowish-brown, stiff-pube-

scence c. 5-11 cm long and 1 mm thick; bracts ovate-acute, mem-

branous, with dense pubescence outside, c. 2-3 by 1-2 mm. Male

flowers in 3-flowered dichasial clusters or solitary along the rachis

perianth 4—7-lobed, with brownish-pubescence outside, glabrescent;

stamens 4~7; filaments c. 2 mm long, glabrous; anthers c. 1 X 1

mm. Female inflorescence c. 1-3 cm long, 2-3 mm thick, with

dense stiff-pubescence, carrying 1-3 flowers. Female flower:

perianth thick-coriaceous, with dense brownish-pubescence out-

side, 5—6-lobed; staminodes 0; styles 3-5, hairy at base, recurved,

c. 2-3 by 0.5-1 mm. Immature cupule cup-shaped, obconical, c.

0.7—1 cm deep and 1-1.5 cm in diameter, rim thick, base attenuate;

lamellae c. 6-8, denticulate or entire, with brownish-pubescence on

both surfaces. Fruit ovoid-conical, with dense brownish tomentum,

glabrescent, c. 1-1.2 cm long and 0.8-1 cm in diameter; apex

rounded or attenuate; base convex.

ae CU

404 Gardens’ Bulletin, Singapore — XXII (1968)

Ecology: in ridge or montane forests, at c. 500-2600 m alt.

Fruiting between March and May.

Distribution: endemic in Mt. Kinabalu, Borneo.

Notes: In vegetative characters Q. kinabaluensis is closely allied

to lowii, but differs from the latter by its thick-rimed cupule and

different type of pubescence.

14. Quercus valdinervosa Soepadmo, spec. et stat. nov., Fig. 14,

Map XII.

Quercus mespilifolia Wali. ex A.DC. var. borneensis Heine in

Fedde’s Rep. 54: 225. 1951. — Lectotype: Clemens 31317, Mt.

Kinabalu, North Borneo, June fl. (Holotype: K: isotypes: A, B,

K, L, SING, UC); paratype: Anderson S. 4547, Baram, path to

the summit of Mt. Mulu, Sarawak, alt. c. 1900-2100 m, fr. 2-VII-

1961 (A, K, L, SAR, SING).

Tree c. 20-35 m tall, trunk c. 30-60 cm in diameter. Bark

smooth, grey. Innovations with dense brownish, stiff, simple- or

stellate-pubescence. Branchlets glabrous, lenticellate. Buds ovoid-

conical; scales arranged in four vertical rows. Stipules linear-

acute, tomentose outside, c. 1-1.5 cm long, 1-2 mm wide, caducous.

Petiole c. 1-2.5 cm long, glabrous, flattened on the adaxial side.

Leaves elliptic or obovate-oblong, c. 8-15 by 3-6 cm, thick-coria-

ceous; upper surface glabrous, lower surface with dense simpie

pubescence, soon becomes completely glabrous; base acute or

attenuate-rounded, margin remotely serrulate in the apical half,

apex 0.5-1 cm acuminate. Midrib and lateral nerves prominent

beneath, flattened or impressed above; reticulation dense, parallel,

prominent beneath. Lateral nerves c. 10-15 pairs, dense, parallel

and straight, arcuating near the margin, forming c. 60° angle with

the midrib. Male inflorescence c. 5-10 cm; bracts lanceolate or

linear-acute, membranous, c. 3-5 by 2-3 mm. Male flowers in

3-flowered dichasial clusters along the rachis; perianth 4—6-lobed,

with dense tomentum outside; stamens 5-9; filaments c. 3-4 mm

long, glabrous; anthers c. 1 x 1 mm; pistillode 0. Female inflores-

cence c. 1-1.5 cm long, with dense tomentum, carrying 4—6 female

flowers; bracts ovate or linear, c. 2-3 by 1 mm. Female flowers:

perianth 6-lobed, with dense tomentum outside; staminodes 0:

Styles 3-6, c. 1-2 mm long, tomentose at base. Mature cupule cup-

shaped, obconical, base attenuate, c. 1.5-1.7 cm in depth and

1-2.5 cm in diameter, covering c. 1/3-1/6 part of the fruit; lamel-

lae 7-8, rim crenate and thick, with dense tomentum outside. Fruit

cylindrical-conical or ellipsoid, apex acute, base convex, c. 3—-3.7

cm in length, 1.3-1.7 cm in diameter, tomentose.

Ecology: in submontane to montane forests, at c. 1300-2300

m alt. Fruiting between February and August.

Distribution: Borneo (Mt. Kinabalu, Brunei, Sarawak, W.

Kutei).

Soepadmo — Quercus subgen. Cyclobalanopsis 405

Fig. 14: Quercus valdinervosa

1 after Ashton Brun. 2378; after Meijer SAN. 29134.

| aii

406 | Gardens’ Bulletin, Singapore — XXII (1968 }

15. Quercus gemelliflora Bl., Fig. 15, Map XIII.

Quercus gemelliflora Blume in Verh. Bat. Genootsch. 9: 223, t.

6. 1823, p.p., excl. fructu; Bijdr.: 523. 1825; Fl. Javae Cupul.: 30,

t. 17. 1829; Mus. Bot. Lugd. Bat. 7: 295. 1850; Endlicher, Gen.

Pl., Suppl. 4, 2: 28. 1847; Miquel, Fl. Ind. Bat. J, 1: 854. 1856;

Suppl. /: 854. 1860; Ann. Mus. Lugd. Bat. 7: 111. 1863; A. DeCan-

dolle, Prodr. /6, 2: 88. 1864; Wenzig in Jahrb. K6n. Bot. Gart..

Berlin 4: 228. 1886; King in Ann. Roy. Bot. Gard. Calc. 2: 88.

1889; Koorders & Valeton, Bijdr. Booms. Java 10: 24. 1904;

Koorders, Exk. 2: 61. 1912; Atl, t. 41. 1913; A. Camus, Les.

Chénes /, 363. 1938; Atl. 2, t. 234. 1936; Backer & Bakhuizen van

den Brink Jr., Fl. Java 2: 5. 1965. — Cyclobalanopsis gemelliflora

(BI.) Oersted in Vidensk. Medd. naturh. For. Kjobn. 8: 77. 1867. —

Type: Blume s.n., Mt. Salak, Java, st. (Holotype: L).

Quercus turbinata Blume, Bijdr.: 523. 1825; Fl. Javae Cupul.:

31, pl. 18. 1829; Mus. Bot. Lugd. Bat. 7: 302. 1850; Miquel, Fl.

Ind. Bat. 7, 1: 855. 1856; Ann. Mus. Bot. Lugd. Bat. J: 114. 1863;

Scheffer in Nat. Tijdschr. Ned. Ind. 37: 361. 1870; lc. 32: 417.

1872; Koorders, l.c.: 60, f. 10. 1912; A. Camus, lc.: 204. 1938;

Atl. J, t. 6, f. 6-16. 1934. — Cyclobalanopsis turbinata (Bl.)

Schottky in Bot. Jahrb. 47; 648. 1912. — Quercus merkusii

Endlicher, Gen. Pl., Suppl. 4, 2: 28. 1847; A. DeCandolle, l.c.:

98. 1864; — Cyclobalanopsis merkusii (Endl.) Oersted, l.c.: 79.

1867. — Quercus lineata Bl. var.’merkusii Wenzig in Jahrb. K6n.

Bot. Gart. Berlin 4: 232. 1886; King, l.c.: 32, pl. 26, f. 2. 1889. —

Type: Blume s.n., Mt. Salak, Java, yfr. (Holotype: L).

Quercus horsfieldii Miquel, |.c.: 856. 1856; Suppl.: 869. 1860;

A. DeCandolle, I.c.: 99. 1864. — Cyclobalanopsis horsfieldii (Miq.)

Oersted, l.c.: 78. 1867. — Type: Horsfield 11, Bangka, yfr. (Holo-

type: U; isotype: K); syn. nov.

Quercus horsfieldii Mig. var. longifolia Miquel, Fl. Ind. Bat.

1, 1: 869. 1856. — Type: Diepenhorst 42, Priaman, Sumatra, st.

(Holotype: U); syn. nov.

Quercus semiserrata (non Roxb.) King, Lc.: 28. 1889, p.p.,

quoad specim. ex Bangka et Sumatra; Koorders & Valeton, l.c.:

25. 1904: Koorders, l.c.: 61. 1912; Lc. t. 55. 1913; A. Camus, L.c.:

186. 1938.

Quercus turbinata Bl. var. crassilamellata Gamble in J. As. Soc.

Bengal 75: 410. 1915; Ridley, Fl. Mal. Pen. 3: 373. 1924. —

Quercus crassilamellata (Gamble) A. Camus in Bull. Mus. Paris,

Ser. II, 3: 689. 1931; Les Chénes /; 226. 1938; Atl. J, pl. 10, f. 1-7.

1934. Type: Wray 1532, Kampong Ulu Batang Padang, Perak,

Malaya, at c. 1600 m alt., fr. (Holotype: K; isotype: SING); syn.

nov.

Tree c. 20-30 m tall, trunk c. 20-60 cm in diameter. Buttresses

up to 1 m tall. Bark smooth or finely fissured, greyish-brown.

Young branchlets and buds with dense tomentum; older branchlets

glabrous, lenticellate. Stipules linear-acute, c. 0.5-1 by 0.1-0.2 cm,

with dense tomentum outside, caducous. Petiole c. 1-3 cm long

407

ICN

Es. 1964,

Fig. 15: Quercus gemellifiora .

1 after Rastini s.n.; 2 after Biinnermeijer 1880; 3 after Teijsmann s.n.

408 Gardens’ Bulletin, Singapore — XXII (1968 );

and 0.1-0.2 cm thick, with dense tomentum, glabrescent. Leaves:

thin-coriaceous, elliptic-lanceolate or elliptic-oblong, c. 5-15 by-

2-5.5 cm; base attenuate-acute, asymmetrical, margin remotely

serrulate in the apical half, apex shortly acuminate; upper surface:

glabrous, lower surface with dense pubescence, soon glabrescent.

Midrib and lateral nerves prominent beneath, flattened or slightly

raised above, especially the midrib; reticulation obscure on both:

surfaces. Lateral nerves c. 8-10 pairs, parallel, straight, arcuating.

towards the margin, forming c. 60° angle with the midrib. Male

inflorescence c. 6 cm long, with dense brownish tomentum; out-.

side, caducous. Male flowers solitary or in 3-flowered dichasial

clusters along the rachis; perianth 3-6-lobed; stamens 3-6; fila-.

ments c. 2 mm, hairy at base; anthers c. 0.5-1 mm long; pistillode:

0. Female inflorescence carrying c. 2-7 flowers, with dense brownish

pubescence; bracts ovate-acute, c. 1-2 by 1 mm. Female flowers:

perianth 4-6-lobed, with dense tomentum; staminodes 0-6; styles.

recurved, c. 1-2 mm long, tomentose at base. Young cupule tur-

binate-obconical or cylindrical-globose, covering the entire or the

greater part of the fruit, base attenuate; lamellae c. 5—7, rim thick,

entire or denticulate, with dense tomentum. Mature cupule cup-

shaped, c. 1.5-2 cm deep and 1.5-2.5 cm in diameter, covering

c. 4-1/3 part of the fruit, rim c. 2 mm thick; lamellae c. 7-8, with

dense tomentum. Ripe fruit conical or cylindrical, c. 2-5.5 cm

long and 1-2 cm in diameter, apex acute, base rounded, with

dense tomentum, glabrescent.

Ecology: in lowland to submontane forests, at c. 100-1600 m

alt. Flowering and fruiting the whole year round.

Distribution: Malaya (Kedah, Penang, Perak Pahang, Negri

Sembilan, Malacca, Johore), Sumatra (Atjeh, Sibolangit,

Tapanuli, Padang, Pajakumbuh, Mt. Sago, Indragiri, Pema-

tang Siantar, Banjuasin-Palembang), Bangka, Borneo (San-

dakan, Mt. Kinabalu, Sarawak, W. Kutei, Martapura and.

Vicinity), Java.

16. Quercus lineata BI., Fig. 16, Map XIV.

Quercus lineata Blume, Bijdr.: 523. 1825; Fl. Javae Cupul.:

32, pl. 19. 1829; Mus. Bot. Lugd. Bat. 7: 302. 1850; Miquel, FI.

Ind. Bat. 7, 1: 855. 1856; in Ann. Mus. Bot. Lugd. Bat. /: 114.

1863; A. DeCandolle, Prodr. 16, 2: 98. 1864; Wenzig in Jahrb.

Kon. Bot. Gart. Berlin 4: 232. 1886; Koorders & Valeton, Bijdr.

Booms. Java 10: 20. 1904; Koorders, Exk. 2: 60. 1912; Atl.

Baumart. Java J: pl. 58. 1913; A. Camus, Les Chénes /: 207.

1938; Atl. J: t. 6, f. 1-5. 1934; Backer & Bakhuizen van den

Brink Jr., Fl. Java 2: 5. 1965. Cyclobalanopsis lineata (B1.)

Oersted in Vidensk. Medd. naturh. For. Kjébn. 8: 78. 1867; in Lib-

mann. Chénes Am. Trop. J: 20. 1869. — Quercus lineata Bl. var.

typica King in Ann. Roy. Bot. Gard. Calc. 2: 33, pl. 26, f. 1. 1889.

— Type: Blume s.n., Java, fl. (Holotype: L; isotypes: A, BO).

Quercus polyneura Miquel, Pl. Jungh. J: 11. 1851. — Quercus

lineata Bl. var. heterochroa Miquel, l.c.: 855. 1856. — Type:

Junghuhn 9, Priangan, Java, yfr. (Holotype: U).

-Soepadmo — Quercus subgen. Cyclobalanopsis 409

Quercus oxyrhyncha Miquel, Fl. Ind. Bat., Suppl. 7: 347. 1860

-—Quercus lineata Bl. var. oxyrhyncha (Miq.) von Seemen in Bull.

Dép. Agr. Ind. Néerl. 7: 4. 1906. — Type: Teijsmann H.B. 676,

Alahan Pandjang, Sumatra, st. (Holotype: VU).

Quercus hendersoniana A. Camus in Bull. Mus. Paris, Sér. L.,

4: 123. 1932; Les Chénes J: 210. 1938; Atl. 7: t. 6, f. 6-8. 1934. —

“Type: Henderson SFN. 18053, Batu Brinchang, Pahang, Malaya,

alt. c. 1600 m, Nov. fr. (Holotype: SING).

Tree c. 20-30 m tall, trunk c. 20-60 cm in diameter. Bark

‘smooth grey; inner bark fibrous, reddish-brown. Young branch-

lets dark-grey with sparse lenticels, with dense stiff-pubescence,

glabrescent; older branchlets glabrous, lenticellate. Buds globose

or ovoid-conical, densely puberulous, glabrescent. Stipules linear-

acute, c. 1-1.5 by 0.1-0.2 mm, with dense pubescence outside,

caducous. Petiole c. 1-2 cm long, shallowly furrowed on the

adaxial side, with dense pubescence, glabrescent. Leaves thin-

coriaceous, ovate-elliptic or ovate-lanceolate, c. 5-16 by 2-6

cm, base attenuate-acute, sometimes asymmetrical, margin re-

motely serrulate in the apical half, apex acute, acuminate or c.

0.5-1.5 cm caudate; upper surface with sparse pubescence,

especially on the midrib and lateral nerves, lower surface with

dense, adpressed, simple-pubescence. Midrib and lateral nerves

prominent beneath, slightly so above; reticulation prominent be-

low, obscure above, dense, parallel. Lateral nerves c. 10-20 pairs,

straight, parallel, dense, arcuating near the margin but not

anastomosing; forming c. 45°-60° angle with the midrib. Male

inflorescence c. 5-10 cm long, with dense pubescence; bracts

ovate-acute, c. 2-3 by 2 mm, membranous, densely sericeous

outside, glabrescent. Male flowers solitary or in 3-flowered

dichasial clusters along the rachis; perianth 5—6-lobed, with dense

tomentum outside; stamens normally 6; filaments c. 0.5-1 mm

long, glabrous; anthers c. 0.5 by 0.5 mm; pistillode reduced to

a cluster of stiff, simple-hairs. Female inflorescence c. 1.5—2 cm

long, with dense pubescence, carrying c. 5-6 flowers. Bracts

-Ovate-acute, c. 1-2 mm, with dense tomentum. Female flowers:

perianth 5—6-lobed; staminodes 0; styles 3 recurved, c. 1-3 mm

long. Young cupule ovoid-globose, densely sericeous, lamellae

. 3-4, thick, the 2 lower ones denticulate, the rest entire.

Mature cupule cup-shaped, obconical, c. 1—-1.2 cm deep, 2-2.5

cm in diameter, attenuating towards the base, with dense tomen-

tum outside, rim thick; lamellae c. 8-10, free. Mature fruit

conical-cylindrical, c. 2-3 cm in length, 1-2 cm in diameter, with

dense tomentum; apex attenuate-rounded, base convex.

Ecology: in submontane to montane forests, at c. 1000-2000

alt. Fruiting between August and April.

Distribution: Malaya (Pahang), Sumatra (Lake Toba vicinity,

Tapanuli, Alahan Pandjang), Borneo (Mt. Kinabalu), Java

(Mt. Karang, Mt. Malabar, Mt. Salak, Mt. Tangkuban Prahu,

Mt. Papandajan, Tjidatar, Tjigenteng, Mt. Telomojo,

Madiun).

Garaens’ Bulletin, Singapore — XXI] (1968 )

410

Fig. 16: Quercus lineata

la-b after Chew & Corner RSNB. 4561; after Chew & Corner RSNB.

4927.

Soepadmo — Quercus subgen. Cyclobalanopsis 41}

17. Quercus steenisii Soepadmo, Map III.

Quercus steenisii Soepadmo in Gard. Bull. Singapore 2/, 3:

389, f. 5. 1966.— Type: van Steenis 8607, Mt. Losir, Atjeh, N.

Sumatra, alt. c. 3300-3460 m. Febr. fr. (Holotype: BO; insotypes:

K, L, SING); paratype: van Steenis 8379, Putjuk Angasan, Atjeh,

N. Sumatra, alt. c. 2000 m, Jan. fl. (BO, K, L, PNH, SING).

Tree c. 15 m tall, trunk c. 60 cm in diameter. Young branchlets

with dense brownish-stiff-pubescence; older branchlets glabrous,

lenticellate. Buds ovoid-ellipsoid, c. 1-1.5 by 0.5 cm. Stipules

linear with blunt tip, c. 0.5-1 by 0.1-0.2 cm, with dense pub-

escence, caducous. Petiole c. 0.2-0.5 cm long, thickened at base,

flattened on the adaxial side, with dense pubescence, glabrescent.

Leaves thick-coriaceous, ovate-elliptic or ovate-orbicular, c, 3-8

by 2-5 cm, base rounded, obtuse, or cordate, margin entire or

remotely serrulate in the apical half, apex rounded emarginate

or bluntly acute; upper surface of the young leaf with dense

brownish, stiff-simple-pubescence, glabrescent, lower surface with

thick layer of brownish, woolly pubescence, glabrescent. Midrib

and lateral nerves prominent beneath, keeled or flattened above;

reticulation prominent beneath, parallel, dense. Lateral nerves c.

6-10 pairs, straight, parallel, arcuating towards the margin, form-

ing c. 45°-60° angle with the midrib. Male inflorescence c. 5

cm long, with dense pubescence; bracts ovate-obtuse, c. 0.2 by

0.1 cm, densely pubescent. Male flowers in 3-flowered dichasial

clusters; perianth 4~6-lobed; stamens 4-6; filaments glabrous,

c. 1-1.3 mm long; anthers c. 1 by 0.5-1 mm; pistillode reduced

to a cluster of stiff, simple-hairs. Female inflorescence not known.

Young infructescence c. 1.5-3 cm long, with spares lenticels,

with brownish, simple-pubescence; carrying 2-7 young fruits;

bracts ovate-acute, caducous. Perianth of the female flowers (seen

in the young fruit) thick-coriaceous, 5-6-lobed, with dense pub-

escence outside; staminodes 0-6, rudimentary; styles 3-4, re-

curved, c. 1-2 mm long. Young cupule ovoid-globose, with

dense pubescence ouside; lamellae thick, c. 3-4, the 2 lower

ones denticulate, the others entire. Mature cupule cup-shaped,

obconical, or patelliform, c. 0.7-1 cm in depth, 1.2-2.5 cm in

diameter, enclosing c. 4-4 part of the fruit; lamellae c. 7-8,

more or less recurved, with dense brownish pubescence. Fruit

globose or ovoid, with dense brownish-tomentum, glabrescent,

c. 1-2 cm in length and in diameter, apex rounded- or depressed-

umbonate, base convex or flat.

Ecology: in montane forest, at c. 2500-3500 m alt. Flowering

in January 28, 1937; fruiting in February 2, 1937.

Distribution: so far only known from Mt. Losir, Atjeh, N.

Sumatra.

412 Gardens’ Bulletin, Singapore — XXII (1968 }

18. Quercus treubiana von Seemen, Fig. 17, Map XV.

Quercus treubiana von Seemen in Bull. Dép. Agr. Ind. Néerl..

I: 3. 1906; in Fedde’s Rep. 3: 173. 1907; Merrill in J. Str..

Br. Roy. As. Soc., Spec. No.: 216. 1921; A, Camus, Les Chénes-

I: 368. 1938; Alt. 7: t. 33, f. 15-17. 1934. — Cyclobalanopsis

treubiana (v. Seem.) Schottky in Bot. Jahrb. 47: 648. 1912. —

Type: Hallier 2915, Mt. Liang Gagang, Borneo, fr. (Holotype:

L; isotypes: K, SAR).

Tree c. 30 m tall, trunk c. 60 cm in diameter, bole irregular..

Buttresses up to 1 m tall. Bark rough,, peeling off profusely

into rectangular pieces, rusty; inner bark c. 1.5 cm thick, ridged.

Innovations with dense brownish-simple-pubescence. Older bran-

chlets glabrous, lenticellate. Buds ovoid-ellipsoid; scales linear-

acute. Stipules linear-acute, c. 5-10 mm long, caducous, densely

pubescent outside. Petiole c. 0.5-1.5 cm long, flattened on the

adaxial side, with dense pubescence, glabrescent. Leaves thin-

chartaceous, linear-lanceolate or elliptic-lanceolate, c. 3-10 by

1-3 cm; base attenuate-acute, sometimes asymmetrical, margin

remotely serrulate in the apical half, apex sharply acute or

0.5-1 cm acuminate; lower surface with dense simple-pubescence,.

glabrescent, upper surface glabrous, except for the midrib and

lateral nerves. Midrib and lateral nerves more or less prominent

on both surfaces; reticulation obscure on both surfaces. Lateral

nerves c. 5-10 pairs, straight, ‘parallel, arcuating towards the

margin, forming c. 45°-60° angle with the midrib. Male infior-

escence c. 3 cm long, rachis c. 0.1 cm thick, with dense pub-

escence; bracts and bracteoles linear-acute, densely sericeous

outside, glabrescent. Male flowers solitary or in 3-flowered dichas-

ial clusters along the rachis; perianth 5—6-lobed, the lobes densely

sericeous outside; stamens 5—6; filament c. 2 mm long, puberulous

at base; anthers c. 0.5-1 mm in length; pistillode reduced to

a cluster of stiff-simple hairs. Female inflorescence not known.

Young infructescence c. 1-2 cm long, rachis c. 1-2 mm thick,

with dense woolly pubescence, glabrescent, carrying 2-5 young

fruits; bracts ovate-acute, c. 1-1.5 by 0.5-1 mm, with dense

tomentum outside. Perianth of the female flowers (seen on the

young fruit) 5—6-lobed; styles 3, recurved, c. 2 mm long, with

dense tomentum at base. Young cupule ovoid-globose or obconi-

cal, lamellae c. 6, with dense tomentum outside. Mature cupule

cup-shaped, subglobose, base attenuate-rounded, c. 1.5—2 cm deep,

1-2 cm in diameter, covering c. +4 part of the fruit; lamellae

10-12, thick, densely sericeous outside. Mature fruit cylindrical-

globose, c. 2-3 cm long and 1.5—2 cm in diameter; apex depressed-

umbonate; base convex.

Ecology: in low ridges to montane forests at c. 600-2000 m

alt. Fruiting in July-August. |

Distribution: Sumatra (Palembang), Borneo (Mt. Kinabalu,

Mt. Liang Gagang, Mt. Beratus, Mt. Palimasan).

‘Soepadmo — Quercus subgen. Cyclobalanopsis 413

Esky.

Fig. 17: Quercus treubiana

la-b after Tikau SAN: 34643; 2a-f after Kostermans 7442; 3 after

Clemens 30465.

414 Gardens’ Bulletin, Singapore — XXII (1968)

19. Quercus elmeri Merr., Fig. 18, Map XVI.

Quercus elmeri Merrill in Univ. Calif. Publ. Bot. /5: 43.

1925: A. Camus, Les Chénes, /: 194. 1938; Atl. J: t. 4, f. 9-12.

1934. — Type: Elmer 21213, Tawao, N. Borneo, fr. Oct. 1923

{Holotype: K; isotypes: A, L).

Tree c. 18-40 m tall, trunk c. 25-60 cm in diameter. Bark

greyish-brown, cankered with longitudinal rows of lenticels or

scaly; inner bark c. 0.5-1.5 cm _ thick, fibrous, reddish-brown.

Buttresses narrow, up to 1.3-3 m tall. Innovations with dense

rufous-tomentum. Older branchlets glabrous, lenticellate. Stipules

linear-acute, c. 5-7 by 1 mm, with dense tomentum outside,

glabrescent. Buds subglobose, c. 3-5 by 4-5 mm. Petiole c.

1-3 cm long, terete or flattened on the adaxial side. Leaves

thin-coriaceous, elliptic-lanceolate or elliptic-oblong, c. 5—14 by

1-5 cm; upper surface glabrous, shining, lower surface with

dense rufous-tomentum, glabrescent: base attenuate-acute, usually

asymmetrical, margin remotely serrulate near the acute or sharply

acuminate apex. Midrib and lateral nerves strongly prominent

beneath, flattened or slightly raised above; reticulation fine, paral-

lel, obscure on both surfaces. Lateral nerves c. 5-12 pairs,

parallel, arcuating towards the margin, forming c. 60°-70° angle

with the midrib. Inflorescence not known. Young infructescence

c. 1-2 cm long, carrying c. 1-5 young fruits; bracts linear-acute,

c. 1-2 mm long, with dense tomentum outside. Perianth of the

female flower (seen in the young fruit) 6—9-lobed; staminodes 0:

styles 3-5, c. 2-3 mm long, recurved. Young cupule ovoid-conical,

lamellae c. 3-4. Mature cupule flattened cup-shaped, or patelliform,

c. 0.5-0.7 cm deep, 2—2.5 cm in diameter, covering c. 4-4 part

of the fruit; lamellae thick, c. 5-7; rim denticulate base truncate

or rounded. Mature fruit ovoid-conical or conical-cylindrical, c.

2-3 cm in length and 1.5-2 cm in diameter, densely sericeous,

glabrescent; base truncate or convex, apex rounded — or depressed-

umbonate.

Ecology: in lowland to submontane forests, at c. 30-1300 m

alt. Fruiting in September-April.

Distribution: Malaya (Pahang, Selangor), Sumatra (Agam,

Pajakumbuh, Indragiri), Borneo (Bukit Kalong, Ranau; Mt.

Kinabalu, Tawao, Sarawak, Samarinda vicinity).

maa i ict cml tl mamma ali A LI

‘Soepadmo — Quercus subgen. Cyclobalanopsis 415

Fig. 18: Quercus elmeri

la-b after Singh SAN. 28310: 2a-d after Elmer 2/213; 3 after Anderson

et alia S. 15374.

416 Gardens’ Bulletin, Singapore — XXII (1968)

List of identifications

(The number following the colon corresponds to the number

of species as treated in this paper).

Abbe et alia 9957: 13; Achmat 1094, 1199, 1501: 6; Alvins 1723,

HS. 14675: 15; Ashton Brun. 2378: 14; Backer 25956: 5; Bakhui-

zen v.d. Brink 4451, 6465: 15; Bangham 859: 4; bb. 2865: 2; 3102:

5; 3106: 4; 3126: 15; 3832: 16; 4177: 15; 4640: 16; 4784:

5; 5219: | 2;. 5326, 54715 S624 Sie ee

3882: 193: 6521: 5: (6572: . Ws.0873S. 2. 10389: 415;-F2i0e:

7; 14468: 16; 15952; 6; 16103: 7; 18994: 2; 20387: 15; 20389:

7; 20855: 2; 22382, 24758: 15; 29598, 30134: 7; 31703, 32190,

32254: 2; 33943: 15; 34224, 34346, 34392, 34439: 7; bb. Ja.

3029: 15; P.T.P; ‘740: 7: S.W.K. ell: as; F-o37 ] Fe.

21¥: 15;°604:° 18: Beecari P.B. 1385: he 2243; 2551: B27,

2919: 7; Berkhout 1057: 7; Biinnemeijer 1880: 15; 2048: 2;

Borssum Waalkes 1760: 15; Burkill 2858: 16; Burkill & Holttum

7753: 16; 8661: 19; Clemens 10963: 8; 11211: 2; 30465: 15;

30813: 14; 30948, 30964, 30967, 30976, 31097: 8; 31238:

14; 31264: 15; 31317, 31459: 14; 32364, 32448, 32933: 8;

34492: 12; 34493: 8; 34494: 16; 40231, 40390, 40533, 40534,

40536, 40637, 40699, 40951: 8; 50215: 14; 51254: 15; 51658:

2; Curtis 434: 4; 905: 15; Docters van Leeuwen 11673: 6;

Ellbert.56, 57, 58: 15; Elmer 13219: 11; Z121ISs5N9; Pymiert

3215: 15; 3624: 14; 4511: 15; 4731: 2: FBaS858-V il. Forbes

572: 7; 585, 2719, 2753, 3834a: 2; Fox 14536: 2; Foxworthy

B.S. 566: : 11; Fuchs & Collenette 21665: 1; Grashoff 142, 160,

230: 2; 800, 842: 7; 907, 1101: 15; Hallier f. 2628, 2864:

2; 2915: 18; 2950: 8; 3349: 2; Hamid-5445: 15; Han®@ 231:

15; Haviland 1772: 12; Herb. Sing. 14534: 7; Horsfield 11: 15;

Hotta 14732: 14; Ichlas 102: 15; Jacobs 5114: 2; Junghuhn

9, 63: 16; 70:15; Kadir 63: 7; Kalshoven.10: 15; KEP. 22574,

29828, 31071: 5; 34029, 34031: 15; 38048: 7; 71649: 8; 99590:

2; Koorders 1407, 1408, 1410, 1418, 1435, 1439, 1440, 1468,

1472, 1475: 15; 1487, 1488: 5; 1503, 1526: 15; 1533: 5; 10940,

10941, 10946, 11729, 11928, 11931, 11932, 11933: 15; 12453:

16; 13859, 13955, 14051, 14118: 15; 14184: 5; 14930: 15;

15342:~'5; 23863, "25716, “20502. ‘26070; 207842" 15:-277i Ta;

29204, 29247, 32762: 715;"33299:"-5; 33415; 33737: 15; 33749:

16;. 33762:.. 153.3714), 371460: 16:,38164. 38155. 3e15)-

38484: 5; 38624, 38689, 38695, 38698: 15; 38784: 5; 38801:

15; 39430: 5; 39608: 15; 40101: 16; Korthals HB. 7949: 2;

Kostermans §. 92: 2; 4078, 4180; 7; 4433: 6; 4474, 6368: 7;

6530: 6; 7442: 18; 8963: 11: 20235, 12795:" fF. issn: | is;

Kostermans & Anta 99: 15; 152: 7; 419, 463, 514, 801: 15;

1240, 1324, 1326: 7; Krukoff 4016: 15; Kunstler (King’s col-

lector) 3723, 8258: 4; Labohm 1130: 2; Lands 27204: 16;

Lorzing 11449: 15; 15661: 16; Meijer 3253: 16; 5879: 15;

8687: 5; Meijer &Amiruddin 23: 19; Monterie 51: 16; Nur

Soepadmo — Quercus subgen. Cyclobalanopsis 417

7378;--15; Poore-AS:- 14900: ,.16;:Poore 855: 15; 1315: 5; 1321:

16; 1334: 15; 1348, 1378: 5; Purseglove 4263: 19; Rajab 482,

483: 5; 607: 16; Richards 1885: 10; Ridley 6443: 2; RSNB.

490: 2; 4248: 19; 4363: 8; 4427: 2; 4434: 1; 4451: 13;

4500: 8; 4533: 2; 4534, 4564: 16; 4792: 14; 4893, 4927: 16;

4976: 19; 7002: 14; 7096: 16; 7148: 13; 8411: 2; SAN. 362la:

15; 16188, 16565: 6; 16683, 16811: 4; 19129: 15; 19868: 7;

202197 2220751: 6bd; 20732: . 2; 2097329 13321088: «83; 25342:

19; 24051: 18; 24106, 24121: 8; 24353: 15; 25119: 7; 25336:

135:28216:, J: 28307: , 15; 28310:.,.19; 28808:, 8;,.28919:, 7:

28991: 16; 29060: 8; 29131: 14; 29134: 14; 30869: 16; 31936:

15; 32253: 11; 33131: 16; 33714: 2; 33949: 14; 34504: 8;

34643: 18; 34742: 7; 38069: 16; 38608, 39135: 8; 41832:

16; 42096, 44309, 44478: 8; 48115: 13; 49459, 49749: 15;

51434, 51438: 8; 53861: 15; SAR. 154: 4; 1065, 1070: 12;

2999: 10; 3919, 3946: 3; 4504, 4543, 4547: 14; 4580: 19;

46612. £2:- 75353: “Srei2615202> 12622: . 19; 12624,, 13182; 13315:

2s 13962, Wad65n40- SCs 14: 153743, «15554: ) 4;,,16315:

2s 200d. 20021. 14: 20M ae 2; 2011S: 3;.20121:. 9;,,.20135:

2 20226: 9: 20601, Z080Z:.. 11; ..22462:. 6;,.22650:; 2: Seuveur

4, fae 7; OFW.; 10252: 3: 11798: 15: 18053: ; 16;.20191:. 2;

Coote 3551) 10; 50268, 27006: 8; ZIfS4:. b>; 31253: . 16:

31798: 5; 31973: 15; 35936: 19; 36275: 15; 36398: 2; Soekaria

107: 2; Strugnell 20308: 15; Symington 37664: 7; Teijsmann

HB. 676: 16; 7595: 15: 7638, 7642, 7643: 7: 7649: 2; 3882,

21080: 7; Toha 2057: 2; van Steenis 904: 2; 8264: 16; 8379,

8607: 17; Verhoef 104: 2; van de Vreeden 93: 7; Watson 5817:

15: Winckel 970: 5; Wind 43: 16; Wray 1532: 15.

418 Gardens’ Bulletin, Singapore — XXI1 (1968 }

__ References

. ARMSTRONG, J. M. & A. P. WyLigz in Nature 205: 1340-1341. 1965.

2. BAILLON, H. Nat. Hist. Pl. 6: 220. 1877.

BARNETT, E. C. in Trans. & Proc. Bot. Soc. Edinburgh 33, 3; 327-343.

1942; l.c. 34, 1: 159-204. 1944.

BERRIDGE, E. M. in Ann. Bot. 28: 509-526. 1914.

Brett, D. W. in New Phytol. 63, 1: 96-118. 1964.

BuRKILL, I. H. Dict. econ. Prod. Mal. Pen. 2: 1849-1859. 1935.

CELAKOVSKY, L. in Bot. Centralbl. 30, 1: 10-12. 1887; in Jahrb. Wiss.

Bot. 27: 128-162. 1890; in Oest. bot. Zeitschr. 43: 272. 1893.

8. Corner, E. J. H. in Gard. Bull. S.S. 70: 279-280. 1939; Wayside

Trees of Malaya /: 290-305. 1940.

9. CuTLerR, D. F. in Kew Bull. 77, 3: 401-409. 1964.

10. Daria Torre, C. G. & H. Harms, Gen. Siphon. : 117-119. 1907.

11. DuFrFrecp. J. W. in Am. J. Bot. 27, 9: 787-788. 1940.

12. Etcuuer, A. W. Bliithendiagramme 2: 20-30. 1878.

13. ForRMAN, L. L. in Kew Bull. /7, 3: 381-396. 1964.

14. Guimpu, V. in Rev. Bot. Appl. & Agr. Trop. 9: 176-179. 1929.

15. Hance, H. F. in J. Bot. /2: 240-243. 1874.

16. HANDEL-MazzeEtTTI1, H. Symb. Sinicae 7: 27-52. 1929.

17. HENDERSON, M. R. in Gard. Bull. S.S. 5: 76-78. 1930.

18. HyeLmavist, H. in Bot. Notis., Suppl. 2, 1: 1-171. 1948.

19. Hosseus, C. C. in Beih. Bot. Centralbl. 28, 2: 284-385. 1911.

20. HuTcHINSON, J. Fam. fi. Pl. 7: 192-193. 1959.

21. JARETZKY, R. in Planta /0: 120-136. 1930.

22. JAYNES, R. A. in Forest Sc. 8, 4: 372-377. 1962.

23. LanGpon, L. M. in Bot. Gaz. 107: 301-327. 1939; 108 : 350-371. 1947.

24. Lemee, A. Dict. descr. syn. Gen. Pl. Phan. 4: 127, 765. 1932: 5:

713-714. 1934; 7: 182-183. 1939.

25. LENDNER, A. in Bull. Soc. Bot. Genéve, Sér. II, 8: 161-166. 1916.

26. Meccuior, H. in Engler’s Syll. Pfi.fam. 2: 49-51. 1964.

27. MUuLLER, C. H. in Miscel. Publ. U.S. Dept. Agr. 477: 1-216. 1942;

in Chronica Bot. 7: 12-14. 1942.

28. Onwl, J. Fl. Jap. : 377-378. 1965S.

29. PALIBIN, J. in Bull. Soc. Bot. Genéve, Sér. II, 7 : 359. 1909.

30. PRANTL, K. in Bot. Jahrb. 8: 321-336. 1887; in Engler & Prantl, Nat.

Pfl.fam. 3, 1: 47-58. 1894.

31. ReHDeR, A. in J. Arm. Arb. J: 122-136. 1919. 1.c. 10: 132-139. 1929.

32. ———_—_——_—— & E.. H.. WILSon in Sargent, Pl. Wils. 3, 2: 190-237.

1916.

33. RoxpurGH, W. Pl. Corom. 3: 94. 1819; Fl. Ind. 3: 634, 639, 640. 1832.

34. Sax, H. J. in J. Am. Arb. 11: 220-223, 1930.

35. Suara, T. in Bot. Mag. Tokyo 45; 353-355. 1931.

36. TRELEASE, W. in Mem. Nat. Acad. Sc. 20: 1-255. 1924.

37. TROLL, W. in Ber. Deutsch. Bot. Ges. 44: 290-295. 1926.

38. YAMAZAKI, R. in Jap. J. Genet. /2: 101. 1936.

PS 9 ae

“JT snaianOG snues 94} JO uONNQU\sIp Jo adues ajyewrxoidde :] dep

419

Soepadmo — Quercus subgen. Cyclobalanopsis

420 Gardens’ Bulletin, Singapore — XXII (1968)

AUSTRALIA

MAP E : DisTRiavrion or Quercus L. susgen. CyccosacanoPsis

COERSTED) SCHNEIDER 1th VIALESIA.

A = Quercus PsEvodo-vERTICILLATA £, SOEPADVO

MAP Hf : Disrrraurion oF 14 @ = _n- 3 Wivea KING

a = san s7Teenwisu EZ. So0€PAaH0

eS EE

ap IZ : DisTRiBuTion oF @. ARGENTATA KORTH.

Map X : DISTRIBUTION OF @. GAwARVENSIS E, SoEPMDNO

422 Gardens’ Bulletin, Singapore — XXII (1968)

= Q. o1vocarra Kor7v.

MAP YW: DISTRIBUTION OF

i 2 = Q. PERCORIACEA £.SOEPADYO

HAP Wl : DisTRievriow oF Q. SUMATRAMA E. SOEPADATO

Soepadmo — Quercus subgen. Cyclobalanopsis 423

MAP VT : Disrarauriow of @Q. Suasearcea A. Canus

MAP IX : DISTRIBUTION OF @. Louw KING

424 Gardens’ Bulletin, Singapore — XXII (1968)

Q. curysorarewa A.tartws

Q. MERRILL. SEENEN

A =: Q. Keranvgasensis E, Soepado

MAP XT ¢ DisTRiavT7/0Ow OF 1

@ = @. xmasatvensis Z. So&PADNO

425:

110° Ee,

HOP. ae Sas

. a mee

MNS a we Sew 2

110°,

MAP XL: DISTRIBUTION OF Q. GEMNELLIFLORA Ble

426 Gardens’ Bulletin, Singapore-— XXII (1968)

NoPE. Ln Le sah

10° €,

MaP 2K: Disraravrion oF @. 7TREVEANA v. SEExen/

Soepadmo — Quercus subgen. Cyclobalanopsis 427

Noe. gel

110° €,

Map XV7: DisTR/BUTION OF QD, ELMER (TERR.

Some Studies of Malayan Agarophytic and

Alginophytic Seaweeds

H. M. BURKILL

Botanic Gardens, Singapore

L. H. GREENWOOD-BARTON

Tropical Products Institute, London

and

P. C. CROWTHER

Tropical Products Institute, London

The social, as distinct from the industrial, uses of the Rhodo-

phyceae in Malaya have been comprehensively listed by I. H.

BURKILL (1935) and in Malesia by J. S. ZANEVELD (1959),

but the actual agar content of Malayan agarophytic species has

nowhere been reported. Similarly the same authors have recorded

(loc. cit.) the social uses of the Phaeophyceae, and some of the

traditional industrial uses such as sources of potash, soda and

iodine. The alginic acid content of Malayan alginophytes has not

been recorded. The object of this paper is to put on record some

values for agarophytes in the first part, and for alginophytes in

the second. The field work was undertaken by the first author:

the analyses were done at the Tropical Products Institute,

London.

Part I — Agarophytes

Japan has long been considered, and rightly so, the main

agar-producing country of the world, though shortages of supply

during World War II led the western nations to, search for

alternative domestic sources of supply. Of the agarophytes

exploited in Japan, Gelidium spp. are the most important. This

genus is represented in the Malayan algal flora but it has not

been found in adequate abundance to make collecting practicable.

The genus Gracilaria is well represented, and at many points

around Singapore and the western coast of the Malay Peninsula

there are beds of G. lichenoides, the source of Bengal isinglass.

At certain times of the year this species is thrown up on the

beaches as jetsam in quantity. This, and some collected in the

shallow sublittoral, is used domestically for making agar-agar

sweetmeats: some is collected for feeding to pigs and ducks

(though its food value has not been investigated), but very

little, if any, is collected for industrial use.

430 Gardens’ Bulletin, Singapore — XXII (1968)

Besides, Gracilaria, there are Eucheuma spp. growing on the

littoral and sublittoral fringe, which are used in a similar and

small way, but never in quantity for commercial exploitation.

This situation is in spite of the fact that Singapore handles a

considerable entrepdt tonnage of agarophytic seaweeds (largely

Gracilaria and Eucheuma) from neighbouring countries.

Samples of Gracilaria and Eucheuma from Singapore and the

Johore River estuary have been collected for chemical analysis

together with a sample from a commercial entrepot consignment

from Indonesia for comparison.

Table I gives: the results of these analyses. All three local

samples of Gracilaria lichenoides had agar contents slightly

superior to that of the Indonesian entrepd6t sample. In respect

of gel strength, both Singapore samples produced gels equal to

the commercial sample, though that from the Johore River gave

a somewhat softer gel. The ‘Difco’ agar against which these

were compared is a standard, high quality, hard agar. The results

from the local collections of Gracilaria lichenoides are considered

to be highly satisfactory.

The Johore River station is a few imiles up the tidal estuary

from its mouth in the East Johore Straits. Pulau Tekong and

Teluk Paku both stand to seaward ot this point. The water at

the Johore River station is of reduc2d salinity and differences

of quality may be due to this.

Eucheuma muricatum yielded an agaroid, that is, an agar

extract which would not gel. This quality is already known for

E. muricatum which finds use in Japan as a diluent for high

grade agars.

In extension of this information on Malayan agar resources,

it seems appropriate to record here previously unpublished data

(T.P.I. archives) on some 1940 analyses made by the Imperial

Institute, London. These analyses were made in the general

search for sources of agar other than from Japan at the beginning

of World War II. Samples were submitted from Penang, Malacca,

Pulau Besar and Pulau Upeh off the Malacca coast, and Singapore.

Samples from Indonesia and Sout!: India were at the same

time examined. All were labelled as Gracilaria lichenoides except

that from Penang which was stated to be Gracilaria sp. The

results are given in Table II.

The hot water soluble matter in these tests compared favourably

with solubilities from different species used in Japan for the

preparation of commercial agar, and they will be seen to be

similar to those recorded in Table I.

Gelation tests, comparing these samples with commercial

Japanese agar, showed the Indian sample to be superior. All

the others except that from Pulau Upeh were good or fair and

were deemed to be promising as sources of commercial agar.

431

‘[O1]UOD & SB posn UOIeIJUBDUOS

OWeS OY} JO IvBe OIG, JO UONNIOS eB JO OSRUDdIOd B sv PossoJdx9d IvSe P3}dRI}X9 JO (SISBq JO}}eLU AIP) Uonnyos jusdJed ¢/"9 JO YISUoIs [BH I

"JORI]X9 I[QNOS 19} P]OS sso] 3OBI}X9 BIQNTOS JOIVM JOH{ y

| |

1Z ras 9I gue. iz & - “4 jouuod Jo yuoosod se i yIsuos [9H

5 0°67 T'6€ V8¢ SIP & ~¥ — "7 a » }U9}U09 IvBWy

6°0L 1°88 SIS 6EP 9°89 « 4 ¥ a ge ae Oe eee

OVE 16S Cun 8°¢ 8°97 = = "* 4OB1]X9 a[QnjOs Ja}BM PlOD

9°8b tir SPI Az QLE ee oe oe ee oe ot a usV

i s10desuig JOALY B10 YOF o10desulsg ds1odesuis Ayeo07]

nuey nejng Suoyoy neing | nye NAL,

[eioxewU09

soplousyoij win} eoTiNuT | soprousyary soplousyoly soprousyol| so1oadg

BIIR]INVID eunoyong | PIIE]IOVID BLILPIOVIDH BlIRIOVID

| (yISUaTs [OD 1d29x9) JUBIOMAIP JO jUSdIOd sv passoidxe soinsi{

SHLAHdOUVOV TVOOT AWOS AO SASATVNV

| ATaVL

ie]

iS)

ie)

mae

oa)

432 Gardens’ Bulletin, Singapore — XXII (1968)

TABLE II

Analyses of some local agarophytes

(Bull. Imp. Inst. 1941, 39, 16 and hitherto unpublished results).

Hot water soluble | Soluble chlorides

Origin extract as NaCl

Penang id t bil 67.1 4.9

Malacca, mainland .. a 59.6 3.8

Malacca, P. Besar im ae 64.1 3.8

Malacca, P. Upeh at Pe 51.4 3.9

Singapore .. i “ 46.8 0.2

Indonesia .. a 4: 42.6 | 0.2

India ae =” wa 57.3 6.3

Penang sample=Gracilaria sp. All other samples= Gracilaria lichenoides.

Part 2 — Alginophytes

The presence of algin is a feature of the larger brown seaweeds,

but the most important genera which meet the world’s demand

for alginic acid are not represented in the Malayan algal flora:

Macrocystis, Laminaria, Ascophyllum, Eisenia, Ecklonia and

others occur in more temperate regions. Newton (1951) records

the use of Sargassum ringgoldianum in Japan, and Hoppe and

Schmidt (1962) refer to the presence of Sargassum species in

the Singapore algal flora without mentioning any specifically.

These authors quote alginic acid contents for S. myriocystum,

22 percent, S. plagiophyllum, 27 percent, and S. wightii, 25 per-

cent, on the Ceylon and Indian coast. The genus Sargassum

from the rather fragmentary collecting that has so far been

done, is clearly well represented in Malayan waters (see Grunow,

1915-16). When further systematic collecting and taxonomic

research has been undertaken, this vast genus will surely be

found to be much richer here than is at present known. Cystoseira

and Turbinaria of the family Sargassaceae also occur around

Malaya.

The Sargassums at Raffles Light

Around Singapore, at least, and probably at many points

along the western Malayan seaboard, the Sargassum beds are a

conspicuous feature of the algal ecosystem of the littoral and

immediate sublittoral coral reefs. For the present exercise, the

beds at Raffles Light (Pulau Satumu) were chosen because of

ease of access and because of facilities available in the marine

laboratory of the University of Singapore Zoology Department

situated there.

Burkill et al. — Malayan agaro- and alginophytes 433

The island lies at the western limit of the Straits of Singapore

at the southern end of the Straits of Malacca. It is of lateritic

rock, barely one acre in extent. It has an intertidal littoral fringe

of coral, rocks and small sandy pools covering about eight acres.

The spring tide range is about 10 feet. The upper limit of the

sargassum is about the midlittoral. The lower limit is at about

two feet below low water spring tide level, which also marks

the edge of the reef where it plunges steeply into deep water.

Some studies of the littoral ecology of this island have been

published by Purchon and Enoch (1954). They record only the

occurrence of Sargassum siliquosum J. Ag. and Turbinaria ornata

J. Ag. of the Sargassaceae, but there is plainly a greater representa-

tion than these two, especially of the genus Sargassum.

Observations recorded in this paper were made in 1960/61 on

Sargassum glaucescens, S. swartzii and S. assimile, and in 1962/63

on S. glaucescens, S. myriocystum and S. swartzii. Vouchers for

all the assays are laid in the Singapore Herbarium. These four

species occupy marked niches on the coral shelf, the most

obvious characteristic of their positions being the degree of tidal

inundation and of exposure of the tidal flow. Tidal predictions

(H.M.S.O. 1962) record peak water movement of over 34 knots.

S. swartzii is predominant on those rocks most exposed to this

movement, and hence it is mainly at the lowest littoral level

though it does occur scattered here and there with S. glaucescens

and S. assimile. These two latter species are freely intermixed with

one another and cover the whole of the lower littoral coral shelf

subject to tidal flow. S. myriocystum occurs only in sheltered

water. It tends to be silt covered and is mainly at mid-tide

level and is therefore inundated the least of the four species.

S. siliquosum, referred to above was not sampled because of its

inabundance relative to these other species. It occurs on the lower

littoral in a small sheltered part.

Growth Cycle

The species are deciduous: S. glaucescens, S. swartzii and

S. assimile completely shed their secondary branches towards

the end of the northeast monsoon period which is approximately

November to March. In 1961 and 1963, these species were

entirely broken down to stubble — the holdfast and brief primary

stem, a few basal leaves and a few incipient secondary branches

— by March. In 1962 breakage was complete by February.

S. myriocystum maintains some secondary branches the year

round, though the old fertile branches are eventually shed. It

appears that the secondary branches of this species, unlike the

branches of the other three, may last longer than one annual

cycle. |

This timing is not necessarily the same elsewhere, and it must

be considered to be a characteristic of the locality. For instance,

S. glaucescens is common on the lower littoral of the Labrador

434 Gardens’ Bulletin, Singapore — XXII (1968)

beach by the western entrance to Singapore docks, and some

10 miles distant. On this beach it is protected both from the

northeast monsoon and also from extreme tidal flow. The mature

growth of this species persists for much longer than it does at

Raffles Light so that when it eventually does break off, the new

secondary branches have grown appreciably and the rocks do not

assume the same bare fallow appearance.

Regeneration at Raffles Light is slow at first and the reef is

bare for some months except in those areas occupied by

S. myriocystem. The sargassum of the lowest level grows first and

most quickly, regeneration spreading upwards so that by July

the reef is clothed again in its mantle. Mature stature is attained

by August-September, and the plants become fertile soon after,

there being some variation between species. Observation on these

points are given in Table III.

TABLE III

Sargassum Growth Cycie at Raffles Light at approximately the

mid-lower littoral level

(a) average length of plants in inches.

(b) state of fertility: s = sterile; f = slightly fertile; ff = moderately

fertile; fff = very fertile.

Species | S. glaucescens |S. myriocystum| SS. swartzii S. assimile

Dae | @ | ® |, @ |®| @ |O)| @ | &

| | | |

20-3-60 x Seria «ee 3 s ‘

27-6-60 ea | 6 s 4

6-9-60 | 15 S | aif | 24 ff

8-10-60 15 ri4 | 24 fff

5-11-60 18 ff;s;4 | as -

21-12-60 24 ff | 36 fff

28-1-61 co fea ee ee eee rr? oe fff

3-3-61 c* duws (ok pomianhees fiikina ss vG

| Persis-

| ting | |

18-5-62 6 5934 9 Ss 6 s

10-6-62 9 s 12 S a oe

3-762 12 s 12 s 4 S

1-8-62 18 s 15 S 18 f

3-9-62 24 S 15 s 18 f

1-10-62 24 S e i Wee 24 f of

1-11-62 24 f salad aa 24 f CL” PETE

30-11-62 | 24 ff 24 f 24 is! eae

28-12-62 24 fff 24 f 24 T. ef “

2-2-63 ig*> | ser) ee PORE 12* f 5 #

25-2-63 | ..T ee sg | fff | T z. POV -

Persis- /

ting | | |

| |

; *End of season breakage occurring. End of season breakage completed.

In the areas dominated by S. glaucescens and S. assimile there-

is an underlayer of Cystoseira prolifera which is exposed when

the sargassum shed its branches, and this too breaks up soon

after.

Burkill et al. — Malayan agaro- and alginophytes 435.

Chemical analysis

Sampling of the material for chemical analysis was carried out

during the growth cycles of 1960/61 and 1962/63, the secondary

branches being cut from the primary stem, which with the

holdfast remained in situ. The harvested material was rinsed in

fresh water, oven-dried and despatched to the Tropical Products.

Institute, London, where the analyses were made.

The initial samplings of March to June 1960 were made on

plants little more than stubble, and the species concerned were

not separated, but they were mainly S. glaucescens and S. assimile

mixed. Subsequent samplings were made on _ these species.

separately. The samplings of 1962/63 were carried out on

S. glaucescens, S. myriocystum and S. swartzii separately.

Alginic acid and mineral ash contents on a percentage basis of

dry matter are shown in histogram form in Figures I and II, and

the annual results averaged out over species are at Table IV.

Alginic acid contents cbserved rank favourably with the contents.

of other species from elsewhere which are exploited commercially.

TABLE IV

AVERAGE ALGINIC ACID AND ASH CONTENTS OF

SARGASSUM SPP.

ALGINIC ACID AS PERCENT ASH AS PERCENT OF

OF DRY WEIGHT DRY WEIGHT

Species —| ——--- a —

1960-61 1962-63 | 1960-61 | 1962-63

Sept.—Jan. _ May—Feb. | Sept.—Jan. | May—Feb.

S. glaucescens 4 23.0 25.6 26.2 23.6

S. myriocystum a iy 20.9 A 30.2

S. swartzii 0 as 24.6 2. 21.5

S. assimile * 23.7 . 26.6

Between species, in respect of alginic acid content S. myriocystum

at 20.9 percent is markedly inferior to S. glaucescens at 25.6.

percent and to S. swartzii at 24.6 percent. It is probably also

inferior to S. assimile at 23.7 percent recorded, however, in a

different, though probably a poorer alginic acid producing year.

(See below.) There appears to be little difference in alginic acid

content between S. glaucescens, S. swartzii and S. assimile.

Mineral ash contents follow a similar, but converse pattern.

These are looked at in more detail below.

=~,

—

nal

ina]

“we

Sy)

436

19/1/62

09/2t/te

09/tt/S

09/9T/8

09/6/9

19/1/62

09/2t/tz

09/Tt/S

09/ot/8

09/6/9

09/9/Lz

09/$/62

09/¢/z

09/€/9z

“Date of sampling

o*€z

2*te

€*22

€°S2

S*ve

T*0z

9°t2

Tbe

6°G2

6°22

1°62

T'te

L°z2

S*Lt

“Value

RSS IS WZLLLLELELLALLLLLULLLALE.

NSS MM|/“/AZXZ_LLLLLLLz™_

WLM MI

MMA YYHWUWW//// U4:

NGG SABA MALL lle Ll

WWW WCW lili,

RSW SB zaauun{qxaazZ

MO QA WW

SSS —DMM{@MKM{u“ramma@e&EXHzzL#“d

MOM AIAN l/l l/l,

CLLLLLLLLLLLD

KS CQL,

RQ Wii sisllulblditth,

PyOY

NSS GM AA)UMMXZ: tt LLL

& P) 3 mn wo 2, 2 8

oTUTaTY

Wey

“ ”

Ter0UTy

L°6T

z°ve

0°0€

S*92

~value

glaucescens assinile

glaucescens —

assimile

mixed

-Specieas

Fig. I. Alginic Acid and Mineral Ash contents of Sargassum species at

Raffles Light, 1960/61 ( as percentage of dry weight).

f9/e/z

t9/et/oe

29/11/ot

9/t1/t

z9/o1/t

29/6/6

29/0/1

29/L/t

29/9/01

29/¢/at

t9/2/se

t/z/e

29/e1/ee

z9/Tt/oe

29/1t/t

29/9t/t

29/6/¢

29/9/1

29/L/¢

29/9/ot

29/6/et

t9/e/e

29/et/ee

29/tt/ot

29/tt/i

z9/ot/t

29/6/€

29/8/t

29/L/€

29/9/ot

z9/Sfat

- sampling

Date of

S62

L've

Vor

ste

6rte

orte

cree

6°6t

6°02

S*at

e*ul

orle

ree

ore

e*be

ore

Lelt

grtz

g°se

6°be

prse

s*te

vee

v°6z

9°0e

v°62

beet

L*zz

Value

| SENET SWI

MSSSASSS W/L

ESO MM May,

SIMOSMQ yyw

SSSA ZAZA ZA

EX X SRO IMs VLE.

KOSS ASS ZZ

SSS UMMM

SSM ILL,

INNS ALLL

MINS

SSNS WLLL TTI

WSS) LM TU

I SIAR L/L

SWS YL J

SSIS. TULL

SNOT ZI

ISSSSSISSIS ITU

RSS ILL ULL

WWII WLLL

— — ater ml

Sen

Gilly

ne =a WML LULL

SVL

SWIMM

ONS LUMA LLL

GW ”"”0°? DELLA

WKY MMM LLL

QQ (/ GW)

a : a 8 . 3 8 & 8

$ *

prey opurity qoy porous

Got

g*0e

ar92

orte

9°6t

ore

€*6

vor

ole

Grre

Gree

eee

Lee

e°re

pee

g*lz

9°82

proe

G*re

ore

gree

29%

S*se

eete

e*ot

o*6t

Brot

eee

tet

Sr2e

S*be

Value

Fyriccystam

inic Acid and Mineral Ash contents of Sargassum species at

Alg

Raffles Light

glaucescens

Species

, 1962/63 (as percentage of dry weight).

Fig. II.

Burkill et al. — Malayan agaro- and alginophytes 437

DATA.

Percentage dry weight

Ash Alginic acid

Sargassum indet. Raffles Le

S.glaucesceas Raffles Le

S. nyriocystum Raffles Le

Pe Undan

P. Nangka

S. swartzii Raffles L.

S. assimile Raffles L.

C. prolifera P. Hantu

Raffles L.

Turbinaria spe P. Hantu

Padina spe P. Hantu

Fig. III. Relationship of Ash Content to Alginic acid content for

species, localities and years

Ash as percentage of dry weight

| 10 20 30

: Alginic Acid as percentage of dry weights.

438 Gardens’ Bulletin, Singapore — XXII (1968 )

Variation between years in alginic acid content and ash content

has been reported by Lunde (Chapman, 1950) in Laminaria

digitata. Such variation is demonstrated here in respect of

S. glaucescens which was recorded in both seasons, the 1962/3

season being better for alginic acid production, with a lower

mineral ash content, than the 1960/61 season. Indeed, some

annual variation in any harvested crop is a natural expectation.

Lunde for L. digitata and Black for L. cloustonii (Chapman

loc. cit.) both recorded an annual period of peak for alginic acid

content. Such a situation is not apparent in either of the two

years over which the sargassum was examined. While there has

been monthly variation, as is shown in Figures I and Il, the

variation has been irregular. Similar but converse irregularities

are also shown for the mineral ash content. The causes of these

abrupt changes are not understood, but that there is some relation-

ship between the alginic acid and mineral ash contents is shown

below. The irregularities however smooth out over the months

and there is no apparent peak period.

TABLE V

| PERCENTAGE OF

; DRY WEIGHT

Species Locality Date

Ash Alginic acid

Sargassum myriocystum | Pulau Undan 24-61 28.8 | 25.4

Malacca

Pulau Nangka 30-4-61 32.4 19.7

Malacca

‘Sargassum swarizii .. | Raffles Light 28-1-61 32.8 22.9

Singapore

‘Sargassum ‘E’ .. | Pulau Nangka 30-4-61 27.1 |

Malacca

-Cystoseira prolifera .. | Pulau Hantu 14-11-59 59.0 9.5

Sir gapore

Raffles Light 3-3-61 21.1 24.6

Singapore

Padina sp. .. | Pulau Hantu 14-11-59 65.2 5.0

Singapore

Turbinaria sp. .. | Pulau Hantu 14-11-59 45.6 12.9

Singapore

Burkili et al. — Malayan agaro- and alginophytes 439

A few other miscellaneous samples of various brown seaweeds

have also been analysed, and these are recorded in Table V.

The sargassum analyses are similar to those of the main part of

this record. §. myriocystum at Pulau Undan, in the Straits of

Malacca and 100 miles distant, ranks highest in alginic acid

content of the observations for this species, though the record

from Pulau Nangka, some four miles off from Pulau Undan,

is only identical with the poorer ones from Raffles Light. Of

particular interest is the record of Cystoseira prolifera taken at

Raffles Light on 3rd March, 1961, which ranks equal in alginic

acid with the best of the sargassums. This species is nowhere

abundant enough to be a major element in the algal flora, and

so would not contribute greatly in any exploitation, but the great

disparity between the alginic acid content of this sample and of

C. prolifera from Pulau Hantu, a Singapore inshore island, is one

of many pointers to the effect of locality (perhaps salinity) that

merits investigation. Black, Richardson and Walker (1959) record

variation of Scottish samples of Laminaria cloustonii within a

_ location and between locations of differing salinity, which is in

line with our findings. Their observation on the variability between

individuals is based on plants of large size, whereas our material

being relatively small and slender did not lend itself to individual

assessment. Variation is a reasonable expectation, but it is not

possible to express an opinion whether our minimum sample of

20 plants (more often a sample contained about 30 plants) was

adequate to smooth out sampling errors. Black, et al. required

120 plants of this very bulky material which varied in age between

4 and 10 years. Our material was all under one year old. It is a

moot point how far we may draw comparisons and inferences

of this sort on materials so differing in morphology, taxonomic

status, and environment.

Weight of wet sargassum in relation to yield of alginic acid.

Yield per unit area has not been observed. It would anyhow

vary between localities, between tide levels in the littoral and

sublittoral, and between species harvested. The weight of wet

sargassum to yield of alginic acid is however an important practical

consideration in harvesting. Some observations on this were taken

from the first one dozen samples of the 1960/61 season and are

summarised in Table VI. The fresh samples were weighed wet

but drip-free, i.e., when all the free surface water had drained

off the sample spread on a wire mesh frame. Related material

dry weights and alginic acid percentages on a dry weight basis

gave data for a calculation of alginic acid on a wet weight basis.

The sargassum at harvesting contained between 8.4 and 11.6

per cent of dry matter and between 1.9 and 2.6 per cent of alginic

acid. There is a suggestion that as the plant grows from ‘stubble’

to maturity there is a decline in dry weight percentage.

440 Gardens’ Bulletin, Singapore — XXII (1968)

TABLE VI

YIELD OF DRY MATTER AND ALGINIC ACID IN RELATION

TO SARGASSUM WET WEIGHTS AT SAMPLING

" Dry matter as | Alginic acid as

Sargassum Species Date —| » of wet weight % of wet weight

28-3-60 11.2 2.0

Mixed (mainly S. glaucescens 2-5-60 11.5 2.6

and S. assimtiule) “

29-5-€0 19.3 2.2

| | 27-6-60 8.3 2.4

6-9-60 8.7 2.0

8-10-60 10.1 / 2.6

S. glaucescens

21-12-60 | 8.8 | 2.1

6-9-60 | 10.6 2.6

8-10-60 | 9.8 / 2.5

S. assimile | ;

| 5-11-60 | 9.3 / 2.1

4 5-11-60 BP i o7! 2.2

21-12-60 | + ay 1.9

Relationship between alginic acid content and ash content

Our records very plainly show a strong negative correlation

between alginic acid and ash contents: when the alginic acid

content is high the ash content is low, and vice versa. Monthly

variations also show this, but what is also noteworthy is that

different genera fall into this pattern. It the values for alginic

acid and ash content contained in Figures I and II and Tables IV

and V are averaged for species and localities and the averages are

plotted against one another, a near straight line regression is

obtained, as is shown in Figure III, for ash contents between

20 and 65 per cent and for alginic acid contents between 5 and 27

per cent. We can offer no explanation for the significance of this

fact, but it is one calling for further investigation. It may be that

this is a mechanism, as Black, et al. (loc. cit.) suggest for a similar

relationship between mannitol and ash in L. cloustonii for the

maintenance of a constant osmotic pressure within the plant.

Possibilities of exploitation

Any plans for possible utilisation must be based on a sound

policy of conservation which permits regeneration. Many aspects

other than the taxonomic need to be studied, such as the ecology

and autecology, and more especially the phenology of the species

Burkill et al. — Malayan agaro- and alginophytes 44]

to be harvested is all-important. The findings presented in this

paper are but an introduction to a large field awaiting investigation

before any exploitation is attempted. Inferences however may be

drawn in respect of the species harvested at Raffles Light. The

information in Table II indicates the need to limit harvesting of

S. glaucescens, S. swartzii and S. assimile to between November

and January when the plants attain a state of full maturity. Since

the alginic acid content analyses give no apparent optimum over

the growth cycle, this may be the only criterion. However since

periodicity between localities varies, each locality will need a

separate assessment. Any harvesting of S. myriocystum at Raffles

Light would have to be deferred till February on account of later

maturing.

Acknowledgements

We gratefully acknowledge the help of the following persons:

Dr. George Papenfuss, University of California for determination

of the four sargassa comprising the main part of the alginic acid

investigation: Captain J. A. L. Pavitt, 0.B.£., Master-Attendant,

Singapore, for facilities at Raffles Lighthouse; succeeding

Professors of Zoology, University of Singapore for the use of the

marine laboratory at Raffles Lighthouse; Che Kadim bin Tassim,

Plant Collector, Singapore Botanic Gardens, who undertook much

of the sampling in 1962; and Mr. T. P. A. Mellon, Scientific

Assistant, Tropical Products Institute, for help with the analyses.

We also record receiving helpful comment from Mr. E. Booth

of the Scottish Seaweed Institute on this paper while in preparation.

References

Back, W. A. P., W. D. RICHARDSON & F. T. WALKER: 1959. Chemical

and growth gradients of Laminaria cloustonii Edm. (L.

hyperborea Fosl.). Econ. Proc. Roy. Dublin Soc. 4(8).

BurKILL, I. H.: 1935. A Dictionary of ihe Economic Products of the

Malay Peninsula. Oxford University Press.

‘CHAPMAN, V. J.: 1950. Seaweeds and their Uses. Methuen & Co. Ltd.

London.

GrRUNOW, A.: 1915-16. Verhandl. Zool. Bot. Gesellschaft, Wien, 65

and 66.

Hoppe, H. A. and O. J. ScHmipT: 1962. Meeresalgen als moderne

Industrieprodukte Botanica Marina 3, Supplement.

H. M. S. O.: sm Tide Tables for Malaya, North Borneo and. Sarawak,

1963.

NEwrTon, L.: 1951. Seaweed Utilisation. Sampson Low, London.

PURCHON, R.D. and J. ENocH 1954. Zonation of the Marine Fauna and

Flora on a rocky shore near Singapore. Bull, Raffles Mus.

Sing. 25, 47-65. ,

ZANEVELD, J. S.: 1959. The utilisation of marine algae in tropical S

and SE Asia. Econ. Bot. 13, 89

Plate I. Drift of Gracilaria spp. at Pasir Ris, Singapore, December 1964,

on the East Johore Straits, and in close proximity to Teluk Paku,

Pulau Tekong and Johore River — see Table I.

Plate II. Drift of Gracilaria spp. at Pasir Ris. December 1964.

Plate III. Sargassum stubble at Raffles Light, April 1963.

Plate IV. Sargassum in young growth at Raffles Light, June 1963. Reef

at low water.

Plate V. Sargassum in young growth at Raffles Light, June 1963.

Plate VI. Sargassum at mature growth at Raffles Light, December 1960.

myricoystum J.G. Ag. and S.

glaucescens J.G. Ag. at Raffles Light, December 1960.

Plate VII. Mature growth of mixed S.

New Species of Dipterocarpaceae, Sterculiaceae

and Monimiaceae, and a re-interpretation of

Tetranthera rumphii BI.

A. J. G. H. KOSTERMANS

Herbarium Bogoriense

DIPTEROCARPACEAE

Hopea celtidifolia Kosterm., spec. nov.

Arbor ramulis minutissime dense adpresse stellato pilosis, foliis

chartaceis vel rigide chartaceis glabris ovato-ellipticis acuminatis

basi in petiolem gracilem contractis supra nitida laevia nervo

mediano prominulis costis vix prominulis subtus sublaevia nervo

mediano prominentibus nervis lateralis inferiore parte erecto-

-patentibus superiore parte subito verticalibus totis nervum basalem

‘costae parallelum quasi nervaturam palmatum aemulantibus nux

elongato ovoideus acutis alae majoribus oblanceolatis obtusis basim

versus sensim attenuatis glabris alae minoribus elongato ovatis

acutis.

Typus: Versteegh BW 4854 (L).

Tree, up to 28 m high, free bole 24 m, diam. 43 cm; buttresses

up to 2 m high, out | m; bark brown, not fissured, rather strongly

peeling; living bark brown or lightbrown; sapwood yellow or

yellowish brown; heartwood yellowish brown or yellow. Branches

smooth, glabrous; branchlets slender, densely, minutely adpressed

stellate pilose. Leaves chartaceous to thickly chartaceous, glabrous,

ovate-elliptic, 2.5 x 6 — 4 x 10 cm, abruptly acuminate, acumen

obtuse, up to 5 mm long, base contracted into the slender, up to

1 cm long, above slightly, narrowly canaliculate petiole; upper

surface smooth, midrib prominulous, ribs faint, prominulous;

lower surface rather smooth, midrib prominent, lateral nerves ca

5 pairs, slender, erect-patent, rather straight, at about halfway the

margin abruptly perpendicular and anastomosing with the former

and next one, forming two pseudo-basal nerves, that reach up to

the leaf tip, at the outside some more arcuate, faint nerves;

secondary nerves dense, parallel, faint. Inflorescence unknown. Nut

glabrous, elongate-ovoid, acute, up to 1 cm long. The two longest

wings oblanceolate, glabrous, obtuse, up to 4 cm long, the widest

part 1 cm, 7-9 ribbed, reticulate, gradually narrowed at base; the

short wings half, respectively 4 the nut length, acute.

W. IRIAN: S.E. part, E. Digul, Iwur R., alt. 400 m, Agathis

forest on clayey soil, ster. BW 8533 (L); along Digul R. near

Wage, alt 15 m, Nov., fr., Versteegh BW 4854 (L); Muju Distr..

Opka, 10 km N.E. of Ninati, alt. 50 m, BW 6441 (L); 5 km N. of

Ninati, alt. 50 m, ster.. BW 6430 (L).

444 Gardens’ Bulletin, Singapore — XXII (1968)

STERCULIACEAE

Scaphium burkillfilii Kosterm., spec. nov.

Arbor magna ramulis crassis glabris lenticellatis foliis alternanti-

bus glabris coriaceis ellipticis acuminatis basi rotundatis supra

nitida laevia nerviis primariis gracilibus prominulis subtus opaca

dense prominule reticulata nervo mediano prominentibus costis

prominentibus basalibus 2 vel 4, majoribus strictis usque ad 4

foliorum lamina adscendentibus, minoribus arcuatis submarginalis

costis caeteribus utrinque 3-4 suberectis curvatis marginem versus

arcuatim evanescentibus; petiolis perlongis glabris; infructescentiis

magnis vix ramosis perdense minutissime stellato pilosis fructus

coriaceis Scaphiformibus acutis extus laxe minutissime stellato

pilosis, intus pernitidis griseis glabrescentibus semen globosum

magnum.

Typus: Ilias Paie S 15575 (SING).

Large tree, 50 cm in diam.; branchlets cylindrical, stout, glabrous,

glossy, grey, with tiny lenticels. Leaves spirally arranged, coriaceous,

glabrous, elliptical, 11 x 24 — 12 x 37 cm, apex shortly acu-

minate, base rounded; upper surface glossy, smooth, main nerves

filiformous, prominulous; lower surface dull, densely, promi-

nulously reticulate, midrib prominent, basal nerves prominent, 2

pairs, the lower pair almost marginal, reaching 1/5 the blade

length, the inner ones straight, reaching % of the leaf length; lateral

nerves 3-4 pairs, towards margin running out arcuately. Petiole

16-21 cm long, glabrous, thickened at both ends, glabrous.

Infructencence hardly branched, up to 25 cm long, densely,

minutely rusty stellate pilose; fruit boat-shaped, acute, up to 7 cm

long and (opened) 2 cm wide, leathery; outside sparsely, minutely

stellate hairy, inside glossy, grey, glabrescent; seed globose, 3 cm

diam.; seedcoat 4 mm thick; peduncle thick, densely pilose, 5-7

mm |.

Outstanding by its leafshape, the thick, leathery capsules and

the enormous thick-walled seed. As the tree was collected in a

swampy area, one is tempted to ascribe the leathery fruit and the

thick-walled seed to the conditions of water dispersal, as opposed

to the other species of Scaphium where the paperthin capsules and

light seed are wind dispersed.

The species is named after Mr. Humphrey M. Burkill, Director

of the Botanic Garden, Singapore, under whose directorship the

library and herbarium facilities have been greatly extended and

improved and whose valuable cooperation to help the Herbarium

Bogoriense in the difficult period of 1957 -— 1967 is herewith

gratefully acknowledged.

SARAWAK: Bintulu Distr., Segan For. Res., swamp forest;

fruit chocolate coloured, Nov., fr., Ilias Paie S 15575 (SING) (also

distributed to K, L, SAN).

Sen din

: NS ~

PF Sontrbien bun Lill la

Eativm, a

{TUR OY SARAWAR

Plate I. Holo-type sheet of Scaphium burkillfilii Kosterm. deposited in the

Singapore Herbarium, Accession No. 518135.

Kostermans — New species 445

MONIMIACEAE

Steganthera suberoso-alata Kosterm., spec. nov.

Arbor parva vel mediocris ramis suberoso-alatis griseo albis

ramulis minutissime laxe adpresse pilosis vel sublanuginosis foliis

chartaceis ellipticis vel ovato-ellipticis obscure longe acuminatis

basi rotundatis supra glabra nervo mediano prominulo costis

filiformibus subimpressis subtus pallidiora sparse minutissime (sub

lente) sublanuginosa vel adpresso pilosis nervo mediano prominen-

tibus costis utrinque 7-8 prominentibus arcuatis marginem versus

arcuatim conjunctis rete laxe prominule petiolis incrassatis brevis

dense minutissime adpresse pilosis vel sublanuginosis. Inflorescentiis

rami vel caulifloris aggregatis dense minutissime griseo sublanu-

ginosis alabastris semi-globosis adpresse pilosis pedicellis longis

gracilis.

Typus: Hunt 2513 (K).

Tree or shrublike tree, 5-15 m high; branches with conspicuous

_ grey-white corky wings. Branchlets minutely, rather sparsely

(denser towards apex) sublanuginose or adpressed pilose. Leaves

chartaceis, elliptical or ovate-elliptical, 4 x 18 — 8 x 27 cm,

gradually acuminate, base rounded; upper surface glabrous, midrib

prominulous, ribs and secondary veins slightly impressed, lower

surface paler with sparse adpressed or subadpressed, tiny hairs

(denser on nerves), midrib prominent, lateral nerves 7-8 pairs,

prominent, arcuate, near the margin arcuately anastomosing; reti-

culation lax, prominulous. Petioles swollen, densely, minutely

sublanuginose, up to 5 (— 8) mm long. Inflorescences on old wood

on accrescences or on the branches, fasciculate, up to 6 cm long,

minutely, grey sublanuginose; flower buds semiglobose, sparsely,

minutely adpressed pilose.

The species is characterized by the corky wings. There is a slight

discrepancy in the tomentum, the type specimen has bent hairs

(sublanuginose), in the other specimens the hairs on the branchlets

and leaves are straight and adpressed. Vernac. name: U-uni-alakau.

SOLOMON ISLANDS: Kolombangara, alt. 800 m, open rain-

forest with much moss, Sept., buds, Hunt 2513 (K); Guadalcanal,

E. slopes of Mt. Gallego, July, fr., Dennis 2142 (K), tree 10 m,

carpel ellipsoid, smooth, 9 x 13 mm, stalk 2 x 3 mm. like the

receptacle minutely, adpressed pilose; ibid., Popomanasiu, Vunvu-

lukama, alt. 1500 m, Oct., fr., Corner 151; fr. with yellow-ochre

carpel stalks and receptacle; E. San Cristobal, 8-11 miles inland,

alt. 500 m, Warahito R., July, fr., Whitmore 6229 (K), peduncles

swollen, bright orange, receptacle 3 cm diam.; Santa Ysabel,

Cockatoo Anchorage, forest on ultrabasic rock near sea, Sept.,

fl. Hunt 2617 (K).

446 Gardens’ Bulletin, Singapore — XXII (1968)

MYRISTICACEAE

Myristica rumphii (Bl.) Kosterm., comb. nov.

Tetranthera rumphii Blume (basionym), Mus. bot. Lugd. bat.

1 (24): 382. 1851; cf. Kostermans, Bibl. Laur. 1424 ro 203. 1964;

typus: sine coll., Moluccas (L); syn.: Litsea rumphii (Bl.) Vill.;

cf. Kostermans, lc. 874 no 402.

The species is most likely conspecific with Myristica hollrungit

Warb.

On some species of Kuglena Ehr. from Singapore

(De Species Eugienae EKhr. ex Singapore.)

B. V. SKVORTZOV

Botanical Institute of Sao Paulo, Brazil

Abstract

Six forms of Euglena Ehr., collected by the author in the Lake of

the Botanic Gardens in 1962, are described. They are (1) E. downiae sp.

nov., named in honour of Mrs. V. V. Down a British resident of

Singapore and an old friend of the author’s family, (2) E. prowsei sp.

nov., named in honour of Dr. G. A. Prowse, Director of the Freshwater

Fish Culture Research Institute, Malacca, (3) E. proxima Dang., var.

tropica, var. nov., (4) E. clara Skuja var. singaporensis var. nov., and

(5) E. variabilis Klebs.

On 25th July 1962 the author arrived in Singapore on a journey

from China to Brazil, and, thanks to the kindness of Mrs. V.V.

Down, he was enabled to visit the Singapore Botanic Gardens.

and her own private garden where material was collected for his

future studies. Samples of water plants and pond mud were taken

for cultivation of algae by addition of pepton at the Cryptogamic

Laboratory of the Botanic Institute, Sao Paulo, Brazil. The culture

was found to contain the material described below.

The holotype of Euglena downiae, E. prowsei, E. proxima var.

tropica, and E. clara var. singaporensis are preserved at the

Cryptogamic Laboratory of the Botanical Institute of Sao Paulo,

Brazil (S.P.).

}. Euglena downiae sp. nov. Plate I, figs. 1-4.

Differt a Euglena proxima Dang. (vide Popova, 1955, p. 143,

figs. 50, 1-3) periplasto non striato, granulis paramylaceis, non

annulatis, ecoloratis, flagellis, fere 4-4 cellulae aequilongis.

Cellula fusiformis, metabolica, anteriore plus minusve attenuata

oblique obtusa, posteriore recta vel obliqua, breviter acuta,

18-30-47-55-63 x 12-15-25 u. Periplastus non striatus. Flagellum

breve 4 cellulae usque aequilongum. Stigma rubrum oblongum.

‘Gula elongata vel ovalis. Nucleus medianus, vel supra vel infra

medium situs. Chromatophora disciforma numerosa vel nulla.

Granula paramylacea ovalia vel bacilliformia, non annulata.

Habitat: Singapore, Horto Botanico, in lacu, legit B. Skvortzov,

July 1962 — S.P.

Dedicavi in honorem Dominae V.V. Down, Singapore.

Cell fusiform, metabolic, 18-63 x 12-25 yu; anteriorly more or

less narrowed, obliquely obtuse, posteriorly straight or oblique,

shortly acute. Periplast non striate. Flagellum small up to half the

448 Gardens’ Bulletin, Singapore — XXII (1968)

length of the cell. Stigma red, oblong. Gullet elongate or oval.

Nucleus situated in the middle or above or below the middle.

Chromatophores absent, or when present many, disc-shaped.

Granules paramylaceous, oval or bacillus-shaped, not annulated.

Differs from E. proxima (vide Popova, 1955 p. 143, figs. 50,

1-3) in the periplast being non-striate, granules paramylaceous

not coloured and not annulated, flagella nearly half or 4 the

length of the cell.

I have dedicated this species to Mrs. V. V. Down, Singapore.

2. Euglena prowsei sp. nov. Plate I, fig. 5.

Differt a E. minuta Prescott et E. vivida Playfair cellulis sine

pyrenoidibus.

Cellula late fusiformis utrinque apiculatis, plus minusve

attenuatis. Flagellum cellulae longius. Stigma oblongum, rubrum.

Chromatophorum laminatum viride sine pyrenoidibus. Granulis

paramylaceis mumerosis ovoideis, utrinque chromatophoris.

Nucleus magnus posterior.

Habitat: Singapore, Horto Botanico, in lacu cum plantis

aquaticis, legit B. Skvortzov, August 1962 — S. P.

Dedicavi hauc speciem ad Dom. Dr. G. A. Prowse, Director

of Tropical Fish Culture Research Institute, Batu Berendam,

Malacca.

Cells broadly fusiform, apiculate at both ends, more or less

narrowed. Flagellum longer than the cell. Stigma red, oblong,

anterior. Chromatophore laminated, green without pyrenoids.

Granules paramylaceous, numerous, ovoid coloured on both sides.

Nucleus large, posterior.

Differs from E. minuta and E. vivida in the absence of pyrenoids

in the cells.

3. Euglena proxima Dang. var. tropica var. nov. Plate LI, fig. 6.

Differt a E. proxima Dang. (vide Popova, 1955, p. 143, fig. 50,

1-3) granulis haematochrome in parte posteriore cellulae instructis.

Cellula late fusiformis, 25-35 x 20-22 u. Periplasto non striato.

Stigma ovale. Chromatophorum disciforme sine pyrenoidibus.

Granulis paramylaceis, annulatis; eis haematochromis numerosis

parte posteriore cellulae. Nucleus posterior. Flagellum cellula

longius.

Habitat: Singapore, Horto Botanico, in lacu, legit B. Skvortzov,

July 1962 — SP.

Cells broadly fusiform, 25-35 x 20-22 yu. Periplast non striate.

Stigma oval. Chromatophores disc-shaped, without any pyrenoids.

Granules paramylaceous, annulated; haematochromes numerous in

the posterior. Nucleus posterior. Flagellum longer than the cell.

Differs from E. proxima Dang. (Vide Popova, 1955, p. 143,

figs. 50, 1-3) in the presence of haematochromic granules in the

posterior of the cell.

Skvortzov — Euglena 449

4. Euglena clara Skuja in Popova (1955), Plate I, figs. 7-8.

Cells metabolic, fusiform, anteriorly shortly attenuate, obliquely

rounded at apex, posteriorly gradually attenuate, acute or obtuse,

25-37-40 x 10-20 ». Chromatophores in oblong plates with

sheathed pyrenoids. Paramylaceous granules bacilliform. Haemato-

chrome granules numerous in the middle of cell. Flagellum % of

the cell-length. Eyespot oblong, red. Gullet oblong, nucleus dis-

tinct.

Differs from the type form in having hyaline periplast and

haematochromic granules.

Habitat: Singapore, Botanic Gardens, in a pond collected by

B. Skvortzov, July 1962. Reported from Europe. |

5. Euglena clara Skuja var. singaporensis var. nov. Plate I, fig. 9.

Differt a E. clara var. clara granulis paramylaceis majoribus,

ellipsoides.

Cellula ca. 30 » longa. Granula paramylacea eis var. clarae

majora, ellipsoidea.

Cells about 30 » long. Paramylaceous granules ellipsoidal,

longer than those in the type form.

Habitat: Singapore, Botanic Gardens, in pond collected with

the type form by B. Skvortzov, July 1962. Type in Herb. S.P.

6. Euglena variablilis Klebs in Popova, 1955. Plate I, fig. 10-12.

Cells ovoid or short ellipsoid, rarely fusiform, 25-35 x 7.20 u.

Periplast not seriated. Flagellum about cell length. Eyesport red,

large, oblong. Chromatophores in large, roundish or oval plates

without pyrenoids. Paramylaceous granules small, oval, numerous.

Habitat: Singapore, Botanic Gardens, in a pond, collected by

B. Skvortzov, July 1962, together with E. clara. Reported from

Europe, central and eastern parts of Asia.

Acknowledgements

_ The writer acknowledges the help of Mrs. V. V. Down and the

Singapore Immigration authorities in granting a four-hour landing

pass, and wishes to express his gratitude to the Director of the

Botanical Institute of Sao Paulo, Brazil, the leaders of the

Cryptogamic Section, Dr. Oswaldo Fidalgo, Dr. Joao Salvador

Furtado and Dr. Carlos Eduardo de Mattos Bicula and the

Secretary of the Institute Marlene Ocana Orlando for facilities

for the conduct of this and the author’s other algological research

‘ work. He also acknowledges the help of Dr. C. X. Furtado of the

Botanic Gardens, Singapore in the preparation of the Latin

diagnoses, and of the Director, Mr. H. M. Burkill, for publishing

this note in The Gardens’ Bulletin, Singapore.

References.

PRINGSHEIM, E. G.: Contribution towards a Monograph of the Genus

Euglena, Leipzig. 1956.

Gospics, M.: Genus Euglena. Madison. 1953.

Popova, T. G.: Euglenophyta. Mosqua. 1955.

HUBER-PESTALOZZI, G.: Euglenophyceen. Stuttgart. 1955.

450 Gardens’ Bulletin, Singapore — XXII (1968)

Plate I. Fig. 1-4, E. downiae; Fig. 5, E. prowsei; Fig. 6, E. proxima var.

tropica; Fig. 7-8, E. clara; Fig. 9, E. clara var. Singaporensis; Fig.

10-12, E. variabilis.

On a new species of the genus Collodictyon Carter,

a colourless flagellata new to the Hongkong flora.

(De specie et genere Collodictyon Carter nova ad

flora Hongkongensis.)

B. V. SKVORTZOV

Instituto de Botanica, Sao Paulo, Brasil

Abstract

While the author was travelling from China to Brasil in 1962 via

Hongkong, he visited The Peak, a mountain near the city, and collected

samples of mosses. These were cultivated with the addition of pepton for

flagellata at the Cryptogamic Laboratory of the Botanical Institute, Sao

Paulo. Amongst the numerous aerial diatoms present in the culture a

large colourless flagellate was found which was feeding on both the diatoms

and green algae. This flagellate belongs to the genus Collodictyon pro-

posed by Carter many years ago from India. During the past 20 years

it. has been found in many parts of Europe. The Hongkong specimens

are here described as Collodictyon hongkongense sp. nov.

The genus Collodictyon was referred by A. Pascher and E.

Lemmermann (1914) to the family Tetramitaceae, order Proto-

mastiginae. Pascher in 1927 transferred it to the colourless

Volvocales. In the latest revision, made by Russian algologists of

Charkov University (Dedusenko-Shtegoleva, et al., 1959),

Collodictyon is placed near to the colourless genera Polytomella

Aragao and Cyromitus Skuja of fam. Polystomellaceae (Aragao)

Volvocales.

Studying the description of the genus Collodictyon in different

books, the present author has come to the conclusion that the

descriptions were made from different species which seem to be

from different genera. For instance, in the description of 1914

(Pascher & Lemmermann) the cells of Collodictyon are stated to

be strongly metabolic: in the revision of 1959 (Dedusenko-

Shtegoleva ef al.), it is written that the cells have a distinct

periplast.

The Hongkong specimens are not similar to Collodictyon

triciliatum Carter as given by Pascher and Lemmermann.

Collodictyon hongkongense sp. nov. Figs. 1-7.

Differt a C. triciliato membranis non metabolicis, vacuolis con-

tractilibus nullis vel indistinctis, nucleo indistincto, forma monadis

diversa, posteriore acuta vel bifurcata.

Cellula solitaris, libere natans, vix vel_non metabolica, holozoice

nutrita, applanata, ambitu variabilis ovata, oblonga vel fere

fusiformis, aut triangularis, 10—-15—18-26, 37-40 u longa; anteriore

acuta, truncato-rotundata, plerumque depressa, cum nucleo indis-

tincto, supra medium sito posteriore utrinsecus angustata, apice

452 Gardens’ Bulletin, Singapore — XXII (1968)

Collodictyon hongkongense,

Skvortzov — Collodictyon 453

acuta or caudata, oblique truncata vel bilobata; ventre carinis 2

paulo altis longitudinaliter praedita; dorso fere applanata.

Periplastus nullus vel tenuissimus, superficie rugosus, nunquam

laevis; cytoplasma hyalinum, vacuolis multis cum granulis olei

leicosinique copiosis instructa; vacuolis contractilibus non visis.

Pseudopodia posteriora tantum visa, vix metabolica, simplicia,

extensibilia. Flagella 4, tenuissima, vix visibilia, aequalia, monade

aequilonga vel eo paulo longiora. Proles divisione cellulae motae

longitudinali aucta.

Habitat: Hongkong, in monte ““The Peak’’ dicto, inter muscos

rupestres, legit Skvortzov. 15—VIII-1962; holotypus ex speciminibus

in Lab. Crypt., Sancto Paulo, Brazil, cultis, et ibidem asservatus

est.

Differs from C. triciliato in the cells being of different shape

and non-metabolic and contractile vacuoles being absent or indis-

tinct, indistinct nucleus and acute or bifurcate posterior end.

Cells solitary, free swimming, hardly or non-metabolic, holozoi-

cally nourished, flattened, varying in shape from ovate, oblong or

almost fusiform or triangular, 10-40 wu long; anteriorly acute,

truncately rounded, often depressed, with an indistinct nucleus

seated above the middle, contractile vacuoles not seen; posteriorly

narrowed on both sides, acute or caudate at apex or obliquely

truncate or bilobed; ventrally provided with little raised ridges;

dorsally more or less flattened. Periplast absent or very slender

with its surface rugose, never smooth; plasma transparent, having

many vacuoles abounding in oil and leicosin granules; contractile

vacuoles not seen. Pseudopodia seen only in the posterior, hardly

metabolic, simple, extensible (?). Flagella 4, very slender, hardly

visible, equal, as long as the cell or slightly longer. Offspring

produced by longitudinal division of free moving cells. Locomotion

is by creeping or rotation, and is not rapid. Feeding holozoic,

diatom fustules, or more rarely cells of Chlorophyta adhering to

the periplast are covered over.

Habitat: The Peak, Hongkong, in mosses growing in rocks,

collected by B. Skvortzov, 15 August 1962; holotype cultivated

from this material at the Cryptogamic Laboratory of the Instituto

de Botanica, Sao Paulo, Brasil and deposited there.

Acknowledgements

The author expresses his gratitude to the Director of the

Botanical Institute, Sao Paulo, Dr. Alcides Ribeiro Teixera and

to the leaders of the Cryptogamic’Section, Dr. Oswaldo Fidalgo,

Dr. Joao Salvador Furtado, and Dr. Carlos Eduardo de Mattos

Bicudo, and to the secretary Marlene Ocana Orlando for facilities

for the conduct of this and his other algological research work,

and for the assistance of Dr. C. X. Furtado of the Botanic Gardens,

Singapore, in the preparation of the Latin diagnosis.

454 Gardens’ Bulletin, Singapore —XXII (1968)

Literature

Fritscu, F. E.: The structure and reproduction of the algae, Vol. L

Cambridge. 1935.

DDEDUSENKO-SHTEGOLEVA, N. T., MATVIENKO, A. M. and SCHKORBATEV,

L. A.: Chlorophyta; Volvocineae. Mosqua, 1959.

PASCHER, A. und LEMMERMANN, E.: Flagellata I in Die Susswasserfiora

Deut., Oster. und der Schwiez. Heft. 1914. Jena.

PASCHER, A.: Volvocales-Phytomonadinae. Flagellatae IV. Chlorophyceae

I. Heft 4. 1927. Jena.

New genera of primitive green flagellata from

Hongkong and Sao Paulo, Brazil.

B. V. SKVORTZOV

Instituto de Botanica, Sao Paulo, Brazil

Abstract

Three new genera of primitive green flagellata are described: Protoch-

roomonas, Angulomonas and Protoaceromonas, belonging to Fam.

Pyramydomonadaceae Pascher, Ord. Polyblepharidales and Class Vol-

vocineae. One new species of the first named genus, two new species.

of the second and three mew species of the last-named genus are

described and illustrated together with a key to the genera.

Materials

The collecting localities (and the abbreviations used in this

paper) of the material studied and the dates of collection were:

Habitat I (Hab. I): Sao Paulo, Brazil, Parque do Estado, in

temperate swamp; col. B. Skvortzov. 20 May 1966.

Habitat II (Hab. Il): /bid, on surface of the soil with a growth

of Oscillatoria; col. B. Skvortzov. 2 May 1966.

- Habitat III (Hab. HI): bid, on surface of soil with polluted

spring water; col. B. Skvortzov. 17 May 1966.

Habitat [IV (Hab. IV): Aberdeen, Hongkong, in a rivulet; col.

S.T. Chan, 16 March 1966.

Key to genera

Ceil naked, with a spinulose or rugose periplast; flagella 2,

connate; cells of varying shapes

Genus 1: Protoaceromonas

—

Cell naked, periplast with spines or wrinkles 2a. Flagella 2,

connate; cells triangular or ellipsoid in front view but much

depressed, short lanceolate, trapezoid or reniform in lateral

view.

Genus 2: Angulomonas

2b. Flagella 2, very remote from one another never connate.

Genus 3: Protochroomonas

Description of genera and species

I. Protoaceromonas gen. nov.

Monades solitariae, libere natantes, vix vel paulo applanatae,

metabolicae. Periplastus tenuissimus vel nullus, rugosus vel minute

spinosus, luteolus vel hyalinus, stigma vel vacuolam contractilem

includens. Flagella 2, connata, fere 14 cellulis longiora, similia

natantia; chromatophora parietalia ut videtur, viridia, insignia cum

pyrenoidibus nudis; motu rotanti, rapidissimo.

456 Gardens’ Bulletin, Singapore — XXII (1968)

Differt ab Angulomonas gen. nov. periplasto rugose vel spinuloso

et a Protochroomonas gen. nov. flagellis connatis.

Typus generis: Protoaceromonas spinosa sp. nov.

Cells solitary, freely swimming, hardly or little flattened,

metabolic. Periplast slender or none, rugose or minutely spinose,

yellowish or hyaline. Flagella 2, connate, almost 14 longer than

the cell, similar, swimming. Eye-spot or contractile vacuole

present. Chromatophores apparently parietal, green, large, with

naked pyrenoids; movement rapid, rotational.

Differs from Angulomonas gen. nov. by its rugose or spinulose

periplast and from Protochroomonas gen. nov. by its connate

flagella.

Species 3. Distribution: Hongkong, Brazil.

Specific Key

1. Cell suborbicular in outline, angled, anteriorly depressed,

5-6-7 p long; periplast slender, rugose or spinulose; eye-spot or

contractile vacuole present.

1. P. spinosa Hab. I

2. Cell suboval, 12-13 x 7-8 yw; periplast rugose; eye-spot not

present.

2. P. rugosa Hab. Ill

3. Cell orbicular, naked, rugose or denticulate, 9-10 » in diam.;

contractile vacuole not seen.

3. P. orbicularis Hab. IV

1. Protoaceromonas spinosa sp. nov. Figs. 1 and 2.

Cellula ambitu suborbicularis, angulata, anteriore depressa,

‘periplasto tenuissimo rugoso vel minute spinoso, 5-6-7 » longa

‘cum stigmate vel vacuola contractili: Hab. I. Legit B. V. Skvortzov.

Holotypus — S. P.

2. Protoaceromonas rugosa sp. nov. Fig. 3.

Cellula subovalis, periplasto rugoso; stigmate nullo; 12-13 x

‘7-8 uw: Hab. IL. Legit B. V. Skvortzov. Holotypus — S. P.

3. Protoaceromonas orbicularis sp. nov. Fig. 4.

Cellula orbicularis, nuda, rugosa vel minute dentata, 9-10 p» in

‘diam.; vacuola contractilis non visa: Hab. IV. Legis B. V.

‘Skvortzov. Holotypus — S. P.

II. Angulomonas gen. nov.

Monades solitariae, libere natantes, non metabolicae, ambitu fere

triangulares, ventraliter applanatae; periplastus distinctus cum

vacuola contractilis; chromatophorum pallide viride, parietale,

perforatum granulosum; pyrenoides praesens vel nulla; stigma

nullum; nucleus centralis; flagella 2, natantia.

Differt a Phyllocardium Korsch. cellulis ambitu triangularibus a

Protoaceromonas gen. nov. cellulis ventraliter applanatis.

Skvortzov — Green flagellata 457

Fig. I and 2. Protoaceromonas spinosa gen. et sp. nov.

3. Protoaceromonas rugosa sp. nov.

4. Protoaceromonas orbicularis sp. nov.

5. Angulomonas triquetra gen. et sp. nov.

6. Angulomonas stagnalis sp. nov.

7. Protochroomonas granulata gen. et sp. nov.

458 Gardens’ Bulletin, Singapore — XXII (1968)

Typus generis: Angulomonas triquetra sp. nov.

Cells solitary, freely swimming, not metabolic, almost triangular

in outline, flattened on the ventral side; periplast distinct;

chromatophore pale green, parietal perforated, granulose; pyrenoids

none or present; contractile vacuole present; eye-spot none, nucleus

central; flagella 2, swimming.

Differs from Phyllcocardium Korsch. in its triangular cells, and

from Protoaceromonas gen. nov. in its cells being ventrally

flattened.

Species 2. Distribution: Brazil. Hab. II.

Key to the species

1. Cells triquetrous in outline, ellipsoid in transverse (?) section,

more or less acute at both ends, 5—6 » long; pyrenoids present.

1. A. triquetra

2. Cells triquetrous, in transverse section almost three angled,

9-10 » long; pyrenoids indistinct.

2. A. stagnalis

Description of the Species

1. Angulomonas triquetra sp. nov. Fig. 5.

Cellula ambitu triquetra, in sectione transversali ellipsoidea,

apice utrinque plus minus acutis, 56 u longa: Hab. II. Legit B. V.

Skvortzov. Holotypus — S. P.

2. Angvlomonas stagnalis sp. nov. Fig. 6.

Cellula triquetra, in sectione transversali fere triangulata, 9-10 u

longa, pyrenoidibus indistinctis: Hab. II. Legit B. V. Skvortzov.

Holotypus — S. P.

Il]. Protochroomonas gen. nov.

Monades solitariae, libere natantes, non applanatae, cum vacuola

contractili; periplasto nullo vel tenuissimo, minute rugoso;

chromatophorum parietale, granulosum pallide viride, granulatum,

granulis amylaceis; pyrenoides et stigma nulla; nucleus centralis;

flagella 2, natantia, similia, 14 monade longiora, inter se remote

sita.

Differt a Mesostigma Lauter. cellulis non applanatis, periplasto

tenuissimo vel nullo et a Caecomonas gen. nov. Fam. Heteromas-

tigiaceae Korsch. Class Volvocineae (vide manuscriptum authorum

1955) flagellis similibus.

Typus generis: Protochroomonas granulata sp. nov.

Cells solitary, free swimming, not flattened; periplast none or

slender and minutely rugose; chromatophore parietal, pale green.

with paramylaceous granules; pyrenoids and eye-spot absent:

vacuoles contractile present, nucleus central; flagella 2, swimming,

similar, remotely seated from each other, 14 times longer than

the cell.

Skvortzov — Green flagellata 459

Differs from Mesostigma Lauter. in non-flattened cells, periplast

being thin or absent, and from Caecomonas gen. nov. ms. in

flagella being similar.

Species 1. Distribution: Brazil. Hab. I.

1. Protochreomonas granulata sp. nov. Fig. 7.

Ceilula 8-9-10 » in diam.; flagella 2, remote sita, natantia.

Hab. I. Legit B. V. Skvortzov. Holotypus — S. P.

Cell 8-9-10 » in diam., flagella 2, very remote from one an-

other, swimming.

Acknowledgements

This study was made in the Cryptogamic Section of the Botanical

Institute, Sao Paulo, Brazil. The author expresses his gratitude to

the Director of the Institute, Dr. Alcides Ribeiro Teixeira, and

to the leaders of the Cryptogamic Section, Dr. Oswaldo Fidalgo,

Dr. Joao Salvador Furtado, Dr. Carlos Eduardo de Mattos Bicudo,

and the Secretary Marlene Ocana Orlando for facilities in carrying

out this work. The material from Hongkong was received in 1966

for cultivation and study through the kindness of Dr. R. C. Stephens

and Dr. T. Widdowson of the Department of Botany, University

of Hongkong. The author thanks Mr. H. M. Burkill and Dr. C. X.

Furtado of the Singapore Botanic Gardens for help in editing

this paper. :

Literature

Fritscu, F. E.: The structure and reproduction of the algae. Vol. I.

Cambridge. 1935.

PaAScHER, A.: Volvocales-Phytomonadinae. Jena. 1927.

DEDUSENKO-SHTEGOLEVA, N. T., MATVIENKO, A. M., SCHKORBATOV, L. A.

CHLOROPHYTA: Volvocineae. Mosqua. 1959.

’* kin

™

LL dali lismid saniaeligt aad

ee,

in if pigiipnr + conn rstay "y

il } I i forme bits fi dis Olgas alulls De

zuqmioloH wesnovad ¥ at 2 at =

yy RSs sis Bhi al « [0-5 iw om

eee yi

, *

a Ci yorathent eal ith

: | oy

“3 f i a J as . - ~ 9 to a — >

f 70] 2 I hw a

_ : } y - Hi 1] i f ik 75 we dy oie rT — s

eathin afl how 2id Share

+ vous haw aseeislt a

Foliar sclereids in some species of Raphidophora:

and Scindapsus

HARDIAL SINGH

Botanic Gardens, Singapore

The taxonomic value of the foliar sclereids has been emphasised

by many early workers and this character has often been utilised

to distinguish between suborders of a family, a species within

the genus and also to assess the taxonomic position of varieties.

Engler (1908, 1920) in the family Araceae, distinguished the

subfamily Pothoideae from the Monsteroideae in that sclereids

were present in the stems, leaves and roots of only the members.

of the Monsteroideae. Again he further distinguished the tribe

Monstereae from the Spathiphylleae in that numerous sclereids

_ were present in the leaves and petioles of members of the

Monstereae when compared with those of Spathiphylleae. Very

few studies have been made in Monocots either to identify or

to delimit the species of different taxa. Recently, Nicolson (1960)

and Rao (1964) have studied sclereids in some species of

Raphidophora and Scindapsus. The present paper records the

observations of a continued study on other Malesian species of

the two above mentioned genera.

Materials and Methods

Materials for the present investigation were derived entirely

from herbarium specimens, except for Scindapsus hederaceus

where fresh material was used from the specimen growing in the

Botanic Gardens.

The following specimens were used in the study:

1. R. acuminata Merr. Ramos M. & Edano G. 75472 at Philippines.

Sept. 1928. SING.

2. R. batoensis Engl. Boden-Kloss C, 14713 at Sipora, Sumatra.

Oct. 1924. SING.

3. R. beccarii Engl. Furtado C. X., No. D at Perak, Malaya.

June 1937. SING.

4. R. cornerii Furtado. Corner E. J. 30441 at Ulu Kajang, Malaya.

Nov. 1935. SING.

R. falcata Ridl. 2? Haniff at ? Penang, Malaya 192 SING.

6. R. foraminifera Engl. Burkill I. H. and Haniff 12331 at Perak,

Malaya. June 1924. SING.

7. R. hollrungii Carr C. E. 13237 at Papua. July 1935. SING.

R. korthalsii Schott. Ridley H. N. at Malang, Borneo. 1893..

SING

9, SR huanseleri Hook. Burkill I. H. 6267 at Ipoh, Malaya. Sept. 1920..

SING.

10. R. lobbi Schott. Hardial 2 at Pahang, Malaya. June 1964. SING.

11. R. megasperma Engl.—at Buitenzorg.

12. R. minor Hook. Gordon H. S. 33280 at Perak, Malaya. Sept..

1937. SING.

462 Gardens’ Bulletin, Singapore — XXII (1968)

13. R. montana (Bl.) Schott. Purseglove J. W. 4473 at Bau, Sarawak.

Sept. 1955. SING.

14. R. octovulata v.A.v.R. — at Buitenzorg. 1927. SING.

15. R. palladivenia v.A.v.R.— at Buitenzorg. 1922. SING.

16. R. pilosula v.A.v.R.—at Buitenzorg. 1920. SING.

17. R. sylvestris (Bl.) Engl. Nur Md. at Kuala Selangor, Malaya.

Oct. 1937. SING.

1. S. beccarii Engl. Sinclair J. & Kiah 40804 at Trengganu, Malaya.

Sept. 1955. SING.

2. §. curranii Engl. & Krause J. & M. S. Clemens 219617 at Tenompok.

B. N. Borneo. May 1932. SING.

3. S. havilandii Ridl. Haviland 20019 at Saribos, Borneo. July

1892. SING.

4. S. treubii Engl. Burkill I. H. 9031 at Bukit Mertajam, Malaya.

Oct. 1922. SING.

5. S. hederaceus Schott. Growing in the Botanic Gardens —

Singapore.

The procedures outlined previously by Foster (1955) and Arnott

(1959) were followed to obtain the leaf clearings and macerations.

‘Cleared leaf material was used to study the density and mode

of distribution of the sclereids while macerated material was

helpful for study of the form and size of the individual sclereids.

All cleared and macerated material was stained with Safranin.

Transverse and paradermal sections of leaves were cut to study

the structure of the leaf and the positional relationship of the

sclereids, and these were stained with Safranin and Fast Green.

To obtain the length of the sclereid. the two distant points of

the arms of the sclereid oriented in opposite directions were

measured, while for the width, the widest portion of the central

axis was measured. In a single species, though a certain form of

sclereid predominated, there were also a few others which did

not exhibit this typical form but were a variation of this. Measure-

ments were only taken of those sclereids exhibiting the basic

Shape typical of that species. The number of sclereids present

in a given microscope field were counted from the leaf clearings

of different species to assess their density of distribution. The

figures recorded in all cases are an average of ten measurements

or counts.

Observations

Of the twenty-two species investigated, sclereids are present in

the mesophyll tissues of all the species except in R. beccarii,

R. corneri, R. korthalsii and R. minor. Sclereids found in each

of the various species are basically either of the LINEAR or the

H-SHAPED type, though relatively few sclereids not exhibiting

this typical form were also present but these were mere variations

of either the linear or the H-shaped condition.

Hardial Singh — foliar sclereids 463

1. LINEAR sclereids were seen to be predominant in eight

of the twenty-two species investigated, namely R. falcata, R. lobbii,

R. octovulata, R. palladivenia, S. beccarii (Figs. 23-24), S. curranit,

S. treubii and S. hederaceus.

In six of the above mentioned species, the linear sclereids

within each species did not vary obviously in their size. However

in R. falcata and S. hederaceus (Fig. 30) two distinct sizes of

linear sclereids were seen, one type being about half the length

and width of the larger sclereids. (See Table I). The variations of

the linear sclereid were that in some, a notch was seen to arise

around the middle portions of the sclereids, (Figs. 23-24), and

in some others this protruberance was much longer resulting in

almost a ‘Y’-shaped sclereid (Fig. 25). In R. lobbii, R. octovulata,

S. beccarii, S. treubii, and S. hederaceus, the linear sclereid

predominated but an H-shaped sclereid was also found to be

present but its occurrence was extremely rare and accounted for

about 1 percent of the sclereids present. :

2. H-SHAPED sclereids are predominant in ten of the species

investigated namely R. acuminata (Figs. 1-2), R. batoensis, R.

foraminifera (Figs. 5-6), R. hollrungii, R. kunstleri (Figs. 8-9),

R. megasperma (Figs. 14-15), R. montana, R. pilosula (Fig. 16),

R. sylvestris (Figs. 19-20) and S. havilandii. Besides the ideal

H-shaped condition, tremendous variation of the H-shaped sclereid

was seen, even in one and the same species. Figures 8-13 show

the range of sclereid forms seen in R. kunstleri. Linear and

Y-shaped sclereids were also seen to be present in species in

which the H-shaped sclereid condition predominated but their

occurrence was extremely rare and accounted only for about

1-2 percent of the total sclereids present. In R. hollrungii and

R. kunstleri, the linear and ‘Y’-shaped sclereids accounted for

about 25 percent of the total sclereids present. Such sclereids were

seen in R. acuminata (Figs. 3-4), R. foraminifera (Fig. 7), R.

kunstleri (Figs. 11-13), and R. sylvestris (Figs. 21-22).

In most of the species, the sclereids were rather sparsely

distributed and only in R. acuminata, R. hollrungii, R. lobbii,

R. montana, R. octovulata, R. pilosula and R. sylvestris were

they densely distributed. All species of Scindapsus had sparsely

distributed sclereids. The sclereids were, in all the species

investigated, orientated in the tangential plane to the leaf surface

and were again diffuse in the distribution.

A very striking feature noticed was the presence of druses

within the mesophyll cells of all the species investigated. These

were best seen in the paradermal sections.

464 Gardens’ Bulletin, Singapore — XXII (1968 }

Discussion and Summary

The presence of H-shaped and linear sclereids in species of

both the genera, namely Raphidophora and Scindapsus shows their

close relationship and thus justifies Engler’s placing both these

genera in the tribe Monstereae of the subfamily Monsteroideae.

Considering the linear sclereid as the basic form, one can suggest

a possible evolutionary trend resulting in the “Y’ and ‘H’-shaped

sclereids, all these three forms and their intermediates being found

at times in one and the same species: (Figs. 31-40).

Foliar sclereids are seen present in eighteen of the twenty-two

species investigated presently. Sclereids found in the various species

were either of the two basic forms, linear or H-shaped, though

variations of these basic forms were also seen to be present but

in extremely small numbers.

Linear sclereids were predominant in eight of the species while

the other ten had the H-shaped sclereids predominanting. All the

species of Scindapsus investigated had sparsely distributed sclereids

and only seven of the seventeen species of Raphidophora exhibited

a dense distribution of sclereids.

Druses are commonly present in the mesophyll cells of all the

species investigated.

Literature cited

1. ENGLER, A., 1908: Das Pflanzenreich. Heft 37. w. Engelmann.

Liepzig.

2. ENGLER, A., 1920: Das Pflanzenreich. Heft 74. w. Engelmann. Liepzig.

Liepzig.

3. NICOLSON, D., 1960: Amer. J. Bot., 47, 598.

Rao, T. A. 1964: Proc. Ind. Acad. Sci. Lx, 66.

Hardial Singh — foliar sclereids 465

TABLE I

SCLEREID

8 a iatiiess Density | Predomi-

Length | Width | Av. No./ nant

‘ field Type

1. Raphidophora acumi-

nata .. nf? 185 1,105 49 Dense-16 | H-shaped

2. R. batoensis 85 589 54 Few - 6 | H-shaped

3. R. beccarii 114 —

4. R. corneri 284 —

5. R. falcata 200 1,278 32 Few - 6 Linear

781 16

6. R. foraminifera 71 525 45 Few - 3 | H-shaped

7. R. hollrungii 128 909 60 Few - 9 | H-shaped

8. R. korthalsii 128 —

9. R. kunstleri 170 33 57 Few - 2 | H-shaped

10. R. lobbii 114 1,266 13 Dense-22 | Linear

11. R. megasperma 170 477 40 Few - 7 | H-shaped

12. R. minor 85 —

13. R. montana 355 312 ae Dense-36 | H-shape d

14. R. octovulata 128 2,800 24 Dense-28 | Linear

15. R. palladivenia 71 1,355 20 Few - 2 | Linear

16. R. pilosula 184 630 61 Few — 9 | H-shaped

17. R. sylvestris 213 540 50 Dense-32 | H-shaped

18. Scindapsus _ beccarii 170 852 16 Few — 8 | Linear

19. S. curranii 142 1,662 57 ee 2 | Linear

20. S. havilandii 128 568 60 Few - 1 | H-shaped

21. S. trevbii 170 2,130 67 Few — 5 | Linear

22. S. hederaceus 284 1,846 64 Few —- 3 | Linear

1,136 32

* All measurements are in microns and each figure represents an average

of ten measurements.

466 Gardens’ Bulletin, Singapore — XXII (1968)

1 2 3 4

16 17

Pde)

| 22

Plate I. Sclereids forms: Figs.

1-4 — R. acuminata, Figs. 5-7 — R.

foraminifera, Figs. 8-13 — R. kunstleri, Figs. 14-15 — R. megasperma,

Figs. 16-18 — R. pilosula, Figs. 19-22 — R. sylvestris, Figs. 23-25,

S. beccarii. Camera lucida drawings x 60.

lvestris x

Em. 27k.

ike. lobbn & 100, Fig. 29 .— Ris

30 — S. hederaceus x 33.

: Fig. 26 — R. acuminata x 100

hollrungii x 100, Fig. 28 —,

Fig.

’

Plate II. Sclereid forms

100

‘(oneUUeIseIq) Op-Ie ‘ssi

‘PIOJO[OS IVOUT] dISeq, OY} WoO, pus ArIevuOTIN[OAD ga[QGissod VY ‘[]] 2d

——_—_— —— a

467

2o ee 1c

foliar sclereids

—

Book Review

THOMAS, A.: Gardening in Hot Countries

Faber & Faber, London, £1/10/—.

In writing a book on the subject of gardening in hot countries

it is not possible to cover fully those aspects of horticulture which

are peculiar or typical of each and every country. Even within

the tropics there is a tremendous range in the climates and soils

which to a great extent determine the types of gardening methods

practised. In Singapore for instance where the soil is predominantly

Clay the traditional Chinese method of growing pot plants in

burnt clay supplemented with dried cow dung and rotted soya

bean cake is very typical of Malaysia and obviously the best

form of pot plant culture in the region. One can also quote other

examples which are typical of gardening of a particular country.

Nevertheless the publication is an excellent and painstaking

effort on the part of the author to cover as much ground as one

possibly can on the various forms of gardening as practised in the

hot countries particularly in the African region.

The author has presented very vividly the difficulties and

problems the gardener has to face in the tropics and many of his

suggestions and ideas as a result of long experience of gardening

in the tropics are very practical and would apply equally to the

‘hot countries, be they in Africa, Asia, Australia or America.

A great deal of the book is spent on the precise descriptions of

the many tropical plants. This is most useful, as the gardener

‘must know his plants before he can grow them. Photographs to

illustrate some of the plants are well taken but some of the

captions need looking into. For example Plumieria obtusifolia

-should read as Plumeria obtusifolia and Renanthera coccinea is

in fact Arachnis maingayi.

I am sure this book will prove of immense value to both the

-gardening enthusiasts and horticulturists alike.

A. G. ALPHONSO

MLIA A

2Pamonu!

a1 asdeit 4 1sdet ‘ .

insy lo iosjdue sdi ao.dood 2 gabiw ah

. fl to dioogas sears yllu) 1sve0 Of sidizzog ton. 1 ai tt”

va. woo yvieva ban dogs to Ia: 1 20 ry

Homily off ni sgna1 avobasmsy s 2f sis) Igo >

4] sq) orl) onterisieb iustxe Jeo R ot dots -

fing ora ororlw sonpteni 10? Moqegaie al .b soem 7

miwota Io berm seen) Innoilther oda

\ agnuh weo bend cdiiw bornsesiqqua vals ire

ode bow sievelaM Io feoiecvi ry zi sno need:

joo ont ai swwitlue Joaky Jog Io anot -

netien to Isotqv) ova soidw ital

ce po

: y ce ai sorediidua 4) ssisdtesvelt iad 3

(ouit2s t8V sorljun set Yo hag 6 oe

grinobieg I) tcmol aonsy apt . uss \idizzog -

nig qeciilA ort oi yhsluoine zotames jen >.

av bstnsend ed 70% ws oft

eat, tsaoineg od) ensidouy

io ile s ee esobt baa snot eATQUE

WV SIs Z2qor sds nt

LA. A, Boi ni yort od .2siviavoo-jod

‘. 4. :

od alt jo lasb }

g *

a¢ el nr +

2 : inalq isorqox

eg ee ae

HW SIG angi Sf} te emoe on veal ‘

LET } 3} WwmetcyR i of Age ; Q pa idool- besn 2a0i8 age *

Pe. ‘ oi

ma brs ailoliaido pivactsitl 2s best aa

WYRM zipwlam mg ot a ;

ot suley sensi Io svoiq Uliw say ehh one f ms }

ee apes baa 2

INDEX, VOL. XXII

(a) Name of authors in CAPITALS.

(b) New taxa and binomials in bold.

Agarophytic seaweeds, 429.

Alginophytic seaweeds, 429.

ALLEN, B. M., 41, 173.

Allowoodsonia Mef., 23.

whitmorei Mef., 23.

Alsophila latebrosa var. ornata Ridl.,

Alstonia vitiensis Seem. var. whit-

morei Mef., 24.

Analectis speciosa Vahl, 166.

Ancistrocladus tectorius (Lour.) Merr.,

Lis:

Angulomonas Skvortz., 456.

stagnalis Skvortz., 458.

triquetra Skvortz., 458.

Anisoptera reticulata Ashton, 260.

ASHTON, P. S., 259.

Aspidium decurrens Presl, 177.

hirtipes Blume, 173. :

mamillosum C. Chr., 177.

pteropus Kze.,177.

Azoala leerii T. & B., 224.

Balibai Blanco, 163.

Book review, Thomas: Gardening in

Hot Countries, 469.

Buchanania spp., 2.

Burckella sorei Royen, 33.

Burkill’s Dictionary of Economic

Products — reprinting, 353.

BURKILL, H. M., 71, 107, 429.

Burkill, I. H. — a bibliography, 71.

Burkill, I. H.—a biography, 67.

Calophyllum cerasiferum Vesque, 13.

euryphyllum Laut., 12.

kajewskii A. C. Smith, 9.

macrophyllum Scheff., 12.

paludosum T. C. White, 13.

peekelii Laut., 11.

sil Laut., 12.

solomonense A. C. Smith, 9.

soulattri Burm., 13.

vitiense Turrill, 12.

Campnosperma brassii Mertr., 4.

brevipetiolata Volk., 4.

Caruthersia mollis Megf., 25.

Chelonespumum spp:, 35.

Chlorogonium aberdeenii Skvortz.,

188.

astigmatae Skvortz., 188.

chanii Skvortz., 188.

euchlorum Ehr., 189.

stentoni Skvortz., 189.

subtropicale Skvortz., 188.

widdowsonii Skvortz., 188.

Collodictyon hongkongense Skvortz.,

451.

CROWTHER, P. C., 429.

Cyathea alleniae Holtt., 45.

alternans (Wall.) Presl, 49.

borneensis Copel., 45.

brunonis (J. Sm.) Wall., 48.

burbidgei sens. Holtt., 50.

contaminans (Wall.) Copel., 49.

excavata Holtt., 46.

gigantea (Wall.) Holtt., 48.

glabra (Bl.) Copel., 48.

hymenodes Mett., 44.

incisoserrata Copel., 46.

kingii (Clarke) Copel., 47.

latebrosa (Wall.) Copel., 47.

latebrosa var. indusiata Holtt., 44.

lurida (Bl.) Copel., 47.

moluccana R. Br., 48.

obscura (Scort.) Copel., 50. -

obtusata Rosenst., 45.

polypoda Bak., 50.

recommutata Copel., 47.

squamulata (Bl.) Copel., 51.

sumatrana Baker, 44.

trichodesma (Scort.) Copel., 50.

tripinnata Copel., 48.

Cyclobalanopsis gemelliflora (Bl.)

Oerst., 406.

horsfieldii (Miq.) Oerst., 406.

lineata (Bl.) Oerst., 408.

lowii (King) Schott., 394.

merkusii (Endl.) Oerst., 406.

merrillii (V. Seem.) Schott., 397.

oidocarpa (Korth.) Oerst., 390.

sericea (Scheff.) Schott., 392.

treubiana (V. Seem.) Schott., 412.

turbinata (Bl.) Schott., 406.

Cyclosorus papilio (Hope) Ching, 180.

Cystoseira, alginate content, 437.

472 Garden's Bulletin, Singapore — XXII (1968)

Diploknema grandiflora (Ridl.) H. J.

Lam, 215.

Dipterocarpaceae, Notes on, 259.

Dipterocarpus cuspidatus Ashton, 261.

Dryobalanops abnormis Sloot., 262.

Dryopteris hirtipes (BI.) O.K., 173.

var. exinvolucrata (C. B. Clarke)

B. Allen, 174.

Endospermum malaccense M.A., 8.

medullosum L. S. Smith, 8.

Eucheuma, agar production, 430.

Eugenia cincta (Merr. & Perry) Whit.,

16.

myriadena (Merr. & Perry) Whit.,

16.

onesima (Merr. & Perry) Whit., 17.

Euglena clara Skuja, 449.

var. singaporensis Skvortz., 449.

downiae Skvortz., 447.

prowsei Skvortz., 448.

ne Dang. var. tropica Skvortz.,

variabilis Klebs, 449.

Flagellata, 187.

Freycinetia, 129.

almonoguiensis Kaneh., 144.

angustissima Ridl., 134.

archboldiana Merr. & Perry, 135.

arborea Gaud., 130.

arfakiana Martelli, 135.

arnotti Gaud., 130.

brassii Merr. & Perry, 135.

boninensis Nakai, 147.

carolinensis Kaneh., 140.

crucigera Kaneh, 136.

cyrtocarpa Kaneh., 132.

distigmata B. C. Stone, 131.

elegantela B. C. Stone, 137.

fibrosa Mart., 136.

funicularis (Savigny) Merr., 132.

inouei Kaneh., 135.

kanehirae B. C. Stone, 136.

kiekie B. C. Stone, 131.

klossii Ridley, 131.

mariannensis Merr., 140.

var. microsyncarpia Hosok., 144.

monticola F. B. H. Brown, 131.

perryana B. C. Stone, 133.

platyphylla Kaneh., 136.

polyclada Merr. & Perry, 134.

ponapensis Mart., 143.

radicans Gaud., 137.

salamauensis Merr. & Perry, 133.

Freycinetia cont.:

scandens sens. Hook. & Arn., 130.

sogerensis Rendle, 133.

sogerensis, 136.

spinellosa Kaneh., 137.

stenodonta Merr. & Perry, 133.

stenophylla Warb., 134.

strobilacea Bl., 132.

tafaensis Merr. & Perry, 137.

villalobosii Mart., 144.

Gilliland, H. B., obituary notice, 107.

Gmelina dalrympleana (F. Muell.)

Lam, 20.

moluccana (Bl.) Backer, 18.

salomonensis Bakh., 18.

Gnetum gnemon L., 29.

latifolium BIL. 29.

Gracilaria, agar production, 429.

GREEN, §S., 53.

GREENWOOD-BaARTON, L. H., 429.

HARDIAL SINGH, 193, 461.

Hasskarlia leucacantha Walpers, 236.

‘ Hocttum, R. E., 41, 67.

Hopea aequalis Ashton, 271.

altocollina Ashton, 272.

andersoni Ashton, 272.

ssp. andersoni, 273.

ssp. basalticola Ashton, 273.

bullatifolia Ashton, 274.

celtidifolia Kosterm., 443.

centipeda Ashton, 274.

depressinerva Ashton, 275.

enicosanthoides Ashton, 276.

kerangasensis Ashton, 277.

laxa Sym., 271.

longirostrata Ashton, 277.

megacarpa Ashton, 278.

mesucides Ashton, 279.

pachycarpa (Heim) Sym., 271.

pterogota Ashton, 280.

tenuivervula Ashton, 281.

Isonandra microphylla de Vriese, 224.

KENG, H., 113.

Kentrochosia Ltb. & K. Schum., 26.

Keura odorifera Forssk., 236.

Kopsia Blume, 26.

KOSTERMANS, A. J. G. H., 443.

Index

Lastrea hirtipes (Bl.) Moore, 173.

Litsaea luzonica Blanco, 163.

Madhuca decipiens J. Sinclair, 215.

dubardii H. J. Lam, 1925, 218.

dubardii H. J. Lam, 1927, 217.

selangorica (K. & G.) J. Sinclair,

Zt.

sericea (Miq.) H. J. Lam, 220.

MARKGRAF, F., 23.

eee superficiale (Bl.) Ching,

Munir, A. A., 153.

Myriophyllum, Notes on, 229.

Myriophyllum tuberculatum Roxb.,

230.

(Bl.) Kosterm.,

Mpyristica rumphii

446.

Nepenthes spp., 53.

Nephrodium decurrens (Presl) Bak.,

Li:

hirtipes (Bl.) Hook., 173.

var. exinvolucrata C. B. Clarke,

174.

papilio Hope, 180.

Ochrosia sciadophylla Megf., 26.

Padina, alginate content, 437.

Pagiantha koroana Mef. var. salom-

onensis Mef., 28.

Palaquium masuui Royen, 37.

Pandanaceae, Studies on, 231.

Pandanus fascicularis Lamk., 236.

eucanthus Hassk., 236.

odoratissimus Linn. f., 236.

odorifera (Forssk.) Kuntze, 236.

verus Rumph., 236.

Payena acuminata (Blume) Pierre, 221.

grandiflora Ridley, 215.

havilandii K. & G., 226.

lanceolata Ridl., 224.

leerii (T. & B.) Kurz, 224.

longipedicellata Brace, 226.

lowiana Pierre, 224.

lucida (G. Don) A. DC., 222.

microphylla (de Vriese) Pierre, 224.

obscura Burck, 226.

pseudoterminalis H. J. Lam, 225.

ridleyi Gaud., 220.

selangorica K. & G., 217,

seriacea Miq., 220.

473

PHILLIPS, J., 107.

Plant diseases in Sarawak, 123.

Pleopeltis nigricans (v.A.v.R.) v.A.

v.R.., :

rupestris var. nigricans V.A.V.R.,

176.

superficialis (Bl.) Bedd., 176.

Polypodium nigricans v.A.v.R., 176.

normale Don var. sumatrana Bk.,

176.

superficiale Blume, 176.

Pometia spp., 17.

Protoaceromonas Skvortz., 455.

orbicularis Skvortz., 456.

rugosa Skvortz., 456.

spinosa Skvortz., 456.

Protochroomonas Skvortz., 458.

granulata Skvortz., 459.

Quercus subgen. Cyclobalanopsis, a

revision, 355.

subgen. Cyclobalanopsis (Oersted)

Schneider, 379.

arbutifolia Warb., 397.

argentata Korth, 384.

brevistyla A. Camus, 376, 390.

chrysotrichia A. Camus, 397.

or eg sera (Gamble) A. Camus,

elmeri Merr., 414.

gaharuensis Soepadmo, 376, 389.

gemelliflora Bl., 378, 406.

hendersoniana A. Camus, 378, 409.

horsfieldii Miq., 406.

var. longifolia Miq., 406.

kerangasensis Soepad., 399.

kinabaluensis Soepad., 401.

lineata BI., 408.

var. heterochroa Miq., 408.

var. merkusii Wenz., 406.

var. oxyrhyncha (Miq.) v. Seem.,

409.

lineata S. Moore, 392.

lowii King, 376, 394.

merkusii Endl., 406.

merrillii von Seem., 397.

mespilifolia Wall. var. borneensis

Heine, 377, 404.

nivea King, 387.

oidocarpa Korth., 376, 390.

oidocarpa Hook. f., 389.

oidocarpa Merrill, 392.

oxyrhyncha Migq., 409.

percoriacea Soepad., 396

‘pinanga Blume, 384.

polyneura Migq., 408.

pseudoverticillata Soepadmo, 384.

semiserrata King, 406.

semiserrata Koord. & Val., 390.

semiserrata Roxb., 376.

474 Garden's Bulletin, Singapore — XXII (1968)

Quercus cont.:

sericea Scheff., 392

steenisii Soepad., 411.

subsericea A. Camus, 392.

sumatrana Soepadmo, 392.

treubiana von Seem., 412.

turbinata Blume, 406.

var. crassilamellata Gamble, 406.

valdinervosa Soepad., 404.

velutina Lindl., 379.

wilhelminae von Seem., 386.

Retinodendropsis Heim, 262.

Sagena decurrens (Presl) Houlst., 177.

mamillosa (C. Chr.) Moore, 177.

pteropus (Kze.) Bedd., 177.

Sapotaceae, Notes on, 213.

Sargassum, alginate production, 432.

Scaphium burkillfilii Kosterm., 444.

Schizomeria brassii Mattf., 5.

floribunda Schl., 5.

pulleana O. C. Schm., 5.

serrata Hochr., 5.

whitei Mattf., 5.

Sclereids, foliar, 461.

in Fagraea, 193.

Sczegleewia luconiensis Turcz., 163.

Shorea, Sec. Muticae Brandis, Ssec.

Muticae, 301.

Sec. Richetioides Heim, 288.

Ssec. Polyandrae Ashton, 288.

Ssec. Richetioides, 288.

agami Ashton, 285.

ssp. agami, 285.

ssp. diminuta Ashton. 285.

alutacea Ashton, 288.

bakoensis Ashton, 289.

brunnescens Ashton, 283.

carapae Ashton, 294.

cordata Ashton, 285.

cuspidata Ashton, 290.

foraminifera Ashton, 295.

grandiflora Brandis, 293.

hemsleyana King, 293.

ssp. grandiflora (Brandis) Ashton,

293.

ssp. hemsleyana, 293.

iliasii Ashton, 291.

johorensis Foxw., 294.

leptoclados Foxw., 294.

lunduensis Ashton, 284.

macrobalanos Ashton, 292.

meadiana Sym., 282.

mujongensis Ashton, 292.

pallidifolia Ashton, 296.

polyandra Ashton, 286.

praestans Ashton, 297,

Shorea cont.:

pubistyla Ashton, 297.

rotundifolia Ashton, 299.

sagittata Ashton, 299.

scrobiculata Burck, 282.

uliginosa Foxw., 294.

SINCLAIR, J., 213, 229. :

SKvorTzov, B. V., 187, 451, 455.

SOEPADMO, E., 355.

Steganthera suberosa-alata Kosterm..,

445

STONE, B. C., 129, 231.

Symphorema Roxb., 153.

cumingianum Briq., 163.

glabrum Hassk., 163.

involucratum Roxb., 165.

inneinernants Roxb. sens. Wall.,

168.

involucratum Spreng, 163.

luzonicum (Blanco) F. Vill., 162.

luzoniense (Turcz.) Benth., 163.

polyandrum Wight, 168.

Synaedrys sericea (Scheff.) Koidz.,

392.

Syzygium cinctum Merr. & Perry, 16.

onesimum Merr. & Perry, 17.

myriadenum Merr. & Perry, 16.

’ Tectaria decurrens (Presl) Copel., 177.

Tree ferns, 41.

Tristiropsis acutangula Radlk., 17.

dentata Radlk., 18.

obtusangula Radlk., 18.

Turbinaria, alginate content, 437.

TURNER, G. J., 123.

VAN ROYEN, P., 33.

Vatica aerea Sloot, 263. 264.

badiifolia Ashton, 266.

brunigii Ashton, 267.

compressa Ashton, 267.

congesta Ashton, 268.

globosa Ashton, 269.

mindanensis Foxw., 263.

oblongifolia Hook. f., 264.

ssp. crassilobata Ashton, 265.

ssp. elliptifolia Ashton, 265.

ssp. oblongifolia, 264.

ssp. multinervosa Ashton, 265.

ssp. Selakoensis Ashton, 264.

odorata (Griff.) Sym., 263.

ssp. mindanensis (Foxw.) Ashton,

263.

retata Ashton, 270.

rynchocarpa Ashton, 270.

sorsogonensis Foxw., 264.

WHITMoRE, T. C., 1.

OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

1. Annual Reports.

Many from 1909 onward remain available.

Prices variable.

2. The Agricultural Bulletin of the Malay Peninsula (Series 1.

Only Nos. 3, 5, 7, 8 and 9 available at 20 cents each.

3. The Agricultvral Bulletin of the Straits and F.M.S. (Series II).

Vols. 1-10, 1901-1911, monthly issues.

All parts available, except Vols. 1, 2, 5 (7), 6 (11) and 9.

Price: $5 per volume, Cents 50 per part.

4. The Gardens’ Bulletin, Straits Settlements (Series III).

Vols. 1-11, 1912-1947.

Vols. 1 (1-5) January-May 1912 is issued under title of

Agricultural Bulletin of the Straits & F.M.S.

All parts available, except Vol. 1 No. 2, 10, Vol. 2, Vol.

3 No. 1-3 Vol. 6, Vol. 10 pt. 2 and Vol. 11 pt. 3.

Prices variable.

5. The Gardens’ Bulletin, Singapore (Series IV).

Vols. 12-21, 1949 —*.

All parts available except 13 (1). Prices variable. Issues

are irregular.

Subscription will be entered against deposit.

Available on exchange.

6. Materials for a Flora of the Malay Peninsula, Monocotyledons.

Parts 1, 2 and 3 remain available.

7. (a) Malayan Orchid Hybrids by M. R. Henderson and G. H.

Addison.

Price $21.

(5) Malayan Orchid Hybrids, Supplement I by G. H. Addison.

Price $15 (Photo-lithographic reprint).

8. A Revised Flora of Malaya.

(a) Vol. 1, Orchids, by R. E. Holttum. Price $20.

(b) Vol. 2, Ferns, by R. E. Holttum. Price $20.

9. Wayside Trees of Malaya by E. J. H. Corner, 2 Vols.

Price $25.

{tems 1-6 can be obtained from the Director, Botanic Gardens,

Singapore.

Items 7-9 can be obtained from the Government Printer, Govern-

ment Printing Office, Singapore.

Prices quoted are in Singapore Dollars

Overseas postage is extra

0097 —S00—€/68